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<input type="email" aria-label="Sender Email Address" placeholder="email@example.com" id="email-from" class="email-from" pattern="^[a-zA-Z0-9_.+-]+@[a-zA-Z0-9-]+\.[a-zA-Z0-9-.]+$" maxlength="256">
</div>
<div class="action-panel-control-wrap">
<label for="email-citation-format" class="action-panel-label">
Format:
</label>
<select id="email-citation-format" name="citation-format" class="action-panel-selector email-citation-format">
<option selected="selected" value="summary">Summary</option>
<option value="summary-text">Summary (text)</option>
<option value="abstract">Abstract</option>
<option value="abstract-text">Abstract (text)</option>
</select>
</div>
<div class="include-supplemental-container">
<input type="checkbox" aria-label="Include MeSH and other data" name="include-supplemental" id="email-include-supplemental" class="email-include-supplemental">
<label for="email-include-supplemental" class="email-include-supplemental-label">MeSH and other data</label>
</div>
<div class="form-field recaptcha ">
<div class="g-recaptcha" id="id-recaptcha" data-sitekey="6LfsWHMdAAAAAClKbtOpjQ2pMjgsGxvv7NdZW9uI"></div>
</div>
<div id="captcha-error-message" class="usa-input-error-message captcha-validation-message" role="alert"></div>
<div class="action-panel-actions">
<button class="action-panel-submit"
type="submit"
data-loading-label="Sending..."
data-ga-category="save_share"
data-ga-action="email"
data-ga-label="send">
Send email
</button>
<button class="action-panel-cancel"
aria-label="Close 'Email citations' panel"
ref="linksrc=close_email_panel"
aria-controls="email-action-panel"
aria-expanded="false"
data-ga-category="save_share"
data-ga-action="email"
data-ga-label="cancel">
Cancel
</button>
</div>
<input type="hidden" name="email-search-details" value="" />
<input type="hidden" name="email-search-details-hash" value="0e42663a6c3bd85498fcb88798998fed7bfdc45d457db35281e41afe13cc0524" />
</form>
</div>
</div>
<div id="collections-action-panel"
class="collections-action-panel action-panel in-progress-dots-panel"
aria-hidden="true"
data-collections-open-panel-enabled="false"
data-collections-open-panel-url-hash="#open-collections-panel">
<div class="inner-wrap">
<h3 class="action-panel-heading">
Add to Collections
</h3>
<form id="collections-action-panel-form"
class="collections-action-panel-form action-panel-content action-form action-panel-smaller-selectors"
data-existing-collections-url="/list-existing-collections/"
data-add-to-existing-collection-url="/add-to-existing-collection/"
data-create-and-add-to-new-collection-url="/create-and-add-to-new-collection/"
data-get-article-ids-by-search-url="/get-article-ids-by-search/"
data-myncbi-max-collection-name-length="100"
data-add-to-collection-max-amount="1000"
data-collections-root-url="https://www.ncbi.nlm.nih.gov/myncbi/collections/">
<input type="hidden" name="csrfmiddlewaretoken" value="Ldy4s1yJl4SC3LPHmAOjZ2IPJTbKRjHJ20i95R15VFF6XcVxZTrltaethXTWKUf3">
<div class="choice-group" role="radiogroup">
<ul class="radio-group-items">
<li>
<input type="radio"
id="collections-action-panel-new"
class="collections-new"
name="collections"
value="new"
data-ga-category="save_share"
data-ga-action="collections"
data-ga-label="collections_radio_new">
<label for="collections-action-panel-new">Create a 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="[^&quot;&amp;=&lt;&gt;\/]*" title="The following characters are not allowed in the Name field: &quot;&amp;=&lt;&gt;/"
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="Ldy4s1yJl4SC3LPHmAOjZ2IPJTbKRjHJ20i95R15VFF6XcVxZTrltaethXTWKUf3">
<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="Ldy4s1yJl4SC3LPHmAOjZ2IPJTbKRjHJ20i95R15VFF6XcVxZTrltaethXTWKUf3">
<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="[^&quot;&amp;=&lt;&gt;\/]*" title="The following characters are not allowed in the Name field: &quot;&amp;=&lt;&gt;/">
</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="Ldy4s1yJl4SC3LPHmAOjZ2IPJTbKRjHJ20i95R15VFF6XcVxZTrltaethXTWKUf3">
<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="Ldy4s1yJl4SC3LPHmAOjZ2IPJTbKRjHJ20i95R15VFF6XcVxZTrltaethXTWKUf3">
<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="[^&quot;&amp;=&lt;&gt;\/]*" title="The following characters are not allowed in the Name field: &quot;&amp;=&lt;&gt;/">
</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="38809974">
<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"
href="https://www.science.org/doi/abs/10.1126/sciadv.adn2789?url_ver=Z39.88-2003&amp;rfr_id=ori:rid:crossref.org&amp;rfr_dat=cr_pub 0pubmed"
target="_blank"
rel="noopener"
ref="linksrc=fulltextorjournal_fulltext&amp;is_pmc=False&amp;PrId=4487&amp;itool=Abstract-def&amp;log$=linkouticon&amp;uid=38809974&amp;db=pubmed&amp;nlmid=101653440"
title="See full text options at Atypon"
data-ga-category="full_text"
data-ga-action="Atypon"
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><img src="https://cdn.ncbi.nlm.nih.gov/corehtml/query/egifs/https:--www.science.org-pb-assets-images-pubmedoa-egp_authorchoice.jpg" alt="Atypon full text link"><span class="text">
Atypon
</span></a><a class="link-item
pmc
"
href="https://pmc.ncbi.nlm.nih.gov/articles/pmid/38809974/"
target="_blank"
rel="noopener"
ref="linksrc=fulltextorjournal_fulltext&amp;is_pmc=True&amp;PrId=3494&amp;itool=Abstract-def&amp;log$=linkouticon&amp;uid=38809974&amp;db=pubmed&amp;nlmid=101653440"
title="Free full text at PubMed Central"
data-ga-category="full_text"
data-ga-action="PMC"
data-ga-label="38809974"
><span class="text">
Free PMC article
</span></a>
</div>
</div>
<div class="short-view">
<a href="#" class="full-text-links-button full-text-links-dialog-trigger">
Full text links
</a>
</div>
</div>
<div class="actions-buttons sidebar"><h3 class="title">Actions</h3><div class="inner-wrap"><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="/38809974/citations/"
data-citation-style="nlm"
data-pubmed-format-link="/38809974/export/"><span class="button-label">Cite</span></button><link type="text/css" href="ncbi-overlay-block/src/overlay-block.css"><div class="collections-button-container" data-article-id="38809974" data-article-db="pubmed"><button class="collections-button collections-dialog-trigger"
aria-label="Save article in MyNCBI collections."
data-ga-category="collections_button"
data-ga-action="click"
data-ga-label="collections_button"
data-collections-open-dialog-enabled="false"
data-collections-open-dialog-url="https://account.ncbi.nlm.nih.gov/?back_url=https%3A%2F%2Fpubmed.ncbi.nlm.nih.gov%2F38809974%2F%23open-collections-dialog"
data-in-collections="false"><span class="button-label">Collections</span></button><div class="overlay" role="dialog"><div id="collections-action-dialog"
class="dialog collections-dialog"
aria-hidden="true"><div class="title">Add to Collections</div><div class="collections-action-panel action-panel"><form id="collections-action-dialog-form"
class="collections-action-panel-form action-panel-content action-form action-panel-smaller-selectors"
data-existing-collections-url="/list-existing-collections/"
data-add-to-existing-collection-url="/add-to-existing-collection/"
data-create-and-add-to-new-collection-url="/create-and-add-to-new-collection/"
data-myncbi-max-collection-name-length="100"
data-collections-root-url="https://www.ncbi.nlm.nih.gov/myncbi/collections/"><input type="hidden" name="csrfmiddlewaretoken" value="Ldy4s1yJl4SC3LPHmAOjZ2IPJTbKRjHJ20i95R15VFF6XcVxZTrltaethXTWKUf3"><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="[^&quot;&amp;=&lt;&gt;\/]*" title="The following characters are not allowed in the Name field: &quot;&amp;=&lt;&gt;/"
maxlength="100"
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<h1 class="heading-title">
A molecular switch controls assembly of bacterial focal adhesions
</h1>
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<h1 class="heading-title">
A molecular switch controls assembly of bacterial focal adhesions
</h1>
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><sup class="key">1</sup> Laboratoire d&#x27;Ingénierie des Systèmes Macromoléculaires (LISM), Institut de Microbiologie de la Méditerranée (IMM), CNRS - Aix-Marseille Université UMR7255, 31 Chemin Joseph Aiguier CS70071, 13402 Marseille Cedex 20, France.</li>
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><sup class="key">2</sup> Laboratoire de Chimie Bactérienne (LCB), Institut de Microbiologie de la Méditerranée (IMM), Turing Center for Living Systems, CNRS - Aix-Marseille Université UMR7283, 31 Chemin Joseph Aiguier CS70071, 13402 Marseille Cedex 20, France.</li>
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><sup class="key">3</sup> Institut de Microbiologie de la Méditerranée (IMM), CNRS - Aix-Marseille Université UMR7283, 31 Chemin Joseph Aiguier CS70071, 13402 Marseille Cedex 20, France.</li>
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><sup class="key">4</sup> National Center for Biotechnology Information, National Library of Medicine, National Institutes of Health, Bethesda, MD 20894, USA.</li>
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><sup class="key">5</sup> Department of Microbiology and Molecular Medicine, Faculty of Medicine/Centre Médical Universitaire, University of Geneva, 1211 Genève 4, Switzerland.</li>
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Cell motility universally relies on spatial regulation of focal adhesion complexes (FAs) connecting the substrate to cellular motors. In bacterial FAs, the Adventurous gliding motility machinery (Agl-Glt) assembles at the leading cell pole following a Mutual gliding-motility protein (MglA)-guanosine 5'-triphosphate (GTP) gradient along the cell axis. Here, we show that GltJ, a machinery membrane protein, contains cytosolic motifs binding MglA-GTP and AglZ and recruiting the MreB cytoskeleton to initiate movement toward the lagging cell pole. In addition, MglA-GTP binding triggers a conformational shift in an adjacent GltJ zinc-finger domain, facilitating MglB recruitment near the lagging pole. This prompts GTP hydrolysis by MglA, leading to complex disassembly. The GltJ switch thus serves as a sensor for the MglA-GTP gradient, controlling FA activity spatially.
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<p> Fig. 1.. Dynamics of the <i> M. xanthus… </i> </p>
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<p> Fig. 1.. Dynamics of the <i> M. xanthus </i> motility complex. </p>
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<div class="figure-caption-contents"><p> ( <b> A </b> ) Proposed architecture of… </p></div>
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Fig. 1.. Dynamics of the <i>M. xanthus</i> motility complex.
</strong>
<div class="figure-caption-contents">(<b>A</b>) Proposed architecture of the motility complex and mechanism of propulsion. (ai) Proposed structure of the trafficking motor complex in nonadhered mobile clusters. The motility complex is assembled at the leading cell pole by recruitment of the IM motors (GltJ, GltG, and AglRQS) to the cytosolic platform formed by the AglZ, MglA, and MreB proteins. This complex uses the proton-motive force to move directionally to the lagging cell pole in a counterclockwise rotational trajectory, until it becomes immobilized at bFA. (aii) Interaction between the IM motor and a complex of OM proteins containing the CglB adhesin is proposed to connect the trafficking complex to the underlying substratum and form bFAs. The motility complex becomes fixed as OM adhesive complex treadmill through the IM motor via proton-motive force (pmf)-driven interaction cycles. The cell moves, rotating along its long axis in the clockwise direction. (<b>B</b>) <i>Myxococcus</i> bFAs containing AglZ-NG imaged by epifluorescence. Fixed clusters are formed at the leading cell pole and either disperse when they reach the lagging cell pole (black arrow) or disperse along the cell body (orange arrows). White arrow indicates the direction of movement. Pictures were taken every 30 s. Scale bar, 2 μm.</div>
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<p> Fig. 2.. The GltJ protein is essential… </p>
</strong>
</div>
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<strong class="figure-label">
<p> Fig. 2.. The GltJ protein is essential for bFA formation. </p>
</strong>
<div class="figure-caption-contents"><p> ( <b> A </b> ) Domain organization… </p></div>
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<strong class="figure-label">
Fig. 2.. The GltJ protein is essential for bFA formation.
</strong>
<div class="figure-caption-contents">(<b>A</b>) Domain organization of GltJ and predicted topology in the cell envelope. (<b>B</b>) Single-cell speeds of <i>gltJ</i> and <i>aglZ</i> mutant strains. Each data point corresponds to the mean of the speeds obtained for single cells across four technical replicates, an experiment repeated three times for each strain. The statistics were obtained applying a Welchs <i>t</i> test. Significance is assumed for <i>P</i> &lt; 0.1. NS, not significant. (<b>C</b>) Mean square displacement (MSD) of single cells of <i>gltJ</i> and <i>aglZ</i> mutant strains. The MSD was calculated as a proxy for directed single cell movements across a time period of 30 min (see Materials and Methods). Each data point corresponds to the mean of the MSD obtained for single cells across four technical replicates, an experiment repeated four times for each strain. The statistics were obtained as in (B). (<b>D</b>) Localization of NG-GltJ in a motile cell. Shown is a time-lapse (one image every 4 s) and derived kymograph of the same cell (one image every 1 s). The black arrows point to fixed clusters. Scale bar, 2 μm. (<b>E</b>) GltJ polar localization and localization to bFAs. Example images of NG fusions of GltJ and mutants are shown for each fusion. Scale bar, 4 μm. (<b>F</b>) Formation of GltJ clusters in cells expressing GltJ variants. Shown is the total number of GltJ-NG clusters (polar and intracellular) per cell scored in <i>n</i> replicates across four independent experiments. The statistics were obtained applying a Wilcoxon test. Significance is assumed for <i>P</i> &lt; 0.1. (<b>G</b>) Polar determinants in the GltJ Nt1-222 region. NG fusions to various motifs of the GltJ Nt1-222 region were expressed in a <i>gltJ</i> mutant background and tested for localization. Example images are shown for each construct. Scale bars, 3 μm.</div>
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<p> Fig. 3.. GYF <sup> GltJ </sup> interacts with the… </p>
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<p> Fig. 3.. GYF <sup> GltJ </sup> interacts with the PRS of AglZ through a noncanonical binding interface. </p>
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<div class="figure-caption-contents"><p> (… </p></div>
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<strong class="figure-label">
Fig. 3.. GYF<sup>GltJ</sup> interacts with the PRS of AglZ through a noncanonical binding interface.
</strong>
<div class="figure-caption-contents">(<b>A</b>) Domain organization of AglZ. (<b>B</b>) ITC binding assay with GYF<sup>GltJ</sup> and PRS<sup>AglZ</sup>. DP, differential power. (<b>C</b>) Representative solution structure of the GYF<sup>GltJ</sup> domain (NMR ensembles of the 20 conformers with the lowest energy are shown in fig. S6B). Secondary structures are labeled. The protein residues involved in the PRS<sup>AglZ</sup> binding are displayed in blue. (<b>D</b>) <sup>1</sup>H,<sup>15</sup>N heteronuclear single quantum coherence (HSQC) spectra, representing all H-N correlations of 0.1 mM <sup>15</sup>N-labeled GYF<sup>GltJ</sup>-free (red) and upon titration of increasing concentrations of PRS<sup>AglZ</sup> are shown (dark to light blue). For comparison, selected resonances of GYF<sup>GltJ</sup> showing the strongest CSPs upon PRS<sup>AglZ</sup> addition are reported. ppm, parts per million. (<b>E</b>) bFA activity in GltJ and AglZ mutants. For each strain, the number of active bFAs was determined by tracking individual bFAs in single cells in <i>n</i> replicates across four independent experiments. The statistics were obtained applying a Wilcoxon test. Significance is assumed for <i>P</i> &lt; 0.1. (<b>F</b>) The GltJ N-terminal region is required for bFA formation. Scale bar, 4 μm. (<b>G</b>) The AglZ PRS motif is required for bFA formation but not for polar localization. Scale bar, 4 μm.</div>
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<p> Fig. 4.. MglA and ZnR <sup> GltJ </sup> interact… </p>
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<p> Fig. 4.. MglA and ZnR <sup> GltJ </sup> interact with Linker <sup> GltJ </sup> . </p>
</strong>
<div class="figure-caption-contents"><p> ( <b> A </b> ) Overlay of the <sup> 1… </sup> </p></div>
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<strong class="figure-label">
Fig. 4.. MglA and ZnR<sup>GltJ</sup> interact with Linker<sup>GltJ</sup>.
</strong>
<div class="figure-caption-contents">(<b>A</b>) Overlay of the <sup>1</sup>H,<sup>15</sup>N HSQCs of <sup>15</sup>N-labeled Linker-GYF<sup>GltJ</sup> free (blue) and bound to MglA-GTP (magenta). Left: Inset shows expanded central part of Linker-GYF<sup>GltJ</sup> spectra. Peaks disappearing upon MglA-GTP interaction are labeled. (<b>B</b>) Relative peak intensities (<i>I</i>/<i>I</i><sub>0</sub>) of NMR signals of Linker-GYF<sup>GltJ</sup> in complex with MglA-GTP. Resonances experiencing strong peak intensity decrease are indicated by magenta bars. Bars with magenta dashed edges indicate disappearing resonances upon interaction. The absence of bars indicates residues that could not be assigned, and P letters specify proline position. Dashed line represents 1σ from the average <i>I</i>/<i>I</i><sub>0</sub>. The main perturbed region is framed in gray, and the corresponding sequence is shown below the plot. (<b>C</b>) Representative solution structure of the ZnR<sup>GltJ</sup> domain. Secondary structures are labeled as well as the cysteine side chains chelating the zinc ion (orange sphere). The protein residues involved in Linker<sup>GltJ</sup> binding are displayed in blue. (<b>D</b>) <sup>1</sup>H,<sup>15</sup>N-HSQC series of <sup>15</sup>N-labeled ZnR<sup>GltJ</sup>-free (salmon) and upon addition of an increasing amount of Linker-GYF<sup>GltJ</sup> (blue shades). Insets show selected resonances experiencing strong chemical shift variations in the presence of the Linker-GYF<sup>GltJ</sup> with, in addition, the corresponding resonance in the <sup>1</sup>H,<sup>15</sup>N HSQC of the Nt1-222 construct (green peaks). Residues experiencing CSPs are reported on ZnR<sup>GltJ</sup> structure [shown in (C)]. (<b>E</b>) Combined CSPs between <sup>1</sup>H,<sup>15</sup>N resonances of the free and ZnR<sup>GltJ</sup>-bound state of Linker-GYF<sup>GltJ</sup> (at 1.7 molar ratio). The absence of bars indicates residues that could not be assigned, and P letters specify the position of a proline. Dashed line represents 1σ from the average CSPs. The main region perturbed upon MglA-GTP interaction with Linker-GYF<sup>GltJ</sup> is framed in gray as in (B). Residues within this region experiencing CSPs upon ZnR<sub>GltJ</sub> binding are colored salmon in the sequence.</div>
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<p> Fig. 5.. MglA-GTP competes with ZnR <sup> GltJ… </sup> </p>
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<strong class="figure-label">
<p> Fig. 5.. MglA-GTP competes with ZnR <sup> GltJ </sup> for Linker <sup> GltJ </sup> binding. </p>
</strong>
<div class="figure-caption-contents"><p> ( <b> A </b> ) Spectrum region showing… </p></div>
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<strong class="figure-label">
Fig. 5.. MglA-GTP competes with ZnR<sup>GltJ</sup> for Linker<sup>GltJ</sup> binding.
</strong>
<div class="figure-caption-contents">(<b>A</b>) Spectrum region showing the overlay of <sup>1</sup>H,<sup>15</sup>N HSQC spectrum of <sup>15</sup>N-labeled Nt1-222<sup>GS8998</sup> (mint), a modified Nt1-222 in which the D89-V98 region has been substituted by a poly-GS sequence, with the spectrum of free ZnR<sup>GltJ</sup> (salmon). (<b>B</b>) Same spectrum region as in (A) showing the overlay of <sup>1</sup>H,<sup>15</sup>N HSQC spectra of Nt1-222<sup>GS8998</sup> in absence (mint) and in presence of MglA-GTP (pink). (<b>C</b>) Normalized peak intensity ratio (<i>I</i>/<i>I</i><sub>0</sub>) analysis of Nt1-222/MglA-GTP interaction. Bars with purple dashed edges indicate peaks disappearing upon MglA-GTP interaction. Above the plot is shown the ZnR and GYF domain limits within Nt1-222. (<b>D</b>) Spectrum region of the <sup>1</sup>H,<sup>15</sup>N HSQCs of <sup>15</sup>N-labeled Nt1-222 free (green) and after addition of 1 (magenta), 2 (light purple), and 3 (purple) molar ratios of MglA-GTP. <sup>1</sup>H,<sup>15</sup>N HSQC of free ZnR<sup>GltJ</sup> (salmon) is also superimposed. Resonances shown correspond to ZnR<sup>GltJ</sup> [same resonances shown in (A) and (B)]. (<b>E</b>) Combined CSPs of Nt1-222 bound to MglA-GTP. (<b>F</b> to <b>H</b>) Disappearance over time of <sup>1</sup>H,<sup>15</sup>N HSQC cross-peaks from cysteines C8 (open circles) and C31 (filled circles) of Zn<sup>2+</sup>-ZnR<sup>GltJ</sup> (F), Zn<sup>2+</sup>Nt1-222 (G), and Zn<sup>2+</sup>Nt1-222 in the presence of MglA-GTP (H) upon incubation with Cd<sup>2+</sup>. (<b>I</b>) Metal exchange rates derived from the experiments shown in (F), (G), and (H). The statistics were obtained applying a Students <i>t</i> test. Significance is assumed for <i>P</i> &lt; 0.05.</div>
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<p> Fig. 6.. The Linker <sup> GltJ </sup> region and… </p>
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<strong class="figure-label">
<p> Fig. 6.. The Linker <sup> GltJ </sup> region and the ZnR <sup> GltJ </sup> domain are functional in vivo. </p>
</strong>
<div class="figure-caption-contents"><p> ( <b> A… </b> </p></div>
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<strong class="figure-label">
Fig. 6.. The Linker<sup>GltJ</sup> region and the ZnR<sup>GltJ</sup> domain are functional in vivo.
</strong>
<div class="figure-caption-contents">(<b>A</b>) AglZ-NG forms highly stable bFAs in a GltJ<sup>ΔZnR</sup>-expressing strain. Shown is a kymograph of a single cell with images captured every 1 s for 120 s. Scale bar, 1 μm. (<b>B</b>) AglZ-containing bFA stability in GltJ mutantexpressing strains. For each strain, the stability of individual bFAs was tracked in single cells. For each strain, the mean cluster stability was determined for <i>n</i> replicates across four independent experiments. The statistics were obtained applying a Wilcoxon test. Significance is assumed for <i>P</i> &lt; 0.1. (<b>C</b>) NG-GltJ<sup>ΔZnR</sup> forms highly stable clusters. Shown is a kymograph of a single cell with images captured every 1 s for 120 s. Scale bar, 1 μm. (<b>D</b>) bFA stability in NG-GltJ and NG-GltJ<sup>ΔZnR</sup> strains. For each strain, the stability of individual bFAs was tracked in single cells. For each strain, the mean cluster stability was determined for <i>n</i> replicates across four independent experiments. The statistics were obtained applying a Wilcoxon test. Significance is assumed for <i>P</i> &lt; 0.1.</div>
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<p> Fig. 7.. ZnR <sup> GltJ </sup> interacts with MglB. </p>
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<div class="figure-caption-contents"><p> (… </p></div>
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<p> Fig. 7.. ZnR <sup> GltJ </sup> interacts with MglB. </p>
</strong>
<div class="figure-caption-contents"><p> ( <b> A </b> ) Combined CSPs between <sup> 1 </sup> H, <sup> 15… </sup> </p></div>
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<strong class="figure-label">
Fig. 7.. ZnR<sup>GltJ</sup> interacts with MglB.
</strong>
<div class="figure-caption-contents">(<b>A</b>) Combined CSPs between <sup>1</sup>H,<sup>15</sup>N resonances of the free ZnR<sup>GltJ</sup> and upon binding to Linker-GYF<sup>GltJ</sup> (blue bars) or MglB (blue-green bars). The secondary structures of ZnR<sup>GltJ</sup> are indicated above the plot. (<b>B</b>) Spectrum regions of <sup>1</sup>H,<sup>15</sup>N HSQC series of <sup>15</sup>N-labeled ZnR<sup>GltJ</sup> free (salmon) and upon addition of increasing amounts of MglB (green shades) up to four molar excess. (<b>C</b>) AlphaFold structure p4rediction of the MglB/ZnR<sup>GltJ</sup> complex. (<b>D</b>) Crystal structure of MglAguanosine 5-<i>O</i>-(3-thiotriphosphate) (GTP-γ-S) bound to MglB homodimer [Protein Data Bank (PDB) ID: 6IZW] (<i>28</i>). (<b>E</b>) AlphaFold structure prediction of the MglA/MglB dimer/ZnR<sup>GltJ</sup> complex. The GTP-γ-S position was inferred by homology to the structure of MglAGTP-γ-S/MglB dimer shown in (D). (<b>F</b>) Initial rate data for Pi release at different concentration of GTP in presence of MglA (orange), MglA + MglB (blue), MglA + MglB + ZnR<sup>GltJ</sup> (red), MglA + MglB<sup>ΔHCt</sup> (green), and MglA + MglB<sup>ΔHCt</sup> + ZnR<sup>GltJ</sup> (purple). (<b>G</b>) Kinetic parameters table. Initial rate data were fitted to the Michaelis-Menten equation. The assays were performed using constant concentration of proteins (MglA, 2 μM; MglB and MglB<sup>ΔHCt</sup>, 4 μM; ZnR<sup>GltJ</sup>, 8 μM) and a range of GTP concentrations (0.5 to 100 μM).</div>
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<p> Fig. 8.. Dynamic assembly of the Agl-Glt… </p>
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<p> Fig. 8.. Dynamic assembly of the Agl-Glt complex in motile <i> Myxococcus </i> cells. </p>
</strong>
<div class="figure-caption-contents"><p> The upper cartoon… </p></div>
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Fig. 8.. Dynamic assembly of the Agl-Glt complex in motile <i>Myxococcus</i> cells.
</strong>
<div class="figure-caption-contents">The upper cartoon depicts the proposed activation/deactivation mechanism of bFAs via connection of the Agl-Glt machinery to the cytoplasmic platform by the GltJ three motifs N-terminal region. For clarity purposes, additional interactions such as MglA/AglZ and MglA/MreB have been omitted. Although it is clear that these interactions are important for bFA assembly, how they integrate with the switch remains to be determined. The lower cartoon represents the cellular location of GltJ-containing complexes for each step of the proposed model. The MglA-MglB proposed concentration gradient within the cell is represented as an orange to green color gradient along the cell axis. 1. When the Agl-Glt complex is inactive and not connected to AglZ and MglA, the switch is OFF, and the ZnR domain is in the closed state through interaction with the Linker motif. 2. At the leading cell pole, the Agl-Glt complex becomes activated via the independent docking of the GYF<sup>GltJ</sup> domain to AglZ and the Linker motif to MglA-GTP, and the ZnR switch is in the free ON conformation. The complex can thus move toward the lagging cell pole. 3. In the front of the cell where MglA concentrations are high, motility IM complexes form bFAs when they interact with the OM complex and thus adhere to the underlying substratum. 4. At the back of the cell where MglB concentration is higher possibly because of the MglA-GTP gradient, MglB is recruited by the ZnR<sup>GltJ</sup> and exerts its GAP activity to convert MglA to the GDP-bound state that cannot bind the Linker motif and thus dissociates from GltJ. Ultimately, this regulation leads to the release of the cytoplasmic platform and back to step 1.</div>
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