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<input type="email" aria-label="Sender Email Address" placeholder="email@example.com" id="email-from" class="email-from" pattern="^[a-zA-Z0-9_.+-]+@[a-zA-Z0-9-]+\.[a-zA-Z0-9-.]+$" maxlength="256">
</div>
<div class="action-panel-control-wrap">
<label for="email-citation-format" class="action-panel-label">
Format:
</label>
<select id="email-citation-format" name="citation-format" class="action-panel-selector email-citation-format">
<option selected="selected" value="summary">Summary</option>
<option value="summary-text">Summary (text)</option>
<option value="abstract">Abstract</option>
<option value="abstract-text">Abstract (text)</option>
</select>
</div>
<div class="include-supplemental-container">
<input type="checkbox" aria-label="Include MeSH and other data" name="include-supplemental" id="email-include-supplemental" class="email-include-supplemental">
<label for="email-include-supplemental" class="email-include-supplemental-label">MeSH and other data</label>
</div>
<div class="form-field recaptcha ">
<div class="g-recaptcha" id="id-recaptcha" data-sitekey="6LfsWHMdAAAAAClKbtOpjQ2pMjgsGxvv7NdZW9uI"></div>
</div>
<div id="captcha-error-message" class="usa-input-error-message captcha-validation-message" role="alert"></div>
<div class="action-panel-actions">
<button class="action-panel-submit"
type="submit"
data-loading-label="Sending..."
data-ga-category="save_share"
data-ga-action="email"
data-ga-label="send">
Send email
</button>
<button class="action-panel-cancel"
aria-label="Close 'Email citations' panel"
ref="linksrc=close_email_panel"
aria-controls="email-action-panel"
aria-expanded="false"
data-ga-category="save_share"
data-ga-action="email"
data-ga-label="cancel">
Cancel
</button>
</div>
<input type="hidden" name="email-search-details" value="" />
<input type="hidden" name="email-search-details-hash" value="0e42663a6c3bd85498fcb88798998fed7bfdc45d457db35281e41afe13cc0524" />
</form>
</div>
</div>
<div id="collections-action-panel"
class="collections-action-panel action-panel in-progress-dots-panel"
aria-hidden="true"
data-collections-open-panel-enabled="false"
data-collections-open-panel-url-hash="#open-collections-panel">
<div class="inner-wrap">
<h3 class="action-panel-heading">
Add to Collections
</h3>
<form id="collections-action-panel-form"
class="collections-action-panel-form action-panel-content action-form action-panel-smaller-selectors"
data-existing-collections-url="/list-existing-collections/"
data-add-to-existing-collection-url="/add-to-existing-collection/"
data-create-and-add-to-new-collection-url="/create-and-add-to-new-collection/"
data-get-article-ids-by-search-url="/get-article-ids-by-search/"
data-myncbi-max-collection-name-length="100"
data-add-to-collection-max-amount="1000"
data-collections-root-url="https://www.ncbi.nlm.nih.gov/myncbi/collections/">
<input type="hidden" name="csrfmiddlewaretoken" value="mGlYfTNPuMHtJ20gfpHdjZ5YpwRmIwTlDt53SJgb4nuXDt66SIkfN7BCXAzyB7rF">
<div class="choice-group" role="radiogroup">
<ul class="radio-group-items">
<li>
<input type="radio"
id="collections-action-panel-new"
class="collections-new"
name="collections"
value="new"
data-ga-category="save_share"
data-ga-action="collections"
data-ga-label="collections_radio_new">
<label for="collections-action-panel-new">Create a new collection</label>
</li>
<li>
<input type="radio"
id="collections-action-panel-existing"
class="collections-existing"
name="collections"
value="existing"
checked="true"
data-ga-category="save_share"
data-ga-action="collections"
data-ga-label="collections_radio_existing">
<label for="collections-action-panel-existing">Add to an existing collection</label>
</li>
</ul>
</div>
<div class="controls-wrapper">
<div class="action-panel-control-wrap new-collections-controls">
<label for="collections-action-panel-add-to-new" class="action-panel-label required-field-asterisk">
Name your collection:
</label>
<input
type="text"
name="add-to-new-collection"
id="collections-action-panel-add-to-new"
class="collections-action-add-to-new"
pattern="[^&quot;&amp;=&lt;&gt;\/]*" title="The following characters are not allowed in the Name field: &quot;&amp;=&lt;&gt;/"
maxlength="100"
data-ga-category="save_share"
data-ga-action="create_collection"
data-ga-label="non_favorties_collection">
<div class="collections-new-name-too-long usa-input-error-message selection-validation-message">
Name must be less than 100 characters
</div>
</div>
<div class="action-panel-control-wrap existing-collections-controls">
<label for="collections-action-panel-add-to-existing" class="action-panel-label">
Choose a collection:
</label>
<select id="collections-action-panel-add-to-existing"
class="action-panel-selector collections-action-add-to-existing"
data-ga-category="save_share"
data-ga-action="select_collection"
data-ga-label="($('#collections-action-add-to-existing').val() === 'Favorites') ? 'Favorites' : 'non_favorites_collection'">
</select>
<div class="collections-retry-load-on-error usa-input-error-message selection-validation-message">
Unable to load your collection due to an error<br>
<a href="#">Please try again</a>
</div>
</div>
</div>
<div class="action-panel-actions">
<button class="action-panel-submit"
type="submit"
data-loading-label="Adding..."
data-pinger-ignore
data-ga-category="save_share"
data-ga-action="collections"
data-ga-label="add">
Add
</button>
<button class="action-panel-cancel"
aria-label="Close 'Add to Collections' panel"
ref="linksrc=close_collections_panel"
aria-controls="collections-action-panel"
aria-expanded="false"
data-ga-category="save_share"
data-ga-action="collections"
data-ga-label="cancel">
Cancel
</button>
</div>
</form>
</div>
</div>
<div id="bibliography-action-panel"
class="bibliography-action-panel action-panel in-progress-dots-panel"
aria-hidden="true"
data-bibliography-open-panel-enabled="false"
data-bibliography-open-panel-url-hash="#open-bibliography-panel">
<div class="inner-wrap">
<h3 class="action-panel-heading">
Add to My Bibliography
</h3>
<form id="bibliography-action-panel-form"
class="bibliography-action-panel-form action-panel-content action-form action-panel-smaller-selectors"
data-add-to-bibliography-max-amount="100"
data-add-to-bibliography-batch-size="10"
data-bibliography-delegates-url="/list-bibliography-delegates/"
data-add-to-bibliography-url="/add-to-bibliography/"
data-get-article-ids-by-search-url="/get-article-ids-by-search/"
data-mybib-root-url="https://www.ncbi.nlm.nih.gov/myncbi/collections/mybibliography/">
<input type="hidden" name="csrfmiddlewaretoken" value="mGlYfTNPuMHtJ20gfpHdjZ5YpwRmIwTlDt53SJgb4nuXDt66SIkfN7BCXAzyB7rF">
<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="mGlYfTNPuMHtJ20gfpHdjZ5YpwRmIwTlDt53SJgb4nuXDt66SIkfN7BCXAzyB7rF">
<div class="action-panel-control-wrap">
<label for="saved-search-name" class="action-panel-label saved-search-name-label required-field-asterisk">
Name of saved search:
</label>
<input maxlength="200"
type="text"
name="saved-search-name"
id="saved-search-name"
class="saved-search-name"
value=""
required
pattern="[^&quot;&amp;=&lt;&gt;\/]*" title="The following characters are not allowed in the Name field: &quot;&amp;=&lt;&gt;/">
</div>
<div class="action-panel-control-wrap">
<label for="saved-search-term" class="action-panel-label required-field-asterisk">
Search terms:
</label>
<textarea name="saved-search-term" id="saved-search-term" class="saved-search-term" required></textarea>
</div>
<div class="test-search-term-wrap">
<a href="#" class="try-search-term">Test search terms</a>
</div>
<div class="choice-group action-panel-extra-margin-top">
<span class="action-panel-label" id="fieldset-label">
Would you like email updates of new search results?
</span>
<fieldset id="saved-search-alert" aria-describedby="fieldset-label">
<legend class="usa-sr-only">Saved Search Alert Radio Buttons</legend>
<ul class="radio-group-items">
<li>
<input type="radio" id="saved-search-alert-yes" class="saved-search-alert-yes" name="saved-search-alert" value="yes" checked>
<label for="saved-search-alert-yes" class="action-panel-label">Yes</label>
</li>
<li>
<input aria-label="No radio input" type="radio" id="saved-search-alert-no" class="saved-search-alert-no" name="saved-search-alert" value="no">
<label for="saved-search-alert-no" class="action-panel-label">No</label>
</li>
</ul>
</fieldset>
</div>
<div class="alert-schedule-wrap">
<div class="action-panel-control-wrap">
<label class="action-panel-label">
Email:
</label>
<span aria-label="Email address" id="saved-search-email" class="action-panel-label"><span class="action-panel-label-bold"></span> (<a class="myncbi-account-settings" href="https://www.ncbi.nlm.nih.gov/account/settings/">change</a>)</span>
</div>
<div class="action-panel-control-wrap action-panel-extra-margin-top">
<label for="saved-search-frequency" class="action-panel-label">
Frequency:
</label>
<select id="saved-search-frequency" class="no-border-panel-selector saved-search-frequency">
<option value="monthly">Monthly</option>
<option value="weekly">Weekly</option>
<option value="daily">Daily</option>
</select>
</div>
<div class="action-panel-control-wrap saved-search-monthly-additional">
<label for="saved-search-monthly-on-day" class="action-panel-label">
Which day?
</label>
<select id="saved-search-monthly-on-day" class="no-border-panel-selector">
<option value="Sunday">The first Sunday</option>
<option value="Monday">The first Monday</option>
<option value="Tuesday">The first Tuesday</option>
<option value="Wednesday">The first Wednesday</option>
<option value="Thursday">The first Thursday</option>
<option value="Friday">The first Friday</option>
<option value="Saturday">The first Saturday</option>
<option value="day">The first day</option>
<option value="weekday">The first weekday</option>
</select>
</div>
<div class="action-panel-control-wrap saved-search-weekly-additional">
<label for="saved-search-weekly-on-day" class="action-panel-label">
Which day?
</label>
<select id="saved-search-weekly-on-day" class="no-border-panel-selector saved-search-weekly-on-day">
<option value="Sunday">Sunday</option>
<option value="Monday">Monday</option>
<option value="Tuesday">Tuesday</option>
<option value="Wednesday">Wednesday</option>
<option value="Thursday">Thursday</option>
<option value="Friday">Friday</option>
<option value="Saturday">Saturday</option>
</select>
</div>
<div class="action-panel-control-wrap">
<label for="saved-search-report" class="action-panel-label">
Report format:
</label>
<select id="saved-search-report" class="no-border-panel-selector saved-search-report">
<option value="DocSum">Summary</option>
<option value="DocSumText">Summary (text)</option>
<option value="Abstract">Abstract</option>
<option value="AbstractText">Abstract (text)</option>
<option value="MEDLINE">PubMed</option>
</select>
</div>
<div class="action-panel-control-wrap">
<label for="saved-search-amount" class="action-panel-label">
Send at most:
</label>
<select id="saved-search-amount" class="no-border-panel-selector saved-search-amount">
<option value="1">1 item</option>
<option value="5" selected>5 items</option>
<option value="10">10 items</option>
<option value="20">20 items</option>
<option value="50">50 items</option>
<option value="100">100 items</option>
<option value="200">200 items</option>
</select>
</div>
<div>
<input type="checkbox" id="saved-search-send-if-no-result" class="saved-search-send-if-no-result" name="saved-search-send-if-no-result">
<label for="saved-search-send-if-no-result" class="action-panel-label smaller-checkbox">
Send even when there aren't any new results
</label>
</div>
<div class="action-panel-control-wrap option-text-in-email-wrap">
<label for="saved-search-email-text" class="action-panel-label">
Optional text in email:
</label>
<textarea name="saved-search-email-text"
id="saved-search-email-text"
class="saved-search-email-text"></textarea>
</div>
</div>
<div class="action-panel-actions">
<button class="action-panel-submit"
type="submit"
data-loading-label="Saving..."
data-ga-category="save_share"
data-ga-action="alert"
data-ga-label="save">
Save
</button>
<button class="action-panel-cancel"
aria-label="Close 'Your saved search' panel"
ref="linksrc=close_saved_search_panel"
aria-controls="saved-search-action-panel"
aria-expanded="false"
data-ga-category="save_share"
data-ga-action="alert"
data-ga-label="cancel">
Cancel
</button>
</div>
</form>
</div>
</div>
<div id="citation-manager-action-panel" class="citation-manager-action-panel action-panel" aria-hidden="true">
<div class="inner-wrap">
<h2 class="action-panel-heading">
Create a file for external citation management software
</h2>
<form id="citation-manager-action-panel-form"
class="action-panel-content action-form"
action="/results-export-ids/"
data-by-search-action="/results-export-search-data/"
data-by-ids-action="/results-export-ids/"
method="post"
data-by-search-method="post"
data-by-ids-method="post">
<input type="hidden" name="csrfmiddlewaretoken" value="mGlYfTNPuMHtJ20gfpHdjZ5YpwRmIwTlDt53SJgb4nuXDt66SIkfN7BCXAzyB7rF">
<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="mGlYfTNPuMHtJ20gfpHdjZ5YpwRmIwTlDt53SJgb4nuXDt66SIkfN7BCXAzyB7rF">
<div class="action-panel-control-wrap">
<label for="rss-name" class="action-panel-label required-field-asterisk">
Name of RSS Feed:
</label>
<input maxlength="200"
placeholder="Name"
type="text"
name="rss-name"
id="rss-name"
class="rss-name"
value=''
required
pattern="[^&quot;&amp;=&lt;&gt;\/]*" title="The following characters are not allowed in the Name field: &quot;&amp;=&lt;&gt;/">
</div>
<div class="rss-limit-wrap">
<div class="action-panel-control-wrap action-panel-extra-margin-top">
<label for="rss-limit" class="action-panel-label">
Number of items displayed:
</label>
<select id="rss-limit" class="no-border-panel-selector rss-limit">
<option value="5">5</option>
<option value="10">10</option>
<option value="15" selected="selected">15</option>
<option value="20">20</option>
<option value="50">50</option>
<option value="100">100</option>
</select>
</div>
</div>
<div class="action-panel-actions">
<button class="action-panel-submit"
type="submit"
data-loading-label="Creating..."
data-ga-category="save_share"
data-ga-action="alert"
data-ga-label="save">
Create RSS
</button>
<button class="action-panel-cancel"
aria-label="Close 'Your RSS' panel"
ref="linksrc=close_rss_panel"
aria-controls="rss-action-panel"
aria-expanded="false"
data-ga-category="save_share"
data-ga-action="alert"
data-ga-label="cancel">
Cancel
</button>
</div>
<div class="action-panel-control-wrap rss-link-copy-wrap">
<label for="rss-link" class="usa-sr-only">RSS Link</label>
<input placeholder="Your RSS Feed Link" type="text" name="rss-link" id="rss-link" class="rss-link" title="RSS Link">
<button
type="button"
disabled
class="rss-link-copy-button disabled"
data-ga-category="save_share"
data-ga-action="rss"
data-ga-label="copy">
Copy
</button>
</div>
</form>
</div>
</div>
</div>
</div>
<div class="article-page" id="article-page" data-article-pmid="35380452">
<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://journals.asm.org/doi/10.1128/aem.00240-22?url_ver=Z39.88-2003&amp;rfr_id=ori:rid:crossref.org&amp;rfr_dat=cr_pub 0pubmed"
target="_blank"
rel="noopener"
ref="linksrc=fulltextorjournal_fulltext&amp;is_pmc=False&amp;PrId=4487&amp;itool=Abstract-def&amp;log$=linkouticon&amp;uid=35380452&amp;db=pubmed&amp;nlmid=7605801"
title="See full text options at Atypon"
data-ga-category="full_text"
data-ga-action="Atypon"
data-ga-label="35380452"
><img src="https://cdn.ncbi.nlm.nih.gov/corehtml/query/egifs/https:--journals.asm.org-pb-assets-pubmed-pubmed_AEM.png" alt="Atypon full text link"><span class="text">
Atypon
</span></a><a class="link-item
pmc
"
href="https://pmc.ncbi.nlm.nih.gov/articles/pmid/35380452/"
target="_blank"
rel="noopener"
ref="linksrc=fulltextorjournal_fulltext&amp;is_pmc=True&amp;PrId=3494&amp;itool=Abstract-def&amp;log$=linkouticon&amp;uid=35380452&amp;db=pubmed&amp;nlmid=7605801"
title="Free full text at PubMed Central"
data-ga-category="full_text"
data-ga-action="PMC"
data-ga-label="35380452"
><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="/35380452/citations/"
data-citation-style="nlm"
data-pubmed-format-link="/35380452/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="35380452" 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%2F35380452%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="mGlYfTNPuMHtJ20gfpHdjZ5YpwRmIwTlDt53SJgb4nuXDt66SIkfN7BCXAzyB7rF"><div class="choice-group" role="radiogroup"><ul class="radio-group-items"><li><input type="radio"
id="collections-action-dialog-new"
class="collections-new"
name="collections"
value="new"
data-ga-category="collections_button"
data-ga-action="click"
data-ga-label="collections_radio_new"><label for="collections-action-dialog-new">Create a new collection</label></li><li><input type="radio"
id="collections-action-dialog-existing"
class="collections-existing"
name="collections"
value="existing"
checked="true"
data-ga-category="collections_button"
data-ga-action="click"
data-ga-label="collections_radio_existing"><label for="collections-action-dialog-existing">Add to an existing collection</label></li></ul></div><div class="controls-wrapper"><div class="action-panel-control-wrap new-collections-controls"><label for="collections-action-dialog-add-to-new" class="action-panel-label required-field-asterisk">
Name your collection:
</label><input
type="text"
name="add-to-new-collection"
id="collections-action-dialog-add-to-new"
class="collections-action-add-to-new"
pattern="[^&quot;&amp;=&lt;&gt;\/]*" title="The following characters are not allowed in the Name field: &quot;&amp;=&lt;&gt;/"
maxlength="100"
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<h1 class="heading-title">
Annulment of Bacterial Antagonism Improves Plant Beneficial Activity of a Bacillus velezensis Consortium
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Annulment of Bacterial Antagonism Improves Plant Beneficial Activity of a Bacillus velezensis Consortium
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</a></li></ul></div></div></div><span class="period">. </span><span class="cit">2022 Apr 26;88(8):e0024022.</span>
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doi: 10.1128/aem.00240-22.
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Epub 2022 Apr 5.
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<span class="authors-list-item "><a class="full-name"
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data-ga-label="Jiahui Shao">Jiahui Shao</a><sup class="equal-contrib-container"><span class="author-sup-separator">&nbsp;</span><a class="equal-contrib" title="Contributed equally" href="#short-view-equal-contrib-explanation" ref="linksrc=author_equal_contrib">#</a></sup><sup class="affiliation-links"><span class="author-sup-separator">&nbsp;</span><a class="affiliation-link" title="Jiangsu Provincial Key Lab of Solid Organic Waste Utilization, Jiangsu Collaborative Innovation Center of Solid Organic Wastes, Educational Ministry Engineering Center of Resource-Saving Fertilizers, The Key Laboratory of Plant Immunity, Nanjing Agricultural Universitygrid.27871.3b, Nanjing, Jiangsu, People&#x27;s Republic of China." href="#short-view-affiliation-1" ref="linksrc=author_aff">
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</a></sup><span class="comma">,&nbsp;</span></span><span class="authors-list-item "><a class="full-name"
href="/?term=Liu+Y&amp;cauthor_id=35380452"
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data-ga-label="Yan Liu">Yan Liu</a><sup class="equal-contrib-container"><span class="author-sup-separator">&nbsp;</span><a class="equal-contrib" title="Contributed equally" href="#short-view-equal-contrib-explanation" ref="linksrc=author_equal_contrib">#</a></sup><sup class="affiliation-links"><span class="author-sup-separator">&nbsp;</span><a class="affiliation-link" title="Jiangsu Provincial Key Lab of Solid Organic Waste Utilization, Jiangsu Collaborative Innovation Center of Solid Organic Wastes, Educational Ministry Engineering Center of Resource-Saving Fertilizers, The Key Laboratory of Plant Immunity, Nanjing Agricultural Universitygrid.27871.3b, Nanjing, Jiangsu, People&#x27;s Republic of China." href="#short-view-affiliation-1" ref="linksrc=author_aff">
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</a></sup><span class="comma">,&nbsp;</span></span><span class="authors-list-item "><a class="full-name"
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data-ga-label="Jiyu Xie">Jiyu Xie</a><sup class="affiliation-links"><span class="author-sup-separator">&nbsp;</span><a class="affiliation-link" title="Jiangsu Provincial Key Lab of Solid Organic Waste Utilization, Jiangsu Collaborative Innovation Center of Solid Organic Wastes, Educational Ministry Engineering Center of Resource-Saving Fertilizers, The Key Laboratory of Plant Immunity, Nanjing Agricultural Universitygrid.27871.3b, Nanjing, Jiangsu, People&#x27;s Republic of China." href="#short-view-affiliation-1" ref="linksrc=author_aff">
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</a></sup><span class="comma">,&nbsp;</span></span><span class="authors-list-item "><a class="full-name"
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data-ga-label="Polonca Štefanič">Polonca Štefanič</a><sup class="affiliation-links"><span class="author-sup-separator">&nbsp;</span><a class="affiliation-link" title="Biotechnical Faculty, University of Ljubljana, Ljubljana, Slovenia." href="#short-view-affiliation-2" ref="linksrc=author_aff">
2
</a></sup><span class="comma">,&nbsp;</span></span><span class="authors-list-item "><a class="full-name"
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data-ga-label="Yu Lv">Yu Lv</a><sup class="affiliation-links"><span class="author-sup-separator">&nbsp;</span><a class="affiliation-link" title="Jiangsu Provincial Key Lab of Solid Organic Waste Utilization, Jiangsu Collaborative Innovation Center of Solid Organic Wastes, Educational Ministry Engineering Center of Resource-Saving Fertilizers, The Key Laboratory of Plant Immunity, Nanjing Agricultural Universitygrid.27871.3b, Nanjing, Jiangsu, People&#x27;s Republic of China." href="#short-view-affiliation-1" ref="linksrc=author_aff">
1
</a></sup><span class="comma">,&nbsp;</span></span><span class="authors-list-item "><a class="full-name"
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data-ga-label="Ben Fan">Ben Fan</a><sup class="affiliation-links"><span class="author-sup-separator">&nbsp;</span><a class="affiliation-link" title="Co-Innovation Center for Sustainable Forestry in Southern China, College of Forestry, Nanjing Forestry University, Nanjing, People&#x27;s Republic of China." href="#short-view-affiliation-3" ref="linksrc=author_aff">
3
</a></sup><span class="comma">,&nbsp;</span></span><span class="authors-list-item "><a class="full-name"
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</a></sup><span class="comma">,&nbsp;</span></span><span class="authors-list-item "><a class="full-name"
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data-ga-label="Ruifu Zhang">Ruifu Zhang</a><sup class="affiliation-links"><span class="author-sup-separator">&nbsp;</span><a class="affiliation-link" title="Jiangsu Provincial Key Lab of Solid Organic Waste Utilization, Jiangsu Collaborative Innovation Center of Solid Organic Wastes, Educational Ministry Engineering Center of Resource-Saving Fertilizers, The Key Laboratory of Plant Immunity, Nanjing Agricultural Universitygrid.27871.3b, Nanjing, Jiangsu, People&#x27;s Republic of China." href="#short-view-affiliation-1" ref="linksrc=author_aff">
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data-ga-label="Qirong Shen">Qirong Shen</a><sup class="affiliation-links"><span class="author-sup-separator">&nbsp;</span><a class="affiliation-link" title="Jiangsu Provincial Key Lab of Solid Organic Waste Utilization, Jiangsu Collaborative Innovation Center of Solid Organic Wastes, Educational Ministry Engineering Center of Resource-Saving Fertilizers, The Key Laboratory of Plant Immunity, Nanjing Agricultural Universitygrid.27871.3b, Nanjing, Jiangsu, People&#x27;s Republic of China." href="#short-view-affiliation-1" ref="linksrc=author_aff">
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</a></sup><span class="comma">,&nbsp;</span></span><span class="authors-list-item "><a class="full-name"
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data-ga-label="Zhihui Xu">Zhihui Xu</a><sup class="affiliation-links"><span class="author-sup-separator">&nbsp;</span><a class="affiliation-link" title="Jiangsu Provincial Key Lab of Solid Organic Waste Utilization, Jiangsu Collaborative Innovation Center of Solid Organic Wastes, Educational Ministry Engineering Center of Resource-Saving Fertilizers, The Key Laboratory of Plant Immunity, Nanjing Agricultural Universitygrid.27871.3b, Nanjing, Jiangsu, People&#x27;s Republic of China." href="#short-view-affiliation-1" ref="linksrc=author_aff">
1
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<li data-affiliation-id="short-view-affiliation-1"
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><sup class="key">1</sup> Jiangsu Provincial Key Lab of Solid Organic Waste Utilization, Jiangsu Collaborative Innovation Center of Solid Organic Wastes, Educational Ministry Engineering Center of Resource-Saving Fertilizers, The Key Laboratory of Plant Immunity, Nanjing Agricultural Universitygrid.27871.3b, Nanjing, Jiangsu, People&#x27;s Republic of China.</li>
<li data-affiliation-id="short-view-affiliation-2"
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><sup class="key">2</sup> Biotechnical Faculty, University of Ljubljana, Ljubljana, Slovenia.</li>
<li data-affiliation-id="short-view-affiliation-3"
id="short-view-affiliation-3"
><sup class="key">3</sup> Co-Innovation Center for Sustainable Forestry in Southern China, College of Forestry, Nanjing Forestry University, Nanjing, People&#x27;s Republic of China.</li>
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<div class="equal-contrib-explanation" id="short-view-equal-contrib-explanation"><sup class="key">#</sup> Contributed equally.</div>
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<p>
<i>Bacillus</i> sp. strains that are beneficial to plants are widely used in commercial biofertilizers and biocontrol agents for sustainable agriculture. Generally, functional <i>Bacillus</i> strains are applied as single-strain communities since the principles of synthetic microbial consortia constructed with <i>Bacillus</i> strains remain largely unclear. Here, we demonstrated that the mutual compatibility directly affects the survival and function of two-member consortia composed of Bacillus velezensis SQR9 and FZB42 in the rhizosphere. A mutation in the global regulator Spo0A of SQR9 markedly reduced the boundary phenotype (appearance of a visible boundary line at the meeting point of two swarms) with wild-type FZB42, and the combined use of the SQR9(△<i>spo0A</i>) mutant and FZB42 improved biofilm formation, root colonization, and the production of secondary metabolites that are beneficial to plants. Furthermore, alleviation of antagonistic interactions of two-member <i>Bacillus</i> consortia improved its beneficial effects to cucumber in a greenhouse experiment. Our results provide evidence that social interactions among bacteria could be an influencing factor for achieving a desired community-level function. <b>IMPORTANCE</b> Bacillus velezensis is one of the most widely applied bacteria in biofertilizers in China and Europe. Additionally, the molecular mechanisms of plant growth promotion and disease suppression by representative model strains are well established, such as <i>B. velezensis</i> SQR9 and FZB42. However, it remains extremely challenging to design efficient consortia based on these model strains. Here, we showed that swarm encounter phenotype is one of the major determinants that affects the performance of two-member <i>Bacillus</i> consortia <i>in vitro</i> and in the rhizosphere. Deletion in global regulatory gene <i>spo0A</i> of SQR9 reduced the strength of boundary formation with FZB42 and resulted in the improved plant growth promotion performance of the dual consortium. This knowledge provides new insights into efficient probiotics consortia design in <i>Bacillus</i> spp.
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</strong>
Bacillus velezensis; biofilm formation; mutual compatibility; plant growth-promoting; social interactions; synthetic microbial consortia.
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<p xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:p1="http://pubmed.gov/pub-one">The authors declare no conflict of interest.</p>
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<a class="figure-link"
href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/2503/9040612/a50424cb69a6/aem.00240-22-f001.jpg"
itemprop="contentUrl"
aria-describedby="figure-caption-0"
role="button"
data-image-width="696"
data-image-height="623"
data-image-alt="FIG 1"
data-pmc-id="PMC9040612"
data-figure-id="F1">
<img class="figure-thumb" itemprop="thumbnail"
id="article-image-0"
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/2503/9040612/d34a2d5c661c/aem.00240-22-f001.gif"
alt="FIG 1" />
</a>
<meta itemprop="width" itemtype="http://schema.org/ImageObject" content="696">
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<div class="caption-wrap">
<strong class="figure-label">
<p> FIG 1 </p>
</strong>
<div class="figure-caption-contents"><p> Regulator Spo0A in <i> B. velezensis… </i> </p></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG 1 </p>
</strong>
<div class="figure-caption-contents"><p> Regulator Spo0A in <i> B. velezensis </i> SQR9 influences the mutual compatibility between <i> B. velezensis… </i> </p></div>
</div>
<figcaption id="figure-caption-0" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
<strong class="figure-label">
FIG 1
</strong>
<div class="figure-caption-contents">Regulator Spo0A in <i>B. velezensis</i> SQR9 influences the mutual compatibility between <i>B. velezensis</i> SQR9 and FZB42. (A) Different candidate gene deletions of <i>B. velezensis</i> SQR9 showed varied multicellular swarming encounter phenotypes with wild-type <i>B. velezensis</i> FZB42. Results are representative of three experiments. For each photograph, <i>B. velezensis</i> SQR9 or its derivative strains were spotted onto the surface of the swarming agar (B-medium) at a similar distance apart, with either wild-type <i>B. velezensis</i> FZB42 or SQR9 spotted onto the lower portion of plates. (B) Growth compatibility assays for coculture of <i>B. velezensis</i> SQR9+FZB42 and <i>B. velezensis</i> SQR9(△<i>spo0A</i>)+FZB42. (C) The ratio of SQR9/FZB42 or SQR9(△<i>spo0A</i>)/FZB42 cells during coculture at 10-, 15-, 20-, 25-, 30-, and 35-h time points. For each column, the area occupied by light blue indicates the proportion of FZB42 cells in a different population.</div>
</figcaption>
</figure>
<figure class="figure-item "
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data-slide-index="1"
data-label-slug="fig-2">
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itemprop="contentUrl"
aria-describedby="figure-caption-1"
role="button"
data-image-width="675"
data-image-height="1292"
data-image-alt="FIG 2"
data-pmc-id="PMC9040612"
data-figure-id="F2">
<img class="figure-thumb" itemprop="thumbnail"
id="article-image-1"
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/2503/9040612/b09d0de7580e/aem.00240-22-f002.gif"
alt="FIG 2" />
</a>
<meta itemprop="width" itemtype="http://schema.org/ImageObject" content="675">
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<div class="caption-wrap">
<strong class="figure-label">
<p> FIG 2 </p>
</strong>
<div class="figure-caption-contents"><p> Pellicle biofilm formation and cucumber… </p></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG 2 </p>
</strong>
<div class="figure-caption-contents"><p> Pellicle biofilm formation and cucumber root colonization by monoculture and coculture of <i> Bacillus… </i> </p></div>
</div>
<figcaption id="figure-caption-1" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
<strong class="figure-label">
FIG 2
</strong>
<div class="figure-caption-contents">Pellicle biofilm formation and cucumber root colonization by monoculture and coculture of <i>Bacillus</i> strains. SQR9 and FZB42 indicate wild-type <i>B. velezensis</i> SQR9 and FZB42, respectively, and SQR9(△<i>spo0A</i>) indicates the <i>spo0A</i> mutant of <i>B. velezensis</i> SQR9. (A) Growth curves of <i>B. velezensis</i> FZB42 and <i>B. velezensis</i> SQR9 and its mutant strains in liquid MSgg medium. (B) Floating pellicle formation by monoculture and coculture of <i>Bacillus</i> strains. (C and D) OD<sub>600</sub> of solubilized crystal violet from the microtiter plate assay at 24 and 48h, respectively. (E and F) The <i>Bacillus</i> populations of monoculture and coculture colonizing cucumber seeding roots at 2- and 4-day time points. For coculture treatments in B, C, D and E, the ratio of dark gray (SQR9 or <i>spo0A</i> mutant) and light blue (FZB42) indicates the proportion of different cells in the population, dark gray represents the cells of strain SQR9 or its <i>spo0A</i> mutant, and light blue represents the cells of FZB42 wild type. Error bars indicate the standard deviations from the results from six independent experiments. Different letters above the bars indicate significant differences (<i>P</i> &lt; 0.05).</div>
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</figure>
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itemscope itemtype="http://schema.org/ImageObject"
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data-label-slug="fig-3">
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itemprop="contentUrl"
aria-describedby="figure-caption-2"
role="button"
data-image-width="625"
data-image-height="329"
data-image-alt="FIG 3"
data-pmc-id="PMC9040612"
data-figure-id="F3">
<img class="figure-thumb" itemprop="thumbnail"
id="article-image-2"
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/2503/9040612/325ade4cc8f5/aem.00240-22-f003.gif"
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</a>
<meta itemprop="width" itemtype="http://schema.org/ImageObject" content="625">
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<div class="caption-wrap">
<strong class="figure-label">
<p> FIG 3 </p>
</strong>
<div class="figure-caption-contents"><p> Qualitative (A) and quantitative (B)… </p></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG 3 </p>
</strong>
<div class="figure-caption-contents"><p> Qualitative (A) and quantitative (B) analysis of the distance between the fungal mycelium… </p></div>
</div>
<figcaption id="figure-caption-2" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
<strong class="figure-label">
FIG 3
</strong>
<div class="figure-caption-contents">Qualitative (A) and quantitative (B) analysis of the distance between the fungal mycelium (F. oxysporum) and the wall of an Oxford cup filled with the supernatants of monoculture or coculture of <i>Bacillus</i> strains. (A) Representative image of the distance between the fungal mycelium (F. oxysporum) and the wall of an Oxford cup filled with the supernatants of monoculture or coculture of <i>Bacillus</i> strains. Error bars in B indicate the standard deviations from the results from six independent experiments. SQR9 and FZB42 indicate wild-type <i>B. velezensis</i> SQR9 and FZB42, respectively, and SQR9(△<i>spo0A</i>) indicates an <i>spo0A</i> mutant of <i>B. velezensis</i> SQR9. Different letters above the bars indicate significant differences (<i>P</i> &lt; 0.05).</div>
</figcaption>
</figure>
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itemscope itemtype="http://schema.org/ImageObject"
itemprop="associatedMedia"
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data-label-slug="fig-4">
<a class="figure-link"
href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/2503/9040612/0d4bf1fed066/aem.00240-22-f004.jpg"
itemprop="contentUrl"
aria-describedby="figure-caption-3"
role="button"
data-image-width="696"
data-image-height="389"
data-image-alt="FIG 4"
data-pmc-id="PMC9040612"
data-figure-id="F4">
<img class="figure-thumb" itemprop="thumbnail"
id="article-image-3"
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/2503/9040612/63ad3f981fe1/aem.00240-22-f004.gif"
alt="FIG 4" />
</a>
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<div class="figure-caption figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="caption" aria-hidden="true">
<div class="caption-wrap">
<strong class="figure-label">
<p> FIG 4 </p>
</strong>
<div class="figure-caption-contents"><p> Reversed-phase HPLC chromatograms of lipopeptides… </p></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG 4 </p>
</strong>
<div class="figure-caption-contents"><p> Reversed-phase HPLC chromatograms of lipopeptides (bacillomycin D and fengycin) produced by monoculture and… </p></div>
</div>
<figcaption id="figure-caption-3" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
<strong class="figure-label">
FIG 4
</strong>
<div class="figure-caption-contents">Reversed-phase HPLC chromatograms of lipopeptides (bacillomycin D and fengycin) produced by monoculture and coculture of <i>Bacillus</i> strains. For the better comparison, we used liquid chromatographic peaks of lipopeptides produced by <i>B. velezensis</i> FZB42 as an internal reference (red lines). Liquid chromatographic peaks in black indicate different samples, as follows: A, lipopeptides produced by <i>B. velezensis</i> SQR9; B, lipopeptides produced by coculture of <i>B. velezensis</i> SQR9 and FZB42; C, lipopeptides produced by <i>spo0A</i> mutant of <i>B. velezensis</i> SQR9; D, lipopeptides produced by coculture of <i>B. velezensis</i> SQR9(△<i>spo0A</i>) and wild-type FZB42.</div>
</figcaption>
</figure>
<figure class="figure-item "
itemscope itemtype="http://schema.org/ImageObject"
itemprop="associatedMedia"
data-slide-index="4"
data-label-slug="fig-5">
<a class="figure-link"
href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/2503/9040612/7ca485a8806c/aem.00240-22-f005.jpg"
itemprop="contentUrl"
aria-describedby="figure-caption-4"
role="button"
data-image-width="800"
data-image-height="884"
data-image-alt="FIG 5"
data-pmc-id="PMC9040612"
data-figure-id="F5">
<img class="figure-thumb" itemprop="thumbnail"
id="article-image-4"
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/2503/9040612/1ab267d6e675/aem.00240-22-f005.gif"
alt="FIG 5" />
</a>
<meta itemprop="width" itemtype="http://schema.org/ImageObject" content="800">
<meta itemprop="height" itemtype="http://schema.org/ImageObject" content="884">
<div class="figure-caption figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="caption" aria-hidden="true">
<div class="caption-wrap">
<strong class="figure-label">
<p> FIG 5 </p>
</strong>
<div class="figure-caption-contents"><p> Production of IAA and acetoin… </p></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG 5 </p>
</strong>
<div class="figure-caption-contents"><p> Production of IAA and acetoin in monoculture and coculture of <i> Bacillus </i> strains. (A… </p></div>
</div>
<figcaption id="figure-caption-4" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
<strong class="figure-label">
FIG 5
</strong>
<div class="figure-caption-contents">Production of IAA and acetoin in monoculture and coculture of <i>Bacillus</i> strains. (A and C) IAA production of monoculture and coculture of <i>Bacillus</i> strains was determined at 24 and 48h, respectively. (B and D) Acetoin production in monoculture and coculture of <i>Bacillus</i> strains at 24- and 48-h time points, respectively. Error bars indicate the standard deviations from the results from six independent experiments. SQR9 and FZB42 indicate wild-type <i>B. velezensis</i> SQR9 and FZB42, respectively, and SQR9(△<i>spo0A</i>) indicates an <i>spo0A</i> mutant of <i>B. velezensis</i> SQR9. Different letters above the bars indicate significant differences (<i>P</i> &lt; 0.05).</div>
</figcaption>
</figure>
<figure class="figure-item "
itemscope itemtype="http://schema.org/ImageObject"
itemprop="associatedMedia"
data-slide-index="5"
data-label-slug="fig-6">
<a class="figure-link"
href="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/2503/9040612/4853151503c2/aem.00240-22-f006.jpg"
itemprop="contentUrl"
aria-describedby="figure-caption-5"
role="button"
data-image-width="660"
data-image-height="573"
data-image-alt="FIG 6"
data-pmc-id="PMC9040612"
data-figure-id="F6">
<img class="figure-thumb" itemprop="thumbnail"
id="article-image-5"
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/2503/9040612/54030d6e2fe6/aem.00240-22-f006.gif"
alt="FIG 6" />
</a>
<meta itemprop="width" itemtype="http://schema.org/ImageObject" content="660">
<meta itemprop="height" itemtype="http://schema.org/ImageObject" content="573">
<div class="figure-caption figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="caption" aria-hidden="true">
<div class="caption-wrap">
<strong class="figure-label">
<p> FIG 6 </p>
</strong>
<div class="figure-caption-contents"><p> Greenhouse experiment indicated that the… </p></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> FIG 6 </p>
</strong>
<div class="figure-caption-contents"><p> Greenhouse experiment indicated that the combined used of SQR9(△ <i> spo0A </i> ) and FZB42… </p></div>
</div>
<figcaption id="figure-caption-5" class="figure-caption-full figure-caption-text" itemtype="http://schema.org/ImageObject" itemprop="description">
<strong class="figure-label">
FIG 6
</strong>
<div class="figure-caption-contents">Greenhouse experiment indicated that the combined used of SQR9(△<i>spo0A</i>) and FZB42 significantly improved the growth of cucumber under F. oxysporum pathogen stress. SQR9 and FZB42 indicate wild-type <i>B. velezensis</i> SQR9 and FZB42, respectively, and SQR9(△<i>spo0A</i>) indicates an <i>spo0A</i> mutant of <i>B. velezensis</i> SQR9. (A) Representative image of the cucumber plants inoculated with monoculture and coculture of <i>Bacillus</i> strains before harvest. (B and C) The effect of different treatments on cucumber plant height and shoot dry weight, respectively. (D) <i>Bacillus</i> populations of monoculture and coculture colonizing cucumber roots after harvest. For coculture treatments [SQR9+FZB42+F. oxysporum and SQR9(△<i>spo0A</i>)+FZB42+F. oxysporum], the ratio of dark gray (SQR9 or <i>spo0A</i> mutant) and light blue (FZB42) indicates the proportion of different cells in the population; dark gray represents the cells of SQR9 or its <i>spo0A</i> mutant; and light blue represents the cells of FZB42. Error bars indicate the standard deviations from the results from six independent experiments. Different letters above the bars indicate significant differences (<i>P</i> &lt; 0.05).</div>
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