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Comparative genomics to explore phylogenetic relationship, cryptic sexual potential and host specificity of Rhynchosporium species on grasses.

Identifieur interne : 001959 ( Main/Exploration ); précédent : 001958; suivant : 001960

Comparative genomics to explore phylogenetic relationship, cryptic sexual potential and host specificity of Rhynchosporium species on grasses.

Auteurs : Daniel Penselin [Allemagne] ; Martin Münsterkötter [Allemagne] ; Susanne Kirsten [Allemagne] ; Marius Felder [Allemagne] ; Stefan Taudien [Allemagne] ; Matthias Platzer [Allemagne] ; Kevin Ashelford [Royaume-Uni] ; Konrad H. Paskiewicz [Royaume-Uni] ; Richard J. Harrison [Royaume-Uni] ; David J. Hughes [Royaume-Uni] ; Thomas Wolf [Allemagne] ; Ekaterina Shelest [Allemagne] ; Jenny Graap [Allemagne] ; Jan Hoffmann [Allemagne] ; Claudia Wenzel [Allemagne, Suisse] ; Nadine Wöltje [Allemagne] ; Kevin M. King [Royaume-Uni] ; Bruce D L. Fitt [Royaume-Uni] ; Ulrich Güldener [Allemagne] ; Anna Avrova [Royaume-Uni] ; Wolfgang Knogge [Allemagne]

Source :

RBID : pubmed:27875982

Descripteurs français

English descriptors

Abstract

BACKGROUND

The Rhynchosporium species complex consists of hemibiotrophic fungal pathogens specialized to different sweet grass species including the cereal crops barley and rye. A sexual stage has not been described, but several lines of evidence suggest the occurrence of sexual reproduction. Therefore, a comparative genomics approach was carried out to disclose the evolutionary relationship of the species and to identify genes demonstrating the potential for a sexual cycle. Furthermore, due to the evolutionary very young age of the five species currently known, this genus appears to be well-suited to address the question at the molecular level of how pathogenic fungi adapt to their hosts.

RESULTS

The genomes of the different Rhynchosporium species were sequenced, assembled and annotated using ab initio gene predictors trained on several fungal genomes as well as on Rhynchosporium expressed sequence tags. Structures of the rDNA regions and genome-wide single nucleotide polymorphisms provided a hypothesis for intra-genus evolution. Homology screening detected core meiotic genes along with most genes crucial for sexual recombination in ascomycete fungi. In addition, a large number of cell wall-degrading enzymes that is characteristic for hemibiotrophic and necrotrophic fungi infecting monocotyledonous hosts were found. Furthermore, the Rhynchosporium genomes carry a repertoire of genes coding for polyketide synthases and non-ribosomal peptide synthetases. Several of these genes are missing from the genome of the closest sequenced relative, the poplar pathogen Marssonina brunnea, and are possibly involved in adaptation to the grass hosts. Most importantly, six species-specific genes coding for protein effectors were identified in R. commune. Their deletion yielded mutants that grew more vigorously in planta than the wild type.

CONCLUSION

Both cryptic sexuality and secondary metabolites may have contributed to host adaptation. Most importantly, however, the growth-retarding activity of the species-specific effectors suggests that host adaptation of R. commune aims at extending the biotrophic stage at the expense of the necrotrophic stage of pathogenesis. Like other apoplastic fungi Rhynchosporium colonizes the intercellular matrix of host leaves relatively slowly without causing symptoms, reminiscent of the development of endophytic fungi. Rhynchosporium may therefore become an object for studying the mutualism-parasitism transition.


DOI: 10.1186/s12864-016-3299-5
PubMed: 27875982
PubMed Central: PMC5118889


Affiliations:


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<name sortKey="Ashelford, Kevin" sort="Ashelford, Kevin" uniqKey="Ashelford K" first="Kevin" last="Ashelford">Kevin Ashelford</name>
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<nlm:affiliation>Institute of Medical Genetics, Cardiff University, Cardiff, UK.</nlm:affiliation>
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<name sortKey="Paskiewicz, Konrad H" sort="Paskiewicz, Konrad H" uniqKey="Paskiewicz K" first="Konrad H" last="Paskiewicz">Konrad H. Paskiewicz</name>
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<nlm:affiliation>Exeter Sequencing Service, Biosciences, University of Exeter, Exeter, UK.</nlm:affiliation>
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<name sortKey="Harrison, Richard J" sort="Harrison, Richard J" uniqKey="Harrison R" first="Richard J" last="Harrison">Richard J. Harrison</name>
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<name sortKey="Graap, Jenny" sort="Graap, Jenny" uniqKey="Graap J" first="Jenny" last="Graap">Jenny Graap</name>
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<nlm:affiliation>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale, Germany.</nlm:affiliation>
<country xml:lang="fr">Allemagne</country>
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<name sortKey="Hoffmann, Jan" sort="Hoffmann, Jan" uniqKey="Hoffmann J" first="Jan" last="Hoffmann">Jan Hoffmann</name>
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<name sortKey="Wenzel, Claudia" sort="Wenzel, Claudia" uniqKey="Wenzel C" first="Claudia" last="Wenzel">Claudia Wenzel</name>
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<nlm:affiliation>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale, Germany.</nlm:affiliation>
<country xml:lang="fr">Allemagne</country>
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<nlm:affiliation>Present address: Food Quality and Nutrition, Agroscope, Bern, Switzerland.</nlm:affiliation>
<country xml:lang="fr">Suisse</country>
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<name sortKey="Woltje, Nadine" sort="Woltje, Nadine" uniqKey="Woltje N" first="Nadine" last="Wöltje">Nadine Wöltje</name>
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<nlm:affiliation>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale, Germany.</nlm:affiliation>
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<name sortKey="King, Kevin M" sort="King, Kevin M" uniqKey="King K" first="Kevin M" last="King">Kevin M. King</name>
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<nlm:affiliation>Biological Chemistry and Crop Protection, Rothamsted Research, Harpenden, Hertfordshire, UK.</nlm:affiliation>
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<name sortKey="Fitt, Bruce D L" sort="Fitt, Bruce D L" uniqKey="Fitt B" first="Bruce D L" last="Fitt">Bruce D L. Fitt</name>
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<name sortKey="Guldener, Ulrich" sort="Guldener, Ulrich" uniqKey="Guldener U" first="Ulrich" last="Güldener">Ulrich Güldener</name>
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<nlm:affiliation>Department of Genome-Oriented Bioinformatics, Technische Universität München, Wissenschaftszentrum Weihenstephan, Freising, Germany.</nlm:affiliation>
<country xml:lang="fr">Allemagne</country>
<wicri:regionArea>Department of Genome-Oriented Bioinformatics, Technische Universität München, Wissenschaftszentrum Weihenstephan, Freising</wicri:regionArea>
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<orgName type="university">Université technique de Munich</orgName>
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<name sortKey="Avrova, Anna" sort="Avrova, Anna" uniqKey="Avrova A" first="Anna" last="Avrova">Anna Avrova</name>
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<nlm:affiliation>Cell and Molecular Sciences, The James Hutton Institute, Invergowrie, Dundee, Scotland.</nlm:affiliation>
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<name sortKey="Knogge, Wolfgang" sort="Knogge, Wolfgang" uniqKey="Knogge W" first="Wolfgang" last="Knogge">Wolfgang Knogge</name>
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<nlm:affiliation>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale, Germany. wknogge@ipb-halle.de.</nlm:affiliation>
<country xml:lang="fr">Allemagne</country>
<wicri:regionArea>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale</wicri:regionArea>
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<name sortKey="Felder, Marius" sort="Felder, Marius" uniqKey="Felder M" first="Marius" last="Felder">Marius Felder</name>
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<name sortKey="Taudien, Stefan" sort="Taudien, Stefan" uniqKey="Taudien S" first="Stefan" last="Taudien">Stefan Taudien</name>
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<nlm:affiliation>Genomic Analysis, Leibniz Institute on Aging, Fritz Lipmann Institute, Jena, Germany.</nlm:affiliation>
<country xml:lang="fr">Allemagne</country>
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<name sortKey="Platzer, Matthias" sort="Platzer, Matthias" uniqKey="Platzer M" first="Matthias" last="Platzer">Matthias Platzer</name>
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<nlm:affiliation>Genomic Analysis, Leibniz Institute on Aging, Fritz Lipmann Institute, Jena, Germany.</nlm:affiliation>
<country xml:lang="fr">Allemagne</country>
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<name sortKey="Ashelford, Kevin" sort="Ashelford, Kevin" uniqKey="Ashelford K" first="Kevin" last="Ashelford">Kevin Ashelford</name>
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<nlm:affiliation>Institute of Medical Genetics, Cardiff University, Cardiff, UK.</nlm:affiliation>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea>Institute of Medical Genetics, Cardiff University, Cardiff</wicri:regionArea>
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<name sortKey="Paskiewicz, Konrad H" sort="Paskiewicz, Konrad H" uniqKey="Paskiewicz K" first="Konrad H" last="Paskiewicz">Konrad H. Paskiewicz</name>
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<nlm:affiliation>Exeter Sequencing Service, Biosciences, University of Exeter, Exeter, UK.</nlm:affiliation>
<country xml:lang="fr">Royaume-Uni</country>
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<name sortKey="Harrison, Richard J" sort="Harrison, Richard J" uniqKey="Harrison R" first="Richard J" last="Harrison">Richard J. Harrison</name>
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<nlm:affiliation>NIAB EMR, East Malling, UK.</nlm:affiliation>
<country xml:lang="fr">Royaume-Uni</country>
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<name sortKey="Hughes, David J" sort="Hughes, David J" uniqKey="Hughes D" first="David J" last="Hughes">David J. Hughes</name>
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<nlm:affiliation>Applied Bioinformatics, Rothamsted Research, Harpenden, Hertfordshire, UK.</nlm:affiliation>
<country xml:lang="fr">Royaume-Uni</country>
<wicri:regionArea>Applied Bioinformatics, Rothamsted Research, Harpenden, Hertfordshire</wicri:regionArea>
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<name sortKey="Wolf, Thomas" sort="Wolf, Thomas" uniqKey="Wolf T" first="Thomas" last="Wolf">Thomas Wolf</name>
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<name sortKey="Graap, Jenny" sort="Graap, Jenny" uniqKey="Graap J" first="Jenny" last="Graap">Jenny Graap</name>
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<nlm:affiliation>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale, Germany.</nlm:affiliation>
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<name sortKey="Hoffmann, Jan" sort="Hoffmann, Jan" uniqKey="Hoffmann J" first="Jan" last="Hoffmann">Jan Hoffmann</name>
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<nlm:affiliation>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale, Germany.</nlm:affiliation>
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<name sortKey="Wenzel, Claudia" sort="Wenzel, Claudia" uniqKey="Wenzel C" first="Claudia" last="Wenzel">Claudia Wenzel</name>
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<nlm:affiliation>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale, Germany.</nlm:affiliation>
<country xml:lang="fr">Allemagne</country>
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<nlm:affiliation>Present address: Food Quality and Nutrition, Agroscope, Bern, Switzerland.</nlm:affiliation>
<country xml:lang="fr">Suisse</country>
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<name sortKey="Woltje, Nadine" sort="Woltje, Nadine" uniqKey="Woltje N" first="Nadine" last="Wöltje">Nadine Wöltje</name>
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<nlm:affiliation>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale, Germany.</nlm:affiliation>
<country xml:lang="fr">Allemagne</country>
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<author>
<name sortKey="King, Kevin M" sort="King, Kevin M" uniqKey="King K" first="Kevin M" last="King">Kevin M. King</name>
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<nlm:affiliation>Biological Chemistry and Crop Protection, Rothamsted Research, Harpenden, Hertfordshire, UK.</nlm:affiliation>
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<name sortKey="Fitt, Bruce D L" sort="Fitt, Bruce D L" uniqKey="Fitt B" first="Bruce D L" last="Fitt">Bruce D L. Fitt</name>
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<name sortKey="Guldener, Ulrich" sort="Guldener, Ulrich" uniqKey="Guldener U" first="Ulrich" last="Güldener">Ulrich Güldener</name>
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<nlm:affiliation>Department of Genome-Oriented Bioinformatics, Technische Universität München, Wissenschaftszentrum Weihenstephan, Freising, Germany.</nlm:affiliation>
<country xml:lang="fr">Allemagne</country>
<wicri:regionArea>Department of Genome-Oriented Bioinformatics, Technische Universität München, Wissenschaftszentrum Weihenstephan, Freising</wicri:regionArea>
<wicri:noRegion>Freising</wicri:noRegion>
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<region type="district" nuts="2">District de Haute-Bavière</region>
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<name sortKey="Avrova, Anna" sort="Avrova, Anna" uniqKey="Avrova A" first="Anna" last="Avrova">Anna Avrova</name>
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<nlm:affiliation>Cell and Molecular Sciences, The James Hutton Institute, Invergowrie, Dundee, Scotland.</nlm:affiliation>
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<placeName>
<region type="country">Écosse</region>
</placeName>
<wicri:cityArea>Cell and Molecular Sciences, The James Hutton Institute, Invergowrie, Dundee</wicri:cityArea>
</affiliation>
</author>
<author>
<name sortKey="Knogge, Wolfgang" sort="Knogge, Wolfgang" uniqKey="Knogge W" first="Wolfgang" last="Knogge">Wolfgang Knogge</name>
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<nlm:affiliation>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale, Germany. wknogge@ipb-halle.de.</nlm:affiliation>
<country xml:lang="fr">Allemagne</country>
<wicri:regionArea>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale</wicri:regionArea>
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<series>
<title level="j">BMC genomics</title>
<idno type="eISSN">1471-2164</idno>
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<date when="2016" type="published">2016</date>
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<term>Amino Acid Sequence (MeSH)</term>
<term>Ascomycota (classification)</term>
<term>Ascomycota (genetics)</term>
<term>Ascomycota (metabolism)</term>
<term>DNA, Intergenic (MeSH)</term>
<term>Gene Expression Regulation, Fungal (MeSH)</term>
<term>Genes, Fungal (MeSH)</term>
<term>Genome, Fungal (MeSH)</term>
<term>Genomics (methods)</term>
<term>Host Specificity (MeSH)</term>
<term>Multigene Family (MeSH)</term>
<term>Phylogeny (MeSH)</term>
<term>Poaceae (microbiology)</term>
<term>Secondary Metabolism (genetics)</term>
</keywords>
<keywords scheme="KwdFr" xml:lang="fr">
<term>ADN intergénique (MeSH)</term>
<term>Ascomycota (classification)</term>
<term>Ascomycota (génétique)</term>
<term>Ascomycota (métabolisme)</term>
<term>Famille multigénique (MeSH)</term>
<term>Gènes fongiques (MeSH)</term>
<term>Génome fongique (MeSH)</term>
<term>Génomique (méthodes)</term>
<term>Métabolisme secondaire (génétique)</term>
<term>Phylogenèse (MeSH)</term>
<term>Poaceae (microbiologie)</term>
<term>Régulation de l'expression des gènes fongiques (MeSH)</term>
<term>Spécificité d'hôte (MeSH)</term>
<term>Séquence d'acides aminés (MeSH)</term>
</keywords>
<keywords scheme="MESH" type="chemical" xml:lang="en">
<term>DNA, Intergenic</term>
</keywords>
<keywords scheme="MESH" qualifier="classification" xml:lang="en">
<term>Ascomycota</term>
</keywords>
<keywords scheme="MESH" qualifier="genetics" xml:lang="en">
<term>Ascomycota</term>
<term>Secondary Metabolism</term>
</keywords>
<keywords scheme="MESH" qualifier="génétique" xml:lang="fr">
<term>Ascomycota</term>
<term>Métabolisme secondaire</term>
</keywords>
<keywords scheme="MESH" qualifier="metabolism" xml:lang="en">
<term>Ascomycota</term>
</keywords>
<keywords scheme="MESH" qualifier="methods" xml:lang="en">
<term>Genomics</term>
</keywords>
<keywords scheme="MESH" qualifier="microbiologie" xml:lang="fr">
<term>Poaceae</term>
</keywords>
<keywords scheme="MESH" qualifier="microbiology" xml:lang="en">
<term>Poaceae</term>
</keywords>
<keywords scheme="MESH" qualifier="métabolisme" xml:lang="fr">
<term>Ascomycota</term>
</keywords>
<keywords scheme="MESH" qualifier="méthodes" xml:lang="fr">
<term>Génomique</term>
</keywords>
<keywords scheme="MESH" xml:lang="en">
<term>Amino Acid Sequence</term>
<term>Gene Expression Regulation, Fungal</term>
<term>Genes, Fungal</term>
<term>Genome, Fungal</term>
<term>Host Specificity</term>
<term>Multigene Family</term>
<term>Phylogeny</term>
</keywords>
<keywords scheme="MESH" xml:lang="fr">
<term>ADN intergénique</term>
<term>Ascomycota</term>
<term>Famille multigénique</term>
<term>Gènes fongiques</term>
<term>Génome fongique</term>
<term>Phylogenèse</term>
<term>Régulation de l'expression des gènes fongiques</term>
<term>Spécificité d'hôte</term>
<term>Séquence d'acides aminés</term>
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<front>
<div type="abstract" xml:lang="en">
<p>
<b>BACKGROUND</b>
</p>
<p>The Rhynchosporium species complex consists of hemibiotrophic fungal pathogens specialized to different sweet grass species including the cereal crops barley and rye. A sexual stage has not been described, but several lines of evidence suggest the occurrence of sexual reproduction. Therefore, a comparative genomics approach was carried out to disclose the evolutionary relationship of the species and to identify genes demonstrating the potential for a sexual cycle. Furthermore, due to the evolutionary very young age of the five species currently known, this genus appears to be well-suited to address the question at the molecular level of how pathogenic fungi adapt to their hosts.</p>
</div>
<div type="abstract" xml:lang="en">
<p>
<b>RESULTS</b>
</p>
<p>The genomes of the different Rhynchosporium species were sequenced, assembled and annotated using ab initio gene predictors trained on several fungal genomes as well as on Rhynchosporium expressed sequence tags. Structures of the rDNA regions and genome-wide single nucleotide polymorphisms provided a hypothesis for intra-genus evolution. Homology screening detected core meiotic genes along with most genes crucial for sexual recombination in ascomycete fungi. In addition, a large number of cell wall-degrading enzymes that is characteristic for hemibiotrophic and necrotrophic fungi infecting monocotyledonous hosts were found. Furthermore, the Rhynchosporium genomes carry a repertoire of genes coding for polyketide synthases and non-ribosomal peptide synthetases. Several of these genes are missing from the genome of the closest sequenced relative, the poplar pathogen Marssonina brunnea, and are possibly involved in adaptation to the grass hosts. Most importantly, six species-specific genes coding for protein effectors were identified in R. commune. Their deletion yielded mutants that grew more vigorously in planta than the wild type.</p>
</div>
<div type="abstract" xml:lang="en">
<p>
<b>CONCLUSION</b>
</p>
<p>Both cryptic sexuality and secondary metabolites may have contributed to host adaptation. Most importantly, however, the growth-retarding activity of the species-specific effectors suggests that host adaptation of R. commune aims at extending the biotrophic stage at the expense of the necrotrophic stage of pathogenesis. Like other apoplastic fungi Rhynchosporium colonizes the intercellular matrix of host leaves relatively slowly without causing symptoms, reminiscent of the development of endophytic fungi. Rhynchosporium may therefore become an object for studying the mutualism-parasitism transition.</p>
</div>
</front>
</TEI>
<pubmed>
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<PMID Version="1">27875982</PMID>
<DateCompleted>
<Year>2017</Year>
<Month>08</Month>
<Day>25</Day>
</DateCompleted>
<DateRevised>
<Year>2019</Year>
<Month>10</Month>
<Day>08</Day>
</DateRevised>
<Article PubModel="Electronic">
<Journal>
<ISSN IssnType="Electronic">1471-2164</ISSN>
<JournalIssue CitedMedium="Internet">
<Volume>17</Volume>
<Issue>1</Issue>
<PubDate>
<Year>2016</Year>
<Month>Nov</Month>
<Day>22</Day>
</PubDate>
</JournalIssue>
<Title>BMC genomics</Title>
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<ArticleTitle>Comparative genomics to explore phylogenetic relationship, cryptic sexual potential and host specificity of Rhynchosporium species on grasses.</ArticleTitle>
<Pagination>
<MedlinePgn>953</MedlinePgn>
</Pagination>
<Abstract>
<AbstractText Label="BACKGROUND">The Rhynchosporium species complex consists of hemibiotrophic fungal pathogens specialized to different sweet grass species including the cereal crops barley and rye. A sexual stage has not been described, but several lines of evidence suggest the occurrence of sexual reproduction. Therefore, a comparative genomics approach was carried out to disclose the evolutionary relationship of the species and to identify genes demonstrating the potential for a sexual cycle. Furthermore, due to the evolutionary very young age of the five species currently known, this genus appears to be well-suited to address the question at the molecular level of how pathogenic fungi adapt to their hosts.</AbstractText>
<AbstractText Label="RESULTS">The genomes of the different Rhynchosporium species were sequenced, assembled and annotated using ab initio gene predictors trained on several fungal genomes as well as on Rhynchosporium expressed sequence tags. Structures of the rDNA regions and genome-wide single nucleotide polymorphisms provided a hypothesis for intra-genus evolution. Homology screening detected core meiotic genes along with most genes crucial for sexual recombination in ascomycete fungi. In addition, a large number of cell wall-degrading enzymes that is characteristic for hemibiotrophic and necrotrophic fungi infecting monocotyledonous hosts were found. Furthermore, the Rhynchosporium genomes carry a repertoire of genes coding for polyketide synthases and non-ribosomal peptide synthetases. Several of these genes are missing from the genome of the closest sequenced relative, the poplar pathogen Marssonina brunnea, and are possibly involved in adaptation to the grass hosts. Most importantly, six species-specific genes coding for protein effectors were identified in R. commune. Their deletion yielded mutants that grew more vigorously in planta than the wild type.</AbstractText>
<AbstractText Label="CONCLUSION">Both cryptic sexuality and secondary metabolites may have contributed to host adaptation. Most importantly, however, the growth-retarding activity of the species-specific effectors suggests that host adaptation of R. commune aims at extending the biotrophic stage at the expense of the necrotrophic stage of pathogenesis. Like other apoplastic fungi Rhynchosporium colonizes the intercellular matrix of host leaves relatively slowly without causing symptoms, reminiscent of the development of endophytic fungi. Rhynchosporium may therefore become an object for studying the mutualism-parasitism transition.</AbstractText>
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<LastName>Penselin</LastName>
<ForeName>Daniel</ForeName>
<Initials>D</Initials>
<AffiliationInfo>
<Affiliation>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale, Germany.</Affiliation>
</AffiliationInfo>
</Author>
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<LastName>Münsterkötter</LastName>
<ForeName>Martin</ForeName>
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</AffiliationInfo>
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<LastName>Kirsten</LastName>
<ForeName>Susanne</ForeName>
<Initials>S</Initials>
<AffiliationInfo>
<Affiliation>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale, Germany.</Affiliation>
</AffiliationInfo>
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<LastName>Felder</LastName>
<ForeName>Marius</ForeName>
<Initials>M</Initials>
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<Affiliation>Genomic Analysis, Leibniz Institute on Aging, Fritz Lipmann Institute, Jena, Germany.</Affiliation>
</AffiliationInfo>
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<ForeName>Stefan</ForeName>
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</AffiliationInfo>
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<ForeName>Kevin</ForeName>
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<LastName>Hughes</LastName>
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</AffiliationInfo>
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<LastName>Wolf</LastName>
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<AffiliationInfo>
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</AffiliationInfo>
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<ForeName>Ekaterina</ForeName>
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<LastName>Wenzel</LastName>
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<Affiliation>Department of Stress and Developmental Biology, Leibniz Institute of Plant Biochemistry, Halle/Saale, Germany.</Affiliation>
</AffiliationInfo>
<AffiliationInfo>
<Affiliation>Present address: Food Quality and Nutrition, Agroscope, Bern, Switzerland.</Affiliation>
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<LastName>King</LastName>
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<LastName>Avrova</LastName>
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<Affiliation>Cell and Molecular Sciences, The James Hutton Institute, Invergowrie, Dundee, Scotland.</Affiliation>
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