An RXLR effector secreted by Phytophthora parasitica is a virulence factor and triggers cell death in various plants.
Identifieur interne : 000583 ( Main/Exploration ); précédent : 000582; suivant : 000584An RXLR effector secreted by Phytophthora parasitica is a virulence factor and triggers cell death in various plants.
Auteurs : Guiyan Huang [République populaire de Chine] ; Zhirou Liu [République populaire de Chine] ; Biao Gu [République populaire de Chine] ; Hong Zhao [République populaire de Chine] ; Jinbu Jia [République populaire de Chine] ; Guangjin Fan [République populaire de Chine] ; Yuling Meng [République populaire de Chine] ; Yu Du [République populaire de Chine] ; Weixing Shan [République populaire de Chine]Source :
- Molecular plant pathology [ 1364-3703 ] ; 2019.
Descripteurs français
- KwdFr :
- MESH :
- microbiologie : Maladies des plantes, Tabac.
- pathogénicité : Phytophthora infestans.
- Virulence.
English descriptors
- KwdEn :
- MESH :
- microbiology : Plant Diseases, Tobacco.
- pathogenicity : Phytophthora infestans.
- Virulence.
Abstract
RXLR effectors encoded by Phytophthora species play a central role in pathogen-plant interactions. An understanding of the biological functions of RXLR effectors is conducive to the illumination of the pathogenic mechanisms and the development of disease control strategies. However, the virulence function of Phytophthora parasitica RXLR effectors is poorly understood. Here, we describe the identification of a P. parasitica RXLR effector gene, PPTG00121 (PpE4), which is highly transcribed during the early stages of infection. Live cell imaging of P. parasitica transformants expressing a full-length PpE4 (E4FL)-mCherry protein indicated that PpE4 is secreted and accumulates around haustoria during plant infection. Silencing of PpE4 in P. parasitica resulted in significantly reduced virulence on Nicotiana benthamiana. Transient expression of PpE4 in N. benthamiana in turn restored the pathogenicity of the PpE4-silenced lines. Furthermore, the expression of PpE4 in both N. benthamiana and Arabidopsis thaliana consistently enhanced plant susceptibility to P. parasitica. These results indicate that PpE4 contributes to pathogen infection. Finally, heterologous expression experiments showed that PpE4 triggers non-specific cell death in a variety of plants, including tobacco, tomato, potato and A. thaliana. Virus-induced gene silencing assays revealed that PpE4-induced cell death is dependent on HSP90, NPK and SGT1, suggesting that PpE4 is recognized by the plant immune system. In conclusion, PpE4 is an important virulence RXLR effector of P. parasitica and recognized by a wide range of host plants.
DOI: 10.1111/mpp.12760
PubMed: 30320960
PubMed Central: PMC6637884
Affiliations:
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populaire de Chine</country><wicri:regionArea>College of Life Sciences, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation></author><author><name sortKey="Liu, Zhirou" sort="Liu, Zhirou" uniqKey="Liu Z" first="Zhirou" last="Liu">Zhirou Liu</name><affiliation wicri:level="1"><nlm:affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation><affiliation wicri:level="1"><nlm:affiliation>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation></author><author><name sortKey="Gu, Biao" sort="Gu, Biao" uniqKey="Gu B" first="Biao" last="Gu">Biao Gu</name><affiliation wicri:level="1"><nlm:affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation><affiliation wicri:level="1"><nlm:affiliation>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation></author><author><name sortKey="Zhao, Hong" sort="Zhao, Hong" uniqKey="Zhao H" first="Hong" last="Zhao">Hong Zhao</name><affiliation wicri:level="1"><nlm:affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation><affiliation wicri:level="1"><nlm:affiliation>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation></author><author><name sortKey="Jia, Jinbu" sort="Jia, Jinbu" uniqKey="Jia J" first="Jinbu" last="Jia">Jinbu Jia</name><affiliation wicri:level="1"><nlm:affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation><affiliation wicri:level="1"><nlm:affiliation>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation><affiliation wicri:level="1"><nlm:affiliation>Institute of Plant and Food Science, Department of Biology, Southern University of Science and Technology, Shenzhen, 518055, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>Institute of Plant and Food Science, Department of Biology, Southern University of Science and Technology, Shenzhen, 518055</wicri:regionArea><wicri:noRegion>518055</wicri:noRegion></affiliation></author><author><name sortKey="Fan, Guangjin" sort="Fan, Guangjin" uniqKey="Fan G" first="Guangjin" last="Fan">Guangjin Fan</name><affiliation wicri:level="1"><nlm:affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation><affiliation wicri:level="1"><nlm:affiliation>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation></author><author><name sortKey="Meng, Yuling" sort="Meng, Yuling" uniqKey="Meng Y" first="Yuling" last="Meng">Yuling Meng</name><affiliation wicri:level="1"><nlm:affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation><affiliation wicri:level="1"><nlm:affiliation>College of Agronomy, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Agronomy, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation></author><author><name sortKey="Du, Yu" sort="Du, Yu" uniqKey="Du Y" first="Yu" last="Du">Yu Du</name><affiliation wicri:level="1"><nlm:affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation><affiliation wicri:level="1"><nlm:affiliation>College of Horticulture, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Horticulture, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation></author><author><name sortKey="Shan, Weixing" sort="Shan, Weixing" uniqKey="Shan W" first="Weixing" last="Shan">Weixing Shan</name><affiliation wicri:level="1"><nlm:affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation><affiliation wicri:level="1"><nlm:affiliation>College of Agronomy, Northwest A&F University, Yangling, Shaanxi, 712100, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>College of Agronomy, Northwest A&F University, Yangling, Shaanxi, 712100</wicri:regionArea><wicri:noRegion>712100</wicri:noRegion></affiliation></author></analytic><series><title level="j">Molecular plant pathology</title><idno type="eISSN">1364-3703</idno><imprint><date when="2019" type="published">2019</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass><keywords scheme="KwdEn" xml:lang="en"><term>Phytophthora infestans (pathogenicity)</term><term>Plant Diseases (microbiology)</term><term>Tobacco (microbiology)</term><term>Virulence (MeSH)</term></keywords><keywords scheme="KwdFr" xml:lang="fr"><term>Maladies des plantes (microbiologie)</term><term>Phytophthora infestans (pathogénicité)</term><term>Tabac (microbiologie)</term><term>Virulence (MeSH)</term></keywords><keywords scheme="MESH" qualifier="microbiologie" xml:lang="fr"><term>Maladies des plantes</term><term>Tabac</term></keywords><keywords scheme="MESH" qualifier="microbiology" xml:lang="en"><term>Plant Diseases</term><term>Tobacco</term></keywords><keywords scheme="MESH" qualifier="pathogenicity" xml:lang="en"><term>Phytophthora infestans</term></keywords><keywords scheme="MESH" qualifier="pathogénicité" xml:lang="fr"><term>Phytophthora infestans</term></keywords><keywords scheme="MESH" xml:lang="en"><term>Virulence</term></keywords><keywords scheme="MESH" xml:lang="fr"><term>Virulence</term></keywords></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en">RXLR effectors encoded by Phytophthora species play a central role in pathogen-plant interactions. An understanding of the biological functions of RXLR effectors is conducive to the illumination of the pathogenic mechanisms and the development of disease control strategies. However, the virulence function of Phytophthora parasitica RXLR effectors is poorly understood. Here, we describe the identification of a P. parasitica RXLR effector gene, PPTG00121 (PpE4), which is highly transcribed during the early stages of infection. Live cell imaging of P. parasitica transformants expressing a full-length PpE4 (E4FL)-mCherry protein indicated that PpE4 is secreted and accumulates around haustoria during plant infection. Silencing of PpE4 in P. parasitica resulted in significantly reduced virulence on Nicotiana benthamiana. Transient expression of PpE4 in N. benthamiana in turn restored the pathogenicity of the PpE4-silenced lines. Furthermore, the expression of PpE4 in both N. benthamiana and Arabidopsis thaliana consistently enhanced plant susceptibility to P. parasitica. These results indicate that PpE4 contributes to pathogen infection. Finally, heterologous expression experiments showed that PpE4 triggers non-specific cell death in a variety of plants, including tobacco, tomato, potato and A. thaliana. Virus-induced gene silencing assays revealed that PpE4-induced cell death is dependent on HSP90, NPK and SGT1, suggesting that PpE4 is recognized by the plant immune system. In conclusion, PpE4 is an important virulence RXLR effector of P. parasitica and recognized by a wide range of host plants.</div></front></TEI><pubmed><MedlineCitation Status="MEDLINE" Owner="NLM"><PMID Version="1">30320960</PMID><DateCompleted><Year>2020</Year><Month>04</Month><Day>27</Day></DateCompleted><DateRevised><Year>2020</Year><Month>04</Month><Day>27</Day></DateRevised><Article PubModel="Print-Electronic"><Journal><ISSN IssnType="Electronic">1364-3703</ISSN><JournalIssue CitedMedium="Internet"><Volume>20</Volume><Issue>3</Issue><PubDate><Year>2019</Year><Month>03</Month></PubDate></JournalIssue><Title>Molecular plant pathology</Title><ISOAbbreviation>Mol Plant Pathol</ISOAbbreviation></Journal><ArticleTitle>An RXLR effector secreted by Phytophthora parasitica is a virulence factor and triggers cell death in various plants.</ArticleTitle><Pagination><MedlinePgn>356-371</MedlinePgn></Pagination><ELocationID EIdType="doi" ValidYN="Y">10.1111/mpp.12760</ELocationID><Abstract><AbstractText>RXLR effectors encoded by Phytophthora species play a central role in pathogen-plant interactions. An understanding of the biological functions of RXLR effectors is conducive to the illumination of the pathogenic mechanisms and the development of disease control strategies. However, the virulence function of Phytophthora parasitica RXLR effectors is poorly understood. Here, we describe the identification of a P. parasitica RXLR effector gene, PPTG00121 (PpE4), which is highly transcribed during the early stages of infection. Live cell imaging of P. parasitica transformants expressing a full-length PpE4 (E4FL)-mCherry protein indicated that PpE4 is secreted and accumulates around haustoria during plant infection. Silencing of PpE4 in P. parasitica resulted in significantly reduced virulence on Nicotiana benthamiana. Transient expression of PpE4 in N. benthamiana in turn restored the pathogenicity of the PpE4-silenced lines. Furthermore, the expression of PpE4 in both N. benthamiana and Arabidopsis thaliana consistently enhanced plant susceptibility to P. parasitica. These results indicate that PpE4 contributes to pathogen infection. Finally, heterologous expression experiments showed that PpE4 triggers non-specific cell death in a variety of plants, including tobacco, tomato, potato and A. thaliana. Virus-induced gene silencing assays revealed that PpE4-induced cell death is dependent on HSP90, NPK and SGT1, suggesting that PpE4 is recognized by the plant immune system. In conclusion, PpE4 is an important virulence RXLR effector of P. parasitica and recognized by a wide range of host plants.</AbstractText><CopyrightInformation>© 2018 The Authors. Molecular Plant Pathology published by BSPP and John Wiley & Sons Ltd.</CopyrightInformation></Abstract><AuthorList CompleteYN="Y"><Author ValidYN="Y"><LastName>Huang</LastName><ForeName>Guiyan</ForeName><Initials>G</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>College of Life Sciences, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Liu</LastName><ForeName>Zhirou</ForeName><Initials>Z</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Gu</LastName><ForeName>Biao</ForeName><Initials>B</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Zhao</LastName><ForeName>Hong</ForeName><Initials>H</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Jia</LastName><ForeName>Jinbu</ForeName><Initials>J</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>Institute of Plant and Food Science, Department of Biology, Southern University of Science and Technology, Shenzhen, 518055, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Fan</LastName><ForeName>Guangjin</ForeName><Initials>G</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>College of Plant Protection, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Meng</LastName><ForeName>Yuling</ForeName><Initials>Y</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>College of Agronomy, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Du</LastName><ForeName>Yu</ForeName><Initials>Y</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>College of Horticulture, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Shan</LastName><ForeName>Weixing</ForeName><Initials>W</Initials><Identifier Source="ORCID">0000-0001-7286-4041</Identifier><AffiliationInfo><Affiliation>State Key Laboratory of Crop Stress Biology for Arid Areas, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>College of Agronomy, Northwest A&F University, Yangling, Shaanxi, 712100, China.</Affiliation></AffiliationInfo></Author></AuthorList><Language>eng</Language><PublicationTypeList><PublicationType UI="D016428">Journal Article</PublicationType><PublicationType UI="D013485">Research Support, Non-U.S. Gov't</PublicationType></PublicationTypeList><ArticleDate DateType="Electronic"><Year>2018</Year><Month>11</Month><Day>22</Day></ArticleDate></Article><MedlineJournalInfo><Country>England</Country><MedlineTA>Mol Plant Pathol</MedlineTA><NlmUniqueID>100954969</NlmUniqueID><ISSNLinking>1364-3703</ISSNLinking></MedlineJournalInfo><CitationSubset>IM</CitationSubset><MeshHeadingList><MeshHeading><DescriptorName UI="D055750" MajorTopicYN="N">Phytophthora infestans</DescriptorName><QualifierName UI="Q000472" MajorTopicYN="Y">pathogenicity</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D010935" MajorTopicYN="N">Plant Diseases</DescriptorName><QualifierName UI="Q000382" MajorTopicYN="Y">microbiology</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D014026" MajorTopicYN="N">Tobacco</DescriptorName><QualifierName UI="Q000382" MajorTopicYN="N">microbiology</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D014774" MajorTopicYN="N">Virulence</DescriptorName></MeshHeading></MeshHeadingList><KeywordList Owner="NOTNLM"><Keyword MajorTopicYN="Y">Phytophthora parasitica</Keyword><Keyword MajorTopicYN="Y">RXLR effector</Keyword><Keyword MajorTopicYN="Y">cell 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last="Jia">Jinbu Jia</name><name sortKey="Jia, Jinbu" sort="Jia, Jinbu" uniqKey="Jia J" first="Jinbu" last="Jia">Jinbu Jia</name><name sortKey="Jia, Jinbu" sort="Jia, Jinbu" uniqKey="Jia J" first="Jinbu" last="Jia">Jinbu Jia</name><name sortKey="Liu, Zhirou" sort="Liu, Zhirou" uniqKey="Liu Z" first="Zhirou" last="Liu">Zhirou Liu</name><name sortKey="Liu, Zhirou" sort="Liu, Zhirou" uniqKey="Liu Z" first="Zhirou" last="Liu">Zhirou Liu</name><name sortKey="Meng, Yuling" sort="Meng, Yuling" uniqKey="Meng Y" first="Yuling" last="Meng">Yuling Meng</name><name sortKey="Meng, Yuling" sort="Meng, Yuling" uniqKey="Meng Y" first="Yuling" last="Meng">Yuling Meng</name><name sortKey="Shan, Weixing" sort="Shan, Weixing" uniqKey="Shan W" first="Weixing" last="Shan">Weixing Shan</name><name sortKey="Shan, Weixing" sort="Shan, Weixing" uniqKey="Shan W" first="Weixing" last="Shan">Weixing Shan</name><name sortKey="Zhao, Hong" sort="Zhao, Hong" uniqKey="Zhao H" first="Hong" last="Zhao">Hong 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