Dion, William https://orcid.org/0000-0002-7697-9433
Tao, Yuren
Chambers, Maci
Zhao, Shanshan
Arbuckle, Riley K.
Sun, Michelle
Kubra, Syeda
Schaich, Matthew A.
Nie, Yuhang
Ye, Megan
Jamal, Imran
Larsen, Mads B. https://orcid.org/0000-0001-5361-0898
Camarco, Daniel
Ickes, Eleanor https://orcid.org/0009-0005-4561-7228
Wang, Haokun H. https://orcid.org/0009-0003-6279-8003
DuPont, C.
Wang, Bingjie https://orcid.org/0000-0001-5056-7476
Liu, Silvia
Pi, Shaohua https://orcid.org/0000-0002-7549-8929
Van Houten, Bennett
Chen, Bill B.
Chen, Yuanyuan
Chen, Xu https://orcid.org/0000-0001-8926-3090
Zhu, Bokai https://orcid.org/0000-0003-0827-5757
Funding for this research was provided by:
U.S. Department of Health & Human Services | NIH | National Institute on Aging (1R21AG071893, F31 AG080998, R01AG074273, R01AG078185)
Richard King Mellon Foundation
Foundation Fighting Blindness (Translational Research Program Award)
U.S. Department of Health & Human Services | NIH | National Institute of General Medical Sciences (1DP2GM140924)
AFAR/Hevolution foundation
U.S. Department of Health & Human Services | NIH | National Heart, Lung, and Blood Institute (T32 HL082610, 1R35HL139860)
the Diana Jacobs Kalman/AFAR Scholarship for Research in the Biology of Aging through the American Federation for Aging Research
U.S. Department of Health & Human Services | NIH | National Institute of Environmental Health Sciences (F32ES034982, R35ES031638)
U.S. Department of Health & Human Services | NIH | National Institute of Diabetes and Digestive and Kidney Diseases (P30DK120531)
U.S. Department of Health & Human Services | NIH | NIH Office of the Director (S10OD028483, S10OD032158)
U.S. Department of Health & Human Services | NIH | National Eye Institute (R01 EY030991, P30 EY08098)
Article History
Received: 3 April 2024
Accepted: 11 July 2025
First Online: 5 August 2025
Competing interests
: B.B.C. is Co-founder of Koutif Therapeutic Inc., Co-founder and VP of Drug Discovery for Generian Pharmaceuticals and Co-founder and C.S.O. of Coloma Therapeutics Inc. The other authors declare no competing interests.