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Acknowledgements
We thank Dr. Jack Kaplan (University of Illinois College of Medicine), Dr. Hirohide Saito (Kyoto University),
and Dr. Glen K. Andrews (University of Kansas Medical Center) for the gifts of MDCK FLp-In T-Rex cells,
IRES-GFP plasmid, and Zip4 and Zip5 cDNA, respectively, and Dr. Tokuji Tsuji, Yukari Matsuo, and Sayaka
Arimoto for technical assistance.
Author contributions
T.K. designed the study; S.N., Y.N., H.Y., N.Y., H.Y., and T.K. collected, analyzed, and interpreted the data; T.K.
and T.S. provided technical assistance; T.K. drafted the manuscript. All authors reviewed the manuscript.
Funding
This work was supported by a Grant-in-Aid for Scientific Research on Innovative Areas “Integrated Bio-metal
Science” (MEXT KAKENHI Grant Number JP19H05768) from the Ministry of Education, Culture, Sports,
Science, and Technology, Japan, a Grant-in-Aid for Scientific Research (B) (JSPS KAKENHI Grant Number
JP19H02883) from the Japan Society for the Promotion of Science, the Kyoto Innovative Medical Technology
Research and Development Support System, the Mitsubishi Foundation, the Kieikai Research Foundation, and
the Tojuro Iijima Foundation for Food Science and Technology (to T. K.).
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/s41598-022-10925-2.
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