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||
</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%2F21316585%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%2F21316585%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="T2qoPImWZC8fSfFbtwcfUgxqEaTyllcDaPatsyPizdVJMGL16PPhoo34ceBKeWKX">
|
||
</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="T2qoPImWZC8fSfFbtwcfUgxqEaTyllcDaPatsyPizdVJMGL16PPhoo34ceBKeWKX">
|
||
|
||
|
||
|
||
<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="T2qoPImWZC8fSfFbtwcfUgxqEaTyllcDaPatsyPizdVJMGL16PPhoo34ceBKeWKX">
|
||
|
||
<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: 21316585 - PubMed</span>
|
||
<input type="hidden" name="email-subject" value="1 selected item: 21316585"/>
|
||
<input type="hidden" name="email-subject-hash" value="b64cd7852a48410f22e58e44d509ed119294fca76ee48658effa992f175b773a"/>
|
||
</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="T2qoPImWZC8fSfFbtwcfUgxqEaTyllcDaPatsyPizdVJMGL16PPhoo34ceBKeWKX">
|
||
|
||
|
||
|
||
<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="T2qoPImWZC8fSfFbtwcfUgxqEaTyllcDaPatsyPizdVJMGL16PPhoo34ceBKeWKX">
|
||
|
||
|
||
|
||
<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="T2qoPImWZC8fSfFbtwcfUgxqEaTyllcDaPatsyPizdVJMGL16PPhoo34ceBKeWKX">
|
||
|
||
<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="T2qoPImWZC8fSfFbtwcfUgxqEaTyllcDaPatsyPizdVJMGL16PPhoo34ceBKeWKX">
|
||
|
||
|
||
|
||
<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="T2qoPImWZC8fSfFbtwcfUgxqEaTyllcDaPatsyPizdVJMGL16PPhoo34ceBKeWKX">
|
||
|
||
<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="21316585">
|
||
<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://linkinghub.elsevier.com/retrieve/pii/S1534-5807(11)00002-5"
|
||
target="_blank"
|
||
rel="noopener"
|
||
|
||
ref="linksrc=fulltextorjournal_fulltext&is_pmc=False&PrId=3048&itool=Abstract-def&log$=linkouticon&uid=21316585&db=pubmed&nlmid=101120028"
|
||
title="See full text options at Elsevier Science"
|
||
|
||
data-ga-category="full_text"
|
||
data-ga-action="Elsevier Science"
|
||
data-ga-label="21316585"
|
||
><img src="https://cdn.ncbi.nlm.nih.gov/corehtml/query/egifs/https:--linkinghub.elsevier.com-ihub-images-celloa.png" alt="Elsevier Science full text link"><span class="text">
|
||
|
||
Elsevier Science
|
||
|
||
</span></a><a class="link-item
|
||
pmc
|
||
"
|
||
href="https://pmc.ncbi.nlm.nih.gov/articles/pmid/21316585/"
|
||
target="_blank"
|
||
rel="noopener"
|
||
|
||
ref="linksrc=fulltextorjournal_fulltext&is_pmc=True&PrId=3494&itool=Abstract-def&log$=linkouticon&uid=21316585&db=pubmed&nlmid=101120028"
|
||
title="Free full text at PubMed Central"
|
||
|
||
data-ga-category="full_text"
|
||
data-ga-action="PMC"
|
||
data-ga-label="21316585"
|
||
><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="/21316585/citations/"
|
||
data-citation-style="nlm"
|
||
data-pubmed-format-link="/21316585/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="21316585" 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%2F21316585%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="T2qoPImWZC8fSfFbtwcfUgxqEaTyllcDaPatsyPizdVJMGL16PPhoo34ceBKeWKX"><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
|
||
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="sidebar-actions-display-format"><option value="abstract">Abstract</option><option value="pubmed">PubMed</option><option value="pmid">PMID</option></select></div></div></div></div></div></div>
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</a></li></ul></div></div></div><span class="period">. </span><span class="cit">2011 Feb 15;20(2):163-76.</span>
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Wnt signaling gradients establish planar cell polarity by inducing Vangl2 phosphorylation through Ror2
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Wnt signaling gradients establish planar cell polarity by inducing Vangl2 phosphorylation through Ror2
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It is fundamentally important that signaling gradients provide positional information to govern morphogenesis of multicellular organisms. Morphogen gradients can generate different cell types in specific spatial order at distinct threshold concentrations. However, it is largely unknown whether and how signaling gradients also control cell polarities by acting as global cues. Here, we show that Wnt signaling gradient provides directional information to a field of cells. Vangl2, a core component in planar cell polarity, forms Wnt-induced receptor complex with Ror2 to sense Wnt dosages. Wnts dose-dependently induce Vangl2 phosphorylation of serine/threonine residues and Vangl2 activities depend on its levels of phosphorylation. In the limb bud, Wnt5a signaling gradient controls limb elongation by establishing PCP in chondrocytes along the proximal-distal axis through regulating Vangl2 phosphorylation. Our studies have provided new insight to Robinow syndrome, Brachydactyly Type B1, and spinal bifida which are caused by mutations in human ROR2, WNT5A, or VANGL.
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data-image-alt="Figure 1"
|
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data-pmc-id="PMC3062198"
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data-figure-id="F1">
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<img class="figure-thumb" itemprop="thumbnail"
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src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/8fba/3062198/5d05521d581f/nihms-263516-f0001.gif"
|
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alt="Figure 1" />
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<strong class="figure-label">
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<p> Figure 1. Ror2 and Vangl2 mediate Wnt5a… </p>
|
||
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<strong class="figure-label">
|
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<p> Figure 1. Ror2 and Vangl2 mediate Wnt5a signaling in embryonic development </p>
|
||
</strong>
|
||
|
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|
||
<div class="figure-caption-contents"><p> (A) The limb length… </p></div>
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<strong class="figure-label">
|
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Figure 1. Ror2 and Vangl2 mediate Wnt5a signaling in embryonic development
|
||
</strong>
|
||
|
||
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|
||
<div class="figure-caption-contents">(A) The limb length from the elbow to the distal tip is indicated by a line and quantified in the right column (two-tail <i>t</i>-test, all *<i>p</i> values <0.01). Error bars are ± SD, n=3 (B) Sections of distal limbs showing chondrocyte organization in the newly formed cartilage (schematized in the right panel). Wheat germ agglutinin (red) stains chondrocyte extracellular matrix; DAPI (blue) stains the nucleus. (C) Statistical analysis of length-to-width ratio of chondrocytes shown in Figure 1B (two-tail <i>t</i>-test, all *<i>p</i> values <0.01). Error bars are ± SD, n>100. L, length; W, width. (D) Neural tube was open (red arrows) in >50% of the <i>Ror2</i><sup>−/−</sup>; <i>Vangl2</i><sup><i>Lp</i>/+</sup> embryo. (E) The <i>Ror2</i><sup>−/−</sup>; <i>Vangl2</i><sup>−/−</sup> and the <i>Wnt5a</i><sup>−/−</sup> embryos exhibited identical phenotypes of loss of distal digits (circles), shortened frontonasal processes (arrows) and tails (arrowheads). The limb was enlarged in the lower panel. (F) Skeletal preparations stained by alcian blue and alizarin red of forelimbs. See also Figure S1.</div>
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|
||
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data-figure-id="F2">
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<img class="figure-thumb" itemprop="thumbnail"
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src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/8fba/3062198/482b489254fc/nihms-263516-f0002.gif"
|
||
alt="Figure 2" />
|
||
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|
||
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||
<strong class="figure-label">
|
||
<p> Figure 2. Ror2 and Vangl2 together mediate… </p>
|
||
</strong>
|
||
|
||
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||
|
||
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||
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||
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|
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|
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|
||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 2. Ror2 and Vangl2 together mediate Wnt5a inhibition of the Wnt/β-catenin signaling </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><p> (A) <i> Sox9… </i> </p></div>
|
||
|
||
|
||
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||
|
||
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|
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|
||
<figcaption id="figure-caption-1" 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 2. Ror2 and Vangl2 together mediate Wnt5a inhibition of the Wnt/β-catenin signaling
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents">(A) <i>Sox9</i> expression detected by whole mount <i>in situ</i> hybridization. Arrows indicate lack of chondrocyte differentiation in the distal limb mesenchymal cells of the <i>Ror2</i><sup>−/−</sup>; <i>Vangl2</i><sup>−/−</sup> and <i>Wnt5a</i><sup>−/−</sup> embryos. (B) X-gal staining of limb buds showed endogenous staining in the joint (black arrows) and ectopic staining in the distal limb of the <i>Ror2</i><sup>−/−</sup>; <i>Vangl2</i><sup>−/−</sup>; <i>TopGal</i> embryo (red arrows). (C) The Wnt/β-catenin signaling activity was measured by the TOPFLASH luciferase assay in MEF cells. The ability of Wnt5a to inhibit Wnt3a-induced Wnt/β-catenin activity in wild type MEFs (two-tail <i>t</i> test *<i>p</i>=0.0013) is impaired in the <i>Ror2</i><sup>−/−</sup> and <i>Ror2</i><sup>−/−</sup>; <i>Vangl2</i><sup>−/−</sup> MEFs (two-tail <i>t</i> test, *<i>p</i>=0.0015). Error bars are ± SD, n=3. The control and Wnt3a conditioned medium was mixed with that of Wnt5a at a 1:1 ratio. See also Figure S2.</div>
|
||
|
||
|
||
|
||
|
||
</figcaption>
|
||
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||
|
||
</figure>
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||
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|
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href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/8fba/3062198/243ad64a2719/nihms-263516-f0003.jpg"
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||
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|
||
data-image-alt="Figure 3"
|
||
data-pmc-id="PMC3062198"
|
||
data-figure-id="F3">
|
||
<img class="figure-thumb" itemprop="thumbnail"
|
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|
||
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/8fba/3062198/e137daa39998/nihms-263516-f0003.gif"
|
||
alt="Figure 3" />
|
||
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||
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|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 3. Wnt5a induces Ror2 and Vangl2… </p>
|
||
</strong>
|
||
|
||
|
||
|
||
|
||
|
||
|
||
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|
||
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|
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||
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|
||
|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 3. Wnt5a induces Ror2 and Vangl2 complex formation </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><p> (A) Fluorescent immunohistochemistry of Ror2 (red)… </p></div>
|
||
|
||
|
||
|
||
|
||
</div>
|
||
|
||
<figcaption id="figure-caption-2" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
|
||
|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
Figure 3. Wnt5a induces Ror2 and Vangl2 complex formation
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents">(A) Fluorescent immunohistochemistry of Ror2 (red) and Vangl2 (green) (viewed by projected Z-stack) showing asymmetrical Vangl2 localization at E12.5, not E11.5 (arrows). Selected limb fields are schematized in the right panel. (B) Co-IP of Ror2 and Vangl2 in HEK293T cells. Wnt5a strongly increased their interaction. (C) MDCK cells expressing YFPVangl2 and Ror2-CFP were assayed for fluorescence resonance energy transfer efficiency (N-FRET) with (0.13 s.d. ± 0.02) or without Wnt5a (0.11 s.d. ± 0.05), Kolmogorov-Smirnov two-sample test (two-tail): <i>p</i>=0.029. (D) A <i>Loop-tail</i> mutant Vangl2<sup>Lp</sup> (S464N) protein failed to form complex with Ror2 in HEK293T cells, even in the presence of Wnt5a. See also Figure S3.</div>
|
||
|
||
|
||
|
||
|
||
</figcaption>
|
||
|
||
|
||
</figure>
|
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|
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|
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|
||
data-image-alt="Figure 4"
|
||
data-pmc-id="PMC3062198"
|
||
data-figure-id="F4">
|
||
<img class="figure-thumb" itemprop="thumbnail"
|
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|
||
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/8fba/3062198/d2ab6af75ad8/nihms-263516-f0004.gif"
|
||
alt="Figure 4" />
|
||
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|
||
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|
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|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 4. Wnt5a is required for Vangl2… </p>
|
||
</strong>
|
||
|
||
|
||
|
||
|
||
|
||
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||
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|
||
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|
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|
||
|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 4. Wnt5a is required for Vangl2 asymmetrical localization </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><p> (A) Digit chondrocytes of E12.5 embryos… </p></div>
|
||
|
||
|
||
|
||
|
||
</div>
|
||
|
||
<figcaption id="figure-caption-3" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
|
||
|
||
|
||
|
||
|
||
|
||
|
||
<strong class="figure-label">
|
||
Figure 4. Wnt5a is required for Vangl2 asymmetrical localization
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents">(A) Digit chondrocytes of E12.5 embryos is shown by Sox9 expression (red) (upper panel). The selected regions (boxed) are enlarged and shown with Vangl2 (green) and β-catenin (red) co-staining (projected Z-stacks) (middle panel). The boxed regions are shown in lower panel with separate staining of Vangl2 and β-catenin. Vangl2 asymmetrical localization (arrow) is completely lost in the <i>Wnt5a</i><sup>−/−</sup> limb. In the <i>Ror2</i><sup>−/−</sup> mutant limb, Vangl2 asymmetrical localization was lost in the more mature cartilage (see Fig. S4A), but still detectable in the distal limb bud with reduced intensity (arrow). Vangl2 is still localized to the cell membrane in <i>Wnt5a</i><sup>−/−</sup> mutants (arrowhead). D, distal; P, proximal. (B) Projected Z-stack pictures of Vangl2 staining (green) from distal (D), middle (M) or proximal (P) parts of E12.5 wild type or <i>Ror2</i><sup>−/−</sup> embryonic distal limbs (schematized in left panel). The number of cells with Vangl2 asymmetrical localization is progressively reduced from distal to proximal parts and much reduced in the <i>Ror2</i><sup>−/−</sup> mutant. Quantified results were shown in lower panel. Error bars are ± SD, n=2. See also Figure S4.</div>
|
||
|
||
|
||
|
||
|
||
</figcaption>
|
||
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||
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||
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|
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|
||
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|
||
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|
||
<img class="figure-thumb" itemprop="thumbnail"
|
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|
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src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/8fba/3062198/82c0a462d540/nihms-263516-f0005.gif"
|
||
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|
||
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|
||
|
||
|
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||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 5. Wnts act through Ror2 to… </p>
|
||
</strong>
|
||
|
||
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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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||
|
||
<strong class="figure-label">
|
||
<p> Figure 5. Wnts act through Ror2 to induce Vangl2 phosphorylation </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><p> (A) Mobility shift of Vangl2… </p></div>
|
||
|
||
|
||
|
||
|
||
</div>
|
||
|
||
<figcaption id="figure-caption-4" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
|
||
|
||
|
||
|
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|
||
|
||
|
||
<strong class="figure-label">
|
||
Figure 5. Wnts act through Ror2 to induce Vangl2 phosphorylation
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents">(A) Mobility shift of Vangl2 expressed in CHO cells on SDS-PAGE gel was enhanced by Wnt5a, but abolished after treatment with calf intestine phosphatase (CIP). (B) CHO cells were transfected to express different mouse <i>Wnts</i>, <i>Ror2</i> and <i>Vangl2</i> in the indicated combination. Vangl2 phosphorylation was analyzed by immunoblot. Arrows indicate hyperphosphorylated (upper), hypophosphorylated (middle) and unphosphorylated (lower) Vangl2 forms. (C) MEF cells were treated with recombinant Wnt5a protein for 2 hours and Vangl2 phosphorylation was analyzed. (D) Mouse Vangl2 with two phosphorylation clusters. Founder sites of Cluster I and II are underlined and shown in bold. Mutant Vangl2 with various serine/threonine (S/T) to alanine (A) variants are shown. (E) S84 and S82 are founder sites for Cluster I phosphorylation. Phosphorylation of Cluster II sites still occurred when all Cluster I sites were mutated. (F) S sites in Cluster II of <i>Vangl2</i> were individually mutated to A. Wnt5a-induced Vangl2 phosphorylation was progressively more reduced from S20A to S5A. (G) Mutating all three founder sites, S5, 82, 84, is sufficient to completely abolish both basal level and Wnt5a-induced phosphorylation in CHO cells. (H) Vangl2 phosphorylation is induced by CK1δ not CK1ε and partially blocked by CKI inhibitor D4476 (100uM) in CHO cells. See also Figure S5.</div>
|
||
|
||
|
||
|
||
|
||
</figcaption>
|
||
|
||
|
||
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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-image-alt="Figure 6"
|
||
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|
||
data-figure-id="F6">
|
||
<img class="figure-thumb" itemprop="thumbnail"
|
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|
||
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/8fba/3062198/25a06b40ec22/nihms-263516-f0006.gif"
|
||
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|
||
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|
||
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|
||
|
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|
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|
||
|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 6. Wnt-induced Vangl2 phosphorylation is required… </p>
|
||
</strong>
|
||
|
||
|
||
|
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|
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|
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|
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|
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|
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|
||
|
||
<strong class="figure-label">
|
||
<p> Figure 6. Wnt-induced Vangl2 phosphorylation is required for its function </p>
|
||
</strong>
|
||
|
||
|
||
|
||
<div class="figure-caption-contents"><p> (A) CHO cells expressing wild… </p></div>
|
||
|
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Figure 6. Wnt-induced Vangl2 phosphorylation is required for its function
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</strong>
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<div class="figure-caption-contents">(A) CHO cells expressing wild type or the indicated mutant <i>Wnt5a</i> were cocultured with <i>Vangl2</i>- and <i>Ror2</i>- expressing CHO cells at a ratio of 4:1. (B) Ror2-BDB (W749X) and Ror2-RS (Q502X) mutants failed to enhance Wnt5a-induced Vangl2 phosphorylation in <i>Ror2</i><sup>−/−</sup> MEFs. The kinase-dead Ror2 (Ror2-KD) showed similar activity as the wild type Ror2. (C) Effects of <i>Vangl2 Lp</i> mutations (D255E and S464N) and the control S464A mutation on Vangl2 phosphorylation were analyzed by immunoblot in CHO cells. (D) Analysis of endogenous Vangl2 protein levels and phosphorylation by immunoblot in the brain of the <i>Vangl2<sup>Lp/Lp</sup></i> embryo. (E) Analysis of wild type Vangl2 phosphorylation in CHO cells when co-expressed with the D255E or S464N <i>Lp</i> mutant <i>Vangl2</i>. (F) Wild type, S84A,S5~17A∷S76~84A (all phospho-mutant) or S5,82,84E (phospho-mimicking) mouse <i>Vangl2</i> mRNA was injected to offsprings from the zebrafish <i>tri</i><sup><i>m209</i>/+</sup> × <i>tri</i><sup><i>m209</i>/+</sup> matings at different doses. The phenotypes of injected zebrafish embryos were classified into four groups (group A, B, C and D) as shown by the representative pictures. (G) Statistical analysis of results of wild type (a, d), S84A (b, e), S5~17A∷S76~84A (c, f) and S5,82,84E (g) mouse <i>Vangl2</i> mRNA injection. Numbers of injected zebrafish embryos of the indicated genotypes are indicated above the bars. The percentage of rescued <i>tri</i><sup><i>m209/m209</i></sup> emrbyos (Group A and B) was shown on the right of each panel. (H) Analysis of Vangl2 proteins in the injected fish embryos. HA-tagged wild type and mutant Vangl2 proteins synthesized from different amounts of injected mRNA in wild type embryos were shown in left panel. Quantified results by densitometer were shown in the right panel. Error bars are ± SD, n=3. See also Figure S6.</div>
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data-figure-id="F7">
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<p> Figure 7. Wnt5a gradient controls pattern formation… </p>
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<p> Figure 7. Wnt5a gradient controls pattern formation by regulating Vangl2 phosphorylation in a dose-dependent manner </p>
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<strong class="figure-label">
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Figure 7. Wnt5a gradient controls pattern formation by regulating Vangl2 phosphorylation in a dose-dependent manner
|
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</strong>
|
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|
||
<div class="figure-caption-contents">(A) The E11.5 mouse limb bud was separated into distal, middle and proximal parts (schematics) for immunoblot analysis of Vangl2 phosphorylation. (B) Distal limb bud tissues of E11.5 wild type, <i>Ror2</i><sup>−/−</sup> and <i>Wnt5a</i><sup>−/−</sup> embryos, as shown by the schematic diagram, were subject to immunoblot analysis using Vangl2 or phospho-specific (pS82/84) Vangl2 antibodies. The Vangl2 in the <i>Ror2</i><sup>−/−</sup> and <i>Wnt5a</i><sup>−/−</sup> mutant limb was shifted to hypophosphorylated forms (lower arrows) without altering S84 and S82 phosphorylation. (C) <i>Wnt5a</i>-expressing CHO cells were cocultured with CHO cells expressing <i>Vangl2+Ror2</i> at increasing ratio. The last boxed lane was extracts from <i>Wnt5a</i>, <i>Ror2</i> and <i>Vangl2</i> co-transfected CHO cells that showed the most robust induction of Vangl2 phosphorylation. (D) Recombinant Wnt5a proteins induced Vangl2 phosphorylation in a dose-dependent manner in CHO cells after 2-hour treatment. Vangl2 phosphorylation was saturated by 500ng/ml of Wnt5a. (E, F) A model of a Wnt5a gradient controlling P-D limb elongation by providing a global directional cue. (E) Wnt5a induces Ror2 and Vangl2 complex formation and as a result, different numbers of S and T residues of Vangl2 in the two clusters were phosphorylated depending on the Wnt5a dosages. (F) Wnt5a is expressed in a gradient (orange) in the developing limb bud and this Wnt5a gradient is translated into an activity gradient of Vangl2 by inducing different levels of Vangl2 phosphorylation (blue). The Vangl2 activity gradient then induces asymmetrical Vangl2 localization (blue) and downstream polarized events. See also Figure S7.</div>
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A new piece to the unsolved planar cell polarity puzzle.
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<div class="docsum-citation full-citation">
|
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<span class="docsum-authors full-authors">Grumolato L, Weber U, Mlodzik M.</span>
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<span class="docsum-authors short-authors">Grumolato L, et al.</span>
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<span class="docsum-journal-citation full-journal-citation">Dev Cell. 2011 Feb 15;20(2):146-7. doi: 10.1016/j.devcel.2011.01.013.</span>
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<span class="docsum-journal-citation short-journal-citation">Dev Cell. 2011.</span>
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<span class="citation-part">PMID: <span class="docsum-pmid">21316583</span></span>
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Afzal AR, Rajab A, Fenske CD, Oldridge M, Elanko N, Ternes-Pereira E, Tuysuz B, Murday VA, Patton MA, Wilkie AO, et al. Recessive Robinow syndrome, allelic to dominant brachydactyly type B, is caused by mutation of ROR2. Nat Genet. 2000;25:419–422.
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Receptor Tyrosine Kinase-like Orphan Receptors / metabolism*
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|
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href="/?term=%22Receptor+Tyrosine+Kinase-like+Orphan+Receptors%2Fmetabolism%22%5BMAJR%5D&sort=date&sort_order=desc"
|
||
data-href="/?term=%22Receptor+Tyrosine+Kinase-like+Orphan+Receptors%2Fmetabolism%22%5BMAJR%5D&sort=date&sort_order=desc"
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Search in MeSH
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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 major"
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Signal Transduction*
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aria-label="Dropdown menu for keyword Signal Transduction*"><div class="title">Actions</div><div class="content"><ul class="keyword-actions-links"><li><a class="search-in-pubmed-link dropdown-block-link"
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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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Wnt Proteins / antagonists & inhibitors
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</button><div id="keyword-actions-mesh-terms-13"
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aria-label="Dropdown menu for keyword Wnt Proteins / antagonists & inhibitors"><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 major"
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Wnt Proteins / metabolism*
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aria-label="Dropdown menu for keyword Wnt Proteins / metabolism*"><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 Wnt-5a Protein"
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Wnt-5a Protein
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hidden
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aria-label="Dropdown menu for keyword Wnt-5a Protein"><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-href="/?term=%22Wnt-5a+Protein%22%5BMeSH%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&mesh="Wnt-5a Protein"[MeSH]"
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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><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 beta Catenin / metabolism"
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beta Catenin / metabolism
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</button><div id="keyword-actions-mesh-terms-16"
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hidden
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aria-label="Dropdown menu for keyword beta Catenin / metabolism"><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-href="/?term=%22beta+Catenin%2Fmetabolism%22%5BMeSH%5D&sort=date&sort_order=desc"
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Add to Search
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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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|
||
|
||
<div id="substances" class="substances keywords-section">
|
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<h2 class="title">
|
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Substances
|
||
</h2>
|
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|
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<ul class="keywords-list"><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 Ltap protein, mouse"
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tabindex="0"
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aria-expanded="false"
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data-pinger-ignore>
|
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Ltap protein, mouse
|
||
</button><div id="keyword-actions-substances-0"
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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 Ltap 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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href="/?term=%22Ltap+protein%2C+mouse%22%5Bnm%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Ltap+protein%2C+mouse%22%5Bnm%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&keyword="Ltap protein, mouse"[nm]"
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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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data-search-term=""Ltap protein, mouse"[nm]"
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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 Nerve Tissue Proteins"
|
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tabindex="0"
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aria-controls="keyword-actions-substances-1"
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aria-expanded="false"
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data-pinger-ignore>
|
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Nerve Tissue Proteins
|
||
</button><div id="keyword-actions-substances-1"
|
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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 Nerve Tissue 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=%22Nerve+Tissue+Proteins%22%5Bnm%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Nerve+Tissue+Proteins%22%5Bnm%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&keyword="Nerve Tissue Proteins"[nm]"
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Search in PubMed
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Search in MeSH
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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 Wnt Proteins"
|
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tabindex="0"
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aria-controls="keyword-actions-substances-2"
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aria-expanded="false"
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data-pinger-ignore>
|
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Wnt Proteins
|
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</button><div id="keyword-actions-substances-2"
|
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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 Wnt 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=%22Wnt+Proteins%22%5Bnm%5D&sort=date&sort_order=desc"
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data-href="/?term=%22Wnt+Proteins%22%5Bnm%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&keyword="Wnt Proteins"[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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Search in MeSH
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|
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data-search-term=""Wnt Proteins"[nm]"
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data-ga-category="search"
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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 Wnt-5a Protein"
|
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tabindex="0"
|
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aria-controls="keyword-actions-substances-3"
|
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aria-expanded="false"
|
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data-pinger-ignore>
|
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Wnt-5a Protein
|
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</button><div id="keyword-actions-substances-3"
|
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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 Wnt-5a Protein"><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=%22Wnt-5a+Protein%22%5Bnm%5D&sort=date&sort_order=desc"
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ref="linksrc=search_in_pubmed_keyword_link&keyword="Wnt-5a Protein"[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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Search in MeSH
|
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</a></li><li><a class="add-to-search-link dropdown-block-link"
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data-search-term=""Wnt-5a Protein"[nm]"
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data-ga-category="search"
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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 Wnt5a protein, mouse"
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tabindex="0"
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aria-expanded="false"
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data-pinger-ignore>
|
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Wnt5a protein, mouse
|
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</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 Wnt5a 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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href="/?term=%22Wnt5a+protein%2C+mouse%22%5Bnm%5D&sort=date&sort_order=desc"
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Search in PubMed
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