{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T21:14:15Z","timestamp":1789161255902,"version":"build-2803163510"},"reference-count":72,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,5,21]],"date-time":"2026-05-21T00:00:00Z","timestamp":1779321600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Aging Neurosci."],"abstract":"<jats:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>Partly due to the inaccessibility of olfactory brain regions vulnerable to early Alzheimer\u2019s Disease (AD) for repeated sampling, proteomic networks underlying progressive cognitive decline remain poorly understood. The olfactory mucosa (OM), an accessible part of the olfactory system, reflects central nervous system physiology and pathology, and represents a promising site for biomarker discovery. This study aimed to identify olfactory proteomic markers and pathways associated with performance in the logical memory II recognition (LM II_recog) subtest of the Wechsler Memory Scale among older adults with subjective cognitive complaints.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      Clinical, olfactory, and cognitive assessments were conducted on 108 adults aged 55\u201385\u202fyears from the Washington, DC region. Nasal exfoliates were sampled from the upper nasal cavities, and protein extracts from these samples were analyzed by mass spectrometry (MS). Linear regression with false discovery rate (FDR) correction (\n                      <jats:italic>q<\/jats:italic>\n                      \u202f&amp;lt;\u202f0.1) was used to identify proteins associated with LM II_recog performance, and ingenuity pathway analysis (IPA) was applied to determine functional pathways.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      A total of 137 proteins meeting the FDR\n                      <jats:italic>q<\/jats:italic>\n                      \u202f&amp;lt;\u202f0.1 threshold were found to be linearly correlated with LM II_recog scores. Of the top 10 most significant proteins, six (PLOD1, MFN2, NGFR, PPP2R5E, C4A\/C4B, and ITGAV) have previously been linked to AD and\/or cognitive function, underscoring their potential as biomarkers of cognitive impairment. Ingenuity pathway analysis using the knowledge base machine learning (ML) platform revealed several disease pathways highly represented among the significant proteins. These included Hyperactive Behavior, Neuromuscular Disease, Tauopathy, Behavioral Deficits, Alzheimer\u2019s Disease, Progressive Dementia, Degenerative Dementia, Alzheimer\u2019s or Frontotemporal Dementia, all of which were associated with LM II_recog performance in the elderly population.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Discussion<\/jats:title>\n                    <jats:p>This study demonstrates the feasibility of using OM-derived proteomics to identify molecular signatures associated with cognitive performance and highlights the OM as a potential site for non-invasive biomarker discovery. These findings provide a foundation for future studies integrating OM profiling with established AD biomarkers.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fnagi.2026.1781518","type":"journal-article","created":{"date-parts":[[2026,5,21]],"date-time":"2026-05-21T05:46:35Z","timestamp":1779342395000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":1,"title":["Proteomic profiling of olfactory exfoliates from people with subjective cognitive complaints reveal networks of olfactory biomarkers of cognitive performance"],"prefix":"10.3389","volume":"18","author":[{"given":"Tamal","family":"Sadhukhan","sequence":"first","affiliation":[{"name":"Department of Psychiatry and Behavioral Sciences, Howard University","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Narayan","family":"Rai","sequence":"additional","affiliation":[{"name":"Department of Psychiatry and Behavioral Sciences, Howard University","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria Ma\u00f1anita S.","family":"Hipolito","sequence":"additional","affiliation":[{"name":"Department of Psychiatry and Behavioral Sciences, Howard University","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Myeshia","family":"Shelby","sequence":"additional","affiliation":[{"name":"Department of Psychiatry and Behavioral Sciences, Howard University","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Claudia Ivonne","family":"Mej\u00eda Mondrag\u00f3n","sequence":"additional","affiliation":[{"name":"Department of Psychiatry and Behavioral Sciences, Howard University","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sriparna","family":"Sadhukhan","sequence":"additional","affiliation":[{"name":"Department of Psychiatry and Behavioral Sciences, Howard University","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Abimbola","family":"Idowu","sequence":"additional","affiliation":[{"name":"Department of Psychiatry and Behavioral Sciences, Howard University","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rency S.","family":"Varghese","sequence":"additional","affiliation":[{"name":"Lombardi Comprehensive Cancer Center, Department of Oncology, Georgetown University Medical Center","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Muhammad Salman","family":"Sajid","sequence":"additional","affiliation":[{"name":"Lombardi Comprehensive Cancer Center, Department of Oncology, Georgetown University Medical Center","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Habtom W.","family":"Ressom","sequence":"additional","affiliation":[{"name":"Lombardi Comprehensive Cancer Center, Department of Oncology, Georgetown University Medical Center","place":["Washington, DC, United States"]},{"name":"Department of Oncology, Genomics and Epigenomics Shared Resources, Georgetown University","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Adedoyin","family":"Kalejaiye","sequence":"additional","affiliation":[{"name":"Division of Otolaryngology\/Head and Neck Surgery, Department of Surgery, Howard University","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas O.","family":"Obisesan","sequence":"additional","affiliation":[{"name":"Department of Internal Medicine, Howard University","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Magdalena","family":"Misiak-Christian","sequence":"additional","affiliation":[{"name":"Department of Physiology and Biophysics, Howard University","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Evaristus A.","family":"Nwulia","sequence":"additional","affiliation":[{"name":"Department of Psychiatry and Behavioral Sciences, Howard University","place":["Washington, DC, United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,5,21]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","first-page":"1247","DOI":"10.30773\/pi.2019.0061e","article-title":"Normative data for the logical memory subtest of the Wechsler memory scale-IV in middle-aged and elderly Korean people","volume":"17","author":"Ahn","year":"2020","journal-title":"Psychiatry Investig."},{"key":"ref2","doi-asserted-by":"publisher","first-page":"5695","DOI":"10.1002\/alz.13844","article-title":"Sex and gender differences in cognitive resilience to aging and Alzheimer's disease","volume":"20","author":"Arenaza-Urquijo","year":"2024","journal-title":"Alzheimers Dement."},{"key":"ref3","doi-asserted-by":"publisher","first-page":"462","DOI":"10.1002\/ana.21910","article-title":"Olfactory epithelium amyloid-beta and paired helical filament-tau pathology in Alzheimer disease","volume":"67","author":"Arnold","year":"2010","journal-title":"Ann. Neurol."},{"key":"ref4","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1016\/s0166-4328(01)00297-2","article-title":"Controlling the false discovery rate in behavior genetics research","volume":"125","author":"Benjamini","year":"2001","journal-title":"Behav. Brain Res."},{"key":"ref5","doi-asserted-by":"publisher","first-page":"1149","DOI":"10.1007\/s11064-010-0371-4","article-title":"Calcium signalling and Alzheimer's disease","volume":"36","author":"Berridge","year":"2011","journal-title":"Neurochem. Res."},{"key":"ref6","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1186\/s40035-022-00311-3","article-title":"Olfactory swab sampling optimization for \u03b1-synuclein aggregate detection in patients with Parkinson's disease","volume":"11","author":"Bongianni","year":"2022","journal-title":"Transl Neurodegener"},{"key":"ref7","doi-asserted-by":"publisher","first-page":"450","DOI":"10.1056\/NEJM200008173430701","article-title":"Patterns of brain activation in people at risk for Alzheimer's disease","volume":"343","author":"Bookheimer","year":"2000","journal-title":"N. Engl. J. Med."},{"key":"ref8","doi-asserted-by":"publisher","first-page":"919","DOI":"10.1007\/s10072-022-06500-2","article-title":"RNAseq analysis of olfactory neuroepithelium cytological samples in individuals with down syndrome compared to euploid controls: a pilot study","volume":"44","author":"Brozzetti","year":"2023","journal-title":"Neurol. Sci."},{"key":"ref9","doi-asserted-by":"publisher","first-page":"3200","DOI":"10.3390\/ijms24043200","article-title":"The nerve growth factor receptor (NGFR\/p75(NTR)): a major player in Alzheimer's disease","volume":"24","author":"Bruno","year":"2023","journal-title":"Int. J. Mol. Sci."},{"key":"ref10","doi-asserted-by":"publisher","first-page":"399","DOI":"10.3233\/JAD-121621","article-title":"Gene expression profiling of peripheral blood leukocytes shows consistent longitudinal downregulation of TOMM40 and upregulation of KIR2DL5A, PLOD1, and SLC2A8 among fast progressors in early Alzheimer's disease","volume":"34","author":"Chong","year":"2013","journal-title":"J. Alzheimer's Dis"},{"key":"ref11","doi-asserted-by":"publisher","first-page":"2245","DOI":"10.1038\/s41467\u20137","article-title":"Olfactory cleft biopsy analysis of Alzheimer's disease pathobiology across disease stages","volume":"17","author":"D'Anniballe","year":"2026","journal-title":"Nat. Commun."},{"key":"ref12","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3390\/proteomes6010001","article-title":"Comparative proteomics analysis of urine reveals down-regulation of acute phase response signaling and LXR\/RXR activation pathways in prostate cancer","volume":"6","author":"Davalieva","year":"2017","journal-title":"Proteomes"},{"key":"ref13","doi-asserted-by":"publisher","first-page":"961","DOI":"10.1001\/jamaneurol.2014.803","article-title":"The preclinical Alzheimer cognitive composite: measuring amyloid-related decline","volume":"71","author":"Donohue","year":"2014","journal-title":"JAMA Neurol."},{"key":"ref14","doi-asserted-by":"publisher","first-page":"480","DOI":"10.2500\/ajra.2009.23.3385","article-title":"Regression of polypoid nasal mucosa after systemic corticosteroid therapy: a proteomics study","volume":"23","author":"Farajzadeh Deroee","year":"2009","journal-title":"Am. J. Rhinol. Allergy"},{"key":"ref15","doi-asserted-by":"crossref","first-page":"1349","DOI":"10.1001\/jama.1997.03550160069041","article-title":"Effects of age, sex, and ethnicity on the association between apolipoprotein E genotype and Alzheimer disease. A meta-analysis. APOE and Alzheimer disease Meta analysis consortium","volume":"278","author":"Farrer","year":"1997","journal-title":"JAMA"},{"key":"ref16","doi-asserted-by":"publisher","first-page":"330","DOI":"10.1038\/s41419-017-0023-6","article-title":"Mitofusin 2: from functions to disease","volume":"9","author":"Filadi","year":"2018","journal-title":"Cell Death Dis."},{"key":"ref17","doi-asserted-by":"publisher","first-page":"7209","DOI":"10.1073\/pnas.0811879106","article-title":"Distinct patterns of brain activity in young carriers of the APOE-epsilon4 allele","volume":"106","author":"Filippini","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref18","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1016\/0022-3956(75)90026-6","article-title":"\"Mini-mental state\". A practical method for grading the cognitive state of patients for the clinician","volume":"12","author":"Folstein","year":"1975","journal-title":"J. Psychiatr. Res."},{"key":"ref19","doi-asserted-by":"publisher","first-page":"1622","DOI":"10.1016\/j.bcp.2006.05.017","article-title":"S100A8 and S100A9 in inflammation and cancer","volume":"72","author":"Gebhardt","year":"2006","journal-title":"Biochem. Pharmacol."},{"key":"ref20","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1111\/nyas.13031","article-title":"Vitamin B1 (thiamine) and dementia","volume":"1367","author":"Gibson","year":"2016","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref21","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1002\/ana.20694","article-title":"APOE epsilon4 is not associated with Alzheimer's disease in elderly Nigerians","volume":"59","author":"Gureje","year":"2006","journal-title":"Ann. Neurol."},{"key":"ref22","doi-asserted-by":"publisher","first-page":"2218","DOI":"10.1111\/cns.13965","article-title":"Association of rs2072446 in the NGFR gene with the risk of Alzheimer's disease and amyloid-\u03b2 deposition in the brain","volume":"28","author":"He","year":"2022","journal-title":"CNS Neurosci. Ther."},{"key":"ref23","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1038\/nrneurol.2017.99","article-title":"ER stress and the unfolded protein response in neurodegeneration","volume":"13","author":"Hetz","year":"2017","journal-title":"Nat. Rev. Neurol."},{"key":"ref24","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1016\/j.nbd.2014.08.025","article-title":"Apolipoprotein E: structure and function in lipid metabolism, neurobiology, and Alzheimer's diseases","volume":"72","author":"Huang","year":"2014","journal-title":"Neurobiol. Dis."},{"key":"ref25","author":"Huber","year":"1967"},{"key":"ref26","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1007\/s00405-006-0173-0","article-title":"Normative data for the \"Sniffin' sticks\" including tests of odor identification, odor discrimination, and olfactory thresholds: an upgrade based on a group of more than 3,000 subjects","volume":"264","author":"Hummel","year":"2007","journal-title":"Eur. Arch. Otorrinolaringol."},{"key":"ref27","doi-asserted-by":"publisher","first-page":"4320","DOI":"10.1021\/acs.jproteome.8b00408","article-title":"Identification of missing proteins in human olfactory epithelial tissue by liquid chromatography-tandem mass spectrometry","volume":"17","author":"Hwang","year":"2018","journal-title":"J. Proteome Res."},{"key":"ref28","doi-asserted-by":"publisher","first-page":"513","DOI":"10.1097\/JCP.0000000000000941","article-title":"Digit symbol substitution test: the case for sensitivity over specificity in neuropsychological testing","volume":"38","author":"Jaeger","year":"2018","journal-title":"J. Clin. Psychopharmacol."},{"key":"ref29","doi-asserted-by":"publisher","first-page":"2042","DOI":"10.1002\/alz.12607","article-title":"Protein phosphatase 2A and complement component 4 are linked to the protective effect of APOE \u025b2 for Alzheimer's disease","volume":"18","author":"Jun","year":"2022","journal-title":"Alzheimers Dement."},{"key":"ref30","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1038\/s41467-022-28090-5","article-title":"Breathing coordinates cortico-hippocampal dynamics in mice during offline states","volume":"13","author":"Karalis","year":"2022","journal-title":"Nat. Commun."},{"key":"ref31","doi-asserted-by":"publisher","first-page":"88","DOI":"10.1007\/s40520-025-02946-4","article-title":"Cognitive function in generally healthy adults age 70 years and older in the 5-country DO-HEALTH study: MMSE and MoCA scores by sex, education and country","volume":"37","author":"Kistler-Fischbacher","year":"2025","journal-title":"Aging Clin. Exp. Res."},{"key":"ref32","doi-asserted-by":"publisher","first-page":"523","DOI":"10.1093\/bioinformatics\/btt703","article-title":"Causal analysis approaches in ingenuity pathway analysis","volume":"30","author":"Kr\u00e4mer","year":"2014","journal-title":"Bioinformatics"},{"key":"ref33","doi-asserted-by":"publisher","first-page":"100","DOI":"10.1016\/j.molmed.2016.12.010","article-title":"The olfactory neural epithelium as a tool in neuroscience","volume":"23","author":"Lavoie","year":"2017","journal-title":"Trends Mol. Med."},{"key":"ref34","doi-asserted-by":"publisher","first-page":"518","DOI":"10.1016\/j.coi.2011.05.007","article-title":"Inflammation and mesenchymal stem cell aging","volume":"23","author":"Lepperdinger","year":"2011","journal-title":"Curr. Opin. Immunol."},{"key":"ref35","doi-asserted-by":"publisher","first-page":"392","DOI":"10.1159\/000092808","article-title":"Brief informant screening test for mild cognitive impairment and early Alzheimer's disease","volume":"21","author":"Li","year":"2006","journal-title":"Dement. Geriatr. Cogn. Disord."},{"key":"ref36","doi-asserted-by":"publisher","first-page":"5189","DOI":"10.3390\/jcm12165189","article-title":"Genetics of Alzheimer's disease in the African American population","volume":"12","author":"Logue","year":"2023","journal-title":"J. Clin. Med."},{"key":"ref37","doi-asserted-by":"publisher","first-page":"1671552","DOI":"10.3389\/fnagi.2025.1671552","article-title":"The estrogen-brain interface in neuroinflammation: a multidimensional mechanistic insight","volume":"17","author":"Lu","year":"2025","journal-title":"Front. Aging Neurosci."},{"key":"ref9001","doi-asserted-by":"publisher","first-page":"S871","DOI":"10.1016\/j.euroneuro.2017.08.163","article-title":"SA91 \u2013 Increased predicted C4A expression is associated with cognitive deficit in both schizophrenia and Alzheimer\u2019s disease","volume":"29","author":"McCarthy","year":"2019","journal-title":"Eur. Neuropsychopharmacol."},{"key":"ref38","doi-asserted-by":"publisher","first-page":"e209","DOI":"10.1212\/NXG.0000000000000209","article-title":"Biallelic CHP1 mutation causes human autosomal recessive ataxia by impairing NHE1 function","volume":"4","author":"Mendoza-Ferreira","year":"2018","journal-title":"Neurol Genet"},{"key":"ref39","doi-asserted-by":"publisher","first-page":"169","DOI":"10.4172\/2471\u20132701.1000169","article-title":"Apo E4 alleles and impaired olfaction as predictors of Alzheimer's disease","volume":"3","author":"Misiak","year":"2017","journal-title":"Clin. Exp. Psychol."},{"key":"ref40","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1146\/annurev-neuro-112210-112917","article-title":"How is the olfactory map formed and interpreted in the mammalian brain?","volume":"34","author":"Mori","year":"2011","journal-title":"Annu. Rev. Neurosci."},{"key":"ref41","doi-asserted-by":"publisher","first-page":"431","DOI":"10.1001\/archneur.63.3.431","article-title":"Association of apolipoprotein E genotype and Alzheimer disease in African Americans","volume":"63","author":"Murrell","year":"2006","journal-title":"Arch. Neurol."},{"key":"ref9002","doi-asserted-by":"publisher","first-page":"148945","DOI":"10.1016\/j.gene.2024.148945","article-title":"PPP2R5E: New gene potentially involved in specific learning disorders and myopathy","volume":"933","author":"Musumeci","year":"2025","journal-title":"Gene"},{"key":"ref42","doi-asserted-by":"publisher","first-page":"537","DOI":"10.1002\/jts.22222","article-title":"A pilot study of reduced olfactory bulb volume as a marker of PTSD in childhood trauma-exposed adult HIV-infected patients","volume":"30","author":"Nwulia","year":"2017","journal-title":"J. Trauma. Stress."},{"key":"ref43","first-page":"155","article-title":"Measurement of blood thiamine metabolites for Alzheimer's disease diagnosis","volume-title":"EBioMedicine","author":"Pan","year":"2015"},{"key":"ref44","doi-asserted-by":"publisher","first-page":"612521","DOI":"10.3389\/fgene.2020.612521","article-title":"An integrative computational approach based on expression similarity signatures to identify protein-protein interaction networks in female-specific cancers","volume":"11","author":"Pane","year":"2020","journal-title":"Front. Genet."},{"key":"ref45","doi-asserted-by":"publisher","first-page":"6054","DOI":"10.1038\/s41380-021-01266-z","article-title":"Integrative brain transcriptome analysis links complement component 4 and HSPA2 to the APOE \u03b52 protective effect in Alzheimer disease","volume":"26","author":"Panitch","year":"2021","journal-title":"Mol. Psychiatry"},{"key":"ref46","doi-asserted-by":"publisher","first-page":"pgae296","DOI":"10.1093\/pnasnexus\/pgae296","article-title":"Phishing vulnerability compounded by older age, apolipoprotein E e4 genotype, and lower cognition","volume":"3","author":"Pehlivanoglu","year":"2024","journal-title":"PNAS Nexus"},{"key":"ref47","first-page":"581","article-title":"Memory and MRI-based hippocampal volumes in aging and AD","volume-title":"Neurology","author":"Petersen","year":"2000"},{"key":"ref48","doi-asserted-by":"publisher","first-page":"357","DOI":"10.1080\/13825580902825220","article-title":"Differential memory test sensitivity for diagnosing amnestic mild cognitive impairment and predicting conversion to Alzheimer's disease","volume":"16","author":"Rabin","year":"2009","journal-title":"Neuropsychol. Dev. Cogn. B Aging Neuropsychol. Cogn."},{"key":"ref49","doi-asserted-by":"publisher","first-page":"247","DOI":"10.1016\/j.brainresrev.2003.08.005","article-title":"Emotional tagging of memory formation--in the search for neural mechanisms","volume":"43","author":"Richter-Levin","year":"2003","journal-title":"Brain Res. Brain Res. Rev."},{"key":"ref50","doi-asserted-by":"publisher","first-page":"24497","DOI":"10.1038\/s41598-021-04326-0","article-title":"Long non-coding RNA-associated competing endogenous RNA axes in the olfactory epithelium in schizophrenia: a bioinformatics analysis","volume":"11","author":"Sabaie","year":"2021","journal-title":"Sci. Rep."},{"key":"ref51","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1002\/ar.10047","article-title":"Neural regeneration and the peripheral olfactory system","volume":"269","author":"Schwob","year":"2002","journal-title":"Anat. Rec."},{"key":"ref9003","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1038\/s41536-023-00311-5","article-title":"Nerve growth factor receptor (Ngfr) induces neurogenic plasticity by suppressing reactive astroglial Lcn2\/Slc22a17 signaling in Alzheimer\\u0027s disease","volume":"8","author":"Siddiqui","year":"2023","journal-title":"NPJ Regen Med."},{"key":"ref52","doi-asserted-by":"publisher","first-page":"126","DOI":"10.3390\/jcm9010126","article-title":"Back to the fusion: mitofusin-2 in Alzheimer's disease","volume":"9","author":"Sita","year":"2020","journal-title":"J. Clin. Med."},{"key":"ref53","doi-asserted-by":"publisher","first-page":"1162","DOI":"10.1002\/alr.22743","article-title":"Olfactory cleft mucus proteome in chronic rhinosinusitis: a case-control pilot study","volume":"11","author":"Soler","year":"2021","journal-title":"Int Forum Allergy Rhinol"},{"key":"ref54","first-page":"287","article-title":"Altered expression levels of the protein phosphatase 2A ABalphaC enzyme are associated with Alzheimer disease pathology","volume-title":"J. Neuropathol. Exp. Neurol.","author":"Sontag","year":"2004"},{"key":"ref55","doi-asserted-by":"publisher","first-page":"16","DOI":"10.3389\/fnmol.2014.00016","article-title":"Protein phosphatase 2A dysfunction in Alzheimer's disease","volume":"7","author":"Sontag","year":"2014","journal-title":"Front. Mol. Neurosci."},{"key":"ref56","doi-asserted-by":"publisher","first-page":"280","DOI":"10.1016\/j.jalz.2011.03.003","article-title":"Toward defining the preclinical stages of Alzheimer's disease: recommendations from the National Institute on Aging-Alzheimer's Association workgroups on diagnostic guidelines for Alzheimer's disease","volume":"7","author":"Sperling","year":"2011","journal-title":"Alzheimers Dement."},{"key":"ref57","doi-asserted-by":"publisher","first-page":"178","DOI":"10.1037\/\/0894-4105.14.2.178","article-title":"Longitudinal changes in cognition, gait, and balance in abstinent and relapsed alcoholic men: relationships to changes in brain structure","volume":"14","author":"Sullivan","year":"2000","journal-title":"Neuropsychology"},{"key":"ref58","doi-asserted-by":"publisher","first-page":"e12018","DOI":"10.1002\/trc2.12018","article-title":"Plasma proteomic signatures predict dementia and cognitive impairment","volume":"6","author":"Tanaka","year":"2020","journal-title":"Alzheimers Dement (N Y)"},{"key":"ref59","doi-asserted-by":"publisher","first-page":"41","DOI":"10.14283\/jpad.2022.101","article-title":"The use of episodic memory tests for screening in clinical trials for early Alzheimer's disease: a comparison of the free and cued selective reminding test (FCSRT) and the repeatable battery for the assessment of neuropsychological status (RBANS)","volume":"10","author":"Teng","year":"2023","journal-title":"J. Prev Alzheimers Dis."},{"key":"ref60","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1016\/j.mcn.2017.07.002","article-title":"Downregulation of protein phosphatase 2A by apolipoprotein E: implications for Alzheimer's disease","volume":"83","author":"Theendakara","year":"2017","journal-title":"Mol. Cell. Neurosci."},{"key":"ref61","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1007\/978-1-4939-3106-4_3","article-title":"Preprocessing and analysis of LC-MS-based proteomic data","volume":"1362","author":"Tsai","year":"2016","journal-title":"Methods Mol. Biol."},{"key":"ref62","doi-asserted-by":"publisher","first-page":"416","DOI":"10.4172\/jpb.1000102","article-title":"LC-MS based detection of differential protein expression","volume":"2","author":"Tuli","year":"2009","journal-title":"J Proteomics Bioinform"},{"key":"ref63","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1016\/j.biocel.2004.07.009","article-title":"The role of inflammation in Alzheimer's disease","volume":"37","author":"Tuppo","year":"2005","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"ref64","doi-asserted-by":"publisher","first-page":"1057","DOI":"10.3233\/JAD-215175","article-title":"Mitochondrial fusion suppresses tau pathology-induced neurodegeneration and cognitive decline","volume":"84","author":"Wang","year":"2021","journal-title":"J. Alzheimer's Dis"},{"key":"ref65","doi-asserted-by":"publisher","first-page":"817","DOI":"10.2307\/1912934","article-title":"A heteroskedasticity-consistent covariance matrix estimator and a direct test for heteroskedasticity","volume":"48","author":"White","year":"1980","journal-title":"Econometrica"},{"key":"ref66","doi-asserted-by":"publisher","first-page":"17170","DOI":"10.1038\/s41598-018-35102-2","article-title":"The human olfactory cleft mucus proteome and its age-related changes","volume":"8","author":"Yoshikawa","year":"2018","journal-title":"Sci. Rep."},{"key":"ref67","doi-asserted-by":"publisher","first-page":"1437575","DOI":"10.3389\/fncir.2024.1437575","article-title":"Circuit dynamics of the olfactory pathway during olfactory learning","volume":"18","author":"Zhang","year":"2024","journal-title":"Front Neural Circuits"},{"key":"ref68","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1186\/s13024-016-0069-4","article-title":"A rapid and cost-effective method for genotyping apolipoprotein E gene polymorphism","volume":"11","author":"Zhong","year":"2016","journal-title":"Mol. Neurodegener."},{"key":"ref69","doi-asserted-by":"publisher","first-page":"869","DOI":"10.2147\/NDT.S104886","article-title":"Olfactory dysfunction in Alzheimer's disease","volume":"12","author":"Zou","year":"2016","journal-title":"Neuropsychiatr. Dis. Treat."}],"container-title":["Frontiers in Aging Neuroscience"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fnagi.2026.1781518\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,4]],"date-time":"2026-06-04T14:06:58Z","timestamp":1780582018000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fnagi.2026.1781518\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,21]]},"references-count":72,"alternative-id":["10.3389\/fnagi.2026.1781518"],"URL":"https:\/\/doi.org\/10.3389\/fnagi.2026.1781518","relation":{},"ISSN":["1663-4365"],"issn-type":[{"value":"1663-4365","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,21]]},"article-number":"1781518"}}