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<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="Kgb51y0ObMoBTATe1nMwWUEnS7yUdN2R13VaEotaLnb5N1Z4EGpyq2a1qbg66oAb">
<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="Kgb51y0ObMoBTATe1nMwWUEnS7yUdN2R13VaEotaLnb5N1Z4EGpyq2a1qbg66oAb">
<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="Kgb51y0ObMoBTATe1nMwWUEnS7yUdN2R13VaEotaLnb5N1Z4EGpyq2a1qbg66oAb">
<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="Kgb51y0ObMoBTATe1nMwWUEnS7yUdN2R13VaEotaLnb5N1Z4EGpyq2a1qbg66oAb">
<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="Kgb51y0ObMoBTATe1nMwWUEnS7yUdN2R13VaEotaLnb5N1Z4EGpyq2a1qbg66oAb">
<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="12949097">
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<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://journals.asm.org/doi/10.1128/JB.185.18.5452-5464.2003?url_ver=Z39.88-2003&amp;rfr_id=ori:rid:crossref.org&amp;rfr_dat=cr_pub 0pubmed"
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target="_blank"
rel="noopener"
ref="linksrc=fulltextorjournal_fulltext&amp;is_pmc=True&amp;PrId=3494&amp;itool=Abstract-def&amp;log$=linkouticon&amp;uid=12949097&amp;db=pubmed&amp;nlmid=2985120R"
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Free PMC article
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<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="/12949097/citations/"
data-citation-style="nlm"
data-pubmed-format-link="/12949097/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="12949097" 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%2F12949097%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="Kgb51y0ObMoBTATe1nMwWUEnS7yUdN2R13VaEotaLnb5N1Z4EGpyq2a1qbg66oAb"><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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TodK, a putative histidine protein kinase, regulates timing of fruiting body morphogenesis in Myxococcus xanthus
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In response to starvation, Myxococcus xanthus initiates a developmental program that results in the formation of spore-filled multicellular fruiting bodies. Fruiting body formation depends on the temporal and spatial coordination of aggregation and sporulation. These two processes are induced by the cell surface-associated C signal, with aggregation being induced after 6 h and sporulation being induced once cells have completed the aggregation process. We report the identification of TodK, a putative histidine protein kinase of two-component regulatory systems that is important for the correct timing of aggregation and sporulation. Loss of TodK function results in early aggregation and early, as well as increased levels of, sporulation. Transcription of todK decreases 10-fold in response to starvation independently of the stringent response. Loss of TodK function specifically results in increased expression of a subset of C-signal-dependent genes. Accelerated development in a todK mutant depends on the known components in the C-signal transduction pathway. TodK is not important for synthesis of the C signal. From these results we suggest that TodK is part of a signal transduction system which converges on the C-signal transduction pathway to negatively regulate aggregation, sporulation, and the expression of a subset of C-signal-dependent genes. TodK and the SdeK histidine protein kinase, which is part of a signal transduction system that converges on the C-signal transduction pathway to stimulate aggregation, sporulation, and C-signal-dependent gene expression, act in independent genetic pathways. We suggest that the signal transduction pathways defined by TodK and SdeK act in concert with the C-signal transduction pathway to control the timing of aggregation and sporulation.
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<strong class="figure-label">
<p> FIG. 1. </p>
</strong>
<div class="figure-caption-contents"><p> Developmental phenotypes of <i> todK </i> and… </p></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG. 1. </p>
</strong>
<div class="figure-caption-contents"><p> Developmental phenotypes of <i> todK </i> and <i> dotR </i> mutants. The indicated strains were exposed to… </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">
FIG. 1.
</strong>
<div class="figure-caption-contents">Developmental phenotypes of <i>todK</i> and <i>dotR</i> mutants. The indicated strains were exposed to starvation on CF agar for the indicated periods of time and viewed in a Leica MZ8 stereomicroscope. The genotypes of the strains are indicated below their names. Bar, 0.2 mm.</div>
</figcaption>
</figure>
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data-slide-index="1"
data-label-slug="fig-2">
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href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/dfbf/193762/70c92de9b9b7/jb1830445002.jpg"
itemprop="contentUrl"
aria-describedby="figure-caption-1"
role="button"
data-image-width="440"
data-image-height="428"
data-image-alt="FIG. 2."
data-pmc-id="PMC193762"
data-figure-id="f2">
<img class="figure-thumb" itemprop="thumbnail"
id="article-image-1"
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/dfbf/193762/451a98ffdffa/jb1830445002.gif"
alt="FIG. 2." />
</a>
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<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> FIG. 2. </p>
</strong>
<div class="figure-caption-contents"><p> Physical map of the <i> todK… </i> </p></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG. 2. </p>
</strong>
<div class="figure-caption-contents"><p> Physical map of the <i> todK </i> - <i> dotR </i> region. ORFs are indicated by open… </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">
FIG. 2.
</strong>
<div class="figure-caption-contents">Physical map of the <i>todK</i>-<i>dotR</i> region. ORFs are indicated by open arrows. Arrows indicate the directions of transcription. Coordinates are relative to +1, the transcriptional start site of <i>todK</i>. Below the physical map, the overlap between the <i>todK</i> and <i>dotR</i> stop codons is shown. The hatched arrowhead indicates the site of insertion of mini-Tn<i>5</i>(<i>tet</i>) Ω8846 at position 839. Plasmids containing the indicated DNA fragments of the <i>todK</i>-<i>dotR</i> locus are shown below the map.</div>
</figcaption>
</figure>
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data-label-slug="fig-3">
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itemprop="contentUrl"
aria-describedby="figure-caption-2"
role="button"
data-image-width="591"
data-image-height="519"
data-image-alt="FIG. 3."
data-pmc-id="PMC193762"
data-figure-id="f3">
<img class="figure-thumb" itemprop="thumbnail"
id="article-image-2"
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alt="FIG. 3." />
</a>
<meta itemprop="width" itemtype="http://schema.org/ImageObject" content="591">
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<div class="caption-wrap">
<strong class="figure-label">
<p> FIG. 3. </p>
</strong>
<div class="figure-caption-contents"><p> Sequence analyses of TodK and… </p></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG. 3. </p>
</strong>
<div class="figure-caption-contents"><p> Sequence analyses of TodK and DotR. (A) Domain organization of TodK. In the… </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">
FIG. 3.
</strong>
<div class="figure-caption-contents">Sequence analyses of TodK and DotR. (A) Domain organization of TodK. In the sensor domain the two PAS domains are indicated; in the kinase domain, the conserved H, N, D/F, and G boxes are indicated. The asterisk over the H box indicates the conserved histidine residue, the site of autophosphorylation in other sensor histidine protein kinases. (B) Alignment of transmitter domain of TodK with those in AsgD from <i>M</i>. <i>xanthus</i> (4), SdeK from <i>M</i>. <i>xanthus</i> (12), and ResE from <i>B</i>. <i>subtilis</i> (59). The conserved H, N, D/F, and G boxes are indicated. The asterisk over the H box indicates the conserved histidine residue, the site of autophosphorylation in other sensor histidine protein kinases. Residues on black, dark gray, and light gray backgrounds are conserved 100, 75, and 50%, respectively. The alignment was made using CLUSTALX (65); the presentation was made using GENEDOC (K. B. Nicholas and H. B. Nicholas, Jr., 1997). (C) Alignment of DotR with receiver domains in PhoP from <i>B</i>. <i>subtilis</i> (38), PhoB from <i>C</i>. <i>crescentus</i> (13), and PilG from <i>P</i>. <i>aeruginosa</i> (9). The asterisks indicate the highly conserved signature residues of receiver domains (67). Conserved residues are indicated as in panel B. The alignment and presentation were prepared as for panel B.</div>
</figcaption>
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data-label-slug="fig-4">
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itemprop="contentUrl"
aria-describedby="figure-caption-3"
role="button"
data-image-width="487"
data-image-height="1203"
data-image-alt="FIG. 4."
data-pmc-id="PMC193762"
data-figure-id="f4">
<img class="figure-thumb" itemprop="thumbnail"
id="article-image-3"
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/dfbf/193762/c3f357620ad1/jb1830445004.gif"
alt="FIG. 4." />
</a>
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<div class="caption-wrap">
<strong class="figure-label">
<p> FIG. 4. </p>
</strong>
<div class="figure-caption-contents"><p> Regulation of transcription of <i> todK… </i> </p></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG. 4. </p>
</strong>
<div class="figure-caption-contents"><p> Regulation of transcription of <i> todK </i> . (A) Mapping of the 5 end of… </p></div>
</div>
<figcaption id="figure-caption-3" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
<strong class="figure-label">
FIG. 4.
</strong>
<div class="figure-caption-contents">Regulation of transcription of <i>todK</i>. (A) Mapping of the 5 end of the <i>todK</i> transcript by primer extension analysis. C, T, A, and G show the sequence ladders. Total RNA was isolated from DK1622 and SA1634 that had been starved for the indicated periods of time on TPM starvation agar. The 5 end of the <i>todK</i> transcript is indicated by the arrow. The sequence around the transcriptional start site is indicated to the left. The broken arrow indicates the transcriptional start site (+1). (B) DNA sequence of the <i>todK</i> promoter. The broken arrowindicates +1. The putative 10 and 35 sequences are indicated in boldface. The putative start codon of <i>todK</i> is indicated in boldface together with the putative ribosomal binding site (RBS). The underlining indicates the location of the primer used in the primer extension analyses. (C) Quantitative RT-PCR analysis of the <i>todK</i> transcript. Total RNA was isolated from DK1622 and SA1634 that had been starved for the indicated periods of time on TPM starvation agar. The level of <i>todK</i> transcript detected is expressed as relative units per nanogram of total RNA.</div>
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itemprop="contentUrl"
aria-describedby="figure-caption-4"
role="button"
data-image-width="612"
data-image-height="637"
data-image-alt="FIG. 5."
data-pmc-id="PMC193762"
data-figure-id="f5">
<img class="figure-thumb" itemprop="thumbnail"
id="article-image-4"
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/dfbf/193762/edf1c255e753/jb1830445005.gif"
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</a>
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<strong class="figure-label">
<p> FIG. 5. </p>
</strong>
<div class="figure-caption-contents"><p> Effect of <i> todK </i> mutation on… </p></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG. 5. </p>
</strong>
<div class="figure-caption-contents"><p> Effect of <i> todK </i> mutation on developmental gene expression. (A) Expression of the indicated… </p></div>
</div>
<figcaption id="figure-caption-4" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
<strong class="figure-label">
FIG. 5.
</strong>
<div class="figure-caption-contents">Effect of <i>todK</i> mutation on developmental gene expression. (A) Expression of the indicated <i>lacZ</i> fusions in wild-type DK1622 cells and in <i>todK</i> SA1634 cells. Cells were starved on TPM agar, samples were withdrawn at the indicated time points, and specific activities of β-galactosidase were determined. The experiments were done in triplicate. Error bars indicate standard deviations. Specific activities of β-galactosidase are given as nanomoles of orthonitrophenyl minute<sup>1</sup> milligram of protein<sup>1</sup>. (B) Immunoblot analysis of the <i>csgA</i> proteins in wild-type DK1622 and in <i>todK</i> SA1634 cells. Cells were starved on TPM agar, and total cell lysates were prepared from samples withdrawn at the indicated time points (in hours), separated by sodium dodecyl sulfate-polyacrylamide gel electrophoresis, and reacted with anti-CsgA antibodies (35). The closed arrow indicates the 25-kDa full-length CsgA protein, and the open arrow indicates the 17-kDa CsgA protein.</div>
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itemprop="contentUrl"
aria-describedby="figure-caption-5"
role="button"
data-image-width="491"
data-image-height="392"
data-image-alt="FIG. 6."
data-pmc-id="PMC193762"
data-figure-id="f6">
<img class="figure-thumb" itemprop="thumbnail"
id="article-image-5"
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/dfbf/193762/2755553fe675/jb1830445006.gif"
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</a>
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<strong class="figure-label">
<p> FIG. 6. </p>
</strong>
<div class="figure-caption-contents"><p> Epistasis analysis of the relationship… </p></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG. 6. </p>
</strong>
<div class="figure-caption-contents"><p> Epistasis analysis of the relationship between <i> todK </i> and <i> sdeK </i> . The indicated strains… </p></div>
</div>
<figcaption id="figure-caption-5" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
<strong class="figure-label">
FIG. 6.
</strong>
<div class="figure-caption-contents">Epistasis analysis of the relationship between <i>todK</i> and <i>sdeK</i>. The indicated strains were exposed to starvation on CF agar for the indicated time points and viewed in a Leica MZ8 stereomicroscope. The genotypes of the strains are indicated below their names. Bar, 0.2 mm.</div>
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itemprop="contentUrl"
aria-describedby="figure-caption-6"
role="button"
data-image-width="493"
data-image-height="309"
data-image-alt="FIG. 7."
data-pmc-id="PMC193762"
data-figure-id="f7">
<img class="figure-thumb" itemprop="thumbnail"
id="article-image-6"
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/dfbf/193762/98d9589f5cb6/jb1830445007.gif"
alt="FIG. 7." />
</a>
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<strong class="figure-label">
<p> FIG. 7. </p>
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<div class="figure-caption-contents"><p> Model of the C-signal transduction… </p></div>
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<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG. 7. </p>
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<div class="figure-caption-contents"><p> Model of the C-signal transduction pathway. The signaling event in the pathway is… </p></div>
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<figcaption id="figure-caption-6" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
<strong class="figure-label">
FIG. 7.
</strong>
<div class="figure-caption-contents">Model of the C-signal transduction pathway. The signaling event in the pathway is the interaction between C signal exposed on the surface of one cell and the hypothetical C-signal sensor on a second cell. Starvation results in a RelA- and (p)ppGpp-dependent transcription of the <i>csgA</i> gene and in the accumulation of the intercellular A signal as well as in the induction of <i>sdeK</i> transcription. Moreover, starvation results in a RelA-independent inhibition of <i>todK</i> transcription. The A and E signals induce transcription of <i>fruA</i> (11, 43). C-signal transmission activates <i>csgA</i> transcription, possibly by activating the proteins in the <i>act</i> operon (14), and FruA, conceivably by stimulating the activity of the cognate FruA histidine protein kinase (labeled HPK); subsequently the phosphoryl group is transferred from the kinase to FruA. Phosphorylated FruA—directly or indirectly—alters the activity of the Frz signal transduction system in such a way that aggregation is induced. In addition phosphorylated FruA activates transcription—directly or indirectly—of C-signal-dependent genes, here exemplified by the <i>dev</i> operon. The possible sites of action of the SdeK kinase and the TodK kinase are indicated by the shaded box (see text for explanation). All the components in the cell to the right are also present in the cell to the left; for simplicity, only <i>csgA</i> is shown in the cell to the left.</div>
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