{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T21:17:55Z","timestamp":1788383875560,"version":"build-2803163510"},"reference-count":55,"publisher":"American Chemical Society (ACS)","issue":"5","license":[{"start":{"date-parts":[[2020,5,13]],"date-time":"2020-05-13T00:00:00Z","timestamp":1589328000000},"content-version":"vor","delay-in-days":69,"URL":"https:\/\/pubs.acs.org\/page\/policy\/authorchoice_ccby_termsofuse.html"}],"funder":[{"DOI":"10.13039\/100000057","name":"National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["R01GM097971"],"award-info":[{"award-number":["R01GM097971"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004359","name":"Vetenskapsr\u00e5det","doi-asserted-by":"publisher","award":["2017-02646"],"award-info":[{"award-number":["2017-02646"]}],"id":[{"id":"10.13039\/501100004359","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2020,5,8]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>A promising strategy to limit cholera severity involves blockers mimicking the canonical cholera toxin ligand (CT) ganglioside GM1. However, to date the efficacies of most of these blockers have been evaluated in noncellular systems that lack ligands other than GM1. Importantly, the CT B subunit (CTB) has a noncanonical site that binds fucosylated structures, which in contrast to GM1 are highly expressed in the human intestine. Here we evaluate the capacity of norbornene polymers displaying galactose and\/or fucose to block CTB binding to immobilized protein-linked glycan structures and also to primary human and murine small intestine epithelial cells (SI ECs). We show that the binding of CTB to human SI ECs is largely dependent on the noncanonical binding site, and interference with the canonical site has a limited effect while the opposite is observed with murine SI ECs. The galactose\u2013fucose polymer blocks binding to fucosylated glycans but not to GM1. However, the preincubation of CT with the galactose\u2013fucose polymer only partially blocks toxic effects on cultured human enteroid cells, while preincubation with GM1 completely blocks CT-mediated secretion. Our results support a model whereby the binding of fucose to the noncanonical site places CT in close proximity to scarcely expressed galactose receptors such as GM1 to enable binding via the canonical site leading to CT internalization and intoxication. Our finding also highlights the importance of complementing CTB binding studies with functional intoxication studies when assessing the efficacy inhibitors of CT.<\/jats:p>","DOI":"10.1021\/acsinfecdis.0c00009","type":"journal-article","created":{"date-parts":[[2020,3,5]],"date-time":"2020-03-05T13:34:48Z","timestamp":1583415288000},"page":"1192-1203","source":"Crossref","is-referenced-by-count":26,"title":["Fucose-Galactose\nPolymers Inhibit Cholera Toxin Binding\nto Fucosylated Structures and Galactose-Dependent Intoxication of\nHuman Enteroids"],"prefix":"10.1021","volume":"6","author":[{"given":"Jakob","family":"Cervin","sequence":"first","affiliation":[{"name":"University of Gothenburg , , ,","place":["Gothenburg, Sweden, 405 30"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4237-4612","authenticated-orcid":true,"given":"Andrew","family":"Boucher","sequence":"additional","affiliation":[{"name":"University of Gothenburg , , ,","place":["Gothenburg, Sweden, 405 30"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9152-1367","authenticated-orcid":true,"given":"Gyusaang","family":"Youn","sequence":"additional","affiliation":[{"name":"Stony Brook University , , , , ,","place":["Stony Brook, New York, United States, 11794-3400"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Per","family":"Bjo\u0308rklund","sequence":"additional","affiliation":[{"name":"Sahlgrenska Academy, University of Gothenburg, Sahlgrenska University Hospital\/O\u0308stra , , ,","place":["Gothenburg, Sweden, 416 85"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ville","family":"Wallenius","sequence":"additional","affiliation":[{"name":"Sahlgrenska Academy, University of Gothenburg, Sahlgrenska University Hospital\/O\u0308stra , , ,","place":["Gothenburg, Sweden, 416 85"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lynda","family":"Mottram","sequence":"additional","affiliation":[{"name":"University of Gothenburg , , ,","place":["Gothenburg, Sweden, 405 30"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2835-7760","authenticated-orcid":true,"given":"Nicole S.","family":"Sampson","sequence":"additional","affiliation":[{"name":"Stony Brook University , , , , ,","place":["Stony Brook, New York, United States, 11794-3400"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ulf","family":"Yrlid","sequence":"additional","affiliation":[{"name":"University of Gothenburg , , ,","place":["Gothenburg, Sweden, 405 30"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2020,3,5]]},"reference":[{"issue":"6","key":"2026090216341770900_cit1","doi-asserted-by":"publisher","first-page":"e0003832","DOI":"10.1371\/journal.pntd.0003832","article-title":"Updated global burden\nof cholera in endemic countries","volume":"9","author":"Ali","year":"2015","journal-title":"PLoS Neglected\nTrop. 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Rep."},{"issue":"10","key":"2026090216341770900_cit4","doi-asserted-by":"publisher","first-page":"e970","DOI":"10.1016\/S2214-109X(17)30332-7","article-title":"The political\ndeterminants of the\ncholera outbreak in Yemen","volume":"5","author":"Kennedy","year":"2017","journal-title":"Lancet Glob Health"},{"issue":"6","key":"2026090216341770900_cit5","first-page":"379","article-title":"Role of oral\nrehydration therapy in controlling epidemic of cholera and watery\ndiarrhoea","volume":"101","author":"Sarkar","year":"2003","journal-title":"J. Indian Med. 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J.\nBiochem. Cell Biol."},{"issue":"1","key":"2026090216341770900_cit31","doi-asserted-by":"publisher","first-page":"52","DOI":"10.1039\/C9OB02369K","article-title":"A hybrid polymer\nto target blood\ngroup dependence of cholera toxin","volume":"18","author":"Haksar","year":"2020","journal-title":"Org. Biomol.\nChem."},{"issue":"1","key":"2026090216341770900_cit32","doi-asserted-by":"publisher","first-page":"473","DOI":"10.1146\/annurev-pathol-012414-040438","article-title":"Protein glycosylation in cancer","volume":"10","author":"Stowell","year":"2015","journal-title":"Annu. Rev. Pathol.: Mech. 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Commun."},{"issue":"5","key":"2026090216341770900_cit36","doi-asserted-by":"publisher","first-page":"1210","DOI":"10.1172\/JCI105810","article-title":"Site and\ncharacteristics of electrolyte loss and effect of intraluminal glucose\nin experimental canine cholera","volume":"47","author":"Carpenter","year":"1968","journal-title":"J. Clin. Invest."},{"issue":"12","key":"2026090216341770900_cit37","doi-asserted-by":"publisher","first-page":"2600","DOI":"10.1172\/JCI105942","article-title":"Response\nof the canine duodenum to intraluminal challenge\nwith cholera exotoxin","volume":"47","author":"Carpenter","year":"1968","journal-title":"J. Clin. Invest."},{"issue":"1","key":"2026090216341770900_cit38","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1172\/JCI106217","article-title":"Intestinal fluid and electrolyte transport in human\ncholera","volume":"49","author":"Banwell","year":"1970","journal-title":"J. Clin. 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Immun."},{"issue":"6","key":"2026090216341770900_cit41","doi-asserted-by":"publisher","first-page":"533","DOI":"10.1007\/BF00731304","article-title":"Comparison\nof the glycolipid-binding specificities of cholera toxin and porcine\nEscherichia coli heat-labile enterotoxin: identification of a receptor-active\nnon-ganglioside glycolipid for the heat-labile toxin in infant rabbit\nsmall intestine","volume":"11","author":"Teneberg","year":"1994","journal-title":"Glycoconjugate J."},{"key":"2026090216341770900_cit42","doi-asserted-by":"publisher","first-page":"45270","DOI":"10.1038\/srep45270","article-title":"A primary\nhuman macrophage-enteroid\nco-culture model to investigate mucosal gut physiology and host-pathogen\ninteractions","volume":"7","author":"Noel","year":"2017","journal-title":"Sci. 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Hyg."},{"issue":"12","key":"2026090216341770900_cit48","doi-asserted-by":"crossref","first-page":"6346","DOI":"10.1128\/IAI.67.12.6346-6349.1999","article-title":"Validation and characterization\nof a human volunteer challenge model for cholera by using frozen bacteria\nof the new Vibrio cholerae epidemic serotype, O139","volume":"67","author":"Cohen","year":"1999","journal-title":"Infect. Immun."},{"issue":"6","key":"2026090216341770900_cit49","doi-asserted-by":"publisher","first-page":"791","DOI":"10.1093\/oxfordjournals.aje.a114050","article-title":"Predisposition for\ncholera of individuals with O blood group. Possible\nevolutionary significance","volume":"121","author":"Glass","year":"1985","journal-title":"Am. J. Epidemiol."},{"issue":"10","key":"2026090216341770900_cit50","first-page":"1181","article-title":"The Broad Street pump revisited:\nresponse of volunteers to ingested cholera vibrios","volume":"47","author":"Hornick","year":"1971","journal-title":"Bull. N Y Acad. 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Biotechnol."},{"issue":"13","key":"2026090216341770900_cit53","doi-asserted-by":"publisher","first-page":"7532","DOI":"10.1073\/pnas.96.13.7532","article-title":"Mice lacking complex gangliosides\ndevelop Wallerian degeneration and myelination defects","volume":"96","author":"Sheikh","year":"1999","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"issue":"20","key":"2026090216341770900_cit54","doi-asserted-by":"publisher","first-page":"10662","DOI":"10.1073\/pnas.93.20.10662","article-title":"Mice with\ndisrupted GM2\/GD2 synthase gene lack complex\ngangliosides but exhibit only subtle defects in their nervous system","volume":"93","author":"Takamiya","year":"1996","journal-title":"Proc. Natl. Acad. Sci. U. S. 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