Articles in press and preprints

Williams PA*†, Harder JM*, Guymer C, Wood JPM, Daskalakis E, Chidlow G, Cardozo BH, Foxworth NE, Cochran KE, Ouellette TB, Wheelock CE, Casson RJ, John SWM†. “Oral pyruvate prevents glaucomatous neurodegeneration”. bioRxiv. 2020 (doi: 10.1101/2020.05.02.072215) [pdf]

* Authors contributed equally to this work, † To whom correspondence should be addressed

Tribble JR*, Kokkali E*, Otmani A, Plastino F, Lardner E, Vohra R, Kolko M, Andre H, Morgan JE, Williams PA. “When is a control not a control? Reactive microglia occur throughout the control contralateral visual pathway in experimental glaucoma”. bioRxiv. 2019 (doi: 10.1101/853275) [pdf]

* Authors contributed equally to this work

Petrova V, Pearson CS, Tribble JR, Solano AG, Reid E, Geller HM, Williams PA, Eva R, Fawcett JW. “Protrudin functions as a scaffold in the endoplasmic reticulum to support axon regeneration in the adult central nervous system”. bioRxiv. 2019 (doi: 10.1101/2020.01.15.907550) [pdf]

Hui F, Tang J, Williams PA, McGuinness MB, Hadoux X, Casson RJ, Coote M, Trounce IA, Martin KR, van Wijngaarden P, Crowston JG. “Improvement in inner retinal function in glaucoma in response to nicotinamide (vitamin B3) supplementation: A crossover randomized clinical trial”. medRxiv. 2020 (doi: 10.1101/2020.01.28.20019075) [pdf]



Tribble JR*, Harder JM*, Williams PA†, John SWM†. “Suppression of homeostatic gene expression and increased expression of metabolism-related genes are early features of glaucoma in optic nerve head microglia”. Molecular Brain. 2019 (doi: 10.1101/856427) [pdf]

* Authors contributed equally to this work, † To whom correspondence should be addressed



Tribble JR, Vasalauskaite A, Redmond T, Young RD, Hassan S, Fautsch MP, Sengpiel F, Williams PA, Morgan JE. “Midget retinal ganglion cell dendritic and mitochondrial degeneration is an early feature of human glaucoma”. Brain Communications. 2019 (doi: 10.1093/braincomms/fcz035) [pdf]

Williams PA, Crowston JG. “Targeting Metabolic Vulnerabilities for Neuroprotection in Glaucoma: B3 or not B3? That is the question”. Glaucoma Today. 2019 ( [pdf]

Williams PA, Braine CE, Kizhatil K, Foxworth NE, Tolman NG, Harder JM, Scott RA, Sousa GL, Panitch A, Howell GR, John SWM. “Inhibition of monocyte-like cell extravasation protects from neurodegeneration in DBA/2J glaucoma”. Molecular Neurodegeneration. 2019 (doi: 10.1186/s13024-018-0303-3) [pdf]



Tribble JR, Williams PA, Caterson B, Sengpiel F, Morgan JE. “Digestion of the glycosaminoglycan extracellular matrix by chondroitinase ABC supports retinal ganglion cell dendritic preservation in a rodent model of experimental glaucoma”. Molecular Brain. 2018 (doi: 10.1186/s13041-018-0412-5) [pdf]

Harder JM, Williams PA, Soto I, Foxworth NE, Fernandes KA, Freeburg NF, Howell GR, Libby R, John SWM. “Jnk2 deficiency increases the rate of glaucomatous neurodegeneration in ocular hypertensive DBA/2J mice”. Cell Death & Disease. 2018 (doi: 10.1038/s41419-018-0705-8) [pdf]

Choquet H, Paylakhi S, Kneeland SC, Thai KK, Hoffmann TJ, Yin J, Kvale MN, Banda Y, Tolman NG, Williams PA, Schaefer C, Melles R, Risch N, John SWM, Nair KS, Jorgenson E. “A multiethnic genome-wide association study of primary open-angle glaucoma identifies novel risk loci”. Nature Communications. 2018 (doi: 10.1038/s41467-018-04555-4) [pdf]

Williams PA, Harder JM, Cardozo BH, Foxworth NE, John SWM. “Nicotinamide treatment robustly protects from inherited mouse glaucoma”. Communicative & Integrative Biology. 2018 (doi: 10.1080/19420889.2017.1356956[pdf]



Williams PA, Harder JM, John SWM. “Glaucoma as a metabolic optic neuropathy: making the case for nicotinamide treatment in glaucoma”. Journal of Glaucoma. 2017 (doi: 10.1097/IJG.0000000000000767[pdf]

Williams PA, John SWM. “Glaucoma dialogue – 71509. Vitamin B3 modulates mitochondrial vulnerability and prevents glaucoma in aged mice”. International Glaucoma Review. 2017 (

Williams PA, Braine CE, Foxworth NE, Cochran KE, John SWM. “GlyCAM1 negatively modulates monocyte entry into the optic nerve head and contributes to radiation-based protection in glaucoma”. Journal of Neuroinflammation. 2017 (doi: 10.1186/s12974-017-0868-8[pdf]

Harder JM, Braine CE, Williams PA, Zhu X, MacNicoll KH, Sousa GL, Buchanan RA, Smith RS, Libby RT, Howell GR, John SWM. “Early immune responses are independent of RGC dysfunction in glaucoma with complement component C3 being protective”. PNAS. 2017 (doi: 10.1073/pnas.1608769114[pdf]

Williams PA, Harder JM, Foxworth NE, Cardozo BH, John SWM. “Nicotinamide and WLDS act together to prevent neurodegeneration in glaucoma”. Frontiers in Neuroscience. 2017 (doi: 10.3389/fnins.2017.00232[pdf]

Williams PA, Harder JM, Foxworth NE, Cochran KE, Philip VM, Porciatti V, Smithies O, John SWM. “Vitamin B3 modulates mitochondrial vulnerability and prevents glaucoma in aged mice”. Science. 2017 (doi: 10.1126/science.aal0092[pdf]

Perspective in Science [link]

Clinical Implications of Basic Research in NEJM [link]

Williams PA, Marsh-Armstrong N, Howell GR. “Neuroinflammation in glaucoma: A new opportunity”. Experimental Eye Research. 2017 (doi: 10.1016/j.exer.2017.02.014[pdf]



Williams PA*, Tribble JR*, Pepper KW, Cross SD, Morgan BP, Morgan JE, John SWM, Howell GR. “Inhibition of the classical pathway of the complement cascade prevents early dendritic and synaptic degeneration in glaucoma”. Molecular Neurodegeneration. 2016 (doi: 10.1186/s13024-016-0091-6[pdf]

* Authors contributed equally to this work



Fernandes KA, Harder JM, Williams PA, Rausch RL, Kiernan AE, Nair KS, Anderson MG, John SWM, Howell GR, Libby RT. “Using genetic mouse models to gain insight into glaucoma: Past results and future possibilities”. Experimental Eye Research. 2015 (doi: 10.1016/j.exer.2015.06.019[pdf]

Lee ST, Williams PA, Braine CE, Lin DT, John SWM, Irazoqui PP. “A miniature, fiber-coupled, wireless, deep-brain optogenetic stimulator”. Transactions on Neural Systems and Rehabilitation Engineering. 2015 (doi: 10.1109/TNSRE.2015.2391282[pdf]



Williams PA, Howell GR, Barbay JM, Braine CE, Sousa GL, John SWM, Morgan JE. “Retinal ganglion cell dendritic atrophy in DBA/2J glaucoma”. PLoS One. 2013 (doi: 10.1371/journal.pone.0072282[pdf]

Williams PA, Thirgood RA, Littlewood E, Votruba M, Oliphant H, Good MA, Williams J, Morgan JE. “Retinal ganglion cell dendritic degeneration in a mouse model of Alzheimer’s disease”. Neurobiology of Aging. 2013 (doi: 10.1016/j.neurobiolaging.2013.01.006[pdf]



Williams PA, Piechota M, Von Ruhland C, Taylor E, Morgan JE, Votruba M. “Opa1 is essential for retinal ganglion cell synaptic architecture and connectivity”. Brain. 2012 (doi: 10.1093/brain/awr330[pdf]



Barnard AR, Issa PC, Perganta G, Williams PA, Davies V, Sekaran S, Votruba M, MacLaren RE. “Specific deficits in visual electrophysiology in a mouse model of dominant optic atrophy”. Experimental Eye Research. 2011 (doi: 10.1016/j.exer.2011.07.004[pdf]

Williams PA, Morgan JE, Votruba M. “Mouse models of dominant optic atrophy: What do they tell us about the pathophysiology of visual loss?”. Vision Research. 2011 (doi: 10.1016/j.visres.2010.08.031[pdf]



Williams PA, Morgan JE, Votruba, M. “Opa1 deficiency in a mouse model of dominant optic atrophy leads to retinal ganglion cell dendropathy”. Brain. 2010 (doi: 10.1093/brain/awq218[pdf]