{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T15:21:24Z","timestamp":1790868084945,"version":"4.1.0"},"reference-count":43,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2010,11,26]],"date-time":"2010-11-26T00:00:00Z","timestamp":1290729600000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/www.springer.com\/tdm"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Calcif Tissue Int"],"published-print":{"date-parts":[[2011,1]]},"DOI":"10.1007\/s00223-010-9435-3","type":"journal-article","created":{"date-parts":[[2010,11,25]],"date-time":"2010-11-25T04:26:38Z","timestamp":1290659198000},"page":"75-86","source":"Crossref","is-referenced-by-count":28,"title":["Osteoclasts Lacking Rac2 Have Defective Chemotaxis and Resorptive Activity"],"prefix":"10.1007","volume":"88","author":[{"given":"Takashi","family":"Itokowa","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mei-ling","family":"Zhu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nancy","family":"Troiano","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jessica","family":"Bian","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tustomu","family":"Kawano","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Karl","family":"Insogna","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2010,11,26]]},"reference":[{"key":"9435_CR1","first-page":"308","volume":"17","author":"GD Roodman","year":"1996","unstructured":"Roodman GD (1996) Advances in bone biology: the osteoclast. Endocr Rev 17:308\u2013332","journal-title":"Endocr Rev"},{"key":"9435_CR2","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1210\/en.140.1.273","volume":"140","author":"CK Lea","year":"1999","unstructured":"Lea CK, Sarma U, Flanagan AM (1999) Macrophage colony stimulating-factor transcripts are differentially regulated in rat bone-marrow by gender hormones. Endocrinology 140:273\u2013279","journal-title":"Endocrinology"},{"key":"9435_CR3","doi-asserted-by":"crossref","first-page":"1850","DOI":"10.1172\/JCI4561","volume":"102","author":"S Srivastava","year":"1998","unstructured":"Srivastava S, Weitzmann MN, Kimble RB, Rizzo M, Zahner M, Milbrandt J, Ross FP, Pacifici R (1998) Estrogen blocks M-CSF gene expression and osteoclast formation by regulating phosphorylation of Egr-1 and its interaction with Sp-1. J Clin Invest 102:1850\u20131859","journal-title":"J Clin Invest"},{"key":"9435_CR4","doi-asserted-by":"crossref","first-page":"1279","DOI":"10.1172\/JCI8672","volume":"105","author":"S Cenci","year":"2000","unstructured":"Cenci S, Weitzmann MN, Gentile MA, Aisa MC, Pacifici R (2000) M-CSF neutralization and egr-1 deficiency prevent ovariectomy-induced bone loss. J Clin Invest 105:1279\u20131287","journal-title":"J Clin Invest"},{"key":"9435_CR5","doi-asserted-by":"crossref","first-page":"1507","DOI":"10.1002\/jbmr.5650081214","volume":"8","author":"EC Weir","year":"1993","unstructured":"Weir EC, Horowitz MC, Baron R, Centrella M, Kacinski BM, Insogna KL (1993) Macrophage colony-stimulating factor release and receptor expression in bone cells. J Bone Miner Res 8:1507\u20131518","journal-title":"J Bone Miner Res"},{"key":"9435_CR6","doi-asserted-by":"crossref","first-page":"1401","DOI":"10.1006\/bbrc.1993.2199","volume":"195","author":"J Owens","year":"1993","unstructured":"Owens J, Chambers TJ (1993) Macrophage colony-stimulating factor (M-CSF) induces migration in osteoclasts in vitro. Biochem Biophys Res Commun 195:1401\u20131407","journal-title":"Biochem Biophys Res Commun"},{"key":"9435_CR7","doi-asserted-by":"crossref","first-page":"1733","DOI":"10.1084\/jem.178.5.1733","volume":"178","author":"K Fuller","year":"1993","unstructured":"Fuller K, Owens JM, Jagger CJ, Wilson A, Moss R, Chambers TJ (1993) Macrophage colony-stimulating factor stimulates survival and chemotactic behavior in isolated osteoclasts. J Exp Med 178:1733\u20131744","journal-title":"J Exp Med"},{"key":"9435_CR8","doi-asserted-by":"crossref","first-page":"2129","DOI":"10.1210\/en.141.6.2129","volume":"141","author":"A Grey","year":"2000","unstructured":"Grey A, Chen Y, Paliwal I, Carlberg K, Insogna K (2000) Evidence for a functional association between phosphatidylinositol 3-kinase and c-src in the spreading response of osteoclasts to colony-stimulating factor-1. Endocrinology 141:2129\u20132138","journal-title":"Endocrinology"},{"key":"9435_CR9","doi-asserted-by":"crossref","first-page":"1290","DOI":"10.1016\/j.bone.2006.06.012","volume":"39","author":"H Sakai","year":"2006","unstructured":"Sakai H, Chen Y, Itokawa T, Yu KP, Zhu ML, Insogna K (2006) Activated c-Fms recruits Vav and Rac during CSF-1-induced cytoskeletal remodeling and spreading in osteoclasts. Bone 39:1290\u20131301","journal-title":"Bone"},{"key":"9435_CR10","doi-asserted-by":"crossref","first-page":"1086","DOI":"10.1126\/science.8493552","volume":"260","author":"TP Stossel","year":"1993","unstructured":"Stossel TP (1993) On the crawling of animal cells. Science 260:1086\u20131094","journal-title":"Science"},{"key":"9435_CR11","doi-asserted-by":"crossref","first-page":"1102","DOI":"10.1126\/science.286.5442.1102","volume":"286","author":"AR Horwitz","year":"1999","unstructured":"Horwitz AR, Parsons JT (1999) Cell migration\u2014movin\u2019 on. Science 286:1102\u20131103","journal-title":"Science"},{"issue":"Pt 6","key":"9435_CR12","doi-asserted-by":"crossref","first-page":"707","DOI":"10.1242\/jcs.110.6.707","volume":"110","author":"WE Allen","year":"1997","unstructured":"Allen WE, Jones GE, Pollard JW, Ridley AJ (1997) Rho, Rac and Cdc42 regulate actin organization and cell adhesion in macrophages. J Cell Sci 110(Pt 6):707\u2013720","journal-title":"J Cell Sci"},{"key":"9435_CR13","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1038\/nm1194","volume":"11","author":"R Faccio","year":"2005","unstructured":"Faccio R, Teitelbaum SL, Fujikawa K, Chappel J, Zallone A, Tybulewicz VL, Ross FP, Swat W (2005) Vav3 regulates osteoclast function and bone mass. Nat Med 11:284\u2013290","journal-title":"Nat Med"},{"key":"9435_CR14","doi-asserted-by":"crossref","first-page":"2245","DOI":"10.1359\/JBMR.050816","volume":"20","author":"A Fukuda","year":"2005","unstructured":"Fukuda A, Hikita A, Wakeyama H, Akiyama T, Oda H, Nakamura K, Tanaka S (2005) Regulation of osteoclast apoptosis and motility by small GTPase binding protein Rac1. J Bone Miner Res 20:2245\u20132253","journal-title":"J Bone Miner Res"},{"key":"9435_CR15","doi-asserted-by":"crossref","first-page":"260","DOI":"10.1359\/jbmr.071013","volume":"23","author":"Y Wang","year":"2008","unstructured":"Wang Y, Lebowitz D, Sun C, Thang H, Grynpas MD, Glogauer M (2008) Identifying the relative contributions of rac1 and rac2 to osteoclastogenesis. J Bone Miner Res 23:260\u2013270","journal-title":"J Bone Miner Res"},{"key":"9435_CR16","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1016\/S1074-7613(00)80019-9","volume":"10","author":"AW Roberts","year":"1999","unstructured":"Roberts AW, Kim C, Zhen L, Lowe JB, Kapur R, Petryniak B, Spaetti A, Pollock JD, Borneo JB, Bradford GB, Atkinson SJ, Dinauer MC, Williams DA (1999) Deficiency of the hematopoietic cell\u2013specific Rho family GTPase Rac2 is characterized by abnormalities in neutrophil function and host defense. Immunity 10:183\u2013196","journal-title":"Immunity"},{"key":"9435_CR17","doi-asserted-by":"crossref","first-page":"1983","DOI":"10.1210\/en.2004-0770","volume":"146","author":"E Knopp","year":"2005","unstructured":"Knopp E, Troiano N, Bouxsein M, Sun BH, Lostritto K, Gundberg C, Dziura J, Insogna K (2005) The effect of aging on the skeletal response to intermittent treatment with parathyroid hormone. Endocrinology 146:1983\u20131990","journal-title":"Endocrinology"},{"key":"9435_CR18","doi-asserted-by":"crossref","first-page":"888","DOI":"10.1210\/endo-114-3-888","volume":"114","author":"KL Insogna","year":"1984","unstructured":"Insogna KL, Stewart AF, Vignery AM, Weir EC, Namnum PA, Baron RE, Kirkwood JM, Deftos LM, Broadus AE (1984) Biochemical and histomorphometric characterization of a rat model for humoral hypercalcemia of malignancy. Endocrinology 114:888\u2013896","journal-title":"Endocrinology"},{"key":"9435_CR19","doi-asserted-by":"crossref","first-page":"1704","DOI":"10.1359\/jbmr.060726","volume":"21","author":"S-K Lee","year":"2006","unstructured":"Lee S-K, Kadono Y, Okada F, Claire Jacquin C, Koczon-Jaremko B, Gloria Gronowicz G, Adams DJ, Aguila HL, Choi Y, Lorenzo JA (2006) T lymphocyte deficient mice lose trabecular bone mass with ovariectomy. J Bone Miner Res 21:1704\u20131712","journal-title":"J Bone Miner Res"},{"key":"9435_CR20","doi-asserted-by":"crossref","first-page":"8117","DOI":"10.1074\/jbc.M410076200","volume":"280","author":"JR Harrison","year":"2005","unstructured":"Harrison JR, Huang YF, Wilson KA, Kelly PL, Adams DJ, Gronowicz GA, Clark SH (2005) Col1a1 promoter-targeted expression of p20 CCAAT enhancer-binding protein beta (C\/EBPbeta), a truncated C\/EBPbeta isoform, causes osteopenia in transgenic mice. J Biol Chem 280:8117\u20138124","journal-title":"J Biol Chem"},{"key":"9435_CR21","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1002\/(SICI)1098-2795(199701)46:1<104::AID-MRD16>3.0.CO;2-2","volume":"46","author":"K Insogna","year":"1997","unstructured":"Insogna K, Tanaka S, Neff L, Horne W, Levy J, Baron R (1997) Role of c-Src in cellular events associated with colony-stimulating factor-1-induced spreading in osteoclasts. Mol Reprod Dev 46:104\u2013108","journal-title":"Mol Reprod Dev"},{"key":"9435_CR22","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1016\/S0076-6879(97)82110-6","volume":"282","author":"T Suda","year":"1997","unstructured":"Suda T, Jimi E, Nakamura I, Takahashi N (1997) Role of 1 alpha,25-dihydroxyvitamin D3 in osteoclast differentiation and function. Methods Enzymol 282:223\u2013235","journal-title":"Methods Enzymol"},{"key":"9435_CR23","doi-asserted-by":"crossref","first-page":"2476","DOI":"10.1172\/JCI119790","volume":"100","author":"KL Insogna","year":"1997","unstructured":"Insogna KL, Sahni M, Grey AB, Tanaka S, Horne WC, Neff L, Mitnick M, Levy JB, Baron R (1997) Colony-stimulating factor-1 induces cytoskeletal reorganization and c-src-dependent tyrosine phosphorylation of selected cellular proteins in rodent osteoclasts. J Clin Invest 100:2476\u20132485","journal-title":"J Clin Invest"},{"key":"9435_CR24","doi-asserted-by":"crossref","first-page":"S136","DOI":"10.1016\/j.bone.2009.01.299","volume":"44","author":"J Quinn","year":"2009","unstructured":"Quinn J, Tam S, Sims N, Saleh H, McGregor N, Poulton I, Walker E, Scott J, Kemp B, Gillespie M (2009) Mice lacking AMP-activated kinase (AMPK) subunits \u03b21 or \u03b22 have low bone mass, while AICAR acts AMPK-independently to increase osteoclast formation. Bone 44:S136","journal-title":"Bone"},{"key":"9435_CR25","doi-asserted-by":"crossref","first-page":"808","DOI":"10.1210\/en.136.2.808","volume":"136","author":"E Jimi","year":"1995","unstructured":"Jimi E, Shuto T, Koga T (1995) Macrophage colony-stimulating factor and interleukin-1alpha maintain the survival of osteoclast-like cells. Endocrinology 136:808\u2013811","journal-title":"Endocrinology"},{"key":"9435_CR26","doi-asserted-by":"crossref","first-page":"1504","DOI":"10.1126\/science.289.5484.1504","volume":"289","author":"SL Teitelbaum","year":"2000","unstructured":"Teitelbaum SL (2000) Bone resorption by osteoclasts. Science 289:1504\u20131508","journal-title":"Science"},{"key":"9435_CR27","doi-asserted-by":"crossref","first-page":"49343","DOI":"10.1074\/jbc.M103642200","volume":"276","author":"SE Lee","year":"2001","unstructured":"Lee SE, Chung WJ, Kwak HB, Chung CH, Kwack KB, Lee ZH, Kim HH (2001) Tumor necrosis factor-alpha supports the survival of osteoclasts through the activation of Akt and ERK. J Biol Chem 276:49343\u201349349","journal-title":"J Biol Chem"},{"key":"9435_CR28","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1083\/jcb.148.2.333","volume":"148","author":"T Miyazaki","year":"2000","unstructured":"Miyazaki T, Katagiri H, Kanegae Y, Takayanagi H, Sawada Y, Yamamoto A, Pando MP, Asano T, Verma IM, Oda H, Nakamura K, Tanaka S (2000) Reciprocal role of ERK and NF-kappaB pathways in survival and activation of osteoclasts. J Cell Biol 148:333\u2013342","journal-title":"J Cell Biol"},{"key":"9435_CR29","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1016\/j.bbrc.2004.05.020","volume":"319","author":"K Sakamoto","year":"2004","unstructured":"Sakamoto K, Arnolds DE, Ekberg I, Thorell A, Goodyear LJ (2004) Exercise regulates Akt and glycogen synthase kinase-3 activities in human skeletal muscle. Biochem Biophys Res Commun 319:419\u2013425","journal-title":"Biochem Biophys Res Commun"},{"key":"9435_CR30","doi-asserted-by":"crossref","first-page":"431","DOI":"10.1530\/eje.0.1440431","volume":"144","author":"S Palacio","year":"2001","unstructured":"Palacio S, Felix R (2001) The role of phosphoinositide 3-kinase in spreading osteoclasts induced by colony-stimulating factor-1. Eur J Endocrinol 144:431\u2013440","journal-title":"Eur J Endocrinol"},{"key":"9435_CR31","doi-asserted-by":"crossref","first-page":"3427","DOI":"10.1038\/sj.onc.1202595","volume":"17","author":"K Sugihara","year":"1998","unstructured":"Sugihara K, Nakatsuji N, Nakamura K, Nakao K, Hashimoto R, Otani H, Sakagami H, Kondo H, Nozawa S, Aiba A, Katsuki M (1998) Rac1 is required for the formation of three germ layers during gastrulation. Oncogene 17:3427\u20133433","journal-title":"Oncogene"},{"key":"9435_CR32","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1126\/science.1088485","volume":"302","author":"Y Gu","year":"2003","unstructured":"Gu Y, Filippi MD, Cancelas JA, Siefring JE, Williams EP, Jasti AC, Harris CE, Lee AW, Prabhakar R, Atkinson SJ, Kwiatkowski DJ, Williams DA (2003) Hematopoietic cell regulation by Rac1 and Rac2 guanosine triphosphatases. Science 302:445\u2013449","journal-title":"Science"},{"key":"9435_CR33","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1126\/science.1089709","volume":"302","author":"MJ Walmsley","year":"2003","unstructured":"Walmsley MJ, Ooi SK, Reynolds LF, Smith SH, Ruf S, Mathiot A, Vanes L, Williams DA, Cancro MP, Tybulewicz VL (2003) Critical roles for Rac1 and Rac2 GTPases in B cell development and signaling. Science 302:459\u2013462","journal-title":"Science"},{"key":"9435_CR34","doi-asserted-by":"crossref","first-page":"5043","DOI":"10.4049\/jimmunol.169.9.5043","volume":"169","author":"S Li","year":"2002","unstructured":"Li S, Yamauchi A, Marchal CC, Molitoris JK, Quilliam LA, Dinauer MC (2002) Chemoattractant-stimulated Rac activation in wild-type and Rac2-deficient murine neutrophils: preferential activation of Rac2 and Rac2 gene dosage effect on neutrophil functions. J Immunol 169:5043\u20135051","journal-title":"J Immunol"},{"key":"9435_CR35","doi-asserted-by":"crossref","first-page":"5763","DOI":"10.1128\/MCB.25.13.5763-5776.2005","volume":"25","author":"S Corbetta","year":"2005","unstructured":"Corbetta S, Gualdoni S, Albertinazzi C, Paris S, Croci L, Consalez GG, de Curtis I (2005) Generation and characterization of Rac3 knockout mice. Mol Cell Biol 25:5763\u20135776","journal-title":"Mol Cell Biol"},{"key":"9435_CR36","doi-asserted-by":"crossref","first-page":"793","DOI":"10.1083\/jcb.200503152","volume":"170","author":"R Pankov","year":"2005","unstructured":"Pankov R, Endo Y, Even-Ram S, Araki M, Clark K, Cukierman E, Matsumoto K, Yamada KM (2005) A Rac switch regulates random versus directionally persistent cell migration. J Cell Biol 170:793\u2013802","journal-title":"J Cell Biol"},{"key":"9435_CR37","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1083\/jcb.152.2.361","volume":"152","author":"I Nakamura","year":"2001","unstructured":"Nakamura I, Lipfert L, Rodan GA, Le TD (2001) Convergence of \u03b1v\u03b23 integrin- and macrophage colony stimulating factor-mediated signals on phospholipase C\u03b3 in prefusion osteoclasts. J Cell Biol 152:361\u2013373","journal-title":"J Cell Biol"},{"key":"9435_CR38","doi-asserted-by":"crossref","first-page":"1259","DOI":"10.1242\/jcs.00997","volume":"117","author":"CM Wells","year":"2004","unstructured":"Wells CM, Walmsley M, Ooi S, Tybulewicz V, Ridley AJ (2004) Rac1-deficient macrophages exhibit defects in cell spreading and membrane ruffling but not migration. J Cell Sci 117:1259\u20131268","journal-title":"J Cell Sci"},{"key":"9435_CR39","doi-asserted-by":"crossref","first-page":"2749","DOI":"10.1242\/jcs.03024","volume":"119","author":"AP Wheeler","year":"2006","unstructured":"Wheeler AP, Wells CM, Smith SD, Vega FM, Henderson RB, Tybulewicz VL, Ridley AJ (2006) Rac1 and Rac2 regulate macrophage morphology but are not essential for migration. J Cell Sci 119:2749\u20132757","journal-title":"J Cell Sci"},{"key":"9435_CR40","doi-asserted-by":"crossref","first-page":"41661","DOI":"10.1074\/jbc.M306491200","volume":"278","author":"D Pradip","year":"2003","unstructured":"Pradip D, Peng X, Durden DL (2003) Rac2 specificity in macrophage integrin signaling: potential role for Syk kinase. J Biol Chem 278:41661\u201341669","journal-title":"J Biol Chem"},{"key":"9435_CR41","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1016\/S0171-9335(99)80058-2","volume":"78","author":"S Razzouk","year":"1999","unstructured":"Razzouk S, Lieberherr M, Cournot G (1999) Rac-GTPase, osteoclast cytoskeleton and bone resorption. Eur J Cell Biol 78:249\u2013255","journal-title":"Eur J Cell Biol"},{"key":"9435_CR42","doi-asserted-by":"crossref","first-page":"1622","DOI":"10.1172\/JCI116032","volume":"90","author":"BF Boyce","year":"1992","unstructured":"Boyce BF, Yoneda T, Lowe C, Soriano P, Mundy GR (1992) Requirement of pp60c-src expression for osteoclasts to form ruffled borders and resorb bone in mice. J Clin Invest 90:1622\u20131627","journal-title":"J Clin Invest"},{"key":"9435_CR43","doi-asserted-by":"crossref","first-page":"4009","DOI":"10.1210\/en.2008-0034","volume":"149","author":"T Kawano","year":"2008","unstructured":"Kawano T, Troiano N, Adams DJ, Wu JJ, Sun BH, Insogna K (2008) The anabolic response to parathyroid hormone is augmented in Rac2 knockout mice. Endocrinology 149:4009\u20134015","journal-title":"Endocrinology"}],"container-title":["Calcified Tissue International"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/link.springer.com\/content\/pdf\/10.1007\/s00223-010-9435-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/link.springer.com\/article\/10.1007\/s00223-010-9435-3\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/link.springer.com\/content\/pdf\/10.1007\/s00223-010-9435-3","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,11,14]],"date-time":"2021-11-14T03:19:39Z","timestamp":1636859979000},"score":1,"resource":{"primary":{"URL":"http:\/\/link.springer.com\/10.1007\/s00223-010-9435-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2010,11,26]]},"references-count":43,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2011,1]]}},"alternative-id":["9435"],"URL":"https:\/\/doi.org\/10.1007\/s00223-010-9435-3","relation":{},"ISSN":["0171-967X","1432-0827"],"issn-type":[{"value":"0171-967X","type":"print"},{"value":"1432-0827","type":"electronic"}],"subject":[],"published":{"date-parts":[[2010,11,26]]}}}