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This work was supported by the National Natural Science Foundation of China (Grant No. 31970948 and No. 31600859 to W.G.; No. 81871035 and No.82071378 to P.L.; No. 82101556 to X.Z.), the Research Fund for International Senior Scientists (Grant No. 82150710558 to J.C.), Zhejiang Provincial Natural Science Foundation (Grant No. LZ19H090001 to P.L.; No.LQ22H090013 to X.Z.), the Project of State Key Laboratory of Ophthalmology, Optometry and Vision Science, Wenzhou Medical University (J01-20190101 to J.C.), La Caixa Foundation (LCF/PR/HP17/52190001 to R.C.), Centro 2020 (CENTRO-01-0246-FEDER-000010 to R.C.) and FCT (POCI-01-0145- FEDER-03127, UIDB/04539/2020 and IF/01492/2015 to R.C.).