{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,30]],"date-time":"2026-10-30T03:12:46Z","timestamp":1793329966865,"version":"4.5.0"},"reference-count":90,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,10,6]],"date-time":"2025-10-06T00:00:00Z","timestamp":1759708800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,10,6]],"date-time":"2025-10-06T00:00:00Z","timestamp":1759708800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/100000864","name":"Michael J. Fox Foundation for Parkinson's Research","doi-asserted-by":"publisher","award":["021129"],"award-info":[{"award-number":["021129"]}],"id":[{"id":"10.13039\/100000864","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000024","name":"Gouvernement du Canada | Canadian Institutes of Health Research","doi-asserted-by":"publisher","award":["MFE-152571"],"award-info":[{"award-number":["MFE-152571"]}],"id":[{"id":"10.13039\/501100000024","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000024","name":"Gouvernement du Canada | Canadian Institutes of Health Research","doi-asserted-by":"publisher","award":["Canada Research Chair (Tier 1) in Parkinson\u2019s disease"],"award-info":[{"award-number":["Canada Research Chair (Tier 1) in Parkinson\u2019s disease"]}],"id":[{"id":"10.13039\/501100000024","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Canada First Research Excellence Fund, Healthy Brain, Healthy Lives awarded to McGill University"},{"name":"PROTEO"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Commun Biol"],"abstract":"<jats:title>Abstract<\/jats:title>\n          <jats:p>Synucleinopathies are characterized by the accumulation and propagation of \u03b1-synuclein (\u03b1-syn) aggregates throughout the brain, leading to neuronal dysfunction and death. In this study, we used an unbiased FACS-based genome-wide CRISPR\/Cas9 knockout screening to identify genes that regulate the entry and accumulation of \u03b1-syn preformed fibrils (PFFs) in cells. We identified key genes and pathways specifically implicated in \u03b1-syn PFFs intracellular accumulation, including heparan sulfate proteoglycans (HSPG) biosynthesis and Golgi trafficking. All confirmed hits affected heparan sulfate (HS), a post-translational modification known to act as a receptor for proteinaceous aggregates including \u03b1-syn and tau. Intriguingly, deletion of <jats:italic>SLC39A9<\/jats:italic> and <jats:italic>C3orf58<\/jats:italic> genes, encoding respectively a Golgi-localized exporter of Zn<jats:sup>2+<\/jats:sup>, and the Golgi-localized putative kinase DIPK2A, specifically impaired the uptake of \u03b1-syn PFFs, by preventing the binding of PFFs to the cell surface. Mass spectrometry-based analysis of HS chains in <jats:italic>SLC39A9<\/jats:italic>\n            <jats:sup>-\/-<\/jats:sup> and <jats:italic>C3orf58<\/jats:italic>\n            <jats:sup>-\/-<\/jats:sup> cells indicated major defects in HS homeostasis. Additionally, Golgi accumulation of NDST1, a prime HSPG biosynthetic enzyme, was detected in <jats:italic>C3orf58<\/jats:italic>\n            <jats:sup>-\/-<\/jats:sup> cells. Interestingly, <jats:italic>C3orf58<\/jats:italic>\n            <jats:sup>-\/-<\/jats:sup> human iPSC-derived microglia and dopaminergic neurons exhibited a strong reduction in their ability to internalize \u03b1-syn PFFs. Altogether, our data identifies new modulators of HSPGs that regulate \u03b1-syn PFFs cell surface binding and uptake.<\/jats:p>","DOI":"10.1038\/s42003-025-08786-2","type":"journal-article","created":{"date-parts":[[2025,10,6]],"date-time":"2025-10-06T12:09:26Z","timestamp":1759752566000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":9,"title":["Novel regulators of heparan sulfate proteoglycans modulate cellular uptake of \u03b1-synuclein fibrils"],"prefix":"10.1038","volume":"8","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2111-1071","authenticated-orcid":false,"given":"Beno\u00eet","family":"Vanderperre","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amitha","family":"Muraleedharan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1285-6087","authenticated-orcid":false,"given":"Marie-France","family":"Dorion","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fr\u00e9d\u00e9rique","family":"Larroquette","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8658-944X","authenticated-orcid":false,"given":"Esther","family":"Del Cid Pellitero","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nishani","family":"Rajakulendran","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carol X.-Q.","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Roxanne","family":"Larivi\u00e8re","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Charlotte","family":"Michaud-Tardif","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas","family":"Goiran","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rony","family":"Chidiac","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Damien","family":"Lipuma","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Graham","family":"MacLeod","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5396-7587","authenticated-orcid":false,"given":"Rhalena","family":"Thomas","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhangjie","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wolfgang E.","family":"Reintsch","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1785-9615","authenticated-orcid":false,"given":"Wen","family":"Luo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3737-268X","authenticated-orcid":false,"given":"Irina","family":"Shlaifer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2803-3704","authenticated-orcid":false,"given":"Fuming","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ke","family":"Xia","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zachary","family":"Steinhart","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2219-5833","authenticated-orcid":false,"given":"Robert J.","family":"Linhardt","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6543-3371","authenticated-orcid":false,"given":"Jean-Fran\u00e7ois","family":"Trempe","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jian","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas M.","family":"Durcan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7241-9044","authenticated-orcid":false,"given":"Stephane","family":"Angers","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5520-6239","authenticated-orcid":false,"given":"Edward A.","family":"Fon","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,10,6]]},"reference":[{"key":"8786_CR1","doi-asserted-by":"publisher","unstructured":"Jellinger, K. A. Synucleinopathies. in Encyclopedia of Movement Disorders (eds. Kompoliti, K. & Metman, L. V.) 203\u2013207 https:\/\/doi.org\/10.1016\/B978-0-12-374105-9.00291-4 (Academic Press, Oxford, 2010).","DOI":"10.1016\/B978-0-12-374105-9.00291-4"},{"key":"8786_CR2","doi-asserted-by":"publisher","first-page":"445","DOI":"10.1007\/s004010050918","volume":"96","author":"K Wakabayashi","year":"1998","unstructured":"Wakabayashi, K. et al. Accumulation of alpha-synuclein\/NACP is a cytopathological feature common to Lewy body disease and multiple system atrophy. Acta Neuropathol. 96, 445\u2013452 (1998).","journal-title":"Acta Neuropathol."},{"key":"8786_CR3","doi-asserted-by":"publisher","first-page":"1128","DOI":"10.1093\/brain\/awt037","volume":"136","author":"M Masuda-Suzukake","year":"2013","unstructured":"Masuda-Suzukake, M. et al. Prion-like spreading of pathological -synuclein in brain. Brain 136, 1128\u20131138 (2013).","journal-title":"Brain"},{"key":"8786_CR4","doi-asserted-by":"publisher","DOI":"10.1007\/s11910-014-0495-z","volume":"14","author":"M Goedert","year":"2014","unstructured":"Goedert, M., Falcon, B., Clavaguera, F. & Tolnay, M. Prion-like mechanisms in the pathogenesis of tauopathies and synucleinopathies. Curr. Neurol. Neurosci. Rep. 14, 495 (2014).","journal-title":"Curr. Neurol. Neurosci. Rep."},{"key":"8786_CR5","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1038\/nm1746","volume":"14","author":"J-Y Li","year":"2008","unstructured":"Li, J.-Y. et al. Lewy bodies in grafted neurons in subjects with Parkinson\u2019s disease suggest host-to-graft disease propagation. Nat. Med. 14, 501\u2013503 (2008).","journal-title":"Nat. Med."},{"key":"8786_CR6","doi-asserted-by":"publisher","first-page":"20051","DOI":"10.1073\/pnas.0908005106","volume":"106","author":"KC Luk","year":"2009","unstructured":"Luk, K. C. et al. Exogenous alpha-synuclein fibrils seed the formation of Lewy body-like intracellular inclusions in cultured cells. Proc. Natl. Acad. Sci. USA 106, 20051\u201320056 (2009).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"8786_CR7","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1016\/j.neuron.2011.08.033","volume":"72","author":"LA Volpicelli-Daley","year":"2011","unstructured":"Volpicelli-Daley, L. A. et al. Exogenous alpha-synuclein fibrils induce Lewy body pathology leading to synaptic dysfunction and neuron death. Neuron 72, 57\u201371 (2011).","journal-title":"Neuron"},{"key":"8786_CR8","doi-asserted-by":"publisher","first-page":"975","DOI":"10.1084\/jem.20112457","volume":"209","author":"KC Luk","year":"2012","unstructured":"Luk, K. C. et al. Intracerebral inoculation of pathological alpha-synuclein initiates a rapidly progressive neurodegenerative alpha-synucleinopathy in mice. J. Exp. Med. 209, 975\u2013986 (2012).","journal-title":"J. Exp. Med."},{"key":"8786_CR9","doi-asserted-by":"publisher","first-page":"949","DOI":"10.1126\/science.1227157","volume":"338","author":"KC Luk","year":"2012","unstructured":"Luk, K. C. et al. Pathological alpha-synuclein transmission initiates Parkinson-like neurodegeneration in nontransgenic mice. Science 338, 949\u2013953 (2012).","journal-title":"Science"},{"key":"8786_CR10","doi-asserted-by":"publisher","first-page":"6838","DOI":"10.1523\/JNEUROSCI.5699-09.2010","volume":"30","author":"E Emmanouilidou","year":"2010","unstructured":"Emmanouilidou, E. et al. Cell-produced alpha-synuclein is secreted in a calcium-dependent manner by exosomes and impacts neuronal survival. J. Neurosci. J. Soc. Neurosci. 30, 6838\u20136851 (2010).","journal-title":"J. Neurosci. J. Soc. Neurosci."},{"key":"8786_CR11","doi-asserted-by":"publisher","DOI":"10.1038\/srep42984","volume":"7","author":"BV Dieriks","year":"2017","unstructured":"Dieriks, B. V. et al. -synuclein transfer through tunneling nanotubes occurs in SH-SY5Y cells and primary brain pericytes from Parkinson\u2019s disease patients. Sci. Rep. 7, 42984 (2017).","journal-title":"Sci. Rep."},{"key":"8786_CR12","doi-asserted-by":"publisher","first-page":"2120","DOI":"10.15252\/embj.201593411","volume":"35","author":"S Abounit","year":"2016","unstructured":"Abounit, S. et al. Tunneling nanotubes spread fibrillar \u03b1-synuclein by intercellular trafficking of lysosomes. EMBO J. 35, 2120\u20132138 (2016).","journal-title":"EMBO J."},{"key":"8786_CR13","doi-asserted-by":"publisher","first-page":"326","DOI":"10.1096\/fj.10-164624","volume":"25","author":"KM Danzer","year":"2011","unstructured":"Danzer, K. M. et al. Heat-shock protein 70 modulates toxic extracellular \u03b1-synuclein oligomers and rescues trans-synaptic toxicity. FASEB J. 25, 326\u2013336 (2011).","journal-title":"FASEB J."},{"key":"8786_CR14","doi-asserted-by":"publisher","first-page":"765","DOI":"10.1038\/ncb3372","volume":"18","author":"J-G Lee","year":"2016","unstructured":"Lee, J.-G., Takahama, S., Zhang, G., Tomarev, S. I. & Ye, Y. Unconventional secretion of misfolded proteins promotes adaptation to proteasome dysfunction in mammalian cells. Nat. Cell Biol. 18, 765\u2013776 (2016).","journal-title":"Nat. Cell Biol."},{"key":"8786_CR15","doi-asserted-by":"publisher","first-page":"835","DOI":"10.1016\/j.celrep.2015.12.075","volume":"14","author":"SH Oh","year":"2016","unstructured":"Oh, S. H. et al. Mesenchymal Stem Cells Inhibit Transmission of \u03b1-Synuclein by Modulating Clathrin-Mediated Endocytosis in a Parkinsonian Model. Cell Rep. 14, 835\u2013849 (2016).","journal-title":"Cell Rep."},{"key":"8786_CR16","doi-asserted-by":"publisher","unstructured":"Zhang, Q. et al. A myosin-7B\u2013dependent endocytosis pathway mediates cellular entry of \u03b1-synuclein fibrils and polycation-bearing cargos. Proc. Natl. Acad. Sci. https:\/\/doi.org\/10.1073\/pnas.1918617117 (2020).","DOI":"10.1073\/pnas.1918617117"},{"key":"8786_CR17","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-53038-z","volume":"9","author":"A Hud\u00e1k","year":"2019","unstructured":"Hud\u00e1k, A. et al. Contribution of syndecans to cellular uptake and fibrillation of \u03b1-synuclein and tau. Sci. Rep. 9, 16543 (2019).","journal-title":"Sci. Rep."},{"key":"8786_CR18","doi-asserted-by":"publisher","first-page":"E3138","DOI":"10.1073\/pnas.1301440110","volume":"110","author":"BB Holmes","year":"2013","unstructured":"Holmes, B. B. et al. Heparan sulfate proteoglycans mediate internalization and propagation of specific proteopathic seeds. Proc. Natl. Acad. Sci. USA110, E3138\u2013E3147 (2013).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"8786_CR19","doi-asserted-by":"publisher","first-page":"111102","DOI":"10.1016\/j.celrep.2022.111102","volume":"40","author":"A Bayati","year":"2022","unstructured":"Bayati, A. et al. Rapid macropinocytic transfer of \u03b1-synuclein to lysosomes. Cell Rep. 40, 111102 (2022).","journal-title":"Cell Rep."},{"key":"8786_CR20","doi-asserted-by":"publisher","first-page":"aah3374","DOI":"10.1126\/science.aah3374","volume":"353","author":"X Mao","year":"2016","unstructured":"Mao, X. et al. Pathological \u03b1-synuclein transmission initiated by binding lymphocyte-activation gene 3. Science 353, aah3374 (2016).","journal-title":"Science"},{"key":"8786_CR21","doi-asserted-by":"publisher","first-page":"e3000318","DOI":"10.1371\/journal.pbio.3000318","volume":"17","author":"M Birol","year":"2019","unstructured":"Birol, M., Wojcik, S. P., Miranker, A. D. & Rhoades, E. Identification of N-linked glycans as specific mediators of neuronal uptake of acetylated \u03b1-Synuclein. PLOS Biol. 17, e3000318 (2019).","journal-title":"PLOS Biol."},{"key":"8786_CR22","doi-asserted-by":"publisher","first-page":"1450","DOI":"10.1016\/j.cell.2018.07.002","volume":"174","author":"P Zhang","year":"2018","unstructured":"Zhang, P. et al. Heparan Sulfate Organizes Neuronal Synapses through Neurexin Partnerships. Cell 174, 1450\u20131464.e23 (2018).","journal-title":"Cell"},{"key":"8786_CR23","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1016\/j.nbd.2017.04.011","volume":"103","author":"F Cavaliere","year":"2017","unstructured":"Cavaliere, F. et al. In vitro \u03b1-synuclein neurotoxicity and spreading among neurons and astrocytes using Lewy body extracts from Parkinson disease brains. Neurobiol. Dis. 103, 101\u2013112 (2017).","journal-title":"Neurobiol. Dis."},{"key":"8786_CR24","doi-asserted-by":"publisher","first-page":"11835","DOI":"10.1523\/JNEUROSCI.0983-17.2017","volume":"37","author":"J Rostami","year":"2017","unstructured":"Rostami, J. et al. Human astrocytes transfer aggregated alpha-synuclein via tunneling nanotubes. J. Neurosci. 37, 11835\u201311853 (2017).","journal-title":"J. Neurosci."},{"key":"8786_CR25","doi-asserted-by":"publisher","first-page":"517","DOI":"10.1002\/ana.23747","volume":"72","author":"EC Freundt","year":"2012","unstructured":"Freundt, E. C. et al. Neuron-to-neuron transmission of \u03b1-synuclein fibrils through axonal transport. Ann. Neurol. 72, 517\u2013524 (2012).","journal-title":"Ann. Neurol."},{"key":"8786_CR26","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-017-08720-5","volume":"7","author":"E Ihse","year":"2017","unstructured":"Ihse, E. et al. Cellular internalization of alpha-synuclein aggregates by cell surface heparan sulfate depends on aggregate conformation and cell type. Sci. Rep. 7, 9008 (2017).","journal-title":"Sci. Rep."},{"key":"8786_CR27","doi-asserted-by":"publisher","first-page":"a004952","DOI":"10.1101\/cshperspect.a004952","volume":"3","author":"S Sarrazin","year":"2011","unstructured":"Sarrazin, S., Lamanna, W. C. & Esko, J. D. Heparan sulfate proteoglycans. Cold Spring Harb. Perspect. Biol. 3, a004952 (2011).","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"8786_CR28","doi-asserted-by":"publisher","first-page":"20008","DOI":"10.1074\/jbc.M801652200","volume":"283","author":"P Carlsson","year":"2008","unstructured":"Carlsson, P., Presto, J., Spillmann, D., Lindahl, U. & Kjell\u00e9n, L. Heparin\/Heparan Sulfate Biosynthesis processive formation of N-sulfated domains. J. Biol. Chem. 283, 20008\u201320014 (2008).","journal-title":"J. Biol. Chem."},{"key":"8786_CR29","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1146\/annurev-biochem-060713-035314","volume":"83","author":"D Xu","year":"2014","unstructured":"Xu, D. & Esko, J. D. Demystifying Heparan Sulfate\u2013Protein Interactions. Annu. Rev. Biochem. 83, 129\u2013157 (2014).","journal-title":"Annu. Rev. Biochem."},{"key":"8786_CR30","doi-asserted-by":"publisher","DOI":"10.1186\/s13059-014-0554-4","volume":"15","author":"W Li","year":"2014","unstructured":"Li, W. et al. MAGeCK enables robust identification of essential genes from genome-scale CRISPR\/Cas9 knockout screens. Genome Biol. 15, 554 (2014).","journal-title":"Genome Biol."},{"key":"8786_CR31","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-018-05191-8","volume":"9","author":"J Dubail","year":"2018","unstructured":"Dubail, J. et al. SLC10A7 mutations cause a skeletal dysplasia with amelogenesis imperfecta mediated by GAG biosynthesis defects. Nat. Commun. 9, 3087 (2018).","journal-title":"Nat. Commun."},{"key":"8786_CR32","doi-asserted-by":"publisher","DOI":"10.15252\/embr.201745453","volume":"19","author":"T Di Mattia","year":"2018","unstructured":"Di Mattia, T. et al. Identification of MOSPD2, a novel scaffold for endoplasmic reticulum membrane contact sites. EMBO Rep. 19, e45453 (2018).","journal-title":"EMBO Rep."},{"key":"8786_CR33","doi-asserted-by":"publisher","first-page":"168","DOI":"10.1016\/j.ajhg.2011.06.008","volume":"89","author":"A Zimprich","year":"2011","unstructured":"Zimprich, A. et al. A mutation in VPS35, encoding a subunit of the retromer complex, causes late-onset Parkinson disease. Am. J. Hum. Genet. 89, 168\u2013175 (2011).","journal-title":"Am. J. Hum. Genet."},{"key":"8786_CR34","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1038\/nrc3179","volume":"12","author":"M Sadelain","year":"2012","unstructured":"Sadelain, M., Papapetrou, E. P. & Bushman, F. D. Safe harbours for the integration of new DNA in the human genome. Nat. Rev. Cancer 12, 51\u201358 (2012).","journal-title":"Nat. Rev. Cancer"},{"key":"8786_CR35","doi-asserted-by":"publisher","first-page":"18","DOI":"10.1016\/j.mcn.2018.09.003","volume":"93","author":"A Ansell-Schultz","year":"2018","unstructured":"Ansell-Schultz, A., Reyes, J. F., Samuelsson, M. & Hallbeck, M. Reduced retromer function results in the accumulation of amyloid-beta oligomers. Mol. Cell. Neurosci. 93, 18\u201326 (2018).","journal-title":"Mol. Cell. Neurosci."},{"key":"8786_CR36","doi-asserted-by":"publisher","first-page":"10826","DOI":"10.1074\/jbc.RA117.000378","volume":"293","author":"BE Stopschinski","year":"2018","unstructured":"Stopschinski, B. E. et al. Specific glycosaminoglycan chain length and sulfation patterns are required for cell uptake of tau versus \u03b1-synuclein and \u03b2-amyloid aggregates. J. Biol. Chem. 293, 10826\u201310840 (2018).","journal-title":"J. Biol. Chem."},{"key":"8786_CR37","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1016\/j.isci.2018.12.011","volume":"11","author":"JS Park","year":"2019","unstructured":"Park, J. S. et al. A FACS-based genome-wide CRISPR screen reveals a requirement for COPI in Chlamydia trachomatis Invasion. iScience 11, 71\u201384 (2019).","journal-title":"iScience"},{"key":"8786_CR38","doi-asserted-by":"publisher","first-page":"4280","DOI":"10.1091\/mbc.e15-06-0361","volume":"26","author":"CH Woo","year":"2015","unstructured":"Woo, C. H. et al. Conserved function of the lysine-based KXD\/E motif in Golgi retention for endomembrane proteins among different organisms. Mol. Biol. Cell 26, 4280\u20134293 (2015).","journal-title":"Mol. Biol. Cell"},{"key":"8786_CR39","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.00432-17","volume":"91","author":"A Tanaka","year":"2017","unstructured":"Tanaka, A. et al. Genome-wide screening uncovers the significance of N-Sulfation of Heparan sulfate as a host cell factor for Chikungunya virus infection. J. Virol. 91, e00432-17 (2017).","journal-title":"J. Virol."},{"key":"8786_CR40","doi-asserted-by":"publisher","first-page":"797","DOI":"10.1080\/15548627.2019.1637199","volume":"16","author":"X Tian","year":"2020","unstructured":"Tian, X. et al. DIPK2A promotes STX17- and VAMP7-mediated autophagosome-lysosome fusion by binding to VAMP7B. Autophagy 16, 797\u2013810 (2020).","journal-title":"Autophagy"},{"key":"8786_CR41","doi-asserted-by":"publisher","first-page":"771","DOI":"10.1042\/BCJ20160976","volume":"474","author":"A Bareja","year":"2017","unstructured":"Bareja, A., Hodgkinson, C. P., Payne, A. J., Pratt, R. E. & Dzau, V. J. HASF (C3orf58) is a novel ligand of the insulin-like growth factor 1 receptor. Biochem. J. 474, 771\u2013780 (2017).","journal-title":"Biochem. J."},{"key":"8786_CR42","doi-asserted-by":"publisher","first-page":"534","DOI":"10.1177\/0022034509338452","volume":"88","author":"MS Takatalo","year":"2009","unstructured":"Takatalo, M. S., Tummers, M., Thesleff, I. & R\u00f6nnholm, R. Novel Golgi Protein, GoPro49, Is a Specific Dental Follicle Marker. J. Dent. Res. 88, 534\u2013538 (2009).","journal-title":"J. Dent. Res."},{"key":"8786_CR43","doi-asserted-by":"publisher","unstructured":"Wells, M. F. et al. Genome-wide screens in accelerated human stem cell-derived neural progenitor cells identify Zika virus host factors and drivers of proliferation. bioRxiv 476440 https:\/\/doi.org\/10.1101\/476440 (2018).","DOI":"10.1101\/476440"},{"key":"8786_CR44","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-018-24904-z","volume":"8","author":"JN Rauch","year":"2018","unstructured":"Rauch, J. N. et al. Tau Internalization is Regulated by 6-O Sulfation on Heparan Sulfate Proteoglycans (HSPGs). Sci. Rep. 8, 6382 (2018).","journal-title":"Sci. Rep."},{"key":"8786_CR45","doi-asserted-by":"publisher","first-page":"599","DOI":"10.1093\/glycob\/cwh067","volume":"14","author":"K Mani","year":"2004","unstructured":"Mani, K. et al. The heparan sulfate-specific epitope 10E4 is NO-sensitive and partly inaccessible in glypican-1. Glycobiology 14, 599\u2013607 (2004).","journal-title":"Glycobiology"},{"key":"8786_CR46","doi-asserted-by":"publisher","first-page":"36267","DOI":"10.1074\/jbc.274.51.36267","volume":"274","author":"F Safaiyan","year":"1999","unstructured":"Safaiyan, F. et al. Selective effects of sodium chlorate treatment on the sulfation of Heparan sulfate*. J. Biol. Chem. 274, 36267\u201336273 (1999).","journal-title":"J. Biol. Chem."},{"key":"8786_CR47","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1016\/j.matbio.2013.10.004","volume":"35","author":"HC Christianson","year":"2014","unstructured":"Christianson, H. C. & Belting, M. Heparan sulfate proteoglycan as a cell-surface endocytosis receptor. Matrix Biol. 35, 51\u201355 (2014).","journal-title":"Matrix Biol."},{"key":"8786_CR48","doi-asserted-by":"publisher","first-page":"1515","DOI":"10.1016\/j.cell.2015.11.015","volume":"163","author":"T Hart","year":"2015","unstructured":"Hart, T. et al. High-resolution CRISPR screens reveal fitness genes and genotype-specific cancer liabilities. Cell 163, 1515\u20131526 (2015).","journal-title":"Cell"},{"key":"8786_CR49","doi-asserted-by":"publisher","first-page":"1142","DOI":"10.1271\/bbb.80910","volume":"73","author":"W Matsuura","year":"2009","unstructured":"Matsuura, W. et al. SLC39A9 (ZIP9) regulates zinc homeostasis in the secretory pathway: characterization of the ZIP subfamily I protein in vertebrate cells. Biosci. Biotechnol. Biochem. 73, 1142\u20131148 (2009).","journal-title":"Biosci. Biotechnol. Biochem."},{"key":"8786_CR50","doi-asserted-by":"publisher","first-page":"2708","DOI":"10.3390\/ijms18122708","volume":"18","author":"T Takagishi","year":"2017","unstructured":"Takagishi, T., Hara, T. & Fukada, T. Recent advances in the role of SLC39A\/ZIP zinc transporters in vivo. Int. J. Mol. Sci. 18, 2708 (2017).","journal-title":"Int. J. Mol. Sci."},{"key":"8786_CR51","doi-asserted-by":"publisher","first-page":"e66427","DOI":"10.1371\/journal.pone.0066427","volume":"8","author":"M Dudkiewicz","year":"2013","unstructured":"Dudkiewicz, M., Lenart, A. & Paw\u0142owski, K. A novel predicted calcium-regulated kinase family implicated in neurological disorders. PloS One 8, e66427 (2013).","journal-title":"PloS One"},{"key":"8786_CR52","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1016\/j.jprot.2019.01.015","volume":"196","author":"Y Li","year":"2019","unstructured":"Li, Y. et al. Sensitive profiling of cell surface proteome by using an optimized biotinylation method. J. Proteom. 196, 33\u201341 (2019).","journal-title":"J. Proteom."},{"key":"8786_CR53","doi-asserted-by":"publisher","first-page":"333","DOI":"10.1016\/j.cels.2020.03.003","volume":"10","author":"R Pourhaghighi","year":"2020","unstructured":"Pourhaghighi, R. et al. BraInMap elucidates the macromolecular connectivity landscape of mammalian brain. Cell Syst. 10, 333\u2013350.e14 (2020).","journal-title":"Cell Syst."},{"key":"8786_CR54","doi-asserted-by":"publisher","first-page":"733","DOI":"10.1093\/genetics\/155.2.733","volume":"155","author":"S Abdelilah-Seyfried","year":"2000","unstructured":"Abdelilah-Seyfried, S. et al. A gain-of-function screen for genes that affect the development of the Drosophila adult external sensory organ. Genetics 155, 733\u2013752 (2000).","journal-title":"Genetics"},{"key":"8786_CR55","doi-asserted-by":"publisher","first-page":"132","DOI":"10.1016\/j.matbio.2022.08.009","volume":"112","author":"J-L Zhang","year":"2022","unstructured":"Zhang, J.-L. et al. Vertebrate extracellular matrix protein hemicentin-1 interacts physically and genetically with basement membrane protein nidogen-2. Matrix Biol. J. Int. Soc. Matrix Biol. 112, 132\u2013154 (2022).","journal-title":"Matrix Biol. J. Int. Soc. Matrix Biol."},{"key":"8786_CR56","doi-asserted-by":"publisher","first-page":"117477","DOI":"10.1016\/j.carbpol.2020.117477","volume":"255","author":"MCZ Meneghetti","year":"2021","unstructured":"Meneghetti, M. C. Z. et al. ER-Golgi dynamics of HS-modifying enzymes via vesicular trafficking is a critical prerequisite for the delineation of HS biosynthesis. Carbohydr. Polym. 255, 117477 (2021).","journal-title":"Carbohydr. Polym."},{"key":"8786_CR57","doi-asserted-by":"publisher","first-page":"765","DOI":"10.1007\/978-1-0716-2639-9_46","volume":"2557","author":"I Mejia","year":"2023","unstructured":"Mejia, I., Chen, Y.-C. & D\u00edaz, B. Analysis of Golgi morphology using immunofluorescence and CellProfiler Software. Methods Mol. Biol. Clifton NJ 2557, 765\u2013784 (2023).","journal-title":"Methods Mol. Biol. Clifton NJ"},{"key":"8786_CR58","doi-asserted-by":"publisher","first-page":"20427","DOI":"10.1074\/jbc.M115.664409","volume":"290","author":"W Dou","year":"2015","unstructured":"Dou, W., Xu, Y., Pagadala, V., Pedersen, L. C. & Liu, J. Role of Deacetylase activity of N-Deacetylase\/N-Sulfotransferase 1 in forming N-sulfated domain in Heparan Sulfate. J. Biol. Chem. 290, 20427\u201320437 (2015).","journal-title":"J. Biol. Chem."},{"key":"8786_CR59","doi-asserted-by":"publisher","first-page":"407","DOI":"10.1016\/j.isci.2019.05.005","volume":"15","author":"T Yamaji","year":"2019","unstructured":"Yamaji, T. et al. A CRISPR screen using subtilase cytotoxin identifies SLC39A9 as a Glycan-regulating factor. iScience 15, 407\u2013420 (2019).","journal-title":"iScience"},{"key":"8786_CR60","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.4599","volume":"6","author":"A Hareza","year":"2018","unstructured":"Hareza, A. et al. Phosphoproteomic insights into processes influenced by the kinase-like protein DIA1\/C3orf58. PeerJ 6, e4599 (2018).","journal-title":"PeerJ"},{"key":"8786_CR61","doi-asserted-by":"publisher","first-page":"351","DOI":"10.1042\/bj3250351","volume":"325","author":"DE Humphries","year":"1997","unstructured":"Humphries, D. E., Sullivan, B. M., Aleixo, M. D. & Stow, J. L. Localization of human heparan glucosaminyl N-deacetylase\/N-sulphotransferase to the trans-Golgi network. Biochem. J. 325, 351\u2013357 (1997).","journal-title":"Biochem. J."},{"key":"8786_CR62","doi-asserted-by":"publisher","first-page":"927","DOI":"10.1038\/s41591-018-0049-z","volume":"24","author":"E Haapaniemi","year":"2018","unstructured":"Haapaniemi, E., Botla, S., Persson, J., Schmierer, B. & Taipale, J. CRISPR\u2013Cas9 genome editing induces a p53-mediated DNA damage response. Nat. Med. 24, 927 (2018).","journal-title":"Nat. Med."},{"key":"8786_CR63","doi-asserted-by":"publisher","first-page":"637","DOI":"10.1016\/j.cell.2020.03.031","volume":"181","author":"M Zhang","year":"2020","unstructured":"Zhang, M. et al. A translocation pathway for vesicle-mediated unconventional protein secretion. Cell 181, 637\u2013652.e15 (2020).","journal-title":"Cell"},{"key":"8786_CR64","doi-asserted-by":"publisher","first-page":"11393","DOI":"10.1073\/pnas.94.21.11393","volume":"94","author":"W Nickel","year":"1997","unstructured":"Nickel, W., Sohn, K., B\u00fcnning, C. & Wieland, F. T. p23, a major COPI-vesicle membrane protein, constitutively cycles through the early secretory pathway. Proc. Natl. Acad. Sci. 94, 11393\u201311398 (1997).","journal-title":"Proc. Natl. Acad. Sci."},{"key":"8786_CR65","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.02160-18","volume":"93","author":"RD Luteijn","year":"2019","unstructured":"Luteijn, R. D. et al. A genome-wide Haploid genetic screen identifies Heparan sulfate-associated genes and the macropinocytosis modulator TMED10 as factors supporting Vaccinia virus infection. J. Virol. 93, e02160-18 (2019).","journal-title":"J. Virol."},{"key":"8786_CR66","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1038\/s41586-019-1831-x","volume":"576","author":"C Ramirez","year":"2019","unstructured":"Ramirez, C., Hauser, A. D., Vucic, E. A. & Bar-Sagi, D. Plasma membrane v-ATPase controls oncogenic Ras-induced macropinocytosis. Nature 576, 477\u2013481 (2019).","journal-title":"Nature"},{"key":"8786_CR67","doi-asserted-by":"publisher","first-page":"E5906","DOI":"10.1073\/pnas.1524532113","volume":"113","author":"CM Buckley","year":"2016","unstructured":"Buckley, C. M. et al. WASH drives early recycling from macropinosomes and phagosomes to maintain surface phagocytic receptors. Proc. Natl. Acad. Sci. 113, E5906\u2013E5915 (2016).","journal-title":"Proc. Natl. Acad. Sci."},{"key":"8786_CR68","doi-asserted-by":"publisher","unstructured":"Li, Z. et al. Cell-Surface RNA forms ternary complex with RNA-binding proteins and Heparan sulfate to recruit immune receptors. 2024.07.23.604836 Preprint at https:\/\/doi.org\/10.1101\/2024.07.23.604836 (2025).","DOI":"10.1101\/2024.07.23.604836"},{"key":"8786_CR69","doi-asserted-by":"publisher","first-page":"778752","DOI":"10.3389\/fonc.2021.778752","volume":"11","author":"C Marques","year":"2021","unstructured":"Marques, C., Reis, C. A., Viv\u00e8s, R. R. & Magalh\u00e3es, A. Heparan sulfate biosynthesis and sulfation profiles as modulators of cancer signalling and progression. Front. Oncol. 11, 778752 (2021).","journal-title":"Front. Oncol."},{"key":"8786_CR70","doi-asserted-by":"publisher","first-page":"5989","DOI":"10.1007\/s00253-018-9064-8","volume":"102","author":"A Prabhu","year":"2018","unstructured":"Prabhu, A., Gadre, R. & Gadgil, M. Zinc supplementation decreases galactosylation of recombinant IgG in CHO cells. Appl. Microbiol. Biotechnol. 102, 5989\u20135999 (2018).","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"8786_CR71","doi-asserted-by":"publisher","first-page":"9527","DOI":"10.1073\/pnas.1900867116","volume":"116","author":"Y Li","year":"2019","unstructured":"Li, Y. et al. Genome-wide CRISPR screen for Zika virus resistance in human neural cells. Proc. Natl. Acad. Sci. 116, 9527\u20139532 (2019).","journal-title":"Proc. Natl. Acad. Sci."},{"key":"8786_CR72","doi-asserted-by":"publisher","first-page":"427","DOI":"10.1093\/brain\/awad298","volume":"147","author":"M-F Dorion","year":"2024","unstructured":"Dorion, M.-F. et al. MerTK is a mediator of alpha-synuclein fibril uptake by human microglia. Brain J. Neurol. 147, 427\u2013443 (2024).","journal-title":"Brain J. Neurol."},{"key":"8786_CR73","doi-asserted-by":"publisher","first-page":"114084","DOI":"10.1016\/j.bcp.2020.114084","volume":"178","author":"C Lanzi","year":"2020","unstructured":"Lanzi, C. & Cassinelli, G. Receptor tyrosine kinases and heparan sulfate proteoglycans: Interplay providing anticancer targeting strategies and new therapeutic opportunities. Biochem. Pharmacol. 178, 114084 (2020).","journal-title":"Biochem. Pharmacol."},{"key":"8786_CR74","doi-asserted-by":"publisher","first-page":"297","DOI":"10.1016\/j.ymeth.2021.09.002","volume":"203","author":"E Deneault","year":"2022","unstructured":"Deneault, E. et al. A streamlined CRISPR workflow to introduce mutations and generate isogenic iPSCs for modeling amyotrophic lateral sclerosis. Methods San. Diego Calif. 203, 297\u2013310 (2022).","journal-title":"Methods San. Diego Calif."},{"key":"8786_CR75","doi-asserted-by":"publisher","first-page":"50","DOI":"10.3390\/mps4030050","volume":"4","author":"CX-Q Chen","year":"2021","unstructured":"Chen, C. X.-Q. et al. A multistep workflow to evaluate newly generated iPSCs and their ability to generate different cell types. Methods Protoc. 4, 50 (2021).","journal-title":"Methods Protoc."},{"key":"8786_CR76","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1186\/s13024-018-0297-x","volume":"13","author":"A McQuade","year":"2018","unstructured":"McQuade, A. et al. Development and validation of a simplified method to generate human microglia from pluripotent stem cells. Mol. Neurodegener. 13, 67 (2018).","journal-title":"Mol. Neurodegener."},{"key":"8786_CR77","doi-asserted-by":"publisher","first-page":"583","DOI":"10.1038\/s41586-021-03819-2","volume":"596","author":"J Jumper","year":"2021","unstructured":"Jumper, J. et al. Highly accurate protein structure prediction with AlphaFold. Nature 596, 583\u2013589 (2021).","journal-title":"Nature"},{"key":"8786_CR78","doi-asserted-by":"publisher","first-page":"243","DOI":"10.1038\/s41587-023-01773-0","volume":"42","author":"M van Kempen","year":"2024","unstructured":"van Kempen, M. et al. Fast and accurate protein structure search with Foldseek. Nat. Biotechnol. 42, 243\u2013246 (2024).","journal-title":"Nat. Biotechnol."},{"key":"8786_CR79","doi-asserted-by":"publisher","first-page":"3338","DOI":"10.1021\/acschembio.6b00511","volume":"11","author":"K Yamada","year":"2016","unstructured":"Yamada, K. et al. Discovery and characterization of allosteric WNK kinase inhibitors. ACS Chem. Biol. 11, 3338\u20133346 (2016).","journal-title":"ACS Chem. Biol."},{"key":"8786_CR80","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.1700344","volume":"3","author":"T Zhang","year":"2017","unstructured":"Zhang, T. et al. Crystal structures of a ZIP zinc transporter reveal a binuclear metal center in the transport pathway. Sci. Adv. 3, e1700344 (2017).","journal-title":"Sci. Adv."},{"key":"8786_CR81","doi-asserted-by":"publisher","first-page":"2719","DOI":"10.1534\/g3.117.041277","volume":"7","author":"T Hart","year":"2017","unstructured":"Hart, T. et al. Evaluation and design of genome-wide CRISPR\/SpCas9 knockout screens. G3 Bethesda Md 7, 2719\u20132727 (2017).","journal-title":"G3 Bethesda Md"},{"key":"8786_CR82","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1007\/978-1-60761-444-9_15","volume":"604","author":"PGA Pedrioli","year":"2010","unstructured":"Pedrioli, P. G. A. Trans-proteomic pipeline: a pipeline for proteomic analysis. Methods Mol. Biol. Clifton NJ 604, 213\u2013238 (2010).","journal-title":"Methods Mol. Biol. Clifton NJ"},{"key":"8786_CR83","doi-asserted-by":"publisher","first-page":"1015","DOI":"10.1038\/nbt1010-1015","volume":"28","author":"G Liu","year":"2010","unstructured":"Liu, G. et al. ProHits: integrated software for mass spectrometry-based interaction proteomics. Nat. Biotechnol. 28, 1015\u20131017 (2010).","journal-title":"Nat. Biotechnol."},{"key":"8786_CR84","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1093\/bioinformatics\/btp616","volume":"26","author":"MD Robinson","year":"2010","unstructured":"Robinson, M. D., McCarthy, D. J. & Smyth, G. K. edgeR: a Bioconductor package for differential expression analysis of digital gene expression data. Bioinforma. Oxf. Engl. 26, 139\u2013140 (2010).","journal-title":"Bioinforma. Oxf. Engl."},{"key":"8786_CR85","unstructured":"Young, M. D., Wakefield, M. J., Oshlack, A. & Smyth, G. K. goseq: Gene Ontology testing for RNA-seq datasets. https:\/\/www.bioconductor.org\/packages\/\/2.10\/bioc\/vignettes\/goseq\/inst\/doc\/goseq.pdf."},{"key":"8786_CR86","doi-asserted-by":"publisher","first-page":"526","DOI":"10.1002\/mds.27642","volume":"34","author":"RN Alcalay","year":"2019","unstructured":"Alcalay, R. N. et al. SMPD1 mutations, activity, and \u03b1-synuclein accumulation in Parkinson\u2019s disease. Mov. Disord. 34, 526\u2013535 (2019).","journal-title":"Mov. Disord."},{"key":"8786_CR87","doi-asserted-by":"publisher","unstructured":"Del Cid Pellitero, E. et al. Quality control characterization of \u03b1-Synuclein preformed Fibrils (PFFs). https:\/\/doi.org\/10.5281\/zenodo.3738340 (2019).","DOI":"10.5281\/zenodo.3738340"},{"key":"8786_CR88","doi-asserted-by":"publisher","unstructured":"Maneca, D.-L. et al. Production of recombinant \u03b1 Synuclein monomers and Preformed Fibrils (PFFs). https:\/\/doi.org\/10.5281\/zenodo.6430401 (2022).","DOI":"10.5281\/zenodo.6430401"},{"key":"8786_CR89","doi-asserted-by":"publisher","first-page":"14234","DOI":"10.1523\/JNEUROSCI.1523-15.2015","volume":"35","author":"M Usenovic","year":"2015","unstructured":"Usenovic, M. et al. Internalized Tau Oligomers cause neurodegeneration by inducing accumulation of pathogenic Tau in human neurons derived from induced Pluripotent stem cells. J. Neurosci. 35, 14234\u201314250 (2015).","journal-title":"J. Neurosci."},{"key":"8786_CR90","doi-asserted-by":"publisher","first-page":"D543","DOI":"10.1093\/nar\/gkae1011","volume":"53","author":"Y Perez-Riverol","year":"2025","unstructured":"Perez-Riverol, Y. et al. The PRIDE database at 20 years: 2025 update. Nucleic Acids Res. 53, D543\u2013D553 (2025).","journal-title":"Nucleic Acids Res."}],"container-title":["Communications Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s42003-025-08786-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-025-08786-2","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-025-08786-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,7]],"date-time":"2025-10-07T00:03:19Z","timestamp":1759795399000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s42003-025-08786-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,10,6]]},"references-count":90,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["8786"],"URL":"https:\/\/doi.org\/10.1038\/s42003-025-08786-2","relation":{},"ISSN":["2399-3642"],"issn-type":[{"value":"2399-3642","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,10,6]]},"assertion":[{"value":"24 December 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"22 August 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"6 October 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"J.L. is a founder and chief scientific officer of Glycan Therapeutics Corp. Z.W. is currently an employee of Glycan Therapeutics Corp. Other authors have no competing interest to declare.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"1426"}}