{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T00:32:46Z","timestamp":1787877166823,"version":"build-2784847793"},"reference-count":76,"publisher":"Springer Science and Business Media LLC","issue":"12","license":[{"start":{"date-parts":[[2025,7,24]],"date-time":"2025-07-24T00:00:00Z","timestamp":1753315200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2025,7,24]],"date-time":"2025-07-24T00:00:00Z","timestamp":1753315200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["2T32-AG 061892"],"award-info":[{"award-number":["2T32-AG 061892"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01AG074569"],"award-info":[{"award-number":["R01AG074569"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01AG074569"],"award-info":[{"award-number":["R01AG074569"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Mol Neurobiol"],"published-print":{"date-parts":[[2025,12]]},"DOI":"10.1007\/s12035-025-05231-7","type":"journal-article","created":{"date-parts":[[2025,7,24]],"date-time":"2025-07-24T14:53:40Z","timestamp":1753368820000},"page":"15759-15778","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Brain Region-Specific Accumulation of Amyloidosis-Associated Proteins in Postmortem Brain Tissues of Alzheimer\u2019s Disease Patients"],"prefix":"10.1007","volume":"62","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3271-1128","authenticated-orcid":false,"given":"Wangchen","family":"Tsering","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jennifer L.","family":"Philips","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Todd E.","family":"Golde","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-9450-4520","authenticated-orcid":false,"given":"Jonathan A.","family":"Villareal","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5633-2149","authenticated-orcid":false,"given":"Stefan","family":"Prokop","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,7,24]]},"reference":[{"key":"5231_CR1","doi-asserted-by":"publisher","first-page":"4195","DOI":"10.1242\/JCS.023820","volume":"123","author":"C Frantz","year":"2010","unstructured":"Frantz C, Stewart KM, Weaver VM (2010) The extracellular matrix at a glance. J Cell Sci 123:4195\u20134200. https:\/\/doi.org\/10.1242\/JCS.023820","journal-title":"J Cell Sci"},{"key":"5231_CR2","doi-asserted-by":"publisher","first-page":"1216","DOI":"10.1126\/SCIENCE.1176009","volume":"326","author":"RO Hynes","year":"2009","unstructured":"Hynes RO (2009) The extracellular matrix: not just pretty fibrils. Science 326:1216\u20131219. https:\/\/doi.org\/10.1126\/SCIENCE.1176009","journal-title":"Science"},{"key":"5231_CR3","doi-asserted-by":"publisher","first-page":"6850","DOI":"10.1111\/febs.15776","volume":"288","author":"NK Karamanos","year":"2021","unstructured":"Karamanos NK, Theocharis AD, Piperigkou Z, Manou D, Passi A, Skandalis SS, Vynios DH, Orian-Rousseau V et al (2021) A guide to the composition and functions of the extracellular matrix. FEBS J 288:6850\u20136912. https:\/\/doi.org\/10.1111\/febs.15776","journal-title":"FEBS J"},{"key":"5231_CR4","doi-asserted-by":"publisher","first-page":"865","DOI":"10.1038\/s41580-024-00767-3","volume":"25","author":"A Naba","year":"2024","unstructured":"Naba A (2024) Mechanisms of assembly and remodelling of the extracellular matrix. Nat Rev Mol Cell Biol 25:865\u2013885. https:\/\/doi.org\/10.1038\/s41580-024-00767-3","journal-title":"Nat Rev Mol Cell Biol"},{"key":"5231_CR5","doi-asserted-by":"publisher","first-page":"103","DOI":"10.1006\/EXNR.1996.0182","volume":"142","author":"DA Dewitt","year":"1996","unstructured":"Dewitt DA, Silver J (1996) Regenerative failure: a potential mechanism for neuritic dystrophy in Alzheimer\u2019s disease. Exp Neurol 142:103\u2013110. https:\/\/doi.org\/10.1006\/EXNR.1996.0182","journal-title":"Exp Neurol"},{"key":"5231_CR6","doi-asserted-by":"publisher","unstructured":"Snow AD, Cummings JA, Lake T (2021) The unifying hypothesis of Alzheimer\u2019s disease: heparan sulfate proteoglycans\/glycosaminoglycans are key as first hypothesized over 30 years ago. Front Aging Neurosci 13. https:\/\/doi.org\/10.3389\/FNAGI.2021.710683","DOI":"10.3389\/FNAGI.2021.710683"},{"key":"5231_CR7","first-page":"456","volume":"133","author":"AD Snow","year":"1988","unstructured":"Snow AD, Mar H, Nochlin D, Kimata K, Kato M, Suzuki S, Hassell J, Wight TN (1988) The presence of heparan sulfate proteoglycans in the neuritic plaques and congophilic angiopathy in Alzheimer\u2019s disease. Am J Pathol 133:456","journal-title":"Am J Pathol"},{"key":"5231_CR8","doi-asserted-by":"publisher","unstructured":"Bai B, Wang X, Li Y, Chen P, Yarbro JM, Beach TG, Peng J (2020) Deep multilayer brain proteomics identifies molecular networks and Netrin\u20101 accumulation in Alzheimer\u2019s disease progression. Alzheimer\u2019s & Dementia 16. https:\/\/doi.org\/10.1002\/alz.037231","DOI":"10.1002\/alz.037231"},{"key":"5231_CR9","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1038\/s41593-021-00999-y","volume":"25","author":"ECB Johnson","year":"2022","unstructured":"Johnson ECB, Carter EK, Dammer EB, Duong DM, Gerasimov ES, Liu Y, Liu J, Betarbet R et al (2022) Large-scale deep multi-layer analysis of Alzheimer\u2019s disease brain reveals strong proteomic disease-related changes not observed at the RNA level. Nat Neurosci 25:213\u2013225. https:\/\/doi.org\/10.1038\/s41593-021-00999-y","journal-title":"Nat Neurosci"},{"key":"5231_CR10","doi-asserted-by":"publisher","first-page":"769","DOI":"10.1038\/s41591-020-0815-6","volume":"26","author":"ECB Johnson","year":"2020","unstructured":"Johnson ECB, Dammer EB, Duong DM, Ping L, Zhou M, Yin L, Higginbotham LA, Guajardo A et al (2020) Large-scale proteomic analysis of Alzheimer\u2019s disease brain and cerebrospinal fluid reveals early changes in energy metabolism associated with microglia and astrocyte activation. Nat Med 26:769\u2013780. https:\/\/doi.org\/10.1038\/s41591-020-0815-6","journal-title":"Nat Med"},{"key":"5231_CR11","doi-asserted-by":"publisher","unstructured":"Levites Y, Dammer EB, Ran Y, Tsering W, Duong D, Abreha M, Gadhavi J, Lolo K, Trejo-Lopez J, Phillips J, Iturbe A, Erquizi A, Moore BD, Ryu D, Natu A, Dillon K, Torrellas J, Moran C, Ladd T, Afroz F, Islam T, Jagirdar J, Funk CC, Robinson M, Rangaraju S, Borchelt DR, Ertekin-Taner N, Kelly JW, Heppner FL, Johnson ECB, McFarland K, Levey AI, Prokop S, Seyfried NT, Golde TE (2024) Integrative proteomics identifies a conserved A\u03b2 amyloid responsome, novel plaque proteins, and pathology modifiers in Alzheimer\u2019s disease. Cell Rep Med 101669. https:\/\/doi.org\/10.1016\/J.XCRM.2024.101669","DOI":"10.1016\/J.XCRM.2024.101669"},{"key":"5231_CR12","doi-asserted-by":"publisher","first-page":"10843","DOI":"10.1073\/pnas.162228299","volume":"99","author":"RB DeMattos","year":"2002","unstructured":"DeMattos RB, O\u2019dell MA, Parsadanian M, Taylor JW, Harmony JAK, Bales KR, Paul SM, Aronow BJ et al (2002) Clusterin promotes amyloid plaque formation and is critical for neuritic toxicity in a mouse model of Alzheimer\u2019s disease. Proc Natl Acad Sci USA 99:10843\u201310848","journal-title":"Proc Natl Acad Sci USA"},{"key":"5231_CR13","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1006\/nbdi.2002.0483","volume":"9","author":"AM Fagan","year":"2002","unstructured":"Fagan AM, Watson M, Parsadanian M, Bales KR, Paul SM, Holtzman DM (2002) Human and murine apoE markedly alters A\u03b2 metabolism before and after plaque formation in a mouse model of Alzheimer\u2019s Disease. Neurobiol Dis 9:305\u2013318. https:\/\/doi.org\/10.1006\/nbdi.2002.0483","journal-title":"Neurobiol Dis"},{"key":"5231_CR14","doi-asserted-by":"publisher","first-page":"247","DOI":"10.1385\/JMN:23:3:247","volume":"23","author":"DM Holtzman","year":"2004","unstructured":"Holtzman DM (2004) In vivo effects of ApoE and clusterin on amyloid-beta metabolism and neuropathology. J Mol Neurosci 23:247\u2013254. https:\/\/doi.org\/10.1385\/JMN:23:3:247","journal-title":"J Mol Neurosci"},{"key":"5231_CR15","doi-asserted-by":"publisher","unstructured":"Bhattarai P, Gunasekaran TI, Belloy ME, Reyes-Dumeyer D, J\u00fclich D, Tayran H, Yilmaz E, Flaherty D, Turgutalp B, Sukumar G, Alba C, McGrath EM, Hupalo DN, Bacikova D, Le Guen Y, Lantigua R, Medrano M, Rivera D, Recio P, Nuriel T, Ertekin-Taner N, Teich AF, Dickson DW, Holley S, Greicius M, Dalgard CL, Zody M, Mayeux R, Kizil C, Vardarajan BN (2024) Rare genetic variation in fibronectin 1 (FN1) protects against APOE\u03b54 in Alzheimer\u2019s disease. Acta Neuropathol 147. https:\/\/doi.org\/10.1007\/S00401-024-02721-1","DOI":"10.1007\/S00401-024-02721-1"},{"key":"5231_CR16","doi-asserted-by":"publisher","first-page":"1243","DOI":"10.1038\/S41591-023-02318-3","volume":"29","author":"F Lopera","year":"2023","unstructured":"Lopera F, Marino C, Chandrahas AS, O\u2019Hare M, Villalba-Moreno ND, Aguillon D, Baena A, Sanchez JS, Vila-Castelar C, Ramirez Gomez L et al (2023) Resilience to autosomal dominant Alzheimer\u2019s disease in a Reelin-COLBOS heterozygous man. Nat Med 29:1243\u20131252. https:\/\/doi.org\/10.1038\/S41591-023-02318-3","journal-title":"Nat Med"},{"key":"5231_CR17","doi-asserted-by":"publisher","first-page":"2104","DOI":"10.1038\/S41593-023-01480-8","volume":"26","author":"MR Nelson","year":"2023","unstructured":"Nelson MR, Liu P, Agrawal A, Yip O, Blumenfeld J, Traglia M, Kim MJ, Koutsodendris N et al (2023) The APOE-R136S mutation protects against APOE4-driven Tau pathology, neurodegeneration and neuroinflammation. Nat Neurosci 26:2104\u20132121. https:\/\/doi.org\/10.1038\/S41593-023-01480-8","journal-title":"Nat Neurosci"},{"issue":"25","key":"5231_CR18","doi-asserted-by":"publisher","first-page":"620263","DOI":"10.1101\/2024.10.25.620263","volume":"10","author":"JM Yarbro","year":"2024","unstructured":"Yarbro JM, Han X, Dasgupta A, Yang K, Liu D, Shrestha HK, Zaman M, Wang Z et al (2024) Human-mouse proteomics reveals the shared pathways in Alzheimer\u2019s disease and delayed protein turnover in the amyloidome. bioRxiv 10(25):620263. https:\/\/doi.org\/10.1101\/2024.10.25.620263","journal-title":"bioRxiv"},{"key":"5231_CR19","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/S40478-023-01688-6\/FIGURES\/7","volume":"11","author":"W Tsering","year":"2023","unstructured":"Tsering W, Hery GP, Phillips JL, Lolo K, Bathe T, Villareal JA, Ruan IY, Prokop S (2023) Transformation of non-neuritic into neuritic plaques during AD progression drives cortical spread of tau pathology via regenerative failure. Acta Neuropathol Commun 11:1\u201320. https:\/\/doi.org\/10.1186\/S40478-023-01688-6\/FIGURES\/7","journal-title":"Acta Neuropathol Commun"},{"key":"5231_CR20","doi-asserted-by":"publisher","first-page":"975","DOI":"10.1016\/j.neuron.2019.12.015","volume":"105","author":"B Bai","year":"2020","unstructured":"Bai B, Wang X, Li Y, Chen PC, Yu K, Dey KK, Yarbro JM, Han X et al (2020) Deep multilayer brain proteomics identifies molecular networks in Alzheimer\u2019s disease progression. Neuron 105:975-991.e7. https:\/\/doi.org\/10.1016\/j.neuron.2019.12.015","journal-title":"Neuron"},{"key":"5231_CR21","doi-asserted-by":"publisher","first-page":"1310094","DOI":"10.3389\/FIMMU.2023.1310094","volume":"14","author":"EE Neumaier","year":"2023","unstructured":"Neumaier EE, Rothhammer V, Linnerbauer M (2023) The role of midkine in health and disease. Front Immunol 14:1310094. https:\/\/doi.org\/10.3389\/FIMMU.2023.1310094","journal-title":"Front Immunol"},{"key":"5231_CR22","doi-asserted-by":"publisher","unstructured":"Ross-Munro E, Kwa F, Kreiner J, Khore M, Miller SL, Tolcos M, Fleiss B, Walker DW (2020) Midkine: the who, what, where, and when of a promising neurotrophic therapy for perinatal brain injury. Front Neurol 11. https:\/\/doi.org\/10.3389\/FNEUR.2020.568814","DOI":"10.3389\/FNEUR.2020.568814"},{"key":"5231_CR23","doi-asserted-by":"publisher","first-page":"377","DOI":"10.1006\/GENO.1997.5199","volume":"48","author":"F Charbonnier","year":"1998","unstructured":"Charbonnier F, P\u00e9rin JP, Mattei MG, Camuzat A, Bonnet F, Gressin L, Alliel PM (1998) Genomic organization of the human SPOCK gene and its chromosomal localization to 5q31. Genomics 48:377\u2013380. https:\/\/doi.org\/10.1006\/GENO.1997.5199","journal-title":"Genomics"},{"key":"5231_CR24","doi-asserted-by":"publisher","first-page":"399","DOI":"10.1111\/JNC.12212","volume":"125","author":"U Hartmann","year":"2013","unstructured":"Hartmann U, H\u00fclsmann H, Seul J, R\u00f6ll S, Midani H, Breloy I, Hechler D, M\u00fcller R et al (2013) Testican-3: a brain-specific proteoglycan member of the BM-40\/SPARC\/osteonectin family. J Neurochem 125:399\u2013409. https:\/\/doi.org\/10.1111\/JNC.12212","journal-title":"J Neurochem"},{"key":"5231_CR25","doi-asserted-by":"publisher","first-page":"479","DOI":"10.1111\/J.1600-0838.2009.00941.X","volume":"19","author":"E Okina","year":"2009","unstructured":"Okina E, Manon-Jensen T, Whiteford JR, Couchman JR (2009) Syndecan proteoglycan contributions to cytoskeletal organization and contractility. Scand J Med Sci Sports 19:479\u2013489. https:\/\/doi.org\/10.1111\/J.1600-0838.2009.00941.X","journal-title":"Scand J Med Sci Sports"},{"key":"5231_CR26","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1016\/j.neuron.2015.11.013","volume":"89","author":"Y Zhang","year":"2016","unstructured":"Zhang Y, Sloan SA, Clarke LE, Caneda C, Plaza CA, Blumenthal PD, Vogel H, Steinberg GK et al (2016) Purification and characterization of progenitor and mature human astrocytes reveals transcriptional and functional differences with mouse. Neuron 89:37\u201353. https:\/\/doi.org\/10.1016\/j.neuron.2015.11.013","journal-title":"Neuron"},{"key":"5231_CR27","doi-asserted-by":"publisher","first-page":"409","DOI":"10.1016\/J.NEUROBIOLAGING.2008.04.009","volume":"31","author":"C Forsell","year":"2010","unstructured":"Forsell C, Bj\u00f6rk BF, Lilius L, Axelman K, Fabre SF, Fratiglioni L, Winblad B, Graff C (2010) Genetic association to the amyloid plaque associated protein gene COL25A1 in Alzheimer\u2019s disease. Neurobiol Aging 31:409\u2013415. https:\/\/doi.org\/10.1016\/J.NEUROBIOLAGING.2008.04.009","journal-title":"Neurobiol Aging"},{"key":"5231_CR28","doi-asserted-by":"publisher","first-page":"1524","DOI":"10.1093\/EMBOJ\/21.7.1524","volume":"21","author":"T Hashimoto","year":"2002","unstructured":"Hashimoto T, Wakabayashi T, Watanabe A, Kowa H, Hosoda R, Nakamura A, Kanazawa I, Arai T et al (2002) CLAC: a novel Alzheimer amyloid plaque component derived from a transmembrane precursor, CLAC-P\/collagen type XXV. EMBO J 21:1524\u20131534. https:\/\/doi.org\/10.1093\/EMBOJ\/21.7.1524","journal-title":"EMBO J"},{"key":"5231_CR29","doi-asserted-by":"publisher","unstructured":"Lim YS, Lee DY, Kim HY, Ok YJ, Hwang S, MooN Y, Yoon S (2021) Descriptive and functional characterization of epidermal growth factor-like domain 8 in mouse cortical thymic epithelial cells by integrated analysis of gene expression signatures and networks. Int J Mol Med 47. https:\/\/doi.org\/10.3892\/IJMM.2020.4837","DOI":"10.3892\/IJMM.2020.4837"},{"key":"5231_CR30","doi-asserted-by":"publisher","unstructured":"Malik R, Beaufort N, Frerich S, Gesierich B, Georgakis MK, Rannikm\u00e4 K, Ferguson AC, Haffner C, Traylor M, Ehrmann M, Sudlow CLM, Dichgans M Whole-exome sequencing reveals a role of HTRA1 and EGFL8 in brain white matter hyperintensities. https:\/\/doi.org\/10.1093\/brain\/awab253","DOI":"10.1093\/brain\/awab253"},{"key":"5231_CR31","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.jalz.2011.10.007","volume":"8","author":"BT Hyman","year":"2012","unstructured":"Hyman BT, Phelps CH, Beach TG, Bigio EH, Cairns NJ, Carrillo MC, Dickson DW, Duyckaerts C et al (2012) National Institute on Aging-Alzheimer\u2019s Association guidelines for the neuropathologic assessment of Alzheimer\u2019s disease. Alzheimers Dement 8:1\u201313. https:\/\/doi.org\/10.1016\/j.jalz.2011.10.007","journal-title":"Alzheimers Dement"},{"key":"5231_CR32","doi-asserted-by":"publisher","first-page":"88","DOI":"10.1212\/WNL.0000000000004058","volume":"89","author":"IG McKeith","year":"2017","unstructured":"McKeith IG, Boeve BF, Dickson DW, Halliday G, Taylor JP, Weintraub D, Aarsland D, Galvin J et al (2017) Diagnosis and management of dementia with Lewy bodies. Neurology 89:88\u2013100. https:\/\/doi.org\/10.1212\/WNL.0000000000004058","journal-title":"Neurology"},{"key":"5231_CR33","doi-asserted-by":"publisher","first-page":"1503","DOI":"10.1093\/BRAIN\/AWZ099","volume":"142","author":"PT Nelson","year":"2019","unstructured":"Nelson PT, Dickson DW, Trojanowski JQ, Jack CR, Boyle PA, Arfanakis K, Rademakers R, Alafuzoff I et al (2019) Limbic-predominant age-related TDP-43 encephalopathy (LATE): consensus working group report. Brain 142:1503\u20131527. https:\/\/doi.org\/10.1093\/BRAIN\/AWZ099","journal-title":"Brain"},{"key":"5231_CR34","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/S40478-024-01770-7","volume":"12","author":"T Bathe","year":"2024","unstructured":"Bathe T, Hery GP, Villareal JAB, Phillips JL, Cohen EM, Sharma RV, Tsering W, Prokop S (2024) Disease and brain region specific immune response profiles in neurodegenerative diseases with pure and mixed protein pathologies. Acta Neuropathologica Communications 12:1\u201325. https:\/\/doi.org\/10.1186\/S40478-024-01770-7","journal-title":"Acta Neuropathologica Communications"},{"key":"5231_CR35","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-017-17204-5","volume":"7","author":"P Bankhead","year":"2017","unstructured":"Bankhead P, Loughrey MB, Fern\u00e1ndez JA, Dombrowski Y, McArt DG, Dunne PD, McQuaid S, Gray RT et al (2017) QuPath: open source software for digital pathology image analysis. Sci Rep 7:1\u20137. https:\/\/doi.org\/10.1038\/s41598-017-17204-5","journal-title":"Sci Rep"},{"key":"5231_CR36","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1111\/J.1365-2818.2006.01706.X","volume":"224","author":"S Bolte","year":"2006","unstructured":"Bolte S, Cordeli\u00e8res FP (2006) A guided tour into subcellular colocalization analysis in light microscopy. J Microsc 224:213\u2013232. https:\/\/doi.org\/10.1111\/J.1365-2818.2006.01706.X","journal-title":"J Microsc"},{"key":"5231_CR37","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/S00401-011-0910-3","volume":"123","author":"TJ Montine","year":"2012","unstructured":"Montine TJ, Phelps CH, Beach TG, Bigio EH, Cairns NJ, Dickson DW, Duyckaerts C, Frosch MP et al (2012) National Institute on Aging-Alzheimer\u2019s Association guidelines for the neuropathologic assessment of Alzheimer\u2019s disease: a practical approach. Acta Neuropathol 123:1\u201311. https:\/\/doi.org\/10.1007\/S00401-011-0910-3","journal-title":"Acta Neuropathol"},{"key":"5231_CR38","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1172\/JCI25410","volume":"116","author":"Y Levites","year":"2006","unstructured":"Levites Y, Das P, Price RW, Rochette MJ, Kostura LA, McGowan EM, Murphy MP, Golde TE (2006) Anti-A\u03b242- and anti-A\u03b240-specific mAbs attenuate amyloid deposition in an Alzheimer disease mouse model. J Clin Investig 116:193. https:\/\/doi.org\/10.1172\/JCI25410","journal-title":"J Clin Investig"},{"key":"5231_CR39","doi-asserted-by":"publisher","first-page":"1791","DOI":"10.1212\/wnl.58.12.1791","volume":"58","author":"DR Thal","year":"2002","unstructured":"Thal DR, R\u00fcb U, Orantes M, Braak H (2002) Phases of A\u03b2-deposition in the human brain and its relevance for the development of AD. Neurology 58:1791\u20131800. https:\/\/doi.org\/10.1212\/wnl.58.12.1791","journal-title":"Neurology"},{"key":"5231_CR40","doi-asserted-by":"publisher","unstructured":"Xia Y, Prokop S, Gorion K-MM, Kim JD, Sorrentino ZA, Bell BM, Manaois AN, Chakrabarty P, Davies P, Giasson BI Tau Ser208 phosphorylation promotes aggregation and reveals neuropathologic diversity in Alzheimer\u2019s disease and other tauopathies. https:\/\/doi.org\/10.1186\/s40478-020-00967-w","DOI":"10.1186\/s40478-020-00967-w"},{"key":"5231_CR41","doi-asserted-by":"publisher","unstructured":"Dammer EB, Ping L, Duong DM, Modeste ES, Seyfried NT, Lah JJ, Levey AI, Johnson ECB (2022) Multi-platform proteomic analysis of Alzheimer\u2019s disease cerebrospinal fluid and plasma reveals network biomarkers associated with proteostasis and the matrisome. Alzheimers Res Ther 14. https:\/\/doi.org\/10.1186\/S13195-022-01113-5","DOI":"10.1186\/S13195-022-01113-5"},{"key":"5231_CR42","doi-asserted-by":"publisher","unstructured":"Drummond E, Kavanagh T, Pires G, Marta-Ariza M, Kanshin E, Nayak S, Faustin A, Berdah V, Ueberheide B, Wisniewski T (2022) The amyloid plaque proteome in early onset Alzheimer\u2019s disease and Down syndrome. Acta Neuropathol Commun 10. https:\/\/doi.org\/10.1186\/S40478-022-01356-1","DOI":"10.1186\/S40478-022-01356-1"},{"key":"5231_CR43","doi-asserted-by":"publisher","first-page":"353","DOI":"10.1126\/science.1072994","volume":"297","author":"J Hardy","year":"1979","unstructured":"Hardy J (1979) Selkoe DJ (2002) The amyloid hypothesis of Alzheimer\u2019s disease: progress and problems on the road to therapeutics. Science 297:353\u2013356. https:\/\/doi.org\/10.1126\/science.1072994","journal-title":"Science"},{"key":"5231_CR44","doi-asserted-by":"publisher","first-page":"595","DOI":"10.15252\/emmm.201606210","volume":"8","author":"DJ Selkoe","year":"2016","unstructured":"Selkoe DJ, Hardy J (2016) The amyloid hypothesis of Alzheimer\u2019s disease at 25 years. EMBO Mol Med 8:595\u2013608. https:\/\/doi.org\/10.15252\/emmm.201606210","journal-title":"EMBO Mol Med"},{"key":"5231_CR45","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/S12035-023-03736-7","volume":"1","author":"W Tsering","year":"2023","unstructured":"Tsering W (2023) Prokop S (2023) Neuritic plaques \u2014 gateways to understanding Alzheimer\u2019s disease. Mol Neurobiol 1:1\u201314. https:\/\/doi.org\/10.1007\/S12035-023-03736-7","journal-title":"Mol Neurobiol"},{"key":"5231_CR46","doi-asserted-by":"publisher","DOI":"10.1111\/JNC.16275","volume":"169","author":"W Tsering","year":"2024","unstructured":"Tsering W, de la Rosa A, Ruan IY, Philips JL, Bathe T, Villareal JA, Prokop S (2024) Preferential clustering of microglia and astrocytes around neuritic plaques during progression of Alzheimer\u2019s disease neuropathological changes. J Neurochem 169:e16275. https:\/\/doi.org\/10.1111\/JNC.16275","journal-title":"J Neurochem"},{"key":"5231_CR47","doi-asserted-by":"publisher","first-page":"5053","DOI":"10.1074\/JBC.M114.600569","volume":"290","author":"CB Jendresen","year":"2015","unstructured":"Jendresen CB, Cui H, Zhang X, Vlodavsky I, Nilsson LNG, Li JP (2015) Overexpression of heparanase lowers the amyloid burden in amyloid-\u03b2 precursor protein transgenic mice. J Biol Chem 290:5053\u20135064. https:\/\/doi.org\/10.1074\/JBC.M114.600569","journal-title":"J Biol Chem"},{"key":"5231_CR48","doi-asserted-by":"publisher","unstructured":"Liu CC, Zhao N, Yamaguchi Y, Cirrito JR, Kanekiyo T, Holtzman DM, Bu G (2016) Neuronal heparan sulfates promote amyloid pathology by modulating brain amyloid-\u03b2 clearance and aggregation in Alzheimer\u2019s disease. Sci Transl Med 8. https:\/\/doi.org\/10.1126\/SCITRANSLMED.AAD3650","DOI":"10.1126\/SCITRANSLMED.AAD3650"},{"key":"5231_CR49","doi-asserted-by":"publisher","first-page":"474","DOI":"10.1093\/JNEN\/NLAA016","volume":"79","author":"L Lorente-Gea","year":"2020","unstructured":"Lorente-Gea L, Garcia B, Martin C, Ordiales H, Garcia-Suarez O, Pina-Batista KM, Merayo-Lloves J, Quiros LM et al (2020) Heparan sulfate proteoglycans undergo differential expression alterations in Alzheimer disease brains. J Neuropathol Exp Neurol 79:474\u2013483. https:\/\/doi.org\/10.1093\/JNEN\/NLAA016","journal-title":"J Neuropathol Exp Neurol"},{"key":"5231_CR50","doi-asserted-by":"publisher","first-page":"C893","DOI":"10.1152\/AJPCELL.00247.2022","volume":"324","author":"I Ozsan McMillan","year":"2023","unstructured":"Ozsan McMillan I, Li J-P, Wang L (2023) Heparan sulfate proteoglycan in Alzheimer\u2019s disease: aberrant expression and functions in molecular pathways related to amyloid-\u03b2 metabolism. Am J Physiol Cell Physiol 324:C893\u2013C909. https:\/\/doi.org\/10.1152\/AJPCELL.00247.2022","journal-title":"Am J Physiol Cell Physiol"},{"key":"5231_CR51","doi-asserted-by":"publisher","unstructured":"Sun Y, Xu S, Jiang M, Liu X, Yang L, Bai Z, Yang Q (2021) Role of the extracellular matrix in Alzheimer\u2019s disease. Front Aging Neurosci 13. https:\/\/doi.org\/10.3389\/FNAGI.2021.707466","DOI":"10.3389\/FNAGI.2021.707466"},{"key":"5231_CR52","doi-asserted-by":"publisher","first-page":"2231","DOI":"10.1111\/J.1742-4658.2005.04647.X","volume":"272","author":"L S\u00f6derberg","year":"2005","unstructured":"S\u00f6derberg L, Dahlqvist C, Kakuyama H, Thyberg J, Ito A, Winblad B, N\u00e4slund J, Tjernberg LO (2005) Collagenous Alzheimer amyloid plaque component assembles amyloid fibrils into protease resistant aggregates. FEBS J 272:2231\u20132236. https:\/\/doi.org\/10.1111\/J.1742-4658.2005.04647.X","journal-title":"FEBS J"},{"key":"5231_CR53","doi-asserted-by":"publisher","unstructured":"Hashimoto T, Fujii D, Naka Y, Kashiwagi-Hakozaki M, Matsuo Y, Matsuura Y, Wakabayashi T, Iwatsubo T (2020) Collagenous Alzheimer amyloid plaque component impacts on the compaction of amyloid-\u03b2 plaques. Acta Neuropathol Commun 8. https:\/\/doi.org\/10.1186\/s40478-020-01075-5","DOI":"10.1186\/s40478-020-01075-5"},{"key":"5231_CR54","doi-asserted-by":"publisher","first-page":"381","DOI":"10.1159\/000363101","volume":"36","author":"A Yamamoto","year":"2014","unstructured":"Yamamoto A, Uchiyama K, Nara T, Nishimura N, Hayasaka M, Hanaoka K, Yamamoto T (2014) Structural abnormalities of corpus callosum and cortical axonal tracts accompanied by decreased anxiety-like behavior and lowered sociability in Spock3-mutant mice. Dev Neurosci 36:381\u2013395. https:\/\/doi.org\/10.1159\/000363101","journal-title":"Dev Neurosci"},{"key":"5231_CR55","doi-asserted-by":"publisher","first-page":"4499","DOI":"10.1002\/ALZ.13867","volume":"20","author":"S Oveisgharan","year":"2024","unstructured":"Oveisgharan S, Yu L, de Paiva LK, Tasaki S, Wang Y, Menon V, Schneider JA, Seyfried NT et al (2024) Proteins linking APOE \u025b4 with Alzheimer\u2019s disease. Alzheimers Dement 20:4499\u20134511. https:\/\/doi.org\/10.1002\/ALZ.13867","journal-title":"Alzheimers Dement"},{"key":"5231_CR56","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1007\/bf00308809","volume":"82","author":"H Braak","year":"1991","unstructured":"Braak H, Braak E (1991) Neuropathological stageing of Alzheimer-related changes. Acta Neuropathol 82:239\u2013259. https:\/\/doi.org\/10.1007\/bf00308809","journal-title":"Acta Neuropathol"},{"key":"5231_CR57","doi-asserted-by":"publisher","first-page":"960","DOI":"10.1097\/NEN.0B013E318232A379","volume":"70","author":"H Braak","year":"2011","unstructured":"Braak H, Thal DR, Ghebremedhin E, Del Tredici K (2011) Stages of the pathologic process in Alzheimer disease: age categories from 1 to 100 years. J Neuropathol Exp Neurol 70:960\u2013969. https:\/\/doi.org\/10.1097\/NEN.0B013E318232A379","journal-title":"J Neuropathol Exp Neurol"},{"key":"5231_CR58","doi-asserted-by":"publisher","first-page":"1707","DOI":"10.1212\/WNL.0000000000004521","volume":"89","author":"LM Besser","year":"2017","unstructured":"Besser LM, Crary JF, Mock C, Kukull WA (2017) Comparison of symptomatic and asymptomatic persons with primary age-related tauopathy. Neurology 89:1707\u20131715. https:\/\/doi.org\/10.1212\/WNL.0000000000004521","journal-title":"Neurology"},{"key":"5231_CR59","doi-asserted-by":"publisher","first-page":"774","DOI":"10.1097\/NEN.0B013E3181AACBE9","volume":"68","author":"PT Nelson","year":"2009","unstructured":"Nelson PT, Abner EL, Schmitt FA, Kryscio RJ, Jicha GA, Santacruz K, Smith CD, Patel E et al (2009) Brains with medial temporal lobe neurofibrillary tangles but no neuritic amyloid plaques are a diagnostic dilemma but may have pathogenetic aspects distinct from Alzheimer disease. J Neuropathol Exp Neurol 68:774\u2013784. https:\/\/doi.org\/10.1097\/NEN.0B013E3181AACBE9","journal-title":"J Neuropathol Exp Neurol"},{"key":"5231_CR60","doi-asserted-by":"publisher","first-page":"66","DOI":"10.1111\/j.1750-3639.2008.00244.x","volume":"20","author":"PT Nelson","year":"2010","unstructured":"Nelson PT, Abner EL, Schmitt FA, Kryscio RJ, Jicha GA, Smith CD, Davis DG, Poduska JW et al (2010) Modeling the association between 43 different clinical and pathological variables and the severity of cognitive impairment in a large autopsy cohort of elderly persons. Brain Pathol 20:66\u201379. https:\/\/doi.org\/10.1111\/j.1750-3639.2008.00244.x","journal-title":"Brain Pathol"},{"key":"5231_CR61","doi-asserted-by":"publisher","first-page":"428","DOI":"10.1016\/J.CELL.2023.11.029","volume":"187","author":"Y Chen","year":"2024","unstructured":"Chen Y, Song S, Parhizkar S, Lord J, Zhu Y, Strickland MR, Wang C, Park J et al (2024) APOE3ch alters microglial response and suppresses A\u03b2-induced tau seeding and spread. Cell 187:428-445.e20. https:\/\/doi.org\/10.1016\/J.CELL.2023.11.029","journal-title":"Cell"},{"key":"5231_CR62","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-018-37476-9","volume":"9","author":"T Letoha","year":"2019","unstructured":"Letoha T, Hud\u00e1k A, Kusz E, Pettk\u00f3-Szandtner A, Domonkos I, J\u00f3svay K, Hofmann-Apitius M, Szil\u00e1k L (2019) Contribution of syndecans to cellular internalization and fibrillation of amyloid-\u03b2(1\u201342). Scientific Reports 9:1\u201317. https:\/\/doi.org\/10.1038\/s41598-018-37476-9","journal-title":"Scientific Reports"},{"key":"5231_CR63","doi-asserted-by":"publisher","first-page":"P1140","DOI":"10.1016\/J.JALZ.2018.06.1550","volume":"14","author":"C-LG Lin","year":"2018","unstructured":"Lin C-LG, Foster JB, Zhao F, Lashley R, Hodgetts K, Lin L, Abreha M, Dammer EB et al (2018) P3\u2013192: novel targets for blocking the uptake of tau oligomers in hiPSC neurons. Alzheimer\u2019s Dementia 14:P1140\u2013P1141. https:\/\/doi.org\/10.1016\/J.JALZ.2018.06.1550","journal-title":"Alzheimer\u2019s Dementia"},{"key":"5231_CR64","doi-asserted-by":"publisher","first-page":"2803","DOI":"10.1093\/BRAIN\/AWAA223","volume":"143","author":"E Drummond","year":"2020","unstructured":"Drummond E, Pires G, MacMurray C, Askenazi M, Nayak S, Bourdon M, Safar J, Ueberheide B et al (2020) Phosphorylated tau interactome in the human Alzheimer\u2019s disease brain. Brain 143:2803\u20132817. https:\/\/doi.org\/10.1093\/BRAIN\/AWAA223","journal-title":"Brain"},{"key":"5231_CR65","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/S13024-022-00572-6","volume":"17","author":"T Kavanagh","year":"2022","unstructured":"Kavanagh T, Halder A, Drummond E (2022) Tau interactome and RNA binding proteins in neurodegenerative diseases. Mol Neurodegen 17:1\u201317. https:\/\/doi.org\/10.1186\/S13024-022-00572-6","journal-title":"Mol Neurodegen"},{"key":"5231_CR66","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/S00401-024-02744-8\/FIGURES\/3","volume":"147","author":"M Thierry","year":"2024","unstructured":"Thierry M, Ponce J, Mart\u00e0-Ariza M, Askenazi M, Faustin A, Leitner D, Pires G, Kanshin E, Drummond E, Ueberheide B, Wisniewski T (2024) The influence of APOE\u03b54 on the pTau interactome in sporadic Alzheimer\u2019s disease. Acta Neuropathol 147:1\u201319. https:\/\/doi.org\/10.1007\/S00401-024-02744-8\/FIGURES\/3","journal-title":"Acta Neuropathol"},{"key":"5231_CR67","doi-asserted-by":"publisher","first-page":"330","DOI":"10.1097\/00005072-198905000-00093","volume":"48","author":"C Joachim","year":"1989","unstructured":"Joachim C, Morris J, Platt D, Selkoe D (1989) Diffuse senile plaques. J Neuropathol Exp Neurol 48:330. https:\/\/doi.org\/10.1097\/00005072-198905000-00093","journal-title":"J Neuropathol Exp Neurol"},{"key":"5231_CR68","first-page":"337","volume":"144","author":"AD Snow","year":"1994","unstructured":"Snow AD, Sekiguchi RT, Nochlin D, Kalaria RN, Kimata K (1994) Heparan sulfate proteoglycan in diffuse plaques of hippocampus but not of cerebellum in Alzheimer\u2019s disease brain. Am J Pathol 144:337","journal-title":"Am J Pathol"},{"key":"5231_CR69","doi-asserted-by":"publisher","first-page":"5143","DOI":"10.1002\/ALZ.13859","volume":"20","author":"CR Jack","year":"2024","unstructured":"Jack CR, Andrews JS, Beach TG, Buracchio T, Dunn B, Graf A, Hansson O, Ho C et al (2024) Revised criteria for diagnosis and staging of Alzheimer\u2019s disease: Alzheimer\u2019s Association Workgroup. Alzheimers Dement 20:5143\u20135169. https:\/\/doi.org\/10.1002\/ALZ.13859","journal-title":"Alzheimers Dement"},{"key":"5231_CR70","doi-asserted-by":"publisher","first-page":"535","DOI":"10.1016\/J.JALZ.2018.02.018","volume":"14","author":"CR Jack","year":"2018","unstructured":"Jack CR, Bennett DA, Blennow K, Carrillo MC, Dunn B, Haeberlein SB, Holtzman DM, Jagust W et al (2018) NIA-AA research framework: toward a biological definition of Alzheimer\u2019s disease. Alzheimers Dement 14:535. https:\/\/doi.org\/10.1016\/J.JALZ.2018.02.018","journal-title":"Alzheimers Dement"},{"key":"5231_CR71","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1016\/J.JALZ.2018.03.002","volume":"14","author":"DS Knopman","year":"2018","unstructured":"Knopman DS, Haeberlein SB, Carrillo MC, Hendrix JA, Kerchner G, Margolin R, Maruff P, Miller DS et al (2018) The National Institute on Aging and the Alzheimer\u2019s Association Research Framework for Alzheimer\u2019s disease: perspectives from the research roundtable. Alzheimers Dement 14:563. https:\/\/doi.org\/10.1016\/J.JALZ.2018.03.002","journal-title":"Alzheimers Dement"},{"key":"5231_CR72","doi-asserted-by":"publisher","first-page":"576","DOI":"10.1016\/J.JALZ.2018.03.004","volume":"14","author":"N Silverberg","year":"2018","unstructured":"Silverberg N, Elliott C, Ryan L, Masliah E, Hodes R (2018) NIA commentary on the NIA-AA research framework: towards a biological definition of Alzheimer\u2019s disease. Alzheimers Dement 14:576\u2013578. https:\/\/doi.org\/10.1016\/J.JALZ.2018.03.004","journal-title":"Alzheimers Dement"},{"key":"5231_CR73","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/S00401-024-02819-6","volume":"148","author":"K Balcomb","year":"2024","unstructured":"Balcomb K, Johnston C, Kavanagh T, Leitner D, Schneider J, Halliday G, Wisniewski T, Sunde M et al (2024) SMOC1 colocalizes with Alzheimer\u2019s disease neuropathology and delays A\u03b2 aggregation. Acta Neuropathologica 148:1\u201323. https:\/\/doi.org\/10.1007\/S00401-024-02819-6","journal-title":"Acta Neuropathologica"},{"key":"5231_CR74","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1016\/J.NEUROBIOLAGING.2021.04.012","volume":"105","author":"BC Carlyle","year":"2021","unstructured":"Carlyle BC, Kandigian SE, Kreuzer J, Das S, Trombetta BA, Kuo Y, Bennett DA, Schneider JA et al (2021) Synaptic proteins associated with cognitive performance and neuropathology in older humans revealed by multiplexed fractionated proteomics. Neurobiol Aging 105:99\u2013114. https:\/\/doi.org\/10.1016\/J.NEUROBIOLAGING.2021.04.012","journal-title":"Neurobiol Aging"},{"key":"5231_CR75","doi-asserted-by":"publisher","unstructured":"Higginbotham L, Ping L, Dammer EB, Duong DM, Zhou M, Gearing M, Hurst C, Glass JD, Factor SA, Johnson ECB, Hajjar I, Lah JJ, Levey AI, Seyfried NT (2020) Integrated proteomics reveals brain-based cerebrospinal fluid biomarkers in asymptomatic and symptomatic Alzheimer\u2019s disease. Sci Adv 6. https:\/\/doi.org\/10.1126\/SCIADV.AAZ9360","DOI":"10.1126\/SCIADV.AAZ9360"},{"key":"5231_CR76","doi-asserted-by":"publisher","unstructured":"Kamalian A, Ho SG, Patel M, Lewis A, Bakker A, Albert M, O\u2019Brien RJ, Moghekar A, Lutz MW (2023) Exploratory assessment of proteomic network changes in cerebrospinal fluid of mild cognitive impairment patients: a pilot study. Biomolecules 13. https:\/\/doi.org\/10.3390\/BIOM13071094","DOI":"10.3390\/BIOM13071094"}],"container-title":["Molecular Neurobiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s12035-025-05231-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s12035-025-05231-7\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s12035-025-05231-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,27]],"date-time":"2025-10-27T13:11:49Z","timestamp":1761570709000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s12035-025-05231-7"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,7,24]]},"references-count":76,"journal-issue":{"issue":"12","published-print":{"date-parts":[[2025,12]]}},"alternative-id":["5231"],"URL":"https:\/\/doi.org\/10.1007\/s12035-025-05231-7","relation":{"has-preprint":[{"id-type":"doi","id":"10.21203\/rs.3.rs-6649354\/v1","asserted-by":"object"}]},"ISSN":["0893-7648","1559-1182"],"issn-type":[{"value":"0893-7648","type":"print"},{"value":"1559-1182","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,7,24]]},"assertion":[{"value":"12 May 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 July 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"24 July 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"All protocols were approved by the University of Florida Institutional Review Board (IRB201600067).","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethical approval"}},{"value":"All the patients or their next-of-kin gave informed consent for the brain donation and use a specimen for research use.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Informed consent"}},{"value":"The authors declare no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Conflict of interest"}}]}}