Marginal zone B cells: virtues of innate-like antibody-producing lymphocytes

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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="VznWqrAUVeeQUj29D1ySTkzMWcJvZT5SmHXmqAO334TubQLhhGpL0IiElGwGWOYa"> <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="VznWqrAUVeeQUj29D1ySTkzMWcJvZT5SmHXmqAO334TubQLhhGpL0IiElGwGWOYa"> <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="VznWqrAUVeeQUj29D1ySTkzMWcJvZT5SmHXmqAO334TubQLhhGpL0IiElGwGWOYa"> <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="VznWqrAUVeeQUj29D1ySTkzMWcJvZT5SmHXmqAO334TubQLhhGpL0IiElGwGWOYa"> <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="23348416"> <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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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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class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Cols+M&cauthor_id=23348416" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Montserrat Cols">Montserrat Cols</a><span class="comma">, </span></span><span class="authors-list-item "><a class="full-name" href="/?term=Puga+I&cauthor_id=23348416" ref="linksrc=author_name_link" data-ga-category="search" data-ga-action="author_link" data-ga-label="Irene Puga">Irene Puga</a></span> </div> </div> <div class="affiliations"> <h3 class="title"> Affiliation </h3> <ul class="item-list"> <li data-affiliation-id="short-view-affiliation-1" id="short-view-affiliation-1" ><sup class="key">1</sup> Catalan Institute for Research and Advanced Studies, Barcelona Biomedical Research Park, Barcelona, Spain. acerutti@imim.es</li> </ul> </div> </div> <ul class="identifiers" id="short-view-identifiers"> <li> <span class="identifier pubmed"> <span class="id-label"> 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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> Protective responses to microorganisms involve the nonspecific but rapid defence mechanisms of the innate immune system, followed by the specific but slow defence mechanisms of the adaptive immune system. Located as sentinels at the interface between the circulation and lymphoid tissue, splenic marginal zone B cells rapidly respond to blood-borne antigens by adopting 'crossover' defensive strategies that blur the conventional boundaries of innate and adaptive immunity. This Review discusses how marginal zone B cells function as innate-like lymphocytes that mount rapid antibody responses to both T cell-dependent and T cell-independent antigens. These responses require the integration of activation signals from germline-encoded and somatically recombined receptors for microorganisms with helper signals from effector cells of the innate and adaptive immune systems. </p> </div> </div> <p class="disclaimer" id="disclaimer"> <a href="/disclaimer/" target="_blank" ga_category="literature_resources" ga_action="disclaimer_link" ga_label="under_abstract">PubMed Disclaimer</a> </p> <div class="conflict-of-interest" id="conflict-of-interest"> <h2 class="title"> Conflict of interest statement </h2> <div class="statement"> <p xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:p1="http://pubmed.gov/pub-one"><bold>Competing interests statement</bold></p> <p xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:p1="http://pubmed.gov/pub-one">The authors declare no competing financial interests.</p> </div> </div> <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/06e1/3652659/7a24d4029c32/nihms456025f1.jpg" itemprop="contentUrl" aria-describedby="figure-caption-0" role="button" data-image-width="762" data-image-height="853" data-image-alt="Figure 1" data-pmc-id="PMC3652659" data-figure-id="F1"> <img class="figure-thumb" itemprop="thumbnail" id="article-image-0" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/06e1/3652659/6b625f2ccc6e/nihms456025f1.gif" alt="Figure 1" /> </a> <meta itemprop="width" itemtype="http://schema.org/ImageObject" content="762"> <meta itemprop="height" itemtype="http://schema.org/ImageObject" content="853"> <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. T cell-independent responses by mouse… </p> </strong> </div> </div> <div class="figure-caption-medium figure-caption-text" aria-hidden="true"> <strong class="figure-label"> <p> Figure 1. T cell-independent responses by mouse MZ B cells </p> </strong> <div class="figure-caption-contents"><p> T cell-independent (Tl) antigens are… </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. T cell-independent responses by mouse MZ B cells </strong> <div class="figure-caption-contents">T cell-independent (Tl) antigens are captured by metallophilic macrophages and marginal zone (MZ) macrophages after entering the splenic MZ via the marginal sinus. Alternatively, Tl antigens are captured by dendritic cells (DCs) and neutrophils in the circulation. Innate response activator (IRA) B cells may enhance the survival and activation of these antigen-capturing cells by releasing granulocyte–macrophage colony-stimulating factor (GM-CSF). Antigen-sampling cells stimulate MZB cells via the B cell receptor (BCR), Toll-like receptors (TLRs) and transmembrane activator and CAML interactor (TACI). TACI delivers signals that induce class-switch recombination and antibody production after ligating B cell-activating factor (BAFF) and a proliferation-inducing ligand (APRIL), which are released by antigen-capturing cells in response to microbial TLR ligands. Antigen-capturing cells, including red pulp DCs and macrophages, also secrete interleukin-6 (IL-6), IL-10, type I interferons (IFNs) and CXC-chemokine ligand 10 (CXCL10), which cooperate with BAFF and APRIL to promote the differentiation and survival of plasmablasts secreting IgM or class-switched IgG. Arrows indicate the path followed by antigens through the spleen. PALS, periarteriolar lymphoid sheath; PRR, pattern-recognition receptor.</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/06e1/3652659/494864ee5020/nihms456025f2.jpg" itemprop="contentUrl" aria-describedby="figure-caption-1" role="button" data-image-width="633" data-image-height="1048" data-image-alt="Figure 2" data-pmc-id="PMC3652659" data-figure-id="F2"> <img class="figure-thumb" itemprop="thumbnail" id="article-image-1" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/06e1/3652659/b5d73e43da6f/nihms456025f2.gif" alt="Figure 2" /> </a> <meta itemprop="width" itemtype="http://schema.org/ImageObject" content="633"> <meta itemprop="height" itemtype="http://schema.org/ImageObject" content="1048"> <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. T cell-independent responses by human… </p> </strong> </div> </div> <div class="figure-caption-medium figure-caption-text" aria-hidden="true"> <strong class="figure-label"> <p> Figure 2. T cell-independent responses by human MZ B cells </p> </strong> <div class="figure-caption-contents"><p> T cell-independent (Tl) antigens are… </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. T cell-independent responses by human MZ B cells </strong> <div class="figure-caption-contents">T cell-independent (Tl) antigens are thought to enter the marginal zone (MZ) through the perifollicular zone. Once in the MZ, they may be captured by neutrophil extracellular trap (NET)-like structures emanating from B cell-helper neutrophils (N<sub>BH</sub> cells). These cells may differentiate from circulating neutrophils as a result of the production of interleukin-10 (IL-10) by perifollicular sinus-lining cells and macrophages in response to microbial Toll-like receptor (TLR) ligands. Antigen capture may also involve reticular cells, macrophages, sinus-lining cells and dendritic cells (DCs). In addition to making Tl antigens available to the B cell receptor (BCR) and TLRs on MZ B cells, antigen-capturing cells release B cell-activating factor (BAFF) and a proliferation-inducing ligand (APRIL), which engage transmembrane activator and CAML interactor (TACI) on MZ B cells. N<sub>BH</sub> cells also release IL-21, thereby inducing class-switch recombination, somatic hypermutation and antibody production in MZ B cells. The generation of plasmablasts secreting IgM or class-switched IgG and IgA involves the production of IL-6, IL-10, IL-21 and CXC-chemokine ligand 10 (CXCL10) by antigen-capturing cells. Arrows indicate the putative path followed by antigens through the spleen. IFN, interferon; PALS, periarteriolar lymphoid sheath; PRR, pattern-recognition receptor.</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/06e1/3652659/33d87689dd94/nihms456025f3.jpg" itemprop="contentUrl" aria-describedby="figure-caption-2" role="button" data-image-width="795" data-image-height="775" data-image-alt="Figure 3" data-pmc-id="PMC3652659" data-figure-id="F3"> <img class="figure-thumb" itemprop="thumbnail" id="article-image-2" src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/06e1/3652659/83a896f51a23/nihms456025f3.gif" alt="Figure 3" /> </a> <meta itemprop="width" itemtype="http://schema.org/ImageObject" content="795"> <meta itemprop="height" itemtype="http://schema.org/ImageObject" content="775"> <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. T cell-dependent responses by mouse… </p> </strong> </div> </div> <div class="figure-caption-medium figure-caption-text" aria-hidden="true"> <strong class="figure-label"> <p> Figure 3. T cell-dependent responses by mouse MZ B cells </p> </strong> <div class="figure-caption-contents"><p> In the extrafollicular T cell-dependent… </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. T cell-dependent responses by mouse MZ B cells </strong> <div class="figure-caption-contents">In the extrafollicular T cell-dependent (TD) pathway, marginal zone (MZ) dendritic cells (DCs) expressing DC inhibitory receptor 2 (DCIR2) capture protein antigens and present them to naive CD4<sup>+</sup> T cells, which subsequently differentiate into T helper 2 (TH2) cells expressing CD40 ligand (CD40L) and interleukin-4 (IL-4). DCIR2<sup>+</sup> DCs also expose antigens to MZ B cells, and signals from the B cell receptor (BCR) convert MZB cells into antigen-presenting cells. By establishing cognate interactions with T<sub>H</sub>2 cells, activated MZ B cells rapidly produce low-affinity IgGl but not IgM. ln an alternative pathway, lipid-reactive MZ B cells activate invariant natural killer T (iNKT) cells by presenting CDld-loaded glycolipids. The expression of CD40L, IL-4 and interferon-γ(IFNγ) by activated iNKT cells induces MZ B cells to differentiate into plasma blasts that secrete low-affinity IgM or IgG. In the follicular TD pathway, MZB cells downregulate receptors for sphin-gosine-1-phosphate (SIP) after capturing complement-decorated protein antigens via CD21 and CD3B. The interruption of MZ-retaining signals from SIP receptors stimulates MZ B cells to enter the follicle in response to CXC-chemokine receptor 5 (CXCRB) signals. In the follicle, MZ B cells deposit antigens on follicular dendritic cells (FDCs) and thereafter upregulate SI P receptor expression to exit the follicle and return to the MZ. Protein antigens displayed by FDCs enhance TD antibody responses by promoting the selection of follicular germinal centre (GC) B cells expressing a high-affinity BCR. This selection also involves cognate interactions between GC B cells and T follicular helper (T<sub>FH</sub>) cells expressing CD40L and IL-21. Ultimately, selected GC B cells differentiate into long-lived memory B cells or plasma cells that produce high-affinity IgG. Alternatively, MZ B cells capture protein antigens with or without the help of metallophilic macrophages and migrate to the border between the periarteriolar lymphoid sheath (PALS) and the follicle to initiate a GC reaction after establishing a cognate interaction with T<sub>FH</sub> cells expressing CD40L, IL-4 or interferon-γ(IFNγ). These T<sub>FH</sub> cells probably emerge from the priming of naive CD4<sup>+</sup>T cells by either MZ B cells or DCs expressing DEC205. A subset of these T<sub>FH</sub> cells can also promote the generation of low-affinity extrafollicular IgM and IgG plasmablasts. CXCL13, CXC-chemokine ligand 13; TCR. T cell receptor; TLR, Toll-like receptor.</div> </figcaption> </figure> </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="/27621420/" ref="article_id=27621420&linksrc=similar_articles_link&ordinalpos=1" data-ga-category="similar_article" data-ga-action="27621420" data-ga-label=""> The soluble pattern recognition receptor PTX3 links humoral innate and adaptive immune responses by helping marginal zone B cells. </a> <div class="docsum-citation full-citation"> <span class="docsum-authors full-authors">Chorny A, Casas-Recasens S, Sintes J, Shan M, Polentarutti N, García-Escudero R, Walland AC, Yeiser JR, Cassis L, Carrillo J, Puga I, Cunha C, Bastos H, Rodrigues F, Lacerda JF, Morais A, Dieguez-Gonzalez R, Heeger PS, Salvatori G, Carvalho A, Garcia-Sastre A, Blander JM, Mantovani A, Garlanda C, Cerutti A.</span> <span class="docsum-authors short-authors">Chorny A, et al.</span> <span class="docsum-journal-citation full-journal-citation">J Exp Med. 2016 Sep 19;213(10):2167-85. doi: 10.1084/jem.20150282. 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role="button" data-search-term=""Animals"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Animals"[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 Antibody Formation*" tabindex="0" aria-controls="keyword-actions-mesh-terms-major-1" aria-expanded="false" data-pinger-ignore> Antibody Formation* </button><div id="keyword-actions-mesh-terms-major-1" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Antibody Formation*"><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=%22Antibody+Formation%22%5BMAJR%5D&sort=date&sort_order=desc" data-href="/?term=%22Antibody+Formation%22%5BMAJR%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Antibody Formation"[MAJR]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Antibody Formation*"> 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=Antibody+Formation" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Antibody Formation*"> 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=""Antibody Formation"[MAJR]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Antibody Formation"[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 Antigens / immunology" tabindex="0" aria-controls="keyword-actions-mesh-terms-2" aria-expanded="false" data-pinger-ignore> Antigens / immunology </button><div id="keyword-actions-mesh-terms-2" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Antigens / 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=%22Antigens%2Fimmunology%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Antigens%2Fimmunology%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Antigens/immunology"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Antigens / 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=Antigens" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Antigens / 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=""Antigens/immunology"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Antigens/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 Antigens / metabolism" tabindex="0" aria-controls="keyword-actions-mesh-terms-3" aria-expanded="false" data-pinger-ignore> Antigens / metabolism </button><div id="keyword-actions-mesh-terms-3" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Antigens / 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=%22Antigens%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Antigens%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Antigens/metabolism"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Antigens / 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=Antigens" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Antigens / 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=""Antigens/metabolism"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Antigens/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 B-Lymphocytes / immunology*" tabindex="0" aria-controls="keyword-actions-mesh-terms-major-4" aria-expanded="false" data-pinger-ignore> B-Lymphocytes / immunology* </button><div id="keyword-actions-mesh-terms-major-4" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword B-Lymphocytes / 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=%22B-Lymphocytes%2Fimmunology%22%5BMAJR%5D&sort=date&sort_order=desc" data-href="/?term=%22B-Lymphocytes%2Fimmunology%22%5BMAJR%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="B-Lymphocytes/immunology"[MAJR]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="B-Lymphocytes / 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=B-Lymphocytes" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="B-Lymphocytes / 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=""B-Lymphocytes/immunology"[MAJR]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""B-Lymphocytes/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 B-Lymphocytes / metabolism" tabindex="0" aria-controls="keyword-actions-mesh-terms-5" aria-expanded="false" data-pinger-ignore> B-Lymphocytes / metabolism </button><div id="keyword-actions-mesh-terms-5" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword B-Lymphocytes / 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=%22B-Lymphocytes%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22B-Lymphocytes%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="B-Lymphocytes/metabolism"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="B-Lymphocytes / 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=B-Lymphocytes" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="B-Lymphocytes / 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=""B-Lymphocytes/metabolism"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""B-Lymphocytes/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 Humans" tabindex="0" aria-controls="keyword-actions-mesh-terms-6" aria-expanded="false" data-pinger-ignore> Humans </button><div id="keyword-actions-mesh-terms-6" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Humans"><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=%22Humans%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Humans%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Humans"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Humans"> 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=Humans" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Humans"> 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=""Humans"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Humans"[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 Immunity, Innate*" tabindex="0" aria-controls="keyword-actions-mesh-terms-major-7" aria-expanded="false" data-pinger-ignore> Immunity, Innate* </button><div id="keyword-actions-mesh-terms-major-7" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Immunity, Innate*"><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=%22Immunity%2C+Innate%22%5BMAJR%5D&sort=date&sort_order=desc" data-href="/?term=%22Immunity%2C+Innate%22%5BMAJR%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Immunity, Innate"[MAJR]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Immunity, Innate*"> 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=Immunity%2C+Innate" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Immunity, Innate*"> 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=""Immunity, Innate"[MAJR]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Immunity, Innate"[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 Immunoglobulin Class Switching" tabindex="0" aria-controls="keyword-actions-mesh-terms-8" aria-expanded="false" data-pinger-ignore> Immunoglobulin Class Switching </button><div id="keyword-actions-mesh-terms-8" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Immunoglobulin Class Switching"><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=%22Immunoglobulin+Class+Switching%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Immunoglobulin+Class+Switching%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Immunoglobulin Class Switching"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Immunoglobulin Class Switching"> 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=Immunoglobulin+Class+Switching" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Immunoglobulin Class Switching"> 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=""Immunoglobulin Class Switching"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Immunoglobulin Class Switching"[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 Lymphoid Tissue / cytology" tabindex="0" aria-controls="keyword-actions-mesh-terms-9" aria-expanded="false" data-pinger-ignore> Lymphoid Tissue / cytology </button><div id="keyword-actions-mesh-terms-9" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Lymphoid Tissue / cytology"><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=%22Lymphoid+Tissue%2Fcytology%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Lymphoid+Tissue%2Fcytology%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Lymphoid Tissue/cytology"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Lymphoid Tissue / 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=Lymphoid+Tissue" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Lymphoid Tissue / 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=""Lymphoid Tissue/cytology"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Lymphoid Tissue/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 Lymphoid Tissue / immunology*" tabindex="0" aria-controls="keyword-actions-mesh-terms-major-10" aria-expanded="false" data-pinger-ignore> Lymphoid Tissue / immunology* </button><div id="keyword-actions-mesh-terms-major-10" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Lymphoid Tissue / 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=%22Lymphoid+Tissue%2Fimmunology%22%5BMAJR%5D&sort=date&sort_order=desc" data-href="/?term=%22Lymphoid+Tissue%2Fimmunology%22%5BMAJR%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Lymphoid Tissue/immunology"[MAJR]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Lymphoid Tissue / 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=Lymphoid+Tissue" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Lymphoid Tissue / 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=""Lymphoid Tissue/immunology"[MAJR]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Lymphoid Tissue/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 Mutation" tabindex="0" aria-controls="keyword-actions-mesh-terms-11" aria-expanded="false" data-pinger-ignore> Mutation </button><div id="keyword-actions-mesh-terms-11" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Mutation"><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=%22Mutation%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Mutation%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Mutation"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Mutation"> 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=Mutation" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Mutation"> 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=""Mutation"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Mutation"[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 Protein Binding / immunology" tabindex="0" aria-controls="keyword-actions-mesh-terms-12" aria-expanded="false" data-pinger-ignore> Protein Binding / immunology </button><div id="keyword-actions-mesh-terms-12" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Protein Binding / 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=%22Protein+Binding%2Fimmunology%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Protein+Binding%2Fimmunology%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Protein Binding/immunology"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Protein Binding / 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=Protein+Binding" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Protein Binding / 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=""Protein Binding/immunology"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Protein Binding/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 Signal Transduction" tabindex="0" aria-controls="keyword-actions-mesh-terms-13" aria-expanded="false" data-pinger-ignore> Signal Transduction </button><div id="keyword-actions-mesh-terms-13" class="keyword-actions-dropdown dropdown dropdown-container" hidden aria-label="Dropdown menu for keyword Signal Transduction"><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=%22Signal+Transduction%22%5BMeSH%5D&sort=date&sort_order=desc" data-href="/?term=%22Signal+Transduction%22%5BMeSH%5D&sort=date&sort_order=desc" ref="linksrc=search_in_pubmed_keyword_link&mesh="Signal Transduction"[MeSH]" data-ga-category="search" data-ga-action="mesh_term" data-ga-label="Signal Transduction"> 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=Signal+Transduction" data-ga-category="search_mesh" data-ga-action="mesh_term_link" data-ga-label="Signal Transduction"> 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=""Signal Transduction"[MeSH]" data-ga-category="search" data-ga-action="add_to_search" data-ga-label=""Signal Transduction"[MeSH]" href="#"> Add to Search </a></li></ul></div></div></div></li></ul> </div> <div id="substances" class="substances 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Marginal zone B cells: virtues of innate-like antibody-producing lymphocytes - PubMed