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We thank Koichi Masuda (UCSD) for the kind cooperation in microCT analysis. This research is supported by AMED-CREST from AMED (JP15gm0410001, JP17gm0810008), Daiichi Sankyo Foundation of Life Science, JSPS KAKENHI [Grant Number: 26113008, 15H02560, 15K15544], and grants from the National Institutes of Health [Grant Number: AR050631, AR065379] to H.A., the Grant from the National Center for Child Health and Development (Grant Number 25-1), the Uehara memorial foundation, the Nakajima Foundation and JSPS KAKENHI (16K18558) for M.I.

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Author notes

  1. These authors contributed equally: Masafumi Inui and Sho Mokuda.


  1. Department of Systems BioMedicine, National Research Institute for Child Health and Development, Tokyo, Japan

    • Masafumi Inui
    • , Tempei Sato
    • , Moe Tamano
    • , Shuji Takada
    •  & Hiroshi Asahara
  2. Laboratory of Animal Regeneration Systemology, Department of Life Sciences, School of Agriculture, Meiji University, Kanagawa, Japan

    • Masafumi Inui
  3. Department of Molecular Medicine, The Scripps Research Institute, La Jolla, CA, USA

    • Sho Mokuda
    •  & Hiroshi Asahara
  4. Department of Systems BioMedicine, Graduate School of Medical and Dental Sciences, Tokyo Medical and Dental University, Tokyo, Japan

    • Tempei Sato
    •  & Hiroshi Asahara
  5. AMED-CREST, Japan Agency for Medical Research and Development (AMED), Tokyo, Japan

    • Hiroshi Asahara


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M.I., S.M. and T.S. analysed the KO mice, M.T. performed the microinjections. M.I., S.T. and H.A. planned the experiments and wrote the manuscript.

Competing interests

The authors declare no competing interests.

Corresponding authors

Correspondence to Masafumi Inui or Hiroshi Asahara.

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