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|
||
</div>
|
||
|
||
<span class="search-input-link user-guide-link" role="navigation" aria-label="User guide">
|
||
<a href="/help/"
|
||
data-ga-category="featured_link"
|
||
data-ga-action="user_guide">
|
||
User Guide
|
||
</a>
|
||
</span>
|
||
|
||
</div>
|
||
</div>
|
||
|
||
|
||
</form>
|
||
|
||
|
||
<div id="article-page-header">
|
||
<div id="article-top-actions-bar" class="article-top-actions-bar">
|
||
<div class="inner-wrap">
|
||
<div class="actions-container">
|
||
<div class="return-to-search">
|
||
|
||
|
||
</div>
|
||
|
||
<div class="multiple-results-actions " role="region" aria-label="save, email, send to">
|
||
<button id="save-results-panel-trigger"
|
||
type="button"
|
||
class="save-results save-results-panel-trigger"
|
||
aria-expanded="false"
|
||
aria-controls="save-action-panel"
|
||
data-ga-category="save_share"
|
||
data-ga-action="save"
|
||
data-ga-label="open">
|
||
Save
|
||
</button><button id="email-results-panel-trigger"
|
||
type="button"
|
||
class="email-results email-results-panel-trigger"
|
||
aria-expanded="false"
|
||
aria-controls="email-action-panel"
|
||
data-ga-category="save_share"
|
||
data-ga-action="email"
|
||
data-ga-label="open">
|
||
Email
|
||
</button><div class="more-actions dropdown-block"><button id="more-actions-trigger"
|
||
type="button"
|
||
class="trigger more-actions-trigger"
|
||
ref="linksrc=show_moreactions_btn"
|
||
aria-label="Send to"
|
||
aria-controls="more-actions-dropdown">Send to</button><div id="more-actions-dropdown" class="dropdown dropdown-container" hidden><div class="content"><ul class="more-actions-links"><li><a class="clipboard-panel-trigger dropdown-block-link "
|
||
href="#"
|
||
role="button"
|
||
data-ga-category="save_share"
|
||
data-ga-action="clipboard"
|
||
data-ga-label="send">
|
||
Clipboard
|
||
</a></li><li><a role="button" class="dropdown-block-link bibliography-trigger-target" href="https://account.ncbi.nlm.nih.gov/?back_url=https%3A%2F%2Fpubmed.ncbi.nlm.nih.gov%2F20956384%2F%23open-bibliography-panel">My Bibliography</a></li><li><a role="button" class="dropdown-block-link collections-trigger-target" href="https://account.ncbi.nlm.nih.gov/?back_url=https%3A%2F%2Fpubmed.ncbi.nlm.nih.gov%2F20956384%2F%23open-collections-panel">Collections</a></li><li><a role="button" class="citation-manager-panel-trigger dropdown-block-link citation-manager-trigger-target" href="#">Citation manager</a></li></ul></div></div></div>
|
||
</div>
|
||
|
||
|
||
|
||
<div class="display-options" data-format-key="format"
|
||
data-default-format-value="abstract"
|
||
data-current-format-value="abstract" role="region">
|
||
<button
|
||
class="trigger usa-button"
|
||
title="Change format"
|
||
data-ga-category="display_options"
|
||
data-ga-action="display_options_dialog">
|
||
<span class="button-label">Display options</span>
|
||
</button>
|
||
|
||
<div class="dropdown" hidden>
|
||
<div class="title">
|
||
Display options
|
||
</div>
|
||
|
||
<div class="content">
|
||
|
||
<div class="format-container">
|
||
<span class="option-label">
|
||
Format
|
||
</span>
|
||
<select name="format" data-initial-value="abstract" aria-label="Display format" id="article-display-format">
|
||
<option value="abstract">Abstract</option>
|
||
|
||
<option value="pubmed">PubMed</option>
|
||
|
||
<option value="pmid">PMID</option>
|
||
|
||
</select>
|
||
</div>
|
||
|
||
</div>
|
||
</div>
|
||
</div>
|
||
<input type="hidden" name="csrfmiddlewaretoken" value="MguLz4CvjVb1pHOtMWXHFb3NCjA1dwrL33eQcU5RTwYvj8UjpfAJ9jzranid67Z5">
|
||
</div>
|
||
</div>
|
||
</div>
|
||
|
||
<div id="article-panels-container" class="article-panels-container">
|
||
<div id="save-action-panel" class="save-action-panel action-panel" aria-hidden="true">
|
||
<div class="inner-wrap">
|
||
<h2 class="action-panel-heading">
|
||
Save citation to file
|
||
</h2>
|
||
<form id="save-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="MguLz4CvjVb1pHOtMWXHFb3NCjA1dwrL33eQcU5RTwYvj8UjpfAJ9jzranid67Z5">
|
||
|
||
|
||
|
||
<div class="action-panel-control-wrap">
|
||
<label for="save-action-format" class="action-panel-label">
|
||
Format:
|
||
</label>
|
||
<select id="save-action-format" name="results-format" class="action-panel-selector">
|
||
<option value="summary-text">Summary (text)</option>
|
||
<option value="pubmed-text">PubMed</option>
|
||
<option value="pmid">PMID</option>
|
||
<option value="abstract">Abstract (text)</option>
|
||
<option value="csv">CSV</option>
|
||
</select>
|
||
</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="save"
|
||
data-ga-label="save">
|
||
Create file
|
||
</button>
|
||
<button class="action-panel-cancel"
|
||
aria-label="Close 'Save citations to file' panel"
|
||
ref="linksrc=close_save_panel"
|
||
aria-controls="save-action-panel"
|
||
aria-expanded="false"
|
||
data-ga-category="save_share"
|
||
data-ga-action="save"
|
||
data-ga-label="cancel">
|
||
Cancel
|
||
</button>
|
||
</div>
|
||
</form>
|
||
</div>
|
||
</div>
|
||
|
||
|
||
|
||
|
||
<div id="email-action-panel"
|
||
class="email-action-panel action-panel"
|
||
aria-hidden="true"
|
||
data-recaptcha-script-url="https://www.recaptcha.net/recaptcha/enterprise.js?render=explicit&onload=onloadCaptchaCallback"
|
||
data-recaptcha-enabled="true">
|
||
<div class="inner-wrap">
|
||
<h2 class="action-panel-heading">
|
||
Email citation
|
||
</h2>
|
||
<form id="email-action-panel-form"
|
||
class="action-panel-content action-form"
|
||
action="/send-email/"
|
||
data-by-search-action="/results-export-email-by-search-data/"
|
||
data-by-ids-action="/send-email/"
|
||
data-by-search-method="post"
|
||
data-by-ids-method="post"
|
||
method="post"
|
||
>
|
||
<input type="hidden" name="csrfmiddlewaretoken" value="MguLz4CvjVb1pHOtMWXHFb3NCjA1dwrL33eQcU5RTwYvj8UjpfAJ9jzranid67Z5">
|
||
|
||
<div class="action-panel-control-wrap">
|
||
<span class="action-panel-label">
|
||
Subject:
|
||
</span>
|
||
<span class="email-subject" id="email-subject" aria-label="Email subject">1 selected item: 20956384 - PubMed</span>
|
||
<input type="hidden" name="email-subject" value="1 selected item: 20956384"/>
|
||
<input type="hidden" name="email-subject-hash" value="180801b378632cc7fec59231d7282eaf73f1b3b7ab44d0f590c56f8e8df8b36c"/>
|
||
</div>
|
||
|
||
<div class="action-panel-control-wrap">
|
||
<label for="email-to" class="action-panel-label required-field-asterisk">
|
||
To:
|
||
</label>
|
||
<input type="email" aria-label="Recipient Email Address" placeholder="email@example.com" id="email-to" class="email-to" pattern="^[a-zA-Z0-9_.+-]+@[a-zA-Z0-9-]+\.[a-zA-Z0-9-.]+$" maxlength="100" required>
|
||
</div>
|
||
|
||
<div class="action-panel-control-wrap">
|
||
<label for="email-from" class="action-panel-label">
|
||
From:
|
||
</label>
|
||
<input type="email" aria-label="Sender Email Address" placeholder="email@example.com" id="email-from" class="email-from" pattern="^[a-zA-Z0-9_.+-]+@[a-zA-Z0-9-]+\.[a-zA-Z0-9-.]+$" 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="MguLz4CvjVb1pHOtMWXHFb3NCjA1dwrL33eQcU5RTwYvj8UjpfAJ9jzranid67Z5">
|
||
|
||
|
||
|
||
<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="[^"&=<>\/]*" title="The following characters are not allowed in the Name field: "&=<>/"
|
||
maxlength="100"
|
||
data-ga-category="save_share"
|
||
data-ga-action="create_collection"
|
||
data-ga-label="non_favorties_collection">
|
||
<div class="collections-new-name-too-long usa-input-error-message selection-validation-message">
|
||
Name must be less than 100 characters
|
||
</div>
|
||
</div>
|
||
<div class="action-panel-control-wrap existing-collections-controls">
|
||
<label for="collections-action-panel-add-to-existing" class="action-panel-label">
|
||
Choose a collection:
|
||
</label>
|
||
<select id="collections-action-panel-add-to-existing"
|
||
class="action-panel-selector collections-action-add-to-existing"
|
||
data-ga-category="save_share"
|
||
data-ga-action="select_collection"
|
||
data-ga-label="($('#collections-action-add-to-existing').val() === 'Favorites') ? 'Favorites' : 'non_favorites_collection'">
|
||
</select>
|
||
<div class="collections-retry-load-on-error usa-input-error-message selection-validation-message">
|
||
Unable to load your collection due to an error<br>
|
||
<a href="#">Please try again</a>
|
||
</div>
|
||
</div>
|
||
</div>
|
||
|
||
<div class="action-panel-actions">
|
||
<button class="action-panel-submit"
|
||
type="submit"
|
||
data-loading-label="Adding..."
|
||
data-pinger-ignore
|
||
data-ga-category="save_share"
|
||
data-ga-action="collections"
|
||
data-ga-label="add">
|
||
Add
|
||
</button>
|
||
<button class="action-panel-cancel"
|
||
aria-label="Close 'Add to Collections' panel"
|
||
ref="linksrc=close_collections_panel"
|
||
aria-controls="collections-action-panel"
|
||
aria-expanded="false"
|
||
data-ga-category="save_share"
|
||
data-ga-action="collections"
|
||
data-ga-label="cancel">
|
||
Cancel
|
||
</button>
|
||
</div>
|
||
</form>
|
||
</div>
|
||
</div>
|
||
|
||
<div id="bibliography-action-panel"
|
||
class="bibliography-action-panel action-panel in-progress-dots-panel"
|
||
aria-hidden="true"
|
||
data-bibliography-open-panel-enabled="false"
|
||
data-bibliography-open-panel-url-hash="#open-bibliography-panel">
|
||
<div class="inner-wrap">
|
||
<h3 class="action-panel-heading">
|
||
Add to My Bibliography
|
||
</h3>
|
||
<form id="bibliography-action-panel-form"
|
||
class="bibliography-action-panel-form action-panel-content action-form action-panel-smaller-selectors"
|
||
data-add-to-bibliography-max-amount="100"
|
||
data-add-to-bibliography-batch-size="10"
|
||
data-bibliography-delegates-url="/list-bibliography-delegates/"
|
||
data-add-to-bibliography-url="/add-to-bibliography/"
|
||
data-get-article-ids-by-search-url="/get-article-ids-by-search/"
|
||
data-mybib-root-url="https://www.ncbi.nlm.nih.gov/myncbi/collections/mybibliography/">
|
||
|
||
<input type="hidden" name="csrfmiddlewaretoken" value="MguLz4CvjVb1pHOtMWXHFb3NCjA1dwrL33eQcU5RTwYvj8UjpfAJ9jzranid67Z5">
|
||
|
||
|
||
|
||
<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="MguLz4CvjVb1pHOtMWXHFb3NCjA1dwrL33eQcU5RTwYvj8UjpfAJ9jzranid67Z5">
|
||
|
||
<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="MguLz4CvjVb1pHOtMWXHFb3NCjA1dwrL33eQcU5RTwYvj8UjpfAJ9jzranid67Z5">
|
||
|
||
|
||
|
||
<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="MguLz4CvjVb1pHOtMWXHFb3NCjA1dwrL33eQcU5RTwYvj8UjpfAJ9jzranid67Z5">
|
||
|
||
<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="20956384">
|
||
<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://rupress.org/jcb/article-lookup/doi/10.1083/jcb.201004108"
|
||
target="_blank"
|
||
rel="noopener"
|
||
|
||
ref="linksrc=fulltextorjournal_fulltext&is_pmc=False&PrId=7898&itool=Abstract-def&log$=linkouticon&uid=20956384&db=pubmed&nlmid=0375356"
|
||
title="See full text options at Silverchair Information Systems"
|
||
|
||
data-ga-category="full_text"
|
||
data-ga-action="Silverchair Information Systems"
|
||
data-ga-label="20956384"
|
||
><img src="https://cdn.ncbi.nlm.nih.gov/corehtml/query/egifs/https:--rupress.org-images-pubmed-jcb_free.png" alt="Silverchair Information Systems full text link"><span class="text">
|
||
|
||
Silverchair Information Systems
|
||
|
||
</span></a><a class="link-item
|
||
pmc
|
||
"
|
||
href="https://pmc.ncbi.nlm.nih.gov/articles/pmid/20956384/"
|
||
target="_blank"
|
||
rel="noopener"
|
||
|
||
ref="linksrc=fulltextorjournal_fulltext&is_pmc=True&PrId=3494&itool=Abstract-def&log$=linkouticon&uid=20956384&db=pubmed&nlmid=0375356"
|
||
title="Free full text at PubMed Central"
|
||
|
||
data-ga-category="full_text"
|
||
data-ga-action="PMC"
|
||
data-ga-label="20956384"
|
||
><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="/20956384/citations/"
|
||
data-citation-style="nlm"
|
||
data-pubmed-format-link="/20956384/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="20956384" 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%2F20956384%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="MguLz4CvjVb1pHOtMWXHFb3NCjA1dwrL33eQcU5RTwYvj8UjpfAJ9jzranid67Z5"><div class="choice-group" role="radiogroup"><ul class="radio-group-items"><li><input type="radio"
|
||
id="collections-action-dialog-new"
|
||
class="collections-new"
|
||
name="collections"
|
||
value="new"
|
||
data-ga-category="collections_button"
|
||
data-ga-action="click"
|
||
data-ga-label="collections_radio_new"><label for="collections-action-dialog-new">Create a new collection</label></li><li><input type="radio"
|
||
id="collections-action-dialog-existing"
|
||
class="collections-existing"
|
||
name="collections"
|
||
value="existing"
|
||
checked="true"
|
||
data-ga-category="collections_button"
|
||
data-ga-action="click"
|
||
data-ga-label="collections_radio_existing"><label for="collections-action-dialog-existing">Add to an existing collection</label></li></ul></div><div class="controls-wrapper"><div class="action-panel-control-wrap new-collections-controls"><label for="collections-action-dialog-add-to-new" class="action-panel-label required-field-asterisk">
|
||
Name your collection:
|
||
</label><input
|
||
type="text"
|
||
name="add-to-new-collection"
|
||
id="collections-action-dialog-add-to-new"
|
||
class="collections-action-add-to-new"
|
||
pattern="[^"&=<>\/]*" title="The following characters are not allowed in the Name field: "&=<>/"
|
||
maxlength="100"
|
||
data-ga-category="collections_button"
|
||
data-ga-action="create_collection"
|
||
data-ga-label="non_favorties_collection"><div class="collections-new-name-too-long usa-input-error-message selection-validation-message">
|
||
Name must be less than 100 characters
|
||
</div></div><div class="action-panel-control-wrap existing-collections-controls"><label for="collections-action-dialog-add-to-existing" class="action-panel-label">
|
||
Choose a collection:
|
||
</label><select id="collections-action-dialog-add-to-existing"
|
||
class="action-panel-selector collections-action-add-to-existing"
|
||
data-ga-category="collections_button"
|
||
data-ga-action="select_collection"
|
||
data-ga-label="($('.collections-action-add-to-existing').val() === '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="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" data-format-key="format"
|
||
data-default-format-value="abstract"
|
||
data-current-format-value="abstract" role="region"><button
|
||
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A mechanism for vertebrate Hedgehog signaling: recruitment to cilia and dissociation of SuFu-Gli protein complexes
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A mechanism for vertebrate Hedgehog signaling: recruitment to cilia and dissociation of SuFu-Gli protein complexes
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<span class="authors-list-item "><span class="full-name">Hanna Tukachinsky</span><span class="citation-separator"> et al.</span></span>
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In vertebrates, Hedgehog (Hh) signaling initiated in primary cilia activates the membrane protein Smoothened (Smo) and leads to activation of Gli proteins, the transcriptional effectors of the pathway. In the absence of signaling, Gli proteins are inhibited by the cytoplasmic protein Suppressor of Fused (SuFu). It is unclear how Hh activates Gli and whether it directly regulates SuFu. We find that Hh stimulation quickly recruits endogenous SuFu-Gli complexes to cilia, suggesting a model in which Smo activates Gli by relieving inhibition by SuFu. In support of this model, we find that Hh causes rapid dissociation of the SuFu-Gli complex, thus allowing Gli to enter the nucleus and activate transcription. Activation of protein kinase A (PKA), an inhibitor of Hh signaling, blocks ciliary localization of SuFu-Gli complexes, which in turn prevents their dissociation by signaling. Our results support a simple mechanism in which Hh signals at vertebrate cilia cause dissociation of inactive SuFu-Gli complexes, a process inhibited by PKA.
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<div class="figure-caption-contents"><b> Endogenous SuFu is rapidly recruited… </b></div>
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<div class="figure-caption-contents"><b> Endogenous SuFu is rapidly recruited to primary cilia by Hh signaling, paralleling recruitment… </b></div>
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<div class="figure-caption-contents"><b>Endogenous SuFu is rapidly recruited to primary cilia by Hh signaling, paralleling recruitment of endogenous Smo, Gli2, and Gli3-FL.</b> (A) Fluorescence micrographs of cilia from untreated cells or cells treated with Shh. Cilia were detected by staining against acetylated tubulin. Because the anti-GliC antibody detects both Gli2 and Gli3-FL, Gli2<sup>−/−</sup> and Gli3<sup>−/−</sup> MEFs are shown to demonstrate ciliary recruitment of Gli2 and Gli3-FL separately. The tip of the cilium points to the left. (B) Cells were treated with Shh for varying amounts of time, and ciliary recruitment of SuFu, Smo, Gli2, and Gli3-FL was determined. Asterisks indicate p-values for ciliary recruitment at 1 h compared with t = 0 (*, P < 0.05; **, P < 0.01; ***, P < 0.001). P < 0.05 for all later time points. (C) In NIH-3T3 cells stimulated with Shh for 1 h, SuFu and Gli proteins localize at the tip, whereas Smo localizes along the length of cilia. Cilia were stained as in A, and centrioles were stained with anti–γ-tubulin. (D) Endogenous SuFu and Gli proteins colocalize at the tips of primary cilia in SAG-treated NIH-3T3 cells. (left) Cilia costained for endogenous SuFu (rabbit antibody) and Gli (goat antibody). (right) Cilia costained for Smo (rabbit antibody) and Gli (goat antibody). (E) Cilia counts for the experiment in D (left). Endogenous SuFu and Gli colocalize both in the resting and stimulated states of the Hh pathway. (F) Recruitment of SuFu, Smo, and Gli to cilia in response to Hh stimulation does not require new protein synthesis. Ciliary localization was determined in NIH-3T3 cells treated or not with Shh in the presence or absence of CHX. (G) Inhibition of protein synthesis does not block the transcriptional output of the Hh pathway. Transcription of the direct transcriptional targets <i>Gli1</i> and <i>Ptch1</i> was assayed by Q-PCR after 3 and 6 h of stimulation with Shh in the presence or absence of CHX. wt, wild type. Error bars indicate mean ± SD for three independent counts. Bars, 2 µm.</div>
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<div class="figure-caption-contents"><b> Hh-dependent recruitment of SuFu and… </b></div>
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<div class="figure-caption-contents"><b> Hh-dependent recruitment of SuFu and Gli proteins to cilia requires active Smo. </b> (A)…</div>
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|
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|
||
</div>
|
||
|
||
<figcaption id="figure-caption-1" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
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||
<strong class="figure-label">
|
||
Figure 2.
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b>Hh-dependent recruitment of SuFu and Gli proteins to cilia requires active Smo.</b> (A) NIH-3T3 cells were treated with the Smo agonist SAG or with the antagonist Cyc. SuFu and Gli are recruited to cilia by SAG but not by Cyc, although both SAG and Cyc recruit Smo to cilia. The tips of cilia point to the left. Bar, 2 µm. (B) Cilia counts for the experiment in A. (C) Q-PCR assay of Hh pathway target genes for the experiment in A. (D) Maintaining increased levels of SuFu and Gli at cilia is continuously dependent on active Smo. Cyc was added in the presence of Shh to NIH-3T3 cells prestimulated with Shh for 3 h. Ciliary localization was determined before and after 3 h of Shh stimulation and 1 and 3 h after Cyc addition. (E) NIH-3T3 cells were stimulated with Shh for 3 h followed by incubation with the Smo antagonist SANT-1 for 3 h. Ciliary localization of SuFu, Gli, and Smo was measured at the indicated times. P < 0.002 for the recruitment of Smo, SuFu, and Gli by Shh stimulation. P-values for exit from the cilium were calculated relative to ciliary localization after 3 h of Hh stimulation. Asterisks indicate the p-values for ciliary exit (*, P < 0.05; **, P < 0.01; ***, P < 0.001). Error bars indicate mean ± SD.</div>
|
||
|
||
|
||
|
||
|
||
</figcaption>
|
||
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||
|
||
</figure>
|
||
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||
<figure class="figure-item "
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||
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|
||
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data-slide-index="2"
|
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data-label-slug="figure-3">
|
||
<a class="figure-link"
|
||
href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/5a36/2958481/df54cc39b96a/JCB_201004108_GS_Fig3.jpg"
|
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role="button"
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data-image-width="664"
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|
||
data-image-alt="Figure 3."
|
||
data-pmc-id="PMC2958481"
|
||
data-figure-id="fig3">
|
||
<img class="figure-thumb" itemprop="thumbnail"
|
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id="article-image-2"
|
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src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/5a36/2958481/555de519317e/JCB_201004108_GS_Fig3.gif"
|
||
alt="Figure 3." />
|
||
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|
||
|
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|
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<strong class="figure-label">
|
||
<p> Figure 3. </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b> Localization of endogenous SuFu and… </b></div>
|
||
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||
|
||
|
||
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||
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|
||
</div>
|
||
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|
||
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||
|
||
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||
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||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 3. </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b> Localization of endogenous SuFu and Gli to cilia is antagonized by PKA. </b> (A)…</div>
|
||
|
||
|
||
|
||
|
||
</div>
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||
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<figcaption id="figure-caption-2" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
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||
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||
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<strong class="figure-label">
|
||
Figure 3.
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b>Localization of endogenous SuFu and Gli to cilia is antagonized by PKA.</b> (A) Activation of PKA by FSK blocks localization of endogenous SuFu and Gli proteins to cilia. NIH-3T3 cells were treated with or without Shh and FSK. Shh, FSK, or Shh and FSK recruit Smo to the cilium; in contrast, endogenous SuFu and Gli are removed from cilia by FSK, both in the presence and absence of Shh stimulation. (B) Cilia counts for the experiment in A. (C) Q-PCR analysis of the experiment in A. Inhibition of SuFu and Gli ciliary localization by FSK correlates with complete inhibition of the transcriptional output of the Hh pathway. Error bar indicates mean ± SD. (D) FSK inhibits localization of SuFu and Gli to primary cilia in Smo<sup>−/−</sup> MEFs. Percentages shown indicate corresponding ciliary counts. (E) NIH-3T3 cells were incubated with or without Shh (left) or with or without FSK (right) followed by immunoblotting for SuFu, Gli3-FL, GSK3, and α-tubulin. (top) Numbers shown indicate the levels of Gli3-FL in each lane relative to α-tubulin. FSK treatment causes only a slight reduction in Gli3-FL, which is much smaller than the decrease caused by Shh. (F) NIH-3T3 cells stably expressing a Gli1–SuFu fusion were incubated with control media, SAG, or FSK. The Gli1–SuFu fusion localizes to cilia in unstimulated cells, and its localization is increased by SAG. FSK treatment completely blocks ciliary localization of the Gli1–SuFu fusion. Percentages shown indicate ciliary localization of the fusion. Bars, 2 µm.</div>
|
||
|
||
|
||
|
||
|
||
</figcaption>
|
||
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||
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||
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|
||
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||
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|
||
<a class="figure-link"
|
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href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/5a36/2958481/b2362f009782/JCB_201004108_GS_Fig4.jpg"
|
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|
||
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|
||
role="button"
|
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data-image-width="664"
|
||
data-image-height="483"
|
||
data-image-alt="Figure 4."
|
||
data-pmc-id="PMC2958481"
|
||
data-figure-id="fig4">
|
||
<img class="figure-thumb" itemprop="thumbnail"
|
||
id="article-image-3"
|
||
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/5a36/2958481/4f9f83b28dc0/JCB_201004108_GS_Fig4.gif"
|
||
alt="Figure 4." />
|
||
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|
||
|
||
<meta itemprop="width" itemtype="http://schema.org/ImageObject" content="664">
|
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<meta itemprop="height" itemtype="http://schema.org/ImageObject" content="483">
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|
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|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 4. </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b> Gli proteins are required to… </b></div>
|
||
|
||
|
||
|
||
|
||
</div>
|
||
</div>
|
||
|
||
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
|
||
|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 4. </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b> Gli proteins are required to localize SuFu to cilia, but Gli proteins can… </b></div>
|
||
|
||
|
||
|
||
|
||
</div>
|
||
|
||
<figcaption id="figure-caption-3" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
|
||
|
||
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||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
Figure 4.
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b>Gli proteins are required to localize SuFu to cilia, but Gli proteins can localize to cilia in the absence of SuFu.</b> (A) Wild-type and Gli2<sup>−/−</sup> Gli3<sup>−/−</sup> MEFs were incubated with or without Shh. SuFu does not localize to cilia with or without Shh stimulation in Gli2<sup>−/−</sup> Gli3<sup>−/−</sup> MEFs, whereas Smo recruitment is normal. (B) Cilia counts for a time course of ciliary recruitment of Smo, SuFu, and Gli in Gli2<sup>−/−</sup> Gli3<sup>−/−</sup> MEFs stimulated with Shh. (C) SuFu<sup>+/−</sup> and SuFu<sup>−/−</sup> MEFs were stimulated or not with Shh. Endogenous Gli proteins do not localize to cilia with or without Shh stimulation in the absence of SuFu. Recruitment of Smo is normal. (D) Immunoblot of SuFu<sup>−/−</sup> and SuFu<sup>+/−</sup> MEFs stably expressing Gli1HA and treated with the proteasome inhibitor bortezomib. Proteasome inhibition allows SuFu<sup>−/−</sup> cells to accumulate Gli1HA to levels similar to those in the control SuFu<sup>+/−</sup> cells. (E) Stably expressed Gli1HA localizes to ciliary tips in SuFu<sup>−/−</sup> MEFs stimulated with SAG in the presence of bortezomib. Percentages shown indicate corresponding ciliary counts. Bars, 2 µm.</div>
|
||
|
||
|
||
|
||
|
||
</figcaption>
|
||
|
||
|
||
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|
||
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||
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|
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data-label-slug="figure-5">
|
||
<a class="figure-link"
|
||
href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/5a36/2958481/3ade18a1b112/JCB_201004108_GS_Fig5.jpg"
|
||
itemprop="contentUrl"
|
||
aria-describedby="figure-caption-4"
|
||
role="button"
|
||
data-image-width="629"
|
||
data-image-height="784"
|
||
data-image-alt="Figure 5."
|
||
data-pmc-id="PMC2958481"
|
||
data-figure-id="fig5">
|
||
<img class="figure-thumb" itemprop="thumbnail"
|
||
id="article-image-4"
|
||
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/5a36/2958481/d1338e3499fc/JCB_201004108_GS_Fig5.gif"
|
||
alt="Figure 5." />
|
||
</a>
|
||
|
||
<meta itemprop="width" itemtype="http://schema.org/ImageObject" content="629">
|
||
<meta itemprop="height" itemtype="http://schema.org/ImageObject" content="784">
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||
|
||
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|
||
|
||
<div class="caption-wrap">
|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 5. </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b> Biochemical evidence that Hh pathway… </b></div>
|
||
|
||
|
||
|
||
|
||
</div>
|
||
</div>
|
||
|
||
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
|
||
|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 5. </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b> Biochemical evidence that Hh pathway activation causes rapid dissociation of endogenous SuFu–Gli complexes.… </b></div>
|
||
|
||
|
||
|
||
|
||
</div>
|
||
|
||
<figcaption id="figure-caption-4" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
|
||
|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
Figure 5.
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b>Biochemical evidence that Hh pathway activation causes rapid dissociation of endogenous SuFu–Gli complexes.</b> (A–I) Endogenous SuFu–Gli complexes were analyzed by sucrose gradient centrifugation (A–G) and immunoprecipitation (H and I). (A) In untreated NIH-3T3 cells, the majority of endogenous SuFu (54 kD) exists as a monomer of similar size as the kinase GSK3-β (47 kD). A small fraction of SuFu from untreated cells forms a higher molecular mass complex (top, black lines), the level of which quickly drops in cells treated with Shh for 1 h (middle), an effect completely blocked if Smo is inhibited with 200 nM SANT-1 (bottom). The position in the gradient of two size markers run in parallel is shown below the Western blots (aldolase: molecular mass, 158 kD; Stokes radius, 48.1 Å; catalase: molecular mass, 232 kD; Stokes radius, 52.2 Å). (B) In Gli2<sup>−/−</sup> Gli3<sup>−/−</sup> MEFs, only the monomeric SuFu peak is seen by sucrose gradient centrifugation. Hh stimulation of Gli2<sup>−/−</sup> Gli3<sup>−/−</sup> MEFs does not change the size of the SuFu peak, although Smo is recruited to the cilia normally in these cells. (C) As in A, but cells were stimulated or not with SAG, and sucrose gradient fractions were immunoblotted for endogenous SuFu, GSK3, and Gli3-FL. The higher molecular mass SuFu peak overlaps with endogenous Gli3-FL in unstimulated cells. Acute Hh pathway stimulation causes the simultaneous disappearance of the overlapping, higher molecular mass SuFu and Gli3-FL peaks. (D) To prevent dissociation of SuFu from Gli, a direct fusion of Gli1 to SuFu was generated. NIH-3T3 cells stably expressing this Gli1–SuFu fusion were stimulated or not with SAG. The apparent size of the Gli1–SuFu fusion peak does not change upon Hh pathway activation. (E) Treatment of NIH-3T3 cells with SAG causes complete disappearance of the SuFu–Gli complex, which is not reversed by inhibition of the proteasome with bortezomib. In contrast, activation of PKA with FSK completely blocks SuFu–Gli dissociation induced by SAG stimulation. (F) Quantification of the experiment in E. The amount of SuFu in each fraction was measured relative to the amount of SuFu in the input lane. The first fraction represents the top of the sucrose gradient. (G) Mouse SuFu expressed in <i>Xenopus</i> embryos shows the same size distribution as endogenous SuFu in mammalian cultured cells, suggesting that SuFu forms a similar complex with endogenous Gli proteins in <i>Xenopus</i> embryos. (H) NIH-3T3 cells were incubated with or without SAG followed by immunoprecipitation (IP) with anti-SuFu antibodies. The level of Gli3-FL is similar in SAG-treated and untreated cells (left). Gli3-FL coimmunoprecipitates with SuFu only in untreated cells but not in SAG-stimulated cells (right), indicating that acute Hh pathway activation dissociates endogenous Gli3-FL from SuFu. (I) NIH-3T3 cells were incubated with control media, SAG, SAG and bortezomib, and SAG and FSK followed by immunoprecipitation with anti–Gli3-FL antibodies. Gli2<sup>−/−</sup> Gli3<sup>−/−</sup> MEFs were used as negative control (lanes 1 and 6). Endogenous SuFu does not coimmunoprecipitate with Gli3-FL in cells stimulated with SAG, although levels of Gli3-FL decrease only slightly. Proteasome inhibition by bortezomib (sufficient to abolish any decrease in the level of Gli3-FL) does not block dissociation of endogenous SuFu from Gli3-FL. In contrast, SAG-induced dissociation of SuFu from Gli3-FL is completely blocked by FSK.</div>
|
||
|
||
|
||
|
||
|
||
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|
||
|
||
|
||
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|
||
|
||
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|
||
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|
||
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|
||
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|
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|
||
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|
||
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|
||
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|
||
data-figure-id="fig6">
|
||
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|
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|
||
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/5a36/2958481/a3dcc125e128/JCB_201004108_GS_Fig6.gif"
|
||
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|
||
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|
||
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||
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|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 6. </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b> A model for activation of… </b></div>
|
||
|
||
|
||
|
||
|
||
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|
||
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|
||
|
||
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
|
||
|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 6. </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b> A model for activation of Gli proteins during vertebrate Hh signaling. </b> In the…</div>
|
||
|
||
|
||
|
||
|
||
</div>
|
||
|
||
<figcaption id="figure-caption-5" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
|
||
|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
Figure 6.
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><b>A model for activation of Gli proteins during vertebrate Hh signaling.</b> In the resting state of the Hh pathway (left), SuFu forms inactive complexes with Gli2 and Gli3-FL, which are sequestered in the cytoplasm. Without Hh stimulation, SuFu–Gli complexes traffic to the primary cilium at a low level independently of Smo; this basal ciliary trafficking is antagonized by PKA. Hh pathway stimulation (right) leads to the translocation of active Smo to the cilium, which, in turn, recruits SuFu–Gli complexes. Active Smo at cilia causes the dissociation of SuFu from Gli. Monomeric SuFu and Gli leave the cilium followed by Gli nuclear translocation and activation of the transcriptional program of the Hh pathway. PKA antagonizes Hh signaling by blocking ciliary localization of SuFu–Gli complexes, thus preventing coupling between active Smo and dissociation of SuFu–Gli complexes.</div>
|
||
|
||
|
||
|
||
|
||
</figcaption>
|
||
|
||
|
||
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|
||
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||
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Alcedo J., Ayzenzon M., Von Ohlen T., Noll M., Hooper J.E. 1996. The Drosophila smoothened gene encodes a seven-pass membrane protein, a putative receptor for the hedgehog signal. Cell. 86:221–232 10.1016/S0092-8674(00)80094-X
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<div id="substances" class="substances keywords-section">
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Substances
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Hedgehog Proteins
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|
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ref="linksrc=search_in_pubmed_keyword_link&keyword="Kruppel-Like Transcription Factors"[nm]"
|
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Search in PubMed
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</a></li><li><a class="search-in-mesh-link dropdown-block-link"
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|
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|
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Search in MeSH
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</a></li><li><a class="add-to-search-link dropdown-block-link"
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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|
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|
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|
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|
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Receptors, G-Protein-Coupled
|
||
</button><div id="keyword-actions-substances-3"
|
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class="keyword-actions-dropdown dropdown dropdown-container"
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aria-label="Dropdown menu for keyword Receptors, G-Protein-Coupled"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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Add to Search
|
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Repressor Proteins"
|
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tabindex="0"
|
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|
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|
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|
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Repressor Proteins
|
||
</button><div id="keyword-actions-substances-4"
|
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class="keyword-actions-dropdown dropdown dropdown-container"
|
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hidden
|
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aria-label="Dropdown menu for keyword Repressor Proteins"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
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href="/?term=%22Repressor+Proteins%22%5Bnm%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&keyword="Repressor Proteins"[nm]"
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Search in PubMed
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|
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Smo protein, mouse"
|
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tabindex="0"
|
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|
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|
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data-pinger-ignore>
|
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Smo protein, mouse
|
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</button><div id="keyword-actions-substances-5"
|
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class="keyword-actions-dropdown dropdown dropdown-container"
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hidden
|
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aria-label="Dropdown menu for keyword Smo protein, mouse"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Smoothened Receptor"
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tabindex="0"
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|
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Smoothened Receptor
|
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</button><div id="keyword-actions-substances-6"
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hidden
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aria-label="Dropdown menu for keyword Smoothened Receptor"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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data-search-term=""Smoothened Receptor"[nm]"
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data-ga-action="add_to_search"
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Add to Search
|
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</a></li></ul></div></div></div></li><li><div class="keyword-actions dropdown-block"><button class="keyword-actions-trigger trigger keyword-link"
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aria-label="Toggle dropdown menu for keyword Sufu protein, mouse"
|
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tabindex="0"
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|
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data-pinger-ignore>
|
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Sufu protein, mouse
|
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</button><div id="keyword-actions-substances-7"
|
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class="keyword-actions-dropdown dropdown dropdown-container"
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hidden
|
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aria-label="Dropdown menu for keyword Sufu protein, mouse"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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Search in PubMed
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Search in MeSH
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data-search-term=""Sufu protein, mouse"[nm]"
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data-ga-category="search"
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Add to Search
|
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</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 Zinc Finger Protein GLI1"
|
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tabindex="0"
|
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aria-controls="keyword-actions-substances-8"
|
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aria-expanded="false"
|
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data-pinger-ignore>
|
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Zinc Finger Protein GLI1
|
||
</button><div id="keyword-actions-substances-8"
|
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class="keyword-actions-dropdown dropdown dropdown-container"
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hidden
|
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aria-label="Dropdown menu for keyword Zinc Finger Protein GLI1"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
|
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|
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ref="linksrc=search_in_pubmed_keyword_link&keyword="Zinc Finger Protein GLI1"[nm]"
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Search in PubMed
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|
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Search in MeSH
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</a></li><li><a class="add-to-search-link dropdown-block-link"
|
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Add to Search
|
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</a></li></ul></div></div></div></li></ul>
|
||
</div>
|
||
|
||
|
||
|
||
|
||
|
||
|
||
|
||
|
||
|
||
|
||
|
||
|
||
<div id="related-links" class="related-links">
|
||
<h2 class="title">
|
||
Related information
|
||
</h2>
|
||
|
||
<ul class="related-links-list"><li><a title="PubMed links associated with Books" href="https://www.ncbi.nlm.nih.gov/books?linkname=pubmed_books_refs&from_uid=20956384"
|
||
ref="log$=recordlinks&logdbfrom=pubmed&ncbi_uid=20956384&ordinalpos=1&linkpos=1">
|
||
Cited in Books
|
||
</a></li><li><a title="Link to related Genes" href="https://www.ncbi.nlm.nih.gov/gene?linkname=pubmed_gene&from_uid=20956384"
|
||
ref="log$=recordlinks&logdbfrom=pubmed&ncbi_uid=20956384&ordinalpos=1&linkpos=2">
|
||
Gene
|
||
</a></li><li><a title="Link to Gene for the GeneRIF subcategory" href="https://www.ncbi.nlm.nih.gov/gene?linkname=pubmed_gene_rif&from_uid=20956384"
|
||
ref="log$=recordlinks&logdbfrom=pubmed&ncbi_uid=20956384&ordinalpos=1&linkpos=3">
|
||
Gene (GeneRIF)
|
||
</a></li><li><a title="Related information in MedGen" href="https://www.ncbi.nlm.nih.gov/medgen?linkname=pubmed_medgen&from_uid=20956384"
|
||
ref="log$=recordlinks&logdbfrom=pubmed&ncbi_uid=20956384&ordinalpos=1&linkpos=4">
|
||
MedGen
|
||
</a></li><li><a title="Related records in MedGen based on citations in GeneReviews and Medical Genetics Summaries" href="https://www.ncbi.nlm.nih.gov/medgen?linkname=pubmed_medgen_bookshelf_cited&from_uid=20956384"
|
||
ref="log$=recordlinks&logdbfrom=pubmed&ncbi_uid=20956384&ordinalpos=1&linkpos=5">
|
||
MedGen (Bookshelf cited)
|
||
</a></li><li><a title="Link to Protein RefSeqs" href="https://www.ncbi.nlm.nih.gov/protein?linkname=pubmed_protein_refseq&from_uid=20956384"
|
||
ref="log$=recordlinks&logdbfrom=pubmed&ncbi_uid=20956384&ordinalpos=1&linkpos=6">
|
||
Protein (RefSeq)
|
||
</a></li></ul>
|
||
</div>
|
||
|
||
|
||
|
||
|
||
|
||
<div id="grants" class="grants">
|
||
<h2 class="title">
|
||
Grants and funding
|
||
</h2>
|
||
<div class="grants-list"><ul id="grants-head-list"><li class="grant-item"><a
|
||
title="All articles for grant R01 GM092924/GM/NIGMS NIH HHS/United States"
|
||
href="/?term=R01+GM092924%2FGM%2FNIGMS+NIH+HHS%2FUnited+States%5BGrants+and+Funding%5D&sort=date&sort_order=desc"
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>
|
||
R01 GM092924/GM/NIGMS NIH HHS/United States
|
||
</a></li></ul></div>
|
||
</div>
|
||
|
||
|
||
|
||
|
||
<div id="linkout" class="linkout">
|
||
<h2 class="title">
|
||
LinkOut - more resources
|
||
</h2>
|
||
|
||
<ul class="linkout-list"><li><h3 class="linkout-category">Full Text Sources</h3><ul class="linkout-category-links"><li><a
|
||
href="https://europepmc.org/abstract/MED/20956384"
|
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target="_blank"
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Europe PubMed Central
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</a></li><li><a
|
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href="https://pmc.ncbi.nlm.nih.gov/articles/pmid/20956384/"
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target="_blank"
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PubMed Central
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</a></li><li><a
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Silverchair Information Systems
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</a></li></ul></li><li><h3 class="linkout-category">Other Literature Sources</h3><ul class="linkout-category-links"><li><a
|
||
href="https://facultyopinions.com/pubmed/20956384"
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||
target="_blank"
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</a></li></ul></li><li><h3 class="linkout-category">Molecular Biology Databases</h3><ul class="linkout-category-links"><li><a
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target="_blank"
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Mouse Genome Informatics (MGI)
|
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</a></li></ul></li><li><h3 class="linkout-category">Miscellaneous</h3><ul class="linkout-category-links"><li><a
|
||
href="https://assays.cancer.gov/non-CPTAC-3385"
|
||
target="_blank"
|
||
rel="noopener"
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|
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</a></li></ul></li></ul>
|
||
</div>
|
||
|
||
|
||
</main>
|
||
|
||
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