{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,5]],"date-time":"2026-10-05T13:36:18Z","timestamp":1791207378428,"version":"4.1.0"},"reference-count":63,"publisher":"eLife Sciences Publications, Ltd","license":[{"start":{"date-parts":[[2013,9,3]],"date-time":"2013-09-03T00:00:00Z","timestamp":1378166400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/3.0\/"},{"start":{"date-parts":[[2013,9,3]],"date-time":"2013-09-03T00:00:00Z","timestamp":1378166400000},"content-version":"am","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/3.0\/"},{"start":{"date-parts":[[2013,9,3]],"date-time":"2013-09-03T00:00:00Z","timestamp":1378166400000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/3.0\/"},{"start":{"date-parts":[[2013,9,3]],"date-time":"2013-09-03T00:00:00Z","timestamp":1378166400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/3.0\/"},{"start":{"date-parts":[[2013,9,3]],"date-time":"2013-09-03T00:00:00Z","timestamp":1378166400000},"content-version":"am","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/3.0\/"},{"start":{"date-parts":[[2013,9,3]],"date-time":"2013-09-03T00:00:00Z","timestamp":1378166400000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/3.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["R01-GM092917"],"award-info":[{"award-number":["R01-GM092917"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"name":"Pew Biomedical Scholar Program"},{"name":"National Institutes of Health Predoctoral Training Grant","award":["GM007759"],"award-info":[{"award-number":["GM007759"]}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01-GM092917"],"award-info":[{"award-number":["R01-GM092917"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000875","name":"Pew Charitable Trusts","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000875","id-type":"DOI","asserted-by":"publisher"}]},{"name":"National Institutes of Health","award":["GM007759"],"award-info":[{"award-number":["GM007759"]}]}],"content-domain":{"domain":["www.elifesciences.org"],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:p>Nucleation promoting factors (NPFs) initiate branched actin network assembly by activating Arp2\/3 complex, a branched actin filament nucleator. Cellular actin networks contain multiple NPFs, but how they coordinately regulate Arp2\/3 complex is unclear. Cortactin is an NPF that activates Arp2\/3 complex weakly on its own, but with WASP\/N-WASP, another class of NPFs, potently activates. We dissect the mechanism of synergy and propose a model in which cortactin displaces N-WASP from nascent branches as a prerequisite for nucleation. Single-molecule imaging revealed that unlike WASP\/N-WASP, cortactin remains bound to junctions during nucleation, and specifically targets junctions with a \u223c160-fold increased on rate over filament sides. N-WASP must be dimerized for potent synergy, and targeted mutations indicate release of dimeric N-WASP from nascent branches limits nucleation. Mathematical modeling shows cortactin-mediated displacement but not N-WASP recycling or filament recruitment models can explain synergy. Our results provide a molecular basis for coordinate Arp2\/3 complex regulation.<\/jats:p>","DOI":"10.7554\/elife.00884","type":"journal-article","created":{"date-parts":[[2013,9,4]],"date-time":"2013-09-04T11:38:51Z","timestamp":1378294731000},"source":"Crossref","is-referenced-by-count":133,"title":["Mechanism of synergistic activation of Arp2\/3 complex by cortactin and N-WASP"],"prefix":"10.7554","volume":"2","author":[{"given":"Luke A","family":"Helgeson","sequence":"first","affiliation":[{"name":"Institute of Molecular Biology, University of Oregon, Eugene, United States"},{"name":"Department of Chemistry and Biochemistry, University of Oregon, Eugene, United States"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Brad J","family":"Nolen","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biology, University of Oregon, Eugene, United States"},{"name":"Department of Chemistry and Biochemistry, University of Oregon, Eugene, United States"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"4374","published-online":{"date-parts":[[2013,9,3]]},"reference":[{"key":"bib1","doi-asserted-by":"publisher","first-page":"423","DOI":"10.1016\/j.cub.2009.12.056","article-title":"A \u2018Primer\u2019-based mechanism underlies branched actin filament network formation and motility","volume":"20","author":"Achard","year":"2010","journal-title":"Curr Biol"},{"key":"bib2","doi-asserted-by":"publisher","first-page":"841","DOI":"10.1016\/j.cell.2008.04.011","article-title":"Capping protein increases the rate of actin-based motility by promoting filament nucleation by the Arp2\/3 complex","volume":"133","author":"Akin","year":"2008","journal-title":"Cell"},{"key":"bib3","doi-asserted-by":"publisher","first-page":"369","DOI":"10.1242\/jcs.008037","article-title":"Multiple regulatory inputs converge on cortactin to control invadopodia biogenesis and extracellular matrix degradation","volume":"121","author":"Ayala","year":"2008","journal-title":"J Cell Sci"},{"key":"bib4","doi-asserted-by":"publisher","first-page":"7135","DOI":"10.1074\/jbc.M705894200","article-title":"Pathway of actin filament branch formation by Arp2\/3 complex","volume":"283","author":"Beltzner","year":"2008","journal-title":"J Biol Chem"},{"key":"bib6","doi-asserted-by":"publisher","first-page":"1273","DOI":"10.1016\/S0960-9822(00)00749-1","article-title":"Interactions of ADF\/cofilin, Arp2\/3 complex, capping protein and profilin in remodeling of branched actin filament networks","volume":"10","author":"Blanchoin","year":"2000","journal-title":"Curr Biol"},{"key":"bib7","doi-asserted-by":"publisher","first-page":"1276","DOI":"10.1016\/j.cub.2005.06.043","article-title":"Cortactin promotes cell motility by enhancing lamellipodial persistence","volume":"15","author":"Bryce","year":"2005","journal-title":"Curr Biol"},{"key":"bib8","doi-asserted-by":"publisher","first-page":"828","DOI":"10.1016\/j.cell.2008.06.054","article-title":"Coronin 1B antagonizes cortactin and remodels Arp2\/3-containing actin branches in lamellipodia","volume":"134","author":"Cai","year":"2008","journal-title":"Cell"},{"key":"bib10","doi-asserted-by":"publisher","first-page":"16644","DOI":"10.1073\/pnas.0507021102","article-title":"Actin-bound structures of Wiskott\u2013Aldrich syndrome protein (WASP)-homology domain 2 and the implications for filament assembly","volume":"102","author":"Chereau","year":"2005","journal-title":"Proc Natl Acad Sci USA"},{"key":"bib11","doi-asserted-by":"publisher","first-page":"1110","DOI":"10.1038\/ncb1007-1110","article-title":"The many faces of actin: matching assembly factors with cellular structures","volume":"9","author":"Chhabra","year":"2007","journal-title":"Nat Cell Biol"},{"key":"bib12","doi-asserted-by":"publisher","first-page":"901","DOI":"10.1016\/j.cell.2006.12.049","article-title":"Mechanism of actin network attachment to moving membranes: barbed end capture by N-WASP WH2 domains","volume":"128","author":"Co","year":"2007","journal-title":"Cell"},{"key":"bib13","doi-asserted-by":"publisher","first-page":"5018","DOI":"10.1021\/ja809117z","article-title":"On the mechanism of Trolox as antiblinking and antibleaching reagent","volume":"131","author":"Cordes","year":"2009","journal-title":"J Am Chem Soc"},{"key":"bib14","doi-asserted-by":"publisher","first-page":"13726","DOI":"10.1073\/pnas.96.24.13726","article-title":"The kinetic mechanism of myosin V","volume":"96","author":"De La Cruz","year":"1999","journal-title":"Proc Natl Acad Sci USA"},{"key":"bib15","doi-asserted-by":"publisher","first-page":"303","DOI":"10.1002\/cm.20361","article-title":"N-WASP and cortactin are involved in invadopodium-dependent chemotaxis to EGF in breast tumor cells","volume":"66","author":"DesMarais","year":"2009","journal-title":"Cell Motil Cytoskel"},{"key":"bib16","doi-asserted-by":"publisher","first-page":"1000","DOI":"10.1038\/emboj.2011.461","article-title":"How a single residue in individual \u03b2-thymosin\/WH2 domains controls their functions in actin assembly","volume":"31","author":"Didry","year":"2011","journal-title":"EMBO J"},{"key":"bib17","doi-asserted-by":"publisher","DOI":"10.1002\/0471142727.mb1420s92","article-title":"Computer control of microscopes using \u00b5Manager","author":"Edelstein","year":"2010","journal-title":"Curr Protoc Mol Biol"},{"key":"bib18","doi-asserted-by":"publisher","first-page":"e383","DOI":"10.1371\/journal.pbio.0030383","article-title":"Mechanism of filament nucleation and branch stability revealed by the structure of the Arp2\/3 complex at actin branch junctions","volume":"3","author":"Egile","year":"2005","journal-title":"PLOS Biol"},{"key":"bib19","doi-asserted-by":"publisher","first-page":"23852","DOI":"10.1074\/jbc.M803448200","article-title":"Close packing of Listeria monocytogenes ActA, a natively unfolded protein, enhances F-actin assembly without dimerization","volume":"283","author":"Footer","year":"2008","journal-title":"J Biol Chem"},{"key":"bib20","doi-asserted-by":"publisher","first-page":"e1","DOI":"10.1371\/journal.pbio.0060001","article-title":"Distinct roles for Arp2\/3 regulators in actin assembly and endocytosis","volume":"6","author":"Galletta","year":"2008","journal-title":"PLOS Biol"},{"key":"bib21","doi-asserted-by":"publisher","first-page":"861","DOI":"10.1016\/j.cub.2010.03.026","article-title":"GMF is a cofilin homolog that binds Arp2\/3 complex to stimulate filament debranching and inhibit actin nucleation","volume":"20","author":"Gandhi","year":"2010","journal-title":"Curr Biol"},{"key":"bib22","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M112.394361","article-title":"Interactions of isolated C-terminal fragments of neural Wiskott-Aldrich syndrome protein (N-WASP) with actin and Arp2\/3 complex","author":"Gaucher","year":"2012","journal-title":"J Biol Chem"},{"key":"bib23","doi-asserted-by":"publisher","first-page":"40171","DOI":"10.1074\/jbc.M110.139337","article-title":"WAVE forms hetero- and homo-oligomeric complexes at integrin junctions in Drosophila visualized by bimolecular fluorescence complementation","volume":"285","author":"Gohl","year":"2010","journal-title":"J Biol Chem"},{"key":"bib24","doi-asserted-by":"publisher","first-page":"713","DOI":"10.1038\/nrm2026","article-title":"The ARP2\/3 complex: an actin nucleator comes of age","volume":"7","author":"Goley","year":"2006","journal-title":"Nat Rev Mol Cell Biol"},{"key":"bib25","doi-asserted-by":"publisher","first-page":"1079","DOI":"10.1111\/j.1600-0854.2010.01075.x","article-title":"Pak1 phosphorylation enhances cortactin-N-WASP interaction in clathrin-caveolin-independent endocytosis","volume":"11","author":"Grassart","year":"2010","journal-title":"Traffic"},{"key":"bib26","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1083\/jcb.201003014","article-title":"VASP is a processive actin polymerase that requires monomeric actin for barbed end association","volume":"191","author":"Hansen","year":"2010","journal-title":"J Cell Biol"},{"key":"bib27","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.chembiol.2013.03.019","article-title":"Small molecules CK-666 and CK-869 inhibit actin-related protein 2\/3 complex by blocking an activating conformational change","volume":"20","author":"Hetrick","year":"2013","journal-title":"Chem Biol"},{"key":"bib28","doi-asserted-by":"publisher","first-page":"1311","DOI":"10.1083\/jcb.150.6.1311","article-title":"Activation by Cdc42 and PIP2 of Wiskott-Aldrich syndrome protein (WASp) stimulates actin nucleation by Arp2\/3 complex","volume":"150","author":"Higgs","year":"2000","journal-title":"J Cell Biol"},{"key":"bib29","doi-asserted-by":"publisher","first-page":"3067","DOI":"10.1093\/bioinformatics\/btl485","article-title":"COPASI\u2013a COmplex PAthway SImulator","volume":"22","author":"Hoops","year":"2006","journal-title":"Bioinformatics"},{"key":"bib30","doi-asserted-by":"publisher","first-page":"10589","DOI":"10.1074\/jbc.M507470200","article-title":"Actin binding to the central domain of WASP\/Scar proteins plays a critical role in the activation of the Arp2\/3 complex","volume":"281","author":"Kelly","year":"2006","journal-title":"J Biol Chem"},{"key":"bib31","doi-asserted-by":"publisher","first-page":"384","DOI":"10.1016\/S0960-9822(03)00107-6","article-title":"Cortactin interacts with WIP in regulating Arp2\/3 activation and membrane protrusion","volume":"13","author":"Kinley","year":"2003","journal-title":"Curr Biol"},{"key":"bib32","doi-asserted-by":"publisher","first-page":"187","DOI":"10.4161\/cam.5.2.14773","article-title":"Cortactin: a multifunctional regulator of cellular invasiveness","volume":"5","author":"Kirkbride","year":"2011","journal-title":"Cell Adh Migr"},{"key":"bib33","doi-asserted-by":"publisher","first-page":"35557","DOI":"10.1074\/jbc.M405319200","article-title":"Mechanism of Blebbistatin inhibition of myosin ii","volume":"279","author":"Kov\u00e1cs","year":"2004","journal-title":"J Biol Chem"},{"key":"bib34","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1242\/jcs.01586","article-title":"Cortactin regulates cell migration through activation of n-WASP","volume":"118","author":"Kowalski","year":"2005","journal-title":"J Cell Sci"},{"key":"bib35","doi-asserted-by":"publisher","first-page":"1387","DOI":"10.1529\/biophysj.104.047399","article-title":"Real-time measurements of actin filament polymerization by total internal reflection fluorescence microscopy","volume":"88","author":"Kuhn","year":"2005","journal-title":"Biophys J"},{"key":"bib36","doi-asserted-by":"publisher","first-page":"982","DOI":"10.1038\/emboj.2008.34","article-title":"Arp2\/3 complex interactions and actin network turnover in lamellipodia","volume":"27","author":"Lai","year":"2008","journal-title":"EMBO J"},{"key":"bib37","doi-asserted-by":"publisher","first-page":"336","DOI":"10.1038\/nature10879","article-title":"Phase transitions in the assembly of multivalent signalling proteins","volume":"483","author":"Li","year":"2012","journal-title":"Nature"},{"key":"bib38","doi-asserted-by":"publisher","first-page":"17039","DOI":"10.1074\/jbc.M111.219964","article-title":"Mechanism of a concentration-dependent Switch between activation and Inhibition of Arp2\/3 complex by coronin","volume":"286","author":"Liu","year":"2011","journal-title":"J Biol Chem"},{"key":"bib39","doi-asserted-by":"publisher","first-page":"3519","DOI":"10.1529\/biophysj.106.080937","article-title":"Kinetics of the formation and dissociation of actin filament branches mediated by Arp2\/3 complex","volume":"91","author":"Mahaffy","year":"2006","journal-title":"Biophys J"},{"key":"bib40","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1038\/35050590","article-title":"Interaction of WASP\/Scar proteins with actin and vertebrate Arp2\/3 complex","volume":"3","author":"Marchand","year":"2001","journal-title":"Nat Cell Biol"},{"key":"bib41","doi-asserted-by":"publisher","first-page":"826","DOI":"10.1038\/ncb1443","article-title":"Arp2\/3 ATP hydrolysis-catalysed branch dissociation is critical for endocytic force generation","volume":"8","author":"Martin","year":"2006","journal-title":"Nat Cell Biol"},{"key":"bib42","doi-asserted-by":"publisher","first-page":"5269","DOI":"10.1128\/MCB.24.12.5269-5280.2004","article-title":"Erk\/Src phosphorylation of cortactin acts as a switch on-switch off mechanism that controls its ability to activate N-WASP","volume":"24","author":"Martinez-Quiles","year":"2004","journal-title":"Mol Cell Biol"},{"key":"bib43","doi-asserted-by":"publisher","first-page":"426","DOI":"10.1016\/j.molcel.2008.10.012","article-title":"Hierarchical regulation of WASP\/WAVE proteins","volume":"32","author":"Padrick","year":"2008","journal-title":"Mol Cell"},{"key":"bib44","doi-asserted-by":"publisher","first-page":"E472","DOI":"10.1073\/pnas.1100236108","article-title":"Arp2\/3 complex is bound and activated by two WASP proteins","volume":"108","author":"Padrick","year":"2011","journal-title":"Proc Natl Acad Sci USA"},{"key":"bib45","doi-asserted-by":"publisher","first-page":"840","DOI":"10.1016\/j.jmb.2006.03.065","article-title":"Cortactin binding to F-actin revealed by electron microscopy and 3D reconstruction","volume":"359","author":"Pant","year":"2006","journal-title":"J Mol Biol"},{"key":"bib46","doi-asserted-by":"publisher","first-page":"2747","DOI":"10.1083\/jcb.103.6.2747","article-title":"Rate constants for the reactions of ATP- and ADP-actin with the ends of actin filaments","volume":"103","author":"Pollard","year":"1986","journal-title":"J Cell Biol"},{"key":"bib47","doi-asserted-by":"publisher","first-page":"451","DOI":"10.1146\/annurev.biophys.35.040405.101936","article-title":"Regulation of actin filament assembly by Arp2\/3 complex and formins","volume":"36","author":"Pollard","year":"2007","journal-title":"Annu Rev Biophys Biomol Struct"},{"key":"bib48","doi-asserted-by":"publisher","first-page":"1208","DOI":"10.1126\/science.1175862","article-title":"Actin, a central player in cell shape and movement","volume":"326","author":"Pollard","year":"2009","journal-title":"Science"},{"key":"bib49","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1016\/j.jmb.2010.08.040","article-title":"Structure of a longitudinal actin dimer assembled by tandem W domains - implications for actin filament nucleation","volume":"403","author":"Rebowski","year":"2010","journal-title":"J Mol Biol"},{"key":"bib50","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1038\/nrm3492","article-title":"New insights into the regulation and cellular functions of the ARP2\/3 complex","volume":"14","author":"Rotty","year":"2013","journal-title":"Nat Rev Mol Cell Biol"},{"key":"bib51","doi-asserted-by":"publisher","first-page":"90","DOI":"10.1002\/cm.20328","article-title":"The actin-binding domain of cortactin is dynamic and unstructured and affects lateral and longitudinal contacts in F-actin","volume":"66","author":"Shvetsov","year":"2009","journal-title":"Cell Motil Cytoskeleton"},{"key":"bib52","doi-asserted-by":"publisher","first-page":"2092","DOI":"10.1016\/j.cub.2011.11.010","article-title":"Cortactin releases the brakes in actin- based motility by enhancing WASP-VCA detachment from Arp2\/3 branches","volume":"21","author":"Siton","year":"2011","journal-title":"Curr Biol"},{"key":"bib53","doi-asserted-by":"publisher","first-page":"1285","DOI":"10.1073\/pnas.1211164110","article-title":"Pathway of actin filament branch formation by Arp2\/3 complex revealed by single-molecule imaging","volume":"110","author":"Smith","year":"2013","journal-title":"Proc Natl Acad Sci USA"},{"key":"bib54","doi-asserted-by":"publisher","first-page":"267","DOI":"10.1016\/j.semcdb.2013.01.006","article-title":"Activation of nucleation promoting factors for directional actin filament elongation: allosteric regulation and multimerization on the membrane","volume":"24","author":"Suetsugu","year":"2013","journal-title":"Semin Cell Dev Biol"},{"key":"bib55","doi-asserted-by":"publisher","first-page":"E472","DOI":"10.1073\/pnas.1100125108","article-title":"Structural and biochemical characterization of two binding sites for nucleation-promoting factor WASp-VCA on Arp2\/3 complex","volume":"108","author":"Ti","year":"2011","journal-title":"Proc Natl Acad Sci USA"},{"key":"bib56","doi-asserted-by":"publisher","first-page":"26086","DOI":"10.1074\/jbc.M301997200","article-title":"Sequential interaction of actin-related proteins 2 and 3 (Arp2\/3) complex with neural Wiscott-Aldrich syndrome protein (N-WASP) and cortactin during branched actin filament network formation","volume":"278","author":"Uruno","year":"2003","journal-title":"J Biol Chem"},{"key":"bib57","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1038\/35060051","article-title":"Activation of Arp2\/3 complex-mediated actin polymerization by cortactin","volume":"3","author":"Uruno","year":"2001","journal-title":"Nat Cell Biol"},{"key":"bib58","doi-asserted-by":"publisher","first-page":"75","DOI":"10.1016\/j.copbio.2010.10.002","article-title":"Single-molecule approaches to characterizing kinetics of biomolecular interactions","volume":"22","author":"Van Oijen","year":"2011","journal-title":"Curr Opin Biotechnol"},{"key":"bib59","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1016\/0014-5793(95)00062-E","article-title":"An exact mathematical expression for describing competitive binding of two different ligands to a protein molecule","volume":"360","author":"Wang","year":"1995","journal-title":"FEBS Lett"},{"key":"bib60","doi-asserted-by":"publisher","first-page":"1270","DOI":"10.1016\/S0960-9822(02)01035-7","article-title":"Interaction of cortactin and N-WASp with Arp2\/3 complex","volume":"12","author":"Weaver","year":"2002","journal-title":"Curr Biol"},{"key":"bib61","doi-asserted-by":"publisher","first-page":"370","DOI":"10.1016\/S0960-9822(01)00098-7","article-title":"Cortactin promotes and stabilizes Arp2\/3-induced actin filament network formation","volume":"11","author":"Weaver","year":"2001","journal-title":"Curr Biol"},{"key":"bib62","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1083\/jcb.151.1.29","article-title":"Cortactin localization to sites of actin assembly in lamellipodia requires interactions with F-actin and the Arp2\/3 complex","volume":"151","author":"Weed","year":"2000","journal-title":"J Cell Biol"},{"key":"bib63","doi-asserted-by":"publisher","first-page":"6994","DOI":"10.1021\/bi00418a049","article-title":"Rate of binding of tropomyosin to actin filaments","volume":"27","author":"Wegner","year":"1988","journal-title":"Biochemistry"},{"key":"bib64","doi-asserted-by":"publisher","first-page":"1417","DOI":"10.1083\/jcb.120.6.1417","article-title":"Cortactin, an 80\/85-kilodalton Pp60src substrate, is a filamentous actin-binding protein enriched in the cell cortex","volume":"120","author":"Wu","year":"1993","journal-title":"J Cell Biol"},{"key":"bib65","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1083\/jcb.200407076","article-title":"Molecular mechanisms of invadopodium formation: the role of the N-WASP-Arp2\/3 complex pathway and cofilin","volume":"168","author":"Yamaguchi","year":"2005","journal-title":"J Cell Biol"}],"container-title":["eLife"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/cdn.elifesciences.org\/articles\/00884\/elife-00884-v1.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/cdn.elifesciences.org\/articles\/00884\/elife-00884-v1.xml","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/elifesciences.org\/articles\/00884","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,10,11]],"date-time":"2023-10-11T20:40:44Z","timestamp":1697056844000},"score":1,"resource":{"primary":{"URL":"https:\/\/elifesciences.org\/articles\/00884"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2013,9,3]]},"references-count":63,"alternative-id":["10.7554\/eLife.00884"],"URL":"https:\/\/doi.org\/10.7554\/elife.00884","archive":["CLOCKSS"],"relation":{"references":[{"id-type":"accession","id":"2A41","asserted-by":"subject"},{"id-type":"accession","id":"2VCP","asserted-by":"subject"}],"has-review":[{"id-type":"doi","id":"10.7554\/eLife.00884.025","asserted-by":"object"},{"id-type":"doi","id":"10.7554\/eLife.00884.024","asserted-by":"object"}]},"ISSN":["2050-084X"],"issn-type":[{"value":"2050-084X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2013,9,3]]},"assertion":[{"value":"2013-04-29","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2013-07-31","order":1,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2013-09-03","order":2,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e00884"}}