Phytophthora Effectors Modulate Genome-wide Alternative Splicing of Host mRNAs to Reprogram Plant Immunity.
Identifieur interne : 000081 ( Main/Exploration ); précédent : 000080; suivant : 000082Phytophthora Effectors Modulate Genome-wide Alternative Splicing of Host mRNAs to Reprogram Plant Immunity.
Auteurs : Jie Huang [République populaire de Chine] ; Xinyu Lu [République populaire de Chine] ; Hongwei Wu [République populaire de Chine] ; Yuchen Xie [République populaire de Chine] ; Qian Peng [République populaire de Chine] ; Lianfeng Gu [République populaire de Chine] ; Juyou Wu [République populaire de Chine] ; Yuanchao Wang [République populaire de Chine] ; Anireddy S N. Reddy [États-Unis] ; Suomeng Dong [République populaire de Chine]Source :
- Molecular plant [ 1752-9867 ] ; 2020.
Abstract
Alternative splicing (AS) of pre-mRNAs increases transcriptome and proteome diversity, regulates gene expression through multiple mechanisms, and plays important roles in plant development and stress responses. However, the prevalence of genome-wide plant AS changes during infection and the mechanisms by which pathogens modulate AS remain poorly understood. Here, we examined the global AS changes in tomato leaves infected with Phytophthora infestans, the infamous Irish famine pathogen. We show that more than 2000 genes exhibiting significant changes in AS are not differentially expressed, indicating that AS is a distinct layer of transcriptome reprogramming during plant-pathogen interactions. Furthermore, our results show that P. infestans subverts host immunity by repressing the AS of positive regulators of plant immunity and promoting the AS of susceptibility factors. To study the underlying mechanism, we established a luminescence-based AS reporter system in Nicotiana benthamiana to screen pathogen effectors modulating plant AS. We identified nine splicing regulatory effectors (SREs) from 87 P. infestans effectors. Further studies revealed that SRE3 physically binds U1-70K to manipulate the plant AS machinery and subsequently modulates AS-mediated plant immunity. Our study not only unveils genome-wide plant AS reprogramming during infection but also establishes a novel AS screening tool to identify SREs from a wide range of plant pathogens, providing opportunities to understand the splicing regulatory mechanisms through which pathogens subvert plant immunity.
DOI: 10.1016/j.molp.2020.07.007
PubMed: 32693165
Affiliations:
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Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095, China; The Key Laboratory of Plant Immunity, Nanjing Agricultural University, Nanjing 210095, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China; Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095, China; The Key Laboratory of Plant Immunity, Nanjing Agricultural University, Nanjing 210095</wicri:regionArea><wicri:noRegion>Nanjing 210095</wicri:noRegion></affiliation></author><author><name sortKey="Reddy, Anireddy S N" sort="Reddy, Anireddy S N" uniqKey="Reddy A" first="Anireddy S N" last="Reddy">Anireddy S N. Reddy</name><affiliation wicri:level="2"><nlm:affiliation>Colorado State University, Program in Cell and Molecular Biology, Fort Collins, CO 80523, USA.</nlm:affiliation><country xml:lang="fr">États-Unis</country><wicri:regionArea>Colorado State University, Program in Cell and Molecular Biology, Fort Collins, CO 80523</wicri:regionArea><placeName><region type="state">Colorado</region></placeName></affiliation></author><author><name sortKey="Dong, Suomeng" sort="Dong, Suomeng" uniqKey="Dong S" first="Suomeng" last="Dong">Suomeng Dong</name><affiliation wicri:level="1"><nlm:affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China; Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095, China; The Key Laboratory of Plant Immunity, Nanjing Agricultural University, Nanjing 210095, China. Electronic address: smdong@njau.edu.cn.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China; Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095, China; The Key Laboratory of Plant Immunity, Nanjing Agricultural University, Nanjing 210095</wicri:regionArea><wicri:noRegion>Nanjing 210095</wicri:noRegion></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PubMed</idno><date when="2020">2020</date><idno type="RBID">pubmed:32693165</idno><idno type="pmid">32693165</idno><idno type="doi">10.1016/j.molp.2020.07.007</idno><idno type="wicri:Area/Main/Corpus">000178</idno><idno type="wicri:explorRef" wicri:stream="Main" wicri:step="Corpus" wicri:corpus="PubMed">000178</idno><idno type="wicri:Area/Main/Curation">000178</idno><idno type="wicri:explorRef" wicri:stream="Main" wicri:step="Curation">000178</idno><idno type="wicri:Area/Main/Exploration">000178</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en">Phytophthora Effectors Modulate Genome-wide Alternative Splicing of Host mRNAs to Reprogram Plant Immunity.</title><author><name sortKey="Huang, Jie" sort="Huang, Jie" uniqKey="Huang J" first="Jie" last="Huang">Jie Huang</name><affiliation wicri:level="1"><nlm:affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China; Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China; Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095</wicri:regionArea><wicri:noRegion>Nanjing 210095</wicri:noRegion></affiliation></author><author><name sortKey="Lu, Xinyu" sort="Lu, Xinyu" uniqKey="Lu X" first="Xinyu" last="Lu">Xinyu Lu</name><affiliation wicri:level="1"><nlm:affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095</wicri:regionArea><wicri:noRegion>Nanjing 210095</wicri:noRegion></affiliation></author><author><name sortKey="Wu, Hongwei" sort="Wu, Hongwei" uniqKey="Wu H" first="Hongwei" last="Wu">Hongwei Wu</name><affiliation wicri:level="1"><nlm:affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095</wicri:regionArea><wicri:noRegion>Nanjing 210095</wicri:noRegion></affiliation></author><author><name sortKey="Xie, Yuchen" sort="Xie, Yuchen" uniqKey="Xie Y" first="Yuchen" last="Xie">Yuchen Xie</name><affiliation wicri:level="1"><nlm:affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095</wicri:regionArea><wicri:noRegion>Nanjing 210095</wicri:noRegion></affiliation></author><author><name sortKey="Peng, Qian" sort="Peng, Qian" uniqKey="Peng Q" first="Qian" last="Peng">Qian Peng</name><affiliation wicri:level="1"><nlm:affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095</wicri:regionArea><wicri:noRegion>Nanjing 210095</wicri:noRegion></affiliation></author><author><name sortKey="Gu, Lianfeng" sort="Gu, Lianfeng" uniqKey="Gu L" first="Lianfeng" last="Gu">Lianfeng Gu</name><affiliation wicri:level="1"><nlm:affiliation>Basic Forestry and Proteomics Research Center, College of Forestry, Fujian Agriculture and Forestry University, Fuzhou 350002, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>Basic Forestry and Proteomics Research Center, College of Forestry, Fujian Agriculture and Forestry University, Fuzhou 350002</wicri:regionArea><placeName><settlement type="city">Fuzhou</settlement><region type="province">Fujian</region></placeName></affiliation></author><author><name sortKey="Wu, Juyou" sort="Wu, Juyou" uniqKey="Wu J" first="Juyou" last="Wu">Juyou Wu</name><affiliation wicri:level="1"><nlm:affiliation>State Key Laboratory of Crop Genetics and Germplasm Enhancement, College of Horticulture, Nanjing Agricultural University, Nanjing 210095, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>State Key Laboratory of Crop Genetics and Germplasm Enhancement, College of Horticulture, Nanjing Agricultural University, Nanjing 210095</wicri:regionArea><wicri:noRegion>Nanjing 210095</wicri:noRegion></affiliation></author><author><name sortKey="Wang, Yuanchao" sort="Wang, Yuanchao" uniqKey="Wang Y" first="Yuanchao" last="Wang">Yuanchao Wang</name><affiliation wicri:level="1"><nlm:affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China; Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095, China; The Key Laboratory of Plant Immunity, Nanjing Agricultural University, Nanjing 210095, China.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China; Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095, China; The Key Laboratory of Plant Immunity, Nanjing Agricultural University, Nanjing 210095</wicri:regionArea><wicri:noRegion>Nanjing 210095</wicri:noRegion></affiliation></author><author><name sortKey="Reddy, Anireddy S N" sort="Reddy, Anireddy S N" uniqKey="Reddy A" first="Anireddy S N" last="Reddy">Anireddy S N. Reddy</name><affiliation wicri:level="2"><nlm:affiliation>Colorado State University, Program in Cell and Molecular Biology, Fort Collins, CO 80523, USA.</nlm:affiliation><country xml:lang="fr">États-Unis</country><wicri:regionArea>Colorado State University, Program in Cell and Molecular Biology, Fort Collins, CO 80523</wicri:regionArea><placeName><region type="state">Colorado</region></placeName></affiliation></author><author><name sortKey="Dong, Suomeng" sort="Dong, Suomeng" uniqKey="Dong S" first="Suomeng" last="Dong">Suomeng Dong</name><affiliation wicri:level="1"><nlm:affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China; Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095, China; The Key Laboratory of Plant Immunity, Nanjing Agricultural University, Nanjing 210095, China. Electronic address: smdong@njau.edu.cn.</nlm:affiliation><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China; Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095, China; The Key Laboratory of Plant Immunity, Nanjing Agricultural University, Nanjing 210095</wicri:regionArea><wicri:noRegion>Nanjing 210095</wicri:noRegion></affiliation></author></analytic><series><title level="j">Molecular plant</title><idno type="eISSN">1752-9867</idno><imprint><date when="2020" type="published">2020</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en">Alternative splicing (AS) of pre-mRNAs increases transcriptome and proteome diversity, regulates gene expression through multiple mechanisms, and plays important roles in plant development and stress responses. However, the prevalence of genome-wide plant AS changes during infection and the mechanisms by which pathogens modulate AS remain poorly understood. Here, we examined the global AS changes in tomato leaves infected with Phytophthora infestans, the infamous Irish famine pathogen. We show that more than 2000 genes exhibiting significant changes in AS are not differentially expressed, indicating that AS is a distinct layer of transcriptome reprogramming during plant-pathogen interactions. Furthermore, our results show that P. infestans subverts host immunity by repressing the AS of positive regulators of plant immunity and promoting the AS of susceptibility factors. To study the underlying mechanism, we established a luminescence-based AS reporter system in Nicotiana benthamiana to screen pathogen effectors modulating plant AS. We identified nine splicing regulatory effectors (SREs) from 87 P. infestans effectors. Further studies revealed that SRE3 physically binds U1-70K to manipulate the plant AS machinery and subsequently modulates AS-mediated plant immunity. Our study not only unveils genome-wide plant AS reprogramming during infection but also establishes a novel AS screening tool to identify SREs from a wide range of plant pathogens, providing opportunities to understand the splicing regulatory mechanisms through which pathogens subvert plant immunity.</div></front></TEI><pubmed><MedlineCitation Status="In-Data-Review" Owner="NLM"><PMID Version="1">32693165</PMID><DateRevised><Year>2020</Year><Month>10</Month><Day>07</Day></DateRevised><Article PubModel="Print-Electronic"><Journal><ISSN IssnType="Electronic">1752-9867</ISSN><JournalIssue CitedMedium="Internet"><Volume>13</Volume><Issue>10</Issue><PubDate><Year>2020</Year><Month>Oct</Month><Day>05</Day></PubDate></JournalIssue><Title>Molecular plant</Title><ISOAbbreviation>Mol Plant</ISOAbbreviation></Journal><ArticleTitle>Phytophthora Effectors Modulate Genome-wide Alternative Splicing of Host mRNAs to Reprogram Plant Immunity.</ArticleTitle><Pagination><MedlinePgn>1470-1484</MedlinePgn></Pagination><ELocationID EIdType="pii" ValidYN="Y">S1674-2052(20)30220-3</ELocationID><ELocationID EIdType="doi" ValidYN="Y">10.1016/j.molp.2020.07.007</ELocationID><Abstract><AbstractText>Alternative splicing (AS) of pre-mRNAs increases transcriptome and proteome diversity, regulates gene expression through multiple mechanisms, and plays important roles in plant development and stress responses. However, the prevalence of genome-wide plant AS changes during infection and the mechanisms by which pathogens modulate AS remain poorly understood. Here, we examined the global AS changes in tomato leaves infected with Phytophthora infestans, the infamous Irish famine pathogen. We show that more than 2000 genes exhibiting significant changes in AS are not differentially expressed, indicating that AS is a distinct layer of transcriptome reprogramming during plant-pathogen interactions. Furthermore, our results show that P. infestans subverts host immunity by repressing the AS of positive regulators of plant immunity and promoting the AS of susceptibility factors. To study the underlying mechanism, we established a luminescence-based AS reporter system in Nicotiana benthamiana to screen pathogen effectors modulating plant AS. We identified nine splicing regulatory effectors (SREs) from 87 P. infestans effectors. Further studies revealed that SRE3 physically binds U1-70K to manipulate the plant AS machinery and subsequently modulates AS-mediated plant immunity. Our study not only unveils genome-wide plant AS reprogramming during infection but also establishes a novel AS screening tool to identify SREs from a wide range of plant pathogens, providing opportunities to understand the splicing regulatory mechanisms through which pathogens subvert plant immunity.</AbstractText><CopyrightInformation>Copyright © 2020 The Author. Published by Elsevier Inc. All rights reserved.</CopyrightInformation></Abstract><AuthorList CompleteYN="Y"><Author ValidYN="Y"><LastName>Huang</LastName><ForeName>Jie</ForeName><Initials>J</Initials><AffiliationInfo><Affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China; Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Lu</LastName><ForeName>Xinyu</ForeName><Initials>X</Initials><AffiliationInfo><Affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Wu</LastName><ForeName>Hongwei</ForeName><Initials>H</Initials><AffiliationInfo><Affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Xie</LastName><ForeName>Yuchen</ForeName><Initials>Y</Initials><AffiliationInfo><Affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Peng</LastName><ForeName>Qian</ForeName><Initials>Q</Initials><AffiliationInfo><Affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Gu</LastName><ForeName>Lianfeng</ForeName><Initials>L</Initials><AffiliationInfo><Affiliation>Basic Forestry and Proteomics Research Center, College of Forestry, Fujian Agriculture and Forestry University, Fuzhou 350002, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Wu</LastName><ForeName>Juyou</ForeName><Initials>J</Initials><AffiliationInfo><Affiliation>State Key Laboratory of Crop Genetics and Germplasm Enhancement, College of Horticulture, Nanjing Agricultural University, Nanjing 210095, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Wang</LastName><ForeName>Yuanchao</ForeName><Initials>Y</Initials><AffiliationInfo><Affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China; Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095, China; The Key Laboratory of Plant Immunity, Nanjing Agricultural University, Nanjing 210095, China.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Reddy</LastName><ForeName>Anireddy S N</ForeName><Initials>ASN</Initials><AffiliationInfo><Affiliation>Colorado State University, Program in Cell and Molecular Biology, Fort Collins, CO 80523, USA.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Dong</LastName><ForeName>Suomeng</ForeName><Initials>S</Initials><AffiliationInfo><Affiliation>Department of Plant Pathology, Nanjing Agricultural University, Nanjing 210095, China; Key Laboratory of Integrated Management of Crop Diseases and Pests (Ministry of Education), Nanjing 210095, China; The Key Laboratory of Plant Immunity, Nanjing Agricultural University, Nanjing 210095, China. Electronic address: smdong@njau.edu.cn.</Affiliation></AffiliationInfo></Author></AuthorList><Language>eng</Language><PublicationTypeList><PublicationType UI="D016428">Journal Article</PublicationType></PublicationTypeList><ArticleDate DateType="Electronic"><Year>2020</Year><Month>07</Month><Day>18</Day></ArticleDate></Article><MedlineJournalInfo><Country>England</Country><MedlineTA>Mol Plant</MedlineTA><NlmUniqueID>101465514</NlmUniqueID><ISSNLinking>1674-2052</ISSNLinking></MedlineJournalInfo><CitationSubset>IM</CitationSubset><KeywordList Owner="NOTNLM"><Keyword MajorTopicYN="N">Phytophthora infestans</Keyword><Keyword MajorTopicYN="N">alternative splicing</Keyword><Keyword MajorTopicYN="N">late blight</Keyword><Keyword MajorTopicYN="N">plant immunity</Keyword><Keyword MajorTopicYN="N">splicing regulatory effectors</Keyword><Keyword MajorTopicYN="N">tomato</Keyword></KeywordList></MedlineCitation><PubmedData><History><PubMedPubDate PubStatus="received"><Year>2019</Year><Month>10</Month><Day>30</Day></PubMedPubDate><PubMedPubDate PubStatus="revised"><Year>2020</Year><Month>06</Month><Day>30</Day></PubMedPubDate><PubMedPubDate PubStatus="accepted"><Year>2020</Year><Month>07</Month><Day>15</Day></PubMedPubDate><PubMedPubDate PubStatus="pubmed"><Year>2020</Year><Month>7</Month><Day>22</Day><Hour>6</Hour><Minute>0</Minute></PubMedPubDate><PubMedPubDate PubStatus="medline"><Year>2020</Year><Month>7</Month><Day>22</Day><Hour>6</Hour><Minute>0</Minute></PubMedPubDate><PubMedPubDate PubStatus="entrez"><Year>2020</Year><Month>7</Month><Day>22</Day><Hour>6</Hour><Minute>0</Minute></PubMedPubDate></History><PublicationStatus>ppublish</PublicationStatus><ArticleIdList><ArticleId IdType="pubmed">32693165</ArticleId><ArticleId IdType="pii">S1674-2052(20)30220-3</ArticleId><ArticleId IdType="doi">10.1016/j.molp.2020.07.007</ArticleId></ArticleIdList></PubmedData></pubmed><affiliations><list><country><li>République populaire de Chine</li><li>États-Unis</li></country><region><li>Colorado</li><li>Fujian</li></region><settlement><li>Fuzhou</li></settlement></list><tree><country name="République populaire de Chine"><noRegion><name sortKey="Huang, Jie" sort="Huang, Jie" uniqKey="Huang J" first="Jie" last="Huang">Jie Huang</name></noRegion><name sortKey="Dong, Suomeng" sort="Dong, Suomeng" uniqKey="Dong S" first="Suomeng" last="Dong">Suomeng Dong</name><name sortKey="Gu, Lianfeng" sort="Gu, Lianfeng" uniqKey="Gu L" first="Lianfeng" last="Gu">Lianfeng Gu</name><name sortKey="Lu, Xinyu" sort="Lu, Xinyu" uniqKey="Lu X" first="Xinyu" last="Lu">Xinyu Lu</name><name sortKey="Peng, Qian" sort="Peng, Qian" uniqKey="Peng Q" first="Qian" last="Peng">Qian Peng</name><name sortKey="Wang, Yuanchao" sort="Wang, Yuanchao" uniqKey="Wang Y" first="Yuanchao" last="Wang">Yuanchao Wang</name><name sortKey="Wu, Hongwei" sort="Wu, Hongwei" uniqKey="Wu H" first="Hongwei" last="Wu">Hongwei Wu</name><name sortKey="Wu, Juyou" sort="Wu, Juyou" uniqKey="Wu J" first="Juyou" last="Wu">Juyou Wu</name><name sortKey="Xie, Yuchen" sort="Xie, Yuchen" uniqKey="Xie Y" first="Yuchen" last="Xie">Yuchen Xie</name></country><country name="États-Unis"><region name="Colorado"><name sortKey="Reddy, Anireddy S N" sort="Reddy, Anireddy S N" uniqKey="Reddy A" first="Anireddy S N" last="Reddy">Anireddy S N. 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HfdIndexSelect -h $EXPLOR_AREA/Data/Main/Exploration/RBID.i -Sk "pubmed:32693165" \
| HfdSelect -Kh $EXPLOR_AREA/Data/Main/Exploration/biblio.hfd \
| NlmPubMed2Wicri -a PlantPathoEffV1
| This area was generated with Dilib version V0.6.38. |  |