{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T17:14:23Z","timestamp":1790874863306,"version":"4.1.0"},"reference-count":59,"publisher":"Public Library of Science (PLoS)","issue":"2","license":[{"start":{"date-parts":[[2012,2,22]],"date-time":"2012-02-22T00:00:00Z","timestamp":1329868800000},"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.0032081","type":"journal-article","created":{"date-parts":[[2012,2,22]],"date-time":"2012-02-22T22:12:48Z","timestamp":1329948768000},"page":"e32081","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":43,"title":["Receptor-Specific Mechanisms Regulate Phosphorylation of AKT at Ser473: Role of RICTOR in \u03b21 Integrin-Mediated Cell Survival"],"prefix":"10.1371","volume":"7","author":[{"given":"Anjum","family":"Riaz","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kathrin Stephanie","family":"Zeller","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Staffan","family":"Johansson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2012,2,22]]},"reference":[{"key":"ref1","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1016\/j.bbrc.2005.01.125","article-title":"Integrin alpha8beta1-fibronectin interactions promote cell survival via PI3 kinase pathway.","volume":"329","author":"E Farias","year":"2005","journal-title":"Biochem Biophys Res Commun"},{"key":"ref2","doi-asserted-by":"crossref","first-page":"738137","DOI":"10.1155\/2011\/738137","article-title":"Integrin signaling, cell survival, and anoikis: distinctions, differences, and differentiation.","volume":"2011","author":"PH Vachon","year":"2011","journal-title":"J Signal Transduct"},{"key":"ref3","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1042\/BST0340647","article-title":"Signalling through Class I PI3Ks in mammalian cells.","volume":"34","author":"PT Hawkins","year":"2006","journal-title":"Biochem Soc Trans"},{"key":"ref4","doi-asserted-by":"crossref","first-page":"710","DOI":"10.1126\/science.279.5351.710","article-title":"Protein kinase B kinases that mediate phosphatidylinositol 3,4,5-trisphosphate-dependent activation of protein kinase B.","volume":"279","author":"L Stephens","year":"1998","journal-title":"Science"},{"key":"ref5","doi-asserted-by":"crossref","first-page":"567","DOI":"10.1126\/science.277.5325.567","article-title":"Dual role of phosphatidylinositol-3,4,5-trisphosphate in the activation of protein kinase B.","volume":"277","author":"D Stokoe","year":"1997","journal-title":"Science"},{"key":"ref6","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1016\/S0014-5793(96)01370-1","article-title":"Molecular basis for the substrate specificity of protein kinase B; comparison with MAPKAP kinase-1 and p70 S6 kinase.","volume":"399","author":"DR Alessi","year":"1996","journal-title":"FEBS Lett"},{"key":"ref7","doi-asserted-by":"crossref","first-page":"31858","DOI":"10.1074\/jbc.C100271200","article-title":"Regulation of Akt\/PKB activation by tyrosine phosphorylation.","volume":"276","author":"R Chen","year":"2001","journal-title":"J Biol Chem"},{"key":"ref8","doi-asserted-by":"crossref","first-page":"38021","DOI":"10.1074\/jbc.M203387200","article-title":"Direct identification of tyrosine 474 as a regulatory phosphorylation site for the Akt protein kinase.","volume":"277","author":"NM Conus","year":"2002","journal-title":"J Biol Chem"},{"key":"ref9","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1016\/S0960-9822(06)00122-9","article-title":"Characterization of a 3-phosphoinositide-dependent protein kinase which phosphorylates and activates protein kinase Balpha.","volume":"7","author":"DR Alessi","year":"1997","journal-title":"Curr Biol"},{"key":"ref10","doi-asserted-by":"crossref","first-page":"E187","DOI":"10.1152\/ajpendo.00011.2005","article-title":"PDK2: the missing piece in the receptor tyrosine kinase signaling pathway puzzle.","volume":"289","author":"LQ Dong","year":"2005","journal-title":"Am J Physiol Endocrinol Metab"},{"key":"ref11","doi-asserted-by":"crossref","first-page":"27462","DOI":"10.1074\/jbc.M102940200","article-title":"Regulation of protein kinase B\/Akt-serine 473 phosphorylation by integrin-linked kinase: critical roles for kinase activity and amino acids arginine 211 and serine 343.","volume":"276","author":"S Persad","year":"2001","journal-title":"J Biol Chem"},{"key":"ref12","doi-asserted-by":"crossref","first-page":"41189","DOI":"10.1074\/jbc.M406731200","article-title":"Identification of a PKB\/Akt hydrophobic motif Ser-473 kinase as DNA-dependent protein kinase.","volume":"279","author":"J Feng","year":"2004","journal-title":"J Biol Chem"},{"key":"ref13","doi-asserted-by":"crossref","first-page":"4029","DOI":"10.1074\/jbc.M410344200","article-title":"Full activation of PKB\/Akt in response to insulin or ionizing radiation is mediated through ATM.","volume":"280","author":"JG Viniegra","year":"2005","journal-title":"J Biol Chem"},{"key":"ref14","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1124\/mol.107.044743","article-title":"AKT is activated in an ataxia-telangiectasia and Rad3-related-dependent manner in response to temozolomide and confers protection against drug-induced cell growth inhibition.","volume":"74","author":"S Caporali","year":"2008","journal-title":"Mol Pharmacol"},{"key":"ref15","doi-asserted-by":"crossref","first-page":"951","DOI":"10.1016\/j.cellsig.2004.01.008","article-title":"Regulation of protein kinase B\/Akt activity and Ser473 phosphorylation by protein kinase Calpha in endothelial cells.","volume":"16","author":"C Partovian","year":"2004","journal-title":"Cell Signal"},{"key":"ref16","doi-asserted-by":"crossref","first-page":"1098","DOI":"10.1126\/science.1106148","article-title":"Phosphorylation and regulation of Akt\/PKB by the rictor-mTOR complex.","volume":"307","author":"DD Sarbassov","year":"2005","journal-title":"Science"},{"key":"ref17","doi-asserted-by":"crossref","first-page":"30025","DOI":"10.1074\/jbc.M803053200","article-title":"In vivo analysis of protein kinase B (PKB)\/Akt regulation in DNA-PKcs-null mice reveals a role for PKB\/Akt in DNA damage response and tumorigenesis.","volume":"283","author":"B Surucu","year":"2008","journal-title":"J Biol Chem"},{"key":"ref18","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1016\/j.molcel.2008.02.024","article-title":"PKBalpha\/Akt1 acts downstream of DNA-PK in the DNA double-strand break response and promotes survival.","volume":"30","author":"L Bozulic","year":"2008","journal-title":"Mol Cell"},{"key":"ref19","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1158\/0008-5472.CAN-05-2304","article-title":"Preferential dependence of breast cancer cells versus normal cells on integrin-linked kinase for protein kinase B\/Akt activation and cell survival.","volume":"66","author":"AA Troussard","year":"2006","journal-title":"Cancer Res"},{"key":"ref20","doi-asserted-by":"crossref","first-page":"22374","DOI":"10.1074\/jbc.M303083200","article-title":"Conditional knock-out of integrin-linked kinase demonstrates an essential role in protein kinase B\/Akt activation.","volume":"278","author":"AA Troussard","year":"2003","journal-title":"J Biol Chem"},{"key":"ref21","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1038\/emboj.2009.376","article-title":"The ILK\/PINCH\/parvin complex: the kinase is dead, long live the pseudokinase!","volume":"29","author":"SA Wickstrom","year":"2010","journal-title":"EMBO J"},{"key":"ref22","doi-asserted-by":"crossref","first-page":"e12356","DOI":"10.1371\/journal.pone.0012356","article-title":"Integrin-linked kinase is a functional Mn2+-dependent protein kinase that regulates glycogen synthase kinase-3beta (GSK-3beta) phosphorylation.","volume":"5","author":"M Maydan","year":"2010","journal-title":"PLoS One"},{"key":"ref23","doi-asserted-by":"crossref","first-page":"21886","DOI":"10.1074\/jbc.M111.240093","article-title":"Biochemical, proteomic, structural, and thermodynamic characterizations of integrin-linked kinase (ILK): cross-validation of the pseudokinase.","volume":"286","author":"K Fukuda","year":"2011","journal-title":"J Biol Chem"},{"key":"ref24","doi-asserted-by":"crossref","first-page":"1296","DOI":"10.1016\/j.cub.2004.06.054","article-title":"Rictor, a novel binding partner of mTOR, defines a rapamycin-insensitive and raptor-independent pathway that regulates the cytoskeleton.","volume":"14","author":"DD Sarbassov","year":"2004","journal-title":"Curr Biol"},{"key":"ref25","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1016\/j.cell.2006.08.033","article-title":"SIN1\/MIP1 maintains rictor-mTOR complex integrity and regulates Akt phosphorylation and substrate specificity.","volume":"127","author":"E Jacinto","year":"2006","journal-title":"Cell"},{"key":"ref26","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1016\/j.devcel.2006.08.013","article-title":"Multiallelic disruption of the rictor gene in mice reveals that mTOR complex 2 is essential for fetal growth and viability.","volume":"11","author":"C Shiota","year":"2006","journal-title":"Dev Cell"},{"key":"ref27","doi-asserted-by":"crossref","first-page":"40406","DOI":"10.1074\/jbc.M508361200","article-title":"mTOR.RICTOR is the Ser473 kinase for Akt\/protein kinase B in 3T3-L1 adipocytes.","volume":"280","author":"RC Hresko","year":"2005","journal-title":"J Biol Chem"},{"key":"ref28","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1016\/j.yjmcc.2007.10.016","article-title":"Regulation of Akt\/PKB activity by P21-activated kinase in cardiomyocytes.","volume":"44","author":"K Mao","year":"2008","journal-title":"J Mol Cell Cardiol"},{"key":"ref29","doi-asserted-by":"crossref","first-page":"6169","DOI":"10.1074\/jbc.C800210200","article-title":"DNA-dependent protein kinase-mediated phosphorylation of protein kinase B requires a specific recognition sequence in the C-terminal hydrophobic motif.","volume":"284","author":"J Park","year":"2009","journal-title":"J Biol Chem"},{"key":"ref30","doi-asserted-by":"crossref","first-page":"1356","DOI":"10.1038\/ncb1795","article-title":"Scaffolding function of PAK in the PDK1-Akt pathway.","volume":"10","author":"M Higuchi","year":"2008","journal-title":"Nat Cell Biol"},{"key":"ref31","doi-asserted-by":"crossref","first-page":"1106","DOI":"10.1016\/j.bbamcr.2010.05.007","article-title":"P21-activated kinase 1 stimulates colon cancer cell growth and migration\/invasion via ERK- and AKT-dependent pathways.","volume":"1803","author":"N Huynh","year":"2010","journal-title":"Biochim Biophys Acta"},{"key":"ref32","doi-asserted-by":"crossref","first-page":"5839","DOI":"10.1038\/onc.2010.318","article-title":"A Rac-Pak signaling pathway is essential for ErbB2-mediated transformation of human breast epithelial cancer cells.","volume":"29","author":"LE Arias-Romero","year":"2010","journal-title":"Oncogene"},{"key":"ref33","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1016\/j.yexcr.2007.08.018","article-title":"EGFR and beta1 integrins utilize different signaling pathways to activate Akt.","volume":"314","author":"T Velling","year":"2008","journal-title":"Exp Cell Res"},{"key":"ref34","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1038\/379091a0","article-title":"Regulation of cell adhesion and anchorage-dependent growth by a new beta 1-integrin-linked protein kinase.","volume":"379","author":"GE Hannigan","year":"1996","journal-title":"Nature"},{"key":"ref35","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1038\/nrc1524","article-title":"Integrin-linked kinase: a cancer therapeutic target unique among its ILK.","volume":"5","author":"G Hannigan","year":"2005","journal-title":"Nat Rev Cancer"},{"key":"ref36","doi-asserted-by":"crossref","first-page":"1007","DOI":"10.1083\/jcb.152.5.1007","article-title":"Drosophila integrin-linked kinase is required at sites of integrin adhesion to link the cytoskeleton to the plasma membrane.","volume":"152","author":"CG Zervas","year":"2001","journal-title":"J Cell Biol"},{"key":"ref37","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1042\/BC20070109","article-title":"A tale of two Paks.","volume":"100","author":"LE Arias-Romero","year":"2008","journal-title":"Biol Cell"},{"key":"ref38","doi-asserted-by":"crossref","first-page":"36609","DOI":"10.1074\/jbc.M502306200","article-title":"Role of group A p21-activated kinases in activation of extracellular-regulated kinase by growth factors.","volume":"280","author":"A Beeser","year":"2005","journal-title":"J Biol Chem"},{"key":"ref39","doi-asserted-by":"crossref","first-page":"44113","DOI":"10.1074\/jbc.M405323200","article-title":"Human intestinal epithelial cell survival and anoikis. Differentiation state-distinct regulation and roles of protein kinase B\/Akt isoforms.","volume":"279","author":"G Dufour","year":"2004","journal-title":"J Biol Chem"},{"key":"ref40","doi-asserted-by":"crossref","first-page":"C580","DOI":"10.1152\/ajpcell.00375.2009","article-title":"The Akt isoforms are present at distinct subcellular locations.","volume":"298","author":"SA Santi","year":"2010","journal-title":"Am J Physiol Cell Physiol"},{"key":"ref41","doi-asserted-by":"crossref","first-page":"1352","DOI":"10.1016\/j.bcp.2008.07.023","article-title":"Anoikis: a necessary death program for anchorage-dependent cells.","volume":"76","author":"P Chiarugi","year":"2008","journal-title":"Biochem Pharmacol"},{"key":"ref42","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1042\/BJ20081056","article-title":"The life of a cell: apoptosis regulation by the PI3K\/PKB pathway.","volume":"415","author":"V Duronio","year":"2008","journal-title":"Biochem J"},{"key":"ref43","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1016\/j.cellsig.2007.07.010","article-title":"PKB and the mitochondria: AKTing on apoptosis.","volume":"20","author":"A Parcellier","year":"2008","journal-title":"Cell Signal"},{"key":"ref44","doi-asserted-by":"crossref","first-page":"660","DOI":"10.1124\/mol.62.3.660","article-title":"Akt activation induced by lysophosphatidic acid and sphingosine-1-phosphate requires both mitogen-activated protein kinase kinase and p38 mitogen-activated protein kinase and is cell-line specific.","volume":"62","author":"LM Baudhuin","year":"2002","journal-title":"Mol Pharmacol"},{"key":"ref45","doi-asserted-by":"crossref","first-page":"C1927","DOI":"10.1152\/ajpcell.00400.2006","article-title":"LPA2 (EDG4) mediates Rho-dependent chemotaxis with lower efficacy than LPA1 (EDG2) in breast carcinoma cells.","volume":"292","author":"M Chen","year":"2007","journal-title":"Am J Physiol Cell Physiol"},{"key":"ref46","doi-asserted-by":"crossref","first-page":"322","DOI":"10.1016\/j.chembiol.2008.03.005","article-title":"An isoform-selective, small-molecule inhibitor targets the autoregulatory mechanism of p21-activated kinase.","volume":"15","author":"SW Deacon","year":"2008","journal-title":"Chem Biol"},{"key":"ref47","doi-asserted-by":"crossref","first-page":"975","DOI":"10.2741\/1592","article-title":"AKT\/PKB signaling mechanisms in cancer and chemoresistance.","volume":"10","author":"D Kim","year":"2005","journal-title":"Front Biosci"},{"key":"ref48","doi-asserted-by":"crossref","first-page":"178","DOI":"10.1016\/j.ccr.2009.08.011","article-title":"A molecular link between AKT regulation and chemotherapeutic response.","volume":"16","author":"CC Dibble","year":"2009","journal-title":"Cancer Cell"},{"key":"ref49","doi-asserted-by":"crossref","first-page":"1227","DOI":"10.1016\/S1097-2765(02)00550-6","article-title":"Molecular mechanism for the regulation of protein kinase B\/Akt by hydrophobic motif phosphorylation.","volume":"9","author":"J Yang","year":"2002","journal-title":"Mol Cell"},{"key":"ref50","first-page":"19","article-title":"Physiological regulation of Akt activity and stability.","volume":"2","author":"Y Liao","year":"2010","journal-title":"Am J Transl Res"},{"key":"ref51","doi-asserted-by":"crossref","first-page":"1838","DOI":"10.1016\/j.cellsig.2010.07.011","article-title":"PI3-kinase p110alpha mediates beta1 integrin-induced Akt activation and membrane protrusion during cell attachment and initial spreading.","volume":"22","author":"KS Zeller","year":"2010","journal-title":"Cell Signal"},{"key":"ref52","doi-asserted-by":"crossref","first-page":"1618","DOI":"10.1158\/0008-5472.CAN-07-5869","article-title":"Rictor and integrin-linked kinase interact and regulate Akt phosphorylation and cancer cell survival.","volume":"68","author":"PC McDonald","year":"2008","journal-title":"Cancer Res"},{"key":"ref53","doi-asserted-by":"crossref","first-page":"3207","DOI":"10.1073\/pnas.97.7.3207","article-title":"Inhibition of integrin-linked kinase (ILK) suppresses activation of protein kinase B\/Akt and induces cell cycle arrest and apoptosis of PTEN-mutant prostate cancer cells.","volume":"97","author":"S Persad","year":"2000","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref54","doi-asserted-by":"crossref","first-page":"e14614","DOI":"10.1371\/journal.pone.0014614","article-title":"Ablation of Akt2 induces autophagy through cell cycle arrest, the downregulation of p70S6K, and the deregulation of mitochondria in MDA-MB231 cells.","volume":"6","author":"SA Santi","year":"2011","journal-title":"PLoS One"},{"key":"ref55","doi-asserted-by":"crossref","first-page":"5765","DOI":"10.1158\/0008-5472.CAN-11-0368","article-title":"The Tumor Suppressive MicroRNA miR-218 Targets the mTOR Component Rictor and Inhibits AKT Phosphorylation in Oral Cancer.","volume":"71","author":"A Uesugi","year":"2011","journal-title":"Cancer Res"},{"key":"ref56","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1016\/S1097-2765(05)00012-2","article-title":"14-3-3 proteins and survival kinases cooperate to inactivate BAD by BH3 domain phosphorylation.","volume":"6","author":"SR Datta","year":"2000","journal-title":"Mol Cell"},{"key":"ref57","doi-asserted-by":"crossref","first-page":"19080","DOI":"10.1074\/jbc.273.30.19080","article-title":"Activation of Akt\/protein kinase B by G protein-coupled receptors. A role for alpha and beta gamma subunits of heterotrimeric G proteins acting through phosphatidylinositol-3-OH kinasegamma.","volume":"273","author":"C Murga","year":"1998","journal-title":"J Biol Chem"},{"key":"ref58","doi-asserted-by":"crossref","first-page":"949","DOI":"10.1002\/jcb.20094","article-title":"G protein mediated signaling pathways in lysophospholipid induced cell proliferation and survival.","volume":"92","author":"J Radeff-Huang","year":"2004","journal-title":"J Cell Biochem"},{"key":"ref59","doi-asserted-by":"crossref","first-page":"5996","DOI":"10.1128\/MCB.00114-08","article-title":"Freud-1\/Aki1, a novel PDK1-interacting protein, functions as a scaffold to activate the PDK1\/Akt pathway in epidermal growth factor signaling.","volume":"28","author":"A Nakamura","year":"2008","journal-title":"Mol Cell Biol"}],"container-title":["PLoS ONE"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/dx.plos.org\/10.1371\/journal.pone.0032081","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2018,10,22]],"date-time":"2018-10-22T16:50:56Z","timestamp":1540227056000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0032081"}},"subtitle":[],"editor":[{"given":"Neil A.","family":"Hotchin","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2012,2,22]]},"references-count":59,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2012,2,22]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0032081","relation":{},"ISSN":["1932-6203"],"issn-type":[{"value":"1932-6203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2012,2,22]]}}}