{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,15]],"date-time":"2026-04-15T14:35:55Z","timestamp":1776263755871,"version":"3.50.1"},"reference-count":59,"publisher":"Oxford University Press (OUP)","issue":"2","license":[{"start":{"date-parts":[[2007,5,17]],"date-time":"2007-05-17T00:00:00Z","timestamp":1179360000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"name":"Ministerio de Educaci\u00f3n y Ciencia","award":["SAF2005-02119"],"award-info":[{"award-number":["SAF2005-02119"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2007,8,1]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>The \u03b14\u03b21 integrin is expressed on thymocytes and mediates cell attachment to its ligands CS-1\/fibronectin (CS-1\/FN) and VCAM-1 in the thymus. The chemokine CCL25 is highly expressed in the thymus, where it binds to its receptor CCR9 on thymocytes promoting migration and activation. We show here that \u03b14\u03b21 and CCR9 are coexpressed mainly on double- and single-positive thymocytes and that CCL25 strongly stimulates CD4+CD8+ and CD4+CD8\u2212 adhesion to CS-1\/FN and VCAM-1. CCL25 rapidly activated the GTPases Rac and Rap1 on thymocytes, and this activation was required for stimulation of adhesion, as detected using the CCR9+\/\u03b14\u03b21+ human T cell line Molt-4. To study the role on CCL25-stimulated adhesion of the Rac downstream effector Wiskott-Aldrich syndrome protein family verproline-homologous protein 2 (WAVE2) as well as of Rap1-GTP-interacting proteins, regulator of adhesion and cell polarization enriched in lymphoid tissues (RAPL) and Rap1-GTP-interacting adapter molecule (RIAM), we knocked down their expression and tested transfectant attachment to \u03b14\u03b21 ligands. We found that WAVE2 and RAPL but not RIAM were required for efficient triggering by CCL25 of T cell adhesion to CS-1\/FN and VCAM-1. Although Rac and Rap1 activation was required during early steps of T cell adhesion stimulated by CCL25, WAVE2 was needed for the development of actin-dependent T cell spreading subsequent to adhesion strengthening but not during initial \u03b14\u03b21-ligand interactions. These results suggest that regulation by CCL25 of adhesion of thymocyte subpopulations mediated by \u03b14\u03b21 could contribute to control their trafficking in the thymus during maturation, and identify Rac-WAVE2 and Rap1-RAPL as pathways whose activation is required in inside-out signaling, leading to stimulated adhesion.<\/jats:p>","DOI":"10.1189\/jlb.1206726","type":"journal-article","created":{"date-parts":[[2007,5,18]],"date-time":"2007-05-18T17:11:07Z","timestamp":1179508267000},"page":"380-391","source":"Crossref","is-referenced-by-count":30,"title":["Intracellular signaling required for CCL25-stimulated T cell adhesion mediated by the integrin \u03b14\u03b21"],"prefix":"10.1093","volume":"82","author":[{"given":"Marisa","family":"Parmo-Caba\u00f1as","sequence":"first","affiliation":[{"name":"Department of Molecular and Cellular Physiopathology, Centro de Investigaciones Biol\u00f3gicas (CSIC) , Madrid , Spain"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"David","family":"Garc\u00eda-Bernal","sequence":"additional","affiliation":[{"name":"Department of Molecular and Cellular Physiopathology, Centro de Investigaciones Biol\u00f3gicas (CSIC) , Madrid , Spain"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Rosa","family":"Garc\u00eda-Verdugo","sequence":"additional","affiliation":[{"name":"Department of Molecular and Cellular Physiopathology, Centro de Investigaciones Biol\u00f3gicas (CSIC) , Madrid , Spain"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Leonor","family":"Kremer","sequence":"additional","affiliation":[{"name":"Protein Tools Unit, Centro Nacional de Biotecnolog\u00eda (CSIC) , Madrid , Spain"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Gabriel","family":"M\u00e1rquez","sequence":"additional","affiliation":[{"name":"Department of Immunology and Oncology, Centro Nacional de Biotecnolog\u00eda (CSIC) , Madrid , Spain"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Joaquin","family":"Teixid\u00f3","sequence":"additional","affiliation":[{"name":"Department of Molecular and Cellular Physiopathology, Centro de Investigaciones Biol\u00f3gicas (CSIC) , Madrid , Spain"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"286","published-online":{"date-parts":[[2007,5,17]]},"reference":[{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"1722","DOI":"10.1172\/JCI117519","article-title":"The pathophysiologic role of \u03b1 4 integrins in vivo","volume":"94","author":"Lobb","year":"1994","journal-title":"J. Clin. Invest."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1034\/j.1600-065X.2002.18611.x","article-title":"\u03b14 Integrins and the immune response","volume":"186","author":"Rose","year":"2002","journal-title":"Immunol. Rev."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"3517","DOI":"10.4049\/jimmunol.149.11.3517","article-title":"Expression of VLA-4 on thymocytes. Maturation stage-associated transition and its correlation with their capacity to adhere to thymic stromal cells","volume":"149","author":"Sawada","year":"1992","journal-title":"J. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"4206","DOI":"10.1182\/blood.V86.11.4206.bloodjournal86114206","article-title":"Differential expression of integrins on human thymocyte subpopulations","volume":"86","author":"Mojcik","year":"1995","journal-title":"Blood"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"1573","DOI":"10.1084\/jem.179.5.1573","article-title":"Constitutive activation of integrin \u03b1 4 \u03b2 1 defines a unique stage of human thymocyte development","volume":"179","author":"Salomon","year":"1994","journal-title":"J. Exp. Med."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"5685","DOI":"10.1073\/pnas.88.13.5685","article-title":"Adhesion of immature thymocytes to thymic stromal cells through fibronectin molecules and its significance for the induction of thymocyte differentiation","volume":"88","author":"Utsumi","year":"1991","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"73","DOI":"10.4049\/jimmunol.153.1.73","article-title":"Modulation of fibronectin and thymic stromal cell-dependent thymocyte maturation by retinoic acid","volume":"153","author":"Meco","year":"1994","journal-title":"J. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1084\/jem.184.1.215","article-title":"Cell adhesion and migration are regulated at distinct stages of thymic T cell development: the roles of fibronectin, VLA4, and VLA5","volume":"184","author":"Crisa","year":"1996","journal-title":"J. Exp. Med."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"2461","DOI":"10.1182\/blood.V89.7.2461","article-title":"Vascular cell adhesion molecule-1 is expressed by cortical thymic epithelial cells and mediates thymocyte adhesion. Implications for the function of \u03b14\u03b21 (VLA4) integrin in T-cell development","volume":"89","author":"Salomon","year":"1997","journal-title":"Blood"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1146\/annurev.immunol.18.1.217","article-title":"The biology of chemokines and their receptors","volume":"18","author":"Rossi","year":"2000","journal-title":"Annu. Rev. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1038\/84298","article-title":"Chemokines: immunology\u2019s high impact factors","volume":"2","author":"Mackay","year":"2001","journal-title":"Nat. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1038\/84209","article-title":"Chemokines and disease","volume":"2","author":"Gerard","year":"2001","journal-title":"Nat. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1038\/nri722","article-title":"Chemokine receptors: multifaceted therapeutic targets","volume":"2","author":"Proudfoot","year":"2002","journal-title":"Nat. Rev. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1016\/S1074-7613(00)80531-2","article-title":"TECK: a novel CC chemokine specifically expressed by thymic dendritic cells and potentially involved in T cell development","volume":"7","author":"Vicari","year":"1997","journal-title":"Immunity"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1002\/1521-4141(200001)30:1<262::AID-IMMU262>3.0.CO;2-0","article-title":"The chemokine TECK is expressed by thymic and intestinal epithelial cells and attracts double- and single-positive thymocytes expressing the TECK receptor CCR9","volume":"30","author":"Wurbel","year":"2000","journal-title":"Eur. J. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"761","DOI":"10.1084\/jem.192.5.761","article-title":"Lymphocyte CC chemokine receptor 9 and epithelial thymus-expressed chemokine (TECK) expression distinguish the small intestinal immune compartment: Epithelial expression of tissue-specific chemokines as an organizing principle in regional immunity","volume":"192","author":"Kunkel","year":"2000","journal-title":"J. Exp. Med."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"5671","DOI":"10.4049\/jimmunol.162.10.5671","article-title":"Cutting edge: identification of the orphan chemokine receptor GPR-9-6 as CCR9, the receptor for the chemokine TECK","volume":"162","author":"Zaballos","year":"1999","journal-title":"J. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"1241","DOI":"10.1084\/jem.190.9.1241","article-title":"Human G protein-coupled receptor GPR-9-6\/CC chemokine receptor 9 is selectively expressed on intestinal homing T lymphocytes, mucosal lymphocytes, and thymocytes and is required for thymus-expressed chemokine-mediated chemotaxis","volume":"190","author":"Zabel","year":"1999","journal-title":"J. Exp. Med."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"2533","DOI":"10.1182\/blood.V94.7.2533.419k37_2533_2536","article-title":"TECK, an efficacious chemoattractant for human thymocytes, uses GPR-9-6\/CCR9 as a specific receptor","volume":"94","author":"Youn","year":"1999","journal-title":"Blood"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"850","DOI":"10.1182\/blood.V97.4.850","article-title":"Expression of CCR9 \u03b2-chemokine receptor is modulated in thymocyte differentiation and is selectively maintained in CD8(+) T cells from secondary lymphoid organs","volume":"97","author":"Carramolino","year":"2001","journal-title":"Blood"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"134","DOI":"10.4049\/jimmunol.168.1.134","article-title":"Characterization of CCR9 expression and CCL25\/thymus-expressed chemokine responsiveness during T cell development: CD3(high)CD69+ thymocytes and \u03b3\u03b4TCR+ thymocytes preferentially respond to CCL25","volume":"168","author":"Uehara","year":"2002","journal-title":"J. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"1182","DOI":"10.1038\/ni1275","article-title":"Immune cell migration in inflammation: present and future therapeutic targets","volume":"6","author":"Luster","year":"2005","journal-title":"Nat. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"546","DOI":"10.1038\/nri1646","article-title":"Intracellular signaling controlling integrin activation in lymphocytes","volume":"5","author":"Kinashi","year":"2005","journal-title":"Nat. Rev. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"3223","DOI":"10.1091\/mbc.e04-12-1049","article-title":"Vav1 and Rac control chemokine-promoted T lymphocyte adhesion mediated by the integrin {\u03b1}4{\u03b2}1","volume":"16","author":"Garcia-Bernal","year":"2005","journal-title":"Mol. Biol. Cell"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1083\/jcb.200301133","article-title":"Rap1 translates chemokine signals to integrin activation, cell polarization, and motility across vascular endothelium under flow","volume":"161","author":"Shimonaka","year":"2003","journal-title":"J. Cell Biol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"1045","DOI":"10.1038\/ni1111","article-title":"Crucial functions of the Rap1 effector molecule RAPL in lymphocyte and dendritic cell trafficking","volume":"5","author":"Katagiri","year":"2004","journal-title":"Nat. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"6932","DOI":"10.1093\/emboj\/17.23.6932","article-title":"WAVE, a novel WASP-family protein involved in actin reorganization induced by Rac","volume":"17","author":"Miki","year":"1998","journal-title":"EMBO J."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"296","DOI":"10.1006\/bbrc.1999.0894","article-title":"Identification of two human WAVE\/SCAR homologues as general actin regulatory molecules which associate with the Arp2\/3 complex","volume":"260","author":"Suetsugu","year":"1999","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"790","DOI":"10.1038\/nature00859","article-title":"Mechanism of regulation of WAVE1-induced actin nucleation by Rac1 and Nck","volume":"418","author":"Eden","year":"2002","journal-title":"Nature"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1016\/j.tcb.2004.04.007","article-title":"Regulation of actin dynamics by WASP and WAVE family proteins","volume":"14","author":"Stradal","year":"2004","journal-title":"Trends Cell Biol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"585","DOI":"10.1016\/j.devcel.2004.07.021","article-title":"RIAM, an Ena\/VASP and profilin ligand, interacts with Rap1-GTP and mediates Rap1-induced adhesion","volume":"7","author":"Lafuente","year":"2004","journal-title":"Dev. Cell"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1016\/j.ceb.2005.02.009","article-title":"Linking Rap to cell adhesion","volume":"17","author":"Bos","year":"2005","journal-title":"Curr. Opin. Cell Biol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"741","DOI":"10.1038\/ni950","article-title":"RAPL, a Rap1-binding molecule that mediates Rap1-induced adhesion through spatial regulation of LFA-1","volume":"4","author":"Katagiri","year":"2003","journal-title":"Nat. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"2696","DOI":"10.1074\/jbc.271.5.2696","article-title":"A region of the integrin VLA \u03b1 4 subunit involved in homotypic cell aggregation and in fibronectin but not vascular cell adhesion molecule-1 binding","volume":"271","author":"Munoz","year":"1996","journal-title":"J. Biol. Chem."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"27224","DOI":"10.1016\/S0021-9258(18)46972-4","article-title":"Integrin \u03b1 4 \u03b2 1-mediated melanoma cell adhesion and migration on vascular cell adhesion molecule-1 (VCAM-1) and the alternatively spliced IIICS region of fibronectin","volume":"269","author":"Mould","year":"1994","journal-title":"J. Biol. Chem."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1091\/mbc.e02-05-0275","article-title":"Rapid up-regulation of \u03b14 integrin-mediated leukocyte adhesion by transforming growth factor-\u03b21","volume":"14","author":"Bartolome","year":"2003","journal-title":"Mol. Biol. Cell"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"1978","DOI":"10.1182\/blood-2002-10-3190","article-title":"Transforming growth factor-\u03b21 down-regulates expression of chemokine stromal cell-derived factor-1: functional consequences in cell migration and adhesion","volume":"102","author":"Wright","year":"2003","journal-title":"Blood"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"703","DOI":"10.1016\/S1074-7613(01)00235-7","article-title":"The GTPase Rac-1 controls cell fate in the thymus by diverting thymocytes from positive to negative selection","volume":"15","author":"Gomez","year":"2001","journal-title":"Immunity"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1038\/ni765","article-title":"Rap1A positively regulates T cells via integrin activation rather than inhibiting lymphocyte signaling","volume":"3","author":"Sebzda","year":"2002","journal-title":"Nat. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"1801","DOI":"10.1242\/jcs.114.10.1801","article-title":"WASP and WAVE family proteins: key molecules for rapid rearrangement of cortical actin filaments and cell movement","volume":"114","author":"Takenawa","year":"2001","journal-title":"J. Cell Sci."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"749","DOI":"10.1038\/sj.emboj.7600084","article-title":"Sra-1 and Nap1 link Rac to actin assembly driving lamellipodia formation","volume":"23","author":"Steffen","year":"2004","journal-title":"EMBO J."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.cub.2005.11.036","article-title":"The WAVE2 complex regulates actin cytoskeletal reorganization and CRAC-mediated calcium entry during T cell activation","volume":"16","author":"Nolz","year":"2006","journal-title":"Curr. Biol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1016\/j.coi.2005.04.003","article-title":"Vav-family proteins in T-cell signaling","volume":"17","author":"Tybulewicz","year":"2005","journal-title":"Curr. Opin. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"3111","DOI":"10.1182\/blood.V99.9.3111","article-title":"Signaling through ZAP-70 is required for CXCL12-mediated T-cell transendothelial migration","volume":"99","author":"Ticchioni","year":"2002","journal-title":"Blood"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"131","DOI":"10.3109\/15419069509081282","article-title":"Vascular cell adhesion molecule (VCAM)-Ig fusion protein defines distinct affinity states of the very late antigen-4 (VLA-4) receptor","volume":"3","author":"Jakubowski","year":"1995","journal-title":"Cell Adhes. Commun."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"28740","DOI":"10.1074\/jbc.270.48.28740","article-title":"\u03b1 4 \u03b2 1 Integrin-dependent cell adhesion is regulated by a low affinity receptor pool that is conformationally responsive to ligand","volume":"270","author":"Yednock","year":"1995","journal-title":"J. Biol. Chem."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"1704","DOI":"10.1126\/science.1092053","article-title":"Cell migration: integrating signals from front to back","volume":"302","author":"Ridley","year":"2003","journal-title":"Science"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"859","DOI":"10.1038\/nri1223","article-title":"Cell migration and the control of post-natal T-cell lymphopoiesis in the thymus","volume":"3","author":"Petrie","year":"2003","journal-title":"Nat. Rev. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"338","DOI":"10.1038\/ni1323","article-title":"Thymic microenvironments for T cell differentiation and selection","volume":"7","author":"Ladi","year":"2006","journal-title":"Nat. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1016\/j.coi.2005.02.001","article-title":"Controlling the thymic microenvironment","volume":"17","author":"Gray","year":"2005","journal-title":"Curr. Opin. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"3652","DOI":"10.1002\/eji.200425248","article-title":"Homing of immature thymocytes to the subcapsular microenvironment within the thymus is not an absolute requirement for T cell development","volume":"34","author":"Benz","year":"2004","journal-title":"Eur. J. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"7006","DOI":"10.1073\/pnas.0602024103","article-title":"A multistep adhesion cascade for lymphoid progenitor cell homing to the thymus","volume":"103","author":"Scimone","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"2626","DOI":"10.1182\/blood.V98.9.2626","article-title":"Mice lacking the CCR9 CC-chemokine receptor show a mild impairment of early T- and B-cell development and a reduction in T-cell receptor \u03b3\u03b4(+) gut intraepithelial lymphocytes","volume":"98","author":"Wurbel","year":"2001","journal-title":"Blood"},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"2811","DOI":"10.4049\/jimmunol.168.6.2811","article-title":"A role for CCR9 in T lymphocyte development and migration","volume":"168","author":"Uehara","year":"2002","journal-title":"J. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1038\/ncb1105","article-title":"Abi1 is essential for the formation and activation of a WAVE2 signaling complex","volume":"6","author":"Innocenti","year":"2004","journal-title":"Nat. Cell Biol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"547","DOI":"10.1016\/j.ceb.2003.08.003","article-title":"Integrin avidity regulation: are changes in affinity and conformation underemphasized?","volume":"15","author":"Carman","year":"2003","journal-title":"Curr. Opin. Cell Biol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1083\/jcb.200206011","article-title":"Integrin activation takes shape","volume":"158","author":"Liddington","year":"2002","journal-title":"J. Cell Biol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"350","DOI":"10.4049\/jimmunol.175.1.350","article-title":"Integrin regulation by RhoA in thymocytes","volume":"175","author":"Vielkind","year":"2005","journal-title":"J. Immunol."},{"key":"2023033006514933800_","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1083\/jcb.200404068","article-title":"Rap1 promotes cell spreading by localizing Rac guanine nucleotide exchange factors","volume":"167","author":"Arthur","year":"2004","journal-title":"J. Cell Biol."}],"container-title":["Journal of Leukocyte Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.wiley.com\/onlinelibrary\/tdm\/v1\/articles\/10.1189%2Fjlb.1206726","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/academic.oup.com\/jleukbio\/article-pdf\/82\/2\/380\/49637488\/jlb0380.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jleukbio\/article-pdf\/82\/2\/380\/49637488\/jlb0380.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,5,11]],"date-time":"2023-05-11T22:21:24Z","timestamp":1683843684000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/jleukbio\/article\/82\/2\/380\/6975806"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2007,5,17]]},"references-count":59,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2007,5,17]]},"published-print":{"date-parts":[[2007,8,1]]}},"URL":"https:\/\/doi.org\/10.1189\/jlb.1206726","relation":{},"ISSN":["0741-5400","1938-3673"],"issn-type":[{"value":"0741-5400","type":"print"},{"value":"1938-3673","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2007,8]]},"published":{"date-parts":[[2007,5,17]]}}}