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<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="r6hGahnBfUl7EczPhuNCQYZMUyuzr4MDIT1LN7QXPv8ByDFFUNqEk6vqsCcLkFkX">
<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="r6hGahnBfUl7EczPhuNCQYZMUyuzr4MDIT1LN7QXPv8ByDFFUNqEk6vqsCcLkFkX">
<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="r6hGahnBfUl7EczPhuNCQYZMUyuzr4MDIT1LN7QXPv8ByDFFUNqEk6vqsCcLkFkX">
<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="r6hGahnBfUl7EczPhuNCQYZMUyuzr4MDIT1LN7QXPv8ByDFFUNqEk6vqsCcLkFkX">
<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="r6hGahnBfUl7EczPhuNCQYZMUyuzr4MDIT1LN7QXPv8ByDFFUNqEk6vqsCcLkFkX">
<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="33219166">
<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/S0021-9258(17)50424-X"
target="_blank"
rel="noopener"
ref="linksrc=fulltextorjournal_fulltext&amp;is_pmc=False&amp;PrId=3048&amp;itool=Abstract-def&amp;log$=linkouticon&amp;uid=33219166&amp;db=pubmed&amp;nlmid=2985121R"
title="See full text options at Elsevier Science"
data-ga-category="full_text"
data-ga-action="Elsevier Science"
data-ga-label="33219166"
><img src="https://cdn.ncbi.nlm.nih.gov/corehtml/query/egifs/https:--linkinghub.elsevier.com-ihub-images-elsevieroa.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/33219166/"
target="_blank"
rel="noopener"
ref="linksrc=fulltextorjournal_fulltext&amp;is_pmc=True&amp;PrId=3494&amp;itool=Abstract-def&amp;log$=linkouticon&amp;uid=33219166&amp;db=pubmed&amp;nlmid=2985121R"
title="Free full text at PubMed Central"
data-ga-category="full_text"
data-ga-action="PMC"
data-ga-label="33219166"
><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="/33219166/citations/"
data-citation-style="nlm"
data-pubmed-format-link="/33219166/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="33219166" 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%2F33219166%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="r6hGahnBfUl7EczPhuNCQYZMUyuzr4MDIT1LN7QXPv8ByDFFUNqEk6vqsCcLkFkX"><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"
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"
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There is nothing quite like the excitement of discovery in science-of finding something no one else knew and seeing a story unfold. One has to be part of an emerging picture to feel the elation. These moments in a lifetime are few and far between, but they fuel enthusiasm and keep one going. They are embedded in struggles and joys of everyday life, years of establishing what Louis Pasteur called "the prepared mind," working with mentors, trainees, and colleagues, failures and successes. This article recalls 1) how I got to be a biochemist; 2) my contributions as an educator and researcher, especially regarding meprin metalloproteases; and 3) my participation in communities of science. Perhaps my reflections will help an aspiring scientist see how fulfilling a career in science can be.
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academic paths; becoming a biochemist; collaborations; mentorship; meprins; metalloprotease; metalloproteases.
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id="article-image-1"
src="https://cdn.ncbi.nlm.nih.gov/pmc/blobs/93f1/7681016/5b0fd1f7e9a0/SB-JBCJ200689F002.gif"
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<strong class="figure-label">
<p> Figure 2. </p>
</strong>
<div class="figure-caption-contents"><b> Mentors and colleagues. </b> <i> A </i> ,…</div>
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</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> Figure 2. </p>
</strong>
<div class="figure-caption-contents"><b> Mentors and colleagues. </b> <i> A </i> , Rob Beynon (colleague), John Bird (my graduate school…</div>
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Figure 2.
</strong>
<div class="figure-caption-contents"><b>Mentors and colleagues.</b>
<i>A</i>, Rob Beynon (colleague), John Bird (my graduate school advisor), and me at a Symposium on Intracellular Protein Catabolism, in Buchenberg, GDR, 1986. <i>B</i>, Rollo Park (Chair of Physiology at Vanderbilt University in the 1960s), me, and Jane Harting Park (my postdoctoral advisor) at the ASBMB centennial celebration in 2006. <i>C</i>, Gaylen Bradley (my husband) and me 1995 in Hershey, Pennsylvania. <i>D</i>, Alan Barrett (my sabbatical mentor) and me at Strangeways Research Laboratory in 1978 in Cambridge, UK.</div>
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<strong class="figure-label">
<p> Figure 3. </p>
</strong>
<div class="figure-caption-contents"><b> Domain and oligomeric structure of… </b></div>
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<strong class="figure-label">
<p> Figure 3. </p>
</strong>
<div class="figure-caption-contents"><b> Domain and oligomeric structure of meprins α and β. </b> Domains are as follows.…</div>
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Figure 3.
</strong>
<div class="figure-caption-contents"><b>Domain and oligomeric structure of meprins α and β.</b> Domains are as follows. <i>S</i>, signal sequence; <i>Pro</i>, pro-sequence; <i>Protease</i>, catalytic domain; <i>MAM</i>, meprin, A5 protein, protein tyrosine phosphatase µ; <i>TRAF</i>, TRAF homology; <i>I</i>, inserted; <i>EGF</i>, epidermal growth factor-like; <i>TM</i>, transmembrane-spanning; <i>C</i>, cytoplasmic domain. During maturation, the meprin α subunit is cleaved in the I domain, separating the subunit from the membrane. As a result, three isoforms of meprin exist: membrane-bound meprin B (a homodimer of β subunits), membrane-bound meprin A (heterotetramers of α and β subunits, found in ratios of α<sub>2</sub>β<sub>2</sub> and α<sub>3</sub>β<sub>1</sub>), and secreted meprin A (homomeric multimers of α subunit dimers). The secreted forms of meprin α dimers tend to self-associate and form large multimers (16 MDa) (25) (see Fig. 5).</div>
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<strong class="figure-label">
<p> Figure 4. </p>
</strong>
<div class="figure-caption-contents"><b> Research groups. </b> <i> A </i> , at…</div>
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<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> Figure 4. </p>
</strong>
<div class="figure-caption-contents"><b> Research groups. </b> <i> A </i> , at MCV/VCU, circa 1983; Rob Beynon, Cheryl Garganta, Malcolm…</div>
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Figure 4.
</strong>
<div class="figure-caption-contents"><b>Research groups.</b>
<i>A</i>, at MCV/VCU, circa 1983; Rob Beynon, Cheryl Garganta, Malcolm McKay, Diane Rosin, Debbie Bellingham (holding a kidney-shaped cake), Dennis Unger, and Judith Bond. <i>B</i>, Virginia Tech and Penn State College of Medicine (PSUCOM) trainees circa 1994; <i>standing</i>: Weiping Jiang, Bruce Doll, Nicolina Stambolieva (visiting scientist), Petra Marchand, Judith Bond, Gary Johnson, and Jie Tang; <i>squatting in front</i>: Janet (Dietrich) Kumar and Jill Bankus. <i>C</i>, PSUCOM circa 2001; <i>standing</i>: Lourdes Norman (McKay), James Villa, Gail Matters, Jackie Crisman, Susan Ishmael, Xiaojing Wang, Jeremy Hengst, and Faoud Ishmael; <i>sitting</i>: Greg Bertenshaw, Judith Bond, and John Bylander. <i>D</i>, PSUCOM circa 2008; John Bylander, Judith Bond, a rotating graduate student, Jialing Bao, Moige Ongeri, Renee (Dusheck) Yura, Tim Keiffer, Sanjita Banerjee, Gail Matters, and Ge Jin.</div>
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<strong class="figure-label">
<p> Figure 5. </p>
</strong>
<div class="figure-caption-contents"><b> Secreted structures of meprin A… </b></div>
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<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> Figure 5. </p>
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<div class="figure-caption-contents"><b> Secreted structures of meprin A homooligomers. </b> Shown are electron micrographs of negatively stained…</div>
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Figure 5.
</strong>
<div class="figure-caption-contents"><b>Secreted structures of meprin A homooligomers.</b> Shown are electron micrographs of negatively stained forms of meprin A. <i>Row 1</i>, closed rings and crescents; <i>row 2</i>, tubes; <i>row 3</i>, spirals. (40).</div>
</figcaption>
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<strong class="figure-label">
<p> Figure 6. </p>
</strong>
<div class="figure-caption-contents"><b> Membrane-bound forms of meprin B.… </b></div>
</div>
</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> Figure 6. </p>
</strong>
<div class="figure-caption-contents"><b> Membrane-bound forms of meprin B. </b> Immunofluorescent localization of meprin β on the brush…</div>
</div>
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Figure 6.
</strong>
<div class="figure-caption-contents"><b>Membrane-bound forms of meprin B.</b> Immunofluorescent localization of meprin β on the brush border membrane of the mouse ileum. <i>Green</i>, meprin β; <i>blue</i>, nuclei (41).</div>
</figcaption>
</figure>
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<img class="figure-thumb" itemprop="thumbnail"
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<strong class="figure-label">
<p> Figure 7. </p>
</strong>
<div class="figure-caption-contents"><b> The I domain of meprin… </b></div>
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</div>
<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> Figure 7. </p>
</strong>
<div class="figure-caption-contents"><b> The I domain of meprin α </b> . Shown is a diagrammatic representation of…</div>
</div>
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Figure 7.
</strong>
<div class="figure-caption-contents"><b>The I domain of meprin α</b>. Shown is a diagrammatic representation of the biosynthesis of meprin subunits. Meprin α subunits (<i>red</i>) transfected into COS-1 cells were secreted (<i>A</i>), whereas meprin β subunits were transported to the cell surface and remained membrane-bound (<i>B</i>). The I domain of meprin α is indicated by the egg-shaped structure between the bulk of the protein (<i>rectangle</i> containing the pro-protease-MAM-TRAF domains) and the EGF-like-TM-cytoplasmic domains. <i>C</i>, if the I domain is deleted from meprin α, the protein does not leave the endoplasmic reticulum. <i>D</i>, if the I domain is inserted into the β transcript, then the proteolytically truncated β subunit is secreted into the medium. (20).</div>
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<strong class="figure-label">
<p> Figure 8. </p>
</strong>
<div class="figure-caption-contents"><b> Active sites of the meprin… </b></div>
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<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> Figure 8. </p>
</strong>
<div class="figure-caption-contents"><b> Active sites of the meprin </b> α <b> and </b> β <b> subunits. </b> Shown are homology models…</div>
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Figure 8.
</strong>
<div class="figure-caption-contents"><b>Active sites of the meprin</b> α <b>and</b> β <b>subunits.</b> Shown are homology models of the active-site clefts. The unprimed (<i>S</i>) and primed (<i>S</i>) substrate-binding regions are shown. <i>Yellow</i>, catalytic centers. <i>Blue</i> and <i>red</i>, basic and acidic amino acid residues, respectively. Key amino acid residues for determining substrate preferences are shown (46). Mutational analyses established the importance of the Lys-185 residue in the meprin β subunit for its preference to cleave substrates with negatively charged amino acids flanking the scissile bond and for the Tyr-199 residue in the meprin α subunit for its preference for small or hydrophobic amino acids flanking the scissile bond (47).</div>
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<p> Figure 9. </p>
</strong>
<div class="figure-caption-contents"><b> Centennial celebration of the ASBMB… </b></div>
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<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> Figure 9. </p>
</strong>
<div class="figure-caption-contents"><b> Centennial celebration of the ASBMB and the JBC in San Francisco in 2006.… </b></div>
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Figure 9.
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<div class="figure-caption-contents"><b>Centennial celebration of the ASBMB and the JBC in San Francisco in 2006.</b>
<i>A</i>, ASBMB group at the Louise M. Davies Symphony Hall, where a special selection of music was presented for meeting attendees. Shown from <i>left</i> to <i>right</i> are Gaylen Bradley, Bettie Sue Masters, George Carman, Heidi Hamm, Linda Roman, Alasdair Neale (conductor), Judith Bond, Martha Dennis, Ed Dennis, and Herb Tabor. Shown are Judith Bond with JBC Editor-in-Chief Herb Tabor (<i>B</i>), Associate Editor Dick Hanson (<i>C</i>), and Heidi Hamm and Bettie Sue Masters rehearsing for a presentation of “Three Little Presidents Are We” (<i>D</i>).</div>
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<p> Figure 10. </p>
</strong>
<div class="figure-caption-contents"><b> Award ceremony for Virginia's Outstanding… </b></div>
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<p> Figure 10. </p>
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<div class="figure-caption-contents"><b> Award ceremony for Virginia's Outstanding Scientists and Industrialists at the Science Museum of… </b></div>
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Figure 10.
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<div class="figure-caption-contents"><b>Award ceremony for Virginia's Outstanding Scientists and Industrialists at the Science Museum of Virginia in 1988 in Richmond, Virginia.</b></div>
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<p> Figure 11. </p>
</strong>
<div class="figure-caption-contents"><b> Plaque. </b> Given to me by…</div>
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<div class="figure-caption-medium figure-caption-text" aria-hidden="true">
<strong class="figure-label">
<p> Figure 11. </p>
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<div class="figure-caption-contents"><b> Plaque. </b> Given to me by an insightful STEP-UP student in 2010 at PSUCOM,…</div>
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Figure 11.
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<div class="figure-caption-contents"><b>Plaque.</b> Given to me by an insightful STEP-UP student in 2010 at PSUCOM, in Hershey, Pennsylvania.</div>
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