Glycans in the immune system and The Altered Glycan Theory of Autoimmunity: a critical review

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Specific topics covered include: the glycosylation sites of IgE, IgM, IgD, IgE, IgA, and IgG; how glycans can encode "self" identity by functioning as either danger associated molecular patterns (DAMPs) or self-associated molecular patt …"><meta name="keywords" content="pmid:25578468, PMC4340844, doi:10.1016/j.jaut.2014.12.002, Research Support, N.I.H., Extramural, Research Support, Non-U.S. Gov't, Review, Emanual Maverakis, Kyoungmi Kim, Carlito B Lebrilla, Autoimmune Diseases / immunology, Autoimmune Diseases / metabolism, Autoimmune Diseases / pathology, Autoimmunity / immunology*, Binding Sites / immunology, Glycosylation, Humans, Immune System / cytology, Immune System / immunology*, Immune System / metabolism, Immunoglobulins / immunology*, Immunoglobulins / metabolism, Models, Immunological, Polysaccharides / immunology*, Polysaccharides / metabolism, PubMed Abstract, NIH, NLM, NCBI, National Institutes of Health, National Center for Biotechnology Information, National Library of Medicine, MEDLINE"><meta name="robots" content="index,nofollow,noarchive"><meta property="og:title" content="Glycans in the immune system and The Altered Glycan Theory of Autoimmunity: a critical review - PubMed"><meta property="og:url" content="https://pubmed.ncbi.nlm.nih.gov/25578468/"><meta property="og:description" content="Herein we will review the role of glycans in the immune system. 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Specific topics covered include: the glycosylation sites of IgE, IgM, IgD, IgE, IgA, and IgG; how glycans can encode "self" identity by functioning as either danger associated molecular patterns (DAMPs) or self-associated molecular patt …"><meta name="twitter:image" content="https://cdn.ncbi.nlm.nih.gov/pubmed/persistent/pubmed-meta-image-v2.jpg">  <link rel="search" type="application/opensearchdescription+xml" href="https://cdn.ncbi.nlm.nih.gov/pubmed/persistent/opensearch.xml" title="PubMed search">  <noscript> <link rel="stylesheet" type="text/css" href="https://cdn.ncbi.nlm.nih.gov/pubmed/14efe001-aebf-4097-9883-28c967333423/core/no-script.css"> </noscript> <meta name="citation_title" content="Glycans in the immune system and The Altered Glycan Theory of Autoimmunity: a critical review"> <meta name="citation_authors" content="Maverakis E;Kim K;Shimoda M;Gershwin ME;Patel F;Wilken R;Raychaudhuri S;Ruhaak LR;Lebrilla CB;"> <meta name="citation_date" content="2015 Feb"> <meta name="citation_publisher" content="J Autoimmun"> <meta name="citation_journal_title" content="Journal of autoimmunity"> <meta name="citation_pmid" content="25578468"> <meta name="citation_abstract_html_url" content="https://pubmed.ncbi.nlm.nih.gov/25578468/"> <meta name="citation_volume" content="57"> <meta name="citation_doi" content="10.1016/j.jaut.2014.12.002"> <meta name="citation_issn" content="1095-9157"> <link rel="canonical" href="https://pubmed.ncbi.nlm.nih.gov/25578468/" /> </head> <body > <noscript> <div class="no-script-banner" id="no-script-banner"> <div class="warning-message"> <div class="warning-message-text"> This site needs JavaScript to work properly. 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new collection</label> </li> <li> <input type="radio" id="collections-action-panel-existing" class="collections-existing" name="collections" value="existing" checked="true" data-ga-category="save_share" data-ga-action="collections" data-ga-label="collections_radio_existing"> <label for="collections-action-panel-existing">Add to an existing collection</label> </li> </ul> </div> <div class="controls-wrapper"> <div class="action-panel-control-wrap new-collections-controls"> <label for="collections-action-panel-add-to-new" class="action-panel-label required-field-asterisk"> Name your collection: </label> <input type="text" name="add-to-new-collection" id="collections-action-panel-add-to-new" class="collections-action-add-to-new" pattern="[^"&=<>\/]*" title="The following characters are not allowed in the Name field: "&=<>/" maxlength="100" data-ga-category="save_share" data-ga-action="create_collection" data-ga-label="non_favorties_collection"> <div class="collections-new-name-too-long usa-input-error-message selection-validation-message"> Name must be less than 100 characters </div> </div> <div class="action-panel-control-wrap existing-collections-controls"> <label for="collections-action-panel-add-to-existing" class="action-panel-label"> Choose a collection: </label> <select id="collections-action-panel-add-to-existing" class="action-panel-selector collections-action-add-to-existing" data-ga-category="save_share" data-ga-action="select_collection" data-ga-label="($('#collections-action-add-to-existing').val() === 'Favorites') ? 'Favorites' : 'non_favorites_collection'"> </select> <div class="collections-retry-load-on-error usa-input-error-message selection-validation-message"> Unable to load your collection due to an error<br> <a href="#">Please try again</a> </div> </div> </div> <div class="action-panel-actions"> <button class="action-panel-submit" type="submit" data-loading-label="Adding..." data-pinger-ignore data-ga-category="save_share" data-ga-action="collections" data-ga-label="add"> Add </button> <button class="action-panel-cancel" aria-label="Close 'Add to Collections' panel" ref="linksrc=close_collections_panel" aria-controls="collections-action-panel" aria-expanded="false" data-ga-category="save_share" data-ga-action="collections" data-ga-label="cancel"> Cancel </button> </div> </form> </div> </div> <div id="bibliography-action-panel" class="bibliography-action-panel action-panel in-progress-dots-panel" aria-hidden="true" data-bibliography-open-panel-enabled="false" data-bibliography-open-panel-url-hash="#open-bibliography-panel"> <div class="inner-wrap"> <h3 class="action-panel-heading"> Add to My Bibliography </h3> <form id="bibliography-action-panel-form" class="bibliography-action-panel-form action-panel-content action-form action-panel-smaller-selectors" data-add-to-bibliography-max-amount="100" data-add-to-bibliography-batch-size="10" data-bibliography-delegates-url="/list-bibliography-delegates/" data-add-to-bibliography-url="/add-to-bibliography/" data-get-article-ids-by-search-url="/get-article-ids-by-search/" data-mybib-root-url="https://www.ncbi.nlm.nih.gov/myncbi/collections/mybibliography/"> <input type="hidden" name="csrfmiddlewaretoken" value="DGG6h7WrPt6Y7j1WHwUypst0d68FZnOgUAbiTMp37ygRwpwo2z2ayVyZN3IUaB34"> <div class="action-panel-control-wrap bibliographies-controls"> <div class="choice-group"> <ul class="bibliographies-action-add radio-group-items"> <li> <input name="bibliography" id="my-bibliography" class="my-bibliography" type="radio" checked/> <label for="my-bibliography">My Bibliography</label> </li> </ul> </div> </div> <div class="bibliographies-retry-load-on-error usa-input-error-message selection-validation-message"> Unable to load your delegates due to an error<br> <a href="#">Please try again</a> </div> <div class="action-panel-actions"> <button class="action-panel-submit" type="submit" data-loading-label="Adding..." data-pinger-ignore> Add </button> <button class="action-panel-cancel" aria-label="Close 'Add to bibliography' panel" ref="linksrc=close_bibliography_panel" aria-controls="bibliography-action-panel" aria-expanded="false" data-ga-category="save_share" data-ga-action="mybib" data-ga-label="cancel"> Cancel </button> </div> </form> </div> </div> <div id="saved-search-action-panel" class="saved-search-action-panel action-panel " aria-hidden="true" data-saved-search-open-panel-enabled="false" data-saved-search-open-panel-url-hash="#open-saved-search-panel"> <div class="inner-wrap"> <h2 class="action-panel-heading"> Your saved search </h2> <form id="saved-search-action-panel-form" class="saved-search-action-panel-form action-panel-content action-form" data-create-saved-search-url="/create-saved-search/" data-try-search-terms-url="/try-search-term/" data-saved-search-root-url="https://www.ncbi.nlm.nih.gov/myncbi/searches/"> <input type="hidden" name="csrfmiddlewaretoken" value="DGG6h7WrPt6Y7j1WHwUypst0d68FZnOgUAbiTMp37ygRwpwo2z2ayVyZN3IUaB34"> <div class="action-panel-control-wrap"> <label for="saved-search-name" class="action-panel-label saved-search-name-label required-field-asterisk"> Name of saved search: </label> <input maxlength="200" type="text" name="saved-search-name" id="saved-search-name" class="saved-search-name" value="" required pattern="[^"&=<>\/]*" title="The following characters are not allowed in the Name field: "&=<>/"> </div> <div class="action-panel-control-wrap"> <label for="saved-search-term" class="action-panel-label required-field-asterisk"> Search terms: </label> <textarea name="saved-search-term" id="saved-search-term" class="saved-search-term" required></textarea> </div> <div class="test-search-term-wrap"> <a href="#" class="try-search-term">Test search terms</a> </div> <div class="choice-group action-panel-extra-margin-top"> <span class="action-panel-label" id="fieldset-label"> Would you like email updates of new search results? </span> <fieldset id="saved-search-alert" aria-describedby="fieldset-label"> <legend class="usa-sr-only">Saved Search Alert Radio Buttons</legend> <ul class="radio-group-items"> <li> <input type="radio" id="saved-search-alert-yes" class="saved-search-alert-yes" name="saved-search-alert" value="yes" checked> <label for="saved-search-alert-yes" class="action-panel-label">Yes</label> </li> <li> <input aria-label="No radio input" type="radio" id="saved-search-alert-no" class="saved-search-alert-no" name="saved-search-alert" value="no"> <label for="saved-search-alert-no" class="action-panel-label">No</label> </li> </ul> </fieldset> </div> <div class="alert-schedule-wrap"> <div class="action-panel-control-wrap"> <label class="action-panel-label"> Email: </label> <span aria-label="Email address" id="saved-search-email" class="action-panel-label"><span class="action-panel-label-bold"></span> (<a class="myncbi-account-settings" href="https://www.ncbi.nlm.nih.gov/account/settings/">change</a>)</span> </div> <div class="action-panel-control-wrap action-panel-extra-margin-top"> <label for="saved-search-frequency" class="action-panel-label"> Frequency: </label> <select id="saved-search-frequency" class="no-border-panel-selector saved-search-frequency"> <option value="monthly">Monthly</option> <option value="weekly">Weekly</option> <option value="daily">Daily</option> </select> </div> <div class="action-panel-control-wrap saved-search-monthly-additional"> <label for="saved-search-monthly-on-day" class="action-panel-label"> Which day? </label> <select id="saved-search-monthly-on-day" class="no-border-panel-selector"> <option value="Sunday">The first Sunday</option> <option value="Monday">The first Monday</option> <option value="Tuesday">The first Tuesday</option> <option value="Wednesday">The first Wednesday</option> <option value="Thursday">The first Thursday</option> <option value="Friday">The first Friday</option> <option value="Saturday">The first Saturday</option> <option value="day">The first day</option> <option value="weekday">The first weekday</option> </select> </div> <div class="action-panel-control-wrap saved-search-weekly-additional"> <label for="saved-search-weekly-on-day" class="action-panel-label"> Which day? </label> <select id="saved-search-weekly-on-day" class="no-border-panel-selector saved-search-weekly-on-day"> <option value="Sunday">Sunday</option> <option value="Monday">Monday</option> <option value="Tuesday">Tuesday</option> <option value="Wednesday">Wednesday</option> <option value="Thursday">Thursday</option> <option value="Friday">Friday</option> <option value="Saturday">Saturday</option> </select> </div> <div class="action-panel-control-wrap"> <label for="saved-search-report" class="action-panel-label"> Report format: </label> <select id="saved-search-report" class="no-border-panel-selector saved-search-report"> <option value="DocSum">Summary</option> <option value="DocSumText">Summary (text)</option> <option value="Abstract">Abstract</option> <option value="AbstractText">Abstract (text)</option> <option value="MEDLINE">PubMed</option> </select> </div> <div class="action-panel-control-wrap"> <label for="saved-search-amount" class="action-panel-label"> Send at most: </label> <select id="saved-search-amount" class="no-border-panel-selector saved-search-amount"> <option value="1">1 item</option> <option value="5" selected>5 items</option> <option value="10">10 items</option> <option value="20">20 items</option> <option value="50">50 items</option> <option value="100">100 items</option> <option value="200">200 items</option> </select> </div> <div> <input type="checkbox" id="saved-search-send-if-no-result" class="saved-search-send-if-no-result" name="saved-search-send-if-no-result"> <label for="saved-search-send-if-no-result" class="action-panel-label smaller-checkbox"> Send even when there aren't any new results </label> </div> <div class="action-panel-control-wrap option-text-in-email-wrap"> <label for="saved-search-email-text" class="action-panel-label"> Optional text in email: </label> <textarea name="saved-search-email-text" id="saved-search-email-text" class="saved-search-email-text"></textarea> </div> </div> <div class="action-panel-actions"> <button class="action-panel-submit" type="submit" data-loading-label="Saving..." data-ga-category="save_share" data-ga-action="alert" data-ga-label="save"> Save </button> <button class="action-panel-cancel" aria-label="Close 'Your saved search' panel" ref="linksrc=close_saved_search_panel" aria-controls="saved-search-action-panel" aria-expanded="false" data-ga-category="save_share" data-ga-action="alert" data-ga-label="cancel"> Cancel </button> </div> </form> </div> </div> <div id="citation-manager-action-panel" class="citation-manager-action-panel action-panel" aria-hidden="true"> <div class="inner-wrap"> <h2 class="action-panel-heading"> Create a file for external citation management software </h2> <form id="citation-manager-action-panel-form" class="action-panel-content action-form" action="/results-export-ids/" data-by-search-action="/results-export-search-data/" data-by-ids-action="/results-export-ids/" method="post" data-by-search-method="post" data-by-ids-method="post"> <input type="hidden" name="csrfmiddlewaretoken" value="DGG6h7WrPt6Y7j1WHwUypst0d68FZnOgUAbiTMp37ygRwpwo2z2ayVyZN3IUaB34"> <input name="results-format" type="hidden" value="pubmed"/> <div class="action-panel-actions"> <button class="action-panel-submit" type="submit" data-loading-label="Sending..." data-ga-category="save_share" data-ga-action="citation_manager" data-ga-label="save"> Create file </button> <button class="action-panel-cancel" aria-label="Close 'Send citations to citation manager' panel" ref="linksrc=close_citation_manager_panel" aria-controls="citation-manager-action-panel" aria-expanded="false" data-ga-category="save_share" data-ga-action="citation_manager" data-ga-label="cancel"> Cancel </button> </div> </form> </div> </div> <div id="rss-action-panel" class="rss-action-panel action-panel " aria-hidden="true"> <div class="inner-wrap"> <h2 class="action-panel-heading"> Your RSS Feed </h2> <form id="rss-action-panel-form" class="rss-action-panel-form action-panel-content action-form" data-create-rss-feed-url="/create-rss-feed-url/" data-search-form-term-value=""> <input type="hidden" name="csrfmiddlewaretoken" value="DGG6h7WrPt6Y7j1WHwUypst0d68FZnOgUAbiTMp37ygRwpwo2z2ayVyZN3IUaB34"> <div class="action-panel-control-wrap"> <label for="rss-name" class="action-panel-label required-field-asterisk"> Name of RSS Feed: </label> <input maxlength="200" placeholder="Name" type="text" name="rss-name" id="rss-name" class="rss-name" value='' required pattern="[^"&=<>\/]*" title="The following characters are not allowed in the Name field: "&=<>/"> </div> <div class="rss-limit-wrap"> <div class="action-panel-control-wrap action-panel-extra-margin-top"> <label for="rss-limit" class="action-panel-label"> Number of items displayed: </label> <select id="rss-limit" class="no-border-panel-selector rss-limit"> <option value="5">5</option> <option value="10">10</option> <option value="15" selected="selected">15</option> <option value="20">20</option> <option value="50">50</option> <option value="100">100</option> </select> </div> </div> <div class="action-panel-actions"> <button class="action-panel-submit" type="submit" data-loading-label="Creating..." data-ga-category="save_share" data-ga-action="alert" data-ga-label="save"> Create RSS </button> <button class="action-panel-cancel" aria-label="Close 'Your RSS' panel" ref="linksrc=close_rss_panel" aria-controls="rss-action-panel" aria-expanded="false" data-ga-category="save_share" data-ga-action="alert" data-ga-label="cancel"> Cancel </button> </div> <div class="action-panel-control-wrap rss-link-copy-wrap"> <label for="rss-link" class="usa-sr-only">RSS Link</label> <input placeholder="Your RSS Feed Link" type="text" name="rss-link" id="rss-link" class="rss-link" title="RSS Link"> <button type="button" disabled class="rss-link-copy-button disabled" data-ga-category="save_share" data-ga-action="rss" data-ga-label="copy"> Copy </button> </div> </form> </div> </div> </div> </div> <div class="article-page" id="article-page" data-article-pmid="25578468"> <aside class="page-sidebar"> <div class="inner-wrap"> <div class="full-text-links"> <div class="full-view"> <h3 class="title"> Full text links </h3> <div class="full-text-links-list"> <a class="link-item dialog-focus" 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name="csrfmiddlewaretoken" value="DGG6h7WrPt6Y7j1WHwUypst0d68FZnOgUAbiTMp37ygRwpwo2z2ayVyZN3IUaB34"><div class="choice-group" role="radiogroup"><ul class="radio-group-items"><li><input type="radio" id="collections-action-dialog-new" class="collections-new" name="collections" value="new" data-ga-category="collections_button" data-ga-action="click" data-ga-label="collections_radio_new"><label for="collections-action-dialog-new">Create a new collection</label></li><li><input type="radio" id="collections-action-dialog-existing" class="collections-existing" name="collections" value="existing" checked="true" data-ga-category="collections_button" data-ga-action="click" data-ga-label="collections_radio_existing"><label for="collections-action-dialog-existing">Add to an existing collection</label></li></ul></div><div class="controls-wrapper"><div class="action-panel-control-wrap new-collections-controls"><label for="collections-action-dialog-add-to-new" class="action-panel-label required-field-asterisk"> Name your collection: </label><input type="text" name="add-to-new-collection" id="collections-action-dialog-add-to-new" class="collections-action-add-to-new" pattern="[^"&=<>\/]*" title="The following characters are not allowed in the Name field: "&=<>/" maxlength="100" data-ga-category="collections_button" data-ga-action="create_collection" data-ga-label="non_favorties_collection"><div class="collections-new-name-too-long usa-input-error-message selection-validation-message"> Name must be less than 100 characters </div></div><div class="action-panel-control-wrap existing-collections-controls"><label for="collections-action-dialog-add-to-existing" class="action-panel-label"> Choose a collection: </label><select id="collections-action-dialog-add-to-existing" class="action-panel-selector collections-action-add-to-existing" data-ga-category="collections_button" data-ga-action="select_collection" data-ga-label="($('.collections-action-add-to-existing').val() === 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'Favorites' : 'non_favorites_collection'"></select><div class="collections-retry-load-on-error usa-input-error-message selection-validation-message"> Unable to load your collection due to an error<br><a href="#">Please try again</a></div></div></div><div class="action-panel-actions"><button class="action-panel-submit" type="submit" data-loading-label="Adding..." data-pinger-ignore data-ga-category="collections_button" data-ga-action="click" data-ga-label="add"> Add </button><button class="action-panel-cancel" aria-label="Close 'Add to Collections' panel" ref="linksrc=close_collections_panel" aria-controls="collections-action-panel" aria-expanded="false" data-ga-category="collections_button" data-ga-action="click" data-ga-label="cancel"> Cancel </button></div></form></div></div></div></div><button class="more-actions-button more-actions-dialog-trigger" title="Open dialog with more actions to take" ref="linksrc=more_actions_btn"></button><div class="display-options" 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autoimmunity" data-pinger-ignore> J Autoimmun </button><div id="full-view-journal" class="journal-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for journal Journal of autoimmunity"><div class="title">Actions</div><div class="content"><ul class="journal-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link" href="/?term=%22J+Autoimmun%22%5Bjour%5D&sort=date&sort_order=desc" data-href="/?term=%22J+Autoimmun%22%5Bjour%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_journal_name_link&journal_abbrev=J Autoimmun" data-ga-category="search" data-ga-action="journal_link" data-ga-label="J Autoimmun"> Search in PubMed </a></li><li><a class="search-in-nlm-catalog-link dropdown-block-link" ref ="linksrc=search_in_nlm_catalog_link" href="https://www.ncbi.nlm.nih.gov/nlmcatalog?term=%22J+Autoimmun%22%5BTitle+Abbreviation%5D" data-ga-category="search_catalog" data-ga-action="journal_link" data-ga-label="J Autoimmun"> Search in NLM Catalog </a></li><li><a class="add-to-search-link dropdown-block-link" ref ="linksrc=add_to_search_link" role="button" data-search-term='"J Autoimmun"[jour]' data-ga-category="search" data-ga-action="add_to_search" data-ga-label='"J Autoimmun"[jour]' href="#"> Add to Search </a></li></ul></div></div></div><span class="period">. </span><span class="cit">2015 Feb:57:1-13.</span> </div> <span class="citation-doi"> doi: 10.1016/j.jaut.2014.12.002. </span> <span class="secondary-date"> Epub 2015 Jan 9. </span> </div> <h1 class="heading-title"> Glycans in the immune system and The Altered Glycan Theory of Autoimmunity: a critical review </h1> <div class="inline-authors"> <div class="authors"> <div class="authors-list"> <span class="authors-list-item "><a class="full-name" href="/?term=Maverakis+E&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Emanual Maverakis">Emanual Maverakis</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Dermatology, University of California, Davis School of Medicine, 3301 C Street, Suite 1400, Sacramento, CA 95816, USA. Electronic address: emaverakis@ucdavis.edu." href="#full-view-affiliation-1" ref="linksrc=author_aff"> 1 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Kim+K&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Kyoungmi Kim">Kyoungmi Kim</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Public Health Sciences, Division of Biostatistics, University of California, Davis Medical Center, Sacramento, CA 95816, USA." href="#full-view-affiliation-2" ref="linksrc=author_aff"> 2 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Shimoda+M&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Michiko Shimoda">Michiko Shimoda</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Dermatology, University of California, Davis School of Medicine, 3301 C Street, Suite 1400, Sacramento, CA 95816, USA." href="#full-view-affiliation-3" ref="linksrc=author_aff"> 3 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Gershwin+ME&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="M Eric Gershwin">M Eric Gershwin</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Internal Medicine, Division of Rheumatology, University of California, Davis School of Medicine, Sacramento, CA 95817, USA." href="#full-view-affiliation-4" ref="linksrc=author_aff"> 4 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Patel+F&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Forum Patel">Forum Patel</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Dermatology, University of California, Davis School of Medicine, 3301 C Street, Suite 1400, Sacramento, CA 95816, USA." href="#full-view-affiliation-3" ref="linksrc=author_aff"> 3 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Wilken+R&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Reason Wilken">Reason Wilken</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Dermatology, University of California, Davis School of Medicine, 3301 C Street, Suite 1400, Sacramento, CA 95816, USA." href="#full-view-affiliation-3" ref="linksrc=author_aff"> 3 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Raychaudhuri+S&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Siba Raychaudhuri">Siba Raychaudhuri</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Internal Medicine, Division of Rheumatology, University of California, Davis School of Medicine, Sacramento, CA 95817, USA." href="#full-view-affiliation-4" ref="linksrc=author_aff"> 4 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Ruhaak+LR&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="L Renee Ruhaak">L Renee Ruhaak</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Public Health Sciences, Division of Biostatistics, University of California, Davis Medical Center, Sacramento, CA 95816, USA." href="#full-view-affiliation-2" ref="linksrc=author_aff"> 2 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Lebrilla+CB&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Carlito B Lebrilla">Carlito B Lebrilla</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Chemistry, University of California, Davis, One Shields Avenue, Davis, CA 95616, USA." href="#full-view-affiliation-5" ref="linksrc=author_aff"> 5 </a></sup></span> </div> </div> </div> <div class="short-article-details"> Affiliations <button class="more-details" id="toggle-authors" data-alt-text="Collapse" 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Electronic address: emaverakis@ucdavis.edu.</li> <li data-affiliation-id="full-view-affiliation-2" id="full-view-affiliation-2" ><sup class="key">2</sup> Department of Public Health Sciences, Division of Biostatistics, University of California, Davis Medical Center, Sacramento, CA 95816, USA.</li> <li data-affiliation-id="full-view-affiliation-3" id="full-view-affiliation-3" ><sup class="key">3</sup> Department of Dermatology, University of California, Davis School of Medicine, 3301 C Street, Suite 1400, Sacramento, CA 95816, USA.</li> <li data-affiliation-id="full-view-affiliation-4" id="full-view-affiliation-4" ><sup class="key">4</sup> Department of Internal Medicine, Division of Rheumatology, University of California, Davis School of Medicine, Sacramento, CA 95817, USA.</li> <li data-affiliation-id="full-view-affiliation-5" id="full-view-affiliation-5" ><sup class="key">5</sup> Department of Chemistry, University of California, Davis, One Shields Avenue, Davis, CA 95616, USA.</li> </ul> </div> </div> </div> <ul class="identifiers" id="full-view-identifiers"> <li> <span class="identifier pubmed"> <span class="id-label"> PMID: </span> <strong class="current-id" title="PubMed ID">25578468</strong> </span> </li> <li> <span class="identifier pmc"> <span class="id-label"> PMCID: </span> <a class="id-link" target="_blank" rel="noopener" ref="linksrc=article_id_link&article_id=PMC4340844&id_type=PMC" href="https://pmc.ncbi.nlm.nih.gov/articles/PMC4340844/" data-ga-category="full_text" data-ga-action="PMCID" > PMC4340844 </a> </span> </li> <li> <span class="identifier doi"> <span class="id-label"> DOI: </span> <a class="id-link" target="_blank" rel="noopener" ref="linksrc=article_id_link&article_id=10.1016/j.jaut.2014.12.002&id_type=DOI" href="https://doi.org/10.1016/j.jaut.2014.12.002" data-ga-category="full_text" data-ga-action="DOI" > 10.1016/j.jaut.2014.12.002 </a> </span> </li> </ul> <div class="in-clipboard-label " hidden> Item in Clipboard </div> </div> <div class="short-view" id="short-view-heading"> <div class="publication-type" >Review</div> <h1 class="heading-title"> Glycans in the immune system and The Altered Glycan Theory of Autoimmunity: a critical review </h1> <div class="short-citation"> <span class="authors-list"> <span class="authors-list-item "><span class="full-name">Emanual Maverakis</span><span class="citation-separator"> et al.</span></span> </span> <span class="citation-journal"> J Autoimmun<span class="citation-separator">.</span> </span> <span class="date"> <span class="citation-year">2015 Feb</span><span class="citation-separator">.</span> </span> </div> <div class="more-article-details-section with-display-settings"> <button class="more-article-details" id="toggle-article-details" data-alt-text="Hide details" ref="linksrc=toggle_article_details" aria-controls="full-article-details" aria-expanded="false" data-fetch-url="/25578468/long-authors/" data-id-prefix=""> <span class="title">Show details</span> </button> <div 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href="/?term=Maverakis+E&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Emanual Maverakis">Emanual Maverakis</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Dermatology, University of California, Davis School of Medicine, 3301 C Street, Suite 1400, Sacramento, CA 95816, USA. Electronic address: emaverakis@ucdavis.edu." href="#short-view-affiliation-1" ref="linksrc=author_aff"> 1 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Kim+K&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Kyoungmi Kim">Kyoungmi Kim</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Public Health Sciences, Division of Biostatistics, University of California, Davis Medical Center, Sacramento, CA 95816, USA." href="#short-view-affiliation-2" ref="linksrc=author_aff"> 2 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Shimoda+M&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Michiko Shimoda">Michiko Shimoda</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Dermatology, University of California, Davis School of Medicine, 3301 C Street, Suite 1400, Sacramento, CA 95816, USA." href="#short-view-affiliation-3" ref="linksrc=author_aff"> 3 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Gershwin+ME&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="M Eric Gershwin">M Eric Gershwin</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Internal Medicine, Division of Rheumatology, University of California, Davis School of Medicine, Sacramento, CA 95817, USA." href="#short-view-affiliation-4" ref="linksrc=author_aff"> 4 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Patel+F&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Forum Patel">Forum Patel</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Dermatology, University of California, Davis School of Medicine, 3301 C Street, Suite 1400, Sacramento, CA 95816, USA." href="#short-view-affiliation-3" ref="linksrc=author_aff"> 3 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Wilken+R&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Reason Wilken">Reason Wilken</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Dermatology, University of California, Davis School of Medicine, 3301 C Street, Suite 1400, Sacramento, CA 95816, USA." href="#short-view-affiliation-3" ref="linksrc=author_aff"> 3 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Raychaudhuri+S&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Siba Raychaudhuri">Siba Raychaudhuri</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Internal Medicine, Division of Rheumatology, University of California, Davis School of Medicine, Sacramento, CA 95817, USA." href="#short-view-affiliation-4" ref="linksrc=author_aff"> 4 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Ruhaak+LR&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="L Renee Ruhaak">L Renee Ruhaak</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Public Health Sciences, Division of Biostatistics, University of California, Davis Medical Center, Sacramento, CA 95816, USA." href="#short-view-affiliation-2" ref="linksrc=author_aff"> 2 </a></sup><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Lebrilla+CB&cauthor_id=25578468" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Carlito B Lebrilla">Carlito B Lebrilla</a><sup class="affiliation-links"><span class="author-sup-separator"> </span><a class="affiliation-link" title="Department of Chemistry, University of California, Davis, One Shields Avenue, Davis, CA 95616, USA." href="#short-view-affiliation-5" ref="linksrc=author_aff"> 5 </a></sup></span> </div> </div> <div class="affiliations"> <h3 class="title"> Affiliations </h3> <ul class="item-list"> <li data-affiliation-id="short-view-affiliation-1" id="short-view-affiliation-1" ><sup class="key">1</sup> Department of Dermatology, University of California, Davis School of Medicine, 3301 C Street, Suite 1400, Sacramento, CA 95816, USA. Electronic address: emaverakis@ucdavis.edu.</li> <li data-affiliation-id="short-view-affiliation-2" id="short-view-affiliation-2" ><sup class="key">2</sup> Department of Public Health Sciences, Division of Biostatistics, University of California, Davis Medical Center, Sacramento, CA 95816, USA.</li> <li data-affiliation-id="short-view-affiliation-3" id="short-view-affiliation-3" ><sup class="key">3</sup> Department of Dermatology, University of California, Davis School of Medicine, 3301 C Street, Suite 1400, Sacramento, CA 95816, USA.</li> <li data-affiliation-id="short-view-affiliation-4" id="short-view-affiliation-4" ><sup class="key">4</sup> Department of Internal Medicine, Division of Rheumatology, University of California, Davis School of Medicine, Sacramento, CA 95817, USA.</li> <li data-affiliation-id="short-view-affiliation-5" id="short-view-affiliation-5" ><sup class="key">5</sup> Department of Chemistry, University of California, Davis, One Shields Avenue, Davis, CA 95616, USA.</li> </ul> </div> </div> <ul class="identifiers" id="short-view-identifiers"> <li> <span class="identifier pubmed"> <span class="id-label"> PMID: </span> <strong class="current-id" title="PubMed ID">25578468</strong> </span> </li> <li> <span class="identifier pmc"> <span class="id-label"> PMCID: </span> <a class="id-link" target="_blank" rel="noopener" ref="linksrc=article_id_link&article_id=PMC4340844&id_type=PMC" href="https://pmc.ncbi.nlm.nih.gov/articles/PMC4340844/" data-ga-category="full_text" data-ga-action="PMCID" > PMC4340844 </a> </span> </li> <li> <span class="identifier doi"> <span class="id-label"> DOI: </span> <a class="id-link" target="_blank" rel="noopener" ref="linksrc=article_id_link&article_id=10.1016/j.jaut.2014.12.002&id_type=DOI" href="https://doi.org/10.1016/j.jaut.2014.12.002" data-ga-category="full_text" data-ga-action="DOI" > 10.1016/j.jaut.2014.12.002 </a> </span> </li> </ul> <div class="in-clipboard-label " hidden> Item in Clipboard </div> </div> </div> </header> <div class="actions-buttons inline"><div class="inner-wrap"><button class="full-text-links-button full-text-links-dialog-trigger" title="Open dialog with full text links" ref="linksrc=full_text_links"> Full text links </button><button class="citation-button citation-dialog-trigger" aria-label="Open dialog with citation text in different styles" data-ga-category="save_share" data-ga-action="cite" data-ga-label="open" data-all-citations-url="/25578468/citations/" data-citation-style="nlm" data-pubmed-format-link="/25578468/export/"><span class="button-label">Cite</span></button><button class="more-actions-button more-actions-dialog-trigger" title="Open dialog with more actions to take" ref="linksrc=more_actions_btn"></button><div class="display-options" data-format-key="format" data-default-format-value="abstract" data-current-format-value="abstract" role="region"><button class="trigger usa-button" title="Change format" data-ga-category="display_options" data-ga-action="display_options_dialog"><span class="button-label">Display options</span></button><div class="dropdown" hidden><div class="title"> Display options </div><div class="content"><div class="format-container"><span class="option-label"> Format </span><select name="format" data-initial-value="abstract" aria-label="Display format" id="inline-actions-display-format"><option value="abstract">Abstract</option><option value="pubmed">PubMed</option><option value="pmid">PMID</option></select></div></div></div></div></div></div> <div class="abstract" id="abstract"> <h2 class="title"> Abstract </h2> <div class="abstract-content selected" id="eng-abstract"> <p> Herein we will review the role of glycans in the immune system. Specific topics covered include: the glycosylation sites of IgE, IgM, IgD, IgE, IgA, and IgG; how glycans can encode "self" identity by functioning as either danger associated molecular patterns (DAMPs) or self-associated molecular patterns (SAMPs); the role of glycans as markers of protein integrity and age; how the glycocalyx can dictate the migration pattern of immune cells; and how the combination of Fc N-glycans and Ig isotype dictate the effector function of immunoglobulins. We speculate that the latter may be responsible for the well-documented association between alterations of the serum glycome and autoimmunity. Due to technological limitations, the extent of these autoimmune-associated glycan alterations and their role in disease pathophysiology has not been fully elucidated. Thus, we also review the current technologies available for glycan analysis, placing an emphasis on Multiple Reaction Monitoring (MRM), a rapid high-throughput technology that has great potential for glycan biomarker research. Finally, we put forth The Altered Glycan Theory of Autoimmunity, which states that each autoimmune disease will have a unique glycan signature characterized by the site-specific relative abundances of individual glycan structures on immune cells and extracellular proteins, especially the site-specific glycosylation patterns of the different immunoglobulin(Ig) classes and subclasses. </p> </div> <p> <strong class="sub-title"> Keywords: </strong> Autoimmunity; Glycan; Glycome; Glycosylation; Immunoglobulin. </p> </div> <p class="copyright" id="copyright"> Copyright © 2014 The Authors. Published by Elsevier Ltd.. All rights reserved. </p> <p class="disclaimer" id="disclaimer"> <a href="/disclaimer/" target="_blank" data-ga-category="literature_resources" data-ga-action="disclaimer_link" data-ga-label="under_abstract">PubMed Disclaimer</a> </p> <div class="figures" id="figures"> <h2 class="title"> Figures </h2> <div class="figures-list" id="slides-container" itemscope itemtype="http://schema.org/ImageGallery"> <figure class="figure-item " itemscope itemtype="http://schema.org/ImageObject" itemprop="associatedMedia" data-slide-index="0" data-label-slug="figure-1"> <a class="figure-link" href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/4cfabc6562fe/nihms654899f1.jpg" itemprop="contentUrl" aria-describedby="figure-caption-0" role="button" data-image-width="611" data-image-height="589" data-image-alt="Figure 1" data-pmc-id="PMC4340844" data-figure-id="F1"> <img class="figure-thumb" itemprop="thumbnail" id="article-image-0" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/71a4f9a19cbb/nihms654899f1.gif" alt="Figure 1" /> </a> <meta itemprop="width" itemtype="http://schema.org/ImageObject" content="611"> <meta itemprop="height" itemtype="http://schema.org/ImageObject" content="589"> <div class="figure-caption figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="caption" aria-hidden="true"> <div class="caption-wrap"> <strong class="figure-label"> <p> Figure 1. Immunoglobulin Isotypes and their sites… </p> </strong> </div> </div> <div class="figure-caption-medium figure-caption-text" aria-hidden="true"> <strong class="figure-label"> <p> Figure 1. Immunoglobulin Isotypes and their sites of glycosylation </p> </strong> <div class="figure-caption-contents"><p> Depicted here are the antibody structures… </p></div> </div> <figcaption id="figure-caption-0" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description"> <strong class="figure-label"> Figure 1. Immunoglobulin Isotypes and their sites of glycosylation </strong> <div class="figure-caption-contents">Depicted here are the antibody structures including their sites of glycosylation: IgM [(N-glycans at Asn-46, 209, 272, 279, and 439 (UNIPROT), or CH1-45, CH2-120, CH3-81, <b>CH3-84.4</b>, and CHS-7 (IMGT)]; IgA1 [(N-glycans at Asn-144 and 352 (UNIPROT) or CH2-20, CHS-7 (IMGT)]; IgA2 [(N-glycans at Asn-47, 92, 131, 205, 327 (UNIPROT) or CH1-45.2, CH1-114, CH2-20, CH2-120 and CHS-7 (IMGT)]; IgG [(N-glycans at Asn-180 (IgG1), Asn-176 (IgG2), Asn-227 (IgG3), Asn-177 (IgG4), and Asn-322 (IgG3) (UNIPROT) or <b>CH2-84.4</b> (IgG1-4) and CH3- 79 (IgG3) (IMGT)]; IgD [(N-glycans at Asn-225, 316, and 367 (UNIPROT) or <b>CH2-84.4</b>, CH3-45.4, CH3- 116 (IMGT)], and IgE [(N-glycans at Asn-21, 49, 99, 146, 252, 264, 275 (UNIPROT) or CH1-15.2, CH1-45.2, CH1-118, CH2-38, CH3-38, CH3-77, and <b>CH3-84.4</b> (IMGT)]. Each immunoglobulin is comprised of two heavy chains (blue) and two light chains (purple) that are linked together by disulfide bonds (black lines). IgA, IgD, and IgG have a flexible hinge region that link the Igs’ antigen-binding Fab region to their Fc receptor-binding region. O-glycosylation sites are depicted in yellow and N-glycosylation sites are depicted in brown. The depicted glycans are important for the structural integrity of the antibodies and their effector function [2].</div> </figcaption> </figure> <figure class="figure-item " itemscope itemtype="http://schema.org/ImageObject" itemprop="associatedMedia" data-slide-index="1" data-label-slug="figure-2"> <a class="figure-link" href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/29548ff8a654/nihms654899f2.jpg" itemprop="contentUrl" aria-describedby="figure-caption-1" role="button" data-image-width="600" data-image-height="510" data-image-alt="Figure 2" data-pmc-id="PMC4340844" data-figure-id="F2"> <img class="figure-thumb" itemprop="thumbnail" id="article-image-1" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/c988e3da5461/nihms654899f2.gif" alt="Figure 2" /> </a> <meta itemprop="width" itemtype="http://schema.org/ImageObject" content="600"> <meta itemprop="height" itemtype="http://schema.org/ImageObject" content="510"> <div class="figure-caption figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="caption" aria-hidden="true"> <div class="caption-wrap"> <strong class="figure-label"> <p> Figure 2. A limited number of sugar… </p> </strong> </div> </div> <div class="figure-caption-medium figure-caption-text" aria-hidden="true"> <strong class="figure-label"> <p> Figure 2. A limited number of sugar monomers can create thousands of complex glycans </p> </strong> <div class="figure-caption-contents"><p> Post-translational… </p></div> </div> <figcaption id="figure-caption-1" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description"> <strong class="figure-label"> Figure 2. A limited number of sugar monomers can create thousands of complex glycans </strong> <div class="figure-caption-contents">Post-translational glycan modifications are generally thought to be important for protein folding; steric protection from proteolytic degradation; and regulation of protein-protein interactions. It is estimated that up to 70% of mammalian proteins are glycosylated. The glycans are attached to proteins via “N” or “O” linkages, with N-glycosylations being more common. N-glycans are attached to asparagine (Asn) residues, whereas O-glycans are attached to amino acids serine (Ser) or threonine (Thr). Depicted here is the process of N-glycosylation, which begins in the endoplasmic reticulum (ER) and ends in the Golgi. N-glycans are attached to proteins at specific motifs; Asparagine-X-Serine or Asparagine-X-threonine, where X can be any amino acid except proline. During the process of N-glycosylation, monosaccharides (often donated by UDP or GDP-sugars) are sequentially added to the glycan structure. Initially, two N-Acetylglucosamine residues are added consecutively to Dol in the cytosol. This is followed by the addition of several mannose (Man) residues. After formation of the intermediate (Man5HexNAc2-PP-Dol), the complex is flipped into the ER-lumen. Then, four additional Man residues are added. This is followed by the addition of 3 glucose (Glc) residues, donated by Glc-P-dolichol, to form the Glc3Man9GlcNAc2-PP-dolichol precursor glycan, which is then transferred to an Asn residue on a newly synthesized protein. Glycosidases and glycosyltransferases then modify the precursor glycan to potentially generate over 10 thousand unique structures, which can be separated into three very broad structural categories (High Mannose, Hybrid, and Complex).</div> </figcaption> </figure> <figure class="figure-item " itemscope itemtype="http://schema.org/ImageObject" itemprop="associatedMedia" data-slide-index="2" data-label-slug="figure-3"> <a class="figure-link" href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/6af4713dcc4a/nihms654899f3.jpg" itemprop="contentUrl" aria-describedby="figure-caption-2" role="button" data-image-width="612" data-image-height="462" data-image-alt="Figure 3" data-pmc-id="PMC4340844" data-figure-id="F3"> <img class="figure-thumb" itemprop="thumbnail" id="article-image-2" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/431dd45c22ec/nihms654899f3.gif" alt="Figure 3" /> </a> <meta itemprop="width" itemtype="http://schema.org/ImageObject" content="612"> <meta itemprop="height" itemtype="http://schema.org/ImageObject" content="462"> <div class="figure-caption figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="caption" aria-hidden="true"> <div class="caption-wrap"> <strong class="figure-label"> <p> Figure 3. Asialoglycoprotein receptors </p> </strong> <div class="figure-caption-contents"><p> Sialylated serum proteins… </p></div> </div> </div> <div class="figure-caption-medium figure-caption-text" aria-hidden="true"> <strong class="figure-label"> <p> Figure 3. Asialoglycoprotein receptors </p> </strong> <div class="figure-caption-contents"><p> Sialylated serum proteins and cells are not recognized by asialoglycoprotein receptors… </p></div> </div> <figcaption id="figure-caption-2" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description"> <strong class="figure-label"> Figure 3. Asialoglycoprotein receptors </strong> <div class="figure-caption-contents">Sialylated serum proteins and cells are not recognized by asialoglycoprotein receptors in the liver and are thus protected from uptake and degradation. However, as an initially sialylated molecule ages it progressively loses its sialic acid moieties making it a target for asialoglycoprotein receptors. In the liver these receptors identify desialylated proteins, targeting them for uptake and degradation.</div> </figcaption> </figure> <figure class="figure-item " itemscope itemtype="http://schema.org/ImageObject" itemprop="associatedMedia" data-slide-index="3" data-label-slug="figure-4"> <a class="figure-link" href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/ec97de2306d5/nihms654899f4.jpg" itemprop="contentUrl" aria-describedby="figure-caption-3" role="button" data-image-width="626" data-image-height="432" data-image-alt="Figure 4" data-pmc-id="PMC4340844" data-figure-id="F4"> <img class="figure-thumb" itemprop="thumbnail" id="article-image-3" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/e4cacc5f9f17/nihms654899f4.gif" alt="Figure 4" /> </a> <meta itemprop="width" itemtype="http://schema.org/ImageObject" content="626"> <meta itemprop="height" itemtype="http://schema.org/ImageObject" content="432"> <div class="figure-caption figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="caption" aria-hidden="true"> <div class="caption-wrap"> <strong class="figure-label"> <p> Figure 4. Cutaneous Leukocyte Antigen (CLA) is… </p> </strong> </div> </div> <div class="figure-caption-medium figure-caption-text" aria-hidden="true"> <strong class="figure-label"> <p> Figure 4. Cutaneous Leukocyte Antigen (CLA) is a glycovariant of P-selectin glycoprotein ligand-1 (PSGL-1) </p> </strong> <div class="figure-caption-contents"><p> A)… </p></div> </div> <figcaption id="figure-caption-3" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description"> <strong class="figure-label"> Figure 4. Cutaneous Leukocyte Antigen (CLA) is a glycovariant of P-selectin glycoprotein ligand-1 (PSGL-1) </strong> <div class="figure-caption-contents">A) PSGL-1, present on the surface of T cells, binds to P-selectin, which is upregulated on endothelial cells in the setting of inflammation. PSGL-1 binding to P-selectin helps initiate leukocyte rolling. B) In contrast, PSGL-1 does not bind to E-selectin, which is present on endothelial cells within the skin. C) Skin-homing T cells up-regulate the glycosylation enzyme FucT-VII, which leads to an increase in the number of sialyl-Lewis X moieties on PSGL-1, bestowing it with the capacity to bind to E-selectin. The sialyl-Lewis X-decorated E-selectin-binding glycoform of PSGL-1 is called CLA, which also differs from PSGL-1 in that it occurs as a monomer.</div> </figcaption> </figure> <figure class="figure-item " itemscope itemtype="http://schema.org/ImageObject" itemprop="associatedMedia" data-slide-index="4" data-label-slug="figure-5"> <a class="figure-link" href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/f03994a5e063/nihms654899f5.jpg" itemprop="contentUrl" aria-describedby="figure-caption-4" role="button" data-image-width="603" data-image-height="321" data-image-alt="Figure 5" data-pmc-id="PMC4340844" data-figure-id="F5"> <img class="figure-thumb" itemprop="thumbnail" id="article-image-4" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/0f52269634dc/nihms654899f5.gif" alt="Figure 5" /> </a> <meta itemprop="width" itemtype="http://schema.org/ImageObject" content="603"> <meta itemprop="height" itemtype="http://schema.org/ImageObject" content="321"> <div class="figure-caption figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="caption" aria-hidden="true"> <div class="caption-wrap"> <strong class="figure-label"> <p> Figure 5. Fc receptors can have unique… </p> </strong> </div> </div> <div class="figure-caption-medium figure-caption-text" aria-hidden="true"> <strong class="figure-label"> <p> Figure 5. Fc receptors can have unique Ig specificities </p> </strong> <div class="figure-caption-contents"><p> FcγRI, FcγRIIA, FcγRIIIA are activating Fc… </p></div> </div> <figcaption id="figure-caption-4" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description"> <strong class="figure-label"> Figure 5. Fc receptors can have unique Ig specificities </strong> <div class="figure-caption-contents">FcγRI, FcγRIIA, FcγRIIIA are activating Fc receptors that differ in their affinities for the individual IgG subclasses; the only commonality being that each of them binds best to IgG1. Of the activating receptors, FcγRI is unique in that it can bind with high affinity to monomeric IgG antibodies. In contrast, FcγRIIA and FcγRIIIA are low affinity Fc receptors that bind only to antigen-antibody complexes. Activating signals originating from FcR-Ig interactions can be initiated by intracellular immunoreceptor tyrosine-based activation motifs (ITAMs), either within the Fc receptor or as part of the Fc receptor complex. In contrast, FcγRIIB is a low affinity inhibitory Fc receptor, which has an immunoreceptor tyrosine-based inhibition motif (ITIM) as part of its cytoplasmic domain. FcγRIIIB, found exclusively on neutrophils, is an activating glycosylphosphatidylinositol (GPI)-linked receptor.</div> </figcaption> </figure> <figure class="figure-item " itemscope itemtype="http://schema.org/ImageObject" itemprop="associatedMedia" data-slide-index="5" data-label-slug="figure-6"> <a class="figure-link" href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/49f2e9dda7b0/nihms654899f6.jpg" itemprop="contentUrl" aria-describedby="figure-caption-5" role="button" data-image-width="605" data-image-height="405" data-image-alt="Figure 6" data-pmc-id="PMC4340844" data-figure-id="F6"> <img class="figure-thumb" itemprop="thumbnail" id="article-image-5" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/96bbc6d72f31/nihms654899f6.gif" alt="Figure 6" /> </a> <meta itemprop="width" itemtype="http://schema.org/ImageObject" content="605"> <meta itemprop="height" itemtype="http://schema.org/ImageObject" content="405"> <div class="figure-caption figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="caption" aria-hidden="true"> <div class="caption-wrap"> <strong class="figure-label"> <p> Figure 6. IgG Glycoforms and their inflammatory… </p> </strong> </div> </div> <div class="figure-caption-medium figure-caption-text" aria-hidden="true"> <strong class="figure-label"> <p> Figure 6. IgG Glycoforms and their inflammatory properties </p> </strong> <div class="figure-caption-contents"><p> Ig heavy chain residue CH2-84.4 is post-translationally… </p></div> </div> <figcaption id="figure-caption-5" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description"> <strong class="figure-label"> Figure 6. IgG Glycoforms and their inflammatory properties </strong> <div class="figure-caption-contents">Ig heavy chain residue CH2-84.4 is post-translationally modified with the addition of an N-glycan, depicted here as a blue, green, and yellow Y-shaped structure between the two IgG heavy chains. This glycosylation site is conserved in all IgG subclasses (IgG1-4). To accommodate its CH2-84.4-linked glycan, IgG has a hydrophobic patch (not depicted). The CH2-84.4-linked glycan can be classified broadly as being either G0, G1, or G2. G0 glycans have a higher affinity for FcγRIII and are associated with a variety of autoimmune diseases. G0 glycans terminate with GlcNac residues and thus have zero galactose residues, hence their name. In contrast, G2 glycans terminate with two galactose residues. CH2-84.4 glycans can also be sialylated or fucosylated, which can bestow the antibody with anti-inflammatory properties because these modifications decrease Ig affinity for FcγRIII and also allow the antibody to interact with endogenous lectins on antigen presenting cells, e.g. sialylated antibodies likely bind to DC-SIGN.</div> </figcaption> </figure> <figure class="figure-item tail" itemscope itemtype="http://schema.org/ImageObject" itemprop="associatedMedia" data-slide-index="6" data-label-slug="figure-7"> <a class="figure-link" href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/b903c080cc2c/nihms654899f7.jpg" itemprop="contentUrl" aria-describedby="figure-caption-6" role="button" data-image-width="429" data-image-height="996" data-image-alt="Figure 7" data-pmc-id="PMC4340844" data-figure-id="F7"> <img class="figure-thumb" itemprop="thumbnail" id="article-image-6" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/70d3/4340844/f6dcbe885185/nihms654899f7.gif" alt="Figure 7" /> </a> <meta itemprop="width" itemtype="http://schema.org/ImageObject" content="429"> <meta itemprop="height" itemtype="http://schema.org/ImageObject" content="996"> <div class="figure-caption figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="caption" aria-hidden="true"> <div class="caption-wrap"> <strong class="figure-label"> <p> Figure 7. Multiple Reaction Monitoring (MRM) to… </p> </strong> </div> </div> <div class="figure-caption-medium figure-caption-text" aria-hidden="true"> <strong class="figure-label"> <p> Figure 7. Multiple Reaction Monitoring (MRM) to identify glycopeptide biomarkers of autoimmunity </p> </strong> <div class="figure-caption-contents"><p> A) Without the… </p></div> </div> <figcaption id="figure-caption-6" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description"> <strong class="figure-label"> Figure 7. Multiple Reaction Monitoring (MRM) to identify glycopeptide biomarkers of autoimmunity </strong> <div class="figure-caption-contents">A) Without the need for additional purification, serum or plasma from peripheral blood is digested with trypsin to yield peptide fragments, including glycopeptides. A C18 Nano-LC Chip is then employed to separate the peptides and glycopeptides form one another utilizing Ultra High-Pressure Liquid Chromatography. The separated sample is then ionized using electron spray ionization and analyzed using a triple quadrupole time of flight mass spectrometry (QqQ-MS) using multiple reaction monitoring (MRM). B) MRM requires prior knowledge of the collision-induced dissociation (CID) behavior of the peptides and glycopeptides of interest. This knowledge allows for the appropriate MRM transitions to be developed for QqQ-MS detection. The process also requires a great deal of instrument optimization and knowledge of the peptide and glycopeptide retention times, but once established MRM can rapidly identify peptide and glycopeptides from serum samples with great sensitivity. Depicted here, a tryptic peptide common to the Fc region of all four IgG subclasses (red arrow) is used for absolute quantitation of total IgG. IgG subclass-specific peptides (light green, purple, black, and dark green arrows) are then used for comparison to the common Fc region peptide to determine the relative abundance of the individual IgG subclasses. C) Using a data set of theoretical IgA values, no significant difference between total IgA or IgA subclass-specific titers is seen between healthy controls and patients with two different autoimmune diseases (AutoD1, AutoD2). However, when the data is graphed as the relative abundance of the different IgA subclasses, it becomes clear that patients with AutoD1 have an increase in IgA1 that is highly significant.</div> </figcaption> </figure> </div> <div class="see-all actions-bar"> <button class="show-more" aria-controls="slides-container" data-ga-category="figure" data-ga-action="show_all"> All figures (7) </button> </div>  <div class="pswp figures-dialog" id="figures-dialog" tabindex="-1" role="dialog" aria-hidden="true" data-image-gallery-uid="article-figures" aria-label="Figures Dialog">  <div class="pswp__bg"></div>  <div class="pswp__scroll-wrap">  <div class="pswp__container"> <div class="pswp__item"></div> <div class="pswp__item"></div> <div class="pswp__item"></div> </div>  <div class="pswp__ui pswp__ui--hidden"> <div class="pswp__top-bar">  <div class="pswp__counter"></div> <button class="pswp__button pswp__button--close" title="Close (Esc)"></button> <button class="pswp__button pswp__button--share" title="Share"></button> <button class="pswp__button pswp__button--fs" title="Toggle fullscreen"></button> <button class="pswp__button pswp__button--zoom" title="Zoom in/out"></button>   <div class="pswp__preloader"> <div class="pswp__preloader__icn"> <div class="pswp__preloader__cut"> <div class="pswp__preloader__donut"></div> </div> </div> </div> </div> <div class="pswp__share-modal pswp__share-modal--hidden pswp__single-tap"> <div class="pswp__share-tooltip"></div> </div> <button class="pswp__button pswp__button--arrow--left" title="Previous (arrow left)"> </button> <button class="pswp__button pswp__button--arrow--right" title="Next (arrow right)"> </button> <div class="pswp__caption"> <div class="pswp__caption__center"></div> </div> </div> </div> </div> <div class="figure-dialog-caption"> <div class="figure-caption-link-container"> <a class="figure-caption-link" href="#">See this image and copyright information in PMC</a> </div> <div class="figure-caption-center" data-expand-title="click to show full caption" data-collapse-title="click to hide full caption"> </div> <button class="figure-caption-toggle-btn" aria-label="show full caption" data-expand-text="show full caption" data-collapse-text="hide full caption">  </button> </div> </div> <div class="similar-articles" id="similar"> <h2 class="title"> Similar articles </h2> <ul class="articles-list" id="similar-articles-list"> <li class="full-docsum"> <div class="docsum-content" style="display:inline;"> <a class="docsum-title" href="/27796794/" ref="article_id=27796794&linksrc=similar_articles_link&ordinalpos=1" data-ga-category="similar_article" data-ga-action="27796794" data-ga-label=""> Immunoglobulin Glycosylation Effects in Allergy and Immunity. </a> <div class="docsum-citation full-citation"> <span class="docsum-authors full-authors">Epp A, Sullivan KC, Herr AB, Strait RT.</span> <span class="docsum-authors short-authors">Epp A, et al.</span> <span class="docsum-journal-citation full-journal-citation">Curr Allergy Asthma Rep. 2016 Nov;16(11):79. doi: 10.1007/s11882-016-0658-x.</span> <span class="docsum-journal-citation short-journal-citation">Curr Allergy Asthma Rep. 2016.</span> <span class="citation-part">PMID: <span class="docsum-pmid">27796794</span></span> <span class="publication-type spaced-citation-item citation-part">Review.</span> </div> </div> </li> <li class="full-docsum"> <div class="docsum-content" style="display:inline;"> <a class="docsum-title" href="/34403816/" ref="article_id=34403816&linksrc=similar_articles_link&ordinalpos=2" data-ga-category="similar_article" data-ga-action="34403816" data-ga-label=""> Glycan biomarkers of autoimmunity and bile acid-associated alterations of the human glycome: Primary biliary cirrhosis and primary sclerosing cholangitis-specific glycans. </a> <div class="docsum-citation full-citation"> <span class="docsum-authors full-authors">Maverakis E, Merleev AA, Park D, Kailemia MJ, Xu G, Ruhaak LR, Kim K, Hong Q, Li Q, Leung P, Liakos W, Wan YY, Bowlus CL, Marusina AI, Lal NN, Xie Y, Luxardi G, Lebrilla CB.</span> <span class="docsum-authors short-authors">Maverakis E, et al.</span> <span class="docsum-journal-citation full-journal-citation">Clin Immunol. 2021 Sep;230:108825. doi: 10.1016/j.clim.2021.108825. 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ref="linksrc=add_to_search_link" role="button" data-search-term=""Autoimmune Diseases/metabolism"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Autoimmune Diseases/metabolism"[MeSH]" href="#"> Add to Search </a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link" aria-label="Toggle dropdown menu for keyword Autoimmune Diseases / pathology" tabindex="0" aria-controls="keyword-actions-mesh-terms-2" aria-expanded="false" data-pinger-ignore> Autoimmune Diseases / pathology </button><div id="keyword-actions-mesh-terms-2" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Autoimmune Diseases / pathology"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link" href="/?term=%22Autoimmune+Diseases%2Fpathology%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Autoimmune+Diseases%2Fpathology%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Autoimmune Diseases/pathology"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Autoimmune Diseases / pathology"> Search in PubMed </a></li><li><a class="search-in-mesh-link dropdown-block-link" ref="linksrc=search_in_mesh_link" href="https://www.ncbi.nlm.nih.gov/mesh?term=Autoimmune+Diseases" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Autoimmune Diseases / pathology"> Search in MeSH </a></li><li><a class="add-to-search-link dropdown-block-link" ref="linksrc=add_to_search_link" role="button" data-search-term=""Autoimmune Diseases/pathology"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Autoimmune Diseases/pathology"[MeSH]" href="#"> Add to Search </a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link major" aria-label="Toggle dropdown menu for keyword Autoimmunity / immunology*" tabindex="0" aria-controls="keyword-actions-mesh-terms-major-3" aria-expanded="false" data-pinger-ignore> Autoimmunity / immunology* </button><div id="keyword-actions-mesh-terms-major-3" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Autoimmunity / immunology*"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link" href="/?term=%22Autoimmunity%2Fimmunology%22%5BMAJR%5D&sort=date&sort_order=desc" data-href="/?term=%22Autoimmunity%2Fimmunology%22%5BMAJR%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Autoimmunity/immunology"[MAJR]" data-ga-category="search" data-ga-action="mesh_term" 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dropdown-block-link" ref="linksrc=add_to_search_link" role="button" data-search-term=""Binding Sites/immunology"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Binding Sites/immunology"[MeSH]" href="#"> Add to Search </a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link" aria-label="Toggle dropdown menu for keyword Glycosylation" tabindex="0" aria-controls="keyword-actions-mesh-terms-5" aria-expanded="false" data-pinger-ignore> Glycosylation </button><div id="keyword-actions-mesh-terms-5" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Glycosylation"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link" href="/?term=%22Glycosylation%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Glycosylation%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Glycosylation"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Glycosylation"> Search in PubMed </a></li><li><a class="search-in-mesh-link dropdown-block-link" ref="linksrc=search_in_mesh_link" href="https://www.ncbi.nlm.nih.gov/mesh?term=Glycosylation" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Glycosylation"> Search in MeSH </a></li><li><a class="add-to-search-link dropdown-block-link" ref="linksrc=add_to_search_link" role="button" data-search-term=""Glycosylation"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Glycosylation"[MeSH]" href="#"> Add to Search </a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link" aria-label="Toggle dropdown menu for keyword Humans" tabindex="0" 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data-href="/?term=%22Immune+System%2Fcytology%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Immune System/cytology"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Immune System / cytology"> Search in PubMed </a></li><li><a class="search-in-mesh-link dropdown-block-link" ref="linksrc=search_in_mesh_link" href="https://www.ncbi.nlm.nih.gov/mesh?term=Immune+System" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Immune System / cytology"> Search in MeSH </a></li><li><a class="add-to-search-link dropdown-block-link" ref="linksrc=add_to_search_link" role="button" data-search-term=""Immune System/cytology"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Immune System/cytology"[MeSH]" href="#"> Add to Search </a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link major" aria-label="Toggle dropdown menu for keyword Immune System / immunology*" tabindex="0" aria-controls="keyword-actions-mesh-terms-major-8" aria-expanded="false" data-pinger-ignore> Immune System / immunology* </button><div id="keyword-actions-mesh-terms-major-8" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Immune System / immunology*"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link" href="/?term=%22Immune+System%2Fimmunology%22%5BMAJR%5D&sort=date&sort_order=desc" data-href="/?term=%22Immune+System%2Fimmunology%22%5BMAJR%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Immune System/immunology"[MAJR]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Immune System / immunology*"> Search in PubMed </a></li><li><a class="search-in-mesh-link dropdown-block-link" ref="linksrc=search_in_mesh_link" href="https://www.ncbi.nlm.nih.gov/mesh?term=Immune+System" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Immune System / immunology*"> Search in MeSH </a></li><li><a class="add-to-search-link dropdown-block-link" ref="linksrc=add_to_search_link" role="button" data-search-term=""Immune System/immunology"[MAJR]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Immune System/immunology"[MAJR]" href="#"> Add to Search </a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link" aria-label="Toggle dropdown menu for keyword Immune System / metabolism" tabindex="0" aria-controls="keyword-actions-mesh-terms-9" aria-expanded="false" data-pinger-ignore> Immune System / metabolism </button><div id="keyword-actions-mesh-terms-9" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Immune System / metabolism"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link" href="/?term=%22Immune+System%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Immune+System%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Immune System/metabolism"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Immune System / metabolism"> Search in PubMed </a></li><li><a class="search-in-mesh-link dropdown-block-link" ref="linksrc=search_in_mesh_link" href="https://www.ncbi.nlm.nih.gov/mesh?term=Immune+System" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Immune System / metabolism"> Search in MeSH </a></li><li><a class="add-to-search-link dropdown-block-link" ref="linksrc=add_to_search_link" role="button" data-search-term=""Immune System/metabolism"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Immune System/metabolism"[MeSH]" href="#"> Add to Search </a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link major" aria-label="Toggle dropdown menu for keyword Immunoglobulins / immunology*" tabindex="0" aria-controls="keyword-actions-mesh-terms-major-10" aria-expanded="false" data-pinger-ignore> Immunoglobulins / immunology* </button><div id="keyword-actions-mesh-terms-major-10" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Immunoglobulins / immunology*"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link" href="/?term=%22Immunoglobulins%2Fimmunology%22%5BMAJR%5D&sort=date&sort_order=desc" data-href="/?term=%22Immunoglobulins%2Fimmunology%22%5BMAJR%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Immunoglobulins/immunology"[MAJR]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Immunoglobulins / immunology*"> Search in PubMed </a></li><li><a class="search-in-mesh-link dropdown-block-link" ref="linksrc=search_in_mesh_link" href="https://www.ncbi.nlm.nih.gov/mesh?term=Immunoglobulins" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Immunoglobulins / immunology*"> Search in MeSH </a></li><li><a class="add-to-search-link dropdown-block-link" ref="linksrc=add_to_search_link" role="button" data-search-term=""Immunoglobulins/immunology"[MAJR]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Immunoglobulins/immunology"[MAJR]" href="#"> Add to Search </a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link" aria-label="Toggle dropdown menu for keyword Immunoglobulins / metabolism" tabindex="0" aria-controls="keyword-actions-mesh-terms-11" aria-expanded="false" data-pinger-ignore> Immunoglobulins / metabolism </button><div id="keyword-actions-mesh-terms-11" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Immunoglobulins / metabolism"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link" href="/?term=%22Immunoglobulins%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Immunoglobulins%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Immunoglobulins/metabolism"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Immunoglobulins / metabolism"> Search in PubMed </a></li><li><a class="search-in-mesh-link dropdown-block-link" ref="linksrc=search_in_mesh_link" href="https://www.ncbi.nlm.nih.gov/mesh?term=Immunoglobulins" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Immunoglobulins / metabolism"> Search in MeSH </a></li><li><a class="add-to-search-link dropdown-block-link" ref="linksrc=add_to_search_link" role="button" data-search-term=""Immunoglobulins/metabolism"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Immunoglobulins/metabolism"[MeSH]" href="#"> Add to Search </a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link" aria-label="Toggle dropdown menu for keyword Models, Immunological" tabindex="0" aria-controls="keyword-actions-mesh-terms-12" aria-expanded="false" data-pinger-ignore> Models, Immunological </button><div id="keyword-actions-mesh-terms-12" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Models, Immunological"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link" href="/?term=%22Models%2C+Immunological%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Models%2C+Immunological%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Models, Immunological"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Models, Immunological"> Search in PubMed </a></li><li><a class="search-in-mesh-link dropdown-block-link" ref="linksrc=search_in_mesh_link" href="https://www.ncbi.nlm.nih.gov/mesh?term=Models%2C+Immunological" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Models, Immunological"> Search in MeSH </a></li><li><a class="add-to-search-link dropdown-block-link" ref="linksrc=add_to_search_link" role="button" data-search-term=""Models, Immunological"[MeSH]" data-ga-category="search" 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metabolism" tabindex="0" aria-controls="keyword-actions-mesh-terms-14" aria-expanded="false" data-pinger-ignore> Polysaccharides / metabolism </button><div id="keyword-actions-mesh-terms-14" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Polysaccharides / metabolism"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link" href="/?term=%22Polysaccharides%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Polysaccharides%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Polysaccharides/metabolism"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Polysaccharides / metabolism"> Search in PubMed </a></li><li><a class="search-in-mesh-link dropdown-block-link" ref="linksrc=search_in_mesh_link" href="https://www.ncbi.nlm.nih.gov/mesh?term=Polysaccharides" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Polysaccharides / metabolism"> Search in MeSH </a></li><li><a class="add-to-search-link dropdown-block-link" ref="linksrc=add_to_search_link" role="button" data-search-term=""Polysaccharides/metabolism"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Polysaccharides/metabolism"[MeSH]" href="#"> Add to Search </a></li></ul></div></div></div></li></ul> </div> <div id="substances" class="substances keywords-section"> <h2 class="title"> Substances </h2> <ul class="keywords-list"><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link" aria-label="Toggle dropdown menu for keyword Immunoglobulins" tabindex="0" aria-controls="keyword-actions-substances-0" aria-expanded="false" data-pinger-ignore> Immunoglobulins </button><div id="keyword-actions-substances-0" 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Glycans in the immune system and The Altered Glycan Theory of Autoimmunity: a critical review - PubMed