{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,31]],"date-time":"2026-03-31T20:01:12Z","timestamp":1774987272762,"version":"3.50.1"},"reference-count":85,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,2,24]],"date-time":"2026-02-24T00:00:00Z","timestamp":1771891200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,3,31]],"date-time":"2026-03-31T00:00:00Z","timestamp":1774915200000},"content-version":"vor","delay-in-days":35,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["npj Microgravity"],"DOI":"10.1038\/s41526-026-00565-5","type":"journal-article","created":{"date-parts":[[2026,2,24]],"date-time":"2026-02-24T06:47:31Z","timestamp":1771915651000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Spaceflight stressors impact on mitochondrial function and the risk for development of ocular pathology"],"prefix":"10.1038","volume":"12","author":[{"given":"Daniel P.","family":"LeBlanc","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Brian","family":"Butterfield","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Luis","family":"Jimenez-Chavez","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sara R.","family":"Zwart","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Scott M.","family":"Smith","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Xiao Wen","family":"Mao","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Brandon R.","family":"Macias","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Honglu","family":"Wu","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2026,2,24]]},"reference":[{"key":"565_CR1","first-page":"345","volume":"23","author":"CA Tobias","year":"1952","unstructured":"Tobias, C. A. Radiation hazards in high altitude aviation. J. Aviat Med. 23, 345\u2013372 (1952).","journal-title":"J. Aviat Med."},{"key":"565_CR2","unstructured":"NASA Mission Operations Branch Flight Crew Support Division. Apollo 11 Technical Crew Debriefing. (1969)."},{"key":"565_CR3","doi-asserted-by":"publisher","first-page":"10","DOI":"10.1667\/RR1580.1","volume":"172","author":"LT Chylack","year":"2009","unstructured":"Chylack, L. T. et al. NASA Study of Cataract in Astronauts (NASCA). Report 1: cross-sectional study of the relationship of exposure to space radiation and risk of lens opacity. Radiat. Res. 172, 10\u201320 (2009).","journal-title":"Radiat. Res."},{"key":"565_CR4","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1667\/RR2876.1","volume":"178","author":"LT Chylack","year":"2012","unstructured":"Chylack, L. T. et al. NASCA Report 2: longitudinal study of relationship of exposure to space radiation and risk of lens opacity. Radiat. Res. 178, 25\u201332 (2012).","journal-title":"Radiat. Res."},{"key":"565_CR5","doi-asserted-by":"publisher","first-page":"2058","DOI":"10.1016\/j.ophtha.2011.06.021","volume":"118","author":"TH Mader","year":"2011","unstructured":"Mader, T. H. et al. Optic disc edema, globe flattening, choroidal folds, and hyperopic shifts observed in astronauts after long-duration space flight. Ophthalmology 118, 2058\u20132069 (2011).","journal-title":"Ophthalmology"},{"key":"565_CR6","doi-asserted-by":"publisher","first-page":"2409","DOI":"10.1038\/s41433-023-02522-y","volume":"37","author":"J Ong","year":"2023","unstructured":"Ong, J., Mader, T. H., Gibson, C. R., Mason, S. S. & Lee, A. G. Spaceflight associated neuro-ocular syndrome (SANS): an update on potential microgravity-based pathophysiology and mitigation development. Eye 37, 2409\u20132415 (2023).","journal-title":"Eye"},{"key":"565_CR7","doi-asserted-by":"publisher","first-page":"1409","DOI":"10.1038\/s41433-024-02951-3","volume":"38","author":"E Waisberg","year":"2024","unstructured":"Waisberg, E. et al. Mitochondrial dysfunction in Spaceflight Associated Neuro-Ocular Syndrome (SANS): a molecular hypothesis in pathogenesis. Eye 38, 1409\u20131411 (2024).","journal-title":"Eye"},{"key":"565_CR8","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1097\/ICU.0b013e328349419d","volume":"22","author":"SA Schrier","year":"2011","unstructured":"Schrier, S. A. & Falk, M. J. Mitochondrial disorders and the eye. Curr. Opin. Ophthalmol. 22, 325 (2011).","journal-title":"Curr. Opin. Ophthalmol."},{"key":"565_CR9","doi-asserted-by":"publisher","DOI":"10.1016\/j.phrs.2024.107439","volume":"209","author":"IR Baduini","year":"2024","unstructured":"Baduini, I. R. et al. Type 2 diabetes mellitus and neurodegenerative disorders: the mitochondrial connection. Pharmacol. Res. 209, 107439 (2024).","journal-title":"Pharmacol. Res."},{"key":"565_CR10","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1159\/000316480","volume":"44","author":"SG Jarrett","year":"2010","unstructured":"Jarrett, S. G., Lewin, A. S. & Boulton, M. E. The importance of mitochondria in age-related and inherited eye disorders. Ophthalmic Res. 44, 179\u2013190 (2010).","journal-title":"Ophthalmic Res."},{"key":"565_CR11","doi-asserted-by":"publisher","first-page":"187","DOI":"10.3390\/metabo13020187","volume":"13","author":"DY Shu","year":"2023","unstructured":"Shu, D. Y. et al. Role of oxidative stress in ocular diseases: a balancing act. Metabolites 13, 187 (2023).","journal-title":"Metabolites"},{"key":"565_CR12","doi-asserted-by":"publisher","first-page":"926627","DOI":"10.3389\/fragi.2022.926627","volume":"3","author":"Y Tong","year":"2022","unstructured":"Tong, Y., Zhang, Z. & Wang, S. Role of mitochondria in retinal pigment epithelial aging and degeneration. Front. Aging 3, 926627 (2022).","journal-title":"Front. Aging"},{"key":"565_CR13","doi-asserted-by":"publisher","first-page":"1337","DOI":"10.1177\/11206721231221584","volume":"34","author":"E Waisberg","year":"2024","unstructured":"Waisberg, E. et al. Radiation-induced ophthalmic risks of long duration spaceflight: current investigations and interventions. Eur. J. Ophthalmol. 34, 1337\u20131345 (2024).","journal-title":"Eur. J. Ophthalmol."},{"key":"565_CR14","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1038\/nature03939","volume":"437","author":"RB Horne","year":"2005","unstructured":"Horne, R. B. et al. Wave acceleration of electrons in the Van Allen radiation belts. Nature 437, 227\u2013230 (2005).","journal-title":"Nature"},{"key":"565_CR15","doi-asserted-by":"crossref","unstructured":"Clement, G. & Slenzka, K. Fundamentals of Space Biology: Research on Cells, Animals, and Plants in Space. (Springer, New York, 2006).","DOI":"10.1007\/0-387-37940-1"},{"key":"565_CR16","doi-asserted-by":"crossref","unstructured":"Buckey, J. C. Space Physiology (Oxford University Press, 2006).","DOI":"10.1093\/oso\/9780195137255.001.0001"},{"key":"565_CR17","unstructured":"Report No. 098 \u2013 Guidance on Radiation Received in Space Activities (1989) - NCRP | Bethesda, MD. https:\/\/ncrponline.org\/shop\/reports\/report-no-098-guidance-on-radiation-received-in-space-activities-1989\/ (2018)."},{"key":"565_CR18","doi-asserted-by":"publisher","first-page":"198","DOI":"10.2307\/3578357","volume":"133","author":"DJ Brenner","year":"1993","unstructured":"Brenner, D. J., Medvedovsky, C., Huang, Y. & Worgul, B. V. Accelerated heavy particles and the lens. VIII. Comparisons between the effects of acute low doses of iron ions (190 keV\/microns) and argon ions (88 keV\/microns). Radiat. Res. 133, 198\u2013203 (1993).","journal-title":"Radiat. Res."},{"key":"565_CR19","doi-asserted-by":"publisher","first-page":"W345","DOI":"10.2214\/AJR.13.11242","volume":"203","author":"GD Frey","year":"2014","unstructured":"Frey, G. D. Radiation cataracts: new data and new recommendations. Am. J. Roentgenol. 203, W345\u2013W346 (2014).","journal-title":"Am. J. Roentgenol."},{"key":"565_CR20","doi-asserted-by":"crossref","unstructured":"Tomsia, M. et al. Long-term space missions\u2019 effects on the human organism: what we do know and what requires further research. Front. Physiol. 15, 1284644 (2024).","DOI":"10.3389\/fphys.2024.1284644"},{"key":"565_CR21","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1016\/j.pss.2012.07.014","volume":"74","author":"G Reitz","year":"2012","unstructured":"Reitz, G., Berger, T. & Matthiae, D. Radiation exposure in the moon environment. Planet. Space Sci. 74, 78\u201383 (2012).","journal-title":"Planet. Space Sci."},{"key":"565_CR22","unstructured":"Dishongh, L. Moon to MARS (M2M) LUNAR SURFACE DATA BOOK (National Aeronautics and Space Administration, 2025)."},{"key":"565_CR23","doi-asserted-by":"publisher","first-page":"1244797","DOI":"10.1126\/science.1244797","volume":"343","author":"DM Hassler","year":"2014","unstructured":"Hassler, D. M. et al. Mars\u2019 surface radiation environment measured with the Mars Science Laboratory\u2019s Curiosity rover. Science 343, 1244797 (2014).","journal-title":"Science"},{"key":"565_CR24","doi-asserted-by":"publisher","unstructured":"George, S. P. et al. Space radiation measurements during the Artemis I lunar mission. Nature 1\u20135, https:\/\/doi.org\/10.1038\/s41586-024-07927-7 (2024).","DOI":"10.1038\/s41586-024-07927-7"},{"key":"565_CR25","doi-asserted-by":"publisher","first-page":"A29","DOI":"10.1088\/0952-4746\/29\/2A\/S03","volume":"29","author":"JH Hendry","year":"2009","unstructured":"Hendry, J. H. et al. Human exposure to high natural background radiation: what can it teach us about radiation risks? J Radiol. Prot. 29, A29\u2013A42 (2009).","journal-title":"J Radiol. Prot."},{"key":"565_CR26","doi-asserted-by":"publisher","first-page":"650","DOI":"10.1016\/j.ijrobp.2010.09.056","volume":"79","author":"VSE Jeganathan","year":"2011","unstructured":"Jeganathan, V. S. E., Wirth, A. & MacManus, M. P. Ocular risks from orbital and periorbital radiation therapy: a critical review. Int. J. Radiat. Oncol.*Biol.*Phys. 79, 650\u2013659 (2011).","journal-title":"Int. J. Radiat. Oncol.*Biol.*Phys."},{"key":"565_CR27","doi-asserted-by":"publisher","first-page":"166","DOI":"10.1159\/000533426","volume":"9","author":"Z Oakey","year":"2023","unstructured":"Oakey, Z., Ye\u015filta\u015f, Y. S. & Singh, A. D. Radiation optic neuropathy: management options. Ocul Oncol. Pathol. 9, 166\u2013171 (2023).","journal-title":"Ocul Oncol. Pathol."},{"key":"565_CR28","doi-asserted-by":"publisher","first-page":"3153","DOI":"10.2147\/OPTH.S263291","volume":"14","author":"R Nuzzi","year":"2020","unstructured":"Nuzzi, R. et al. Ocular complications after radiation therapy: an observational study. Clin. Ophthalmol. 14, 3153\u20133166 (2020).","journal-title":"Clin. Ophthalmol."},{"key":"565_CR29","first-page":"465","volume":"55","author":"B Grajewski","year":"2011","unstructured":"Grajewski, B. et al. Airline pilot cosmic radiation and circadian disruption exposure assessment from logbooks and company records. Ann. Occup. Hyg. 55, 465\u2013475 (2011).","journal-title":"Ann. Occup. Hyg."},{"key":"565_CR30","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1016\/S0273-1177(02)00881-5","volume":"31","author":"L Narici","year":"2003","unstructured":"Narici, L. et al. ALTEA: anomalous long term effects in astronauts. A probe on the influence of cosmic radiation and microgravity on the central nervous system during long flights. Adv. Space Res. 31, 141\u2013146 (2003).","journal-title":"Adv. Space Res."},{"key":"565_CR31","first-page":"1","volume":"6","author":"T Akiyama","year":"2020","unstructured":"Akiyama, T. et al. How does spaceflight affect the acquired immune system? npj. Microgravity 6, 1\u20137 (2020).","journal-title":"Microgravity"},{"key":"565_CR32","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41526-021-00158-4","volume":"7","author":"OJ Juhl","year":"2021","unstructured":"Juhl, O. J. et al. Update on the effects of microgravity on the musculoskeletal system. npj Microgravity 7, 1\u201315 (2021).","journal-title":"npj Microgravity"},{"key":"565_CR33","volume":"Vasc 30","author":"S Patel","year":"2020","unstructured":"Patel, S. The effects of microgravity and space radiation on cardiovascular health: From low-Earth orbit and beyond. Int. J. Cardiol. Heart Vasc 30, 100595 (2020).","journal-title":"Int. J. Cardiol. Heart"},{"key":"565_CR34","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1016\/j.survophthal.2012.04.002","volume":"58","author":"G Taibbi","year":"2013","unstructured":"Taibbi, G., Cromwell, R. L., Kapoor, K. G., Godley, B. F. & Vizzeri, G. The effect of microgravity on ocular structures and visual function: a review. Surv. Ophthalmol. 58, 155\u2013163 (2013).","journal-title":"Surv. Ophthalmol."},{"key":"565_CR35","doi-asserted-by":"crossref","unstructured":"Mao, X., Stanbouly, S., Holley, J., Pecaut, M. & Crapo, J. Evidence of spaceflight-induced adverse effects on photoreceptors and retinal function in the mouse eye. Int. J. Mol. Sci. 24, 7362 (2023).","DOI":"10.3390\/ijms24087362"},{"key":"565_CR36","doi-asserted-by":"publisher","first-page":"1263","DOI":"10.3390\/life13061263","volume":"13","author":"EN Grigoryan","year":"2023","unstructured":"Grigoryan, E. N. Impact of microgravity and other spaceflight factors on retina of vertebrates and humans in vivo and in vitro. Life 13, 1263 (2023).","journal-title":"Life"},{"key":"565_CR37","first-page":"633","volume":"12","author":"JE Roberts","year":"2006","unstructured":"Roberts, J. E. et al. Simulated microgravity induced damage in human retinal pigment epithelial cells. Mol. Vis. 12, 633\u2013638 (2006).","journal-title":"Mol. Vis."},{"key":"565_CR38","doi-asserted-by":"publisher","DOI":"10.1038\/s41526-020-0097-9","volume":"6","author":"AG Lee","year":"2020","unstructured":"Lee, A. G. et al. Spaceflight associated neuro-ocular syndrome (SANS) and the neuro-ophthalmologic effects of microgravity: a review and an update. NPJ Microgravity 6, 7 (2020).","journal-title":"NPJ Microgravity"},{"key":"565_CR39","doi-asserted-by":"publisher","first-page":"R496","DOI":"10.1152\/ajpregu.00086.2018","volume":"315","author":"JC Buckey","year":"2018","unstructured":"Buckey, J. C. et al. Microgravity-induced ocular changes are related to body weight. Am. J. Physiol. Regul. Integr. Comp. Physiol. 315, R496\u2013R499 (2018).","journal-title":"Am. J. Physiol. Regul. Integr. Comp. Physiol."},{"key":"565_CR40","doi-asserted-by":"publisher","first-page":"1185","DOI":"10.1016\/j.cell.2020.11.002","volume":"183","author":"WA da Silveira","year":"2020","unstructured":"da Silveira, W. A. et al. Comprehensive multi-omics analysis reveals mitochondrial stress as a central biological hub for spaceflight impact. Cell 183, 1185\u20131201.e20 (2020).","journal-title":"Cell"},{"key":"565_CR41","doi-asserted-by":"publisher","first-page":"522","DOI":"10.1096\/fj.03-0419fje","volume":"18","author":"T Nikawa","year":"2004","unstructured":"Nikawa, T. et al. Skeletal muscle gene expression in space-flown rats. FASEB J. 18, 522\u2013524 (2004).","journal-title":"FASEB J."},{"key":"565_CR42","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-017-15612-1","volume":"7","author":"A Michaletti","year":"2017","unstructured":"Michaletti, A., Gioia, M., Tarantino, U. & Zolla, L. Effects of microgravity on osteoblast mitochondria: a proteomic and metabolomics profile. Sci Rep. 7, 15376 (2017).","journal-title":"Sci Rep."},{"key":"565_CR43","doi-asserted-by":"publisher","first-page":"890","DOI":"10.3109\/10715762.2014.920086","volume":"48","author":"XW Mao","year":"2014","unstructured":"Mao, X. W. et al. Biological and metabolic response in STS-135 space-flown mouse skin. Free Radic. Res. 48, 890\u2013897 (2014).","journal-title":"Free Radic. Res."},{"key":"565_CR44","doi-asserted-by":"publisher","DOI":"10.1126\/science.aau8650","volume":"364","author":"FE Garrett-Bakelman","year":"2019","unstructured":"Garrett-Bakelman, F. E. et al. The NASA twins study: a multidimensional analysis of a year-long human spaceflight. Science 364, eaau8650 (2019).","journal-title":"Science"},{"key":"565_CR45","doi-asserted-by":"publisher","DOI":"10.1038\/srep39015","volume":"6","author":"HP Indo","year":"2016","unstructured":"Indo, H. P. et al. Changes in mitochondrial homeostasis and redox status in astronauts following long stays in space. Sci Rep. 6, 39015 (2016).","journal-title":"Sci Rep."},{"key":"565_CR46","doi-asserted-by":"publisher","DOI":"10.1161\/JAHA.121.022055","volume":"10","author":"M Bisserier","year":"2021","unstructured":"Bisserier, M. et al. Cell-free mitochondrial DNA as a potential biomarker for Astronauts\u2019 Health. J. Am. Heart Assoc. 10, e022055 (2021).","journal-title":"J. Am. Heart Assoc."},{"key":"565_CR47","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41526-024-00406-3","volume":"10","author":"M Murgia","year":"2024","unstructured":"Murgia, M. et al. Spaceflight on the ISS changed the skeletal muscle proteome of two astronauts. npj Microgravity 10, 1\u201313 (2024).","journal-title":"npj Microgravity"},{"key":"565_CR48","doi-asserted-by":"publisher","first-page":"99","DOI":"10.2147\/EB.S9078","volume":"2","author":"M Wong-Riley","year":"2010","unstructured":"Wong-Riley, M. Energy metabolism of the visual system. Eye Brain 2, 99\u2013116 (2010).","journal-title":"Eye Brain"},{"key":"565_CR49","doi-asserted-by":"publisher","first-page":"395","DOI":"10.1017\/S0952523802194028","volume":"19","author":"QV Hoang","year":"2002","unstructured":"Hoang, Q. V., Linsenmeier, R. A., Chung, C. K. & Curcio, C. A. Photoreceptor inner segments in monkey and human retina: mitochondrial density, optics, and regional variation. Vis. Neuroscim 19, 395\u2013407 (2002).","journal-title":"Vis. Neuroscim"},{"key":"565_CR50","doi-asserted-by":"publisher","first-page":"513","DOI":"10.1016\/j.freeradbiomed.2016.07.016","volume":"97","author":"M Kubota","year":"2016","unstructured":"Kubota, M. et al. Mitochondrial oxygen metabolism in primary human lens epithelial cells: Association with age, diabetes and glaucoma. Free Radic. Biol. Med. 97, 513\u2013519 (2016).","journal-title":"Free Radic. Biol. Med."},{"key":"565_CR51","first-page":"2003","volume":"8","author":"C Guo","year":"2013","unstructured":"Guo, C., Sun, L., Chen, X. & Zhang, D. Oxidative stress, mitochondrial damage and neurodegenerative diseases. Neural Regen. Res. 8, 2003\u20132014 (2013).","journal-title":"Neural Regen. Res."},{"key":"565_CR52","doi-asserted-by":"publisher","first-page":"20190829","DOI":"10.1259\/bjr.20190829","volume":"93","author":"N Hamada","year":"2020","unstructured":"Hamada, N., Azizova, T. V. & Little, M. P. An update on effects of ionizing radiation exposure on the eye. Br. J. Radiol. 93, 20190829 (2020).","journal-title":"Br. J. Radiol."},{"key":"565_CR53","doi-asserted-by":"publisher","first-page":"11047","DOI":"10.3390\/ijms222011047","volume":"22","author":"D Averbeck","year":"2021","unstructured":"Averbeck, D. & Rodriguez-Lafrasse, C. Role of mitochondria in radiation responses: epigenetic, metabolic, and signaling impacts. Int. J. Mol. Sci. 22, 11047 (2021).","journal-title":"Int. J. Mol. Sci."},{"key":"565_CR54","doi-asserted-by":"publisher","DOI":"10.1016\/j.exer.2022.109192","volume":"223","author":"RB Richardson","year":"2022","unstructured":"Richardson, R. B. The role of oxygen and the Goldilocks range in the development of cataracts induced by space radiation in US astronauts. Exp. Eye Res. 223, 109192 (2022).","journal-title":"Exp. Eye Res."},{"key":"565_CR55","first-page":"e56793","volume":"16","author":"C Karakosta","year":"2024","unstructured":"Karakosta, C., Samiotaki, M., Panayotou, G., Papaconstantinou, D. S. & Moschos, M. M. Lens cytoskeleton: an update on the etiopathogenesis of human cataracts. Cureus 16, e56793 (2024).","journal-title":"Cureus"},{"key":"565_CR56","unstructured":"Nizami, A. A., Gurnani, B. & Gulani, A. C. Cataract. in StatPearls (StatPearls Publishing, 2024)."},{"key":"565_CR57","doi-asserted-by":"publisher","first-page":"108662","DOI":"10.1016\/j.exer.2021.108662","volume":"209","author":"A Shiels","year":"2021","unstructured":"Shiels, A. & Hejtmancik, J. F. Inherited cataracts: genetic mechanisms and pathways new and old. Exp. Eye Res. 209, 108662 (2021).","journal-title":"Exp. Eye Res."},{"key":"565_CR58","doi-asserted-by":"crossref","first-page":"A56","DOI":"10.3357\/ASEM.86050.2007","volume":"78","author":"JA Jones","year":"2007","unstructured":"Jones, J. A. et al. Cataract formation mechanisms and risk in aviation and space crews. Aviat. Space Environ. Med. 78, A56\u2013A66 (2007).","journal-title":"Aviat. Space Environ. Med."},{"key":"565_CR59","doi-asserted-by":"publisher","first-page":"1102","DOI":"10.1001\/archopht.123.8.1102","volume":"123","author":"V Rafnsson","year":"2005","unstructured":"Rafnsson, V. et al. Cosmic radiation increases the risk of nuclear cataract in airline pilots: a population-based case-control study. Arch. Ophthalmol. 123, 1102\u20131105 (2005).","journal-title":"Arch. Ophthalmol."},{"key":"565_CR60","doi-asserted-by":"publisher","first-page":"895","DOI":"10.3357\/ASEM.3034.2011","volume":"82","author":"AC Chorley","year":"2011","unstructured":"Chorley, A. C., Evans, B. J. W. & Benwell, M. J. Civilian pilot exposure to ultraviolet and blue light and pilot use of sunglasses. Aviat. Space Environ. Med. 82, 895\u2013900 (2011).","journal-title":"Aviat. Space Environ. Med."},{"key":"565_CR61","doi-asserted-by":"publisher","first-page":"416","DOI":"10.1038\/s41433-023-02742-2","volume":"38","author":"E Waisberg","year":"2024","unstructured":"Waisberg, E., Ong, J. & Lee, A. G. Space radiation and the potential for early cataract development. Eye 38, 416\u2013417 (2024).","journal-title":"Eye"},{"key":"565_CR62","doi-asserted-by":"publisher","DOI":"10.1155\/2020\/4703286","volume":"2020","author":"S Furukawa","year":"2020","unstructured":"Furukawa, S. et al. Space radiation biology for \u201cliving in space. Biomed. Res. Int. 2020, 4703286 (2020).","journal-title":"Biomed. Res. Int."},{"key":"565_CR63","doi-asserted-by":"publisher","first-page":"491","DOI":"10.3390\/life4030491","volume":"4","author":"JC Chancellor","year":"2014","unstructured":"Chancellor, J. C., Scott, G. B. I. & Sutton, J. P. Space radiation: the number one risk to astronaut health beyond low earth orbit. Life 4, 491\u2013510 (2014).","journal-title":"Life"},{"key":"565_CR64","doi-asserted-by":"publisher","first-page":"607","DOI":"10.1016\/j.freeradbiomed.2013.07.024","volume":"65","author":"WW-Y Kam","year":"2013","unstructured":"Kam, W. W.-Y. & Banati, R. B. Effects of ionizing radiation on mitochondria. Free Radic. Biol. Med. 65, 607\u2013619 (2013).","journal-title":"Free Radic. Biol. Med."},{"key":"565_CR65","first-page":"1629","volume":"6","author":"J Kaur","year":"2012","unstructured":"Kaur, J., Kukreja, S., Kaur, A., Malhotra, N. & Kaur, R. The oxidative stress in cataract patients. J. Clin. Diagn. Res. 6, 1629\u20131632 (2012).","journal-title":"J. Clin. Diagn. Res."},{"key":"565_CR66","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1016\/j.bbadis.2016.09.020","volume":"1863","author":"L Brennan","year":"2017","unstructured":"Brennan, L., Khoury, J. & Kantorow, M. Parkin elimination of mitochondria is important for maintenance of lens epithelial cell ROS levels and survival upon oxidative stress exposure. Biochim. Biophys. Acta Mol. Basis Dis. 1863, 21\u201332 (2017).","journal-title":"Biochim. Biophys. Acta Mol. Basis Dis."},{"key":"565_CR67","doi-asserted-by":"publisher","first-page":"1034","DOI":"10.1038\/s41433-023-02841-0","volume":"38","author":"M Masalkhi","year":"2024","unstructured":"Masalkhi, M., Ong, J., Waisberg, E. & Lee, A. G. Ocular oxidative changes and antioxidant therapy during spaceflight. Eye 38, 1034\u20131035 (2024).","journal-title":"Eye"},{"key":"565_CR68","doi-asserted-by":"publisher","DOI":"10.1038\/s41526-021-00162-8","volume":"7","author":"X G\u00f3mez","year":"2021","unstructured":"G\u00f3mez, X. et al. Key points for the development of antioxidant cocktails to prevent cellular stress and damage caused by reactive oxygen species (ROS) during manned space missions. NPJ Microgravity 7, 35 (2021).","journal-title":"NPJ Microgravity"},{"key":"565_CR69","doi-asserted-by":"publisher","first-page":"514","DOI":"10.1038\/s41420-024-02277-9","volume":"10","author":"JD Rondeau","year":"2024","unstructured":"Rondeau, J. D. et al. Mitochondria-targeted antioxidant MitoQ radiosensitizes tumors by decreasing mitochondrial oxygen consumption. Cell Death Discov. 10, 514 (2024).","journal-title":"Cell Death Discov."},{"key":"565_CR70","doi-asserted-by":"publisher","DOI":"10.1186\/s12974-019-1627-9","volume":"16","author":"W Zhao","year":"2019","unstructured":"Zhao, W. et al. Elamipretide (SS-31) improves mitochondrial dysfunction, synaptic and memory impairment induced by lipopolysaccharide in mice. J. Neuroinflamm. 16, 230 (2019).","journal-title":"J. Neuroinflamm."},{"key":"565_CR71","doi-asserted-by":"publisher","first-page":"31","DOI":"10.3390\/biology8020031","volume":"8","author":"JT Eells","year":"2019","unstructured":"Eells, J. T. Mitochondrial dysfunction in the aging retina. Biology 8, 31 (2019).","journal-title":"Biology"},{"key":"565_CR72","doi-asserted-by":"publisher","first-page":"2416","DOI":"10.1038\/s41433-023-02523-x","volume":"37","author":"BS Chen","year":"2023","unstructured":"Chen, B. S., Harvey, J. P., Gilhooley, M. J., Jurkute, N. & Yu-Wai-Man, P. Mitochondria and the eye\u2014manifestations of mitochondrial diseases and their management. Eye 37, 2416\u20132425 (2023).","journal-title":"Eye"},{"key":"565_CR73","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-49453-x","volume":"9","author":"EG Overbey","year":"2019","unstructured":"Overbey, E. G. et al. Spaceflight influences gene expression, photoreceptor integrity, and oxidative stress-related damage in the murine retina. Sci Rep. 9, 13304 (2019).","journal-title":"Sci Rep."},{"key":"565_CR74","doi-asserted-by":"publisher","DOI":"10.1016\/j.arr.2023.101909","volume":"87","author":"M Capri","year":"2023","unstructured":"Capri, M. et al. Long-term human spaceflight and inflammaging: does it promote aging? Ageing Res. Rev. 87, 101909 (2023).","journal-title":"Ageing Res. Rev."},{"key":"565_CR75","doi-asserted-by":"publisher","first-page":"340","DOI":"10.1667\/RR3309.1","volume":"180","author":"XW Mao","year":"2013","unstructured":"Mao, X. W. et al. Spaceflight environment induces mitochondrial oxidative damage in ocular tissue. Radiat. Res. 180, 340\u2013350 (2013).","journal-title":"Radiat. Res."},{"key":"565_CR76","doi-asserted-by":"publisher","first-page":"958","DOI":"10.1016\/j.bbrc.2018.04.214","volume":"500","author":"X Jin","year":"2018","unstructured":"Jin, X. et al. Different mitochondrial fragmentation after irradiation with X-rays and carbon ions in HeLa cells and its influence on cellular apoptosis. Biochem. Biophys. Res. Commun. 500, 958\u2013965 (2018).","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"565_CR77","doi-asserted-by":"publisher","first-page":"4959","DOI":"10.1073\/pnas.96.9.4959","volume":"96","author":"L-J Wu","year":"1999","unstructured":"Wu, L.-J. et al. Targeted cytoplasmic irradiation with alpha particles induces mutations in mammalian cells. Proc. Natl. Acad. Sci. USA 96, 4959\u20134964 (1999).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"565_CR78","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-45669-z","volume":"9","author":"K Ariyoshi","year":"2019","unstructured":"Ariyoshi, K. et al. Radiation-induced bystander effect is mediated by mitochondrial DNA in exosome-like vesicles. Sci. Rep. 9, 9103 (2019).","journal-title":"Sci. Rep."},{"key":"565_CR79","doi-asserted-by":"publisher","DOI":"10.1038\/srep46684","volume":"7","author":"DWM Walsh","year":"2017","unstructured":"Walsh, D. W. M. et al. Live cell imaging of mitochondria following targeted irradiation in situ reveals rapid and highly localized loss of membrane potential. Sci. Rep. 7, 46684 (2017).","journal-title":"Sci. Rep."},{"key":"565_CR80","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1667\/RR14977.1","volume":"190","author":"XW Mao","year":"2018","unstructured":"Mao, X. W. et al. Acute effect of low-dose space radiation on mouse retina and retinal endothelial cells. Radiat. Res. 190, 45\u201352 (2018).","journal-title":"Radiat. Res."},{"key":"565_CR81","doi-asserted-by":"publisher","first-page":"39","DOI":"10.3390\/cells13010039","volume":"13","author":"L Wang","year":"2023","unstructured":"Wang, L. et al. Dose-dependent effects of radiation on mitochondrial morphology and clonogenic cell survival in human microvascular endothelial cells. Cells 13, 39 (2023).","journal-title":"Cells"},{"key":"565_CR82","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2120439119","volume":"119","author":"G Barisano","year":"2022","unstructured":"Barisano, G. et al. The effect of prolonged spaceflight on cerebrospinal fluid and perivascular spaces of astronauts and cosmonauts. Proc. Natl. Acad. Sci. USA 119, e2120439119 (2022).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"565_CR83","unstructured":"Smith, S. M., Zwart, S. R., Douglas, G. L. & Heer, M. Human Adaptation to Spaceflight: The Role of Food and Nutrition (Government Publishing Office, 2014)."},{"key":"565_CR84","doi-asserted-by":"publisher","first-page":"78","DOI":"10.3389\/fphys.2019.00078","volume":"10","author":"JJE Janssen","year":"2019","unstructured":"Janssen, J. J. E., Grefte, S., Keijer, J. & de Boer, V. C. J. Mito-nuclear communication by mitochondrial metabolites and its regulation by B-Vitamins. Front. Physiol. 10, 78 (2019).","journal-title":"Front. Physiol."},{"key":"565_CR85","doi-asserted-by":"publisher","first-page":"648958","DOI":"10.3389\/fneur.2021.648958","volume":"12","author":"J Ong","year":"2021","unstructured":"Ong, J., Lee, A. G. & Moss, H. E. Head-down tilt bed rest studies as a terrestrial analog for spaceflight associated neuro-ocular syndrome. Front. Neurol. 12, 648958 (2021).","journal-title":"Front. Neurol."}],"container-title":["npj Microgravity"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41526-026-00565-5","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41526-026-00565-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41526-026-00565-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,31]],"date-time":"2026-03-31T19:17:37Z","timestamp":1774984657000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41526-026-00565-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,24]]},"references-count":85,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["565"],"URL":"https:\/\/doi.org\/10.1038\/s41526-026-00565-5","relation":{},"ISSN":["2373-8065"],"issn-type":[{"value":"2373-8065","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,24]]},"assertion":[{"value":"22 January 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"18 January 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"24 February 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"30"}}