{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,29]],"date-time":"2026-05-29T19:02:44Z","timestamp":1780081364810,"version":"3.54.0"},"reference-count":54,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,2,23]],"date-time":"2026-02-23T00:00:00Z","timestamp":1771804800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,3,27]],"date-time":"2026-03-27T00:00:00Z","timestamp":1774569600000},"content-version":"vor","delay-in-days":32,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"name":"Brain Science and Brain-like Intelligence Technology-National Science and Technology Major Project","award":["2021ZD0200600"],"award-info":[{"award-number":["2021ZD0200600"]}]},{"name":"Brain Science and Brain-like Intelligence Technology-National Science and Technology Major Project","award":["2021ZD0202001"],"award-info":[{"award-number":["2021ZD0202001"]}]},{"name":"CAMS Innovation Fund for Medical Sciences","award":["2021-I2M-1-020"],"award-info":[{"award-number":["2021-I2M-1-020"]}]},{"name":"National High Level Hospital Clinical Research Funding","award":["2022-PUMCH-D-002"],"award-info":[{"award-number":["2022-PUMCH-D-002"]}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["J Neuroinflammation"],"DOI":"10.1186\/s12974-026-03751-0","type":"journal-article","created":{"date-parts":[[2026,2,23]],"date-time":"2026-02-23T14:45:45Z","timestamp":1771857945000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Spatial multiomics dissects the SARS-CoV-2-induced disruption of cell adhesion and immune dynamics in the human hippocampus"],"prefix":"10.1186","volume":"23","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7277-1619","authenticated-orcid":false,"given":"Hui","family":"Wei","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-1927-6595","authenticated-orcid":false,"given":"Zhongman","family":"Jin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Caiyun","family":"Zhu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pengsheng","family":"You","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiaping","family":"Sui","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lian","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nian","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wanwan","family":"Zhu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2660-8092","authenticated-orcid":false,"given":"Qi","family":"Xu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,2,23]]},"reference":[{"key":"3751_CR1","unstructured":"COVID-19 cases | WHO COVID-. 19 dashboard. datadot. https:\/\/data.who.int\/dashboards\/covid19\/cases Cited 2025 Sept 23. Accessed 23 Sept 2025."},{"key":"3751_CR2","doi-asserted-by":"publisher","first-page":"168","DOI":"10.1038\/s41593-020-00758-5","volume":"24","author":"J Meinhardt","year":"2021","unstructured":"Meinhardt J, Radke J, Dittmayer C, Franz J, Thomas C, Mothes R, et al. Olfactory transmucosal SARS-CoV-2 invasion as a port of central nervous system entry in individuals with COVID-19. Nat Neurosci. 2021;24:168\u201375. https:\/\/doi.org\/10.1038\/s41593-020-00758-5.","journal-title":"Nat Neurosci"},{"key":"3751_CR3","doi-asserted-by":"publisher","first-page":"758","DOI":"10.1038\/s41586-022-05542-y","volume":"612","author":"SR Stein","year":"2022","unstructured":"Stein SR, Ramelli SC, Grazioli A, Chung J-Y, Singh M, Yinda CK, et al. SARS-CoV-2 infection and persistence in the human body and brain at autopsy. Nature. 2022;612:758\u201363. https:\/\/doi.org\/10.1038\/s41586-022-05542-y.","journal-title":"Nature"},{"key":"3751_CR4","doi-asserted-by":"publisher","first-page":"416","DOI":"10.1016\/S2215-0366(21)00084-5","volume":"8","author":"M Taquet","year":"2021","unstructured":"Taquet M, Geddes JR, Husain M, Luciano S, Harrison PJ. 6-month neurological and psychiatric outcomes in 236 379 survivors of COVID-19: a retrospective cohort study using electronic health records. Lancet Psychiatry. 2021;8:416\u201327. https:\/\/doi.org\/10.1016\/S2215-0366(21)00084-5.","journal-title":"Lancet Psychiatry"},{"key":"3751_CR5","doi-asserted-by":"publisher","first-page":"e2112131","DOI":"10.1001\/jamanetworkopen.2021.12131","volume":"4","author":"SH-Y Chou","year":"2021","unstructured":"Chou SH-Y, Beghi E, Helbok R, Moro E, Sampson J, Altamirano V, et al. Global incidence of neurological manifestations among patients hospitalized with COVID-19\u2014a report for the GCS-NeuroCOVID consortium and the ENERGY consortium. JAMA Netw Open. 2021;4:e2112131. https:\/\/doi.org\/10.1001\/jamanetworkopen.2021.12131.","journal-title":"JAMA Netw Open"},{"key":"3751_CR6","doi-asserted-by":"publisher","first-page":"683","DOI":"10.1001\/jamaneurol.2020.1127","volume":"77","author":"L Mao","year":"2020","unstructured":"Mao L, Jin H, Wang M, Hu Y, Chen S, He Q, et al. Neurologic manifestations of hospitalized patients with coronavirus disease 2019 in Wuhan, china. JAMA Neurol. 2020;77:683\u201390. https:\/\/doi.org\/10.1001\/jamaneurol.2020.1127.","journal-title":"JAMA Neurol"},{"key":"3751_CR7","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1038\/s41591-024-03309-8","volume":"31","author":"GK Wood","year":"2025","unstructured":"Wood GK, Sargent BF, Ahmad Z-U-A, Tharmaratnam K, Dunai C, Egbe FN, et al. Posthospitalization COVID-19 cognitive deficits at 1 year are global and associated with elevated brain injury markers and gray matter volume reduction. Nat Med. 2025;31:245\u201357. https:\/\/doi.org\/10.1038\/s41591-024-03309-8.","journal-title":"Nat Med"},{"key":"3751_CR8","doi-asserted-by":"publisher","first-page":"111573","DOI":"10.1016\/j.celrep.2022.111573","volume":"41","author":"D Beckman","year":"2022","unstructured":"Beckman D, Bonillas A, Diniz GB, Ott S, Roh JW, Elizaldi SR, et al. SARS-CoV-2 infects neurons and induces neuroinflammation in a non-human primate model of COVID-19. Cell Rep. 2022;41:111573. https:\/\/doi.org\/10.1016\/j.celrep.2022.111573.","journal-title":"Cell Rep"},{"key":"3751_CR9","doi-asserted-by":"publisher","first-page":"4097","DOI":"10.1093\/brain\/awac321","volume":"145","author":"EJ Needham","year":"2022","unstructured":"Needham EJ, Ren AL, Digby RJ, Norton EJ, Ebrahimi S, Outtrim JG, et al. Brain injury in COVID-19 is associated with dysregulated innate and adaptive immune responses. Brain. 2022;145:4097\u2013107. https:\/\/doi.org\/10.1093\/brain\/awac321.","journal-title":"Brain"},{"key":"3751_CR10","doi-asserted-by":"publisher","first-page":"713","DOI":"10.1016\/j.bbi.2024.04.015","volume":"119","author":"MB VanElzakker","year":"2024","unstructured":"VanElzakker MB, Bues HF, Brusaferri L, Kim M, Saadi D, Ratai E-M, et al. Neuroinflammation in post-acute sequelae of COVID-19 (PASC) as assessed by [11\u00a0C]PBR28 PET correlates with vascular disease measures. Brain Behav Immun. 2024;119:713\u201323. https:\/\/doi.org\/10.1016\/j.bbi.2024.04.015.","journal-title":"Brain Behav Immun"},{"key":"3751_CR11","doi-asserted-by":"publisher","first-page":"745","DOI":"10.1007\/s11357-024-01487-4","volume":"47","author":"M Fekete","year":"2025","unstructured":"Fekete M, Lehoczki A, Szappanos \u00c1, Toth A, Mahdi M, S\u00f3tonyi P, et al. Cerebromicrovascular mechanisms contributing to long COVID: implications for neurocognitive health. GeroScience Springer Sci Bus Media LLC. 2025;47:745\u201379. https:\/\/doi.org\/10.1007\/s11357-024-01487-4.","journal-title":"GeroScience Springer Sci Bus Media LLC"},{"key":"3751_CR12","doi-asserted-by":"publisher","first-page":"2555","DOI":"10.1093\/brain\/awac151","volume":"145","author":"MH Lee","year":"2022","unstructured":"Lee MH, Perl DP, Steiner J, Pasternack N, Li W, Maric D, et al. Neurovascular injury with complement activation and inflammation in COVID-19. Brain. 2022;145:2555\u201368. https:\/\/doi.org\/10.1093\/brain\/awac151.","journal-title":"Brain"},{"key":"3751_CR13","doi-asserted-by":"publisher","first-page":"3303","DOI":"10.1007\/s00415-023-11767-2","volume":"270","author":"DD Adingupu","year":"2023","unstructured":"Adingupu DD, Soroush A, Hansen A, Twomey R, Dunn JF. Brain hypoxia, neurocognitive impairment, and quality of life in people post-COVID-19. J Neurol. 2023;270:3303\u201314. https:\/\/doi.org\/10.1007\/s00415-023-11767-2.","journal-title":"J Neurol"},{"key":"3751_CR14","doi-asserted-by":"publisher","first-page":"112189","DOI":"10.1016\/j.celrep.2023.112189","volume":"42","author":"FL Fontes-Dantas","year":"2023","unstructured":"Fontes-Dantas FL, Fernandes GG, Gutman EG, De Lima EV, Antonio LS, Hammerle MB, et al. SARS-CoV-2 spike protein induces TLR4-mediated long-term cognitive dysfunction recapitulating post-COVID-19 syndrome in mice. Cell Rep. 2023;42:112189. https:\/\/doi.org\/10.1016\/j.celrep.2023.112189.","journal-title":"Cell Rep"},{"key":"3751_CR15","doi-asserted-by":"publisher","first-page":"101570","DOI":"10.1016\/j.xcrm.2024.101570","volume":"5","author":"B Lee","year":"2024","unstructured":"Lee B, Choi HN, Che YH, Ko M, Seong HM, Jo MG, et al. SARS-CoV-2 infection exacerbates the cellular pathology of parkinson\u2019s disease in human dopaminergic neurons and a mouse model. Cell Rep Med. 2024;5:101570. https:\/\/doi.org\/10.1016\/j.xcrm.2024.101570.","journal-title":"Cell Rep Med"},{"key":"3751_CR16","doi-asserted-by":"publisher","first-page":"e20202135","DOI":"10.1084\/jem.20202135","volume":"218","author":"E Song","year":"2021","unstructured":"Song E, Zhang C, Israelow B, Lu-Culligan A, Prado AV, Skriabine S, et al. Neuroinvasion of SARS-CoV-2 in human and mouse brain. J Exp Med. 2021;218:e20202135. https:\/\/doi.org\/10.1084\/jem.20202135.","journal-title":"J Exp Med"},{"key":"3751_CR17","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41392-021-00719-9","volume":"6","author":"L Zhang","year":"2021","unstructured":"Zhang L, Zhou L, Bao L, Liu J, Zhu H, Lv Q, et al. SARS-CoV-2 crosses the blood\u2013brain barrier accompanied with basement membrane disruption without tight junctions alteration. Signal Transduct Target Ther. 2021;6:1\u201312. https:\/\/doi.org\/10.1038\/s41392-021-00719-9.","journal-title":"Signal Transduct Target Ther"},{"key":"3751_CR18","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41421-021-00310-0","volume":"7","author":"H Ma","year":"2021","unstructured":"Ma H, Zhu Z, Lin H, Wang S, Zhang P, Li Y, et al. Pyroptosis of syncytia formed by fusion of SARS-CoV-2 spike and ACE2-expressing cells. Cell Discovery. 2021;7:1\u20134. https:\/\/doi.org\/10.1038\/s41421-021-00310-0.","journal-title":"Cell Discovery"},{"key":"3751_CR19","doi-asserted-by":"publisher","first-page":"eadg2248","DOI":"10.1126\/sciadv.adg2248","volume":"9","author":"R Mart\u00ednez-M\u00e1rmol","year":"2023","unstructured":"Mart\u00ednez-M\u00e1rmol R, Giordano-Santini R, Kaulich E, Cho A-N, Przybyla M, Riyadh MA, et al. SARS-CoV-2 infection and viral fusogens cause neuronal and glial fusion that compromises neuronal activity. Sci Adv. 2023;9:eadg2248. https:\/\/doi.org\/10.1126\/sciadv.adg2248.","journal-title":"Sci Adv"},{"key":"3751_CR20","doi-asserted-by":"publisher","first-page":"167280","DOI":"10.1016\/j.jmb.2021.167280","volume":"434","author":"MM Rajah","year":"2022","unstructured":"Rajah MM, Bernier A, Buchrieser J, Schwartz O. The mechanism and consequences of SARS-CoV-2 spike-mediated fusion and syncytia formation. J Mol Biol. 2022;434:167280. https:\/\/doi.org\/10.1016\/j.jmb.2021.167280.","journal-title":"J Mol Biol"},{"key":"3751_CR21","doi-asserted-by":"publisher","first-page":"565","DOI":"10.1038\/s41586-021-03710-0","volume":"595","author":"AC Yang","year":"2021","unstructured":"Yang AC, Kern F, Losada PM, Agam MR, Maat CA, Schmartz GP, et al. Dysregulation of brain and choroid plexus cell types in severe COVID-19. Nature. 2021;595:565\u201371. https:\/\/doi.org\/10.1038\/s41586-021-03710-0.","journal-title":"Nature"},{"key":"3751_CR22","doi-asserted-by":"publisher","first-page":"24","DOI":"10.1038\/s41392-022-01291-6","volume":"8","author":"P-P Zhang","year":"2023","unstructured":"Zhang P-P, He Z-C, Yao X-H, Tang R, Ma J, Luo T, et al. COVID-19-associated monocytic encephalitis (CAME): histological and proteomic evidence from autopsy. Signal Transduct Target Ther. 2023;8:24\u201335. https:\/\/doi.org\/10.1038\/s41392-022-01291-6.","journal-title":"Signal Transduct Target Ther"},{"key":"3751_CR23","doi-asserted-by":"publisher","first-page":"1598","DOI":"10.1038\/s43587-024-00730-z","volume":"4","author":"T Zhang","year":"2024","unstructured":"Zhang T, Li Y, Pan L, Sha J, Bailey M, Faure-Kumar E, et al. Brain-wide alterations revealed by spatial transcriptomics and proteomics in COVID-19 infection. Nat Aging. 2024;4:1598\u2013618. https:\/\/doi.org\/10.1038\/s43587-024-00730-z.","journal-title":"Nat Aging"},{"key":"3751_CR24","doi-asserted-by":"publisher","first-page":"270","DOI":"10.1007\/s12264-018-0306-7","volume":"35","author":"W Qiu","year":"2019","unstructured":"Qiu W, Zhang H, Bao A, Zhu K, Huang Y, Yan X, et al. Standardized operational protocol for human brain banking in China. Neurosci Bull. 2019;35:270\u20136. https:\/\/doi.org\/10.1007\/s12264-018-0306-7.","journal-title":"Neurosci Bull"},{"key":"3751_CR25","doi-asserted-by":"publisher","first-page":"2191","DOI":"10.1001\/jama.2013.281053","volume":"310","author":"World Medical Association","year":"2013","unstructured":"World Medical Association. World medical association declaration of helsinki: ethical principles for medical research involving human subjects. JAMA. 2013;310:2191\u20134. https:\/\/doi.org\/10.1001\/jama.2013.281053.","journal-title":"JAMA"},{"key":"3751_CR26","doi-asserted-by":"publisher","first-page":"452","DOI":"10.1016\/j.neuron.2021.10.036","volume":"110","author":"D Franjic","year":"2022","unstructured":"Franjic D, Skarica M, Ma S, Arellano JI, Tebbenkamp ATN, Choi J, et al. Transcriptomic taxonomy and neurogenic trajectories of adult human, macaque, and pig hippocampal and entorhinal cells. Neuron. 2022;110:452\u2013e46914. https:\/\/doi.org\/10.1016\/j.neuron.2021.10.036.","journal-title":"Neuron"},{"key":"3751_CR27","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1126\/science.abm1741","volume":"377","author":"R Fang","year":"2022","unstructured":"Fang R, Xia C, Close JL, Zhang M, He J, Huang Z, et al. Conservation and divergence of cortical cell organization in human and mouse revealed by MERFISH. Science. 2022;377:56\u201362. https:\/\/doi.org\/10.1126\/science.abm1741.","journal-title":"Science"},{"key":"3751_CR28","doi-asserted-by":"publisher","first-page":"3127","DOI":"10.1002\/cne.24080","volume":"524","author":"S-L Ding","year":"2016","unstructured":"Ding S-L, Royall JJ, Sunkin SM, Ng L, Facer BAC, Lesnar P, et al. Comprehensive cellular-resolution atlas of the adult human brain. J Comp Neurol. 2016;524:3127\u2013481. https:\/\/doi.org\/10.1002\/cne.24080.","journal-title":"J Comp Neurol"},{"key":"3751_CR29","doi-asserted-by":"publisher","first-page":"112420","DOI":"10.1016\/j.biopha.2021.112420","volume":"145","author":"M Farahani","year":"2022","unstructured":"Farahani M, Niknam Z, Mohammadi Amirabad L, Amiri-Dashatan N, Koushki M, Nemati M, et al. Molecular pathways involved in COVID-19 and potential pathway-based therapeutic targets. Biomed Pharmacother. 2022;145:112420. https:\/\/doi.org\/10.1016\/j.biopha.2021.112420.","journal-title":"Biomed Pharmacother"},{"key":"3751_CR30","doi-asserted-by":"publisher","first-page":"2521","DOI":"10.3389\/fimmu.2019.02521","volume":"10","author":"JK Ljungberg","year":"2019","unstructured":"Ljungberg JK, Kling JC, Tran TT, Blumenthal A. Functions of the WNT signaling network in shaping host responses to infection. Front Immunol. 2019;10:2521. https:\/\/doi.org\/10.3389\/fimmu.2019.02521.","journal-title":"Front Immunol"},{"key":"3751_CR31","doi-asserted-by":"publisher","first-page":"250","DOI":"10.1038\/s41423-020-00620-5","volume":"18","author":"C Chhatbar","year":"2021","unstructured":"Chhatbar C, Prinz M. The roles of microglia in viral encephalitis: from sensome to therapeutic targeting. Cell Mol Immunol. 2021;18:250\u20138. https:\/\/doi.org\/10.1038\/s41423-020-00620-5.","journal-title":"Cell Mol Immunol"},{"key":"3751_CR32","doi-asserted-by":"publisher","first-page":"2878","DOI":"10.1038\/s41380-022-01831-0","volume":"28","author":"EA Albornoz","year":"2023","unstructured":"Albornoz EA, Amarilla AA, Modhiran N, Parker S, Li XX, Wijesundara DK, et al. SARS-CoV-2 drives NLRP3 inflammasome activation in human microglia through spike protein. Mol Psychiatry. 2023;28:2878\u201393. https:\/\/doi.org\/10.1038\/s41380-022-01831-0.","journal-title":"Mol Psychiatry"},{"key":"3751_CR33","doi-asserted-by":"publisher","first-page":"4485","DOI":"10.1038\/s41467-023-40228-7","volume":"14","author":"GD de Melo","year":"2023","unstructured":"de Melo GD, Perraud V, Alvarez F, Vieites-Prado A, Kim S, Kergoat L, et al. Neuroinvasion and anosmia are independent phenomena upon infection with SARS-CoV-2 and its variants. Nat Commun. 2023;14:4485. https:\/\/doi.org\/10.1038\/s41467-023-40228-7.","journal-title":"Nat Commun"},{"key":"3751_CR34","doi-asserted-by":"publisher","first-page":"e2200960119","DOI":"10.1073\/pnas.2200960119","volume":"119","author":"F Crunfli","year":"2022","unstructured":"Crunfli F, Carregari VC, Veras FP, Silva LS, Nogueira MH, Antunes ASLM, et al. Morphological, cellular, and molecular basis of brain infection in COVID-19 patients. Proc Natl Acad Sci. 2022;119:e2200960119. https:\/\/doi.org\/10.1073\/pnas.2200960119.","journal-title":"Proc Natl Acad Sci"},{"key":"3751_CR35","doi-asserted-by":"publisher","first-page":"311","DOI":"10.1016\/j.cell.2019.11.039","volume":"180","author":"CCH Cohen","year":"2020","unstructured":"Cohen CCH, Popovic MA, Klooster J, Weil M-T, M\u00f6bius W, Nave K-A, et al. Saltatory conduction along myelinated axons involves a periaxonal nanocircuit. Cell. 2020;180:311\u2013e32215. https:\/\/doi.org\/10.1016\/j.cell.2019.11.039.","journal-title":"Cell"},{"key":"3751_CR36","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1007\/s12035-009-8071-2","volume":"40","author":"O Jahn","year":"2009","unstructured":"Jahn O, Tenzer S, Werner HB. Myelin proteomics: molecular anatomy of an insulating sheath. Mol Neurobiol. 2009;40:55\u201372. https:\/\/doi.org\/10.1007\/s12035-009-8071-2.","journal-title":"Mol Neurobiol"},{"key":"3751_CR37","doi-asserted-by":"publisher","first-page":"6176","DOI":"10.1073\/pnas.1423205112","volume":"112","author":"O Varea","year":"2015","unstructured":"Varea O, Martin-de-Saavedra MD, Kopeikina KJ, Sch\u00fcrmann B, Fleming HJ, Fawcett-Patel JM, et al. Synaptic abnormalities and cytoplasmic glutamate receptor aggregates in contactin associated protein-like 2\/Caspr2 knockout neurons. Proc Natl Acad Sci. 2015;112:6176\u201381. https:\/\/doi.org\/10.1073\/pnas.1423205112.","journal-title":"Proc Natl Acad Sci"},{"key":"3751_CR38","doi-asserted-by":"publisher","first-page":"1189","DOI":"10.1038\/s41564-024-01657-2","volume":"9","author":"E Partiot","year":"2024","unstructured":"Partiot E, Hirschler A, Colomb S, Lutz W, Claeys T, Delalande F, et al. Brain exposure to SARS-CoV-2 virions perturbs synaptic homeostasis. Nat Microbiol. 2024;9:1189\u2013206. https:\/\/doi.org\/10.1038\/s41564-024-01657-2.","journal-title":"Nat Microbiol"},{"key":"3751_CR39","doi-asserted-by":"publisher","first-page":"101044","DOI":"10.1016\/j.eclinm.2021.101044","volume":"39","author":"A Hampshire","year":"2021","unstructured":"Hampshire A, Trender W, Chamberlain SR, Jolly AE, Grant JE, Patrick F, et al. Cognitive deficits in people who have recovered from COVID-19. EClinicalMedicine. 2021;39:101044. https:\/\/doi.org\/10.1016\/j.eclinm.2021.101044.","journal-title":"EClinicalMedicine"},{"key":"3751_CR40","doi-asserted-by":"publisher","unstructured":"Venkataramani V, Winkler F. Cognitive deficits in long covid-19. Phimister EG, editor. N Engl J Med. 2022;387:1813\u20135. https:\/\/doi.org\/10.1056\/NEJMcibr2210069","DOI":"10.1056\/NEJMcibr2210069"},{"key":"3751_CR41","doi-asserted-by":"publisher","first-page":"102434","DOI":"10.1016\/j.eclinm.2024.102434","volume":"68","author":"S Zhao","year":"2024","unstructured":"Zhao S, Martin EM, Reuken PA, Scholcz A, Ganse-Dumrath A, Srowig A, et al. Long COVID is associated with severe cognitive slowing: a multicentre cross-sectional study. eClinicalMedicine. 2024;68:102434. https:\/\/doi.org\/10.1016\/j.eclinm.2024.102434.","journal-title":"eClinicalMedicine"},{"key":"3751_CR42","doi-asserted-by":"publisher","first-page":"556","DOI":"10.1016\/j.bbi.2018.02.003","volume":"69","author":"Y Cheng","year":"2018","unstructured":"Cheng Y, Desse S, Martinez A, Worthen RJ, Jope RS, Beurel E. TNF\u03b1 disrupts blood brain barrier integrity to maintain prolonged depressive-like behavior in mice. Brain Behav Immun. 2018;69:556\u201367. https:\/\/doi.org\/10.1016\/j.bbi.2018.02.003.","journal-title":"Brain Behav Immun"},{"key":"3751_CR43","doi-asserted-by":"publisher","first-page":"142","DOI":"10.1038\/s41420-021-00531-y","volume":"7","author":"Q Liu","year":"2021","unstructured":"Liu Q, Liu X, Zhao D, Ruan X, Su R, Shang X, et al. Pseudogene ACTBP2 increases blood-brain barrier permeability by promoting KHDRBS2 transcription through recruitment of KMT2D\/WDR5 in A\u03b21\u201342 microenvironment. Cell Death Discovery. 2021;7:142\u201354. https:\/\/doi.org\/10.1038\/s41420-021-00531-y.","journal-title":"Cell Death Discovery"},{"key":"3751_CR44","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1186\/s12964-024-01982-3","volume":"23","author":"K Wang","year":"2025","unstructured":"Wang K, Qu H, Hu R, Lass\u00e8gue B, Eaton DC, Song C, et al. Polymerase delta-interacting protein 2 mediates brain vascular permeability by regulating ROS-mediated ZO-1 phosphorylation and localization at the interendothelial border. Cell Commun Signal. 2025;23:9\u201325. https:\/\/doi.org\/10.1186\/s12964-024-01982-3.","journal-title":"Cell Commun Signal"},{"key":"3751_CR45","doi-asserted-by":"publisher","first-page":"e140329","DOI":"10.1172\/jci.insight.140329","volume":"5","author":"KE Remy","year":"2020","unstructured":"Remy KE, Mazer M, Striker DA, Ellebedy AH, Walton AH, Unsinger J, et al. Severe immunosuppression and not a cytokine storm characterizes COVID-19 infections. JCI Insight. 2020;5:e140329. https:\/\/doi.org\/10.1172\/jci.insight.140329.","journal-title":"JCI Insight"},{"key":"3751_CR46","doi-asserted-by":"publisher","first-page":"5859","DOI":"10.1038\/s41467-020-19706-9","volume":"11","author":"W Tian","year":"2020","unstructured":"Tian W, Zhang N, Jin R, Feng Y, Wang S, Gao S, et al. Immune suppression in the early stage of COVID-19 disease. Nat Commun. 2020;11:5859. https:\/\/doi.org\/10.1038\/s41467-020-19706-9.","journal-title":"Nat Commun"},{"key":"3751_CR47","doi-asserted-by":"publisher","first-page":"1895","DOI":"10.1016\/j.cell.2021.01.053","volume":"184","author":"X Ren","year":"2021","unstructured":"Ren X, Wen W, Fan X, Hou W, Su B, Cai P, et al. COVID-19 immune features revealed by a large-scale single-cell transcriptome atlas. Cell. 2021;184:1895\u2013e1913. https:\/\/doi.org\/10.1016\/j.cell.2021.01.053. 19.","journal-title":"Cell"},{"key":"3751_CR48","doi-asserted-by":"publisher","first-page":"118","DOI":"10.1186\/s13073-021-00933-8","volume":"13","author":"JF Fullard","year":"2021","unstructured":"Fullard JF, Lee H-C, Voloudakis G, Suo S, Javidfar B, Shao Z, et al. Single-nucleus transcriptome analysis of human brain immune response in patients with severe COVID-19. Genome Med. 2021;13:118\u201331. https:\/\/doi.org\/10.1186\/s13073-021-00933-8.","journal-title":"Genome Med"},{"key":"3751_CR49","doi-asserted-by":"publisher","first-page":"3939","DOI":"10.1038\/s41380-022-01786-2","volume":"27","author":"AO Samudyata, Oliveira","year":"2022","unstructured":"Samudyata, Oliveira AO, Malwade S, Rufino de Sousa N, Goparaju SK, Gracias J, et al. SARS-CoV-2 promotes microglial synapse elimination in human brain organoids. Mol Psychiatry. 2022;27:3939\u201350. https:\/\/doi.org\/10.1038\/s41380-022-01786-2.","journal-title":"Mol Psychiatry"},{"key":"3751_CR50","doi-asserted-by":"publisher","first-page":"130","DOI":"10.1038\/s41398-024-02846-7","volume":"14","author":"M Geng","year":"2024","unstructured":"Geng M, Shao Q, Fu J, Gu J, Feng L, Zhao L, et al. Down-regulation of MKP-1 in hippocampus protects against stress-induced depression-like behaviors and neuroinflammation. Transl Psychiatry. 2024;14:130\u201340. https:\/\/doi.org\/10.1038\/s41398-024-02846-7.","journal-title":"Transl Psychiatry"},{"key":"3751_CR51","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1186\/s12974-024-03261-x","volume":"21","author":"X Huang","year":"2024","unstructured":"Huang X, Wei P, Fang C, Yu M, Yang S, Qiu L, et al. Compromised endothelial wnt\/\u03b2-catenin signaling mediates the blood-brain barrier disruption and leads to neuroinflammation in endotoxemia. J Neuroinflammation. 2024;21:265. https:\/\/doi.org\/10.1186\/s12974-024-03261-x.","journal-title":"J Neuroinflammation"},{"key":"3751_CR52","doi-asserted-by":"publisher","first-page":"106","DOI":"10.1038\/s41392-025-02142-w","volume":"10","author":"C Xue","year":"2025","unstructured":"Xue C, Chu Q, Shi Q, Zeng Y, Lu J, Li L. Wnt signaling pathways in biology and disease: mechanisms and therapeutic advances. Signal Transduct Target Ther. 2025;10:106. https:\/\/doi.org\/10.1038\/s41392-025-02142-w.","journal-title":"Signal Transduct Target Ther"},{"key":"3751_CR53","doi-asserted-by":"publisher","first-page":"e2111400119","DOI":"10.1073\/pnas.2111400119","volume":"119","author":"C Zeng","year":"2022","unstructured":"Zeng C, Evans JP, King T, Zheng Y-M, Oltz EM, Whelan SPJ, et al. SARS-CoV-2 spreads through cell-to-cell transmission. Proc Natl Acad Sci. 2022;119:e2111400119. https:\/\/doi.org\/10.1073\/pnas.2111400119.","journal-title":"Proc Natl Acad Sci"},{"key":"3751_CR54","doi-asserted-by":"publisher","first-page":"153","DOI":"10.1186\/s13059-024-03303-w","volume":"25","author":"L Atta","year":"2024","unstructured":"Atta L, Clifton K, Anant M, Aihara G, Fan J. Gene count normalization in single-cell imaging-based spatially resolved transcriptomics. Genome Biol. 2024;25:153\u201378. https:\/\/doi.org\/10.1186\/s13059-024-03303-w.","journal-title":"Genome Biol"}],"container-title":["Journal of Neuroinflammation"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12974-026-03751-0","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12974-026-03751-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12974-026-03751-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,27]],"date-time":"2026-03-27T13:48:56Z","timestamp":1774619336000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s12974-026-03751-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,23]]},"references-count":54,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["3751"],"URL":"https:\/\/doi.org\/10.1186\/s12974-026-03751-0","relation":{},"ISSN":["1742-2094"],"issn-type":[{"value":"1742-2094","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,23]]},"assertion":[{"value":"21 November 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"17 February 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"23 February 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Ethical approval for the study was granted by the Institutional Review Board of the Institute of Basic Medical Sciences & School of Basic Medicine, Chinese Academy of Medical Sciences & Peking Union Medical College, and informed consent was obtained from the donors, ensuring that the use of these brain samples adhered strictly to the National Human Brain Bank for Development and Function guidelines and the Declaration of Helsinki. The research protocol was approved by the Institutional Review Board of the Institute of Basic Medical Sciences & School of Basic Medicine, Chinese Academy of Medical Sciences & Peking Union Medical College, China (Approval Number: 2022125).","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"All the authors have consented to the publication of this manuscript.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare no competing interests.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"106"}}