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This work was financially supported by grants from the Creative Research Initiative (2010-0018286),WCU Program(R32-10217),National Research Foundation of Korea (2011-0017611)and the second stage BK21for Chemistry and Medical Sciences of Yonsei University.S.W.P.was supported by the National Research Foundation,

Mid-career Researcher Program(72011-0043).M.H.C.was supported by a Hi Seoul Science/Humanities Fellowship from the Seoul Scholarship Foundation.

Author contributions

J.C.and J-S.S.conceived and designed the experiments.M.H.C.,E.J.L.,M.S.and J-w.K. performed the experiments.S.W.P.provided advice on the in vivo zebrafish experiments. M.H.C.,E.J.L.,J-H.L.,D.Y.,J-S.S.and J.C.wrote the manuscript.

Additional information

Supplementary information is available in the online version of the paper.Reprints and permissions information is available online at /reprints. Correspondence and requests for materials should be addressed to J-S.S.or J.C. Competing?nancial interests

The authors declare no competing financial interests.

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