{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,4]],"date-time":"2026-10-04T23:11:39Z","timestamp":1791155499784,"version":"4.1.0"},"reference-count":61,"publisher":"Public Library of Science (PLoS)","issue":"12","license":[{"start":{"date-parts":[[2011,12,14]],"date-time":"2011-12-14T00:00:00Z","timestamp":1323820800000},"content-version":"unspecified","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS ONE"],"DOI":"10.1371\/journal.pone.0028710","type":"journal-article","created":{"date-parts":[[2011,12,14]],"date-time":"2011-12-14T20:19:38Z","timestamp":1323893978000},"page":"e28710","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":48,"title":["Nephrin Regulates Lamellipodia Formation by Assembling a Protein Complex That Includes Ship2, Filamin and Lamellipodin"],"prefix":"10.1371","volume":"6","author":[{"given":"Madhusudan","family":"Venkatareddy","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Leslie","family":"Cook","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kamal","family":"Abuarquob","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rakesh","family":"Verma","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Puneet","family":"Garg","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2011,12,14]]},"reference":[{"key":"ref1","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1038\/74166","article-title":"NPHS2, encoding the glomerular protein podocin, is mutated in autosomal recessive steroid-resistant nephrotic syndrome.","volume":"24","author":"N Boute","year":"2000","journal-title":"Nat Genet"},{"key":"ref2","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1016\/S1097-2765(00)80057-X","article-title":"Positionally cloned gene for a novel glomerular protein\u2013nephrin\u2013is mutated in congenital nephrotic syndrome.","volume":"1","author":"M Kestila","year":"1998","journal-title":"Mol Cell"},{"key":"ref3","doi-asserted-by":"crossref","first-page":"1481","DOI":"10.1046\/j.1523-1755.1999.00719.x","article-title":"Nephrin localizes to the slit pore of the glomerular epithelial cell.","volume":"56","author":"LB Holzman","year":"1999","journal-title":"Kidney Int"},{"key":"ref4","doi-asserted-by":"crossref","first-page":"1586","DOI":"10.1097\/01.ASN.0000016142.29721.22","article-title":"Nephrin TRAP mice lack slit diaphragms and show fibrotic glomeruli and cystic tubular lesions.","volume":"13","author":"M Rantanen","year":"2002","journal-title":"J Am Soc Nephrol"},{"key":"ref5","doi-asserted-by":"crossref","first-page":"31154","DOI":"10.1074\/jbc.M204806200","article-title":"Determinants of vascular permeability in the kidney glomerulus.","volume":"277","author":"Y Hamano","year":"2002","journal-title":"J Biol Chem"},{"key":"ref6","doi-asserted-by":"crossref","first-page":"20716","DOI":"10.1074\/jbc.M301689200","article-title":"Fyn binds to and phosphorylates the kidney slit diaphragm component Nephrin.","volume":"278","author":"R Verma","year":"2003","journal-title":"J Biol Chem"},{"key":"ref7","doi-asserted-by":"crossref","first-page":"1346","DOI":"10.1172\/JCI27414","article-title":"Nephrin ectodomain engagement results in Src kinase activation, nephrin phosphorylation, Nck recruitment, and actin polymerization.","volume":"116","author":"R Verma","year":"2006","journal-title":"J Clin Invest"},{"key":"ref8","doi-asserted-by":"crossref","first-page":"818","DOI":"10.1038\/nature04662","article-title":"Nck adaptor proteins link nephrin to the actin cytoskeleton of kidney podocytes.","volume":"440","author":"N Jones","year":"2006","journal-title":"Nature"},{"key":"ref9","doi-asserted-by":"crossref","first-page":"9177","DOI":"10.1074\/jbc.M707247200","article-title":"Neph1, a component of the kidney slit diaphragm, is tyrosine-phosphorylated by the Src family tyrosine kinase and modulates intracellular signaling by binding to Grb2.","volume":"283","author":"Y Harita","year":"2008","journal-title":"J Biol Chem"},{"key":"ref10","doi-asserted-by":"crossref","first-page":"556","DOI":"10.1038\/sj.ki.5002691","article-title":"Nephrin mediates actin reorganization via phosphoinositide 3-kinase in podocytes.","volume":"73","author":"J Zhu","year":"2008","journal-title":"Kidney Int"},{"key":"ref11","doi-asserted-by":"crossref","first-page":"8951","DOI":"10.1074\/jbc.M806851200","article-title":"Phosphorylation of Nephrin Triggers Ca2+ Signaling by Recruitment and Activation of Phospholipase C-{gamma}1.","volume":"284","author":"Y Harita","year":"2009","journal-title":"J Biol Chem"},{"key":"ref12","doi-asserted-by":"crossref","first-page":"1188","DOI":"10.1172\/JCI200523371","article-title":"Synaptopodin regulates the actin-bundling activity of alpha-actinin in an isoform-specific manner.","volume":"115","author":"K Asanuma","year":"2005","journal-title":"J Clin Invest"},{"key":"ref13","doi-asserted-by":"crossref","first-page":"2035","DOI":"10.1128\/MCB.01770-07","article-title":"Phosphorylated YDXV motifs and Nck SH2\/SH3 adaptors act cooperatively to induce actin reorganization.","volume":"28","author":"IM Blasutig","year":"2008","journal-title":"Mol Cell Biol"},{"key":"ref14","doi-asserted-by":"crossref","first-page":"9814","DOI":"10.1073\/pnas.0504166102","article-title":"Cell junction-associated proteins IQGAP1, MAGI-2, CASK, spectrins, and alpha-actinin are components of the nephrin multiprotein complex.","volume":"102","author":"S Lehtonen","year":"2005","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref15","doi-asserted-by":"crossref","first-page":"22676","DOI":"10.1074\/jbc.M110.122929","article-title":"Actin-depolymerizing factor cofilin-1 is necessary in maintaining mature podocyte architecture.","volume":"285","author":"P Garg","journal-title":"J Biol Chem"},{"key":"ref16","doi-asserted-by":"crossref","first-page":"8698","DOI":"10.1128\/MCB.00948-07","article-title":"Neph1 cooperates with nephrin to transduce a signal that induces actin polymerization.","volume":"27","author":"P Garg","year":"2007","journal-title":"Mol Cell Biol"},{"key":"ref17","doi-asserted-by":"crossref","first-page":"453","DOI":"10.1016\/S0092-8674(03)00120-X","article-title":"Cellular motility driven by assembly and disassembly of actin filaments.","volume":"112","author":"TD Pollard","year":"2003","journal-title":"Cell"},{"key":"ref18","doi-asserted-by":"crossref","first-page":"62","DOI":"10.3410\/B2-62","article-title":"Roles of ADF\/cofilin in actin polymerization and beyond.","volume":"2","author":"JR Bamburg","year":"2010","journal-title":"F1000 Biol Rep"},{"key":"ref19","doi-asserted-by":"crossref","first-page":"160","DOI":"10.4161\/cam.5.2.14401","article-title":"The filamins: organizers of cell structure and function.","volume":"5","author":"F Nakamura","year":"2011","journal-title":"Cell Adh Migr"},{"key":"ref20","doi-asserted-by":"crossref","first-page":"1011","DOI":"10.1083\/jcb.200707073","article-title":"Structural basis of filamin A functions.","volume":"179","author":"F Nakamura","year":"2007","journal-title":"J Cell Biol"},{"key":"ref21","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1016\/S0955-0674(99)00067-8","article-title":"Focal adhesions - the cytoskeletal connection.","volume":"12","author":"DR Critchley","year":"2000","journal-title":"Curr Opin Cell Biol"},{"key":"ref22","doi-asserted-by":"crossref","first-page":"2260","DOI":"10.1016\/j.biocel.2005.05.003","article-title":"The SH2 domain containing inositol polyphosphate 5-phosphatase-2: SHIP2.","volume":"37","author":"JM Dyson","year":"2005","journal-title":"Int J Biochem Cell Biol"},{"key":"ref23","doi-asserted-by":"crossref","first-page":"1876","DOI":"10.1182\/blood.V89.6.1876","article-title":"The human SHIP gene is differentially expressed in cell lineages of the bone marrow and blood.","volume":"89","author":"SJ Geier","year":"1997","journal-title":"Blood"},{"key":"ref24","doi-asserted-by":"crossref","first-page":"18605","DOI":"10.1074\/jbc.273.29.18605","article-title":"Growth factors and insulin stimulate tyrosine phosphorylation of the 51C\/SHIP2 protein.","volume":"273","author":"T Habib","year":"1998","journal-title":"J Biol Chem"},{"key":"ref25","doi-asserted-by":"crossref","first-page":"1170","DOI":"10.1016\/j.exphem.2003.09.011","article-title":"SHIP, SHIP2, and PTEN activities are regulated in vivo by modulation of their protein levels: SHIP is up-regulated in macrophages and mast cells by lipopolysaccharide.","volume":"31","author":"LM Sly","year":"2003","journal-title":"Exp Hematol"},{"key":"ref26","doi-asserted-by":"crossref","first-page":"1051","DOI":"10.1101\/gad.10.9.1051","article-title":"Inositols do it all.","volume":"10","author":"PW Majerus","year":"1996","journal-title":"Genes Dev"},{"key":"ref27","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1016\/S0959-4388(97)80060-8","article-title":"Phosphoinositides as spatial regulators of membrane traffic.","volume":"7","author":"TF Martin","year":"1997","journal-title":"Curr Opin Neurobiol"},{"key":"ref28","doi-asserted-by":"crossref","first-page":"1065","DOI":"10.1083\/jcb.200104005","article-title":"The SH2-containing inositol polyphosphate 5-phosphatase, SHIP-2, binds filamin and regulates submembraneous actin.","volume":"155","author":"JM Dyson","year":"2001","journal-title":"J Cell Biol"},{"key":"ref29","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/j.chom.2009.12.004","article-title":"Enteropathogenic Escherichia coli recruits the cellular inositol phosphatase SHIP2 to regulate actin-pedestal formation.","volume":"7","author":"K Smith","year":"2010","journal-title":"Cell Host Microbe"},{"key":"ref30","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.chom.2009.12.006","article-title":"Phosphoinositides influence pathogen surfing: EPEC rights the SHIP.","volume":"7","author":"KG Campellone","year":"2010","journal-title":"Cell Host Microbe"},{"key":"ref31","doi-asserted-by":"crossref","first-page":"404","DOI":"10.1046\/j.1523-1755.2003.00097.x","article-title":"Clustering-induced tyrosine phosphorylation of nephrin by Src family kinases.","volume":"64","author":"J Lahdenpera","year":"2003","journal-title":"Kidney Int"},{"key":"ref32","doi-asserted-by":"crossref","first-page":"625","DOI":"10.1016\/S0092-8674(00)81604-9","article-title":"A requirement for caveolin-1 and associated kinase Fyn in integrin signaling and anchorage-dependent cell growth.","volume":"94","author":"KK Wary","year":"1998","journal-title":"Cell"},{"key":"ref33","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.cub.2003.12.033","article-title":"Inducible clustering of membrane-targeted SH3 domains of the adaptor protein Nck triggers localized actin polymerization.","volume":"14","author":"GM Rivera","year":"2004","journal-title":"Curr Biol"},{"key":"ref34","doi-asserted-by":"crossref","first-page":"5095","DOI":"10.1016\/j.bpj.2009.03.046","article-title":"Absence of filamin A prevents cells from responding to stiffness gradients on gels coated with collagen but not fibronectin.","volume":"96","author":"FJ Byfield","year":"2009","journal-title":"Biophys J"},{"key":"ref35","doi-asserted-by":"crossref","first-page":"22676","DOI":"10.1074\/jbc.M110.122929","article-title":"Actin-depolymerizing factor cofilin-1 is necessary in maintaining mature podocyte architecture.","volume":"285","author":"P Garg","year":"2010","journal-title":"J Biol Chem"},{"key":"ref36","doi-asserted-by":"crossref","first-page":"940","DOI":"10.1182\/blood-2002-09-2897","article-title":"SHIP-2 forms a tetrameric complex with filamin, actin, and GPIb-IX-V: localization of SHIP-2 to the activated platelet actin cytoskeleton.","volume":"102","author":"JM Dyson","year":"2003","journal-title":"Blood"},{"key":"ref37","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1038\/ncb838","article-title":"Filamin is essential in actin cytoskeletal assembly mediated by p21-activated kinase 1.","volume":"4","author":"RK Vadlamudi","year":"2002","journal-title":"Nat Cell Biol"},{"key":"ref38","doi-asserted-by":"crossref","first-page":"2762","DOI":"10.1016\/j.yexcr.2008.06.004","article-title":"Filamin A is a novel caveolin-1-dependent target in IGF-I-stimulated cancer cell migration.","volume":"314","author":"D Ravid","year":"2008","journal-title":"Exp Cell Res"},{"key":"ref39","doi-asserted-by":"crossref","first-page":"511","DOI":"10.1083\/jcb.200105148","article-title":"Filamin A, the Arp2\/3 complex, and the morphology and function of cortical actin filaments in human melanoma cells.","volume":"155","author":"LA Flanagan","year":"2001","journal-title":"J Cell Biol"},{"key":"ref40","doi-asserted-by":"crossref","first-page":"3807","DOI":"10.1242\/jcs.00070","article-title":"Src family tyrosine kinases regulate adhesion-dependent tyrosine phosphorylation of 5\u2032-inositol phosphatase SHIP2 during cell attachment and spreading on collagen I.","volume":"115","author":"N Prasad","year":"2002","journal-title":"J Cell Sci"},{"key":"ref41","doi-asserted-by":"crossref","first-page":"8347","DOI":"10.1074\/jbc.274.13.8347","article-title":"The role of phosphoinositide 3-kinase lipid products in cell function.","volume":"274","author":"LE Rameh","year":"1999","journal-title":"J Biol Chem"},{"key":"ref42","doi-asserted-by":"crossref","first-page":"53717","DOI":"10.1074\/jbc.M410654200","article-title":"The phosphoinositol 3,4-bisphosphate-binding protein TAPP1 interacts with syntrophins and regulates actin cytoskeletal organization.","volume":"279","author":"A Hogan","year":"2004","journal-title":"J Biol Chem"},{"key":"ref43","doi-asserted-by":"crossref","first-page":"571","DOI":"10.1016\/j.devcel.2004.07.024","article-title":"Lamellipodin, an Ena\/VASP ligand, is implicated in the regulation of lamellipodial dynamics.","volume":"7","author":"M Krause","year":"2004","journal-title":"Dev Cell"},{"key":"ref44","doi-asserted-by":"crossref","first-page":"854","DOI":"10.1016\/j.cub.2006.03.083","article-title":"MIG-10\/lamellipodin and AGE-1\/PI3K promote axon guidance and outgrowth in response to slit and netrin.","volume":"16","author":"C Chang","year":"2006","journal-title":"Curr Biol"},{"key":"ref45","doi-asserted-by":"crossref","first-page":"1390","DOI":"10.1172\/JCI20402","article-title":"Induction of B7-1 in podocytes is associated with nephrotic syndrome.","volume":"113","author":"J Reiser","year":"2004","journal-title":"J Clin Invest"},{"key":"ref46","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1152\/physrev.00036.2009","article-title":"Regulation of the actin cytoskeleton-plasma membrane interplay by phosphoinositides.","volume":"90","author":"J Saarikangas","year":"2010","journal-title":"Physiol Rev"},{"key":"ref47","doi-asserted-by":"crossref","first-page":"926","DOI":"10.1038\/44860","article-title":"Actin-based motility of vaccinia virus mimics receptor tyrosine kinase signalling.","volume":"401","author":"F Frischknecht","year":"1999","journal-title":"Nature"},{"key":"ref48","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1083\/jcb.130.2.331","article-title":"Actin-based movement of Listeria monocytogenes: actin assembly results from the local maintenance of uncapped filament barbed ends at the bacterium surface.","volume":"130","author":"JB Marchand","year":"1995","journal-title":"J Cell Biol"},{"key":"ref49","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/S0962-8924(98)01418-4","article-title":"Putting E. coli on a pedestal: a unique system to study signal transduction and the actin cytoskeleton.","volume":"9","author":"DL Goosney","year":"1999","journal-title":"Trends Cell Biol"},{"key":"ref50","doi-asserted-by":"crossref","first-page":"856","DOI":"10.1038\/ncb0901-856","article-title":"Enteropathogenic E. coli Tir binds Nck to initiate actin pedestal formation in host cells.","volume":"3","author":"S Gruenheid","year":"2001","journal-title":"Nat Cell Biol"},{"key":"ref51","doi-asserted-by":"crossref","first-page":"1025","DOI":"10.1007\/s00018-009-0213-1","article-title":"CAS proteins in normal and pathological cell growth control.","volume":"67","author":"N Tikhmyanova","year":"2010","journal-title":"Cell Mol Life Sci"},{"key":"ref52","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1016\/j.tcb.2006.03.003","article-title":"p130Cas: a versatile scaffold in signaling networks.","volume":"16","author":"P Defilippi","year":"2006","journal-title":"Trends Cell Biol"},{"key":"ref53","doi-asserted-by":"crossref","first-page":"1416","DOI":"10.1128\/MCB.21.4.1416-1428.2001","article-title":"SH2-containing inositol 5\u2032-phosphatase SHIP2 associates with the p130(Cas) adapter protein and regulates cellular adhesion and spreading.","volume":"21","author":"N Prasad","year":"2001","journal-title":"Mol Cell Biol"},{"key":"ref54","doi-asserted-by":"crossref","first-page":"1853","DOI":"10.1182\/blood-2007-12-127795","article-title":"Beta2 integrin phosphorylation on Thr758 acts as a molecular switch to regulate 14-3-3 and filamin binding.","volume":"112","author":"H Takala","year":"2008","journal-title":"Blood"},{"key":"ref55","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1016\/j.molcel.2006.01.011","article-title":"The molecular basis of filamin binding to integrins and competition with talin.","volume":"21","author":"T Kiema","year":"2006","journal-title":"Mol Cell"},{"key":"ref56","doi-asserted-by":"crossref","first-page":"2421","DOI":"10.1084\/jem.20100433","article-title":"Filamin A regulates focal adhesion disassembly and suppresses breast cancer cell migration and invasion.","volume":"207","author":"Y Xu","year":"2010","journal-title":"J Exp Med"},{"key":"ref57","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1016\/0167-4838(87)90099-9","article-title":"Calpain abolishes the effect of filamin on the actomyosin system in platelets.","volume":"912","author":"T Onji","year":"1987","journal-title":"Biochim Biophys Acta"},{"key":"ref58","first-page":"97","article-title":"SHIP2 phosphoinositol phosphatase positively regulates EGFR-Akt pathway, CXCR4 expression, and cell migration in MDA-MB-231 breast cancer cells.","volume":"34","author":"NK Prasad","year":"2009","journal-title":"Int J Oncol"},{"key":"ref59","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1083\/jcb.151.2.249","article-title":"Role of the PI3K regulatory subunit in the control of actin organization and cell migration.","volume":"151","author":"C Jimenez","year":"2000","journal-title":"J Cell Biol"},{"key":"ref60","doi-asserted-by":"crossref","first-page":"13129","DOI":"10.1074\/jbc.M410289200","article-title":"SH2-containing 5\u2032-inositol phosphatase, SHIP2, regulates cytoskeleton organization and ligand-dependent down-regulation of the epidermal growth factor receptor.","volume":"280","author":"NK Prasad","year":"2005","journal-title":"J Biol Chem"},{"key":"ref61","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1242\/jcs.037564","article-title":"Spontaneous phosphoinositide 3-kinase signaling dynamics drive spreading and random migration of fibroblasts.","volume":"122","author":"MC Weiger","year":"2009","journal-title":"J Cell Sci"}],"container-title":["PLoS ONE"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/dx.plos.org\/10.1371\/journal.pone.0028710","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,12,20]],"date-time":"2021-12-20T10:00:42Z","timestamp":1639994442000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0028710"}},"subtitle":[],"editor":[{"given":"James","family":"Keen","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2011,12,14]]},"references-count":61,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2011,12,14]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0028710","relation":{},"ISSN":["1932-6203"],"issn-type":[{"value":"1932-6203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2011,12,14]]}}}