Pathogenomic Analysis of Wheat Yellow Rust Lineages Detects Seasonal Variation and Host Specificity.
Identifieur interne : 000263 ( Main/Exploration ); précédent : 000262; suivant : 000264Pathogenomic Analysis of Wheat Yellow Rust Lineages Detects Seasonal Variation and Host Specificity.
Auteurs : Vanessa Bueno-Sancho [Royaume-Uni] ; Antoine Persoons [Royaume-Uni] ; Amelia Hubbard [Royaume-Uni] ; Luis Enrique Cabrera-Quio [Royaume-Uni] ; Clare M. Lewis [Royaume-Uni] ; Pilar Corredor-Moreno [Royaume-Uni] ; Daniel C E. Bunting [Royaume-Uni] ; Sajid Ali [Pakistan] ; Soonie Chng [Nouvelle-Zélande] ; David P. Hodson [Éthiopie] ; Ricardo Madariaga Burrows [Chili] ; Rosie Bryson [Allemagne] ; Jane Thomas [Royaume-Uni] ; Sarah Holdgate [Royaume-Uni] ; Diane G O. Saunders [Royaume-Uni]Source :
- Genome biology and evolution [ 1759-6653 ] ; 2017.
Descripteurs français
- KwdFr :
- Basidiomycota (classification), Basidiomycota (génétique), Génome fongique (MeSH), Génomique (méthodes), Génotype (MeSH), Maladies des plantes (microbiologie), Phylogenèse (MeSH), Saisons (MeSH), Spécificité d'hôte (MeSH), Séquençage nucléotidique à haut débit (MeSH), Triticum (microbiologie), Virulence (MeSH), Épidémies de maladies (MeSH).
- MESH :
- génétique : Basidiomycota.
- microbiologie : Maladies des plantes, Triticum.
- méthodes : Génomique.
- classification : Basidiomycota, Génome fongique, Génotype, Phylogenèse, Saisons, Spécificité d'hôte, Séquençage nucléotidique à haut débit, Virulence, Épidémies de maladies.
English descriptors
- KwdEn :
- Basidiomycota (classification), Basidiomycota (genetics), Disease Outbreaks (MeSH), Genome, Fungal (MeSH), Genomics (methods), Genotype (MeSH), High-Throughput Nucleotide Sequencing (MeSH), Host Specificity (MeSH), Phylogeny (MeSH), Plant Diseases (microbiology), Seasons (MeSH), Triticum (microbiology), Virulence (MeSH).
- MESH :
- classification : Basidiomycota.
- genetics : Basidiomycota.
- methods : Genomics.
- microbiology : Plant Diseases, Triticum.
- Disease Outbreaks, Genome, Fungal, Genotype, High-Throughput Nucleotide Sequencing, Host Specificity, Phylogeny, Seasons, Virulence.
Abstract
Recent disease outbreaks caused by (re-)emerging plant pathogens have been associated with expansions in pathogen geographic distribution and increased virulence. For example, in the past two decades' wheat yellow (stripe) rust, Puccinia striiformis f. sp. tritici, has seen the emergence of new races that are adapted to warmer temperatures, have expanded virulence profiles, and are more aggressive than previous races, leading to wide-scale epidemics. Here, we used field-based genotyping to generate high-resolution data on P. striiformis genetics and carried out global population analysis. We also undertook comparative analysis of the 2014 and 2013 UK populations and assessed the temporal dynamics and host specificity of distinct pathogen genotypes. Our analysis revealed that P. striiformis lineages recently detected in Europe are extremely diverse and in fact similar to globally dispersed populations. In addition, we identified a considerable shift in the UK P. striiformis population structure including the first identification of one infamous race known as Kranich. Next, by establishing the genotype of both the pathogen and host within a single infected field sample, we uncovered evidence for varietal specificity for genetic groups of P. striiformis. Finally, we found potential seasonal specificity for certain genotypes of the pathogen with several lineages identified only in samples collected in late spring and into the summer, whereas one lineage was identified throughout the wheat growing season. Our discovery of which wheat varieties are susceptible to which specific P. striiformis isolates, and when those isolates are prevalent throughout the year, represents a powerful tool for disease management.
DOI: 10.1093/gbe/evx241
PubMed: 29177504
PubMed Central: PMC5730935
Affiliations:
Links toward previous steps (curation, corpus...)
Le document en format XML
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Lewis</name><affiliation wicri:level="1"><nlm:affiliation>Crop Genetics, John Innes Centre, Norwich Research Park, United Kingdom.</nlm:affiliation><country xml:lang="fr">Royaume-Uni</country><wicri:regionArea>Crop Genetics, John Innes Centre, Norwich Research Park</wicri:regionArea><wicri:noRegion>Norwich Research Park</wicri:noRegion></affiliation></author><author><name sortKey="Corredor Moreno, Pilar" sort="Corredor Moreno, Pilar" uniqKey="Corredor Moreno P" first="Pilar" last="Corredor-Moreno">Pilar Corredor-Moreno</name><affiliation wicri:level="1"><nlm:affiliation>Crop Genetics, John Innes Centre, Norwich Research Park, United Kingdom.</nlm:affiliation><country xml:lang="fr">Royaume-Uni</country><wicri:regionArea>Crop Genetics, John Innes Centre, Norwich Research Park</wicri:regionArea><wicri:noRegion>Norwich Research Park</wicri:noRegion></affiliation></author><author><name sortKey="Bunting, Daniel C E" sort="Bunting, Daniel C E" uniqKey="Bunting D" first="Daniel C E" last="Bunting">Daniel C E. Bunting</name><affiliation wicri:level="1"><nlm:affiliation>Crop Genetics, John Innes Centre, Norwich Research Park, United Kingdom.</nlm:affiliation><country xml:lang="fr">Royaume-Uni</country><wicri:regionArea>Crop Genetics, John Innes Centre, Norwich Research Park</wicri:regionArea><wicri:noRegion>Norwich Research Park</wicri:noRegion></affiliation></author><author><name sortKey="Ali, Sajid" sort="Ali, Sajid" uniqKey="Ali S" first="Sajid" last="Ali">Sajid Ali</name><affiliation wicri:level="1"><nlm:affiliation>The University of Agriculture, Peshawar, Pakistan.</nlm:affiliation><country xml:lang="fr">Pakistan</country><wicri:regionArea>The University of Agriculture, Peshawar</wicri:regionArea><wicri:noRegion>Peshawar</wicri:noRegion></affiliation></author><author><name sortKey="Chng, Soonie" sort="Chng, Soonie" uniqKey="Chng S" first="Soonie" last="Chng">Soonie Chng</name><affiliation wicri:level="1"><nlm:affiliation>The New Zealand Institute for Plant & Food Research, Lincoln, New Zealand.</nlm:affiliation><country xml:lang="fr">Nouvelle-Zélande</country><wicri:regionArea>The New Zealand Institute for Plant & Food Research, Lincoln</wicri:regionArea><wicri:noRegion>Lincoln</wicri:noRegion></affiliation></author><author><name sortKey="Hodson, David P" sort="Hodson, David P" uniqKey="Hodson D" first="David P" last="Hodson">David P. Hodson</name><affiliation wicri:level="1"><nlm:affiliation>CIMMYT, Addis Ababa, Ethiopia.</nlm:affiliation><country xml:lang="fr">Éthiopie</country><wicri:regionArea>CIMMYT, Addis Ababa</wicri:regionArea><wicri:noRegion>Addis Ababa</wicri:noRegion></affiliation></author><author><name sortKey="Madariaga Burrows, Ricardo" sort="Madariaga Burrows, Ricardo" uniqKey="Madariaga Burrows R" first="Ricardo" last="Madariaga Burrows">Ricardo Madariaga Burrows</name><affiliation wicri:level="1"><nlm:affiliation>National Institute of Agricultural Research, INIA, Chile.</nlm:affiliation><country xml:lang="fr">Chili</country><wicri:regionArea>National Institute of Agricultural Research, INIA</wicri:regionArea><wicri:noRegion>INIA</wicri:noRegion></affiliation></author><author><name sortKey="Bryson, Rosie" sort="Bryson, Rosie" uniqKey="Bryson R" first="Rosie" last="Bryson">Rosie Bryson</name><affiliation wicri:level="1"><nlm:affiliation>BASF SE, Agricultural Centre, Limburgerhof, Germany.</nlm:affiliation><country xml:lang="fr">Allemagne</country><wicri:regionArea>BASF SE, Agricultural Centre, Limburgerhof</wicri:regionArea><wicri:noRegion>Limburgerhof</wicri:noRegion><wicri:noRegion>Limburgerhof</wicri:noRegion><wicri:noRegion>Limburgerhof</wicri:noRegion></affiliation></author><author><name sortKey="Thomas, Jane" sort="Thomas, Jane" uniqKey="Thomas J" first="Jane" last="Thomas">Jane Thomas</name><affiliation wicri:level="1"><nlm:affiliation>ational Institute of Agricultural Botany, Cambridge, United Kingdom.</nlm:affiliation><country xml:lang="fr">Royaume-Uni</country><wicri:regionArea>ational Institute of Agricultural Botany, Cambridge</wicri:regionArea><wicri:noRegion>Cambridge</wicri:noRegion></affiliation></author><author><name sortKey="Holdgate, Sarah" sort="Holdgate, Sarah" uniqKey="Holdgate S" first="Sarah" last="Holdgate">Sarah Holdgate</name><affiliation wicri:level="1"><nlm:affiliation>ational Institute of Agricultural Botany, Cambridge, United Kingdom.</nlm:affiliation><country xml:lang="fr">Royaume-Uni</country><wicri:regionArea>ational Institute of Agricultural Botany, Cambridge</wicri:regionArea><wicri:noRegion>Cambridge</wicri:noRegion></affiliation></author><author><name sortKey="Saunders, Diane G O" sort="Saunders, Diane G O" uniqKey="Saunders D" first="Diane G O" last="Saunders">Diane G O. Saunders</name><affiliation wicri:level="1"><nlm:affiliation>Crop Genetics, John Innes Centre, Norwich Research Park, United Kingdom.</nlm:affiliation><country xml:lang="fr">Royaume-Uni</country><wicri:regionArea>Crop Genetics, John Innes Centre, Norwich Research Park</wicri:regionArea><wicri:noRegion>Norwich Research Park</wicri:noRegion></affiliation></author></analytic><series><title level="j">Genome biology and evolution</title><idno type="eISSN">1759-6653</idno><imprint><date when="2017" type="published">2017</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass><keywords scheme="KwdEn" xml:lang="en"><term>Basidiomycota (classification)</term><term>Basidiomycota (genetics)</term><term>Disease Outbreaks (MeSH)</term><term>Genome, Fungal (MeSH)</term><term>Genomics (methods)</term><term>Genotype (MeSH)</term><term>High-Throughput Nucleotide Sequencing (MeSH)</term><term>Host Specificity (MeSH)</term><term>Phylogeny (MeSH)</term><term>Plant Diseases (microbiology)</term><term>Seasons (MeSH)</term><term>Triticum (microbiology)</term><term>Virulence (MeSH)</term></keywords><keywords scheme="KwdFr" xml:lang="fr"><term>Basidiomycota (classification)</term><term>Basidiomycota (génétique)</term><term>Génome fongique (MeSH)</term><term>Génomique (méthodes)</term><term>Génotype (MeSH)</term><term>Maladies des plantes (microbiologie)</term><term>Phylogenèse (MeSH)</term><term>Saisons (MeSH)</term><term>Spécificité d'hôte (MeSH)</term><term>Séquençage nucléotidique à haut débit (MeSH)</term><term>Triticum (microbiologie)</term><term>Virulence (MeSH)</term><term>Épidémies de maladies (MeSH)</term></keywords><keywords scheme="MESH" qualifier="classification" xml:lang="en"><term>Basidiomycota</term></keywords><keywords scheme="MESH" qualifier="genetics" xml:lang="en"><term>Basidiomycota</term></keywords><keywords scheme="MESH" qualifier="génétique" xml:lang="fr"><term>Basidiomycota</term></keywords><keywords scheme="MESH" qualifier="methods" xml:lang="en"><term>Genomics</term></keywords><keywords scheme="MESH" qualifier="microbiologie" xml:lang="fr"><term>Maladies des plantes</term><term>Triticum</term></keywords><keywords scheme="MESH" qualifier="microbiology" xml:lang="en"><term>Plant Diseases</term><term>Triticum</term></keywords><keywords scheme="MESH" qualifier="méthodes" xml:lang="fr"><term>Génomique</term></keywords><keywords scheme="MESH" xml:lang="en"><term>Disease Outbreaks</term><term>Genome, Fungal</term><term>Genotype</term><term>High-Throughput Nucleotide Sequencing</term><term>Host Specificity</term><term>Phylogeny</term><term>Seasons</term><term>Virulence</term></keywords><keywords scheme="MESH" qualifier="classification" xml:lang="fr"><term>Basidiomycota</term><term>Génome fongique</term><term>Génotype</term><term>Phylogenèse</term><term>Saisons</term><term>Spécificité d'hôte</term><term>Séquençage nucléotidique à haut débit</term><term>Virulence</term><term>Épidémies de maladies</term></keywords></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en">Recent disease outbreaks caused by (re-)emerging plant pathogens have been associated with expansions in pathogen geographic distribution and increased virulence. For example, in the past two decades' wheat yellow (stripe) rust, Puccinia striiformis f. sp. tritici, has seen the emergence of new races that are adapted to warmer temperatures, have expanded virulence profiles, and are more aggressive than previous races, leading to wide-scale epidemics. Here, we used field-based genotyping to generate high-resolution data on P. striiformis genetics and carried out global population analysis. We also undertook comparative analysis of the 2014 and 2013 UK populations and assessed the temporal dynamics and host specificity of distinct pathogen genotypes. Our analysis revealed that P. striiformis lineages recently detected in Europe are extremely diverse and in fact similar to globally dispersed populations. In addition, we identified a considerable shift in the UK P. striiformis population structure including the first identification of one infamous race known as Kranich. Next, by establishing the genotype of both the pathogen and host within a single infected field sample, we uncovered evidence for varietal specificity for genetic groups of P. striiformis. Finally, we found potential seasonal specificity for certain genotypes of the pathogen with several lineages identified only in samples collected in late spring and into the summer, whereas one lineage was identified throughout the wheat growing season. Our discovery of which wheat varieties are susceptible to which specific P. striiformis isolates, and when those isolates are prevalent throughout the year, represents a powerful tool for disease management.</div></front></TEI><pubmed><MedlineCitation Status="MEDLINE" Owner="NLM"><PMID Version="1">29177504</PMID><DateCompleted><Year>2018</Year><Month>05</Month><Day>10</Day></DateCompleted><DateRevised><Year>2018</Year><Month>11</Month><Day>13</Day></DateRevised><Article PubModel="Print"><Journal><ISSN IssnType="Electronic">1759-6653</ISSN><JournalIssue CitedMedium="Internet"><Volume>9</Volume><Issue>12</Issue><PubDate><Year>2017</Year><Month>12</Month><Day>01</Day></PubDate></JournalIssue><Title>Genome biology and evolution</Title><ISOAbbreviation>Genome Biol Evol</ISOAbbreviation></Journal><ArticleTitle>Pathogenomic Analysis of Wheat Yellow Rust Lineages Detects Seasonal Variation and Host Specificity.</ArticleTitle><Pagination><MedlinePgn>3282-3296</MedlinePgn></Pagination><ELocationID EIdType="doi" ValidYN="Y">10.1093/gbe/evx241</ELocationID><Abstract><AbstractText>Recent disease outbreaks caused by (re-)emerging plant pathogens have been associated with expansions in pathogen geographic distribution and increased virulence. For example, in the past two decades' wheat yellow (stripe) rust, Puccinia striiformis f. sp. tritici, has seen the emergence of new races that are adapted to warmer temperatures, have expanded virulence profiles, and are more aggressive than previous races, leading to wide-scale epidemics. Here, we used field-based genotyping to generate high-resolution data on P. striiformis genetics and carried out global population analysis. We also undertook comparative analysis of the 2014 and 2013 UK populations and assessed the temporal dynamics and host specificity of distinct pathogen genotypes. Our analysis revealed that P. striiformis lineages recently detected in Europe are extremely diverse and in fact similar to globally dispersed populations. In addition, we identified a considerable shift in the UK P. striiformis population structure including the first identification of one infamous race known as Kranich. Next, by establishing the genotype of both the pathogen and host within a single infected field sample, we uncovered evidence for varietal specificity for genetic groups of P. striiformis. Finally, we found potential seasonal specificity for certain genotypes of the pathogen with several lineages identified only in samples collected in late spring and into the summer, whereas one lineage was identified throughout the wheat growing season. Our discovery of which wheat varieties are susceptible to which specific P. striiformis isolates, and when those isolates are prevalent throughout the year, represents a powerful tool for disease management.</AbstractText><CopyrightInformation>© The Author 2017. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.</CopyrightInformation></Abstract><AuthorList CompleteYN="Y"><Author ValidYN="Y"><LastName>Bueno-Sancho</LastName><ForeName>Vanessa</ForeName><Initials>V</Initials><AffiliationInfo><Affiliation>Crop Genetics, John Innes Centre, Norwich Research Park, United Kingdom.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Persoons</LastName><ForeName>Antoine</ForeName><Initials>A</Initials><AffiliationInfo><Affiliation>Crop Genetics, John Innes Centre, Norwich Research Park, United Kingdom.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Hubbard</LastName><ForeName>Amelia</ForeName><Initials>A</Initials><AffiliationInfo><Affiliation>ational Institute of Agricultural Botany, Cambridge, United Kingdom.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Cabrera-Quio</LastName><ForeName>Luis Enrique</ForeName><Initials>LE</Initials><AffiliationInfo><Affiliation>Earlham Institute, Norwich Research Park, United Kingdom.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>The Sainsbury Laboratory, Norwich Research Park, United Kingdom.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Lewis</LastName><ForeName>Clare M</ForeName><Initials>CM</Initials><AffiliationInfo><Affiliation>Crop Genetics, John Innes Centre, Norwich Research Park, United Kingdom.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Corredor-Moreno</LastName><ForeName>Pilar</ForeName><Initials>P</Initials><AffiliationInfo><Affiliation>Crop Genetics, John Innes Centre, Norwich Research Park, United Kingdom.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Bunting</LastName><ForeName>Daniel C E</ForeName><Initials>DCE</Initials><AffiliationInfo><Affiliation>Crop Genetics, John Innes Centre, Norwich Research Park, United Kingdom.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Ali</LastName><ForeName>Sajid</ForeName><Initials>S</Initials><AffiliationInfo><Affiliation>The University of Agriculture, Peshawar, Pakistan.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Chng</LastName><ForeName>Soonie</ForeName><Initials>S</Initials><AffiliationInfo><Affiliation>The New Zealand Institute for Plant & Food Research, Lincoln, New Zealand.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Hodson</LastName><ForeName>David P</ForeName><Initials>DP</Initials><AffiliationInfo><Affiliation>CIMMYT, Addis Ababa, Ethiopia.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Madariaga Burrows</LastName><ForeName>Ricardo</ForeName><Initials>R</Initials><AffiliationInfo><Affiliation>National Institute of Agricultural Research, INIA, Chile.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Bryson</LastName><ForeName>Rosie</ForeName><Initials>R</Initials><AffiliationInfo><Affiliation>BASF SE, Agricultural Centre, Limburgerhof, Germany.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Thomas</LastName><ForeName>Jane</ForeName><Initials>J</Initials><AffiliationInfo><Affiliation>ational Institute of Agricultural Botany, Cambridge, United Kingdom.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Holdgate</LastName><ForeName>Sarah</ForeName><Initials>S</Initials><AffiliationInfo><Affiliation>ational Institute of Agricultural Botany, Cambridge, United Kingdom.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Saunders</LastName><ForeName>Diane G O</ForeName><Initials>DGO</Initials><AffiliationInfo><Affiliation>Crop Genetics, John Innes Centre, Norwich Research Park, United Kingdom.</Affiliation></AffiliationInfo></Author></AuthorList><Language>eng</Language><GrantList CompleteYN="Y"><Grant><GrantID>BB/P016855/1 </GrantID><Agency>Biotechnology and Biological Sciences Research Council</Agency><Country>United Kingdom</Country></Grant><Grant><GrantID>BB/P012574/1</GrantID><Agency>Biotechnology and Biological Sciences Research Council</Agency><Country>United Kingdom</Country></Grant></GrantList><PublicationTypeList><PublicationType UI="D016428">Journal Article</PublicationType><PublicationType UI="D013485">Research Support, Non-U.S. Gov't</PublicationType></PublicationTypeList></Article><MedlineJournalInfo><Country>England</Country><MedlineTA>Genome Biol Evol</MedlineTA><NlmUniqueID>101509707</NlmUniqueID><ISSNLinking>1759-6653</ISSNLinking></MedlineJournalInfo><CitationSubset>IM</CitationSubset><MeshHeadingList><MeshHeading><DescriptorName UI="D001487" MajorTopicYN="N">Basidiomycota</DescriptorName><QualifierName UI="Q000145" MajorTopicYN="Y">classification</QualifierName><QualifierName UI="Q000235" MajorTopicYN="Y">genetics</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D004196" MajorTopicYN="N">Disease Outbreaks</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D016681" MajorTopicYN="N">Genome, Fungal</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D023281" MajorTopicYN="N">Genomics</DescriptorName><QualifierName UI="Q000379" MajorTopicYN="Y">methods</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D005838" MajorTopicYN="N">Genotype</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D059014" MajorTopicYN="N">High-Throughput Nucleotide Sequencing</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D058507" MajorTopicYN="Y">Host Specificity</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D010802" MajorTopicYN="N">Phylogeny</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D010935" MajorTopicYN="N">Plant Diseases</DescriptorName><QualifierName UI="Q000382" MajorTopicYN="Y">microbiology</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D012621" MajorTopicYN="N">Seasons</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D014908" MajorTopicYN="N">Triticum</DescriptorName><QualifierName UI="Q000382" MajorTopicYN="Y">microbiology</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D014774" MajorTopicYN="N">Virulence</DescriptorName></MeshHeading></MeshHeadingList><KeywordList Owner="NOTNLM"><Keyword MajorTopicYN="Y">Puccinia striiformis</Keyword><Keyword MajorTopicYN="Y">pathogenomics</Keyword><Keyword MajorTopicYN="Y">plant pathology</Keyword><Keyword MajorTopicYN="Y">population genomics</Keyword><Keyword MajorTopicYN="Y">wheat yellow rust</Keyword></KeywordList></MedlineCitation><PubmedData><History><PubMedPubDate PubStatus="accepted"><Year>2017</Year><Month>11</Month><Day>20</Day></PubMedPubDate><PubMedPubDate 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