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Acknowledgements

We thank Dr. Yamamoto H, Ms. Kosaka K and Ms. Morinaga R for technical assistance. This work was supported by the JSPS KAKENHI Grant Numbers JP17K17814 (H.I.), JP25461342 (K.M.), JP15K08229 (T.I.) and

JP19K09020 (K.M.) and by Research Funding Grant by the president of Shiga University of Medical Science. We

would like to thank Editage (www.edita​ge.com) for English language editing.

Author contributions

H.I. designed the study, performed experiments, analysed data, and wrote the manuscript. Y.I., M.L. and L.S.

performed experiments. K.M. and T.I. designed the study and wrote the manuscript. T.S., J.M., H.S., E.N. and

H.M. made suggestions for the study design and experiments. All authors commented on the manuscript.

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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/s4159​8-020-80742​-y.

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