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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">RXLR-Mediated Entry of <italic>Phytophthora sojae</italic>
Effector <italic>Avr1b</italic>
into Soybean Cells Does Not Require Pathogen-Encoded Machinery<xref ref-type="fn" rid="fn4">[W]</xref>
</title>
<author><name sortKey="Dou, Daolong" sort="Dou, Daolong" uniqKey="Dou D" first="Daolong" last="Dou">Daolong Dou</name>
</author>
<author><name sortKey="Kale, Shiv D" sort="Kale, Shiv D" uniqKey="Kale S" first="Shiv D." last="Kale">Shiv D. Kale</name>
</author>
<author><name sortKey="Wang, Xia" sort="Wang, Xia" uniqKey="Wang X" first="Xia" last="Wang">Xia Wang</name>
</author>
<author><name sortKey="Jiang, Rays H Y" sort="Jiang, Rays H Y" uniqKey="Jiang R" first="Rays H. Y." last="Jiang">Rays H. Y. Jiang</name>
</author>
<author><name sortKey="Bruce, Nathan A" sort="Bruce, Nathan A" uniqKey="Bruce N" first="Nathan A." last="Bruce">Nathan A. Bruce</name>
</author>
<author><name sortKey="Arredondo, Felipe D" sort="Arredondo, Felipe D" uniqKey="Arredondo F" first="Felipe D." last="Arredondo">Felipe D. Arredondo</name>
</author>
<author><name sortKey="Zhang, Xuemin" sort="Zhang, Xuemin" uniqKey="Zhang X" first="Xuemin" last="Zhang">Xuemin Zhang</name>
</author>
<author><name sortKey="Tyler, Brett M" sort="Tyler, Brett M" uniqKey="Tyler B" first="Brett M." last="Tyler">Brett M. Tyler</name>
</author>
</titleStmt>
<publicationStmt><idno type="wicri:source">PMC</idno>
<idno type="pmid">18621946</idno>
<idno type="pmc">2518231</idno>
<idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2518231</idno>
<idno type="RBID">PMC:2518231</idno>
<idno type="doi">10.1105/tpc.107.056093</idno>
<date when="2008">2008</date>
<idno type="wicri:Area/Pmc/Corpus">000283</idno>
<idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000283</idno>
</publicationStmt>
<sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">RXLR-Mediated Entry of <italic>Phytophthora sojae</italic>
Effector <italic>Avr1b</italic>
into Soybean Cells Does Not Require Pathogen-Encoded Machinery<xref ref-type="fn" rid="fn4">[W]</xref>
</title>
<author><name sortKey="Dou, Daolong" sort="Dou, Daolong" uniqKey="Dou D" first="Daolong" last="Dou">Daolong Dou</name>
</author>
<author><name sortKey="Kale, Shiv D" sort="Kale, Shiv D" uniqKey="Kale S" first="Shiv D." last="Kale">Shiv D. Kale</name>
</author>
<author><name sortKey="Wang, Xia" sort="Wang, Xia" uniqKey="Wang X" first="Xia" last="Wang">Xia Wang</name>
</author>
<author><name sortKey="Jiang, Rays H Y" sort="Jiang, Rays H Y" uniqKey="Jiang R" first="Rays H. Y." last="Jiang">Rays H. Y. Jiang</name>
</author>
<author><name sortKey="Bruce, Nathan A" sort="Bruce, Nathan A" uniqKey="Bruce N" first="Nathan A." last="Bruce">Nathan A. Bruce</name>
</author>
<author><name sortKey="Arredondo, Felipe D" sort="Arredondo, Felipe D" uniqKey="Arredondo F" first="Felipe D." last="Arredondo">Felipe D. Arredondo</name>
</author>
<author><name sortKey="Zhang, Xuemin" sort="Zhang, Xuemin" uniqKey="Zhang X" first="Xuemin" last="Zhang">Xuemin Zhang</name>
</author>
<author><name sortKey="Tyler, Brett M" sort="Tyler, Brett M" uniqKey="Tyler B" first="Brett M." last="Tyler">Brett M. Tyler</name>
</author>
</analytic>
<series><title level="j">The Plant Cell</title>
<idno type="ISSN">1040-4651</idno>
<idno type="eISSN">1532-298X</idno>
<imprint><date when="2008">2008</date>
</imprint>
</series>
</biblStruct>
</sourceDesc>
</fileDesc>
<profileDesc><textClass></textClass>
</profileDesc>
</teiHeader>
<front><div type="abstract" xml:lang="en"><p>Effector proteins secreted by oomycete and fungal pathogens have been inferred to enter host cells, where they interact with host resistance gene products. Using the effector protein Avr1b of <italic>Phytophthora sojae</italic>
, an oomycete pathogen of soybean (<italic>Glycine max</italic>
), we show that a pair of sequence motifs, RXLR and dEER, plus surrounding sequences, are both necessary and sufficient to deliver the protein into plant cells. Particle bombardment experiments demonstrate that these motifs function in the absence of the pathogen, indicating that no additional pathogen-encoded machinery is required for effector protein entry into host cells. Furthermore, fusion of the Avr1b RXLR-dEER domain to green fluorescent protein (GFP) allows GFP to enter soybean root cells autonomously. The conclusion that RXLR and dEER serve to transduce oomycete effectors into host cells indicates that the >370 RXLR-dEER–containing proteins encoded in the genome sequence of <italic>P. sojae</italic>
are candidate effectors. We further show that the RXLR and dEER motifs can be replaced by the closely related erythrocyte targeting signals found in effector proteins of <italic>Plasmodium</italic>
, the protozoan that causes malaria in humans. Mutational analysis of the RXLR motif shows that the required residues are very similar in the motifs of <italic>Plasmodium</italic>
and <italic>Phytophthora</italic>
. Thus, the machinery of the hosts (soybean and human) targeted by the effectors may be very ancient.</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">Plant Cell</journal-id>
<journal-id journal-id-type="publisher-id">plantcell</journal-id>
<journal-title>The Plant Cell</journal-title>
<issn pub-type="ppub">1040-4651</issn>
<issn pub-type="epub">1532-298X</issn>
<publisher><publisher-name>American Society of Plant Biologists</publisher-name>
</publisher>
</journal-meta>
<article-meta><article-id pub-id-type="pmid">18621946</article-id>
<article-id pub-id-type="pmc">2518231</article-id>
<article-id pub-id-type="publisher-id">056093</article-id>
<article-id pub-id-type="doi">10.1105/tpc.107.056093</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Research Articles</subject>
</subj-group>
</article-categories>
<title-group><article-title>RXLR-Mediated Entry of <italic>Phytophthora sojae</italic>
Effector <italic>Avr1b</italic>
into Soybean Cells Does Not Require Pathogen-Encoded Machinery<xref ref-type="fn" rid="fn4">[W]</xref>
</article-title>
</title-group>
<contrib-group><contrib contrib-type="author"><name><surname>Dou</surname>
<given-names>Daolong</given-names>
</name>
<xref ref-type="fn" rid="fn1">1</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Kale</surname>
<given-names>Shiv D.</given-names>
</name>
<xref ref-type="fn" rid="fn1">1</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Wang</surname>
<given-names>Xia</given-names>
</name>
</contrib>
<contrib contrib-type="author"><name><surname>Jiang</surname>
<given-names>Rays H.Y.</given-names>
</name>
<xref ref-type="fn" rid="fn2">2</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Bruce</surname>
<given-names>Nathan A.</given-names>
</name>
</contrib>
<contrib contrib-type="author"><name><surname>Arredondo</surname>
<given-names>Felipe D.</given-names>
</name>
</contrib>
<contrib contrib-type="author"><name><surname>Zhang</surname>
<given-names>Xuemin</given-names>
</name>
<xref ref-type="fn" rid="fn3">3</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Tyler</surname>
<given-names>Brett M.</given-names>
</name>
<xref ref-type="corresp" rid="cor1">4</xref>
</contrib>
</contrib-group>
<aff id="N0x1d1bf10N0x44975e8">Virginia Bioinformatics Institute, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061</aff>
<author-notes><fn id="fn1"><label>1</label>
<p>These authors contributed equally to this work.</p>
</fn>
<fn id="fn2"><label>2</label>
<p>Current address: Laboratory of Phytopathology, Wageningen University, NL-6709 PD Wageningen, The Netherlands.</p>
</fn>
<fn id="fn3"><label>3</label>
<p>Current address: Center for Biosystems Research, University of Maryland Biotechnology Institute, 9600 Gudelsky Dr., Rockville, MD 20850.</p>
</fn>
<fn id="cor1"><label>4</label>
<p>Address correspondence to <email>bmtyler@vt.edu</email>
.</p>
</fn>
</author-notes>
<pub-date pub-type="ppub"><month>7</month>
<year>2008</year>
</pub-date>
<volume>20</volume>
<issue>7</issue>
<fpage>1930</fpage>
<lpage>1947</lpage>
<history><date date-type="received"><day>11</day>
<month>10</month>
<year>2007</year>
</date>
<date date-type="rev-recd"><day>6</day>
<month>6</month>
<year>2008</year>
</date>
<date date-type="accepted"><day>27</day>
<month>6</month>
<year>2008</year>
</date>
</history>
<permissions><copyright-statement>Copyright © 2008, American Society of Plant Biologists</copyright-statement>
</permissions>
<abstract><p>Effector proteins secreted by oomycete and fungal pathogens have been inferred to enter host cells, where they interact with host resistance gene products. Using the effector protein Avr1b of <italic>Phytophthora sojae</italic>
, an oomycete pathogen of soybean (<italic>Glycine max</italic>
), we show that a pair of sequence motifs, RXLR and dEER, plus surrounding sequences, are both necessary and sufficient to deliver the protein into plant cells. Particle bombardment experiments demonstrate that these motifs function in the absence of the pathogen, indicating that no additional pathogen-encoded machinery is required for effector protein entry into host cells. Furthermore, fusion of the Avr1b RXLR-dEER domain to green fluorescent protein (GFP) allows GFP to enter soybean root cells autonomously. The conclusion that RXLR and dEER serve to transduce oomycete effectors into host cells indicates that the >370 RXLR-dEER–containing proteins encoded in the genome sequence of <italic>P. sojae</italic>
are candidate effectors. We further show that the RXLR and dEER motifs can be replaced by the closely related erythrocyte targeting signals found in effector proteins of <italic>Plasmodium</italic>
, the protozoan that causes malaria in humans. Mutational analysis of the RXLR motif shows that the required residues are very similar in the motifs of <italic>Plasmodium</italic>
and <italic>Phytophthora</italic>
. Thus, the machinery of the hosts (soybean and human) targeted by the effectors may be very ancient.</p>
</abstract>
</article-meta>
<notes><fn-group><fn><p>The author responsible for distribution of materials integral to the findings presented in this article in accordance with the policy described in the Instructions for Authors (<ext-link ext-link-type="uri" xlink:href="www.plantcell.org">www.plantcell.org</ext-link>
) is: Brett M. Tyler (<email>bmtyler@vt.edu</email>
).</p>
</fn>
<fn id="fn4"><label>[W]</label>
<p>Online version contains Web-only data.</p>
</fn>
<fn><p><ext-link ext-link-type="uri" xlink:href="www.plantcell.org/cgi/doi/10.1105/tpc.107.056093">www.plantcell.org/cgi/doi/10.1105/tpc.107.056093</ext-link>
</p>
</fn>
</fn-group>
</notes>
</front>
</pmc>
</record>
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