About the Authors

Tokiyoshi Matsushita

Roles Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Project administration, Validation, Visualization, Writing – original draft, Writing – review & editing

tokiyoshi.matsushita@santen.com

Affiliations Faculty of Pharmaceutical Sciences, Department of Gene Regulation, Tokyo University of Science, Noda, Chiba, Japan, Product Discovery, Ophthalmology Innovation Center, Santen Pharmaceutical Co., Ltd., Ikoma, Nara, Japan

Akishi Onishi

Roles Conceptualization, Project administration, Validation, Writing – review & editing

Affiliations Laboratory for Retinal Regeneration, RIKEN Center for Biosystems Dynamics Research, Kobe, Hyogo, Japan, Cell and Gene Therapy in Ophthalmology Laboratory, Baton Zone Program, RIKEN, Wako, Saitama, Japan

Takahiro Matsuyama

Roles Data curation, Formal analysis, Investigation, Methodology, Validation

Affiliation Product Discovery, Ophthalmology Innovation Center, Santen Pharmaceutical Co., Ltd., Ikoma, Nara, Japan

Tomohiro Masuda

Roles Data curation, Formal analysis, Investigation, Methodology

Affiliations Laboratory for Retinal Regeneration, RIKEN Center for Biosystems Dynamics Research, Kobe, Hyogo, Japan, Cell and Gene Therapy in Ophthalmology Laboratory, Baton Zone Program, RIKEN, Wako, Saitama, Japan

Yoko Ogino

Roles Project administration

Affiliation Faculty of Pharmaceutical Sciences, Department of Gene Regulation, Tokyo University of Science, Noda, Chiba, Japan

Masaaki Kageyama

Roles Supervision, Writing – review & editing

Affiliation Product Discovery, Ophthalmology Innovation Center, Santen Pharmaceutical Co., Ltd., Ikoma, Nara, Japan

Masayo Takahashi

Roles Supervision

Affiliations Laboratory for Retinal Regeneration, RIKEN Center for Biosystems Dynamics Research, Kobe, Hyogo, Japan, Cell and Gene Therapy in Ophthalmology Laboratory, Baton Zone Program, RIKEN, Wako, Saitama, Japan

Fumiaki Uchiumi

Roles Supervision, Writing – review & editing

Affiliation Faculty of Pharmaceutical Sciences, Department of Gene Regulation, Tokyo University of Science, Noda, Chiba, Japan

Competing Interests

The study was funded by Santen Pharmaceutical Co. Ltd and Tokiyoshi Matsushita, Takahiro Matsuyama, and Masaaki Kageyama are employed by Santen Pharmaceutical Co., Ltd. Tokiyoshi Matsushita, Akishi Onishi, and Masayo Takahashi are co-inventors of a patent related to this work, filed by RIKEN and Santen Pharmaceutical Co., Ltd [WO2022138803A1]. None of these alter our adherence to PLOS ONE policies on sharing data and materials.