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Acknowledgements
We thank Drs. S. Tamaki, and H. Yoshitomi for invaluable comments and discussion, Dr. K. Woltjen for providing GFP-labelled iPS cell line, and Dr. P. Karagiannis for proofreading the manuscript. Preparation of the tissue
slides and staining was supported by the Applied Medical Research (Osaka, Japan). This study was supported
by iPS Academia Japan, Inc., to S.K., by a grant-in-aid for the Acceleration Program for Intractable Disease
Research Utilizing Disease Specific iPS Cells (Japan Agency if) to T.A, S.M, and J.T., the Core Center for iPS Cell
Research (AMED) to S.K. and J.T., the Centers for Clinical Application Research on Specific Disease/Organ (type
B) grants (AMED) to J.T., and the iPS Cell Research Fund to J.T. These funders had no role in the study design,
data collection and analysis, decision to publish, or preparation of the manuscript.
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Author contributions
S.K. and J.T. designed the research and wrote the manuscript. J.S. performed the imaging studies and made
experimental materials. S.N. and M.N. performed the in vitro and in vivo experiments. T.K. and L.S. assisted with
the in vitro experiments. S.S. and A.W. performed the scRNA sequencing experiments. M.F. and Y.J. analyzed the
scRNA sequencing data. S.M. and T.A. advised on the project. All authors provided feedback on the manuscript.
Competing interests The authors declare no competing interests.
Additional information
Supplementary Information The online version contains supplementary material available at https://doi.org/
10.1038/s41598-023-27556-w.
Correspondence and requests for materials should be addressed to S.K. or J.T.
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