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<title xml:lang="en">Horizontal gene transfer and gene dosage drives adaptation to wood colonization in a tree pathogen</title>
<author>
<name sortKey="Dhillon, Braham" sort="Dhillon, Braham" uniqKey="Dhillon B" first="Braham" last="Dhillon">Braham Dhillon</name>
<affiliation>
<nlm:aff id="aff1">Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Feau, Nicolas" sort="Feau, Nicolas" uniqKey="Feau N" first="Nicolas" last="Feau">Nicolas Feau</name>
<affiliation>
<nlm:aff id="aff1">Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Aerts, Andrea L" sort="Aerts, Andrea L" uniqKey="Aerts A" first="Andrea L." last="Aerts">Andrea L. Aerts</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Beauseigle, Stephanie" sort="Beauseigle, Stephanie" uniqKey="Beauseigle S" first="Stéphanie" last="Beauseigle">Stéphanie Beauseigle</name>
<affiliation>
<nlm:aff id="aff1">Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Bernier, Louis" sort="Bernier, Louis" uniqKey="Bernier L" first="Louis" last="Bernier">Louis Bernier</name>
<affiliation>
<nlm:aff id="aff3">Centre d’Étude de la Forêt,
<institution>Université Laval</institution>
, Québec, QC,
<country>Canada</country>
G1V 0A6;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Copeland, Alex" sort="Copeland, Alex" uniqKey="Copeland A" first="Alex" last="Copeland">Alex Copeland</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Foster, Adam" sort="Foster, Adam" uniqKey="Foster A" first="Adam" last="Foster">Adam Foster</name>
<affiliation>
<nlm:aff id="aff4">
<institution>Natural Resources Canada</institution>
, Canadian Forest Service, Laurentian Forestry Centre, Québec, QC,
<country>Canada</country>
G1V 4C7;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Gill, Navdeep" sort="Gill, Navdeep" uniqKey="Gill N" first="Navdeep" last="Gill">Navdeep Gill</name>
<affiliation>
<nlm:aff id="aff5">Department of Botany,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Henrissat, Bernard" sort="Henrissat, Bernard" uniqKey="Henrissat B" first="Bernard" last="Henrissat">Bernard Henrissat</name>
<affiliation>
<nlm:aff id="aff6">UMR 7257 Centre National de la Recherche Scientifique,
<institution>Aix-Marseille University</institution>
, 13288 Marseille,
<country>France</country>
;</nlm:aff>
</affiliation>
<affiliation>
<nlm:aff id="aff7">Department of Biological Sciences,
<institution>King Abdulaziz University</institution>
, Jeddah,
<country>Saudi Arabia</country>
;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Herath, Padmini" sort="Herath, Padmini" uniqKey="Herath P" first="Padmini" last="Herath">Padmini Herath</name>
<affiliation>
<nlm:aff id="aff1">Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Labutti, Kurt M" sort="Labutti, Kurt M" uniqKey="Labutti K" first="Kurt M." last="Labutti">Kurt M. Labutti</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Levasseur, Anthony" sort="Levasseur, Anthony" uniqKey="Levasseur A" first="Anthony" last="Levasseur">Anthony Levasseur</name>
<affiliation>
<nlm:aff id="aff8">Unité de Recherche sur les Maladies Infectieuses et Tropicales Emergentes (URMITE), UM63, CNRS 7278, IRD 198, INSERM U1095, IHU Méditerranée Infection, Aix-Marseille University, 13005 Marseille,
<country>France</country>
;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Lindquist, Erika A" sort="Lindquist, Erika A" uniqKey="Lindquist E" first="Erika A." last="Lindquist">Erika A. Lindquist</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Majoor, Eline" sort="Majoor, Eline" uniqKey="Majoor E" first="Eline" last="Majoor">Eline Majoor</name>
<affiliation>
<nlm:aff id="aff9">Fungal Physiology,
<institution>Centraalbureau voor Schimmelcultures</institution>
<institution>Royal Netherlands Academy of Arts and Sciences Fungal Biodiversity Centre (CBS-KNAW)</institution>
, 3584 CT, Utrecht,
<country>The Netherlands</country>
;</nlm:aff>
</affiliation>
<affiliation>
<nlm:aff id="aff10">Fungal Molecular Physiology,
<institution>Utrecht University</institution>
, 3584 CT, Utrecht,
<country>The Netherlands</country>
;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Ohm, Robin A" sort="Ohm, Robin A" uniqKey="Ohm R" first="Robin A." last="Ohm">Robin A. Ohm</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Pangilinan, Jasmyn L" sort="Pangilinan, Jasmyn L" uniqKey="Pangilinan J" first="Jasmyn L." last="Pangilinan">Jasmyn L. Pangilinan</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Pribowo, Amadeus" sort="Pribowo, Amadeus" uniqKey="Pribowo A" first="Amadeus" last="Pribowo">Amadeus Pribowo</name>
<affiliation>
<nlm:aff wicri:cut="; and" id="aff11">Forest Products Biotechnology and Bioenergy,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Saddler, John N" sort="Saddler, John N" uniqKey="Saddler J" first="John N." last="Saddler">John N. Saddler</name>
<affiliation>
<nlm:aff wicri:cut="; and" id="aff11">Forest Products Biotechnology and Bioenergy,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Sakalidis, Monique L" sort="Sakalidis, Monique L" uniqKey="Sakalidis M" first="Monique L." last="Sakalidis">Monique L. Sakalidis</name>
<affiliation>
<nlm:aff id="aff1">Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="De Vries, Ronald P" sort="De Vries, Ronald P" uniqKey="De Vries R" first="Ronald P." last="De Vries">Ronald P. De Vries</name>
<affiliation>
<nlm:aff id="aff9">Fungal Physiology,
<institution>Centraalbureau voor Schimmelcultures</institution>
<institution>Royal Netherlands Academy of Arts and Sciences Fungal Biodiversity Centre (CBS-KNAW)</institution>
, 3584 CT, Utrecht,
<country>The Netherlands</country>
;</nlm:aff>
</affiliation>
<affiliation>
<nlm:aff id="aff10">Fungal Molecular Physiology,
<institution>Utrecht University</institution>
, 3584 CT, Utrecht,
<country>The Netherlands</country>
;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Grigoriev, Igor V" sort="Grigoriev, Igor V" uniqKey="Grigoriev I" first="Igor V." last="Grigoriev">Igor V. Grigoriev</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Goodwin, Stephen B" sort="Goodwin, Stephen B" uniqKey="Goodwin S" first="Stephen B." last="Goodwin">Stephen B. Goodwin</name>
<affiliation>
<nlm:aff id="aff12">US Department of Agriculture–Agricultural Research Service Crop Production and Pest Control Research Unit,
<institution>Purdue University</institution>
, West Lafayette,
<addr-line>IN</addr-line>
47907-2054</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Tanguay, Philippe" sort="Tanguay, Philippe" uniqKey="Tanguay P" first="Philippe" last="Tanguay">Philippe Tanguay</name>
<affiliation>
<nlm:aff id="aff4">
<institution>Natural Resources Canada</institution>
, Canadian Forest Service, Laurentian Forestry Centre, Québec, QC,
<country>Canada</country>
G1V 4C7;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Hamelin, Richard C" sort="Hamelin, Richard C" uniqKey="Hamelin R" first="Richard C." last="Hamelin">Richard C. Hamelin</name>
<affiliation>
<nlm:aff id="aff1">Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
<affiliation>
<nlm:aff id="aff4">
<institution>Natural Resources Canada</institution>
, Canadian Forest Service, Laurentian Forestry Centre, Québec, QC,
<country>Canada</country>
G1V 4C7;</nlm:aff>
</affiliation>
</author>
</titleStmt>
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<idno type="wicri:source">PMC</idno>
<idno type="pmid">25733908</idno>
<idno type="pmc">4371944</idno>
<idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4371944</idno>
<idno type="RBID">PMC:4371944</idno>
<idno type="doi">10.1073/pnas.1424293112</idno>
<date when="2015">2015</date>
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<analytic>
<title xml:lang="en" level="a" type="main">Horizontal gene transfer and gene dosage drives adaptation to wood colonization in a tree pathogen</title>
<author>
<name sortKey="Dhillon, Braham" sort="Dhillon, Braham" uniqKey="Dhillon B" first="Braham" last="Dhillon">Braham Dhillon</name>
<affiliation>
<nlm:aff id="aff1">Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Feau, Nicolas" sort="Feau, Nicolas" uniqKey="Feau N" first="Nicolas" last="Feau">Nicolas Feau</name>
<affiliation>
<nlm:aff id="aff1">Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Aerts, Andrea L" sort="Aerts, Andrea L" uniqKey="Aerts A" first="Andrea L." last="Aerts">Andrea L. Aerts</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Beauseigle, Stephanie" sort="Beauseigle, Stephanie" uniqKey="Beauseigle S" first="Stéphanie" last="Beauseigle">Stéphanie Beauseigle</name>
<affiliation>
<nlm:aff id="aff1">Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Bernier, Louis" sort="Bernier, Louis" uniqKey="Bernier L" first="Louis" last="Bernier">Louis Bernier</name>
<affiliation>
<nlm:aff id="aff3">Centre d’Étude de la Forêt,
<institution>Université Laval</institution>
, Québec, QC,
<country>Canada</country>
G1V 0A6;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Copeland, Alex" sort="Copeland, Alex" uniqKey="Copeland A" first="Alex" last="Copeland">Alex Copeland</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Foster, Adam" sort="Foster, Adam" uniqKey="Foster A" first="Adam" last="Foster">Adam Foster</name>
<affiliation>
<nlm:aff id="aff4">
<institution>Natural Resources Canada</institution>
, Canadian Forest Service, Laurentian Forestry Centre, Québec, QC,
<country>Canada</country>
G1V 4C7;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Gill, Navdeep" sort="Gill, Navdeep" uniqKey="Gill N" first="Navdeep" last="Gill">Navdeep Gill</name>
<affiliation>
<nlm:aff id="aff5">Department of Botany,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Henrissat, Bernard" sort="Henrissat, Bernard" uniqKey="Henrissat B" first="Bernard" last="Henrissat">Bernard Henrissat</name>
<affiliation>
<nlm:aff id="aff6">UMR 7257 Centre National de la Recherche Scientifique,
<institution>Aix-Marseille University</institution>
, 13288 Marseille,
<country>France</country>
;</nlm:aff>
</affiliation>
<affiliation>
<nlm:aff id="aff7">Department of Biological Sciences,
<institution>King Abdulaziz University</institution>
, Jeddah,
<country>Saudi Arabia</country>
;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Herath, Padmini" sort="Herath, Padmini" uniqKey="Herath P" first="Padmini" last="Herath">Padmini Herath</name>
<affiliation>
<nlm:aff id="aff1">Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Labutti, Kurt M" sort="Labutti, Kurt M" uniqKey="Labutti K" first="Kurt M." last="Labutti">Kurt M. Labutti</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Levasseur, Anthony" sort="Levasseur, Anthony" uniqKey="Levasseur A" first="Anthony" last="Levasseur">Anthony Levasseur</name>
<affiliation>
<nlm:aff id="aff8">Unité de Recherche sur les Maladies Infectieuses et Tropicales Emergentes (URMITE), UM63, CNRS 7278, IRD 198, INSERM U1095, IHU Méditerranée Infection, Aix-Marseille University, 13005 Marseille,
<country>France</country>
;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Lindquist, Erika A" sort="Lindquist, Erika A" uniqKey="Lindquist E" first="Erika A." last="Lindquist">Erika A. Lindquist</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Majoor, Eline" sort="Majoor, Eline" uniqKey="Majoor E" first="Eline" last="Majoor">Eline Majoor</name>
<affiliation>
<nlm:aff id="aff9">Fungal Physiology,
<institution>Centraalbureau voor Schimmelcultures</institution>
<institution>Royal Netherlands Academy of Arts and Sciences Fungal Biodiversity Centre (CBS-KNAW)</institution>
, 3584 CT, Utrecht,
<country>The Netherlands</country>
;</nlm:aff>
</affiliation>
<affiliation>
<nlm:aff id="aff10">Fungal Molecular Physiology,
<institution>Utrecht University</institution>
, 3584 CT, Utrecht,
<country>The Netherlands</country>
;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Ohm, Robin A" sort="Ohm, Robin A" uniqKey="Ohm R" first="Robin A." last="Ohm">Robin A. Ohm</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Pangilinan, Jasmyn L" sort="Pangilinan, Jasmyn L" uniqKey="Pangilinan J" first="Jasmyn L." last="Pangilinan">Jasmyn L. Pangilinan</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Pribowo, Amadeus" sort="Pribowo, Amadeus" uniqKey="Pribowo A" first="Amadeus" last="Pribowo">Amadeus Pribowo</name>
<affiliation>
<nlm:aff wicri:cut="; and" id="aff11">Forest Products Biotechnology and Bioenergy,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Saddler, John N" sort="Saddler, John N" uniqKey="Saddler J" first="John N." last="Saddler">John N. Saddler</name>
<affiliation>
<nlm:aff wicri:cut="; and" id="aff11">Forest Products Biotechnology and Bioenergy,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Sakalidis, Monique L" sort="Sakalidis, Monique L" uniqKey="Sakalidis M" first="Monique L." last="Sakalidis">Monique L. Sakalidis</name>
<affiliation>
<nlm:aff id="aff1">Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="De Vries, Ronald P" sort="De Vries, Ronald P" uniqKey="De Vries R" first="Ronald P." last="De Vries">Ronald P. De Vries</name>
<affiliation>
<nlm:aff id="aff9">Fungal Physiology,
<institution>Centraalbureau voor Schimmelcultures</institution>
<institution>Royal Netherlands Academy of Arts and Sciences Fungal Biodiversity Centre (CBS-KNAW)</institution>
, 3584 CT, Utrecht,
<country>The Netherlands</country>
;</nlm:aff>
</affiliation>
<affiliation>
<nlm:aff id="aff10">Fungal Molecular Physiology,
<institution>Utrecht University</institution>
, 3584 CT, Utrecht,
<country>The Netherlands</country>
;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Grigoriev, Igor V" sort="Grigoriev, Igor V" uniqKey="Grigoriev I" first="Igor V." last="Grigoriev">Igor V. Grigoriev</name>
<affiliation>
<nlm:aff id="aff2">
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Goodwin, Stephen B" sort="Goodwin, Stephen B" uniqKey="Goodwin S" first="Stephen B." last="Goodwin">Stephen B. Goodwin</name>
<affiliation>
<nlm:aff id="aff12">US Department of Agriculture–Agricultural Research Service Crop Production and Pest Control Research Unit,
<institution>Purdue University</institution>
, West Lafayette,
<addr-line>IN</addr-line>
47907-2054</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Tanguay, Philippe" sort="Tanguay, Philippe" uniqKey="Tanguay P" first="Philippe" last="Tanguay">Philippe Tanguay</name>
<affiliation>
<nlm:aff id="aff4">
<institution>Natural Resources Canada</institution>
, Canadian Forest Service, Laurentian Forestry Centre, Québec, QC,
<country>Canada</country>
G1V 4C7;</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Hamelin, Richard C" sort="Hamelin, Richard C" uniqKey="Hamelin R" first="Richard C." last="Hamelin">Richard C. Hamelin</name>
<affiliation>
<nlm:aff id="aff1">Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</nlm:aff>
</affiliation>
<affiliation>
<nlm:aff id="aff4">
<institution>Natural Resources Canada</institution>
, Canadian Forest Service, Laurentian Forestry Centre, Québec, QC,
<country>Canada</country>
G1V 4C7;</nlm:aff>
</affiliation>
</author>
</analytic>
<series>
<title level="j">Proceedings of the National Academy of Sciences of the United States of America</title>
<idno type="ISSN">0027-8424</idno>
<idno type="eISSN">1091-6490</idno>
<imprint>
<date when="2015">2015</date>
</imprint>
</series>
</biblStruct>
</sourceDesc>
</fileDesc>
<profileDesc>
<textClass></textClass>
</profileDesc>
</teiHeader>
<front>
<div type="abstract" xml:lang="en">
<title>Significance</title>
<p>Some of the most damaging tree diseases are caused by pathogens that induce cankers, a stem deformation often lethal. To investigate the cause of this adaptation, we sequenced the genomes of poplar pathogens that do and do not cause cankers. We found a unique cluster of genes that produce secondary metabolites and are co-activated when the canker pathogen is grown on poplar wood and leaves. The gene genealogy is discordant with the species phylogeny, showing a signature of horizontal transfer from fungi associated with wood decay. Furthermore, genes encoding hemicellulose-degrading enzymes are up-regulated on poplar wood chips, with some having been acquired horizontally. We propose that adaptation to colonize poplar woody stems is the result of acquisition of these genes.</p>
</div>
</front>
</TEI>
<pmc article-type="research-article">
<pmc-comment>The publisher of this article does not allow downloading of the full text in XML form.</pmc-comment>
<front>
<journal-meta>
<journal-id journal-id-type="nlm-ta">Proc Natl Acad Sci U S A</journal-id>
<journal-id journal-id-type="iso-abbrev">Proc. Natl. Acad. Sci. U.S.A</journal-id>
<journal-id journal-id-type="hwp">pnas</journal-id>
<journal-id journal-id-type="pmc">pnas</journal-id>
<journal-id journal-id-type="publisher-id">PNAS</journal-id>
<journal-title-group>
<journal-title>Proceedings of the National Academy of Sciences of the United States of America</journal-title>
</journal-title-group>
<issn pub-type="ppub">0027-8424</issn>
<issn pub-type="epub">1091-6490</issn>
<publisher>
<publisher-name>National Academy of Sciences</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="pmid">25733908</article-id>
<article-id pub-id-type="pmc">4371944</article-id>
<article-id pub-id-type="publisher-id">201424293</article-id>
<article-id pub-id-type="doi">10.1073/pnas.1424293112</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Biological Sciences</subject>
<subj-group>
<subject>Genetics</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Horizontal gene transfer and gene dosage drives adaptation to wood colonization in a tree pathogen</article-title>
<alt-title alt-title-type="short">Makings of a poplar pathogen</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Dhillon</surname>
<given-names>Braham</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>a</sup>
</xref>
<xref ref-type="author-notes" rid="fn1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Feau</surname>
<given-names>Nicolas</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>a</sup>
</xref>
<xref ref-type="author-notes" rid="fn1">
<sup>1</sup>
</xref>
<xref ref-type="corresp" rid="cor1">
<sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Aerts</surname>
<given-names>Andrea L.</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>b</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Beauseigle</surname>
<given-names>Stéphanie</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>a</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Bernier</surname>
<given-names>Louis</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>c</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Copeland</surname>
<given-names>Alex</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>b</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Foster</surname>
<given-names>Adam</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>d</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Gill</surname>
<given-names>Navdeep</given-names>
</name>
<xref ref-type="aff" rid="aff5">
<sup>e</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Henrissat</surname>
<given-names>Bernard</given-names>
</name>
<xref ref-type="aff" rid="aff6">
<sup>f</sup>
</xref>
<xref ref-type="aff" rid="aff7">
<sup>g</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Herath</surname>
<given-names>Padmini</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>a</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>LaButti</surname>
<given-names>Kurt M.</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>b</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Levasseur</surname>
<given-names>Anthony</given-names>
</name>
<xref ref-type="aff" rid="aff8">
<sup>h</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Lindquist</surname>
<given-names>Erika A.</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>b</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Majoor</surname>
<given-names>Eline</given-names>
</name>
<xref ref-type="aff" rid="aff9">
<sup>i</sup>
</xref>
<xref ref-type="aff" rid="aff10">
<sup>j</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ohm</surname>
<given-names>Robin A.</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>b</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Pangilinan</surname>
<given-names>Jasmyn L.</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>b</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Pribowo</surname>
<given-names>Amadeus</given-names>
</name>
<xref ref-type="aff" rid="aff11">
<sup>k</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Saddler</surname>
<given-names>John N.</given-names>
</name>
<xref ref-type="aff" rid="aff11">
<sup>k</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Sakalidis</surname>
<given-names>Monique L.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>a</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>de Vries</surname>
<given-names>Ronald P.</given-names>
</name>
<xref ref-type="aff" rid="aff9">
<sup>i</sup>
</xref>
<xref ref-type="aff" rid="aff10">
<sup>j</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Grigoriev</surname>
<given-names>Igor V.</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>b</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Goodwin</surname>
<given-names>Stephen B.</given-names>
</name>
<xref ref-type="aff" rid="aff12">
<sup>l</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Tanguay</surname>
<given-names>Philippe</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>d</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Hamelin</surname>
<given-names>Richard C.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>a</sup>
</xref>
<xref ref-type="aff" rid="aff4">
<sup>d</sup>
</xref>
<xref ref-type="corresp" rid="cor1">
<sup>2</sup>
</xref>
</contrib>
<aff id="aff1">
<sup>a</sup>
Department of Forest and Conservation Sciences,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</aff>
<aff id="aff2">
<sup>b</sup>
<institution>US Department of Energy Joint Genome Institute</institution>
, Walnut Creek,
<addr-line>CA</addr-line>
94598;</aff>
<aff id="aff3">
<sup>c</sup>
Centre d’Étude de la Forêt,
<institution>Université Laval</institution>
, Québec, QC,
<country>Canada</country>
G1V 0A6;</aff>
<aff id="aff4">
<sup>d</sup>
<institution>Natural Resources Canada</institution>
, Canadian Forest Service, Laurentian Forestry Centre, Québec, QC,
<country>Canada</country>
G1V 4C7;</aff>
<aff id="aff5">
<sup>e</sup>
Department of Botany,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4;</aff>
<aff id="aff6">
<sup>f</sup>
UMR 7257 Centre National de la Recherche Scientifique,
<institution>Aix-Marseille University</institution>
, 13288 Marseille,
<country>France</country>
;</aff>
<aff id="aff7">
<sup>g</sup>
Department of Biological Sciences,
<institution>King Abdulaziz University</institution>
, Jeddah,
<country>Saudi Arabia</country>
;</aff>
<aff id="aff8">
<sup>h</sup>
Unité de Recherche sur les Maladies Infectieuses et Tropicales Emergentes (URMITE), UM63, CNRS 7278, IRD 198, INSERM U1095, IHU Méditerranée Infection, Aix-Marseille University, 13005 Marseille,
<country>France</country>
;</aff>
<aff id="aff9">
<sup>i</sup>
Fungal Physiology,
<institution>Centraalbureau voor Schimmelcultures</institution>
<institution>Royal Netherlands Academy of Arts and Sciences Fungal Biodiversity Centre (CBS-KNAW)</institution>
, 3584 CT, Utrecht,
<country>The Netherlands</country>
;</aff>
<aff id="aff10">
<sup>j</sup>
Fungal Molecular Physiology,
<institution>Utrecht University</institution>
, 3584 CT, Utrecht,
<country>The Netherlands</country>
;</aff>
<aff id="aff11">
<sup>k</sup>
Forest Products Biotechnology and Bioenergy,
<institution>The University of British Columbia</institution>
, Vancouver, BC,
<country>Canada</country>
V6T 1Z4; and</aff>
<aff id="aff12">
<sup>l</sup>
US Department of Agriculture–Agricultural Research Service Crop Production and Pest Control Research Unit,
<institution>Purdue University</institution>
, West Lafayette,
<addr-line>IN</addr-line>
47907-2054</aff>
</contrib-group>
<author-notes>
<corresp id="cor1">
<sup>2</sup>
To whom correspondence may be addressed. Email:
<email>feaunico@mail.ubc.ca</email>
or
<email>Richard.Hamelin@RNCan-NRCan.gc.ca</email>
.</corresp>
<fn fn-type="edited-by">
<p>Edited by Ronald R. Sederoff, North Carolina State University, Raleigh, NC, and approved January 27, 2015 (received for review December 22, 2014)</p>
</fn>
<fn fn-type="con">
<p>Author contributions: B.D., N.F., I.V.G., S.B.G., P.T., and R.H. designed research; B.D., N.F., A.L.A., S.B., A.C., A.F., P.H., K.M.L., E.A.L., E.M., R.A.O., J.L.P., A.P., J.N.S., R.P.d.V., and P.T. performed research; J.N.S., I.V.G., S.B.G., P.T., and R.H. contributed new reagents/analytic tools; B.D., N.F., A.F., N.G., B.H., A.L., M.L.S., and P.T. analyzed data; and B.D., N.F., L.B., S.B.G., and R.H. wrote the paper.</p>
</fn>
<fn fn-type="equal" id="fn1">
<p>
<sup>1</sup>
B.D. and N.F. contributed equally to this work.</p>
</fn>
<fn fn-type="conflict">
<p>The authors declare no conflict of interest.</p>
</fn>
</author-notes>
<pub-date pub-type="ppub">
<day>17</day>
<month>3</month>
<year>2015</year>
</pub-date>
<pub-date pub-type="epub">
<day>2</day>
<month>3</month>
<year>2015</year>
</pub-date>
<volume>112</volume>
<issue>11</issue>
<fpage>3451</fpage>
<lpage>3456</lpage>
<self-uri xlink:title="pdf" xlink:href="pnas.201424293.pdf"></self-uri>
<abstract abstract-type="executive-summary">
<title>Significance</title>
<p>Some of the most damaging tree diseases are caused by pathogens that induce cankers, a stem deformation often lethal. To investigate the cause of this adaptation, we sequenced the genomes of poplar pathogens that do and do not cause cankers. We found a unique cluster of genes that produce secondary metabolites and are co-activated when the canker pathogen is grown on poplar wood and leaves. The gene genealogy is discordant with the species phylogeny, showing a signature of horizontal transfer from fungi associated with wood decay. Furthermore, genes encoding hemicellulose-degrading enzymes are up-regulated on poplar wood chips, with some having been acquired horizontally. We propose that adaptation to colonize poplar woody stems is the result of acquisition of these genes.</p>
</abstract>
<abstract>
<p>Some of the most damaging tree pathogens can attack woody stems, causing lesions (cankers) that may be lethal. To identify the genomic determinants of wood colonization leading to canker formation, we sequenced the genomes of the poplar canker pathogen,
<italic>Mycosphaerella populorum</italic>
, and the closely related poplar leaf pathogen,
<italic>M. populicola</italic>
. A secondary metabolite cluster unique to
<italic>M. populorum</italic>
is fully activated following induction by poplar wood and leaves. In addition, genes encoding hemicellulose-degrading enzymes, peptidases, and metabolite transporters were more abundant and were up-regulated in
<italic>M. populorum</italic>
growing on poplar wood-chip medium compared with
<italic>M. populicola</italic>
. The secondary gene cluster and several of the carbohydrate degradation genes have the signature of horizontal transfer from ascomycete fungi associated with wood decay and from prokaryotes. Acquisition and maintenance of the gene battery necessary for growth in woody tissues and gene dosage resulting in gene expression reconfiguration appear to be responsible for the adaptation of
<italic>M. populorum</italic>
to infect, colonize, and cause mortality on poplar woody stems.</p>
</abstract>
<kwd-group>
<kwd>poplar pathogen</kwd>
<kwd>tree disease</kwd>
<kwd>fungal genomics</kwd>
<kwd>
<italic>Septoria</italic>
canker</kwd>
</kwd-group>
<funding-group>
<award-group id="gs1">
<funding-source id="sp1">Genome Canada and Genome Bristish Columbia</funding-source>
<award-id rid="sp1">Genome BC project 2112</award-id>
</award-group>
<award-group id="gs2">
<funding-source id="sp2">Office of Science of the U.S. department of Energy</funding-source>
<award-id rid="sp2">DE-AC02-05CH11231</award-id>
</award-group>
</funding-group>
<counts>
<page-count count="6"></page-count>
</counts>
</article-meta>
</front>
</pmc>
</record>

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