{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T17:03:55Z","timestamp":1790874235381,"version":"4.1.0"},"posted":{"date-parts":[[2026,9,28]]},"group-title":"Cell Biology","reference-count":106,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T00:00:00Z","timestamp":1790553600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"award":["R01AR076924"],"award-info":[{"award-number":["R01AR076924"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"award":["S10OD032364"],"award-info":[{"award-number":["S10OD032364"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2026,9,28]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                <jats:p>\n                  All-\n                  <jats:italic>trans<\/jats:italic>\n                  -retinal (atRAL) is a photoreactive aldehyde generated during visual pigment regeneration, and impaired atRAL clearance contributes to oxidative stress and retinal degeneration. Retinol dehydrogenase 12 (RDH12) reduces atRAL to all-trans-retinol, yet the temporal mechanisms linking its upstream enzymatic activity to downstream cellular stress adaptations remain poorly defined. Here, we utilized stable RDH12-expressing HEK293T cells to match proteomic and lipidomic profiles and performed cell viability assays to map the transition from acute atRAL stress to cellular recovery. RDH12 expression increased cellular tolerance to retinaldehyde-induced cytotoxicity. Acute atRAL exposure engaged the supply arm of the KEAP1-NRF2 program with increased HMOX1, SLC7A11, GCLC, MGST2, and MGST3, while the terminal glutathione peroxidase arm declined. Acute exposure was accompanied by membrane remodeling, indicated by increased glycerophospholipid enzyme abundance and increased stress-signaling ceramides and lysophosphatidylcholines. Concurrently, acute stress induced upregulation of the transferrin receptor (TFRC) and SLC11A2, which influence intracellular iron flux and enable Fenton chemistry that can lead to oxidative stress. During recovery, RDH12-expressing cells exhibited a distinct post-atRAL molecular state characterized by increased abundance of phosphatidylcholines (PCs) and ether-linked PCs, and proteome changes associated with cytoskeletal organization and antioxidant processes. Furthermore, we performed a pharmacological rescue with Ferrostatin-1, which indicated that atRAL-induced cytotoxicity in RDH12-expressing cells is dependent, in part, on lipid radical propagation and lipid peroxidation, supporting ferroptosis-regulated cell death. Collectively, the multiomics and biological findings establish a novel temporal sequence for retinaldehyde toxicity, showing that RDH12 is a critical component of upstream metabolic defense for acute aldehyde clearance, late-stage lipid detoxification, and adaptive membrane lipid homeostasis.\n                <\/jats:p>","DOI":"10.64898\/2026.09.24.754135","type":"posted-content","created":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T16:05:36Z","timestamp":1790611536000},"source":"Crossref","is-referenced-by-count":0,"title":["Multiomic Profiling Links RDH12-Dependent Retinaldehyde Detoxification to Membrane Remodeling and Ferroptosis"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-8636-2616","authenticated-orcid":false,"given":"KanagaVijayan","family":"Dhanabalan","sequence":"first","affiliation":[{"name":"Department of Biological Sciences, Purdue University, West Lafayette, IN, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3566-3384","authenticated-orcid":false,"given":"Miranda R.","family":"Muhoberac","sequence":"additional","affiliation":[{"name":"The James Tarpo Jr. and Margaret Tarpo Department of Chemistry, Purdue University, West Lafayette, IN, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6419-3206","authenticated-orcid":false,"given":"Ramya","family":"Visvanathan","sequence":"additional","affiliation":[{"name":"Proteomics Facility, Bindley Bioscience Center, West Lafayette, IN, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5581-0252","authenticated-orcid":false,"given":"Devender","family":"Arora","sequence":"additional","affiliation":[{"name":"Proteomics Facility, Bindley Bioscience Center, West Lafayette, IN, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2512-925X","authenticated-orcid":false,"given":"Olga","family":"Belyaeva","sequence":"additional","affiliation":[{"name":"Department of Biochemistry and Molecular Genetics, University of Alabama at Birmingham, Birmingham, AL, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6917-4891","authenticated-orcid":false,"given":"Natalia Y.","family":"Kedishvili","sequence":"additional","affiliation":[{"name":"Department of Biochemistry and Molecular Genetics, University of Alabama at Birmingham, Birmingham, AL, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6709-190X","authenticated-orcid":false,"given":"Ramaswamy","family":"Subramanian","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Purdue University, West Lafayette, IN, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5631-512X","authenticated-orcid":false,"given":"Bryon S.","family":"Drown","sequence":"additional","affiliation":[{"name":"The James Tarpo Jr. and Margaret Tarpo Department of Chemistry, Purdue University, West Lafayette, IN, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026100109150772000_2026.09.24.754135v1.1","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1186\/s13024-022-00528-w","article-title":"Cellular stress signaling and the unfolded protein response in retinal degeneration: mechanisms and therapeutic implications","volume":"17","year":"2022","journal-title":"Mol Neurodegeneration"},{"key":"2026100109150772000_2026.09.24.754135v1.2","doi-asserted-by":"publisher","DOI":"10.1021\/cr400107q"},{"key":"2026100109150772000_2026.09.24.754135v1.3","doi-asserted-by":"publisher","DOI":"10.1016\/j.exer.2010.08.013"},{"key":"2026100109150772000_2026.09.24.754135v1.4","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M111.315432"},{"key":"2026100109150772000_2026.09.24.754135v1.5","doi-asserted-by":"publisher","DOI":"10.1016\/j.freeradbiomed.2009.08.005"},{"key":"2026100109150772000_2026.09.24.754135v1.6","doi-asserted-by":"publisher","DOI":"10.1093\/toxsci\/kfu223"},{"key":"2026100109150772000_2026.09.24.754135v1.7","doi-asserted-by":"publisher","DOI":"10.1016\/j.toxlet.2020.01.005"},{"key":"2026100109150772000_2026.09.24.754135v1.8","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M804505200"},{"key":"2026100109150772000_2026.09.24.754135v1.9","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.RA120.015779"},{"key":"2026100109150772000_2026.09.24.754135v1.10","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1400780111"},{"key":"2026100109150772000_2026.09.24.754135v1.11","doi-asserted-by":"crossref","first-page":"100042","DOI":"10.1194\/jlr.TR120000742","article-title":"Bisretinoid phospholipid and vitamin A aldehyde: shining a light","volume":"62","year":"2021","journal-title":"Journal of Lipid Research"},{"key":"2026100109150772000_2026.09.24.754135v1.12","doi-asserted-by":"publisher","DOI":"10.1562\/2004-11-13-IR-371"},{"key":"2026100109150772000_2026.09.24.754135v1.13","doi-asserted-by":"crossref","first-page":"108440","DOI":"10.1016\/j.pharmthera.2023.108440","article-title":"The essential role of docosahexaenoic acid and its derivatives for retinal integrity","volume":"247","year":"2023","journal-title":"Pharmacology & Therapeutics"},{"key":"2026100109150772000_2026.09.24.754135v1.14","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0802607105"},{"key":"2026100109150772000_2026.09.24.754135v1.15","doi-asserted-by":"publisher","DOI":"10.1016\/j.preteyeres.2004.06.002"},{"key":"2026100109150772000_2026.09.24.754135v1.16","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1016\/j.exer.2018.09.012","article-title":"Light-induced generation and toxicity of docosahexaenoate-derived oxidation products in retinal pigmented epithelial cells","volume":"181","year":"2019","journal-title":"Experimental Eye Research"},{"key":"2026100109150772000_2026.09.24.754135v1.17","doi-asserted-by":"crossref","first-page":"104686","DOI":"10.1016\/j.jbc.2023.104686","article-title":"eIF2\u03b1 incites photoreceptor cell and retina damage by all-trans-retinal","volume":"299","year":"2023","journal-title":"Journal of Biological Chemistry"},{"key":"2026100109150772000_2026.09.24.754135v1.18","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1016\/j.freeradbiomed.2024.12.007","article-title":"Inhibition of JNK signaling attenuates photoreceptor ferroptosis caused by all-trans-retinal","volume":"227","year":"2025","journal-title":"Free Radical Biology and Medicine"},{"key":"2026100109150772000_2026.09.24.754135v1.19","doi-asserted-by":"crossref","first-page":"7029","DOI":"10.7150\/ijbs.114187","article-title":"Activation of GSDME by all-trans-retinal increases sensitivity to photoreceptor ferroptosis","volume":"21","year":"2025","journal-title":"International Journal of Biological Sciences"},{"key":"2026100109150772000_2026.09.24.754135v1.20","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M608375200"},{"key":"2026100109150772000_2026.09.24.754135v1.21","doi-asserted-by":"publisher","DOI":"10.1016\/j.exer.2019.107793"},{"key":"2026100109150772000_2026.09.24.754135v1.22","doi-asserted-by":"crossref","first-page":"3342","DOI":"10.1016\/j.cub.2024.06.031","article-title":"RDH12 allows cone photoreceptors to regenerate opsin visual pigments from a chromophore precursor to escape competition with rods","volume":"34","year":"2024","journal-title":"Current Biology"},{"key":"2026100109150772000_2026.09.24.754135v1.23","doi-asserted-by":"crossref","unstructured":"Fahim, A. T. , Bouzia, Z. , Branham, K. H. , Kumaran, N. , Vargas, M. E. , Feathers, K. L. , Perera, N. D. , Young, K. , Khan, N. W. , Heckenlively, J. R. , Webster, A. R. , Pennesi, M. E. , Ali, R. R. , Thompson, D. A. , and Michaelides, M . (2019) Detailed clinical characterisation, unique features and natural history of autosomal recessive RDH12 -associated retinal degeneration. Br J Ophthalmol, bjophthalmol-2018\u2013313580","DOI":"10.1136\/bjophthalmol-2018-313580"},{"key":"2026100109150772000_2026.09.24.754135v1.24","doi-asserted-by":"publisher","DOI":"10.1080\/13816810.2021.1966053"},{"key":"2026100109150772000_2026.09.24.754135v1.25","first-page":"4293","article-title":"Genetics and phenotypes of RPE65 mutations in inherited retinal degeneration","volume":"41","year":"2000","journal-title":"Invest Ophthalmol Vis Sci"},{"key":"2026100109150772000_2026.09.24.754135v1.26","doi-asserted-by":"publisher","DOI":"10.1038\/ng1394"},{"key":"2026100109150772000_2026.09.24.754135v1.27","doi-asserted-by":"publisher","DOI":"10.1167\/iovs.06-0599"},{"key":"2026100109150772000_2026.09.24.754135v1.28","doi-asserted-by":"publisher","DOI":"10.1001\/archopht.126.9.1301"},{"key":"2026100109150772000_2026.09.24.754135v1.29","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M109.020966"},{"key":"2026100109150772000_2026.09.24.754135v1.30","doi-asserted-by":"publisher","DOI":"10.1111\/aos.12146"},{"key":"2026100109150772000_2026.09.24.754135v1.31","doi-asserted-by":"publisher","DOI":"10.1111\/jcmm.16569"},{"key":"2026100109150772000_2026.09.24.754135v1.32","doi-asserted-by":"crossref","first-page":"183","DOI":"10.3390\/metabo11030183","article-title":"Association of Human Plasma Metabolomics with Delayed Dark Adaptation in Age-Related Macular Degeneration","volume":"11","year":"2021","journal-title":"Metabolites"},{"key":"2026100109150772000_2026.09.24.754135v1.33","doi-asserted-by":"crossref","first-page":"2979","DOI":"10.1021\/acs.jproteome.2c00506","article-title":"Metabolic Characterization of Ocular Tissues in Relation to Laser-Induced Choroidal Neovascularization in Rats","volume":"21","year":"2022","journal-title":"J. Proteome Res"},{"key":"2026100109150772000_2026.09.24.754135v1.34","doi-asserted-by":"publisher","DOI":"10.1016\/S0092-8674(00)80602-9"},{"key":"2026100109150772000_2026.09.24.754135v1.35","doi-asserted-by":"publisher","DOI":"10.1007\/s00204-019-02409-6"},{"key":"2026100109150772000_2026.09.24.754135v1.36","unstructured":"36. Sparrow, J. R. , Kim, S. R. , and Wu, Y. (2010) in Retinoids: Methods and Protocols, eds Sun H , Travis GH (Humana Press, Totowa, NJ), pp 315\u2013327."},{"key":"2026100109150772000_2026.09.24.754135v1.37","doi-asserted-by":"publisher","DOI":"10.1167\/iovs.15-18551"},{"key":"2026100109150772000_2026.09.24.754135v1.38","doi-asserted-by":"crossref","first-page":"110280","DOI":"10.1016\/j.exer.2025.110280","article-title":"Involvement of STAT3 activation in ameliorating all-trans-retinal-induced ferroptosis in photoreceptor-derived 661W cells in vitro","volume":"253","year":"2025","journal-title":"Experimental Eye Research"},{"key":"2026100109150772000_2026.09.24.754135v1.39","doi-asserted-by":"publisher","DOI":"10.1021\/bi050226k"},{"key":"2026100109150772000_2026.09.24.754135v1.40","doi-asserted-by":"crossref","first-page":"1366","DOI":"10.3390\/ijms27031366","article-title":"Functional In Vitro Assessment of rAAV-Delivered Retinol Dehydrogenase 12 (RDH12) Activity","volume":"27","year":"2026","journal-title":"International Journal of Molecular Sciences"},{"key":"2026100109150772000_2026.09.24.754135v1.41","doi-asserted-by":"publisher","DOI":"10.1038\/nprot.2017.147"},{"key":"2026100109150772000_2026.09.24.754135v1.42","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkv007"},{"key":"2026100109150772000_2026.09.24.754135v1.43","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M800019200"},{"key":"2026100109150772000_2026.09.24.754135v1.44","doi-asserted-by":"crossref","first-page":"18343","DOI":"10.1021\/acs.analchem.4c02835","article-title":"The Addition of Transcriptomics to the Bead-Enabled Accelerated Monophasic Multi-Omics Method: A Step toward Universal Sample Preparation","volume":"96","year":"2024","journal-title":"Anal. Chem"},{"key":"2026100109150772000_2026.09.24.754135v1.45","doi-asserted-by":"crossref","unstructured":"Muehlbauer, L. K. , Jen, A. , Zhu, Y. , He, Y. , Shishkova, E. , Overmyer, K. A. , and Coon, J. J . (2023) Rapid Multi-Omics Sample Preparation for Mass Spectrometry. Anal. Chem., acs.analchem.2c02042","DOI":"10.1021\/acs.analchem.2c02042"},{"issue":"3f","key":"2026100109150772000_2026.09.24.754135v1.46","first-page":"c1f","article-title":"Step-by-Step Approach to Build Multiple Reaction Monitoring (MRM) Profiling Instrument Acquisition Methods for Class-based Lipid Exploratory Analysis by Mass Spectrometry","volume":"34","year":"2023","journal-title":"J Biomol Tech"},{"key":"2026100109150772000_2026.09.24.754135v1.47","doi-asserted-by":"publisher","DOI":"10.1038\/s41592-019-0638-x"},{"key":"2026100109150772000_2026.09.24.754135v1.48","doi-asserted-by":"publisher","DOI":"10.1002\/pmic.202200092"},{"key":"2026100109150772000_2026.09.24.754135v1.49","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M112.351429"},{"key":"2026100109150772000_2026.09.24.754135v1.50","doi-asserted-by":"crossref","first-page":"EMBJ2021109272","DOI":"10.15252\/embj.2021109272","article-title":"ZDHHC18 negatively regulates cGAS-mediated innate immunity through palmitoylation","volume":"41","year":"2022","journal-title":"EMBO J"},{"key":"2026100109150772000_2026.09.24.754135v1.51","doi-asserted-by":"publisher","DOI":"10.1016\/S0014-5793(02)03588-3"},{"key":"2026100109150772000_2026.09.24.754135v1.52","doi-asserted-by":"crossref","first-page":"524","DOI":"10.1016\/j.cmet.2020.08.002","article-title":"The Role of HIF in Immunity and Inflammation","volume":"32","year":"2020","journal-title":"Cell Metabolism"},{"key":"2026100109150772000_2026.09.24.754135v1.53","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbrc.2009.11.094"},{"key":"2026100109150772000_2026.09.24.754135v1.54","doi-asserted-by":"crossref","first-page":"926","DOI":"10.1038\/s41580-025-00855-y","article-title":"N-glycan-dependent protein maturation and quality control in the ER","volume":"26","year":"2025","journal-title":"Nat Rev Mol Cell Biol"},{"key":"2026100109150772000_2026.09.24.754135v1.55","doi-asserted-by":"crossref","first-page":"571","DOI":"10.1007\/s00424-003-1141-9","article-title":"SLC11 family of H+-coupled metal-ion transporters NRAMP1 and DMT1","volume":"447","year":"2004","journal-title":"Pfl\u00fcgers Archiv European Journal of Physiology"},{"key":"2026100109150772000_2026.09.24.754135v1.56","doi-asserted-by":"publisher","DOI":"10.1093\/jn\/nxx041"},{"key":"2026100109150772000_2026.09.24.754135v1.57","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-39401-9"},{"key":"2026100109150772000_2026.09.24.754135v1.58","doi-asserted-by":"publisher","DOI":"10.1007\/s13238-020-00789-5"},{"key":"2026100109150772000_2026.09.24.754135v1.59","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1038\/s41401-024-01359-9","article-title":"The correlation between mitochondria-associated endoplasmic reticulum membranes (MAMs) and Ca2+ transport in the pathogenesis of diseases","volume":"46","year":"2025","journal-title":"Acta Pharmacol Sin"},{"key":"2026100109150772000_2026.09.24.754135v1.60","doi-asserted-by":"crossref","first-page":"110110","DOI":"10.1016\/j.exer.2024.110110","article-title":"Current insights on mitochondria-associated endoplasmic reticulum membranes (MAMs) and their significance in the pathophysiology of ocular disorders","volume":"248","year":"2024","journal-title":"Experimental Eye Research"},{"key":"2026100109150772000_2026.09.24.754135v1.61","doi-asserted-by":"publisher","DOI":"10.1016\/S1471-4914(01)02103-7"},{"key":"2026100109150772000_2026.09.24.754135v1.62","doi-asserted-by":"publisher","DOI":"10.1016\/j.sbi.2025.102999"},{"key":"2026100109150772000_2026.09.24.754135v1.63","doi-asserted-by":"publisher","DOI":"10.1007\/s12551-018-0400-0"},{"key":"2026100109150772000_2026.09.24.754135v1.64","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M306053200"},{"key":"2026100109150772000_2026.09.24.754135v1.65","doi-asserted-by":"publisher","DOI":"10.1186\/s12915-023-01695-y"},{"key":"2026100109150772000_2026.09.24.754135v1.66","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M113.476358"},{"key":"2026100109150772000_2026.09.24.754135v1.67","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2019.03.012"},{"key":"2026100109150772000_2026.09.24.754135v1.68","doi-asserted-by":"publisher","DOI":"10.1021\/acsptsci.5c00482"},{"key":"2026100109150772000_2026.09.24.754135v1.69","doi-asserted-by":"crossref","first-page":"1149","DOI":"10.3390\/ijms20051149","article-title":"An Updated Review of Lysophosphatidylcholine Metabolism in Human Diseases","volume":"20","year":"2019","journal-title":"IJMS"},{"key":"2026100109150772000_2026.09.24.754135v1.70","doi-asserted-by":"crossref","first-page":"2221","DOI":"10.1038\/s41467-025-57402-8","article-title":"PNPLA6 regulates retinal homeostasis by choline through phospholipid turnover","volume":"16","year":"2025","journal-title":"Nat Commun"},{"key":"2026100109150772000_2026.09.24.754135v1.71","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms6614"},{"key":"2026100109150772000_2026.09.24.754135v1.72","doi-asserted-by":"publisher","DOI":"10.1021\/cr1004106"},{"key":"2026100109150772000_2026.09.24.754135v1.73","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.R117.000582"},{"key":"2026100109150772000_2026.09.24.754135v1.74","doi-asserted-by":"publisher","DOI":"10.1016\/j.tibs.2025.01.007"},{"key":"2026100109150772000_2026.09.24.754135v1.75","doi-asserted-by":"publisher","DOI":"10.4161\/auto.5.4.8390"},{"key":"2026100109150772000_2026.09.24.754135v1.76","doi-asserted-by":"crossref","first-page":"26","DOI":"10.1038\/s41419-020-03297-w","article-title":"Acid sphingomyelinase-dependent autophagic degradation of GPX4 is critical for the execution of ferroptosis","volume":"12","year":"2021","journal-title":"Cell Death Dis"},{"key":"2026100109150772000_2026.09.24.754135v1.77","doi-asserted-by":"crossref","unstructured":"Njie-Mbye, Y. F. , Kulkarni-Chitnis, M. , Opere, C. A. , Barrett, A. , and Ohia, S. E . (2013) Lipid peroxidation: pathophysiological and pharmacological implications in the eye. Front. Physiol. 4,","DOI":"10.3389\/fphys.2013.00366"},{"key":"2026100109150772000_2026.09.24.754135v1.78","doi-asserted-by":"publisher","DOI":"10.1089\/ars.2009.2597"},{"key":"2026100109150772000_2026.09.24.754135v1.79","first-page":"E1195","article-title":"Biochemistry, physiology, and genetics of GPAT","volume":"296","year":"2009","journal-title":"AGPAT, and lipin enzymes in triglyceride synthesis. American Journal of Physiology-Endocrinology And Metabolism"},{"key":"2026100109150772000_2026.09.24.754135v1.80","first-page":"543","article-title":"Formation and function of phosphatidylserine and phosphatidylethanolamine in mammalian cells","volume":"1831","year":"2013","journal-title":"Biochimica et Biophysica Acta (BBA)-Molecular and Cell Biology of Lipids"},{"key":"2026100109150772000_2026.09.24.754135v1.81","doi-asserted-by":"publisher","DOI":"10.1002\/iub.337"},{"key":"2026100109150772000_2026.09.24.754135v1.82","doi-asserted-by":"publisher","DOI":"10.1194\/jlr.R800035-JLR200"},{"key":"2026100109150772000_2026.09.24.754135v1.83","doi-asserted-by":"crossref","first-page":"198","DOI":"10.1038\/s41419-021-03482-5","article-title":"Zinc transporter ZIP7 is a novel determinant of ferroptosis","volume":"12","year":"2021","journal-title":"Cell death & disease"},{"key":"2026100109150772000_2026.09.24.754135v1.84","doi-asserted-by":"crossref","first-page":"101107","DOI":"10.1016\/j.redox.2019.101107","article-title":"NRF2 plays a critical role in mitigating lipid peroxidation and ferroptosis","volume":"23","year":"2019","journal-title":"Redox biology"},{"key":"2026100109150772000_2026.09.24.754135v1.85","doi-asserted-by":"crossref","first-page":"jcs263834","DOI":"10.1242\/jcs.263834","article-title":"Fibronectin matrix assembly at a glance","volume":"138","year":"2025","journal-title":"Journal of cell science"},{"key":"2026100109150772000_2026.09.24.754135v1.86","doi-asserted-by":"publisher","DOI":"10.1038\/nrd3599"},{"key":"2026100109150772000_2026.09.24.754135v1.87","first-page":"321","article-title":"ATF3 and stress responses","volume":"7","year":"1999","journal-title":"Gene Expr"},{"key":"2026100109150772000_2026.09.24.754135v1.88","doi-asserted-by":"publisher","DOI":"10.1016\/j.tcb.2005.11.004"},{"key":"2026100109150772000_2026.09.24.754135v1.89","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1111\/j.1748-1716.2011.02294.x","article-title":"Wnt5a: its signalling, functions and implication in diseases","volume":"204","year":"2012","journal-title":"Acta physiologica"},{"key":"2026100109150772000_2026.09.24.754135v1.90","doi-asserted-by":"publisher","DOI":"10.1038\/nrc2780"},{"key":"2026100109150772000_2026.09.24.754135v1.91","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2012.03.042"},{"key":"2026100109150772000_2026.09.24.754135v1.92","doi-asserted-by":"publisher","DOI":"10.1038\/s41580-020-00324-8"},{"key":"2026100109150772000_2026.09.24.754135v1.93","doi-asserted-by":"publisher","DOI":"10.1038\/nature13148"},{"key":"2026100109150772000_2026.09.24.754135v1.94","doi-asserted-by":"publisher","DOI":"10.1016\/j.mam.2008.05.005"},{"key":"2026100109150772000_2026.09.24.754135v1.95","doi-asserted-by":"crossref","first-page":"2248","DOI":"10.3389\/fmicb.2019.02248","article-title":"From cyanobacteria to human, MAPEG-type glutathione-S-transferases operate in cell tolerance to heat, cold, and lipid peroxidation","volume":"10","year":"2019","journal-title":"Frontiers in Microbiology"},{"key":"2026100109150772000_2026.09.24.754135v1.96","doi-asserted-by":"publisher","DOI":"10.1007\/s00018-016-2223-0"},{"key":"2026100109150772000_2026.09.24.754135v1.97","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0606424103"},{"key":"2026100109150772000_2026.09.24.754135v1.98","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M910191199"},{"key":"2026100109150772000_2026.09.24.754135v1.99","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1038\/nchembio.2239","article-title":"ACSL4 dictates ferroptosis sensitivity by shaping cellular lipid composition","volume":"13","year":"2017","journal-title":"Nature chemical biology"},{"key":"2026100109150772000_2026.09.24.754135v1.100","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M600225200"},{"key":"2026100109150772000_2026.09.24.754135v1.101","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1016\/j.chembiol.2018.11.016","article-title":"Exogenous monounsaturated fatty acids promote a ferroptosis-resistant cell state","volume":"26","year":"2019","journal-title":"Cell chemical biology"},{"key":"2026100109150772000_2026.09.24.754135v1.102","doi-asserted-by":"publisher","DOI":"10.1038\/nrm.2017.107"},{"key":"2026100109150772000_2026.09.24.754135v1.103","first-page":"1442","article-title":"Functions of plasmalogen lipids in health and disease","volume":"1822","year":"2012","journal-title":"Biochimica et Biophysica Acta (BBA)-Molecular Basis of Disease"},{"key":"2026100109150772000_2026.09.24.754135v1.104","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-020-2732-8"},{"key":"2026100109150772000_2026.09.24.754135v1.105","doi-asserted-by":"publisher","DOI":"10.1021\/tx5001046"},{"key":"2026100109150772000_2026.09.24.754135v1.106","doi-asserted-by":"crossref","first-page":"1103","DOI":"10.1186\/s12967-025-07195-7","article-title":"Ferrostatin-1, a ferroptosis inhibitor, mitigates all-trans-retinal-induced retinal pigment epithelium degeneration in mice","volume":"23","year":"2025","journal-title":"Journal of translational medicine"}],"container-title":[],"original-title":[],"link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.64898\/2026.09.24.754135","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T16:20:40Z","timestamp":1790871640000},"score":1,"resource":{"primary":{"URL":"http:\/\/biorxiv.org\/lookup\/doi\/10.64898\/2026.09.24.754135"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,28]]},"references-count":106,"URL":"https:\/\/doi.org\/10.64898\/2026.09.24.754135","relation":{},"subject":[],"published":{"date-parts":[[2026,9,28]]},"subtype":"preprint"}}