{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,6,30]],"date-time":"2026-06-30T22:12:06Z","timestamp":1782857526447,"version":"3.54.5"},"posted":{"date-parts":[[2026,6,29]]},"group-title":"Neuroscience","reference-count":74,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2026,6,29]],"date-time":"2026-06-29T00:00:00Z","timestamp":1782691200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/"}],"funder":[{"name":"Trans-NIH Metabolomics Consortium\u2019s 2021 Call for Applications","award":["(K.J.M., F.M.N.-N)"],"award-info":[{"award-number":["(K.J.M., F.M.N.-N)"]}]},{"name":"NIEHS Intramural Research","award":["ES103363-01 (A.J.)"],"award-info":[{"award-number":["ES103363-01 (A.J.)"]}]},{"name":"Office of the Assistant Secretary of Defense for Health Affairs and the Defense Health Agency J9, Research and Development Directorate","award":["CDMRPL-18-0-VR180205 (K.J.M.)"],"award-info":[{"award-number":["CDMRPL-18-0-VR180205 (K.J.M.)"]}]},{"name":"Fundaci\u00f3n S\u00e9neca, Agencia de Ciencia y Tecnolog\u00eda Regi\u00f3n de Murcia","award":["22395\/SF\/23 (F.M.N.-N)"],"award-info":[{"award-number":["22395\/SF\/23 (F.M.N.-N)"]}]},{"name":"\"Innovate Together\" NEI Intramural grant","award":["(K.J.M.)"],"award-info":[{"award-number":["(K.J.M.)"]}]},{"name":"Intramural Research Program of the National Institutes of Health","award":["(F.M.N.-N, R.M., K.O., A.J., K.J.M.)"],"award-info":[{"award-number":["(F.M.N.-N, R.M., K.O., A.J., K.J.M.)"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2026,6,29]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                <jats:p>\n                  Glaucoma is a group of eye diseases that affects 4 million people in the US and is one of the leading causes of vision loss due to damage to the eye\u2019s optic nerve (ON) which is composed of axons from retinal ganglion cells (RGCs) that transmit visual information to the brain. Injury to the ON often triggers RGC death and subsequent loss of visual function. Despite its increasing prevalence worldwide, effective therapies for glaucoma remain elusive. Notably, the thirteen-lined ground squirrel (TLGS) exhibits intrinsic neuroprotection during hibernation; however, reproducing this protective state pharmacologically has proven challenging. To elucidate the metabolic mechanisms underlying this resilience, we conducted untargeted metabolomic analyses on TLGS retinas at 6 hours, 3 days, and 7 days following ON crush. Retinas from awake and hibernating animals were compared to identify temporal and state-dependent metabolic signatures. Distinct metabolomic profiles were observed in hibernating animals relative to their awake counterparts. Pathway analyses revealed coordinated regulation of amino acid, lipid, and purine metabolism that likely contributes to hibernation-induced resilience. Furthermore, our findings indicate that hibernating TLGS retinas increase exosome biogenesis, prompting\n                  <jats:italic>in vitro<\/jats:italic>\n                  validation using TLGS-derived exosomes, which demonstrated robust neuroprotective and anti-inflammatory effects. Proteomic and transcriptomic characterization of exosomal cargo identified conserved miRNAs, mRNAs, and proteins implicated in redox balance, cytoskeletal stabilization, and stress-response regulation. Collectively, these data support the hypothesis that metabolic reprogramming and exosome-mediated intercellular signaling underlie hibernation-associated neuroprotection. Modulating these pathways may provide a blueprint for novel therapeutic strategies to mitigate neurodegeneration and promote recovery following optic nerve injury.\n                <\/jats:p>\n                <jats:sec>\n                  <jats:title>Graphical Abstract<\/jats:title>\n                  <jats:p>Illustration depicting state-dependent metabolic responses to optic nerve crush (ONC) injury in Thirteen-lined Ground Squirrels (TLGS). In Awake animals, injury triggers enhanced ATP production through the TCA cycle, leading to excessive reactive oxygen species (ROS) generation and subsequent retinal ganglion cell (RGC) death. In contrast, Hibernating animals shift toward lipid metabolism and utilize ATP for the biosynthesis of ceramides and sphingolipids, promoting membrane integrity and exosomal signaling. Additionally, a range of metabolites associated with hibernation-linked neuroprotection are elevated, contributing to enhanced RGC survival.<\/jats:p>\n                  <jats:fig id=\"ufig1\" position=\"float\" orientation=\"portrait\" fig-type=\"figure\">\n                    <jats:graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" xlink:href=\"733742v1_ufig1\" position=\"float\" orientation=\"portrait\"\/>\n                  <\/jats:fig>\n                <\/jats:sec>","DOI":"10.64898\/2026.06.23.733742","type":"posted-content","created":{"date-parts":[[2026,6,29]],"date-time":"2026-06-29T10:05:16Z","timestamp":1782727516000},"source":"Crossref","is-referenced-by-count":0,"title":["Exosomal Profiling Reveals Mechanisms of Hibernation-Associated Neuroprotection"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4121-514X","authenticated-orcid":false,"given":"Francisco M.","family":"Nadal-Nicol\u00e1s","sequence":"first","affiliation":[{"name":"National Eye Institute, NIH, Bethesda, MD, USA"},{"name":"Department of Ophthalmology, Faculty of Medicine, University of Murcia and Biomedical Research Institute of Murcia (IMIB-Pascual Parrilla), Murcia 30120, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rachel","family":"McNeel","sequence":"additional","affiliation":[{"name":"National Eye Institute, NIH, Bethesda, MD, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3607-4768","authenticated-orcid":false,"given":"Kirsten","family":"Overdahl","sequence":"additional","affiliation":[{"name":"Metabolomics Core Facility, National Institute of Environmental Health Sciences, NIH, Durahm, NC, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2228-6308","authenticated-orcid":false,"given":"Alan","family":"Jarmusch","sequence":"additional","affiliation":[{"name":"Metabolomics Core Facility, National Institute of Environmental Health Sciences, NIH, Durahm, NC, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9744-3152","authenticated-orcid":false,"given":"Kiyoharu J.","family":"Miyagishima","sequence":"additional","affiliation":[{"name":"National Eye Institute, NIH, Bethesda, MD, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026063014350624000_2026.06.23.733742v1.1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cbpb.2012.01.008"},{"key":"2026063014350624000_2026.06.23.733742v1.2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2018.03.010"},{"key":"2026063014350624000_2026.06.23.733742v1.3","doi-asserted-by":"crossref","unstructured":"Tang Z , Zhang S , Lee C , Kumar A , Arjunan P , Li Y , Zhang F , Li X : An optic nerve crush injury murine model to study retinal ganglion cell survival. 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