Publications

Tolman N, Li T, Balasubramanian R, Li G, Pfeiffer R, Bupp-Chickering V, Kelly RA, Simón M, Peregrin J, Montgomery C, Jones B, Stamer WD, Qian J, John SWM. Single-cell profiling of trabecular meshwork identifies mitochondrial dysfunction in a glaucoma model that is protected by vitamin B3 treatment. Elife. 2026 Jan 20;14:RP107161. doi: 10.7554/eLife.107161. PMID: 41556506; PMCID: PMC12818872. https://pubmed.ncbi.nlm.nih.gov/41556506/ 

Balasubramanian R, Tolman N, Li T, Hannan A, Bupp-Chickering V, Polanco K, Bhandari A, Zhou S, Simón M, Peregrin J, Montgomery C, Kizhatil K, Qian J, John SWM. Single-Cell Characterization of Anterior Segment Development: Cell Types, Pathways, and Signals Driving Formation of the Trabecular Meshwork and Schlemm's Canal. bioRxiv [Preprint]. 2025 Aug 21:2025.08.15.670438. doi: 10.1101/2025.08.15.670438. PMID: 40894664; PMCID: PMC12393362. https://pubmed.ncbi.nlm.nih.gov/40894664/ 

Kelly RA, Kuhn MS, Reina-Torres E, Balasubramanian R, Perkumas KM, Li G, Takahashi T, John SW, Elliott MH, Overby DR, Stamer WD. Endothelial cell-specific postnatal deletion of Nos3 preserves intraocular pressure homeostasis via macrophage recruitment and NOS2 upregulation. J Clin Invest. 2025 Feb 11;135(7):e183440. doi: 10.1172/JCI183440. PMID: 39932792; PMCID: PMC11957705. https://pubmed.ncbi.nlm.nih.gov/39932792/ 

Liu KC, Grimsrud AO, Suarez MF, Schuman D, De Ieso ML, Kuhn M, Kelly RA, Mathew R, Kalnitsky J, Mack M, Ginhoux F, Bupp-Chickering V, Balasubramanian R, John SWM, Stamer WD, Saban DR. Resident Tissue Macrophages Govern Intraocular Pressure Homeostasis. bioRxiv [Preprint]. 2025 Jan 27:2025.01.25.634888. doi: 10.1101/2025.01.25.634888. PMID: 39975071; PMCID: PMC11838227. https://pubmed.ncbi.nlm.nih.gov/39975071/ 

Kizhatil K, Clark GM, Sunderland DK, Bhandari A, Horbal LJ, Balasubramanian R, John SWM. FYN regulates aqueous humor outflow and IOP through the phosphorylation of VE-CADHERIN. Nat Commun. 2025 Jan 2;16(1):51. doi: 10.1038/s41467-024-55232-8. PMID: 39746990; PMCID: PMC11696269. https://pubmed.ncbi.nlm.nih.gov/39746990/ 

Balasubramanian R, Kizhatil K, Li T, Tolman N, Bhandari A, Clark G, Bupp-Chickering V, Kelly RA, Zhou S, Peregrin J, Simón M, Montgomery C, Stamer WD, Qian J, John SWM. Transcriptomic profiling of Schlemm's canal cells reveals a lymphatic-biased identity and three major cell states. Elife. 2024 Oct 18;13:RP96459. doi: 10.7554/eLife.96459. PMID: 39422453; PMCID: PMC11488849. https://pubmed.ncbi.nlm.nih.gov/39422453/ 

Slavi N, Balasubramanian R, Lee MA, Liapin M, Oaks-Leaf R, Peregrin J, Potenski A, Troy CM, Ross ME, Herrera E, Kosmidis S, John SWM, Mason CA. CyclinD2-mediated regulation of neurogenic output from the retinal ciliary margin is perturbed in albinism. Neuron. 2023 Jan 4;111(1):49-64.e5. doi: 10.1016/j.neuron.2022.10.025. Epub 2022 Nov 8. PMID: 36351424; PMCID: PMC9822872. https://pubmed.ncbi.nlm.nih.gov/36351424/ 

Balasubramanian R, Min X, Quinn PMJ, Giudice QL, Tao C, Polanco K, Makrides N, Peregrin J, Bouaziz M, Mao Y, Wang Q, da Costa BL, Buenaventura D, Wang F, Ma L, Tsang SH, Fabre PJ, Zhang X. Phase transition specified by a binary code patterns the vertebrate eye cup. Sci Adv. 2021 Nov 12;7(46):eabj9846. doi: 10.1126/sciadv.abj9846. Epub 2021 Nov 10. PMID: 34757798; PMCID: PMC8580326. https://pubmed.ncbi.nlm.nih.gov/34757798/ 

Tolman NG, Balasubramanian R, Macalinao DG, Kearney AL, MacNicoll KH, Montgomery CL, de Vries WN, Jackson IJ, Cross SH, Kizhatil K, Nair KS, John SWM. Genetic background modifies vulnerability to glaucoma-related phenotypes in Lmx1b mutant mice. Dis Model Mech. 2021 Feb 19;14(2):dmm046953. doi: 10.1242/dmm.046953. PMID: 33462143; PMCID: PMC7903917. https://pubmed.ncbi.nlm.nih.gov/33462143/ 

Harder JM, Guymer C, Wood JPM, Daskalaki E, Chidlow G, Zhang C, Balasubramanian R, Cardozo BH, Foxworth NE, Deering KE, Ouellette TB, Montgomery C, Wheelock CE, Casson RJ, Williams PA, John SWM. Disturbed glucose and pyruvate metabolism in glaucoma with neuroprotection by pyruvate or rapamycin. Proc Natl Acad Sci U S A. 2020 Dec 29;117(52):33619-33627. doi: 10.1073/pnas.2014213117. Epub 2020 Dec 14. PMID: 33318177; PMCID: PMC7776900. https://pubmed.ncbi.nlm.nih.gov/33318177/ 

Iwai-Takekoshi L, Balasubramanian R, Sitko A, Khan R, Weinreb S, Robinson K, Mason C. Activation of Wnt signaling reduces ipsilaterally projecting retinal ganglion cells in pigmented retina. Development. 2018 Nov 2;145(21):dev163212. doi: 10.1242/dev.163212. PMID: 30254141; PMCID: PMC6240318. https://pubmed.ncbi.nlm.nih.gov/30254141/ 

Zhang Y, Yang Y, Jiang M, Huang SX, Zhang W, Al Alam D, Danopoulos S, Mori M, Chen YW, Balasubramanian R, Chuva de Sousa Lopes SM, Serra C, Bialecka M, Kim E, Lin S, Toste de Carvalho ALR, Riccio PN, Cardoso WV, Zhang X, Snoeck HW, Que J. 3D Modeling of Esophageal Development using Human PSC-Derived Basal Progenitors Reveals a Critical Role for Notch Signaling. Cell Stem Cell. 2018 Oct 4;23(4):516-529.e5. doi: 10.1016/j.stem.2018.08.009. Epub 2018 Sep 20. PMID: 30244870; PMCID: PMC6282026. https://pubmed.ncbi.nlm.nih.gov/30244870/ 

Balasubramanian R, Bui A, Dong X, Gan L. Lhx9 Is Required for the Development of Retinal Nitric Oxide-Synthesizing Amacrine Cell Subtype. Mol Neurobiol. 2018 Apr;55(4):2922-2933. doi: 10.1007/s12035-017-0554-y. Epub 2017 Apr 29. PMID: 28456934; PMCID: PMC5898628. https://pubmed.ncbi.nlm.nih.gov/28456934/ 

Ding Q, Balasubramanian R, Zheng D, Liang G, Gan L. Barhl2 Determines the Early Patterning of the Diencephalon by Regulating Shh. Mol Neurobiol. 2017 Aug;54(6):4414-4420. doi: 10.1007/s12035-016-0001-5. Epub 2016 Jun 27. PMID: 27349434; PMCID: PMC5191999. https://pubmed.ncbi.nlm.nih.gov/27349434/ 

Balasubramanian R, Zhang X. Mechanisms of FGF gradient formation during embryogenesis. Semin Cell Dev Biol. 2016 May;53:94-100. doi: 10.1016/j.semcdb.2015.10.004. Epub 2015 Oct 8. PMID: 26454099; PMCID: PMC4906438. https://pubmed.ncbi.nlm.nih.gov/26454099/ 

Balasubramanian R, Gan L. Development of Retinal Amacrine Cells and Their Dendritic Stratification. Curr Ophthalmol Rep. 2014 Sep 1;2(3):100-106. doi: 10.1007/s40135-014-0048-2. PMID: 25170430; PMCID: PMC4142557. https://pubmed.ncbi.nlm.nih.gov/25170430/ 

Balasubramanian R, Bui A, Xie X, Deng M, Gan L. Generation and characterization of Lhx9-GFPCreER(T2) knock-in mouse line. Genesis. 2014 Sep;52(9):827-832. doi: 10.1002/dvg.22805. Epub 2014 Aug 12. PMID: 25112520; PMCID: PMC4167977. https://pubmed.ncbi.nlm.nih.gov/25112520/ 

Balasubramanian R, Bui A, Ding Q, Gan L. Expression of LIM-homeodomain transcription factors in the developing and mature mouse retina. Gene Expr Patterns. 2014 Jan;14(1):1-8. doi: 10.1016/j.gep.2013.12.001. Epub 2013 Dec 10. PMID: 24333658; PMCID: PMC3921069. https://pubmed.ncbi.nlm.nih.gov/24333658/