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Acknowledgements
The authors would like to thank Dr. Keisuke Okita for providing the pCXLE-gw episomal expression vectors,
Dr. Rina Ogura for supporting western blot experiments, and Dr. Peter Karagiannis for critically reading of the
manuscript. This work was partially supported by the Japan Society for the Promotion of Science (JSPS) through
its Grant-in-Aid for Scientific Research (B) (JSPS KAKENHI Grant Number 21H02979) to K.O. and by the Japan
Agency for Medical Research and Development (AMED) through its research grant “Core Center for iPS Cell
Research (JP22bm0104001), R&D Program of Regenerative/Cellular Medicine and Gene Therapies from the Basic
to Nonclinical Phase (JP22bm1123002), Research Center Network for Realization of Regenerative Medicine” to
K.O. and by the iPS Cell Research Fund.
Author contributions
R.I. performed all the experiments. R.I. and K.O. drafted the manuscript. R.I., A.K., T.T. and K.O. contributed to
the study conception and design. Y. Hirose contributed to the establishment of the screening system. Y. Hatano
provided consultation on the statistical analysis methods. A.K., A.M., Y.K., T.N., and J.F. contributed to the
interpretation and planning of each experiment. S.-I.M. provided consultation on the RNA-seq analysis. Y.N.
and A.O. supervised the screening system. T.T., N.I. and K.O. supervised all aspects of the study. All authors
were involved in the manuscript preparation.
Competing interests K.O. is a founder and member of the scientific advisory boards of iPS Portal, Inc., and a founder and chief
scientific advisor of RegeNephro Co., Ltd. The other authors have no competing interest.
Additional information
Supplementary Information The online version contains supplementary material available at https://doi.org/
10.1038/s41598-023-35875-1.
Correspondence and requests for materials should be addressed to T.T. or K.O.
Reprints and permissions information is available at www.nature.com/reprints.
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