1. Department of Molecular and Cellular Biology, Institute for Frontier Medical Sciences, Kyoto University
2. Department of Health and Environmental Sciences, Graduate School of Medicine and Faculty of Medicine, Kyoto University
3. These two authors equally contributed to this work.
4. CREST, Japan Science Technology Agency
5. Department of Pathology and Biology of Diseases, Kyoto University Graduate School of Medicine
6. Present address: Department of Public Health, Hyogo College of Medicine