{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T21:42:35Z","timestamp":1790804555651,"version":"4.1.0"},"reference-count":55,"publisher":"MDPI AG","issue":"2","license":[{"start":{"date-parts":[[2026,1,8]],"date-time":"2026-01-08T00:00:00Z","timestamp":1767830400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100004837","name":"Ministerio de Ciencia, Innovaci\u00f3n y Universidades","doi-asserted-by":"publisher","award":["PID2020-113921RB-I00; PID2024-155549OB-I00"],"award-info":[{"award-number":["PID2020-113921RB-I00; PID2024-155549OB-I00"]}],"id":[{"id":"10.13039\/501100004837","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Herpes simplex virus type 1 (HSV-1) has been proposed as an environmental risk factor for Alzheimer\u2019s disease (AD). Viral infection of neuronal cells can reproduce hallmark pathological features of AD, including intracellular beta-amyloid (A\u03b2) accumulation, tau hyperphosphorylation, and lysosomal dysfunction. However, the molecular mechanisms underlying these alterations remain unclear, partly due to limitations of existing experimental models. Here, we established both two-dimensional (2D) and three-dimensional (3D) LUHMES neuronal cultures\u2014a human mesencephalic-derived neural cell line that differentiates rapidly into mature neurons\u2014to investigate HSV-1-induced AD-associated markers. Our results demonstrate that HSV-1 infection induces key features of AD, including intracellular accumulation of A\u03b2 peptides and hyperphosphorylation of tau protein. Moreover, we observed disruptions in the autophagy\u2013lysosome pathway, characterized by increased LC3-II levels, reduced cathepsin activity, and impaired lysosomal burden. Notably, these AD-like alterations were reproduced in 3D LUHMES neuronal aggregates, confirming their susceptibility to productive HSV-1 infection. Collectively, these findings indicate that HSV-1 not only triggers AD-like neuropathological markers but also disrupts cellular clearance mechanisms that may contribute to neuronal dysfunction and degeneration. This study validates the 3D LUHMES system as a useful human neuronal model to study virus-induced neurodegeneration and its mechanistic links to AD pathology.<\/jats:p>","DOI":"10.3390\/ijms27020642","type":"journal-article","created":{"date-parts":[[2026,1,8]],"date-time":"2026-01-08T14:32:38Z","timestamp":1767882758000},"page":"642","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Impact of HSV-1 Infection on Alzheimer\u2019s Disease Neurodegeneration Markers: Insights from LUHMES 2D and 3D Neuronal Models"],"prefix":"10.3390","volume":"27","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9513-9619","authenticated-orcid":false,"given":"Mar\u00eda","family":"Mart\u00edn-Rico","sequence":"first","affiliation":[{"name":"Centro de Biologia Molecular Severo Ochoa (CBM), CSIC\u2014Universidad Autonoma de Madrid, 28049 Madrid, Spain"},{"name":"CIBER de Enfermedades Neurodegenerativas, Instituto de Salud Carlos III, 28031 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Blanca","family":"Salgado","sequence":"additional","affiliation":[{"name":"Centro de Biologia Molecular Severo Ochoa (CBM), CSIC\u2014Universidad Autonoma de Madrid, 28049 Madrid, Spain"},{"name":"CIBER de Enfermedades Neurodegenerativas, Instituto de Salud Carlos III, 28031 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"In\u00e9s","family":"Beamonte","sequence":"additional","affiliation":[{"name":"Centro de Biologia Molecular Severo Ochoa (CBM), CSIC\u2014Universidad Autonoma de Madrid, 28049 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Isabel","family":"Sastre","sequence":"additional","affiliation":[{"name":"Centro de Biologia Molecular Severo Ochoa (CBM), CSIC\u2014Universidad Autonoma de Madrid, 28049 Madrid, Spain"},{"name":"CIBER de Enfermedades Neurodegenerativas, Instituto de Salud Carlos III, 28031 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6477-1117","authenticated-orcid":false,"given":"Mar\u00eda J.","family":"Bullido","sequence":"additional","affiliation":[{"name":"Centro de Biologia Molecular Severo Ochoa (CBM), CSIC\u2014Universidad Autonoma de Madrid, 28049 Madrid, Spain"},{"name":"CIBER de Enfermedades Neurodegenerativas, Instituto de Salud Carlos III, 28031 Madrid, Spain"},{"name":"Instituto de Investigacion Sanitaria del Hospital Universitario La Paz\u2014IdiPAZ, 28046 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7098-4709","authenticated-orcid":false,"given":"Jes\u00fas","family":"Aldudo","sequence":"additional","affiliation":[{"name":"Centro de Biologia Molecular Severo Ochoa (CBM), CSIC\u2014Universidad Autonoma de Madrid, 28049 Madrid, Spain"},{"name":"CIBER de Enfermedades Neurodegenerativas, Instituto de Salud Carlos III, 28031 Madrid, Spain"},{"name":"Instituto de Investigacion Sanitaria del Hospital Universitario La Paz\u2014IdiPAZ, 28046 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,1,8]]},"reference":[{"key":"ref_1","unstructured":"(2025, December 04). Dementia. Available online: https:\/\/www.who.int\/news-room\/fact-sheets\/detail\/dementia."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Breijyeh, Z., and Karaman, R. (2020). Comprehensive Review on Alzheimer\u2019s Disease: Causes and Treatment. Molecules, 25.","DOI":"10.3390\/molecules25245789"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"312","DOI":"10.1016\/j.cell.2019.09.001","article-title":"Alzheimer Disease: An Update on Pathobiology and Treatment Strategies","volume":"179","author":"Long","year":"2019","journal-title":"Cell"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"794","DOI":"10.1038\/nn.4017","article-title":"The Case for Rejecting the Amyloid Cascade Hypothesis","volume":"18","author":"Herrup","year":"2015","journal-title":"Nat. Neurosci."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"102615","DOI":"10.1016\/j.arr.2024.102615","article-title":"An Overview on the Impact of Viral Pathogens on Alzheimer\u2019s Disease","volume":"104","author":"AmeliMojarad","year":"2024","journal-title":"Ageing Res. Rev."},{"key":"ref_6","unstructured":"(2025, December 04). Herpes Simplex Virus. Available online: https:\/\/www.who.int\/news-room\/fact-sheets\/detail\/herpes-simplex-virus."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"808","DOI":"10.1016\/j.tim.2020.03.003","article-title":"Herpes Simplex Virus-1 in the Brain: The Dark Side of a Sneaky Infection","volume":"28","author":"Marcocci","year":"2020","journal-title":"Trends Microbiol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"e2550","DOI":"10.1002\/rmv.2550","article-title":"Alzheimer\u2019s Disease and Herpes Viruses: Current Events and Perspectives","volume":"34","author":"Pinchera","year":"2024","journal-title":"Rev. Med. Virol."},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Duarte, L.F., Far\u00edas, M.A., \u00c1lvarez, D.M., Bueno, S.M., Riedel, C.A., and Gonz\u00e1lez, P.A. (2019). Herpes Simplex Virus Type 1 Infection of the Central Nervous System: Insights Into Proposed Interrelationships with Neurodegenerative Disorders. Front. Cell Neurosci., 13.","DOI":"10.3389\/fncel.2019.00046"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"5850","DOI":"10.1007\/s12035-024-04535-4","article-title":"Unveiling the Involvement of Herpes Simplex Virus-1 in Alzheimer\u2019s Disease: Possible Mechanisms and Therapeutic Implications","volume":"62","author":"Chauhan","year":"2024","journal-title":"Mol. Neurobiol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1016\/S0140-6736(96)10149-5","article-title":"Herpes Simplex Virus Type 1 in Brain and Risk of Alzheimer\u2019s Disease","volume":"349","author":"Itzhaki","year":"1997","journal-title":"Lancet"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1007\/s13311-018-0611-x","article-title":"Anti-Herpetic Medications and Reduced Risk of Dementia in Patients with Herpes Simplex Virus Infections-a Nationwide, Population-Based Cohort Study in Taiwan","volume":"15","author":"Tzeng","year":"2018","journal-title":"Neurotherapeutics"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1841","DOI":"10.3233\/JAD-230718","article-title":"Herpes Simplex Viral Infection Doubles the Risk of Dementia in a Contemporary Cohort of Older Adults: A Prospective Study","volume":"97","author":"Vestin","year":"2024","journal-title":"J. Alzheimers Dis."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"e093946","DOI":"10.1136\/bmjopen-2024-093946","article-title":"Association between Herpes Simplex Virus Type 1 and the Risk of Alzheimer\u2019s Disease: A Retrospective Case-Control Study","volume":"15","author":"Liu","year":"2025","journal-title":"BMJ Open"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1002\/path.2449","article-title":"Herpes Simplex Virus Type 1 DNA Is Located within Alzheimer\u2019s Disease Amyloid Plaques","volume":"217","author":"Wozniak","year":"2009","journal-title":"J. Pathol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"430.e19","DOI":"10.1016\/j.neurobiolaging.2010.12.010","article-title":"Herpes Simplex Virus Type I Induces the Accumulation of Intracellular \u03b2-Amyloid in Autophagic Compartments and the Inhibition of the Non-Amyloidogenic Pathway in Human Neuroblastoma Cells","volume":"33","author":"Santana","year":"2012","journal-title":"Neurobiol. Aging"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1020","DOI":"10.1002\/jnr.23003","article-title":"Herpes Simplex Virus Type 1 Induces Nuclear Accumulation of Hyperphosphorylated Tau in Neuronal Cells","volume":"90","author":"Alvarez","year":"2012","journal-title":"J. Neurosci. Res."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"176","DOI":"10.1186\/s12974-024-03166-9","article-title":"Herpes Simplex Virus 1 Accelerates the Progression of Alzheimer\u2019s Disease by Modulating Microglial Phagocytosis and Activating NLRP3 Pathway","volume":"21","author":"Wang","year":"2024","journal-title":"J. Neuroinflammat."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Acu\u00f1a-Hinrichsen, F., Covarrubias-Pinto, A., Ishizuka, Y., Stolzenbach, M.F., Martin, C., Salazar, P., Castro, M.A., Bramham, C.R., and Otth, C. (2021). Herpes Simplex Virus Type 1 Neuronal Infection Triggers the Disassembly of Key Structural Components of Dendritic Spines. Front. Cell Neurosci., 15.","DOI":"10.3389\/fncel.2021.580717"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"926","DOI":"10.1038\/s41580-024-00757-5","article-title":"Mechanisms of Autophagy-Lysosome Dysfunction in Neurodegenerative Diseases","volume":"25","author":"Nixon","year":"2024","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1038\/s41579-018-0003-6","article-title":"Autophagy during Viral Infection\u2014A Double-Edged Sword","volume":"16","author":"Choi","year":"2018","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"619","DOI":"10.1038\/s41556-025-01623-y","article-title":"Proteostasis and Lysosomal Repair Deficits in Transdifferentiated Neurons of Alzheimer\u2019s Disease","volume":"27","author":"Chou","year":"2025","journal-title":"Nat. Cell Biol."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Ripa, I., Andreu, S., L\u00f3pez-Guerrero, J.A., and Bello-Morales, R. (2022). Interplay between Autophagy and Herpes Simplex Virus Type 1: ICP34.5, One of the Main Actors. Int. J. Mol. Sci., 23.","DOI":"10.20944\/preprints202209.0445.v1"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"957","DOI":"10.1111\/j.1471-4159.2011.07255.x","article-title":"Rapid, Complete and Large-Scale Generation of Post-Mitotic Neurons from the Human LUHMES Cell Line","volume":"119","author":"Scholz","year":"2011","journal-title":"J. Neurochem."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"2725","DOI":"10.1007\/s00204-015-1637-z","article-title":"A LUHMES 3D Dopaminergic Neuronal Model for Neurotoxicity Testing Allowing Long-Term Exposure and Cellular Resilience Analysis","volume":"90","author":"Smirnova","year":"2016","journal-title":"Arch. Toxicol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"11.23.1","DOI":"10.1002\/cptx.29","article-title":"3D Differentiation of LUHMES Cell Line to Study Recovery and Delayed Neurotoxic Effects","volume":"73","author":"Harris","year":"2017","journal-title":"Curr. Protoc. Toxicol."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Beliakov, S.V., Blokhin, V., Surkov, S.A., and Ugrumov, M.V. (2023). LUHMES Cells: Phenotype Refinement and Development of an MPP+-Based Test System for Screening Antiparkinsonian Drugs. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24010733"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"945","DOI":"10.1038\/aps.2014.36","article-title":"Cell-Based Assays for Parkinson\u2019s Disease Using Differentiated Human LUHMES Cells","volume":"35","author":"Zhang","year":"2014","journal-title":"Acta Pharmacol. Sin."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"100143","DOI":"10.1016\/j.slasd.2024.01.004","article-title":"3D-Suspension Culture Platform for High Throughput Screening of Neurotoxic Chemicals Using LUHMES Dopaminergic Neurons","volume":"29","author":"Tong","year":"2024","journal-title":"SLAS Discov."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Whisnant, A.W., Clark, S.E., Aguilar-Brise\u00f1o, J.A., Durnell, L.A., Grothey, A., Miller, A.M., Varga, S.M., Meier, J.L., Grose, C., and Sinn, P.L. (2025). A More Rapid Method for Culturing LUHMES-Derived Neurons Provides Greater Cell Numbers and Facilitates Studies of Multiple Viruses. Viruses, 17.","DOI":"10.3390\/v17071001"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"e02210-18","DOI":"10.1128\/JVI.02210-18","article-title":"Lund Human Mesencephalic (LUHMES) Neuronal Cell Line Supports Herpes Simplex Virus 1 Latency In Vitro","volume":"93","author":"Edwards","year":"2019","journal-title":"J. Virol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"e0143823","DOI":"10.1128\/jvi.01438-23","article-title":"A Viral lncRNA Tethers HSV-1 Genomes at the Nuclear Periphery to Establish Viral Latency","volume":"97","author":"Grams","year":"2023","journal-title":"J. Virol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1016\/j.neurobiolaging.2012.03.012","article-title":"Control of A\u03b2 Release from Human Neurons by Differentiation Status and RET Signaling","volume":"34","author":"Scholz","year":"2013","journal-title":"Neurobiol. Aging"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"256","DOI":"10.1111\/jnc.14467","article-title":"Reduced A\u03b2 Secretion by Human Neurons under Conditions of Strongly Increased BACE Activity","volume":"147","author":"Scholz","year":"2018","journal-title":"J. Neurochem."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"815","DOI":"10.3233\/JAD-2012-112000","article-title":"Herpes Simplex Virus Type I Induces an Incomplete Autophagic Response in Human Neuroblastoma Cells","volume":"30","author":"Santana","year":"2012","journal-title":"J. Alzheimers Dis."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"414","DOI":"10.1016\/j.phrs.2018.01.016","article-title":"The Role of Dopaminergic Midbrain in Alzheimer\u2019s Disease: Translating Basic Science into Clinical Practice","volume":"130","author":"Mercuri","year":"2018","journal-title":"Pharmacol. Res."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"e00968-24","DOI":"10.1128\/jvi.00968-24","article-title":"Olfactory and Trigeminal Routes of HSV-1 CNS Infection with Regional Microglial Heterogeneity","volume":"98","author":"Niemeyer","year":"2024","journal-title":"J. Virol."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Salgado, B., Sastre, I., Bullido, M.J., and Aldudo, J. (2023). Herpes Simplex Virus Type 1 Induces AD-like Neurodegeneration Markers in Human Progenitor and Differentiated ReNcell VM Cells. Microorganisms, 11.","DOI":"10.3390\/microorganisms11051205"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"e02179-20","DOI":"10.1128\/JVI.02179-20","article-title":"Herpes Simplex Virus 1 MicroRNA miR-H8 Is Dispensable for Latency and Reactivation In Vivo","volume":"95","author":"Barrozo","year":"2021","journal-title":"J. Virol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"S4","DOI":"10.1186\/scrt365","article-title":"Toward a 3D Model of Human Brain Development for Studying Gene\/Environment Interactions","volume":"4","author":"Hogberg","year":"2013","journal-title":"Stem Cell Res. Ther."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1016\/j.neurobiolaging.2018.03.025","article-title":"The Lysosome System Is Severely Impaired in a Cellular Model of Neurodegeneration Induced by HSV-1 and Oxidative Stress","volume":"68","author":"Kristen","year":"2018","journal-title":"Neurobiol. Aging"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"224","DOI":"10.1002\/jmv.1890330403","article-title":"Latent Herpes Simplex Virus Type 1 in Normal and Alzheimer\u2019s Disease Brains","volume":"33","author":"Jamieson","year":"1991","journal-title":"J. Med. Virol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.neuron.2018.06.030","article-title":"Alzheimer\u2019s Disease-Associated \u03b2-Amyloid Is Rapidly Seeded by Herpesviridae to Protect against Brain Infection","volume":"99","author":"Eimer","year":"2018","journal-title":"Neuron"},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"D\u2019Mello, S.R. (2021). When Good Kinases Go Rogue: GSK3, P38 MAPK and CDKs as Therapeutic Targets for Alzheimer\u2019s and Huntington\u2019s Disease. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22115911"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"849","DOI":"10.3233\/JAD-131706","article-title":"Inflammatory and Neurodegeneration Markers during Asymptomatic HSV-1 Reactivation","volume":"39","author":"Martin","year":"2014","journal-title":"J. Alzheimers Dis."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"De Chiara, G., Piacentini, R., Fabiani, M., Mastrodonato, A., Marcocci, M.E., Limongi, D., Napoletani, G., Protto, V., Coluccio, P., and Celestino, I. (2019). Recurrent Herpes Simplex Virus-1 Infection Induces Hallmarks of Neurodegeneration and Cognitive Deficits in Mice. PLoS Pathog., 15.","DOI":"10.1371\/journal.ppat.1007617"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"eado6430","DOI":"10.1126\/scisignal.ado6430","article-title":"Repetitive Injury Induces Phenotypes Associated with Alzheimer\u2019s Disease by Reactivating HSV-1 in a Human Brain Tissue Model","volume":"18","author":"Cairns","year":"2025","journal-title":"Sci. Signal."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1252","DOI":"10.1038\/s41564-023-01405-y","article-title":"Modelling Viral Encephalitis Caused by Herpes Simplex Virus 1 Infection in Cerebral Organoids","volume":"8","author":"Wyler","year":"2023","journal-title":"Nat. Microbiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"e00111-19","DOI":"10.1128\/JVI.00111-19","article-title":"Modeling Herpes Simplex Virus 1 Infections in Human Central Nervous System Neuronal Cells Using Two- and Three-Dimensional Cultures Derived from Induced Pluripotent Stem Cells","volume":"93","author":"Bloom","year":"2019","journal-title":"J. Virol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"eaay8828","DOI":"10.1126\/sciadv.aay8828","article-title":"A 3D Human Brain-like Tissue Model of Herpes-Induced Alzheimer\u2019s Disease","volume":"6","author":"Cairns","year":"2020","journal-title":"Sci. Adv."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1189","DOI":"10.3233\/JAD-220287","article-title":"Potential Involvement of Varicella Zoster Virus in Alzheimer\u2019s Disease via Reactivation of Quiescent Herpes Simplex Virus Type 1","volume":"88","author":"Cairns","year":"2022","journal-title":"J. Alzheimers Dis."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"5220","DOI":"10.1128\/JVI.76.10.5220-5232.2002","article-title":"Rapid Directional Translocations in Virus Replication","volume":"76","author":"Willard","year":"2002","journal-title":"J. Virol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1016\/0166-0934(82)90034-9","article-title":"Production and Titration of African Swine Fever Virus in Porcine Alveolar Macrophages","volume":"3","author":"Carrascosa","year":"1982","journal-title":"J. Virol. Methods"},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Santana, S., Sastre, I., Recuero, M., Bullido, M.J., and Aldudo, J. (2013). Oxidative Stress Enhances Neurodegeneration Markers Induced by Herpes Simplex Virus Type 1 Infection in Human Neuroblastoma Cells. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0075842"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"581","DOI":"10.4161\/auto.23568","article-title":"Lysosomal Basification and Decreased Autophagic Flux in Oxidatively Stressed Trabecular Meshwork Cells: Implications for Glaucoma Pathogenesis","volume":"9","author":"Porter","year":"2013","journal-title":"Autophagy"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/2\/642\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,10]],"date-time":"2026-01-10T05:14:39Z","timestamp":1768022079000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/2\/642"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,8]]},"references-count":55,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2026,1]]}},"alternative-id":["ijms27020642"],"URL":"https:\/\/doi.org\/10.3390\/ijms27020642","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,8]]}}}