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The common hallmark of all of these diseases is tau pathology, but its connection with disease progress has not been completely understood so far. Therefore, uncovering novel tau-interacting partners and pathology affected molecular pathways can reveal the causes of diseases as well as potential targets for the development of AD treatment. Despite the large number of known tau-interacting partners, a limited number of studies focused on in vivo tau interactions in disease or healthy conditions are available. Here, we applied an in vivo cross-linking approach, capable of capturing weak and transient protein\u2013protein interactions, to a unique transgenic rat model of progressive tau pathology similar to human AD. We have identified 175 potential novel and known tau-interacting proteins by MALDI-TOF mass spectrometry. Several of the most promising candidates for possible drug development were selected for validation by coimmunoprecipitation and colocalization experiments in animal and cellular models. Three proteins, Baiap2, Gpr37l1, and Nptx1, were confirmed as novel tau-interacting partners, and on the basis of their known functions and implications in neurodegenerative or psychiatric disorders, we proposed their potential role in tau pathology.<\/jats:p>","DOI":"10.1021\/acs.jproteome.0c00137","type":"journal-article","created":{"date-parts":[[2020,5,1]],"date-time":"2020-05-01T19:02:50Z","timestamp":1588359770000},"page":"2429-2442","source":"Crossref","is-referenced-by-count":42,"title":["Physiological\nTau Interactome in Brain and Its Link\nto Tauopathies"],"prefix":"10.1021","volume":"19","author":[{"given":"Jakub","family":"Sinsky","sequence":"first","affiliation":[{"name":"Slovak Academy of Sciences , , Dubravska cesta 9 , ,","place":["Bratislava, Slovakia, 84510"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Petra","family":"Majerova","sequence":"additional","affiliation":[{"name":"Slovak Academy of Sciences , , Dubravska cesta 9 , ,","place":["Bratislava, Slovakia, 84510"]},{"name":"AXON Neuroscience R&D Services SE , Dvorakovo nabrezie 10 , ,","place":["Bratislava, Slovakia, 811\u202f02"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrej","family":"Kovac","sequence":"additional","affiliation":[{"name":"Slovak Academy of Sciences , , Dubravska cesta 9 , ,","place":["Bratislava, Slovakia, 84510"]},{"name":"AXON Neuroscience R&D Services SE , Dvorakovo nabrezie 10 , ,","place":["Bratislava, Slovakia, 811\u202f02"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Max","family":"Kotlyar","sequence":"additional","affiliation":[{"name":"Krembil Research Institute, UHN , 60 Leonard Avenue , , ,","place":["Toronto, Ontario, Canada, M5T\u202f0S8"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Igor","family":"Jurisica","sequence":"additional","affiliation":[{"name":"Slovak Academy of Sciences , , Dubravska cesta 9 , ,","place":["Bratislava, Slovakia, 84510"]},{"name":"Krembil Research Institute, UHN , 60 Leonard Avenue , , ,","place":["Toronto, Ontario, Canada, M5T\u202f0S8"]},{"name":"University of Toronto , , 27 King\u2019s College Circle , , ,","place":["Toronto, Ontario, Canada, ON\u202fM5S"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5427-2042","authenticated-orcid":true,"given":"Jozef","family":"Hanes","sequence":"additional","affiliation":[{"name":"Slovak Academy of Sciences , , Dubravska cesta 9 , ,","place":["Bratislava, Slovakia, 84510"]},{"name":"AXON Neuroscience R&D Services SE , Dvorakovo nabrezie 10 , ,","place":["Bratislava, Slovakia, 811\u202f02"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2020,5,1]]},"reference":[{"key":"2026081416154204100_cit1","first-page":"492","article-title":"Building the\nNeuronal Microtubule Cytoskeleton","volume-title":"Neuron","author":"Kapitein","year":"2015"},{"key":"2026081416154204100_cit2","first-page":"693","article-title":"Hydrated Autoclave\nPretreatment Enhances Tau Immunoreactivity in Formalin-Fixed Normal\nand Alzheimer\u2019s Disease Brain Tissues","volume":"64","author":"Shin","year":"1991","journal-title":"Lab. Invest."},{"key":"2026081416154204100_cit3","doi-asserted-by":"publisher","first-page":"813","DOI":"10.1007\/s00702-004-0221-0","article-title":"Post-Translational\nModifications of Tau Protein in Alzheimer\u2019s Disease","volume":"112","author":"Gong","year":"2005","journal-title":"J. Neural Transm."},{"key":"2026081416154204100_cit4","doi-asserted-by":"publisher","first-page":"458","DOI":"10.1016\/j.neuint.2010.12.023","article-title":"Post-Translational\nModifications\nof Tau Protein: Implications for Alzheimer\u2019s Disease","volume":"58","author":"Martin","year":"2011","journal-title":"Neurochem. Int."},{"key":"2026081416154204100_cit5","first-page":"346","article-title":"Post-Translational\nModifications of Tau Protein","volume":"107","author":"Pevalova","year":"2006","journal-title":"Bratisl. Lek.\nListy"},{"key":"2026081416154204100_cit6","first-page":"354","article-title":"Intrinsically\nDisordered Tau Protein in Alzheimer\u2019s Tangles: A Coincidence\nor a Rule?","volume":"107","author":"Skrabana","year":"2006","journal-title":"Bratisl. Lek. Listy"},{"key":"2026081416154204100_cit7","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1007\/BF00308809","article-title":"Neuropathological Stageing\nof Alzheimer-Related Changes","volume":"82","author":"Braak","year":"1991","journal-title":"Acta Neuropathol."},{"key":"2026081416154204100_cit8","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1016\/0197-4580(95)00021-6","article-title":"Staging of Alzheimer\u2019s\nDisease-Related Neurofibrillary Changes","volume":"16","author":"Braak","year":"1995","journal-title":"Neurobiol.\nAging"},{"key":"2026081416154204100_cit9","doi-asserted-by":"publisher","first-page":"656","DOI":"10.2174\/156720510793611592","article-title":"Tau in Alzheimer Disease and Related\nTauopathies","volume":"7","author":"Iqbal","year":"2010","journal-title":"Curr. Alzheimer Res."},{"key":"2026081416154204100_cit10","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1038\/nrneurol.2018.9","article-title":"Imaging\nTau and Amyloid-\u03b2 Proteinopathies in Alzheimer Disease and Other\nConditions","volume":"14","author":"Villemagne","year":"2018","journal-title":"Nat. Rev. Neurol."},{"key":"2026081416154204100_cit11","doi-asserted-by":"publisher","first-page":"277","DOI":"10.1002\/humu.20086","article-title":"The Role of\nTau (MAPT) in Frontotemporal\nDementia and Related Tauopathies","volume":"24","author":"Rademakers","year":"2004","journal-title":"Hum. Mutat."},{"key":"2026081416154204100_cit12","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1016\/0304-3940(90)90829-X","article-title":"Constant Neurofibrillary\nChanges in the Neocortex in\nProgressive Supranuclear Palsy. Basic Differences with Alzheimer\u2019s\nDisease and Aging","volume":"119","author":"Hauw","year":"1990","journal-title":"Neurosci. Lett."},{"key":"2026081416154204100_cit13","doi-asserted-by":"publisher","first-page":"72","DOI":"10.1002\/ana.10793","article-title":"Identification of Amino-Terminally Cleaved\nTau Fragments That Distinguish Progressive Supranuclear Palsy from\nCorticobasal Degeneration","volume":"55","author":"Arai","year":"2004","journal-title":"Ann. Neurol."},{"key":"2026081416154204100_cit14","doi-asserted-by":"publisher","first-page":"736","DOI":"10.1016\/S1474-4422(04)00936-6","article-title":"Corticobasal Degeneration","volume":"3","author":"Mahapatra","year":"2004","journal-title":"Lancet\nNeurol."},{"key":"2026081416154204100_cit15","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1038\/s41586-018-0454-y","article-title":"Structures of Filaments from Pick\u2019s\nDisease Reveal a Novel Tau Protein Fold","volume":"561","author":"Falcon","year":"2018","journal-title":"Nature"},{"key":"2026081416154204100_cit16","doi-asserted-by":"publisher","first-page":"1211","DOI":"10.1016\/S0140-6736(86)92212-9","article-title":"Filamentous\nAggregates in Pick\u2019s Disease, Progressive Supranuclear Palsy,\nand Alzheimer\u2019s Disease Share Antigenic Determinants with Microtubule-Associated\nProtein, Tau","volume":"328","author":"Pollock","year":"1986","journal-title":"Lancet"},{"key":"2026081416154204100_cit17","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1016\/S0072-9752(07)01251-1","article-title":"Argyrophilic Grain Disease","volume":"89","author":"Tolnay","year":"2008","journal-title":"Handbook\nof Clinical Neurology."},{"key":"2026081416154204100_cit18","doi-asserted-by":"publisher","first-page":"4671","DOI":"10.3390\/ijms15034671","article-title":"Tau Protein\nModifications and Interactions: Their Role in Function and Dysfunction","volume":"15","author":"Mietelska-Porowska","year":"2014","journal-title":"Int. J. Mol. Sci."},{"key":"2026081416154204100_cit19","doi-asserted-by":"publisher","first-page":"681","DOI":"10.1042\/BST20120015","article-title":"The Self-Perpetuating Tau Truncation Circle","volume":"40","author":"Zilka","year":"2012","journal-title":"Biochem. Soc. 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Methods"},{"key":"2026081416154204100_cit24","doi-asserted-by":"publisher","first-page":"D754","DOI":"10.1093\/nar\/gkx1098","article-title":"Ensembl 2018","volume":"46","author":"Zerbino","year":"2018","journal-title":"Nucleic\nAcids Res."},{"key":"2026081416154204100_cit25","doi-asserted-by":"publisher","first-page":"D1228","DOI":"10.1093\/nar\/gks1147","article-title":"InnateDB:\nSystems Biology of Innate Immunity and beyond\ue5f8Recent Updates\nand Continuing Curation","volume":"41","author":"Breuer","year":"2013","journal-title":"Nucleic Acids Res."},{"key":"2026081416154204100_cit26","doi-asserted-by":"publisher","first-page":"D767","DOI":"10.1093\/nar\/gkn892","article-title":"Human Protein Reference Database--2009 Update","volume":"37","author":"Keshava\nPrasad","year":"2009","journal-title":"Nucleic Acids Res."},{"key":"2026081416154204100_cit27","doi-asserted-by":"publisher","first-page":"R95","DOI":"10.1186\/gb-2007-8-5-r95","article-title":"Unequal Evolutionary Conservation of Human Protein\nInteractions in Interologous Networks","volume":"8","author":"Brown","year":"2007","journal-title":"Genome\nBiol."},{"key":"2026081416154204100_cit28","doi-asserted-by":"publisher","first-page":"951","DOI":"10.1038\/nbt1103","article-title":"Probabilistic\nModel of the Human Protein-Protein Interaction\nNetwork","volume":"23","author":"Rhodes","year":"2005","journal-title":"Nat. Biotechnol."},{"key":"2026081416154204100_cit29","doi-asserted-by":"publisher","first-page":"42","DOI":"10.1007\/978-3-540-73060-6_4","article-title":"A Context-Specific Network of Protein-DNA\nand Protein-Protein\nInteractions Reveals New Regulatory Motifs in Human B Cells","volume":"4532 LNBI","author":"Lefebvre","year":"2007","journal-title":"In Lecture Notes in Computer Science (including subseries\nLecture Notes in Artificial Intelligence and Lecture Notes in Bioinformatics)"},{"key":"2026081416154204100_cit30","doi-asserted-by":"publisher","first-page":"M111.010629","DOI":"10.1074\/mcp.M111.010629","article-title":"Large-Scale de Novo\nPrediction of Physical Protein-Protein Association","volume":"10","author":"Elefsinioti","year":"2011","journal-title":"Mol. Cell. Proteomics"},{"key":"2026081416154204100_cit31","doi-asserted-by":"publisher","first-page":"556","DOI":"10.1038\/nature11503","article-title":"Structure-Based Prediction of Protein-Protein Interactions\non a Genome-Wide Scale","volume":"490","author":"Zhang","year":"2012","journal-title":"Nature"},{"key":"2026081416154204100_cit32","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1111\/j.1460-9568.2008.06329.x","article-title":"Truncated\nTau Expression Levels Determine\nLife Span of a Rat Model of Tauopathy without Causing Neuronal Loss\nor Correlating with Terminal Neurofibrillary Tangle Load","volume":"28","author":"Koson","year":"2008","journal-title":"Eur. J. Neurosci."},{"key":"2026081416154204100_cit33","doi-asserted-by":"publisher","first-page":"3582","DOI":"10.1016\/j.febslet.2006.05.029","article-title":"Truncated Tau from Sporadic Alzheimer\u2019s\nDisease Suffices to\nDrive Neurofibrillary Degeneration in Vivo","volume":"580","author":"Zilka","year":"2006","journal-title":"FEBS Lett."},{"key":"2026081416154204100_cit34","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1016\/j.brainres.2006.10.085","article-title":"Neurodegeneration\nCaused by Expression of Human Truncated Tau Leads to Progressive Neurobehavioural\nImpairment in Transgenic Rats","volume":"1130","author":"Hrnkova","year":"2007","journal-title":"Brain Res."},{"key":"2026081416154204100_cit35","doi-asserted-by":"publisher","first-page":"724","DOI":"10.1038\/nbt969","article-title":"Time-Controlled\nTranscardiac Perfusion Cross-Linking for the Study of Protein Interactions\nin Complex Tissues","volume":"22","author":"Schmitt-Ulms","year":"2004","journal-title":"Nat. Biotechnol."},{"key":"2026081416154204100_cit36","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1074\/mcp.M700077-MCP200","article-title":"The\nin Vivo Brain Interactome\nof the Amyloid Precursor Protein","volume":"7","author":"Bai","year":"2008","journal-title":"Mol. Cell.\nProteomics"},{"key":"2026081416154204100_cit37","doi-asserted-by":"publisher","first-page":"D1145","DOI":"10.1093\/nar\/gkz984","article-title":"The\nProteomeXchange Consortium in 2020: Enabling \u2018Big Data\u2019\nApproaches in Proteomics","volume":"48","author":"Deutsch","year":"2019","journal-title":"Nucleic Acids Res."},{"key":"2026081416154204100_cit38","doi-asserted-by":"publisher","first-page":"161","DOI":"10.3233\/JAD-2010-101434","article-title":"Hyperphosphorylated Truncated Protein\nTau Induces Caspase-3 Independent Apoptosis-Like Pathway in the Alzheimer\u2019s\nDisease Cellular Model","volume":"23","author":"Zilkova","year":"2011","journal-title":"J. Alzheimer's\nDis."},{"key":"2026081416154204100_cit39","doi-asserted-by":"publisher","first-page":"D581","DOI":"10.1093\/nar\/gky1037","article-title":"IID 2018 Update: Context-Specific\nPhysical Protein-Protein Interactions in Human, Model Organisms and\nDomesticated Species","volume":"47","author":"Kotlyar","year":"2019","journal-title":"Nucleic Acids Res."},{"key":"2026081416154204100_cit40","doi-asserted-by":"publisher","first-page":"3327","DOI":"10.1093\/bioinformatics\/btp595","article-title":"NAViGaTOR: Network\nAnalysis, Visualization and Graphing Toronto","volume":"25","author":"Brown","year":"2009","journal-title":"Bioinformatics"},{"key":"2026081416154204100_cit41","doi-asserted-by":"publisher","first-page":"D479","DOI":"10.1093\/nar\/gkz989","article-title":"PathDIP 4:\nAn Extended Pathway Annotations and Enrichment Analysis Resource for\nHuman, Model Organisms and Domesticated Species","volume":"48","author":"Rahmati","year":"2019","journal-title":"Nucleic Acids Res."},{"key":"2026081416154204100_cit42","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1016\/j.nbt.2019.05.003","article-title":"The Impact\nof Crosslinking and Non-Crosslinking Fixatives on Antigen Retrieval\nand Immunohistochemistry","volume":"52","author":"Stumptner","year":"2019","journal-title":"New Biotechnol."},{"key":"2026081416154204100_cit43","doi-asserted-by":"publisher","first-page":"421","DOI":"10.1016\/j.jasms.2009.11.006","article-title":"Digestion Completeness\nof Microwave-Assisted and Conventional Trypsin-Catalyzed Reactions","volume":"21","author":"Reddy","year":"2010","journal-title":"J. Am. Soc. Mass Spectrom."},{"key":"2026081416154204100_cit44","doi-asserted-by":"publisher","first-page":"699","DOI":"10.1002\/jms.1415","article-title":"Utility of\nFormaldehyde Cross-Linking\nand Mass Spectrometry in the Study of Protein-Protein Interactions","volume":"43","author":"Sutherland","year":"2008","journal-title":"J. Mass Spectrom."},{"key":"2026081416154204100_cit45","first-page":"11.4.1","article-title":"Chemical Cleavage\nof Proteins in Solution","volume-title":"Current Protocols\nin Protein Science","author":"Crimmins","year":"2005"},{"key":"2026081416154204100_cit46","doi-asserted-by":"publisher","first-page":"24","DOI":"10.1159\/000283478","article-title":"Transition\nof Tau Protein from Disordered\nto Misordered in Alzheimer\u2019s Disease","volume":"7","author":"Kovacech","year":"2010","journal-title":"Neurodegener. Dis."},{"key":"2026081416154204100_cit47","doi-asserted-by":"crossref","first-page":"3737","DOI":"10.1242\/jcs.113.21.3737","article-title":"C-Terminal Inhibition of Tau Assembly\nin Vitro and in Alzheimer\u2019s Disease","volume":"113","author":"Abraha","year":"2000","journal-title":"J. Cell Sci."},{"key":"2026081416154204100_cit48","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41467-018-03509-0","article-title":"A Common Mechanism of Proteasome\nImpairment by Neurodegenerative Disease-Associated Oligomers","volume":"9","author":"Thibaudeau","year":"2018","journal-title":"Nat. Commun."},{"key":"2026081416154204100_cit49","doi-asserted-by":"publisher","first-page":"46","DOI":"10.1038\/nm.4011","article-title":"Tau-Driven\n26S Proteasome\nImpairment and Cognitive Dysfunction Can Be Prevented Early in Disease\nby Activating CAMP-PKA Signaling","volume":"22","author":"Myeku","year":"2016","journal-title":"Nat. Med."},{"key":"2026081416154204100_cit50","doi-asserted-by":"publisher","first-page":"D833","DOI":"10.1093\/nar\/gkw943","article-title":"DisGeNET:\nA Comprehensive Platform\nIntegrating Information on Human Disease-Associated Genes and Variants","volume":"45","author":"Pin\u0303ero","year":"2017","journal-title":"Nucleic Acids Res."},{"key":"2026081416154204100_cit51","doi-asserted-by":"publisher","first-page":"8173","DOI":"10.1074\/jbc.M115.641902","article-title":"Co-Immunoprecipitation\nwith Tau Isoform-Specific Antibodies Reveals Distinct Protein Interactions\nand Highlights a Putative Role for 2N Tau in Disease","volume":"291","author":"Liu","year":"2016","journal-title":"J. Biol. Chem."},{"key":"2026081416154204100_cit52","doi-asserted-by":"publisher","first-page":"3845","DOI":"10.1002\/pmic.200400856","article-title":"Identification of Protein-Protein Interactions Usingin\nVivo Cross-Linking and Mass Spectrometry","volume":"4","author":"Vasilescu","year":"2004","journal-title":"Proteomics"},{"key":"2026081416154204100_cit53","doi-asserted-by":"publisher","first-page":"234","DOI":"10.1074\/mcp.M500198-MCP200","article-title":"Identification of VCP\/P97,\nCarboxyl Terminus of Hsp70-Interacting Protein (CHIP), and Amphiphysin\nII Interaction Partners Using Membrane-Based Human Proteome Arrays","volume":"5","author":"Grelle","year":"2006","journal-title":"Mol. Cell. Proteomics"},{"key":"2026081416154204100_cit54","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1126\/scisignal.aad1089","article-title":"Metalloprotease\nCleavage of the N Terminus of the Orphan G Protein-Coupled Receptor\nGPR37L1 Reduces Its Constitutive Activity","volume":"9","author":"Coleman","year":"2016","journal-title":"Sci.\nSignaling"},{"key":"2026081416154204100_cit55","doi-asserted-by":"publisher","first-page":"3000","DOI":"10.1074\/mcp.M115.050724","article-title":"The Human\nTau Interactome: Binding to the Ribonucleoproteome, and Impaired Binding\nof the Proline-to-Leucine Mutant at Position 301 (P301L) to Chaperones\nand the Proteasome","volume":"14","author":"Gunawardana","year":"2015","journal-title":"Mol. Cell. 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