{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T01:54:25Z","timestamp":1792720465892,"version":"4.5.0"},"reference-count":47,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2012,10,2]],"date-time":"2012-10-02T00:00:00Z","timestamp":1349136000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2012,10,2]],"date-time":"2012-10-02T00:00:00Z","timestamp":1349136000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Commun"],"DOI":"10.1038\/ncomms2105","type":"journal-article","created":{"date-parts":[[2012,10,1]],"date-time":"2012-10-01T16:23:25Z","timestamp":1349108605000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":117,"title":["Mst1 regulates integrin-dependent thymocyte trafficking and antigen recognition in the thymus"],"prefix":"10.1038","volume":"3","author":[{"given":"Yoshihiro","family":"Ueda","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Koko","family":"Katagiri","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Takashi","family":"Tomiyama","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kaneki","family":"Yasuda","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Katsuyoshi","family":"Habiro","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tomoya","family":"Katakai","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Susumu","family":"Ikehara","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mitsuru","family":"Matsumoto","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tatsuo","family":"Kinashi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2012,10,2]]},"reference":[{"key":"BFncomms2105_CR1","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1146\/annurev.immunol.21.120601.141107","volume":"21","author":"TK Starr","year":"2003","unstructured":"Starr, T. K., Jameson, S. C. & Hogquist, K. A. Positive and negative selection of T cells. Annu. Rev. Immunol. 21, 139\u2013176 (2003).","journal-title":"Annu. Rev. Immunol."},{"key":"BFncomms2105_CR2","doi-asserted-by":"publisher","first-page":"469","DOI":"10.1038\/nri2989","volume":"11","author":"PE Love","year":"2011","unstructured":"Love, P. E. & Bhandoola, A. Signal integration and crosstalk during thymocyte migration and emigration. Nat. Rev. Immunol. 11, 469\u2013477 (2011).","journal-title":"Nat. Rev. Immunol."},{"key":"BFncomms2105_CR3","doi-asserted-by":"publisher","first-page":"1032","DOI":"10.1038\/ni723","volume":"2","author":"J Derbinski","year":"2001","unstructured":"Derbinski, J., Schulte, A., Kyewski, B. & Klein, L. Promiscuous gene expression in medullary thymic epithelial cells mirrors the peripheral self. Nat. Immunol. 2, 1032\u20131039 (2001).","journal-title":"Nat. Immunol."},{"key":"BFncomms2105_CR4","doi-asserted-by":"publisher","first-page":"1395","DOI":"10.1126\/science.1075958","volume":"298","author":"MS Anderson","year":"2002","unstructured":"Anderson, M. S. et al. Projection of an immunological self shadow within the thymus by the aire protein. Science 298, 1395\u20131401 (2002).","journal-title":"Science"},{"key":"BFncomms2105_CR5","doi-asserted-by":"publisher","first-page":"350","DOI":"10.1038\/ni906","volume":"4","author":"A Liston","year":"2003","unstructured":"Liston, A., Lesage, S., Wilson, J., Peltonen, L. & Goodnow, C. C. Aire regulates negative selection of organ-specific T cells. Nat. Immunol. 4, 350\u2013354 (2003).","journal-title":"Nat. Immunol."},{"key":"BFncomms2105_CR6","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1016\/j.immuni.2005.07.005","volume":"23","author":"MS Anderson","year":"2005","unstructured":"Anderson, M. S. et al. The cellular mechanism of Aire control of T cell tolerance. Immunity 23, 227\u2013239 (2005).","journal-title":"Immunity"},{"key":"BFncomms2105_CR7","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1146\/annurev.immunol.25.022106.141532","volume":"27","author":"D Mathis","year":"2009","unstructured":"Mathis, D. & Benoist, C. Aire. Annu. Rev. Immunol. 27, 287\u2013312 (2009).","journal-title":"Annu. Rev. Immunol."},{"key":"BFncomms2105_CR8","doi-asserted-by":"publisher","first-page":"e160","DOI":"10.1371\/journal.pbio.0030160","volume":"3","author":"CM Witt","year":"2005","unstructured":"Witt, C. M., Raychaudhuri, S., Schaefer, B., Chakraborty, A. K. & Robey, E. A. Directed migration of positively selected thymocytes visualized in real time. PLoS Biol. 3, e160 (2005).","journal-title":"PLoS Biol."},{"key":"BFncomms2105_CR9","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1038\/ni1161","volume":"6","author":"NR Bhakta","year":"2005","unstructured":"Bhakta, N. R., Oh, D. Y. & Lewis, R. S. Calcium oscillations regulate thymocyte motility during positive selection in the three-dimensional thymic environment. Nat. Immunol. 6, 143\u2013151 (2005).","journal-title":"Nat. Immunol."},{"key":"BFncomms2105_CR10","doi-asserted-by":"publisher","first-page":"823","DOI":"10.1038\/ni.1761","volume":"10","author":"M Le Borgne","year":"2009","unstructured":"Le Borgne, M. et al. The impact of negative selection on thymocyte migration in the medulla. Nat. Immunol. 10, 823\u2013830 (2009).","journal-title":"Nat. Immunol."},{"key":"BFncomms2105_CR11","doi-asserted-by":"publisher","first-page":"986","DOI":"10.1016\/j.immuni.2009.09.020","volume":"31","author":"LI Ehrlich","year":"2009","unstructured":"Ehrlich, L. I., Oh, D. Y., Weissman, I. L. & Lewis, R. S. Differential contribution of chemotaxis and substrate restriction to segregation of immature and mature thymocytes. Immunity 31, 986\u2013998 (2009).","journal-title":"Immunity"},{"key":"BFncomms2105_CR12","doi-asserted-by":"publisher","first-page":"503","DOI":"10.1038\/ni.1868","volume":"11","author":"H Phee","year":"2010","unstructured":"Phee, H. et al. Regulation of thymocyte positive selection and motility by GIT2. Nat. Immunol. 11, 503\u2013511 (2010).","journal-title":"Nat. Immunol."},{"key":"BFncomms2105_CR13","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/j.immuni.2005.12.011","volume":"24","author":"H Kurobe","year":"2006","unstructured":"Kurobe, H. et al. CCR7-dependent cortex-to-medulla migration of positively selected thymocytes is essential for establishing central tolerance. Immunity 24, 165\u2013177 (2006).","journal-title":"Immunity"},{"key":"BFncomms2105_CR14","doi-asserted-by":"publisher","first-page":"613","DOI":"10.1002\/eji.200636656","volume":"37","author":"AC Davalos-Misslitz","year":"2007","unstructured":"Davalos-Misslitz, A. C. et al. Generalized multi-organ autoimmunity in CCR7-deficient mice. Eur. J. Immunol. 37, 613\u2013622 (2007).","journal-title":"Eur. J. Immunol."},{"key":"BFncomms2105_CR15","doi-asserted-by":"publisher","first-page":"4351","DOI":"10.1182\/blood-2007-01-070284","volume":"110","author":"AC Davalos-Misslitz","year":"2007","unstructured":"Davalos-Misslitz, A. C., Worbs, T., Willenzon, S., Bernhardt, G. & Forster, R. Impaired responsiveness to T-cell receptor stimulation and defective negative selection of thymocytes in CCR7-deficient mice. Blood 110, 4351\u20134359 (2007).","journal-title":"Blood"},{"key":"BFncomms2105_CR16","doi-asserted-by":"publisher","first-page":"17129","DOI":"10.1073\/pnas.0906956106","volume":"106","author":"T Nitta","year":"2009","unstructured":"Nitta, T., Nitta, S., Lei, Y., Lipp, M. & Takahama, Y. CCR7-mediated migration of developing thymocytes to the medulla is essential for negative selection to tissue-restricted antigens. Proc. Natl Acad. Sci. USA 106, 17129\u201317133 (2009).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"BFncomms2105_CR17","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1038\/nrc2070","volume":"7","author":"K Harvey","year":"2007","unstructured":"Harvey, K. & Tapon, N. The Salvador-Warts-Hippo pathway\u2014an emerging tumour-suppressor network. Nat. Rev. Cancer 7, 182\u2013191 (2007).","journal-title":"Nat. Rev. Cancer"},{"key":"BFncomms2105_CR18","doi-asserted-by":"publisher","first-page":"613","DOI":"10.1038\/nrm2221","volume":"8","author":"LJ Saucedo","year":"2007","unstructured":"Saucedo, L. J. & Edgar, B. A. Filling out the Hippo pathway. Nat. Rev. Mol. Cell. Biol. 8, 613\u2013621 (2007).","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"BFncomms2105_CR19","doi-asserted-by":"publisher","first-page":"919","DOI":"10.1038\/ni1374","volume":"7","author":"K Katagiri","year":"2006","unstructured":"Katagiri, K., Imamura, M. & Kinashi, T. Spatiotemporal regulation of the kinase Mst1 by binding protein RAPL is critical for lymphocyte polarity and adhesion. Nat. Immunol. 7, 919\u2013928 (2006).","journal-title":"Nat. Immunol."},{"key":"BFncomms2105_CR20","doi-asserted-by":"publisher","first-page":"1319","DOI":"10.1038\/emboj.2009.82","volume":"28","author":"K Katagiri","year":"2009","unstructured":"Katagiri, K. et al. Mst1 controls lymphocyte trafficking and interstitial motility within lymph nodes. EMBO J. 28, 1319\u20131331 (2009).","journal-title":"EMBO J."},{"key":"BFncomms2105_CR21","doi-asserted-by":"publisher","first-page":"3865","DOI":"10.4049\/jimmunol.0900678","volume":"183","author":"Y Dong","year":"2009","unstructured":"Dong, Y. et al. A cell-intrinsic role for Mst1 in regulating thymocyte egress. J. Immunol. 183, 3865\u20133872 (2009).","journal-title":"J. Immunol."},{"key":"BFncomms2105_CR22","doi-asserted-by":"publisher","first-page":"e8011","DOI":"10.1371\/journal.pone.0008011","volume":"4","author":"J Choi","year":"2009","unstructured":"Choi, J. et al. Mst1-FoxO signaling protects Naive T lymphocytes from cellular oxidative stress in mice. PLoS One 4, e8011 (2009).","journal-title":"PLoS One"},{"key":"BFncomms2105_CR23","doi-asserted-by":"publisher","first-page":"741","DOI":"10.1084\/jem.20111692","volume":"209","author":"F Mou","year":"2012","unstructured":"Mou, F. et al. The Mst1 and Mst2 kinases control activation of rho family GTPases and thymic egress of mature thymocytes. J. Exp. Med. 209, 741\u2013759 (2012).","journal-title":"J. Exp. Med."},{"key":"BFncomms2105_CR24","doi-asserted-by":"publisher","first-page":"20321","DOI":"10.1073\/pnas.0810773105","volume":"105","author":"D Zhou","year":"2008","unstructured":"Zhou, D. et al. The Nore1B\/Mst1 complex restrains antigen receptor-induced proliferation of naive T cells. Proc. Natl. Acad. Sci. USA 105, 20321\u201320326 (2008).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"BFncomms2105_CR25","doi-asserted-by":"publisher","first-page":"3458","DOI":"10.1182\/blood-2011-09-378364","volume":"119","author":"NT Nehme","year":"2012","unstructured":"Nehme, N. T. et al. MST1 mutations in autosomal recessive primary immunodeficiency characterized by defective naive T cells survival. Blood 119, 3458\u20133468 (2012).","journal-title":"Blood"},{"key":"BFncomms2105_CR26","doi-asserted-by":"publisher","first-page":"3450","DOI":"10.1182\/blood-2011-09-378158","volume":"119","author":"H Abdollahpour","year":"2012","unstructured":"Abdollahpour, H. et al. The phenotype of human STK4 deficiency. Blood 119, 3450\u20133457 (2012).","journal-title":"Blood"},{"key":"BFncomms2105_CR27","doi-asserted-by":"publisher","first-page":"338","DOI":"10.1038\/ni1323","volume":"7","author":"E Ladi","year":"2006","unstructured":"Ladi, E., Yin, X., Chtanova, T. & Robey, E. A. Thymic microenvironments for T cell differentiation and selection. Nat. Immunol. 7, 338\u2013343 (2006).","journal-title":"Nat. Immunol."},{"key":"BFncomms2105_CR28","doi-asserted-by":"publisher","first-page":"1125","DOI":"10.1126\/science.1347662","volume":"255","author":"M Isobe","year":"1992","unstructured":"Isobe, M., Yagita, H., Okumura, K. & Ihara, A. Specific acceptance of cardiac allograft after treatment with antibodies to ICAM-1 and LFA-1. Science 255, 1125\u20131127 (1992).","journal-title":"Science"},{"key":"BFncomms2105_CR29","doi-asserted-by":"publisher","first-page":"1039","DOI":"10.1084\/jem.20041457","volume":"200","author":"AM Gallegos","year":"2004","unstructured":"Gallegos, A. M. & Bevan, M. J. Central tolerance to tissue-specific antigens mediated by direct and indirect antigen presentation. J. Exp. Med. 200, 1039\u20131049 (2004).","journal-title":"J. Exp. Med."},{"key":"BFncomms2105_CR30","doi-asserted-by":"publisher","first-page":"2462","DOI":"10.1182\/blood-2010-06-286393","volume":"118","author":"FX Hubert","year":"2011","unstructured":"Hubert, F. X. et al. Aire regulates the transfer of antigen from mTECs to dendritic cells for induction of thymic tolerance. Blood 118, 2462\u20132472 (2011).","journal-title":"Blood"},{"key":"BFncomms2105_CR31","doi-asserted-by":"publisher","first-page":"742","DOI":"10.1038\/333742a0","volume":"333","author":"P Kisielow","year":"1988","unstructured":"Kisielow, P., Bluthmann, H., Staerz, U. D., Steinmetz, M. & von Boehmer, H. Tolerance in T-cell-receptor transgenic mice involves deletion of nonmature CD4+8+ thymocytes. Nature 333, 742\u2013746 (1988).","journal-title":"Nature"},{"key":"BFncomms2105_CR32","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1038\/nri1032","volume":"3","author":"JA Bluestone","year":"2003","unstructured":"Bluestone, J. A. & Abbas, A. K. Natural versus adaptive regulatory T cells. Nat. Rev. Immunol. 3, 253\u2013257 (2003).","journal-title":"Nat. Rev. Immunol."},{"key":"BFncomms2105_CR33","doi-asserted-by":"publisher","first-page":"775","DOI":"10.1016\/j.cell.2008.05.009","volume":"133","author":"S Sakaguchi","year":"2008","unstructured":"Sakaguchi, S., Yamaguchi, T., Nomura, T. & Ono, M. Regulatory T cells and immune tolerance. Cell 133, 775\u2013787 (2008).","journal-title":"Cell"},{"key":"BFncomms2105_CR34","doi-asserted-by":"publisher","first-page":"618","DOI":"10.1038\/ni.1884","volume":"11","author":"W Ouyang","year":"2010","unstructured":"Ouyang, W. et al. Foxo proteins cooperatively control the differentiation of Foxp3+ regulatory T cells. Nat. Immunol. 11, 618\u2013627 (2010).","journal-title":"Nat. Immunol."},{"key":"BFncomms2105_CR35","doi-asserted-by":"publisher","first-page":"2513","DOI":"10.1084\/jem.20070601","volume":"204","author":"TM McCaughtry","year":"2007","unstructured":"McCaughtry, T. M., Wilken, M. S. & Hogquist, K. A. Thymic emigration revisited. J. Exp. Med. 204, 2513\u20132520 (2007).","journal-title":"J. Exp. Med."},{"key":"BFncomms2105_CR36","doi-asserted-by":"publisher","first-page":"2832","DOI":"10.1002\/1521-4141(200010)30:10<2832::AID-IMMU2832>3.0.CO;2-D","volume":"30","author":"D Amsen","year":"2000","unstructured":"Amsen, D., Kruisbeek, A., Bos, J. L. & Reedquist, K. Activation of the Ras-related GTPase Rap1 by thymocyte TCR engagement and during selection. Eur. J. Immunol. 30, 2832\u20132841 (2000).","journal-title":"Eur. J. Immunol."},{"key":"BFncomms2105_CR37","doi-asserted-by":"publisher","first-page":"154","DOI":"10.1038\/nature02238","volume":"427","author":"TR Mempel","year":"2004","unstructured":"Mempel, T. R., Henrickson, S. E. & Von Andrian, U. H. T-cell priming by dendritic cells in lymph nodes occurs in three distinct phases. Nature 427, 154\u2013159 (2004).","journal-title":"Nature"},{"key":"BFncomms2105_CR38","doi-asserted-by":"publisher","first-page":"585","DOI":"10.1146\/annurev.immunol.24.021605.090620","volume":"26","author":"MD Cahalan","year":"2008","unstructured":"Cahalan, M. D. & Parker, I. Choreography of cell motility and interaction dynamics imaged by two-photon microscopy in lymphoid organs. Annu. Rev. Immunol. 26, 585\u2013626 (2008).","journal-title":"Annu. Rev. Immunol."},{"key":"BFncomms2105_CR39","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1038\/86302","volume":"2","author":"MS Jordan","year":"2001","unstructured":"Jordan, M. S. et al. Thymic selection of CD4+CD25+ regulatory T cells induced by an agonist self-peptide. Nat. Immunol. 2, 301\u2013306 (2001).","journal-title":"Nat. Immunol."},{"key":"BFncomms2105_CR40","doi-asserted-by":"publisher","first-page":"7889","DOI":"10.4049\/jimmunol.175.12.7889","volume":"175","author":"M Marski","year":"2005","unstructured":"Marski, M., Kandula, S., Turner, J. R. & Abraham, C. CD18 is required for optimal development and function of CD4+CD25+ T regulatory cells. J. Immunol. 175, 7889\u20137897 (2005).","journal-title":"J. Immunol."},{"key":"BFncomms2105_CR41","doi-asserted-by":"publisher","first-page":"512","DOI":"10.1038\/ni.1874","volume":"11","author":"M Hinterberger","year":"2010","unstructured":"Hinterberger, M. et al. Autonomous role of medullary thymic epithelial cells in central CD4(+) T cell tolerance. Nat. Immunol. 11, 512\u2013519 (2010).","journal-title":"Nat. Immunol."},{"key":"BFncomms2105_CR42","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1038\/nature02284","volume":"427","author":"M Matloubian","year":"2004","unstructured":"Matloubian, M. et al. Lymphocyte egress from thymus and peripheral lymphoid organs is dependent on S1P receptor 1. Nature 427, 355\u2013360 (2004).","journal-title":"Nature"},{"key":"BFncomms2105_CR43","doi-asserted-by":"publisher","first-page":"546","DOI":"10.1038\/nri1646","volume":"5","author":"T Kinashi","year":"2005","unstructured":"Kinashi, T. Intracellular signaling controlling integrin activation in lymphocytes. Nat. Rev. Immunol. 5, 546\u2013559 (2005).","journal-title":"Nat. Rev. Immunol."},{"key":"BFncomms2105_CR44","doi-asserted-by":"publisher","first-page":"2827","DOI":"10.1084\/jem.20080046","volume":"205","author":"M Yano","year":"2008","unstructured":"Yano, M. et al. Aire controls the differentiation program of thymic epithelial cells in the medulla for the establishment of self-tolerance. J. Exp. Med. 205, 2827\u20132838 (2008).","journal-title":"J. Exp. Med."},{"key":"BFncomms2105_CR45","doi-asserted-by":"publisher","first-page":"417","DOI":"10.1083\/jcb.200301133","volume":"161","author":"M Shimonaka","year":"2003","unstructured":"Shimonaka, M. et al. Rap1 translates chemokine signals to integrin activation, cell polarization, and motility across vascular endothelium under flow. J. Cell Biol. 161, 417\u2013427 (2003).","journal-title":"J. Cell Biol."},{"key":"BFncomms2105_CR46","doi-asserted-by":"publisher","first-page":"741","DOI":"10.1038\/ni950","volume":"4","author":"K Katagiri","year":"2003","unstructured":"Katagiri, K., Maeda, A., Shimonaka, M. & Kinashi, T. RAPL, a Rap1-binding molecule that mediates Rap1-induced adhesion through spatial regulation of LFA-1. Nat. Immunol. 4, 741\u2013748 (2003).","journal-title":"Nat. Immunol."},{"key":"BFncomms2105_CR47","doi-asserted-by":"publisher","first-page":"4359","DOI":"10.4049\/jimmunol.171.8.4359","volume":"171","author":"T Katakai","year":"2003","unstructured":"Katakai, T., Hara, T., Sugai, M., Gonda, H. & Shimizu, A. Th1-biased tertiary lymphoid tissue supported by CXC chemokine ligand 13-producing stromal network in chronic lesions of autoimmune gastritis. J. Immunol. 171, 4359\u20134368 (2003).","journal-title":"J. Immunol."}],"container-title":["Nature Communications"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/ncomms2105","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/ncomms2105.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/ncomms2105.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,1,6]],"date-time":"2023-01-06T00:22:50Z","timestamp":1672964570000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/ncomms2105"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2012,10,2]]},"references-count":47,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2012,1]]}},"alternative-id":["BFncomms2105"],"URL":"https:\/\/doi.org\/10.1038\/ncomms2105","relation":{},"ISSN":["2041-1723"],"issn-type":[{"value":"2041-1723","type":"electronic"}],"subject":[],"published":{"date-parts":[[2012,10,2]]},"assertion":[{"value":"12 March 2012","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"30 August 2012","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 October 2012","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing financial interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"1098"}}