Affinage

WASL

Actin nucleation-promoting factor WASL · UniProt O00401

Length
505 aa
Mass
54.8 kDa
Annotated
2026-06-11
100 papers in source corpus 53 papers cited in narrative 53 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

N-WASP (WASL) is a ubiquitously expressed nucleation-promoting factor that converts upstream signals into branched actin polymerization by binding and activating the Arp2/3 complex through its C-terminal VCA domain (PMID:10219243, PMID:22847007). In the resting state the protein is held in an autoinhibited monomeric conformation by an intramolecular contact between its N-terminal region and the VCA effector domain that occludes the Arp2/3-binding site; this autoinhibition is relieved by active Cdc42 and PI(4,5)P2 acting through a conserved basic sequence near the Cdc42-binding site, which reduces the affinity between the N- and C-termini (PMID:10995436). A diverse set of inputs converge on this conformational switch: direct binding of Cdc42 generated by intersectin-1 GEF activity in a feed-forward loop (PMID:9422512, PMID:11584276), the WIP/Toca-1 module that constitutes the predominant cellular form of N-WASP (PMID:10878810, PMID:15260990), membrane-curvature- and phosphatidylserine-sensitive F-BAR proteins (Toca-1, FBP17, CIP4, amphiphysin 1) (PMID:18923421, PMID:19759398, PMID:20940394), Nck and PI(4,5)P2 acting interdependently (PMID:19917259), and additional activators including WISH, Abi1, IQGAP1 and Abp1 (PMID:11157975, PMID:16155590, PMID:17085436, PMID:17476322). Activity is further tuned by phosphorylation: FAK phosphorylates N-WASP at Tyr256, shifting it from nuclear to cytoplasmic localization and promoting migration (PMID:14676198), while HSP90 binding enhances Src-mediated phosphorylation and protects activated N-WASP from proteasomal degradation (PMID:15791211). Through this machinery N-WASP drives filopodium formation (PMID:9422512, PMID:11331876), endosome/lysosome and pathogen-driven actin comet motility (PMID:10662777, PMID:11584271, PMID:19262673), clathrin-mediated endocytosis and receptor trafficking (PMID:12426380, PMID:16155590, PMID:21610097), dendritic spine and synapse formation (PMID:18430734), invadopodium formation and tumor invasion/metastasis via MT1-MMP delivery and LPAR1 recycling (PMID:15684033, PMID:22389406, PMID:23091069, PMID:31668663), and is genetically required for muscle-cell fusion, Schwann-cell myelination, junctional and barrier integrity, and podocyte foot-process maintenance (PMID:21263026, PMID:24967734, PMID:22736793, PMID:23471198). N-WASP also operates in non-canonical, Arp2/3-independent modes: with nebulin it nucleates unbranched actin at myofibril Z-bands during IGF-1-induced hypertrophy (PMID:21148390), and it stabilizes junctional actin filaments via WIRE without directly activating Arp2/3 (PMID:21785420). In the nucleus, unphosphorylated N-WASP shuttles via an NLS/NES system and joins a PSF-NonO/RNA polymerase II complex where nuclear actin polymerization supports transcription (PMID:12871950, PMID:16767080).

Mechanistic history

Synthesis pass · year-by-year structured walk · 19 steps
  1. 1996 High

    Established N-WASP as a brain actin-regulatory protein whose membrane retention and cortical actin function depend on PIP2 binding and its actin-interacting domain, while also linking it to EGF receptor signaling.

    Evidence PH-domain mutagenesis, VCA deletion, EGFR co-IP and immunofluorescence in COS7 cells

    PMID:8895577

    Open questions at the time
    • Did not define the Arp2/3 connection
    • Mechanism of nuclear vs cortical partitioning unresolved
  2. 1998 High

    Showed that N-WASP acts downstream of active Cdc42 to drive filopodium formation via signal-dependent exposure of its actin-regulatory region, distinguishing it from WASP.

    Evidence Co-expression in cells, cell-free actin assay, Cdc42-binding experiments

    PMID:9422512

    Open questions at the time
    • Did not yet identify Arp2/3 as the effector
    • Molecular basis of autoinhibition not defined
  3. 1999 High

    Identified the VCA-Arp2/3 axis as the core nucleation mechanism and showed full-length activity is enhanced by Cdc42 and PI(4,5)P2, connecting signaling to actin assembly.

    Evidence Xenopus egg extract actin polymerization, VCA-Arp2/3 binding assays, immunodepletion

    PMID:10219243

    Open questions at the time
    • Structural basis of autoinhibition not yet shown
    • How multiple activators are integrated unclear
  4. 2000 High

    Defined the intramolecular autoinhibition mechanism and showed PI(4,5)P2 and Cdc42 relieve it via a basic sequence element, providing the central conformational switch model.

    Evidence In vitro polymerization, hydrodynamics, domain-binding assays, basic-stretch mutants in Xenopus extracts

    PMID:10995436

    Open questions at the time
    • Kinetics of switching in cells not measured
    • Did not address phosphorylation-based regulation
  5. 2000 High

    Demonstrated N-WASP drives Arp2/3-dependent vesicle propulsion (actin comet tails) and that the WH1 domain recruits it to actin-assembly sites via WIP, establishing the WIP partnership.

    Evidence Live imaging in Xenopus eggs, cell-free reconstitution, mutant/co-IP analysis in pathogen motility

    PMID:10662777 PMID:10878810

    Open questions at the time
    • Stoichiometry of the WIP-N-WASP complex not resolved
    • How recruitment integrates with activation unclear
  6. 2001 High

    Genetic knockout established N-WASP as embryonically essential and pathogen-selective (required for Shigella/vaccinia but not Listeria motility), while expanding the activator network (WIP, WISH, intersectin, TC10/GLUT4).

    Evidence Gene targeting, pathogen motility assays, in vitro reconstitution, GEF assays, dominant-negative GLUT4 trafficking

    PMID:11157975 PMID:11331876 PMID:11584271 PMID:11584276 PMID:11694514

    Open questions at the time
    • Functional redundancy with WASP not yet defined
    • How distinct activators are spatially deployed unknown
  7. 2002 Medium

    Extended N-WASP function to membrane trafficking — Golgi-to-ER transport, PIP2-driven endomembrane comets, and syndapin-linked receptor endocytosis — broadening it beyond filopodia.

    Evidence GFP-localization, transport/endocytosis assays, mutant reconstitution, co-IP in cells

    PMID:11907268 PMID:12147689 PMID:12426380

    Open questions at the time
    • Single-lab functional transport claims
    • Direct vs indirect contribution to each trafficking step unresolved
  8. 2003 Medium

    Revealed phosphorylation-based and nuclear regulation: FAK phosphorylates Tyr256 to shift N-WASP cytoplasmic, and Src-family phosphorylation controls NLS/NES shuttling, with nuclear N-WASP repressing HSP90 transcription.

    Evidence In vitro kinase assay, fractionation, LMB treatment, importin co-IP, transcription reporters, ChIP

    PMID:12871950 PMID:14676198

    Open questions at the time
    • Nuclear transcriptional role rests on single-lab data
    • Physiological significance of HSP90 repression unclear
  9. 2004 High

    Established N-WASP/Arp2/3 as the engine of invadopodium formation and matrix degradation in carcinoma cells, and visualized spatially restricted N-WASP activation by FRET, linking conformational state to subcellular site.

    Evidence RNAi, dominant-negative, time-lapse and matrix-degradation assays, conformational FRET biosensor; Toca-1 purification and mDab1 activation

    PMID:15084285 PMID:15260990 PMID:15361067 PMID:15684033

    Open questions at the time
    • In vivo metastatic relevance not yet tested
    • Quantitative link between activation site and invasion output unmeasured
  10. 2005 High

    Defined HSP90 as a stabilizer of activated N-WASP and added Abi1 as a high-affinity activator coupling N-WASP to vesicular transport and EGFR endocytosis.

    Evidence Direct binding, in vitro kinase/polymerization assays, proteasome rescue, nanomolar affinity measurement, RNAi with trafficking readouts

    PMID:15791211 PMID:16155590

    Open questions at the time
    • How HSP90 selectively recognizes the active conformation unclear
    • Crosstalk between WAVE-complex and N-WASP regulation not resolved
  11. 2006 Medium

    Provided evidence for a nuclear N-WASP complex with PSF-NonO and RNA polymerase II in which nuclear actin polymerization supports transcription.

    Evidence Nuclear complex co-IP, RNA Pol II co-IP, transcription reporters, fractionation

    PMID:16767080

    Open questions at the time
    • Single-lab observation
    • Target genes and physiological scope of nuclear function undefined
  12. 2007 High

    Resolved biochemical and structural detail of the effector module — VCA-Arp2/3-actin stoichiometry and the multi-epitope WIP-EVH1 interface — and extended N-WASP function to dendritic spine/synapse formation and T-cell development (redundant with WASP).

    Evidence Crystallography/NMR, hydrodynamics, spectrofluorimetry, RNAi/inhibitor/dominant-negative neuronal assays, double-knockout immunology

    PMID:17229736 PMID:17476322 PMID:17878299 PMID:18430734 PMID:22847007

    Open questions at the time
    • Full-length activated structure not solved
    • Tissue-specific division of labor with WASP not fully mapped
  13. 2008 High

    Showed membrane curvature directly couples to N-WASP activation: F-BAR proteins Toca-1/FBP17 recruit and open the N-WASP-WIP complex on curved phosphatidylserine membranes independent of Cdc42 and PIP2.

    Evidence Defined-lipid in vitro reconstitution with curvature control, mutagenesis; conformation-sensitive antibody in Shigella infection

    PMID:18191793 PMID:18923421

    Open questions at the time
    • Relative contribution of curvature vs GTPase input in cells unquantified
    • Toca-1 recruitment mechanism by pathogen effectors single-lab
  14. 2009 High

    Demonstrated that N-WASP turnover dynamics set the rate of actin-based motility and that Nck and PI(4,5)P2 are reciprocally required for localized activation; amphiphysin 1 added as a BAR-domain activator.

    Evidence FRAP with mutants, motility measurements, Nck KO/RNAi and lipid manipulation, knockout-cytosol and FRET-FLIM

    PMID:19262673 PMID:19759398 PMID:19917259

    Open questions at the time
    • How turnover rate is tuned in physiological contexts unclear
    • Integration of multiple BAR activators not reconciled
  15. 2011 High

    Uncovered Arp2/3-independent functions: with nebulin N-WASP nucleates unbranched actin at Z-bands for muscle hypertrophy, and at the zonula adherens it stabilizes junctional filaments via WIRE without activating Arp2/3.

    Evidence Co-IP, in vitro nucleation with/without Arp2/3, conditional muscle KO; RNAi with Arp2/3-deficient rescue mutant and WIRE binding

    PMID:21148390 PMID:21263026 PMID:21610097 PMID:21785420

    Open questions at the time
    • Structural basis of unbranched nucleation undefined
    • How N-WASP toggles between branched and non-branched modes unknown
  16. 2012 High

    Defined physiological requirements across tissues — muscle fusion, blood-testis barrier, podocyte foot processes, B-cell receptor signaling, endothelial junctions — and established N-WASP-driven invadopodia and MT1-MMP delivery as drivers of breast cancer metastasis in vivo.

    Evidence Multiple tissue-specific conditional knockouts, EM, permeability/filtration assays, TIRF/signaling, shRNA + dominant-negative with in vivo metastasis and MT1-MMP trafficking

    PMID:22389406 PMID:22736793 PMID:23091069 PMID:23212915 PMID:23471198 PMID:24223520 PMID:24967734

    Open questions at the time
    • Mechanistic links between conformational regulation and each tissue phenotype incomplete
    • Some junction/endothelial claims single-lab
  17. 2014 High

    Placed N-WASP genetically downstream of Cdc42 in developmental morphogenesis (pancreatic beta-cell delamination/differentiation) and added the PC1-Pacsin2 complex linking it to directional migration.

    Evidence In vivo double-mutant rescue genetics; yeast two-hybrid, co-IP and migration assays

    PMID:24385601 PMID:24449844

    Open questions at the time
    • Direct effectors mediating delamination not identified
    • PC1-Pacsin2 complex relevance single-lab
  18. 2019 High

    Identified a chemotaxis/mechanotransduction role: N-WASP with SNX18 controls LPAR1 recycling to sustain RhoA-mediated contractility, force generation and pancreatic cancer metastasis.

    Evidence RNAi, receptor trafficking, RhoA activation, traction force microscopy, in vivo metastasis, SNX18 co-IP

    PMID:31668663

    Open questions at the time
    • Generality of LPAR1-recycling role beyond this cancer model untested
  19. 2021 Medium

    Showed that pathogen effectors directly hijack N-WASP: Chlamydia TmeA binds and activates N-WASP for Arp2/3-dependent entry, synergizing with a parallel TarP pathway.

    Evidence Chlamydial gene deletion, proximity labeling, direct binding and actin assays, infection assays

    PMID:33468693

    Open questions at the time
    • Single-lab effector characterization
    • Structural basis of TmeA-N-WASP activation undefined

Open questions

Synthesis pass · forward-looking unresolved questions
  • How the many competing activators, inhibitors, phosphorylation events, BAR proteins and nuclear functions are integrated into a single spatiotemporal output, and how N-WASP switches between branched (Arp2/3) and unbranched nucleation modes, remains unresolved.
  • No unified quantitative model of activator competition
  • Structural basis of mode-switching between branched and unbranched nucleation unknown
  • Physiological scope of nuclear/transcriptional function undefined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0008092 cytoskeletal protein binding 3 GO:0008289 lipid binding 3 GO:0060090 molecular adaptor activity 3 GO:0098772 molecular function regulator activity 3 GO:0140110 transcription regulator activity 2
Localization
GO:0005634 nucleus 3 GO:0005856 cytoskeleton 3 GO:0005886 plasma membrane 3 GO:0031410 cytoplasmic vesicle 3 GO:0005829 cytosol 2 GO:0005794 Golgi apparatus 1
Pathway
R-HSA-1266738 Developmental Biology 3 R-HSA-162582 Signal Transduction 3 R-HSA-1643685 Disease 3 R-HSA-5653656 Vesicle-mediated transport 3 R-HSA-168256 Immune System 2 R-HSA-112316 Neuronal System 1
Complex memberships
N-WASP-WIP complexPSF-NonO/RNA polymerase II nuclear complexnebulin-N-WASP Z-band complex

Evidence

Reading pass · 53 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1996 N-WASP was identified as a 65 kDa brain protein that binds the SH3 domains of Ash/Grb2 and contains a pleckstrin homology (PH) domain and cofilin-homologous region through which it depolymerizes actin filaments. PH domain mutation (C38W) that reduces PIP2 binding and deletion of the VCA actin-binding domain both abolish cortical actin rearrangements and cause predominantly nuclear localization, establishing that PIP2 binding and actin interaction are required for membrane retention and function. EGF treatment induces complex formation of EGF receptors with N-WASP and produces microspikes. Mutagenesis (C38W PH domain mutation, deltaVCA deletion), overexpression in COS7 cells, co-immunoprecipitation with EGF receptor, subcellular localization by immunofluorescence The EMBO journal High 8895577
1998 N-WASP induces extremely long actin microspikes only when co-expressed with active Cdc42, whereas WASP does not, despite structural similarities. In a cell-free system, active Cdc42 stimulates the actin-depolymerizing activity of N-WASP by exposing its actin-depolymerizing region. N-WASP directly interacts with Cdc42 and is required downstream of Cdc42 for filopodium formation. Co-expression in cells, cell-free actin polymerization assay, Cdc42-binding experiments Nature High 9422512
1999 N-WASP is required for Cdc42-stimulated actin polymerization in Xenopus egg extracts. The C terminus of N-WASP (VCA domain) binds directly to the Arp2/3 complex and dramatically stimulates its actin nucleation activity. Full-length N-WASP activity is greatly enhanced by Cdc42 and PI(4,5)P2, linking signal transduction to actin polymerization through an N-WASP/Arp2/3 core mechanism. In vitro actin polymerization assay in Xenopus egg extracts, biochemical binding assays (VCA–Arp2/3 interaction), immunodepletion Cell High 10219243
2000 The N-terminal domain of N-WASP physically interacts with its C-terminal effector (VCA) domain in an intramolecular, autoinhibitory interaction that occludes the Arp2/3-binding site. N-WASP is a monomer in solution. PI(4,5)P2 activates N-WASP through a conserved basic sequence element near the Cdc42-binding site (not the WH1 domain), reducing the affinity between N- and C-termini. Cdc42 similarly relieves autoinhibition. In Xenopus extracts, PI(4,5)P2 acts both as a direct N-WASP activator and as an indirect activator of Cdc42. In vitro actin polymerization assay, sedimentation/gel filtration (monomer determination), domain-binding assays, mutant N-WASP lacking basic stretch in Xenopus extracts The Journal of cell biology High 10995436
2000 N-WASP is recruited to the surface of endosomes and lysosomes that form actin comet tails in Xenopus eggs and in mammalian cell extracts, mediating vesicle propulsion through Arp2/3-complex-dependent actin assembly. Live imaging in Xenopus eggs, cell-free reconstitution, immunofluorescence, electron microscopy, acridine orange staining The Journal of cell biology High 10662777
2000 The WH1 domain (not the polyproline-rich region) of N-WASP mediates its recruitment to sites of actin polymerization during vaccinia virus motility via direct interaction with WASP-interacting protein (WIP). N-WASP and WIP form a functional complex that integrates signaling cascades leading to actin polymerization. In Shigella motility, WIP is recruited by N-WASP. Mutant expression, co-immunoprecipitation, actin comet tail assays in vaccinia/Shigella-infected cells Nature cell biology High 10878810
2001 WIP directly interacts with N-WASP and actin. WIP retards N-WASP/Cdc42-activated actin polymerization mediated by the Arp2/3 complex and stabilizes actin filaments. WIP and N-WASP act as a functional unit in filopodium formation: anti-N-WASP antibody inhibits WIP-induced filopodia, and anti-WIP antibody blocks N-WASP-induced filopodia. In vitro actin polymerization assay (pyrene-actin), direct binding assay, microinjection of antibodies into NIH 3T3 cells Nature cell biology High 11331876
2001 N-WASP knockout mice die before embryonic day 12 with developmental delay. N-WASP is not required for Listeria actin-based motility but is absolutely required for Shigella and vaccinia virus actin-based motility. N-WASP-deficient fibroblasts can still form filopodia and spread via lamellipodia. Gene targeting (homologous recombination), genetic knockout, pathogen actin motility assays, cell spreading/morphology analysis Nature cell biology High 11584271
2001 Intersectin-l (neuronal variant) functions via its DH domain as a GEF for Cdc42. N-WASP binds directly to intersectin-l and upregulates its GEF activity, generating GTP-bound Cdc42 which in turn activates N-WASP, creating a feed-forward activation loop that drives actin assembly via the Arp2/3 complex. GEF activity assay (GDP/GTP exchange), direct binding assay, co-immunoprecipitation, actin polymerization assay, cell-based actin rearrangement analysis Nature cell biology High 11584276
2001 A novel adaptor protein WISH binds N-WASP through its SH3 domain and strongly enhances N-WASP-induced Arp2/3 complex activation independent of Cdc42 in vitro. WISH coexpression with N-WASP induces marked microspike formation even without stimuli; an N-WASP mutant (H208D) that cannot bind Cdc42 still induces microspikes with WISH. In vitro actin polymerization assay, co-immunoprecipitation, overexpression in COS7 cells, N-WASP depletion from brain extracts The Journal of cell biology High 11157975
2001 N-WASP is involved in insulin-stimulated GLUT4 recycling. Insulin causes PI3K-independent cortical localization of N-WASP and Arp3 plus cortical F-actin polymerization in adipocytes. Dominant-inhibitory N-WASP-DeltaWA attenuates cortical F-actin rearrangements and inhibits insulin-stimulated GLUT4 translocation. TC10 (a Cdc42-related GTPase) acts upstream of N-WASP in this pathway; inhibitory TC10 (T31N) blocks cortical N-WASP localization. Dominant-negative expression, immunofluorescence localization, GLUT4 translocation assay in adipocytes, PI3K inhibitor (wortmannin) treatment The Journal of biological chemistry High 11694514
2002 Cdc42 regulates Golgi-to-ER protein transport through N-WASP. Cdc42V12 recruits GFP-N-WASP to the Golgi complex. Coexpression of Cdc42 and N-WASP inhibits retrograde Golgi-to-ER transport; this inhibition requires the Arp2/3-binding WA domain of N-WASP, as the N-WASP(ΔWA) mutant does not inhibit transport. Overexpression, GFP-N-WASP localization imaging, Shiga toxin retrograde transport assay, KDEL receptor redistribution assay, dominant-active Sar1 assay Molecular biology of the cell Medium 11907268
2002 N-WASP is essential for actin assembly at the surface of endomembranes induced by elevated PIP2 levels, leading to actin comet-driven vesicle motility. This process requires WH1 and polyproline domains of N-WASP for vesicle surface recruitment/activation, and Nck, Grb2, and WIP are also recruited. Direct interaction of N-WASP with Cdc42 is not required for reconstitution of vesicle motility. N-WASP-deficient cell reconstitution, N-WASP mutant expression, vesicle motility assay, co-immunoprecipitation The Journal of biological chemistry High 12147689
2002 Syndapins interact with N-WASP through its proline-rich domain and integrate N-WASP functions in receptor-mediated endocytosis. Co-overexpression of syndapins rescues the endocytosis block caused by N-WASP dominant-negative. Depletion of endogenous N-WASP by sequestration to mitochondria or anti-N-WASP antibodies impairs endocytosis. In vivo reconstitution of the syndapin-N-WASP interaction at cellular membranes triggered local actin polymerization. Co-overexpression rescue assay, N-WASP depletion via mitochondrial targeting and antibody microinjection, endocytosis assay, in vivo reconstitution at membranes The EMBO journal High 12426380
2003 FAK directly interacts with N-WASP and phosphorylates it at Tyr256. Phosphorylation of Tyr256 reduces N-WASP interaction with nuclear importin NPI-1, shifting N-WASP from nuclear to cytoplasmic localization. Nuclear localization of N-WASP also depends on being in the open conformation (Cdc42 activation or VCA truncation). Tyr256 phosphorylation promotes cell migration. In vitro kinase assay (FAK phosphorylation of N-WASP), co-immunoprecipitation, subcellular fractionation/localization, co-immunoprecipitation with importin NPI-1, cell migration assay The Journal of biological chemistry High 14676198
2003 N-WASP localizes to the nucleus and its nuclear/cytoplasmic shuttling is controlled by phosphorylation by Src family kinases. Phosphorylated N-WASP is exported from the nucleus via a nuclear export signal (NES) in a leptomycin B-sensitive manner; N-WASP also has a nuclear localization signal (NLS) in its basic region. Unphosphorylated nuclear N-WASP suppresses HSP90 expression by binding heat shock transcription factor (HSTF) and enhancing HSTF association with heat shock element (HSE). Reduced HSP90 in turn decreases Src kinase activity. Subcellular fractionation, leptomycin B treatment (NES validation), NLS/NES identification, co-immunoprecipitation with HSTF, luciferase/transcription reporter assays, chromatin immunoprecipitation The Journal of biological chemistry Medium 12871950
2004 Toca-1 (transducer of Cdc42-dependent actin assembly) was biochemically purified as an essential component of the Cdc42/N-WASP pathway. Toca-1 binds both N-WASP and Cdc42. Toca-1 promotes actin nucleation by activating the N-WASP-WIP complex (the predominant cellular form of N-WASP), and cooperative actions of both N-WASP-WIP and Toca-1 are required for Cdc42-induced actin assembly. Biochemical purification, in vitro actin polymerization assay, binding assays (Toca-1 with N-WASP and Cdc42), Xenopus egg extract reconstitution Cell High 15260990
2004 mDab1 directly binds N-WASP via the PTB domain of mDab1 interacting with the NRFY sequence near the CRIB motif of N-WASP, and directly activates N-WASP to induce Arp2/3-mediated actin polymerization and filopodium formation in cells. This filopodium formation depends on N-WASP activity. Fyn kinase-mediated phosphorylation of mDab1 leads to its Cbl-dependent ubiquitination and loss of filopodium induction, acting as a negative regulatory switch. Direct binding assay (in vitro), in vitro actin polymerization assay, overexpression in COS-7 cells, dominant-negative N-WASP rescue, phosphorylation and ubiquitination assays The Biochemical journal Medium 15361067
2004 N-WASP and the Arp2/3 complex are required for invadopodium formation in metastatic carcinoma cells. N-WASP is activated at the base of invadopodia. Upstream regulators Nck1, Cdc42, and WIP are also necessary. Cofilin is required for stabilization and maturation of long-lived invadopodia. EGF receptor signaling drives invadopodium formation through the N-WASP-Arp2/3 pathway. RNAi, dominant-negative mutant expression, time-lapse microscopy, EGFR kinase inhibitors, matrix degradation assay The Journal of cell biology High 15684033
2004 N-WASP activity is spatially regulated in living cells: N-WASP is activated at the leading edge of lamellipodia and at the base of invadopodia in invasive carcinoma cells, as demonstrated by a FRET biosensor distinguishing active (open) vs. inactive (closed) N-WASP conformations. FRET biosensor (N-WASP conformational sensor) in live cells Current biology Medium 15084285
2005 HSP90 binds directly to N-WASP. Binding alone does not affect basal actin polymerization rate, but HSP90 enhances v-Src-mediated phosphorylation of N-WASP, leading to increased actin polymerization. HSP90 also protects phosphorylated/activated N-WASP from proteasome-dependent degradation, amplifying N-WASP activity. HSP90-N-WASP association is increased proportional to N-WASP activation. Blocking HSP90 binding to N-WASP inhibits podosome formation and neurite extension. Direct binding assay (pull-down), in vitro actin polymerization assay, phosphorylation assay (v-Src), proteasome inhibitor rescue, co-immunoprecipitation at podosomes, wiskostatin/HSP90 inhibitor treatment The EMBO journal High 15791211
2005 Abi1, an essential component of the WAVE protein complex, also binds N-WASP with nanomolar affinity and cooperates with Cdc42 to potently stimulate N-WASP activity in vitro. Abi1 and N-WASP (but not WAVE) regulate actin-based vesicular transport, EGFR endocytosis, and EGFR/TfR cell-surface distribution. In vitro actin polymerization assay, direct binding affinity determination, RNAi knockdown, EGFR endocytosis assay, receptor surface distribution analysis Nature cell biology High 16155590
2006 N-WASP is present in the nucleus within a large protein complex containing PSF-NonO, nuclear actin, and RNA polymerase II. N-WASP interacts with the PSF-NonO complex and couples it to RNA polymerase II to regulate transcription. Nuclear actin polymerization promoted by N-WASP plays an important role in this transcriptional regulation. Co-immunoprecipitation of nuclear complex, RNA polymerase II co-IP, transcription reporter assays, nuclear fractionation Nature cell biology Medium 16767080
2006 IQGAP1 controls co-localization of N-WASP with the Arp2/3 complex in lamellipodia. The C-terminal half of IQGAP1 activates N-WASP by interacting with its BR-CRIB domain in a Cdc42-like manner; the N-terminal half of IQGAP1 antagonizes this by associating with the C-terminal region of IQGAP1 (autoinhibition). Signal-induced relief of IQGAP1 autoinhibition allows it to activate N-WASP for Arp2/3-dependent actin assembly. Quantitative co-localization, IQGAP1 downregulation, co-immunoprecipitation, pull-down with N-WASP domains, kinetic actin polymerization assay The Journal of biological chemistry High 17085436
2007 WASP and N-WASP have combined, partially redundant roles in T cell development. Double knockout (lacking both WASP and N-WASP) in T cells causes thymic hypocellularity, reduced peripheral T cells, impaired DN-to-DP transition with reduced cycling DN3 cells, and decreased SP cell migration. N-WASP single deficiency in T cells is indistinguishable from wild-type. Homologous recombination plus conditional Cre-loxP knockout, RAG-2-deficient blastocyst complementation, flow cytometry, migration assays Proceedings of the National Academy of Sciences High 17878299
2007 N-WASP and the Arp2/3 complex regulate the formation of dendritic spines and synapses in hippocampal neurons. N-WASP localizes to spines and active synapses. RNAi knockdown or wiskostatin treatment decreases spine and excitatory synapse number. Deletion of the C-terminal VCA domain that binds/activates Arp2/3 dramatically decreases spines and synapses. Cdc42 knockdown phenocopies N-WASP knockdown, placing Cdc42 upstream. RNAi knockdown, wiskostatin pharmacological inhibition, VCA deletion mutant, FM4-64 dye loading (functional synapse marking), immunofluorescence co-localization The Journal of biological chemistry High 18430734
2007 Abp1 (F-actin-binding protein) directly interacts with N-WASP and releases N-WASP autoinhibition in cooperation with Cdc42, promoting N-WASP-triggered Arp2/3-mediated actin polymerization. Abp1 knockdown in neurons increases axon length, phenocopying Arp2/3 complex inhibition. Abp1, N-WASP and Arp2/3 colocalize at actin polymerization sites in neurons. Direct interaction assay (in vitro pull-down), in vitro actin polymerization assay, Abp1 RNAi knockdown, N-WASP mutants lacking Abp1 or Cdc42 binding, immunofluorescence PLoS ONE Medium 17476322
2008 EFC/F-BAR domain proteins (Toca-1 and FBP17) activate the N-WASP-WIP complex-mediated actin polymerization depending on membrane curvature in the presence of phosphatidylserine-containing membranes, even in the absence of Cdc42 and PIP2. Toca-1/FBP17 recruit N-WASP-WIP to the membrane and position it spatially via conserved acidic residues near their SH3 domain. In vitro actin polymerization assay with defined lipid vesicles of varying curvature, mutant analysis of acidic residues, N-WASP-WIP recruitment assay The EMBO journal High 18923421
2008 Toca-1 is required in intact mammalian cells for the conversion of N-WASP from a closed (inactive) to an open (active) conformation during Shigella actin tail initiation. N-WASP recruitment to Shigella is dependent on the bacterial IcsA, whereas Toca-1 recruitment is mediated by type III secretion effectors, showing two independently hijacked nodes of the N-WASP actin assembly pathway. Toca-1 RNAi knockdown, conformation-sensitive N-WASP antibody assay, cell infection assays, fluorescence microscopy Cell host & microbe Medium 18191793
2009 N-WASP exchange rate limits the extent of Arp2/3-dependent actin-based motility of vaccinia virus. N-WASP rapidly turns over at the virus surface (FRAP), and its turnover depends on its ability to stimulate Arp2/3 actin polymerization. Disrupting N-WASP interaction with Grb2 or barbed ends increases N-WASP exchange rate and results in faster virus movement. N-WASP thus controls the rate of actin-based motility by regulating actin polymerization extent. FRAP (fluorescence recovery after photobleaching), N-WASP mutant analysis, vaccinia actin motility assay Nature High 19262673
2009 Amphiphysin 1 directly interacts with N-WASP and stimulates N-WASP- and Arp2/3-dependent actin polymerization. Both the SH3 and N-BAR domains of amphiphysin 1 are required for stimulation. Acidic liposome-triggered N-WASP-dependent actin polymerization is strongly impaired in amphiphysin 1 knockout mouse brain cytosol. FRET-FLIM confirmed the association in vivo in Sertoli cells; association is enhanced by phosphatidylserine receptor stimulation. Direct binding assay, in vitro actin polymerization assay, amphiphysin 1 knockout mouse brain cytosol, FRET-FLIM in cells The Journal of biological chemistry High 19759398
2009 Nck and PI(4,5)P2 show reciprocal interdependence in promoting localized N-WASP-mediated actin polymerization. Nck knockdown/knockout suppresses PIP5K-induced actin comets. PI(4,5)P2 is necessary for localized Nck-induced actin polymerization. PI(4,5)P2 and PIP5K are enriched at Nck-induced actin comets. The extent of N-WASP-mediated actin polymerization is modulated by PI(4,5)P2-sensitive N-WASP mutants. Nck RNAi knockdown, Nck knockout cells, PIP5K overexpression, inositol 5-phosphatase coclustering, N-WASP mutant analysis, live-cell imaging of actin comets Molecular cell High 19917259
2010 Nebulin and N-WASP form a complex at Z bands of myofibrils upon IGF-1 stimulation, downstream of PI3K-Akt signaling through inhibition of GSK-3β. Importantly, the nebulin-N-WASP complex promotes unbranched actin filament nucleation from Z bands without Arp2/3 complex, representing a non-canonical Arp2/3-independent function of N-WASP. N-WASP is required for IGF-1-induced muscle hypertrophy. Co-immunoprecipitation, in vitro actin polymerization assay (with and without Arp2/3), IGF-1 stimulation, GSK-3β inhibitor treatment, N-WASP conditional knockout in muscle Science High 21148390
2011 N-WASP regulates the epithelial junctional actin cytoskeleton through a nucleation-independent pathway at the zonula adherens. N-WASP depletion decreases junctional F-actin but does not affect junctional actin nucleation (dominantly mediated by Arp2/3). An N-WASP mutant unable to directly activate Arp2/3 rescues the junctional defect. N-WASP stabilizes newly formed actin filaments via the WIP-family protein WIRE, which binds the N-WASP WH1 domain. RNAi knockdown, rescue with Arp2/3-activation-deficient N-WASP mutant, WIRE binding assay, junctional F-actin quantification Nature cell biology High 21785420
2011 N-WASP is required for membrane wrapping and myelination by Schwann cells. Schwann cell-specific N-WASP knockout mice fail to myelinate (cells arrest at promyelinating stage); a limited number form unusually short internodes with thin myelin and occasional myelin misfoldings. Schwann cells can sort and ensheath axons without N-WASP. Conditional knockout (Schwann cell-specific Cre-loxP), electron microscopy, histological analysis, nerve morphology assessment The Journal of cell biology High 21263026
2012 N-WASP is an essential negative regulator of B cell receptor (BCR) signaling. B-cell-specific N-WASP deletion causes enhanced and prolonged BCR signaling, elevated autoantibodies, increased F-actin at the B-cell surface, enhanced spreading, delayed contraction, inhibition of BCR microcluster merging into central clusters, and blockage of BCR internalization. WASP is activated first upon BCR activation, followed by N-WASP; N-WASP activation is suppressed by Bruton's tyrosine kinase-induced WASP activation and restored by SHIP-mediated WASP inactivation. B-cell-specific conditional N-WASP knockout, TIRF microscopy, actin imaging, BCR cluster analysis, serum autoantibody measurement, signaling assays PLoS biology High 24223520
2012 N-WASP-mediated invadopodium formation is essential for breast cancer invasion, intravasation and lung metastasis in vivo. Both N-WASP shRNA and dominant-negative N-WASP constructs decrease invadopodium formation, extracellular matrix degradation, tumor intravasation, and lung metastasis in a rat mammary tumor model. Stable shRNA knockdown, dominant-negative expression, intravital imaging, lung metastasis counting, in vivo tumor intravasation assay Journal of cell science High 22389406
2012 N-WASP coordinates delivery of MT1-MMP to invasive pseudopods from late endosomes and stabilizes MT1-MMP at the plasma membrane via direct tethering of its cytoplasmic tail to F-actin. N-WASP promotes assembly of elongated pseudopodia required for matrix degradation in 3D. Co-immunoprecipitation (N-WASP with MT1-MMP), live-cell trafficking assays, dominant-negative N-WASP, 3D matrix invasion assay, immunofluorescence The Journal of cell biology High 23091069
2012 N-WASP is required for structural integrity of the blood-testis barrier (BTB). Sertoli cell-specific N-WASP knockout leads to mislocalization of junctional/cytoskeletal elements, disruption of BTB function, and complete spermatogenic arrest. N-WASP-Arp2/3 actin polymerization machinery generates branched-actin arrays at an advanced stage of BTB remodeling, mediating restructuring through endocytic recycling of BTB components. Sertoli cell-specific conditional knockout, electron microscopy, junction protein localization, BTB permeability assay PLoS genetics High 24967734
2012 N-WASP binds p120-catenin through its VCA domain and links p120-catenin to the Arp2-cortical actin polymerization machinery to stabilize endothelial adherens junctions. This interaction requires Tyr256 phosphorylation of N-WASP by FAK. Phosphomimicking Y256D-N-WASP or VCA expression stabilizes junctions and facilitates barrier recovery after thrombin. Co-immunoprecipitation, N-WASP depletion, VCA domain expression, phosphomimetic mutant (Y256D), endothelial permeability assay, actin imaging The Journal of biological chemistry Medium 23212915
2012 N-WASP is required for muscle-cell fusion during mouse skeletal myogenesis. N-WASP-deficient myoblasts fail to fuse but otherwise differentiate normally, maintain motility, and attach normally. N-WASP activity is required in both partners of a fusing myoblast pair. Conditional N-WASP knockout in muscle, primary satellite cell cultures, cell fusion quantification, motility and morphology assays Proceedings of the National Academy of Sciences High 22736793
2013 WIP acts as an essential link between Nck and N-WASP. WIP (or its homolog WIRE) is required for N-WASP recruitment and actin-based motility of vaccinia virus. WIP contains two Nck-binding sites; it is recruited to the virus by the second SH3 domain of Nck. N-WASP's recruitment depends on its interaction with WIP rather than directly with Nck. The first and third SH3 domains of Nck stimulate actin assembly but are not required for WIP-N-WASP recruitment. MEF knockouts (Nck, WIP, N-WASP), vaccinia actin motility assay, co-immunoprecipitation, domain-specific mutant analysis Current biology High 23707428
2013 Cdc42 cooperates with Nck to promote actin tail formation by stabilizing N-WASP beneath vaccinia virus. Cdc42 activation is mediated by the Rho-GEF intersectin-1 (ITSN1), which is recruited to the virus before actin-based motility. Cdc42, ITSN1, and N-WASP function in a feed-forward loop to promote actin polymerization. This pathway also operates in FcγR-mediated phagocytosis. RNAi knockdown, co-immunoprecipitation, vaccinia actin tail assay, phagocytosis assay, genetic epistasis Journal of cell science Medium 24284073
2013 N-WASP is required for stabilization of podocyte foot processes. Podocyte-specific N-WASP knockout mice develop proteinuria and kidney failure. N-WASP-deficient podocytes show impaired dynamic actin reorganization (dorsal ruffle formation). N-WASP-mediated Arp2/3 actin nucleation of branched microfilament networks is specifically required for foot process maintenance. Podocyte-specific conditional N-WASP knockout, electron microscopy of foot processes, proteinuria measurement, primary culture actin dynamics assay Journal of the American Society of Nephrology High 23471198
2014 Cdc42/N-WASP signaling controls β cell delamination and differentiation during pancreatic development. Expression of constitutively active Cdc42 inhibits β cell delamination and differentiation associated with junctional actin and cell-cell junction disassembly. Genetic ablation of N-WASP in constitutively active Cdc42-expressing β cells partially restores both delamination and β cell differentiation, placing N-WASP downstream of Cdc42 in this process. Conditional mouse genetics (Cre-loxP), constitutively active Cdc42 expression, N-WASP conditional knockout, immunofluorescence of junction proteins and differentiation markers Development High 24449844
2014 PC1 (polycystin-1), Pacsin 2, and N-WASP are in the same protein complex. Both PC1 and Pacsin 2 are required for N-WASP/Arp2/3-dependent actin remodeling and directional cell migration in kidney epithelial cells. Yeast two-hybrid, co-immunoprecipitation, PC1/Pacsin2 siRNA knockdown, directional migration assay, actin remodeling assay Human molecular genetics Medium 24385601
2019 N-WASP drives pancreatic cancer metastasis through chemotaxis and matrix remodeling. N-WASP and the endocytic adapter SNX18 promote lysophosphatidic acid (LPA)-induced RhoA-mediated contractility and force generation by controlling LPA receptor (LPAR1) recycling and preventing its degradation. N-WASP-depleted cells do not recognize LPA gradients, showing altered RhoA activation, decreased contractility and traction forces, and reduced metastasis. N-WASP depletion (RNAi), LPAR1 trafficking assay (receptor recycling vs. degradation), RhoA activation assay, traction force microscopy, in vivo metastasis model, co-immunoprecipitation with SNX18 Developmental cell High 31668663
2010 CIP4 (Cdc42 interacting protein-4), an F-BAR protein, interacts with N-WASp in an EGF-dependent manner. CIP4 silencing causes decreased tyrosine phosphorylation of N-WASp at the Src-dependent site Y256, impairs invadopodium formation and gelatin degradation, and reduces migration and invasion. Co-immunoprecipitation, siRNA knockdown of CIP4, invadopodium assay, phospho-Y256 N-WASP Western blot, invasion/migration assays Cancer research Medium 20940394
2010 CIP4 promotes GLUT4 endocytosis by interacting with both N-WASp and Dynamin-2 in an insulin-dependent manner. Knockdown of CIP4 increases surface GLUT4 by decreasing endocytosis. FRET confirmed insulin-dependent subcellular coordination of CIP4-N-WASp and CIP4-Dynamin-2 interactions at the plasma membrane and in cytosol. Co-immunoprecipitation, FRET, siRNA knockdown, GLUT4 surface quantification by flow cytometry, glucose uptake assay Journal of cell science Medium 19509061
2011 N-WASP and CK2 (casein kinase 2) form a complex and co-localize at clathrin-coated vesicles. N-WASP binds to and is phosphorylated by CK2, thereby reducing CK2 kinase activity. Conversely, N-WASP-promoted actin polymerization is decreased by CK2 phosphorylation. Both CK2 and N-WASP knockdown inhibit the initial rate of EGFR clathrin-mediated endocytosis (CME). Full rescue requires reconstitution of the N-WASP-CK2 complex; N-WASP controls F-actin presence at clathrin-coated structures. Co-immunoprecipitation, in vitro kinase assay, CK2/N-WASP knockdown, EGFR endocytosis rate measurement, TIRF microscopy of clathrin-coated structures, F-actin quantification Journal of cell science High 21610097
2007 The VCA domain of N-WASP binds the Arp2/3 complex in a 1:1 stoichiometry even with excess VCA. VCA-Arp2/3 binds one actin in a 1:1:1 complex (latrunculin A-sensitive), with binding of the second actin to VCA weakened in the ternary complex. Each of the two WH2 (V) domains independently binds G-actin in 1:2 complexes. V, VC, and VCA enhance barbed end depolymerization but do not nucleate or sever filaments. Protein crystallography (partial VC-actin crystal structure), hydrodynamic methods, spectrofluorimetry, in vitro actin polymerization/depolymerization assays The Journal of biological chemistry High 22847007
2007 Multiple WIP-binding epitopes (three distinct regions in WIP residues 451-485) are required for functional interaction with the N-WASP EVH1 (WH1) domain. A central polyproline motif occupies the canonical EVH1 binding site in a reversed orientation; flanking hydrophobic contacts (WIP residues 454-459 and 475-478) augment binding. Disruption of any of the three WIP epitopes reduces N-WASP binding in cells. NMR structure determination of WIP-EVH1 complex, binding affinity measurements, site-directed mutagenesis, co-immunoprecipitation in cells The Journal of biological chemistry High 17229736
2021 The Chlamydia trachomatis type III secretion effector TmeA directly activates N-WASP to promote Arp2/3-dependent actin polymerization during chlamydial invasion. TmeA and TarP influence separate but synergistic pathways for chlamydial entry. Chlamydial gene deletion (FRAEM), proximity labeling, direct binding assay, actin polymerization assay, infection assays with TmeA deletion mutants mBio Medium 33468693

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1999 The interaction between N-WASP and the Arp2/3 complex links Cdc42-dependent signals to actin assembly. Cell 1107 10219243
1998 Induction of filopodium formation by a WASP-related actin-depolymerizing protein N-WASP. Nature 568 9422512
2005 Molecular mechanisms of invadopodium formation: the role of the N-WASP-Arp2/3 complex pathway and cofilin. The Journal of cell biology 560 15684033
1996 N-WASP, a novel actin-depolymerizing protein, regulates the cortical cytoskeletal rearrangement in a PIP2-dependent manner downstream of tyrosine kinases. The EMBO journal 548 8895577
2000 Mechanism of N-WASP activation by CDC42 and phosphatidylinositol 4, 5-bisphosphate. The Journal of cell biology 512 10995436
2004 Toca-1 mediates Cdc42-dependent actin nucleation by activating the N-WASP-WIP complex. Cell 368 15260990
2000 Actin-dependent propulsion of endosomes and lysosomes by recruitment of N-WASP. The Journal of cell biology 363 10662777
2001 Endocytic protein intersectin-l regulates actin assembly via Cdc42 and N-WASP. Nature cell biology 304 11584276
2000 A complex of N-WASP and WIP integrates signalling cascades that lead to actin polymerization. Nature cell biology 282 10878810
2001 N-WASP deficiency reveals distinct pathways for cell surface projections and microbial actin-based motility. Nature cell biology 267 11584271
2001 WIP regulates N-WASP-mediated actin polymerization and filopodium formation. Nature cell biology 238 11331876
2012 N-WASP-mediated invadopodium formation is involved in intravasation and lung metastasis of mammary tumors. Journal of cell science 217 22389406
2008 N-wasp and the arp2/3 complex are critical regulators of actin in the development of dendritic spines and synapses. The Journal of biological chemistry 184 18430734
2005 Abi1 regulates the activity of N-WASP and WAVE in distinct actin-based processes. Nature cell biology 182 16155590
2002 Syndapins integrate N-WASP in receptor-mediated endocytosis. The EMBO journal 172 12426380
2008 EFC/F-BAR proteins and the N-WASP-WIP complex induce membrane curvature-dependent actin polymerization. The EMBO journal 152 18923421
2003 Focal adhesion kinase regulation of N-WASP subcellular localization and function. The Journal of biological chemistry 151 14676198
2006 Regulation of RNA-polymerase-II-dependent transcription by N-WASP and its nuclear-binding partners. Nature cell biology 138 16767080
2012 N-WASP coordinates the delivery and F-actin-mediated capture of MT1-MMP at invasive pseudopods. The Journal of cell biology 135 23091069
2005 N-WASP deficiency impairs EGF internalization and actin assembly at clathrin-coated pits. Journal of cell science 133 15985465
2004 Imaging sites of N-wasp activity in lamellipodia and invadopodia of carcinoma cells. Current biology : CB 132 15084285
2001 A novel neural Wiskott-Aldrich syndrome protein (N-WASP) binding protein, WISH, induces Arp2/3 complex activation independent of Cdc42. The Journal of cell biology 127 11157975
2006 IQGAP1 stimulates actin assembly through the N-WASP-Arp2/3 pathway. The Journal of biological chemistry 126 17085436
2002 Regulation of protein transport from the Golgi complex to the endoplasmic reticulum by CDC42 and N-WASP. Molecular biology of the cell 126 11907268
2002 Phosphatidylinositol 4,5-biphosphate (PIP2)-induced vesicle movement depends on N-WASP and involves Nck, WIP, and Grb2. The Journal of biological chemistry 123 12147689
2001 A phosphatidylinositol 3-kinase-independent insulin signaling pathway to N-WASP/Arp2/3/F-actin required for GLUT4 glucose transporter recycling. The Journal of biological chemistry 123 11694514
2009 The rate of N-WASP exchange limits the extent of ARP2/3-complex-dependent actin-based motility. Nature 118 19262673
2011 N-WASP regulates the epithelial junctional actin cytoskeleton through a non-canonical post-nucleation pathway. Nature cell biology 115 21785420
2010 Generation of branched actin networks: assembly and regulation of the N-WASP and WAVE molecular machines. BioEssays : news and reviews in molecular, cellular and developmental biology 115 20091750
2004 Cortactin regulates cell migration through activation of N-WASP. Journal of cell science 111 15585574
2007 Differential regulation of WASP and N-WASP by Cdc42, Rac1, Nck, and PI(4,5)P2. Biochemistry 108 17302440
2010 Cdc42-interacting protein 4 promotes breast cancer cell invasion and formation of invadopodia through activation of N-WASp. Cancer research 92 20940394
2009 N-WASP and cortactin are involved in invadopodium-dependent chemotaxis to EGF in breast tumor cells. Cell motility and the cytoskeleton 91 19373774
2001 N-WASP, WAVE and Mena play different roles in the organization of actin cytoskeleton in lamellipodia. Journal of cell science 88 11282031
2005 Differential regulation of cortactin and N-WASP-mediated actin polymerization by missing in metastasis (MIM) protein. Oncogene 87 15688017
2007 Wiskott Aldrich syndrome protein (WASP) and N-WASP are critical for T cell development. Proceedings of the National Academy of Sciences of the United States of America 85 17878299
2012 The actin regulator N-WASp is required for muscle-cell fusion in mice. Proceedings of the National Academy of Sciences of the United States of America 83 22736793
2015 microRNA miR-142-3p Inhibits Breast Cancer Cell Invasiveness by Synchronous Targeting of WASL, Integrin Alpha V, and Additional Cytoskeletal Elements. PloS one 82 26657485
2007 Regulation of N-WASP and the Arp2/3 complex by Abp1 controls neuronal morphology. PloS one 80 17476322
2005 Spatially distinct binding of Cdc42 to PAK1 and N-WASP in breast carcinoma cells. Molecular and cellular biology 76 15713627
2004 Association of Cdc42/N-WASP/Arp2/3 signaling pathway with Golgi membranes. Traffic (Copenhagen, Denmark) 76 15479449
2007 N-WASP regulates extension of filopodia and processes by oligodendrocyte progenitors, oligodendrocytes, and Schwann cells-implications for axon ensheathment at myelination. Glia 70 17405146
1999 Signalling to actin: the Cdc42-N-WASP-Arp2/3 connection. Chemistry & biology 70 10467124
2010 Nebulin and N-WASP cooperate to cause IGF-1-induced sarcomeric actin filament formation. Science (New York, N.Y.) 69 21148390
2004 Regulation of actin cytoskeleton by mDab1 through N-WASP and ubiquitination of mDab1. The Biochemical journal 69 15361067
2008 Cortactin (CTTN), N-WASP (WASL), and clathrin (CLTC) are present at podosome-like tubulobulbar complexes in the rat testis. Biology of reproduction 68 18799755
2009 The Toca-1-N-WASP complex links filopodial formation to endocytosis. The Journal of biological chemistry 67 19213734
2004 N-WASP and WAVE2 acting downstream of phosphatidylinositol 3-kinase are required for myogenic cell migration induced by hepatocyte growth factor. The Journal of biological chemistry 67 15496413
2011 N-WASP is required for membrane wrapping and myelination by Schwann cells. The Journal of cell biology 65 21263026
2013 WIP provides an essential link between Nck and N-WASP during Arp2/3-dependent actin polymerization. Current biology : CB 64 23707428
2012 Loss of Scar/WAVE complex promotes N-WASP- and FAK-dependent invasion. Current biology : CB 62 23273897
2007 Sorting nexin 9 interacts with dynamin 1 and N-WASP and coordinates synaptic vesicle endocytosis. The Journal of biological chemistry 62 17681954
2006 Tuba stimulates intracellular N-WASP-dependent actin assembly. Journal of cell science 62 16757518
2016 Cdc42 deficiency induces podocyte apoptosis by inhibiting the Nwasp/stress fibers/YAP pathway. Cell death & disease 61 26986510
2012 Wiskott-Aldrich syndrome protein (WASP) and N-WASP are critical for peripheral B-cell development and function. Blood 61 22411869
2013 N-wasp is essential for the negative regulation of B cell receptor signaling. PLoS biology 60 24223520
2012 Claudin-5 is involved in breast cancer cell motility through the N-WASP and ROCK signalling pathways. Journal of experimental & clinical cancer research : CR 60 22559840
2010 The expression of CFL1 and N-WASP in esophageal squamous cell carcinoma and its correlation with clinicopathological features. Diseases of the esophagus : official journal of the International Society for Diseases of the Esophagus 59 20095995
2010 Pak1 phosphorylation enhances cortactin-N-WASP interaction in clathrin-caveolin-independent endocytosis. Traffic (Copenhagen, Denmark) 59 20444238
2002 N-WASP activation by a beta1-integrin-dependent mechanism supports PI3K-independent chemotaxis stimulated by urokinase-type plasminogen activator. Journal of cell science 59 11865026
2013 N-wasp is required for stabilization of podocyte foot processes. Journal of the American Society of Nephrology : JASN 56 23471198
2009 Dynamic interaction of amphiphysin with N-WASP regulates actin assembly. The Journal of biological chemistry 56 19759398
2006 N-WASP involvement in dorsal ruffle formation in mouse embryonic fibroblasts. Molecular biology of the cell 56 17182853
2002 WICH, a novel verprolin homology domain-containing protein that functions cooperatively with N-WASP in actin-microspike formation. Biochemical and biophysical research communications 56 11829459
2010 Cdc42 interaction with N-WASP and Toca-1 regulates membrane tubulation, vesicle formation and vesicle motility: implications for endocytosis. PloS one 55 20730103
2009 The F-BAR protein CIP4 promotes GLUT4 endocytosis through bidirectional interactions with N-WASp and Dynamin-2. Journal of cell science 53 19509061
2013 Cdc42 and the Rho GEF intersectin-1 collaborate with Nck to promote N-WASP-dependent actin polymerisation. Journal of cell science 49 24284073
2003 Translocation of N-WASP by nuclear localization and export signals into the nucleus modulates expression of HSP90. The Journal of biological chemistry 49 12871950
2010 Gas7 functions with N-WASP to regulate the neurite outgrowth of hippocampal neurons. The Journal of biological chemistry 48 20150425
2006 N-WASP inhibitor wiskostatin nonselectively perturbs membrane transport by decreasing cellular ATP levels. American journal of physiology. Cell physiology 48 17092993
2002 How signaling proteins integrate multiple inputs: a comparison of N-WASP and Cdk2. Current opinion in cell biology 48 11891112
2007 Integrin-linked kinase regulates N-WASp-mediated actin polymerization and tension development in tracheal smooth muscle. The Journal of biological chemistry 47 17897939
2008 Bacterial actin assembly requires toca-1 to relieve N-wasp autoinhibition. Cell host & microbe 46 18191793
2005 Interaction of HSP90 to N-WASP leads to activation and protection from proteasome-dependent degradation. The EMBO journal 45 15791211
2001 A role for N-WASP in invasin-promoted internalisation. FEBS letters 45 11734206
1997 Identification of N-WASP homologs in human and rat brain. Gene 45 9322739
2019 N-WASP Control of LPAR1 Trafficking Establishes Response to Self-Generated LPA Gradients to Promote Pancreatic Cancer Cell Metastasis. Developmental cell 43 31668663
2014 Cdc42/N-WASP signaling links actin dynamics to pancreatic β cell delamination and differentiation. Development (Cambridge, England) 43 24449844
2012 Neural Wiskott-Aldrich syndrome protein (N-WASP)-mediated p120-catenin interaction with Arp2-Actin complex stabilizes endothelial adherens junctions. The Journal of biological chemistry 43 23212915
2011 An SK3 channel/nWASP/Abi-1 complex is involved in early neurogenesis. PloS one 43 21464958
2009 Nck- and N-WASP-dependent actin-based motility is conserved in divergent vertebrate poxviruses. Cell host & microbe 43 20006842
2008 Repetitive N-WASP-binding elements of the enterohemorrhagic Escherichia coli effector EspF(U) synergistically activate actin assembly. PLoS pathogens 42 18974829
2007 C. elegans Enabled exhibits novel interactions with N-WASP, Abl, and cell-cell junctions. Current biology : CB 42 17935994
2007 N-WASP is a putative tumour suppressor in breast cancer cells, in vitro and in vivo, and is associated with clinical outcome in patients with breast cancer. Clinical & experimental metastasis 42 17985201
2014 Polycystin-1 regulates actin cytoskeleton organization and directional cell migration through a novel PC1-Pacsin 2-N-Wasp complex. Human molecular genetics 41 24385601
2006 Characterization of TccP-mediated N-WASP activation during enterohaemorrhagic Escherichia coli infection. Cellular microbiology 40 16922863
2020 TRPV4 activates the Cdc42/N-wasp pathway to promote glioblastoma invasion by altering cellular protrusions. Scientific reports 39 32843668
2007 Multiple WASP-interacting protein recognition motifs are required for a functional interaction with N-WASP. The Journal of biological chemistry 39 17229736
2012 Interactions of isolated C-terminal fragments of neural Wiskott-Aldrich syndrome protein (N-WASP) with actin and Arp2/3 complex. The Journal of biological chemistry 37 22847007
2007 N-WASP plays a critical role in fibroblast adhesion and spreading. Biochemical and biophysical research communications 37 17963692
2014 N-wasp is required for structural integrity of the blood-testis barrier. PLoS genetics 36 24967734
2009 A reciprocal interdependence between Nck and PI(4,5)P(2) promotes localized N-WASp-mediated actin polymerization in living cells. Molecular cell 34 19917259
2021 Chlamydia trachomatis TmeA Directly Activates N-WASP To Promote Actin Polymerization and Functions Synergistically with TarP during Invasion. mBio 33 33468693
2001 WASP and N-WASP in human platelets differ in sensitivity to protease calpain. Blood 33 11698281
2012 Nck and Cdc42 co-operate to recruit N-WASP to promote FcγR-mediated phagocytosis. Journal of cell science 32 22454526
2018 Rac1/WAVE2 and Cdc42/N-WASP Participation in Actin-Dependent Host Cell Invasion by Extracellular Amastigotes of Trypanosoma cruzi. Frontiers in microbiology 31 29541069
2011 WAVE2, N-WASP, and Mena facilitate cell invasion via phosphatidylinositol 3-kinase-dependent local accumulation of actin filaments. Journal of cellular biochemistry 31 21769917
2006 SPIN90/WISH interacts with PSD-95 and regulates dendritic spinogenesis via an N-WASP-independent mechanism. The EMBO journal 31 16990791
2014 High mobility group Box-1 inhibits cancer cell motility and metastasis by suppressing activation of transcription factor CREB and nWASP expression. Oncotarget 30 25277185
2011 Interplay between N-WASP and CK2 optimizes clathrin-mediated endocytosis of EGFR. Journal of cell science 30 21610097

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