Serveur d'exploration Covid (26 mars)

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The E Protein Is a Multifunctional Membrane Protein of SARS-CoV

Identifieur interne : 000052 ( Ncbi/Merge ); précédent : 000051; suivant : 000053

The E Protein Is a Multifunctional Membrane Protein of SARS-CoV

Auteurs : Qingfa Wu [République populaire de Chine] ; Yilin Zhang [République populaire de Chine] ; Hong Lü [République populaire de Chine] ; Jing Wang [République populaire de Chine] ; Ximiao He [République populaire de Chine] ; Yong Liu [République populaire de Chine] ; Chen Ye [République populaire de Chine] ; Wei Lin [République populaire de Chine] ; Jianfei Hu [République populaire de Chine] ; Jia Ji [République populaire de Chine] ; Jing Xu [République populaire de Chine] ; Jia Ye [République populaire de Chine] ; Yongwu Hu [République populaire de Chine] ; Wenjun Chen [République populaire de Chine] ; Songgang Li [République populaire de Chine] ; Jun Wang [République populaire de Chine] ; Jian Wang [République populaire de Chine] ; Shengli Bi [République populaire de Chine] ; Huanming Yang [République populaire de Chine]

Source :

RBID : PMC:5172412

Abstract

The E (envelope) protein is the smallest structural protein in all coronaviruses and is the only viral structural protein in which no variation has been detected. We conducted genome sequencing and phylogenetic analyses of SARS-CoV. Based on genome sequencing, we predicted the E protein is a transmembrane (TM) protein characterized by a TM region with strong hydrophobicity and α-helix conformation. We identified a segment (NH2-_L-Cys-A-Y-Cys-Cys-N_-COOH) in the carboxyl-terminal region of the E protein that appears to form three disulfide bonds with another segment of corresponding cysteines in the carboxyl-terminus of the S (spike) protein. These bonds point to a possible structural association between the E and S proteins. Our phylogenetic analyses of the E protein sequences in all published coronaviruses place SARS-CoV in an independent group in Coronaviridae and suggest a non-human animal origin.


Url:
DOI: 10.1016/S1672-0229(03)01017-9
PubMed: 15626343
PubMed Central: 5172412

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PMC:5172412

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<nlm:aff id="aff0005">Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300, China</nlm:aff>
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<settlement type="city">Pékin</settlement>
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<affiliation wicri:level="4">
<nlm:aff id="aff0015">College of Life Sciences, Peking University, Beijing 100871, China</nlm:aff>
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<wicri:regionArea>College of Life Sciences, Peking University, Beijing 100871</wicri:regionArea>
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<settlement type="city">Pékin</settlement>
<region type="capitale">Pékin</region>
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</affiliation>
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<name sortKey="Ji, Jia" sort="Ji, Jia" uniqKey="Ji J" first="Jia" last="Ji">Jia Ji</name>
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<nlm:aff id="aff0005">Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300</wicri:regionArea>
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<settlement type="city">Pékin</settlement>
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<author>
<name sortKey="Xu, Jing" sort="Xu, Jing" uniqKey="Xu J" first="Jing" last="Xu">Jing Xu</name>
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<nlm:aff id="aff0005">Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300</wicri:regionArea>
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<settlement type="city">Pékin</settlement>
</placeName>
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</author>
<author>
<name sortKey="Ye, Jia" sort="Ye, Jia" uniqKey="Ye J" first="Jia" last="Ye">Jia Ye</name>
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<nlm:aff id="aff0005">Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300</wicri:regionArea>
<placeName>
<settlement type="city">Pékin</settlement>
</placeName>
</affiliation>
<affiliation wicri:level="4">
<nlm:aff id="aff0010">James D. Watson Institute of Genome Sciences, Zhijiang Campus, Zhejiang University, Hangzhou 310008, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>James D. Watson Institute of Genome Sciences, Zhijiang Campus, Zhejiang University, Hangzhou 310008</wicri:regionArea>
<placeName>
<settlement type="city">Hangzhou</settlement>
<region type="province">Zhejiang</region>
</placeName>
<orgName type="university">Université de Zhejiang</orgName>
<placeName>
<settlement type="city">Hangzhou</settlement>
<region type="province">Zhejiang</region>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Hu, Yongwu" sort="Hu, Yongwu" uniqKey="Hu Y" first="Yongwu" last="Hu">Yongwu Hu</name>
<affiliation wicri:level="1">
<nlm:aff id="aff0005">Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300</wicri:regionArea>
<placeName>
<settlement type="city">Pékin</settlement>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Chen, Wenjun" sort="Chen, Wenjun" uniqKey="Chen W" first="Wenjun" last="Chen">Wenjun Chen</name>
<affiliation wicri:level="1">
<nlm:aff id="aff0005">Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300</wicri:regionArea>
<placeName>
<settlement type="city">Pékin</settlement>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Li, Songgang" sort="Li, Songgang" uniqKey="Li S" first="Songgang" last="Li">Songgang Li</name>
<affiliation wicri:level="1">
<nlm:aff id="aff0005">Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300</wicri:regionArea>
<placeName>
<settlement type="city">Pékin</settlement>
</placeName>
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<affiliation wicri:level="4">
<nlm:aff id="aff0015">College of Life Sciences, Peking University, Beijing 100871, China</nlm:aff>
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<wicri:regionArea>College of Life Sciences, Peking University, Beijing 100871</wicri:regionArea>
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<settlement type="city">Pékin</settlement>
</placeName>
<orgName type="university">Université de Pékin</orgName>
<placeName>
<settlement type="city">Pékin</settlement>
<region type="capitale">Pékin</region>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Wang, Jun" sort="Wang, Jun" uniqKey="Wang J" first="Jun" last="Wang">Jun Wang</name>
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<nlm:aff id="aff0005">Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300</wicri:regionArea>
<placeName>
<settlement type="city">Pékin</settlement>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Wang, Jian" sort="Wang, Jian" uniqKey="Wang J" first="Jian" last="Wang">Jian Wang</name>
<affiliation wicri:level="1">
<nlm:aff id="aff0005">Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300</wicri:regionArea>
<placeName>
<settlement type="city">Pékin</settlement>
</placeName>
</affiliation>
<affiliation wicri:level="4">
<nlm:aff id="aff0010">James D. Watson Institute of Genome Sciences, Zhijiang Campus, Zhejiang University, Hangzhou 310008, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>James D. Watson Institute of Genome Sciences, Zhijiang Campus, Zhejiang University, Hangzhou 310008</wicri:regionArea>
<placeName>
<settlement type="city">Hangzhou</settlement>
<region type="province">Zhejiang</region>
</placeName>
<orgName type="university">Université de Zhejiang</orgName>
<placeName>
<settlement type="city">Hangzhou</settlement>
<region type="province">Zhejiang</region>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Bi, Shengli" sort="Bi, Shengli" uniqKey="Bi S" first="Shengli" last="Bi">Shengli Bi</name>
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<nlm:aff id="aff0025">Center of Disease Control and Prevention, Beijing 100050, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Center of Disease Control and Prevention, Beijing 100050</wicri:regionArea>
<placeName>
<settlement type="city">Pékin</settlement>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Yang, Huanming" sort="Yang, Huanming" uniqKey="Yang H" first="Huanming" last="Yang">Huanming Yang</name>
<affiliation wicri:level="1">
<nlm:aff id="aff0005">Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300</wicri:regionArea>
<placeName>
<settlement type="city">Pékin</settlement>
</placeName>
</affiliation>
<affiliation wicri:level="4">
<nlm:aff id="aff0010">James D. Watson Institute of Genome Sciences, Zhijiang Campus, Zhejiang University, Hangzhou 310008, China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>James D. Watson Institute of Genome Sciences, Zhijiang Campus, Zhejiang University, Hangzhou 310008</wicri:regionArea>
<placeName>
<settlement type="city">Hangzhou</settlement>
<region type="province">Zhejiang</region>
</placeName>
<orgName type="university">Université de Zhejiang</orgName>
<placeName>
<settlement type="city">Hangzhou</settlement>
<region type="province">Zhejiang</region>
</placeName>
</affiliation>
</author>
</analytic>
<series>
<title level="j">Genomics, Proteomics & Bioinformatics</title>
<idno type="ISSN">1672-0229</idno>
<idno type="eISSN">2210-3244</idno>
<imprint>
<date when="2003">2003</date>
</imprint>
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<front>
<div type="abstract" xml:lang="en">
<p>The E (envelope) protein is the smallest structural protein in all coronaviruses and is the only viral structural protein in which no variation has been detected. We conducted genome sequencing and phylogenetic analyses of SARS-CoV. Based on genome sequencing, we predicted the E protein is a transmembrane (TM) protein characterized by a TM region with strong hydrophobicity and
<italic>α</italic>
-helix conformation. We identified a segment (NH
<sub>2</sub>
-_L-Cys-A-Y-Cys-Cys-N_-COOH) in the carboxyl-terminal region of the E protein that appears to form three disulfide bonds with another segment of corresponding cysteines in the carboxyl-terminus of the S (spike) protein. These bonds point to a possible structural association between the E and S proteins. Our phylogenetic analyses of the E protein sequences in all published coronaviruses place SARS-CoV in an independent group in
<italic>Coronaviridae</italic>
and suggest a non-human animal origin.</p>
</div>
</front>
<back>
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</TEI>
<pmc article-type="research-article">
<pmc-dir>properties open_access</pmc-dir>
<front>
<journal-meta>
<journal-id journal-id-type="nlm-ta">Genomics Proteomics Bioinformatics</journal-id>
<journal-id journal-id-type="iso-abbrev">Genomics Proteomics Bioinformatics</journal-id>
<journal-title-group>
<journal-title>Genomics, Proteomics & Bioinformatics</journal-title>
</journal-title-group>
<issn pub-type="ppub">1672-0229</issn>
<issn pub-type="epub">2210-3244</issn>
<publisher>
<publisher-name>Elsevier</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="pmid">15626343</article-id>
<article-id pub-id-type="pmc">5172412</article-id>
<article-id pub-id-type="publisher-id">S1672-0229(03)01017-9</article-id>
<article-id pub-id-type="doi">10.1016/S1672-0229(03)01017-9</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Invited Article</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>The E Protein Is a Multifunctional Membrane Protein of SARS-CoV</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Wu</surname>
<given-names>Qingfa</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
<xref rid="aff0010" ref-type="aff">2</xref>
<xref rid="fn1" ref-type="fn">*</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Zhang</surname>
<given-names>Yilin</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
<xref rid="fn1" ref-type="fn">*</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname></surname>
<given-names>Hong</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
<xref rid="fn1" ref-type="fn">*</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wang</surname>
<given-names>Jing</given-names>
</name>
<xref rid="aff0015" ref-type="aff">3</xref>
<xref rid="aff0005" ref-type="aff">1</xref>
<xref rid="fn1" ref-type="fn">*</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>He</surname>
<given-names>Ximiao</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Liu</surname>
<given-names>Yong</given-names>
</name>
<xref rid="aff0020" ref-type="aff">4</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ye</surname>
<given-names>Chen</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Lin</surname>
<given-names>Wei</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Hu</surname>
<given-names>Jianfei</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
<xref rid="aff0015" ref-type="aff">3</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ji</surname>
<given-names>Jia</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Xu</surname>
<given-names>Jing</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ye</surname>
<given-names>Jia</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
<xref rid="aff0010" ref-type="aff">2</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Hu</surname>
<given-names>Yongwu</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Chen</surname>
<given-names>Wenjun</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Li</surname>
<given-names>Songgang</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
<xref rid="aff0015" ref-type="aff">3</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wang</surname>
<given-names>Jun</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wang</surname>
<given-names>Jian</given-names>
</name>
<xref rid="aff0005" ref-type="aff">1</xref>
<xref rid="aff0010" ref-type="aff">2</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Bi</surname>
<given-names>Shengli</given-names>
</name>
<xref rid="aff0025" ref-type="aff">5</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Yang</surname>
<given-names>Huanming</given-names>
</name>
<email>yanghm@genomics.org.cn</email>
<xref rid="aff0005" ref-type="aff">1</xref>
<xref rid="aff0010" ref-type="aff">2</xref>
<xref rid="cor1" ref-type="corresp">#</xref>
</contrib>
</contrib-group>
<aff id="aff0005">
<label>1</label>
Beijing Genomics Institute, Chinese Academy of Sciences, Beijing 101300, China</aff>
<aff id="aff0010">
<label>2</label>
James D. Watson Institute of Genome Sciences, Zhijiang Campus, Zhejiang University, Hangzhou 310008, China</aff>
<aff id="aff0015">
<label>3</label>
College of Life Sciences, Peking University, Beijing 100871, China</aff>
<aff id="aff0020">
<label>4</label>
Peking Union Medical College Hospital, Chinese Academy of Medical Sciences, Beijing 100730, China</aff>
<aff id="aff0025">
<label>5</label>
Center of Disease Control and Prevention, Beijing 100050, China</aff>
<author-notes>
<corresp id="cor1">
<label>#</label>
Corresponding author.
<email>yanghm@genomics.org.cn</email>
</corresp>
<fn id="fn1">
<label>*</label>
<p id="ntp0005">These authors contributed equally to this work.</p>
</fn>
</author-notes>
<pub-date pub-type="pmc-release">
<day>28</day>
<month>11</month>
<year>2016</year>
</pub-date>
<pmc-comment> PMC Release delay is 0 months and 0 days and was based on .</pmc-comment>
<pub-date pub-type="ppub">
<month>5</month>
<year>2003</year>
</pub-date>
<pub-date pub-type="epub">
<day>28</day>
<month>11</month>
<year>2016</year>
</pub-date>
<volume>1</volume>
<issue>2</issue>
<fpage>131</fpage>
<lpage>144</lpage>
<permissions>
<copyright-statement>.</copyright-statement>
<copyright-year>2003</copyright-year>
<copyright-holder>Beijing Institute of Genomics, the Chinese Academy of Sciences and the Genetics Society of China</copyright-holder>
<license license-type="CC BY-NC-ND" xlink:href="http://creativecommons.org/licenses/by-nc-nd/4.0/">
<license-p>This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).</license-p>
</license>
</permissions>
<abstract id="ab0005">
<p>The E (envelope) protein is the smallest structural protein in all coronaviruses and is the only viral structural protein in which no variation has been detected. We conducted genome sequencing and phylogenetic analyses of SARS-CoV. Based on genome sequencing, we predicted the E protein is a transmembrane (TM) protein characterized by a TM region with strong hydrophobicity and
<italic>α</italic>
-helix conformation. We identified a segment (NH
<sub>2</sub>
-_L-Cys-A-Y-Cys-Cys-N_-COOH) in the carboxyl-terminal region of the E protein that appears to form three disulfide bonds with another segment of corresponding cysteines in the carboxyl-terminus of the S (spike) protein. These bonds point to a possible structural association between the E and S proteins. Our phylogenetic analyses of the E protein sequences in all published coronaviruses place SARS-CoV in an independent group in
<italic>Coronaviridae</italic>
and suggest a non-human animal origin.</p>
</abstract>
<kwd-group id="keys0005">
<title>Key words</title>
<kwd>SARS</kwd>
<kwd>SARS-CoV</kwd>
<kwd>the E protein</kwd>
<kwd>envelope</kwd>
<kwd>TM region</kwd>
</kwd-group>
</article-meta>
</front>
<floats-group>
<fig id="f0005">
<label>Fig. 1</label>
<caption>
<p>The predicted TM region in the E protein of SARS-CoV by TMHMM.</p>
</caption>
<alt-text id="at0005">Fig. 1</alt-text>
<graphic xlink:href="gr1"></graphic>
</fig>
<fig id="f0010">
<label>Fig. 2</label>
<caption>
<p>Predicted secondary structures in the E protein of SARS-CoV. Software programs: I. PSIpred; II. NNPredict; III. SPLIT; IV. 1-3. Antheprot 5.0.</p>
</caption>
<alt-text id="at0010">Fig. 2</alt-text>
<graphic xlink:href="gr2"></graphic>
</fig>
<fig id="f0015">
<label>Fig. 3-I, 3-II, 3-III</label>
<caption>
<p>Predicted TM regions in the E protein of the three groups of coronaviruses.</p>
</caption>
<alt-text id="at0015">Fig. 3</alt-text>
<graphic xlink:href="gr3"></graphic>
</fig>
<fig id="f0020">
<label>Fig. 4-I, 4-II, 4-III</label>
<caption>
<p>The distribution of GC content (I), charge (II) and hydrophobicity (III) in the E protein of SARS-CoV.</p>
</caption>
<alt-text id="at0020">Fig. 4</alt-text>
<graphic xlink:href="gr4"></graphic>
</fig>
<fig id="f0025">
<label>Fig. 5</label>
<caption>
<p>Multi-alignment of amino acid sequences of the E protein between SARS-CoV and other coronaviruses. AIBV: avian infectious bronchitis virus; BCoV: bovine coronavirus; CcoV: canine coronavirus; FCoV: feline coronavirus; HCoV-229E: human coronavirus 229E; HCoV-OC43: human coronavirus OC43; MHV: murine hepatitis virus; PEDV: porcine epidemic diarrhea virus; PHEV: porcine hemagglutinating encephalomyelitis virus; RCoV: rat coronavirus; TCoV: turkey coronavirus; TGV: transmissible gastroenteritis virus.</p>
</caption>
<alt-text id="at0025">Fig. 5</alt-text>
<graphic xlink:href="gr5"></graphic>
</fig>
<fig id="f0030">
<label>Fig. 6</label>
<caption>
<p>Pairwise similarity between coronaviruses based on the E protein. The numbers indicate the percentage of similarity between each listed coronavirus and BJ01. (HCoV-E: HCoV-229E; HCoV-O: HCoV-OC43.)</p>
</caption>
<alt-text id="at0030">Fig. 6</alt-text>
<graphic xlink:href="gr6"></graphic>
</fig>
<fig id="f0035">
<label>Fig. 7</label>
<caption>
<p>An unrooted phylogenetic tree of the coronaviruses based on the amino acid sequence of the E protein by ClustalW and TreeView.</p>
</caption>
<alt-text id="at0035">Fig. 7</alt-text>
<graphic xlink:href="gr7"></graphic>
</fig>
<fig id="f0040">
<label>Fig. 8</label>
<caption>
<p>The predicted disulfide bonds between the E and S proteins of SARS-CoV.</p>
</caption>
<alt-text id="at0040">Fig. 8</alt-text>
<graphic xlink:href="gr8"></graphic>
</fig>
<table-wrap id="t0005" position="float">
<label>Table 1</label>
<caption>
<p>The Genomic and Biochemical Features of the Entire E Protein and Its Three Subregions</p>
</caption>
<alt-text id="at0045">Table 1</alt-text>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th></th>
<th align="center">TM region</th>
<th align="center">N-terminus</th>
<th align="center">C-terminus</th>
<th align="center">E protein</th>
</tr>
</thead>
<tbody>
<tr>
<td align="center">G+C (%)</td>
<td align="center">42.0</td>
<td align="center">35.7</td>
<td align="center">40.8</td>
<td align="center">40.2</td>
</tr>
<tr>
<td align="center">A</td>
<td align="center">11 (15.9%)</td>
<td align="center">13 (31.0%)</td>
<td align="center">30 (25.0%)</td>
<td align="center">54 (23.4%)</td>
</tr>
<tr>
<td align="center">U</td>
<td align="center">29 (42.1%)</td>
<td align="center">14 (33.3%)</td>
<td align="center">41 (34.2%)</td>
<td align="center">84 (36.4%)</td>
</tr>
<tr>
<td align="center">C</td>
<td align="center">18 (26.1%)</td>
<td align="center">6 (14.3%)</td>
<td align="center">23 (19.2%)</td>
<td align="center">47 (20.3%)</td>
</tr>
<tr>
<td align="center">G</td>
<td align="center">11 (15.9%)</td>
<td align="center">9 (21.4%)</td>
<td align="center">26 (21.6%)</td>
<td align="center">46 (19.9%)</td>
</tr>
<tr>
<td align="center">Total (nt)</td>
<td align="center">69 (100%)</td>
<td align="center">42 (100%)</td>
<td align="center">120 (100%)</td>
<td align="center">231 (100%)</td>
</tr>
<tr>
<td></td>
<td></td>
<td></td>
<td></td>
<td></td>
</tr>
<tr>
<td align="center">Leu</td>
<td align="center">8 (10.6)</td>
<td align="center">1 (1.3)</td>
<td align="center">5 (6.6)</td>
<td align="center">14 (18.4)</td>
</tr>
<tr>
<td align="center">Val</td>
<td align="center">4 (5.3)</td>
<td align="center">2 (2.6)</td>
<td align="center">8 (10.6)</td>
<td align="center">14 (18.4)</td>
</tr>
<tr>
<td align="center">Phe</td>
<td align="center">3 (3.9)</td>
<td align="center">1 (1.3)</td>
<td align="center">0</td>
<td align="center">4 (5.3)</td>
</tr>
<tr>
<td align="center">Ala</td>
<td align="center">3 (3.9)</td>
<td align="center">0</td>
<td align="center">1 (1.3)</td>
<td align="center">4 (5.3)</td>
</tr>
<tr>
<td align="center">Thr</td>
<td align="center">2 (2.7)</td>
<td align="center">2 (2.6)</td>
<td align="center">1 (1.3)</td>
<td align="center">5 (6.6)</td>
</tr>
<tr>
<td align="center">Ile</td>
<td align="center">1 (1.3)</td>
<td align="center">1 (1.3)</td>
<td align="center">1 (1.3)</td>
<td align="center">3 (3.9)</td>
</tr>
<tr>
<td align="center">Asn</td>
<td align="center">1 (1.3)</td>
<td align="center">0</td>
<td align="center">4 (5.3)</td>
<td align="center">5 (6.6)</td>
</tr>
<tr>
<td align="center">Ser</td>
<td align="center">1 (1.3)</td>
<td align="center">2 (2.6)</td>
<td align="center">4 (5.3)</td>
<td align="center">7 (9.2)</td>
</tr>
<tr>
<td align="center">Glu</td>
<td align="center">0</td>
<td align="center">2 (2.6)</td>
<td align="center">1 (1.3)</td>
<td align="center">3 (3.9)</td>
</tr>
<tr>
<td align="center">Tyr</td>
<td align="center">0</td>
<td align="center">1 (1.3)</td>
<td align="center">3 (3.9)</td>
<td align="center">4 (5.3)</td>
</tr>
<tr>
<td align="center">Gly</td>
<td align="center">0</td>
<td align="center">1 (1.3)</td>
<td align="center">1 (1.3)</td>
<td align="center">2 (2.6)</td>
</tr>
<tr>
<td align="center">Met</td>
<td align="center">0</td>
<td align="center">1(1.3)</td>
<td align="center">0</td>
<td align="center">1(1.3)</td>
</tr>
<tr>
<td align="center">Cys</td>
<td align="center">0</td>
<td align="center">0</td>
<td align="center">3 (3.9)</td>
<td align="center">3 (3.9)</td>
</tr>
<tr>
<td align="center">Lys</td>
<td align="center">0</td>
<td align="center">0</td>
<td align="center">2 (2.6)</td>
<td align="center">2 (2.6)</td>
</tr>
<tr>
<td align="center">Arg</td>
<td align="center">0</td>
<td align="center">0</td>
<td align="center">2 (2.6)</td>
<td align="center">2 (2.6)</td>
</tr>
<tr>
<td align="center">Pro</td>
<td align="center">0</td>
<td align="center">0</td>
<td align="center">2 (2.6)</td>
<td align="center">2 (2.6)</td>
</tr>
<tr>
<td align="center">Asp</td>
<td align="center">0</td>
<td align="center">0</td>
<td align="center">1 (1.3)</td>
<td align="center">1 (1.3)</td>
</tr>
<tr>
<td align="center">Total</td>
<td align="center">23 (30.3)</td>
<td align="center">14 (18.4)</td>
<td align="center">39 (51.3)</td>
<td align="center">76 (100)</td>
</tr>
<tr>
<td></td>
<td></td>
<td></td>
<td></td>
<td></td>
</tr>
<tr>
<td align="center">Molecular Weight (a.a.)</td>
<td align="center">2491</td>
<td align="center">1576</td>
<td align="center">4330</td>
<td align="center">8361</td>
</tr>
<tr>
<td align="center">pI</td>
<td align="center">5.52</td>
<td align="center">3.79</td>
<td align="center">8.61</td>
<td align="center">6.01</td>
</tr>
<tr>
<td align="center">Net Charge</td>
<td align="center">0</td>
<td align="center">-2</td>
<td align="center">+2</td>
<td align="center">0</td>
</tr>
<tr>
<td></td>
<td></td>
<td align="center">(-2.7%)</td>
<td align="center">(-2.7%, +5.4%)</td>
<td align="center">(-5.4%, +5.4%)</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="t0010" position="float">
<label>Table 2</label>
<caption>
<p>Codon Usage Frequency of the TM Region and the Entire E Protein</p>
</caption>
<alt-text id="at0050">Table 2</alt-text>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th>a. a.</th>
<th>TM region</th>
<th>E protein</th>
<th>Codon</th>
<th>TM region</th>
<th>E protein</th>
</tr>
</thead>
<tbody>
<tr>
<td rowspan="4" valign="middle">Ala
<hr></hr>
</td>
<td rowspan="4" valign="middle">3 (4.00%)
<hr></hr>
</td>
<td rowspan="4" valign="middle">4 (5.33%)
<hr></hr>
</td>
<td>GCA</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>GCC</td>
<td>1 (1.33%)</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td>GCG</td>
<td>1 (1.33%)</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td>GCU</td>
<td>1 (1.33%)</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="2" valign="middle">Cys
<hr></hr>
</td>
<td rowspan="2" valign="middle">0
<hr></hr>
</td>
<td rowspan="2" valign="middle">3 (4.00%)
<hr></hr>
</td>
<td>UGC</td>
<td>0</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td>UGU</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="2" valign="middle">Asp
<hr></hr>
</td>
<td rowspan="2" valign="middle">0
<hr></hr>
</td>
<td rowspan="2" valign="middle">1 (1.33%)
<hr></hr>
</td>
<td>GAC</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>GAU</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="2" valign="middle">Glu
<hr></hr>
</td>
<td rowspan="2" valign="middle">0
<hr></hr>
</td>
<td rowspan="2" valign="middle">3 (4.00%)
<hr></hr>
</td>
<td>GAA</td>
<td>0</td>
<td>3 (4.00%)</td>
</tr>
<tr>
<td>GAG</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="2" valign="middle">Phe
<hr></hr>
</td>
<td rowspan="2" valign="middle">3 (4.00%)
<hr></hr>
</td>
<td rowspan="2" valign="middle">4 (5.33%)
<hr></hr>
</td>
<td>UUC</td>
<td>2 (2.67%)</td>
<td>3 (4.00%)</td>
</tr>
<tr>
<td>UUU</td>
<td>1 (1.33%)</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="4" valign="middle">Gly
<hr></hr>
</td>
<td rowspan="4" valign="middle">0
<hr></hr>
</td>
<td rowspan="4" valign="middle">2 (2.67%)
<hr></hr>
</td>
<td>GGA</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td>GGC</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>GGG</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>GGU</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="2" valign="middle">His
<hr></hr>
</td>
<td rowspan="2" valign="middle">0
<hr></hr>
</td>
<td rowspan="2" valign="middle">0
<hr></hr>
</td>
<td>CAC</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>CAU</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="3" valign="middle">Ile
<hr></hr>
</td>
<td rowspan="3" valign="middle">1 (1.33%)
<hr></hr>
</td>
<td rowspan="3" valign="middle">3 (4.00%)
<hr></hr>
</td>
<td>AUA</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td>AUC</td>
<td>1 (1.33%)</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td>AUU</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="2" valign="middle">Lys
<hr></hr>
</td>
<td rowspan="2" valign="middle">0
<hr></hr>
</td>
<td rowspan="2" valign="middle">2 (2.67%)
<hr></hr>
</td>
<td>AAA</td>
<td>0</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td>AAG</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="6" valign="middle">Leu
<hr></hr>
</td>
<td rowspan="6" valign="middle">8 (10.67%)
<hr></hr>
</td>
<td rowspan="6" valign="middle">14 (18.67%)
<hr></hr>
</td>
<td>CUA</td>
<td>2 (2.67%)</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td>CUC</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>CUG</td>
<td>0</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td>CUU</td>
<td>5 (6.67%)</td>
<td>6 (8.00%)</td>
</tr>
<tr>
<td>UUA</td>
<td>0</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td>UUG</td>
<td>1 (1.33%)</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td>Met</td>
<td>0</td>
<td>1 (1.33%)</td>
<td>AUG</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="2" valign="middle">Asn
<hr></hr>
</td>
<td rowspan="2" valign="middle">1 (1.33%)
<hr></hr>
</td>
<td rowspan="2" valign="middle">5 (6.67%)
<hr></hr>
</td>
<td>AAC</td>
<td>0</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td>AAU</td>
<td>1 (1.33%)</td>
<td>3 (4.00%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="4" valign="middle">Pro
<hr></hr>
</td>
<td rowspan="4" valign="middle">0
<hr></hr>
</td>
<td rowspan="4" valign="middle">2 (2.67%)
<hr></hr>
</td>
<td>CCA</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td>CCC</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>CCG</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>CCU</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="2" valign="middle">Gln
<hr></hr>
</td>
<td rowspan="2" valign="middle">0
<hr></hr>
</td>
<td rowspan="2" valign="middle">0
<hr></hr>
</td>
<td>CAA</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>CAG</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="6" valign="middle">Arg
<hr></hr>
</td>
<td rowspan="6" valign="middle">0
<hr></hr>
</td>
<td rowspan="6" valign="middle">2 (2.67%)
<hr></hr>
</td>
<td>AGA</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>AGG</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>CGA</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td>CGC</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>CGG</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>CGU</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="6" valign="middle">Ser
<hr></hr>
</td>
<td rowspan="6" valign="middle">1 (1.33%)
<hr></hr>
</td>
<td rowspan="6" valign="middle">7 (9.33%)
<hr></hr>
</td>
<td>AGC</td>
<td>1 (1.33%)</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td>AGU</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td>UCA</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td>UCC</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>UCG</td>
<td>0</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td>UCU</td>
<td>0</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="4" valign="middle">Thr
<hr></hr>
</td>
<td rowspan="4" valign="middle">2 (2.67%)
<hr></hr>
</td>
<td rowspan="4" valign="middle">5 (6.67%)
<hr></hr>
</td>
<td>ACA</td>
<td>1 (1.33%)</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td>ACC</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>ACG</td>
<td>0</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td>ACU</td>
<td>1 (1.33%)</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="4" valign="middle">Val
<hr></hr>
</td>
<td rowspan="4" valign="middle">4 (5.33%)
<hr></hr>
</td>
<td rowspan="4" valign="middle">14 (18.67%) 0
<hr></hr>
</td>
<td>GUA</td>
<td>2 (2.67%)</td>
<td>3 (4.00%)</td>
</tr>
<tr>
<td>GUC</td>
<td>1 (1.33%)</td>
<td>3 (4.00%)</td>
</tr>
<tr>
<td>GUG</td>
<td>1 (1.33%)</td>
<td>2 (2.67%)</td>
</tr>
<tr>
<td>GUU</td>
<td>0</td>
<td>6 (8.00%)</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td>Trp</td>
<td>0</td>
<td>0</td>
<td>UGG</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="2" valign="middle">Tyr
<hr></hr>
</td>
<td rowspan="2" valign="middle">0
<hr></hr>
</td>
<td rowspan="2" valign="middle">4 (5.33%)
<hr></hr>
</td>
<td>UAC</td>
<td>0</td>
<td>4 (5.33%)</td>
</tr>
<tr>
<td>UAU</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td colspan="6">
<hr></hr>
</td>
</tr>
<tr>
<td rowspan="3" valign="middle">STOP</td>
<td rowspan="3" valign="middle">0</td>
<td rowspan="3" valign="middle">1 (1.33%)</td>
<td>UAA</td>
<td>0</td>
<td>1 (1.33%)</td>
</tr>
<tr>
<td>UAG</td>
<td>0</td>
<td>0</td>
</tr>
<tr>
<td>UGA</td>
<td>0</td>
<td>0</td>
</tr>
</tbody>
</table>
</table-wrap>
</floats-group>
</pmc>
<affiliations>
<list>
<country>
<li>République populaire de Chine</li>
</country>
<region>
<li>Pékin</li>
<li>Zhejiang</li>
</region>
<settlement>
<li>Hangzhou</li>
<li>Pékin</li>
</settlement>
<orgName>
<li>Université de Pékin</li>
<li>Université de Zhejiang</li>
</orgName>
</list>
<tree>
<country name="République populaire de Chine">
<noRegion>
<name sortKey="Wu, Qingfa" sort="Wu, Qingfa" uniqKey="Wu Q" first="Qingfa" last="Wu">Qingfa Wu</name>
</noRegion>
<name sortKey="Bi, Shengli" sort="Bi, Shengli" uniqKey="Bi S" first="Shengli" last="Bi">Shengli Bi</name>
<name sortKey="Chen, Wenjun" sort="Chen, Wenjun" uniqKey="Chen W" first="Wenjun" last="Chen">Wenjun Chen</name>
<name sortKey="He, Ximiao" sort="He, Ximiao" uniqKey="He X" first="Ximiao" last="He">Ximiao He</name>
<name sortKey="Hu, Jianfei" sort="Hu, Jianfei" uniqKey="Hu J" first="Jianfei" last="Hu">Jianfei Hu</name>
<name sortKey="Hu, Jianfei" sort="Hu, Jianfei" uniqKey="Hu J" first="Jianfei" last="Hu">Jianfei Hu</name>
<name sortKey="Hu, Yongwu" sort="Hu, Yongwu" uniqKey="Hu Y" first="Yongwu" last="Hu">Yongwu Hu</name>
<name sortKey="Ji, Jia" sort="Ji, Jia" uniqKey="Ji J" first="Jia" last="Ji">Jia Ji</name>
<name sortKey="Li, Songgang" sort="Li, Songgang" uniqKey="Li S" first="Songgang" last="Li">Songgang Li</name>
<name sortKey="Li, Songgang" sort="Li, Songgang" uniqKey="Li S" first="Songgang" last="Li">Songgang Li</name>
<name sortKey="Lin, Wei" sort="Lin, Wei" uniqKey="Lin W" first="Wei" last="Lin">Wei Lin</name>
<name sortKey="Liu, Yong" sort="Liu, Yong" uniqKey="Liu Y" first="Yong" last="Liu">Yong Liu</name>
<name sortKey="Lu, Hong" sort="Lu, Hong" uniqKey="Lu H" first="Hong" last="Lü">Hong Lü</name>
<name sortKey="Wang, Jian" sort="Wang, Jian" uniqKey="Wang J" first="Jian" last="Wang">Jian Wang</name>
<name sortKey="Wang, Jian" sort="Wang, Jian" uniqKey="Wang J" first="Jian" last="Wang">Jian Wang</name>
<name sortKey="Wang, Jing" sort="Wang, Jing" uniqKey="Wang J" first="Jing" last="Wang">Jing Wang</name>
<name sortKey="Wang, Jing" sort="Wang, Jing" uniqKey="Wang J" first="Jing" last="Wang">Jing Wang</name>
<name sortKey="Wang, Jun" sort="Wang, Jun" uniqKey="Wang J" first="Jun" last="Wang">Jun Wang</name>
<name sortKey="Wu, Qingfa" sort="Wu, Qingfa" uniqKey="Wu Q" first="Qingfa" last="Wu">Qingfa Wu</name>
<name sortKey="Xu, Jing" sort="Xu, Jing" uniqKey="Xu J" first="Jing" last="Xu">Jing Xu</name>
<name sortKey="Yang, Huanming" sort="Yang, Huanming" uniqKey="Yang H" first="Huanming" last="Yang">Huanming Yang</name>
<name sortKey="Yang, Huanming" sort="Yang, Huanming" uniqKey="Yang H" first="Huanming" last="Yang">Huanming Yang</name>
<name sortKey="Ye, Chen" sort="Ye, Chen" uniqKey="Ye C" first="Chen" last="Ye">Chen Ye</name>
<name sortKey="Ye, Jia" sort="Ye, Jia" uniqKey="Ye J" first="Jia" last="Ye">Jia Ye</name>
<name sortKey="Ye, Jia" sort="Ye, Jia" uniqKey="Ye J" first="Jia" last="Ye">Jia Ye</name>
<name sortKey="Zhang, Yilin" sort="Zhang, Yilin" uniqKey="Zhang Y" first="Yilin" last="Zhang">Yilin Zhang</name>
</country>
</tree>
</affiliations>
</record>

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