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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Evolution of Disease Defense Genes and Their Regulators in Plants</title>
<author><name sortKey="Zhang, Rongzhi" sort="Zhang, Rongzhi" uniqKey="Zhang R" first="Rongzhi" last="Zhang">Rongzhi Zhang</name>
<affiliation><nlm:aff id="af1-ijms-20-00335">Institute of Crop Science, Shandong Academy of Agricultural Sciences, Key Laboratory of Wheat Biology & Genetic Improvement on North Yellow & Huai River Valley, Ministry of Agriculture, National Engineering Laboratory for Wheat & Maize, Jinan 250100, China;<email>zhangrongzhi1981@126.com</email>
(R.Z.);<email>zsjhappy@163.com</email>
(S.Z.);<email>lgy111@126.com</email>
(G.L.)</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Zheng, Fengya" sort="Zheng, Fengya" uniqKey="Zheng F" first="Fengya" last="Zheng">Fengya Zheng</name>
<affiliation><nlm:aff id="af2-ijms-20-00335">BGI Institute of Applied Agriculture, BGI-Agro, Shenzhen 518083, China;<email>zhengfengya@genomics.cn</email>
</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Wei, Shugen" sort="Wei, Shugen" uniqKey="Wei S" first="Shugen" last="Wei">Shugen Wei</name>
<affiliation><nlm:aff id="af3-ijms-20-00335">Guangxi Botanical Garden of Medicinal Plants, Nanning 530023, China;<email>weishugen2@163.com</email>
</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Zhang, Shujuan" sort="Zhang, Shujuan" uniqKey="Zhang S" first="Shujuan" last="Zhang">Shujuan Zhang</name>
<affiliation><nlm:aff id="af1-ijms-20-00335">Institute of Crop Science, Shandong Academy of Agricultural Sciences, Key Laboratory of Wheat Biology & Genetic Improvement on North Yellow & Huai River Valley, Ministry of Agriculture, National Engineering Laboratory for Wheat & Maize, Jinan 250100, China;<email>zhangrongzhi1981@126.com</email>
(R.Z.);<email>zsjhappy@163.com</email>
(S.Z.);<email>lgy111@126.com</email>
(G.L.)</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Li, Genying" sort="Li, Genying" uniqKey="Li G" first="Genying" last="Li">Genying Li</name>
<affiliation><nlm:aff id="af1-ijms-20-00335">Institute of Crop Science, Shandong Academy of Agricultural Sciences, Key Laboratory of Wheat Biology & Genetic Improvement on North Yellow & Huai River Valley, Ministry of Agriculture, National Engineering Laboratory for Wheat & Maize, Jinan 250100, China;<email>zhangrongzhi1981@126.com</email>
(R.Z.);<email>zsjhappy@163.com</email>
(S.Z.);<email>lgy111@126.com</email>
(G.L.)</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Cao, Peijian" sort="Cao, Peijian" uniqKey="Cao P" first="Peijian" last="Cao">Peijian Cao</name>
<affiliation><nlm:aff id="af4-ijms-20-00335">China Tobacco Gene Research Center, Zhengzhou Tobacco Research Institute of CNTC, Zhengzhou 450001, China</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Zhao, Shancen" sort="Zhao, Shancen" uniqKey="Zhao S" first="Shancen" last="Zhao">Shancen Zhao</name>
<affiliation><nlm:aff id="af2-ijms-20-00335">BGI Institute of Applied Agriculture, BGI-Agro, Shenzhen 518083, China;<email>zhengfengya@genomics.cn</email>
</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="af4-ijms-20-00335">China Tobacco Gene Research Center, Zhengzhou Tobacco Research Institute of CNTC, Zhengzhou 450001, China</nlm:aff>
</affiliation>
</author>
</titleStmt>
<publicationStmt><idno type="wicri:source">PMC</idno>
<idno type="pmid">30650550</idno>
<idno type="pmc">6358896</idno>
<idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6358896</idno>
<idno type="RBID">PMC:6358896</idno>
<idno type="doi">10.3390/ijms20020335</idno>
<date when="2019">2019</date>
<idno type="wicri:Area/Pmc/Corpus">000C27</idno>
<idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000C27</idno>
</publicationStmt>
<sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Evolution of Disease Defense Genes and Their Regulators in Plants</title>
<author><name sortKey="Zhang, Rongzhi" sort="Zhang, Rongzhi" uniqKey="Zhang R" first="Rongzhi" last="Zhang">Rongzhi Zhang</name>
<affiliation><nlm:aff id="af1-ijms-20-00335">Institute of Crop Science, Shandong Academy of Agricultural Sciences, Key Laboratory of Wheat Biology & Genetic Improvement on North Yellow & Huai River Valley, Ministry of Agriculture, National Engineering Laboratory for Wheat & Maize, Jinan 250100, China;<email>zhangrongzhi1981@126.com</email>
(R.Z.);<email>zsjhappy@163.com</email>
(S.Z.);<email>lgy111@126.com</email>
(G.L.)</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Zheng, Fengya" sort="Zheng, Fengya" uniqKey="Zheng F" first="Fengya" last="Zheng">Fengya Zheng</name>
<affiliation><nlm:aff id="af2-ijms-20-00335">BGI Institute of Applied Agriculture, BGI-Agro, Shenzhen 518083, China;<email>zhengfengya@genomics.cn</email>
</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Wei, Shugen" sort="Wei, Shugen" uniqKey="Wei S" first="Shugen" last="Wei">Shugen Wei</name>
<affiliation><nlm:aff id="af3-ijms-20-00335">Guangxi Botanical Garden of Medicinal Plants, Nanning 530023, China;<email>weishugen2@163.com</email>
</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Zhang, Shujuan" sort="Zhang, Shujuan" uniqKey="Zhang S" first="Shujuan" last="Zhang">Shujuan Zhang</name>
<affiliation><nlm:aff id="af1-ijms-20-00335">Institute of Crop Science, Shandong Academy of Agricultural Sciences, Key Laboratory of Wheat Biology & Genetic Improvement on North Yellow & Huai River Valley, Ministry of Agriculture, National Engineering Laboratory for Wheat & Maize, Jinan 250100, China;<email>zhangrongzhi1981@126.com</email>
(R.Z.);<email>zsjhappy@163.com</email>
(S.Z.);<email>lgy111@126.com</email>
(G.L.)</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Li, Genying" sort="Li, Genying" uniqKey="Li G" first="Genying" last="Li">Genying Li</name>
<affiliation><nlm:aff id="af1-ijms-20-00335">Institute of Crop Science, Shandong Academy of Agricultural Sciences, Key Laboratory of Wheat Biology & Genetic Improvement on North Yellow & Huai River Valley, Ministry of Agriculture, National Engineering Laboratory for Wheat & Maize, Jinan 250100, China;<email>zhangrongzhi1981@126.com</email>
(R.Z.);<email>zsjhappy@163.com</email>
(S.Z.);<email>lgy111@126.com</email>
(G.L.)</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Cao, Peijian" sort="Cao, Peijian" uniqKey="Cao P" first="Peijian" last="Cao">Peijian Cao</name>
<affiliation><nlm:aff id="af4-ijms-20-00335">China Tobacco Gene Research Center, Zhengzhou Tobacco Research Institute of CNTC, Zhengzhou 450001, China</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Zhao, Shancen" sort="Zhao, Shancen" uniqKey="Zhao S" first="Shancen" last="Zhao">Shancen Zhao</name>
<affiliation><nlm:aff id="af2-ijms-20-00335">BGI Institute of Applied Agriculture, BGI-Agro, Shenzhen 518083, China;<email>zhengfengya@genomics.cn</email>
</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="af4-ijms-20-00335">China Tobacco Gene Research Center, Zhengzhou Tobacco Research Institute of CNTC, Zhengzhou 450001, China</nlm:aff>
</affiliation>
</author>
</analytic>
<series><title level="j">International Journal of Molecular Sciences</title>
<idno type="eISSN">1422-0067</idno>
<imprint><date when="2019">2019</date>
</imprint>
</series>
</biblStruct>
</sourceDesc>
</fileDesc>
<profileDesc><textClass></textClass>
</profileDesc>
</teiHeader>
<front><div type="abstract" xml:lang="en"><p>Biotic stresses do damage to the growth and development of plants, and yield losses for some crops. Confronted with microbial infections, plants have evolved multiple defense mechanisms, which play important roles in the never-ending molecular arms race of plant–pathogen interactions. The complicated defense systems include pathogen-associated molecular patterns (PAMP) triggered immunity (PTI), effector triggered immunity (ETI), and the exosome-mediated cross-kingdom RNA interference (CKRI) system. Furthermore, plants have evolved a classical regulation system mediated by miRNAs to regulate these defense genes. Most of the genes/small RNAs or their regulators that involve in the defense pathways can have very rapid evolutionary rates in the longitudinal and horizontal co-evolution with pathogens. According to these internal defense mechanisms, some strategies such as molecular switch for the disease resistance genes, host-induced gene silencing (HIGS), and the new generation of RNA-based fungicides, have been developed to control multiple plant diseases. These broadly applicable new strategies by transgene or spraying ds/sRNA may lead to reduced application of pesticides and improved crop yield.</p>
</div>
</front>
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<pmc article-type="review-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Int J Mol Sci</journal-id>
<journal-id journal-id-type="iso-abbrev">Int J Mol Sci</journal-id>
<journal-id journal-id-type="publisher-id">ijms</journal-id>
<journal-title-group><journal-title>International Journal of Molecular Sciences</journal-title>
</journal-title-group>
<issn pub-type="epub">1422-0067</issn>
<publisher><publisher-name>MDPI</publisher-name>
</publisher>
</journal-meta>
<article-meta><article-id pub-id-type="pmid">30650550</article-id>
<article-id pub-id-type="pmc">6358896</article-id>
<article-id pub-id-type="doi">10.3390/ijms20020335</article-id>
<article-id pub-id-type="publisher-id">ijms-20-00335</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Review</subject>
</subj-group>
</article-categories>
<title-group><article-title>Evolution of Disease Defense Genes and Their Regulators in Plants</article-title>
</title-group>
<contrib-group><contrib contrib-type="author"><name><surname>Zhang</surname>
<given-names>Rongzhi</given-names>
</name>
<xref ref-type="aff" rid="af1-ijms-20-00335">1</xref>
<xref ref-type="author-notes" rid="fn1-ijms-20-00335">†</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Zheng</surname>
<given-names>Fengya</given-names>
</name>
<xref ref-type="aff" rid="af2-ijms-20-00335">2</xref>
<xref ref-type="author-notes" rid="fn1-ijms-20-00335">†</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Wei</surname>
<given-names>Shugen</given-names>
</name>
<xref ref-type="aff" rid="af3-ijms-20-00335">3</xref>
<xref ref-type="author-notes" rid="fn1-ijms-20-00335">†</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Zhang</surname>
<given-names>Shujuan</given-names>
</name>
<xref ref-type="aff" rid="af1-ijms-20-00335">1</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Li</surname>
<given-names>Genying</given-names>
</name>
<xref ref-type="aff" rid="af1-ijms-20-00335">1</xref>
</contrib>
<contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0001-9991-423X</contrib-id>
<name><surname>Cao</surname>
<given-names>Peijian</given-names>
</name>
<xref ref-type="aff" rid="af4-ijms-20-00335">4</xref>
<xref rid="c1-ijms-20-00335" ref-type="corresp">*</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Zhao</surname>
<given-names>Shancen</given-names>
</name>
<xref ref-type="aff" rid="af2-ijms-20-00335">2</xref>
<xref ref-type="aff" rid="af4-ijms-20-00335">4</xref>
<xref rid="c1-ijms-20-00335" ref-type="corresp">*</xref>
</contrib>
</contrib-group>
<aff id="af1-ijms-20-00335"><label>1</label>
Institute of Crop Science, Shandong Academy of Agricultural Sciences, Key Laboratory of Wheat Biology & Genetic Improvement on North Yellow & Huai River Valley, Ministry of Agriculture, National Engineering Laboratory for Wheat & Maize, Jinan 250100, China;<email>zhangrongzhi1981@126.com</email>
(R.Z.);<email>zsjhappy@163.com</email>
(S.Z.);<email>lgy111@126.com</email>
(G.L.)</aff>
<aff id="af2-ijms-20-00335"><label>2</label>
BGI Institute of Applied Agriculture, BGI-Agro, Shenzhen 518083, China;<email>zhengfengya@genomics.cn</email>
</aff>
<aff id="af3-ijms-20-00335"><label>3</label>
Guangxi Botanical Garden of Medicinal Plants, Nanning 530023, China;<email>weishugen2@163.com</email>
</aff>
<aff id="af4-ijms-20-00335"><label>4</label>
China Tobacco Gene Research Center, Zhengzhou Tobacco Research Institute of CNTC, Zhengzhou 450001, China</aff>
<author-notes><corresp id="c1-ijms-20-00335"><label>*</label>
Correspondence: <email>PeijianCao@163.com</email>
(P.C.); <email>zhaoshancen@genomics.cn</email>
(S.Z.)</corresp>
<fn id="fn1-ijms-20-00335"><label>†</label>
<p>These authors contributed equally to this work.</p>
</fn>
</author-notes>
<pub-date pub-type="epub"><day>15</day>
<month>1</month>
<year>2019</year>
</pub-date>
<pub-date pub-type="collection"><month>1</month>
<year>2019</year>
</pub-date>
<volume>20</volume>
<issue>2</issue>
<elocation-id>335</elocation-id>
<history><date date-type="received"><day>04</day>
<month>12</month>
<year>2018</year>
</date>
<date date-type="accepted"><day>10</day>
<month>1</month>
<year>2019</year>
</date>
</history>
<permissions><copyright-statement>© 2019 by the authors.</copyright-statement>
<copyright-year>2019</copyright-year>
<license license-type="open-access"><license-p>Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (<ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/">http://creativecommons.org/licenses/by/4.0/</ext-link>
).</license-p>
</license>
</permissions>
<abstract><p>Biotic stresses do damage to the growth and development of plants, and yield losses for some crops. Confronted with microbial infections, plants have evolved multiple defense mechanisms, which play important roles in the never-ending molecular arms race of plant–pathogen interactions. The complicated defense systems include pathogen-associated molecular patterns (PAMP) triggered immunity (PTI), effector triggered immunity (ETI), and the exosome-mediated cross-kingdom RNA interference (CKRI) system. Furthermore, plants have evolved a classical regulation system mediated by miRNAs to regulate these defense genes. Most of the genes/small RNAs or their regulators that involve in the defense pathways can have very rapid evolutionary rates in the longitudinal and horizontal co-evolution with pathogens. According to these internal defense mechanisms, some strategies such as molecular switch for the disease resistance genes, host-induced gene silencing (HIGS), and the new generation of RNA-based fungicides, have been developed to control multiple plant diseases. These broadly applicable new strategies by transgene or spraying ds/sRNA may lead to reduced application of pesticides and improved crop yield.</p>
</abstract>
<kwd-group><kwd>disease resistance gene</kwd>
<kwd>miRNA regulation</kwd>
<kwd>CKRI</kwd>
<kwd>ETI</kwd>
<kwd>PTI</kwd>
<kwd>HIGS</kwd>
<kwd>SIGS</kwd>
</kwd-group>
</article-meta>
</front>
<body><sec id="sec1-ijms-20-00335"><title>1. Introduction</title>
<p>The arms race of plants and host-pathogens seems never to stop, and sometimes the race is very intense. During the evolutionary process, plants have had to evolve multiple immunity mechanisms to survive danger signals in extracellular and intracellular milieus. Plants are able to enhance disease resistance and increase the food security, as well as to balance the resource allocation between growth and development. The prevalent defense mechanisms are categorized into three defense layers: the preliminary defense, pathogen-associated molecular pattern (PAMP) triggered immunity (PTI) [<xref rid="B1-ijms-20-00335" ref-type="bibr">1</xref>
], the secondary defense, effector-triggered immunity (ETI) [<xref rid="B2-ijms-20-00335" ref-type="bibr">2</xref>
], and the additional defense, the exosome-mediated cross-kingdom RNA interference (CKRI) system [<xref rid="B3-ijms-20-00335" ref-type="bibr">3</xref>
]. </p>
<p>It is well-known that PTI functions in basal defense. Using the cell surface-localized pattern recognition receptors (PRR), plants can detect the infection of invaders by recognizing the conserved microbe-associated or pathogen-associated molecular patterns (MAMPs or PAMPs) [<xref rid="B1-ijms-20-00335" ref-type="bibr">1</xref>
]. Plant PRRs are cell surface localized, and always are receptor-like kinases (RLKs) and receptor like proteins (RLPs). RLKs are comprised of extracellular domains, transmembrane domains, and intracellular kinase domains, which are required for transmitting the signals to the downstream defense responses, whereas RLPs are only comprised of the basic conformation without intracellular kinase domain. PTI with broad-spectrum defense is not sufficient to prevent most pathogens, and if plants have defect in PRRs, they often become more susceptible to microbes [<xref rid="B4-ijms-20-00335" ref-type="bibr">4</xref>
,<xref rid="B5-ijms-20-00335" ref-type="bibr">5</xref>
,<xref rid="B6-ijms-20-00335" ref-type="bibr">6</xref>
,<xref rid="B7-ijms-20-00335" ref-type="bibr">7</xref>
]. In turn, pathogens employ kinds of virulence effectors to overcome PTI and establish successful infection, termed effector-triggered immunity. Thus, ETI functions in the second defense of elicitor mediated defenses. </p>
<p>Most of the genes involved in ETI pathway contain intracellular nucleotide-binding site and leucine-rich repeat domains (NBS-LRRs or NLRs), which are typically cytoplasmic receptor proteins. <italic>NBS-LRR</italic>
genes can detect or recognize the polymorphic, strain-specific pathogen-secreted virulence effectors, and then transfer the signals to the downstream of defense genes. Thus, ETI-pathways belong to the species-specific disease resistance, and rapidly co-evolve with their pathogens. Plant species in eudicots and dicots have lots of <italic>NB-LRR</italic>
genes. According to the N-terminal features and functions, the NB-LRR proteins in plants can be termed into two classes with the terminal Toll/interleukin-1receptor (TIR) or coiled-coil (CC)/resistance to powdery mildew8 (RPW8) domains [<xref rid="B8-ijms-20-00335" ref-type="bibr">8</xref>
,<xref rid="B9-ijms-20-00335" ref-type="bibr">9</xref>
,<xref rid="B10-ijms-20-00335" ref-type="bibr">10</xref>
]. The TIR, CC or RPW8 domains are crucial in signaling transmit in cellular targets for effector action or downstream signaling components [<xref rid="B11-ijms-20-00335" ref-type="bibr">11</xref>
]. Although the <italic>NB-LRR</italic>
genes were demonstrated as the ancient and conserved genes in plants, their comparative genomic analyses have shown great structural diversity. For example, the CC domains are prevalent in eudicots and monocots, while the TIR domains are nearly absent in monocots [<xref rid="B12-ijms-20-00335" ref-type="bibr">12</xref>
]. Cross-kingdom RNA interference (CKRI) functions in the third layer, which protects plants by extracellular vesicles transport small RNAs or microRNAs (miRNAs) to microbial pathogens and then silence the virulence genes [<xref rid="B3-ijms-20-00335" ref-type="bibr">3</xref>
].</p>
<p>As one kind of typically small non-coding RNAs, miRNAs function in post-transcriptional gene regulation. Small miRNAs play big roles in a variety of biological processes, such as development, hormone responses and stress adaptations [<xref rid="B13-ijms-20-00335" ref-type="bibr">13</xref>
,<xref rid="B14-ijms-20-00335" ref-type="bibr">14</xref>
,<xref rid="B15-ijms-20-00335" ref-type="bibr">15</xref>
,<xref rid="B16-ijms-20-00335" ref-type="bibr">16</xref>
]. In PTI and ETI pathways, microRNAs as the classical regulators in post-transcript or translation level regulate defense/defense-associated genes [<xref rid="B17-ijms-20-00335" ref-type="bibr">17</xref>
,<xref rid="B18-ijms-20-00335" ref-type="bibr">18</xref>
], which can balance the benefits and costs of their targets. Plants employ miRNAs as shields against the pathogen attacks. MiRNAs respond to virus, bacteria and fungi by negatively regulating of mRNAs, which mainly function in both PTI and ETI. Until now, totally 153 disease resistance genes from PRGdb database [<xref rid="B19-ijms-20-00335" ref-type="bibr">19</xref>
], which involved in the plant immunity to biotic stresses, were validated by experiments in wet labs. Of them, 62.09% (95 from 153) genes, 17.65% (27 from 153) genes, 20.26% (31 from 153) genes were classified as NBS-LRR families, RLP/RLK, and other kinds of genes, respectively (<xref ref-type="fig" rid="ijms-20-00335-f001">Figure 1</xref>
). </p>
<p>In regard to defense genes, studies have shown a number of genes/small RNAs linked to anti-pathogen immunity. Here, we mainly summarize the current knowledge of the defense genes and their evolution paths regulated by miRNAs in plants, and then discuss their potential applications in crop improvements in the last section. </p>
</sec>
<sec id="sec2-ijms-20-00335"><title>2. Three Layers of Defense Mechanisms to Biotic Stresses in Plants</title>
<sec id="sec2dot1-ijms-20-00335"><title>2.1. The First Layer of Defense: Defense Genes in PTI</title>
<p>As one of the most important sensory protein groups, RLKs and RLPs in plants play crucial roles both in cell–cell and the plant–environment communications such as plant–pathogen interaction. In addition, RLKs and RLPs play fundamental roles in plant growth and development. Plants deploy a wide assay of RLKs and RLPs as the first layer of inducible defense to detect microbe- and host- derived molecular patterns (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
A, the first layer) [<xref rid="B63-ijms-20-00335" ref-type="bibr">63</xref>
]. Numbers of RLKs/RLPs have been cloned in plants [<xref rid="B64-ijms-20-00335" ref-type="bibr">64</xref>
]. The best classical example is FLAGELLIN-SENSITIVE2 (FLS2), belonging to RLK family, which have been verified to response to Flagellin fragment flg22 of bacteria in <italic>Arabidopsis</italic>
[<xref rid="B65-ijms-20-00335" ref-type="bibr">65</xref>
], grapevine [<xref rid="B66-ijms-20-00335" ref-type="bibr">66</xref>
], tobacco [<xref rid="B67-ijms-20-00335" ref-type="bibr">67</xref>
], rice [<xref rid="B68-ijms-20-00335" ref-type="bibr">68</xref>
] and tomato [<xref rid="B69-ijms-20-00335" ref-type="bibr">69</xref>
]. As a “molecular glue”, flg22 induces the activity of the heterodimerization complex FLS2-BAK1 (BRI1-ASSOCIATED RECEPTOR KINASE). In different plant species, FLS2 receptors display different affinities for the conserved part of flagellin from different bacteria, which possibly reflect the coevolution with specific-pathogens [<xref rid="B66-ijms-20-00335" ref-type="bibr">66</xref>
]. Except FLS2, EF-TU RECEPTOR (EFR), PEP 1 RECEPTOR (PEPR1), PEPR2, RLP23, RLP30 [<xref rid="B70-ijms-20-00335" ref-type="bibr">70</xref>
], the endogenous AtPep1 [<xref rid="B71-ijms-20-00335" ref-type="bibr">71</xref>
], NLPs [<xref rid="B72-ijms-20-00335" ref-type="bibr">72</xref>
], and SCFE1 [<xref rid="B73-ijms-20-00335" ref-type="bibr">73</xref>
], can also recognize bacterial EF-Tu, respectively. All of them are associated with the regulatory BAK1 that acts as a co-receptor for flg22/EF-Tu/AtPep1/nlp30/SCFE1 of pathogens and are crucial for signaling activation [<xref rid="B74-ijms-20-00335" ref-type="bibr">74</xref>
].</p>
<p>Long chitin oligomers as bivalent ligands, lead to the homodimerization of CHITIN ELICITOR RECEPTOR KINASE 1 (AtCERK1) and generate an active receptor complex in <italic>Arabidopsis</italic>
, which directly trigger chitin-induced immune signaling [<xref rid="B75-ijms-20-00335" ref-type="bibr">75</xref>
]. The chitin perception system in rice is significantly different from the one in <italic>Arabidopsis</italic>
. OsCERK1 dimmer does not bind chitin since the single LysM domain, while the dimer elicitor-binding LysM-RLP (OsCEBiP) can bind the chitin by ligand. The OsCERK1-chitin-OsCEBiP then forms a sandwich-type receptor dimerization for chitin oligomers [<xref rid="B76-ijms-20-00335" ref-type="bibr">76</xref>
].</p>
<p>There are a number of RLKs/RLPs involved in plant immunity, which have been well summarized by Tang et al [<xref rid="B63-ijms-20-00335" ref-type="bibr">63</xref>
]. After plant sensing of pathogen/microbe-associated molecular patterns, these pattern recognition receptors instantly trigger a number of downstream responses, such as the activation of mitogen-activated protein kinases (MAPKs) (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
A, the first layer), which is one of the earliest signaling events [<xref rid="B77-ijms-20-00335" ref-type="bibr">77</xref>
]. By phosphorylation to transmit response signals, MAPKKK actives MKK, and then MKK actives MPK [<xref rid="B78-ijms-20-00335" ref-type="bibr">78</xref>
]. MAPK cascades is involved in multiple signaling defense responses, including the biosynthesis/signaling of plant stress/defense hormones, reactive oxygen species generation, stomatal closure, defense gene activation, phytoalexin biosynthesis, cell wall strengthening, and hypersensitive response (HR) cell death (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
A, the first layer) [<xref rid="B77-ijms-20-00335" ref-type="bibr">77</xref>
]. The activation of MAPK cascades is essential for plant immunity. </p>
<p>In addition, some transcription factors were found to regulate the defense-related genes that involved in signal transduction in rice. For example, a <italic>bZIP</italic>
gene <italic>OsBBI1</italic>
in rice, is a major transcription factor to regulate the resistance spectrum for diverse groups of <italic>M. oryzae</italic>
by altering the first level of innate immunity in host plants [<xref rid="B79-ijms-20-00335" ref-type="bibr">79</xref>
]. WRKY13 as another major regulatory factor was identified to transfer signals from WRKY45 to downstream WRKY42 as functioning WRKY- type transcription factors (TFs) [<xref rid="B80-ijms-20-00335" ref-type="bibr">80</xref>
]. Following the SA-pathway-dependent disease response mechanism, WRKY13 shows correlation of the defense to <italic>M. oryzae</italic>
and <italic>Xoo</italic>
[<xref rid="B81-ijms-20-00335" ref-type="bibr">81</xref>
]. By activation of NPR1 protein, the SA pathway plays a crucial role in the systemic acquired resistance response mechanism (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
A, the first layer) [<xref rid="B82-ijms-20-00335" ref-type="bibr">82</xref>
]. As a result, kinds of genes comprised of cellulase surface disease resistance genes and intracellular transcript factors could function in the complex PTI.</p>
</sec>
<sec id="sec2dot2-ijms-20-00335"><title>2.2. The Second Layer of Defense: The Defense Genes in ETI</title>
<p>In ETI pathway, plants have developed NBS-LRR proteins to recognize effectors and trigger the ETI response [<xref rid="B2-ijms-20-00335" ref-type="bibr">2</xref>
], which can cause programmed cell death together with the downstream of <italic>WRKY</italic>
and lead to hypersensitive response (HR) (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
A, the second layer) [<xref rid="B97-ijms-20-00335" ref-type="bibr">97</xref>
]. <italic>NBS-LRRs</italic>
as an interesting class of disease resistance genes own a larger member in plants. In <xref rid="ijms-20-00335-t001" ref-type="table">Table 1</xref>
, about 1.19–3.48% of total coding genes were defined as <italic>NBS-LRR</italic>
genes. Although <italic>NB-LRR</italic>
genes are abundant in plants, only 93 genes are validated to play important roles in the innate immunity of plants up to now. Of the validated <italic>NBS-LRR</italic>
genes, 65.59% (61 from 93) genes contain the CC domains, while only 19.35% (18 from 93) genes contain the TIR domains, and the others contain only one domain of either NBS, LRR, TIR, CC, or RPW8 (<xref ref-type="fig" rid="ijms-20-00335-f001">Figure 1</xref>
). The verified disease resistance genes with CNL or TNL domains are listed in <xref rid="ijms-20-00335-t002" ref-type="table">Table 2</xref>
. For example, seven CNLs and seven TNLs in <italic>Arabidopsis thaliana</italic>
, eleven CNLs in <italic>Oryza sativa</italic>
, five CNLs and one TNL in <italic>Solanum lycopersicium</italic>
, seven CNLs in <italic>Triticum aestivum</italic>
, three CNLs in <italic>Hodeum vulgare</italic>
had been exemplified by experiments. These defense genes in plants can confer the resistance to fungi, oomycetes, bacteria, viruses, nematodes, and insects. </p>
<p>One type of plant disease can be prevented by several genes (<xref rid="ijms-20-00335-t002" ref-type="table">Table 2</xref>
). For example, the bacterial blight in <italic>Arabidopsis</italic>
caused by <italic>Pseudomonas syringae</italic>
/<italic>Xanthomonas oryzae</italic>
, can be defended by RPM1 (CNL) [<xref rid="B99-ijms-20-00335" ref-type="bibr">99</xref>
], Rps2 (CNL) [<xref rid="B100-ijms-20-00335" ref-type="bibr">100</xref>
], RPS5 (CNL) [<xref rid="B101-ijms-20-00335" ref-type="bibr">101</xref>
], SSI4 (TNL) [<xref rid="B102-ijms-20-00335" ref-type="bibr">102</xref>
], and Rps4 (TNL) genes [<xref rid="B103-ijms-20-00335" ref-type="bibr">103</xref>
]. The downy mildew of cucurbits that caused by <italic>Pseudoperonospora cubensis</italic>
(Oomycetes) in <italic>Arabidopsis</italic>
, can be resisted by RPP13/RPP8 (CNL/CNL) [<xref rid="B104-ijms-20-00335" ref-type="bibr">104</xref>
], RPP1/RPP4 (TNL/TNL) [<xref rid="B105-ijms-20-00335" ref-type="bibr">105</xref>
,<xref rid="B106-ijms-20-00335" ref-type="bibr">106</xref>
], and RPP5 (TNL) [<xref rid="B107-ijms-20-00335" ref-type="bibr">107</xref>
,<xref rid="B108-ijms-20-00335" ref-type="bibr">108</xref>
]. In rice, the famous rice blast disease caused by <italic>Magnaporthe grisea</italic>
or <italic>Magnaporthe oryzae</italic>
, can be defended by 17 CNL type of disease resistance genes including Pi-ta/PIB [<xref rid="B109-ijms-20-00335" ref-type="bibr">109</xref>
], RGA5 [<xref rid="B110-ijms-20-00335" ref-type="bibr">110</xref>
], Pi36/Pi9/Pi2 [<xref rid="B111-ijms-20-00335" ref-type="bibr">111</xref>
,<xref rid="B112-ijms-20-00335" ref-type="bibr">112</xref>
,<xref rid="B113-ijms-20-00335" ref-type="bibr">113</xref>
], Piz-t/Pikm1-TS/Pikm2-TS/Pid3/Pi5-1/Pi5-2/Pit/Pikp-2 [<xref rid="B113-ijms-20-00335" ref-type="bibr">113</xref>
,<xref rid="B114-ijms-20-00335" ref-type="bibr">114</xref>
,<xref rid="B115-ijms-20-00335" ref-type="bibr">115</xref>
,<xref rid="B116-ijms-20-00335" ref-type="bibr">116</xref>
,<xref rid="B117-ijms-20-00335" ref-type="bibr">117</xref>
], Pia [<xref rid="B118-ijms-20-00335" ref-type="bibr">118</xref>
], Pi37 [<xref rid="B119-ijms-20-00335" ref-type="bibr">119</xref>
] and Rpr1 [<xref rid="B120-ijms-20-00335" ref-type="bibr">120</xref>
]. In barley, the powdery mildew caused by <italic>Blumeria graminis</italic>
, can be resistant by CNL type of genes including MLA10 [<xref rid="B121-ijms-20-00335" ref-type="bibr">121</xref>
], MLA1 [<xref rid="B122-ijms-20-00335" ref-type="bibr">122</xref>
], and MLA13 [<xref rid="B123-ijms-20-00335" ref-type="bibr">123</xref>
]. In <italic>Linum usitatissimum</italic>
, flax rust caused by <italic>Melampsora lini</italic>
(Fungal), can be resistant by TNL type of genes including P2 [<xref rid="B124-ijms-20-00335" ref-type="bibr">124</xref>
], L6 [<xref rid="B125-ijms-20-00335" ref-type="bibr">125</xref>
], M [<xref rid="B126-ijms-20-00335" ref-type="bibr">126</xref>
], L [<xref rid="B127-ijms-20-00335" ref-type="bibr">127</xref>
], L1-L11 [<xref rid="B128-ijms-20-00335" ref-type="bibr">128</xref>
,<xref rid="B129-ijms-20-00335" ref-type="bibr">129</xref>
], P [<xref rid="B129-ijms-20-00335" ref-type="bibr">129</xref>
,<xref rid="B130-ijms-20-00335" ref-type="bibr">130</xref>
], and P1 [<xref rid="B124-ijms-20-00335" ref-type="bibr">124</xref>
]. One disease resistance gene can also confer plants resistant to several plant diseases (<xref rid="ijms-20-00335-t003" ref-type="table">Table 3</xref>
). For example, XA1 (CNL) [<xref rid="B131-ijms-20-00335" ref-type="bibr">131</xref>
] in rice, can defense to bacterial blight caused by bacterium of <italic>Pseudomonas syringae</italic>
and <italic>Xanthomonas oryzae</italic>
. Rx2 in <italic>Solanum acaule</italic>
, can defense to potato virus X (Virus) and <italic>Heterodera schachtii</italic>
(Nematode) [<xref rid="B132-ijms-20-00335" ref-type="bibr">132</xref>
].</p>
<p>The disease resistance genes were abundant in the wild resource. In <italic>Triticeae</italic>
for example, the defense genes <italic>Sr31</italic>
and <italic>Sr50</italic>
[<xref rid="B133-ijms-20-00335" ref-type="bibr">133</xref>
] from cereal rye (<italic>Secale cereale</italic>
), can confer the resistance to stem rust disease caused by <italic>Puccinia graminis f. sp. tritici</italic>
(<italic>Pgt</italic>
). <italic>Sr35</italic>
gene from <italic>Triticum monococcum</italic>
confers the resistance to Ug99 Stem Rust Race Group [<xref rid="B134-ijms-20-00335" ref-type="bibr">134</xref>
]. In addition, some non-<italic>NBS-LRR</italic>
genes can also provide the defense to pathogens. For example, Stb6 in wheat can directly interacted with the effector AvrStb6 that produced by wheat pathogen <italic>Zymoseptoria</italic>
<italic>tritici</italic>
[<xref rid="B135-ijms-20-00335" ref-type="bibr">135</xref>
]. The X10 gene, which has four potential transmembrane helices in rice, can be induced by transcription activator–like (TAL) effector AvrXa10. The gene can confer disease resistance to rice bacterial blight by inducing programmed cell death in rice [<xref rid="B136-ijms-20-00335" ref-type="bibr">136</xref>
,<xref rid="B137-ijms-20-00335" ref-type="bibr">137</xref>
].</p>
<p>By introgression or transgene strategy, these defense genes confer the disease resistance in plants. For example, by overexpressing Pm3a/c/d/f/g in wheat, all tested transgenic lines showed the significantly more resistance than their respective non-transformed sister lines in field experiments [<xref rid="B138-ijms-20-00335" ref-type="bibr">138</xref>
]. The T0 and T1 transgenic lines with the <italic>Sr50</italic>
gene were resistant to <italic>Puccinia graminis f. sp. tritici</italic>
(<italic>Pgt</italic>
), while lines without the transgene were susceptible [<xref rid="B133-ijms-20-00335" ref-type="bibr">133</xref>
]. </p>
</sec>
<sec id="sec2dot3-ijms-20-00335"><title>2.3. The Third Layer of Defense: Cross-Kingdom/Organism RNA Interference</title>
<p>It had been demonstrated that plasmodesmata sRNAs can presumably move from cell to cell, and they systemically travel through vasculature [<xref rid="B139-ijms-20-00335" ref-type="bibr">139</xref>
]. Remarkably, sRNAs also move and induce their target gene silencing between interacted organisms and hosts. The phenomenon was defined as cross-kingdom/organism RNA interference (CKRI) [<xref rid="B20-ijms-20-00335" ref-type="bibr">20</xref>
,<xref rid="B93-ijms-20-00335" ref-type="bibr">93</xref>
,<xref rid="B140-ijms-20-00335" ref-type="bibr">140</xref>
,<xref rid="B141-ijms-20-00335" ref-type="bibr">141</xref>
,<xref rid="B142-ijms-20-00335" ref-type="bibr">142</xref>
]. Pathogens can deliver sRNAs into plants. It was recently discovered as a novel class of pathogen effectors (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
A, the third layer). <italic>Botrytis cinerea</italic>
can deliver small RNAs (Bc-sRNAs) to plant cells to silence host immunity genes [<xref rid="B140-ijms-20-00335" ref-type="bibr">140</xref>
]. Such small RNA effectors in <italic>B. cinerea</italic>
are mostly produced by Dicer-like protein 1/2 (Bc-DCL1/2). In reverse, over-expressing sRNAs that target <italic>Bc-DCL1</italic>
and <italic>Bc-DCL2</italic>
in tomato and <italic>Arabidopsis</italic>
, would silence <italic>Bc-DCL</italic>
genes and inhibit fungal growth and pathogenicity. It exemplified bidirectional CKRI and sRNA trafficking between plants and fungi [<xref rid="B93-ijms-20-00335" ref-type="bibr">93</xref>
]. The easy traveling phenomenon suggests naturally occurring small RNAs might exchange each other across cross-kingdom/organism.</p>
<p>Conversely, hosts also can transfer naturally occurring small RNAs into pests or pathogens to attenuate their virulence (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
A, the third layer). Recently, two reports have demonstrated that naturally occurring plant small RNAs might be delivered into pathogens to silence their target genes. In response to the infection of <italic>Verticillium dahliae</italic>
, cotton plants increase the dose of miR159 and miR166 in expression level and then export both to the fungal hyphae for specific silencing. Two genes encoding an <italic>isotrichodermin C-15 hydroxylase</italic>
and a <italic>Ca</italic>
<sup>2+</sup>
<italic>-dependent cysteine protease</italic>
, were targeted by miR159 and miR166, respectively. Both of the target genes are essential for fungal virulence [<xref rid="B20-ijms-20-00335" ref-type="bibr">20</xref>
]. Another example is that host <italic>Arabidopsis</italic>
cells by secreting exosome-like extracellular vesicles can also transfer small RNAs into fungal pathogen <italic>Botrytis cinerea</italic>
. At the infection sites, these sRNA-containing vesicles accumulate and then are taken up by the fungal cells. Delivered host small RNAs induce the silence of fungal genes that is critical for pathogenicity. TAS1c-siR483 target two genes <italic>BC1G_10728</italic>
and <italic>BC1G_10508</italic>
from <italic>B. cinerea</italic>
, and TAS2-siR453 targets <italic>BC1T_08464</italic>
. All of the three genes involving in vesicle trafficking pathways are critical for pathogenicity [<xref rid="B3-ijms-20-00335" ref-type="bibr">3</xref>
]. Of them, <italic>BC1G_10728</italic>
encodes a vacuolar protein sorting 51 and plays a crucial role in <italic>Candida albicans</italic>
virulence [<xref rid="B21-ijms-20-00335" ref-type="bibr">21</xref>
]. Thus, <italic>Arabidopsis</italic>
has adapted exosome-mediated CKRI mechanism as part of its immune responses during the evolutionary arms race with the pathogens [<xref rid="B3-ijms-20-00335" ref-type="bibr">3</xref>
]. </p>
<p>Based on the above description, since only two miRNAs and two small RNAs in plants were identified to function in CKRI, data are inefficient to deduce their evolution among species. Thus, in the next section, we only discussed the evolution of disease resistance genes and their regulator miRNAs in PTI and ETI.</p>
</sec>
</sec>
<sec id="sec3-ijms-20-00335"><title>3. The Regulation of Disease Resistance Genes by Small RNAs</title>
<sec id="sec3dot1-ijms-20-00335"><title>3.1. The First Layer of Defense Regulation: miRNAs Involved in the PTI Pathway</title>
<p>During pathogen infection, plant small RNAs play key roles in gene regulation level. According to the targets of miRNAs that how to respond to the pathogen infection, miRNAs were divided into active and repressed regulation in basal resistance (<xref ref-type="fig" rid="ijms-20-00335-f001">Figure 1</xref>
A, <xref rid="ijms-20-00335-t003" ref-type="table">Table 3</xref>
). In the positive regulation, overexpression of miRNAs conferred the resistance to defense diseases in plants. For example, miR393 in <italic>Arabidopsis</italic>
, was discovered to contribute to the antibacterial resistance by negatively targeting the transcripts of the <italic>F-box auxin receptors TIR1</italic>
[<xref rid="B22-ijms-20-00335" ref-type="bibr">22</xref>
]. Repressing auxin signaling through miR393 overexpression increases bacterial resistance; conversely, augmenting auxin signaling through over-expressing a <italic>TIR</italic>
enhances susceptibility to virulent <italic>Pto DC3000</italic>
. miR444/<italic>OsMADS</italic>
directly monitors <italic>OsRDR1</italic>
transcription, and involves in the rice antiviral response [<xref rid="B23-ijms-20-00335" ref-type="bibr">23</xref>
]. Overexpression of miR444 enhanced rice resistance against rice stripe virus (<italic>RSV</italic>
) infection by diminishes the repressive roles of <italic>OsMADS23</italic>
, <italic>OsMADS27a,</italic>
and <italic>OsMADS57</italic>
and concomitant by the up-regulation of <italic>OsRDR1</italic>
expression. Thus, miR444 can indirectly activate the <italic>OsRDR1</italic>
-dependent antiviral RNA-silencing pathway. Over-expression of osa-miR171b conferred less susceptibility to rice stripe virus infection by regulating the target <italic>OsSCL6</italic>
. <italic>OsSCL6-IIa/b/c</italic>
was down-regulated or up-regulated in plants, where osa-miR171b was over-expressed or interfered [<xref rid="B24-ijms-20-00335" ref-type="bibr">24</xref>
].</p>
<p>In the negative regulation, overexpression their target genes could confer the resistance to pathogens in plants. miR169 suppresses the expression of <italic>NFYA</italic>
in immunity against the infection of bacterial wilt <italic>Ralstonia solanacearum</italic>
[<xref rid="B25-ijms-20-00335" ref-type="bibr">25</xref>
] and the blast fungus <italic>Magnaporthe oryzae in Arabidopsis</italic>
and rice, respectively [<xref rid="B26-ijms-20-00335" ref-type="bibr">26</xref>
]. The transgenic lines of over-expressing miR169a, became hyper-susceptible to pathogens. MiR156 and miR395 regulate apple resistance to Leaf Spot Disease [<xref rid="B27-ijms-20-00335" ref-type="bibr">27</xref>
]. In apple, Md-miR156ab and Md-miR395 suppress <italic>MdWRKYN1</italic>
and <italic>MdWRKY26</italic>
expression, which decreases the expression of some pathogenesis-related genes, and results in susceptibility to <italic>Alternaria alternaria f. sp. mali</italic>
. In <italic>Arabidopsis</italic>
, miR396/<italic>GRF</italic>
module mediates innate immunity against <italic>P. cucumerina</italic>
infection without growth costs. Reduced activity of miR396 (MIM396 plants) was found to improve broad resistance to necrotrophic and hemibiotrophic fungal pathogens [<xref rid="B28-ijms-20-00335" ref-type="bibr">28</xref>
]. MiR319/<italic>TCP</italic>
module involves in the rice blast disease. Increasing expression level of rice miR319 or decreasing expression level of its target <italic>TCP21</italic>
, <italic>LIPOXYGENASE2</italic>
(<italic>LOX2</italic>
) and <italic>LOX5</italic>
can facilitate rice ragged stunt virus (<italic>RRSV</italic>
) infection [<xref rid="B29-ijms-20-00335" ref-type="bibr">29</xref>
], which caused the decreased endogenous jasmonic acid (JA) [<xref rid="B30-ijms-20-00335" ref-type="bibr">30</xref>
]. Inhibiting ath-miR773 activity accompanied with up-regulation of its target gene <italic>METHYLTRANSFERASE 2</italic>
increased resistance to hemibiotrophic (<italic>Fusarium oxysporum</italic>
, <italic>Colletototrichum higginianum</italic>
) and necrotrophic (<italic>Plectosphaerrella cucumerina</italic>
) fungal pathogens in <italic>Arabidopsis</italic>
[<xref rid="B31-ijms-20-00335" ref-type="bibr">31</xref>
]. By regulating the transcription of <italic>GhMKK6</italic>
gene in cotton, ghr-miR5272a involved in the immune response. Over-expressing ghr-miR5272a increased sensitivity to <italic>Fusarium oxysporum</italic>
by decreasing the expression of <italic>GhMKK6</italic>
and the followed disease-resistance genes, which lead a similar phenotype to <italic>GhMKK6</italic>
-silenced cotton [<xref rid="B32-ijms-20-00335" ref-type="bibr">32</xref>
]. In addition, miRNAs could also be involved in the resistance to nematode invasion. For example, miR827 in <italic>Arabidopsis</italic>
down-regulated the expression of <italic>NITROGEN LIMITATION ADAPTATION</italic>
(<italic>NLA</italic>
) gene. It suppressed the basal defense pathway by enhancing susceptibility to the cyst nematode <italic>Heterodera schachtii</italic>
[<xref rid="B33-ijms-20-00335" ref-type="bibr">33</xref>
].</p>
<p>Except these miRNAs indirectly regulation the PTI pathway, a few of miRNAs were predicted to directly regulate the receptor-like genes. For example, when osa-miR159a.1 was repressed, the expression of <italic>OsLRR-RLK2</italic>
was induced, which is responded to <italic>Xanthomonas oryzae pv. Oryzae</italic>
in rice [<xref rid="B31-ijms-20-00335" ref-type="bibr">31</xref>
]. In future, some miRNAs regulation of pattern recognition receptors (PRR) genes may be validated by experiments.</p>
</sec>
<sec id="sec3dot2-ijms-20-00335"><title>3.2. The Second Layer of Defense Regulation: The Defense Signal Small RNAs in ETI</title>
<p>In addition to the basal defense, miRNAs are also involved in ETI pathway to directly and indirectly regulate the disease resistance genes (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
A & <xref rid="ijms-20-00335-t003" ref-type="table">Table 3</xref>
). MiR393*, the complementary strand of miR393 within the sRNA duplex, by targeting a protein trafficking gene <italic>Membrin 12</italic>
promote the secretion of antimicrobial PR proteins, which functions in ETI during infection of <italic>Pseudomonas syringae pv. Tomato</italic>
in <italic>Arabidopsis</italic>
[<xref rid="B34-ijms-20-00335" ref-type="bibr">34</xref>
]. The miR863-3p is induced by the bacterial pathogen <italic>Pseudomonas syringae</italic>
. During early infection, miR863-3p silences two negative regulators of plant defense, namely <italic>atypical receptor-like pseudokinase1</italic>
(<italic>ARLPK1</italic>
) and <italic>ARLPK2</italic>
, both of which trigger immunity through mRNA degradation. Later during infection, miR863-3p silences <italic>SERRATE</italic>
, and positively regulates defense. And <italic>SERRATE</italic>
is essential for miR863-3p accumulation by a negative feedback loop. Thus, miR863-3p targets both negative and positive regulators of immunity through two modes of action to fine-tune in the timing and amplitude of defense responses [<xref rid="B35-ijms-20-00335" ref-type="bibr">35</xref>
].</p>
<p>High expression of plant <italic>NBS-LRR</italic>
defense genes is often lethal to plant cells, which is associated with the fitness costs. Thus, plants develop several mechanisms to regulate the transcript level of <italic>NBS-LRR</italic>
genes. One of the key mechanism is the suppression of regulation network in microRNAs and <italic>NBS-LRRs</italic>
, which may play a crucial role in plant-microbe interactions by sRNA silencing mechanism [<xref rid="B18-ijms-20-00335" ref-type="bibr">18</xref>
]. <italic>NBS-LRR</italic>
genes confer defense against the pathogen infections in gene dosage dynamic expression level by multiple duplications and diversification, while miRNAs minimized the cost of gene copies by inhibiting their expression [<xref rid="B36-ijms-20-00335" ref-type="bibr">36</xref>
]. One miRNA can regulate dozens to hundreds of <italic>NBS-LRRs</italic>
by targeting the similar motif sites [<xref rid="B37-ijms-20-00335" ref-type="bibr">37</xref>
], which make it more economical to balance the benefits and costs of these copies in genome. Until now, a few of miRNAs had been validated to be involved in the regulation of <italic>NBS-LRR</italic>
genes. </p>
<p>The regulation between miRNAs and <italic>CC-NB-LRR</italic>
or <italic>TIR-NB-LRR</italic>
gene classes was mostly characterized in eudicots. In most of the post-transcriptional regulation networks, the miRNA can trigger the 21-nt phased siRNA generation in <italic>NB-LRR</italic>
transcripts, which were processed by RNA-dependent RNA polymerase 6 (RDR6) and DICER-LIKE 4 (DCL4) [<xref rid="B38-ijms-20-00335" ref-type="bibr">38</xref>
]. For example, in <italic>Brassica</italic>
miR1885 were validated to induce by Turnip Mosaic Virus (<italic>TuMV</italic>
) infection, which cleaved <italic>TIR-NB-LRR</italic>
class genes [<xref rid="B39-ijms-20-00335" ref-type="bibr">39</xref>
]. In Tobacco, by cleaving <italic>TIR-NB-LRR</italic>
immune receptors, both of nta-miR6020 and nta-miR6019 provide resistance to Tobacco mosaic virus (<italic>TMV</italic>
) [<xref rid="B40-ijms-20-00335" ref-type="bibr">40</xref>
,<xref rid="B41-ijms-20-00335" ref-type="bibr">41</xref>
]. In tomato, sl-miR5300 and sl-miR482f controlled NB domain-containing proteins in mRNA stability and translation level, which involved in plant immunity [<xref rid="B42-ijms-20-00335" ref-type="bibr">42</xref>
]. In <italic>Arabidopsis</italic>
, miR472 modulated the disease resistance genes mediated by RDR6 silencing pathway [<xref rid="B43-ijms-20-00335" ref-type="bibr">43</xref>
]. In <italic>Medicago</italic>
, miR2109, miR482/miR2118 and miR1507 were found to influence <italic>NB-LRR</italic>
gene family [<xref rid="B37-ijms-20-00335" ref-type="bibr">37</xref>
]. In legumes, miR482, miR1507, miR1510, and miR2109 suppressed <italic>NB-LRR</italic>
gene class with CC or TIR domains, which were proposed to function in the regulation of defense response or host specificity during rhizobium colonization [<xref rid="B38-ijms-20-00335" ref-type="bibr">38</xref>
,<xref rid="B44-ijms-20-00335" ref-type="bibr">44</xref>
]. In addition, miR482/miR2118, miR946, miR950, miR951, miR1311, miR1312, miR3697, miR3701, and miR3709 were also mediated to generate phased siRNAs by targeting <italic>NBS-LRR</italic>
gene class in Norway Spruce [<xref rid="B45-ijms-20-00335" ref-type="bibr">45</xref>
]. In monocots, miR2009 (also named miR9863 in miRBase) was first predicted in wheat to target the <italic>Mla</italic>
alleles [<xref rid="B46-ijms-20-00335" ref-type="bibr">46</xref>
]. In barley, the miR9863 family was confirmed to trigger response to the <italic>Mla</italic>
alleles [<xref rid="B47-ijms-20-00335" ref-type="bibr">47</xref>
].</p>
</sec>
</sec>
<sec id="sec4-ijms-20-00335"><title>4. The Evolution of Defense Genes</title>
<sec id="sec4dot1-ijms-20-00335"><title>4.1. The Evolution of Defense Gene in PTI</title>
<p>In land plants, RLKs expanded extensively and fulfilled these diverse roles including perceive growth hormones, environmental/danger signals derived from pathogens [<xref rid="B143-ijms-20-00335" ref-type="bibr">143</xref>
]. In <italic>Arabidopsis</italic>
, 44 RLK subgroups were defined, and leucine-rich repeat receptor-like kinases (LRR-RLK) belong to the largest receptor-like kinase family and are focused by researchers [<xref rid="B144-ijms-20-00335" ref-type="bibr">144</xref>
]. According to characters of unique basic gene structures and protein motif compositions, plant <italic>LRR-RLKs</italic>
constitute 19 subfamilies, most of which were derived from the common ancestors in land plants. The proportions of <italic>LRR-RLK</italic>
genes in Lycophytes and moss genome are 0.30% and 0.36%, respectively, while the proportions of <italic>LRR-RLK</italic>
genes in angiosperms are 0.67–1.39% [<xref rid="B145-ijms-20-00335" ref-type="bibr">145</xref>
], which indicated the special expansion of defense genes in angiosperm genomes. <italic>LRR-RLK</italic>
involved in the defense/resistance-related genes was less conserved than that involved in development. Defense-associated <italic>LRR-RLKs</italic>
undergone many duplication events, and most of them were massively lineage-specific expansion mainly by tandem duplication [<xref rid="B143-ijms-20-00335" ref-type="bibr">143</xref>
,<xref rid="B144-ijms-20-00335" ref-type="bibr">144</xref>
]. These discoveries provide important resources for future functional research for these critical signaling genes in PTI.</p>
</sec>
<sec id="sec4dot2-ijms-20-00335"><title>4.2. The Evolution of Defense Gene in ETI</title>
<p><italic>NBS-LRR</italic>
genes as a class of ancient and conserved genes have been detected in gymnosperms, angiosperm plants and animals to ensure immunity [<xref rid="B12-ijms-20-00335" ref-type="bibr">12</xref>
,<xref rid="B146-ijms-20-00335" ref-type="bibr">146</xref>
,<xref rid="B147-ijms-20-00335" ref-type="bibr">147</xref>
]. However, comparative genomic analyses have demonstrated that <italic>NBS-LRR</italic>
genes have a great structural diversity in plants and animals. For example, TIR domains were established in the ancestor plants conifers and mosses, and also in animals shared functionality regarding innate immunity [<xref rid="B148-ijms-20-00335" ref-type="bibr">148</xref>
,<xref rid="B149-ijms-20-00335" ref-type="bibr">149</xref>
,<xref rid="B150-ijms-20-00335" ref-type="bibr">150</xref>
]. TIR genes specially expanded in dicot genomes, but are absent or at least rare in monocot genomes [<xref rid="B8-ijms-20-00335" ref-type="bibr">8</xref>
,<xref rid="B147-ijms-20-00335" ref-type="bibr">147</xref>
,<xref rid="B151-ijms-20-00335" ref-type="bibr">151</xref>
,<xref rid="B152-ijms-20-00335" ref-type="bibr">152</xref>
,<xref rid="B153-ijms-20-00335" ref-type="bibr">153</xref>
]. For <italic>NBS-LRR</italic>
genes, tandem duplication in genome is the major expansion mechanism in plants. More than 60% of <italic>NBS-LRR</italic>
genes organized in a general pattern of clusters in plant genomes (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
B) [<xref rid="B98-ijms-20-00335" ref-type="bibr">98</xref>
]. During whole genome duplication, biased deletions happened in the duplicated paralogous blocks with <italic>NB-LRR</italic>
genes, and it could be possibly compensated by their local tandem duplication mechanism (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
B). </p>
<p>The miRNAs typically target highly duplicated <italic>NBS-LRRs</italic>
, and families of heterogeneous <italic>NBS-LRRs</italic>
were rarely targeted by miRNAs in <italic>Brassicaceae</italic>
and <italic>Poaceae</italic>
genomes [<xref rid="B18-ijms-20-00335" ref-type="bibr">18</xref>
]. miRNAs/<italic>NBS-LRR</italic>
-genes interactions drove functional diploidization of structurally retained <italic>NBS-LRR</italic>
genes duplicates by suppression regulation (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
B) [<xref rid="B98-ijms-20-00335" ref-type="bibr">98</xref>
]. Evolutionary shuffling events such as diploidization and tandem duplication, leaded to copy number variations and presence absence variations in the synteny collapse of <italic>NBS-LRR</italic>
genes [<xref rid="B154-ijms-20-00335" ref-type="bibr">154</xref>
,<xref rid="B155-ijms-20-00335" ref-type="bibr">155</xref>
,<xref rid="B156-ijms-20-00335" ref-type="bibr">156</xref>
,<xref rid="B157-ijms-20-00335" ref-type="bibr">157</xref>
]. In addition, the polymorphisms often exist in a population [<xref rid="B158-ijms-20-00335" ref-type="bibr">158</xref>
]. A contrasted conservation of <italic>NBS-LRR</italic>
genes was observed with only 23.8% for monocots and 6.6% for dicots. Thus, <italic>NBS-LRR</italic>
genes as one of the most plastic gene family in plants have less conservation such as synteny erosion or alternatively loss in plants compared with the other coding protein genes [<xref rid="B98-ijms-20-00335" ref-type="bibr">98</xref>
].</p>
</sec>
</sec>
<sec id="sec5-ijms-20-00335"><title>5. The Evolution of microRNAs in the Defense Pathway</title>
<sec id="sec5dot1-ijms-20-00335"><title>5.1. The Evolution of miRNAs in PTI</title>
<p>In the PTI pathway, most of miRNAs were very conserved and directly/indirectly involve multiple biological processes in the development and abiotic/biotic stresses. All of the MiR169, miR171, miR393, miR395, and miR396 were ancient miRNAs present in both dicots and monocots [<xref rid="B48-ijms-20-00335" ref-type="bibr">48</xref>
]. miR444 was specific in monocots [<xref rid="B49-ijms-20-00335" ref-type="bibr">49</xref>
], whereas miR773 and miR5272 were lineage-specific in <italic>Arabidopsis</italic>
and <italic>Medicago</italic>
. The miRNAs conserved in plants mostly regulate the important transcript factors. These transcript factors tend to involve multiple biological processes. Take miR169 and miR396 for example, miR169/<italic>NFYA</italic>
in <italic>Arabidopsis</italic>
indirectly affected lateral root initiation [<xref rid="B50-ijms-20-00335" ref-type="bibr">50</xref>
], nitrogen-starvation [<xref rid="B51-ijms-20-00335" ref-type="bibr">51</xref>
], drought stress [<xref rid="B52-ijms-20-00335" ref-type="bibr">52</xref>
], and biotic stress [<xref rid="B25-ijms-20-00335" ref-type="bibr">25</xref>
,<xref rid="B26-ijms-20-00335" ref-type="bibr">26</xref>
]. In <italic>Arabidopsis</italic>
roots, miR396/GRF regulates the switch between stem cells and transit-amplifying cells [<xref rid="B53-ijms-20-00335" ref-type="bibr">53</xref>
], which affects rice yield by shaping inflorescence architecture [<xref rid="B54-ijms-20-00335" ref-type="bibr">54</xref>
], and biotic stresses [<xref rid="B28-ijms-20-00335" ref-type="bibr">28</xref>
]. </p>
<p>Both of the miRNA/target regulation and their function are very conserved in plants. MiR169/<italic>NFYA</italic>
module influences the <italic>Ralstonia solanacearum</italic>
pathogenicity in <italic>Arabidopsis</italic>
[<xref rid="B25-ijms-20-00335" ref-type="bibr">25</xref>
] and the resistance to <italic>M. oryzae</italic>
strains in rice [<xref rid="B26-ijms-20-00335" ref-type="bibr">26</xref>
]. In addition, these conserved miRNAs’ targets were expanded except for their classical miRNA/target model. For example, miRNA156 regulates of the SQUAMOSA-PROMOTER BINDING PROTEIN-LIKE (SPL) family involve in the timing of vegetative and reproductive phase change, which is highly conserved among phylogenetically distinct plant species [<xref rid="B55-ijms-20-00335" ref-type="bibr">55</xref>
]. miR395 by targeting a high-affinity sulphate transporter and three ATP sulfurylases involved in the sulfate homeostasis, is also conserved in plants [<xref rid="B56-ijms-20-00335" ref-type="bibr">56</xref>
,<xref rid="B57-ijms-20-00335" ref-type="bibr">57</xref>
]. Differently, both miR156 and miR395 regulate apple resistance to leaf spot disease by targeting <italic>WRKY</italic>
. Thus, miRNAs involved in PTI pathway, are conserved in PTI defense pathway and in plant development such as miR393 <italic>vs TIR</italic>
in auxin signal pathway [<xref rid="B22-ijms-20-00335" ref-type="bibr">22</xref>
] and miR319 <italic>vs. TCL</italic>
in JA pathway [<xref rid="B29-ijms-20-00335" ref-type="bibr">29</xref>
]. Only few of miRNAs were reported to potentially regulating the <italic>RLK</italic>
/<italic>RLP</italic>
by osa-miR159a.1 [<xref rid="B58-ijms-20-00335" ref-type="bibr">58</xref>
], MiR5638 and miR1315 [<xref rid="B59-ijms-20-00335" ref-type="bibr">59</xref>
]. Genes involved in the PTI pathway were relatively conserved compared to these genes involved in ETI pathway. Thus, most of their regulator miRNAs were also conserved miRNAs or neofunctionalization of miRNAs in plants.</p>
</sec>
<sec id="sec5dot2-ijms-20-00335"><title>5.2. The Evolution of miRNAs in ETI</title>
<p>Although there are many miRNAs regulated <italic>NB-LRR</italic>
genes, the conservation level of miRNAs is lower than the development associated miRNAs or PTI-associated miRNAs. In the eudicots and monocots, there is no conserved miRNAs targeting the <italic>NB-LRR</italic>
genes. Lineage- or species-specific disease resistance-associated miRNAs were continually present and accompanies the continually varied pathogens. And some miRNAs with similar sequences had obvious functional diversity. miR482/miR2118 in eudicots mostly targeted <italic>NB-LRR</italic>
genes, however, it only initiated the generation of 21-nt phased siRNAs in rice, and most of the target transcripts were noncoding sequences and specifically expressed in the rice stamen and the maize premeiotic and meiotic anther [<xref rid="B60-ijms-20-00335" ref-type="bibr">60</xref>
,<xref rid="B61-ijms-20-00335" ref-type="bibr">61</xref>
,<xref rid="B62-ijms-20-00335" ref-type="bibr">62</xref>
]. It clearly concluded that miR2118 initiated the phased siRNA in male reproductive organs. Therefore, a functional switch occurred in miR482/miR2118 between eudicots and dicots. Their expression level also varies in the lineage-related species. Tae-miR3117 was predicted to target the numbers of <italic>NBS-LRRs</italic>
with higher expression in the tetraploid and hexaploid <italic>Triticum</italic>
seedlings, while it had lower expression levels in <italic>Aegilops tauschii</italic>
(not published data). And in rice, maize, and sorghum, miR3117 also displayed lower expression levels.</p>
<p>Diverse miRNAs, as negative transcriptional regulators, inhibit <italic>NBS-LRRs</italic>
in plants. The highly duplicated <italic>NBS-LRRs</italic>
were typically targeted by miRNAs (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
B), while families of heterogeneous <italic>NBS-LRR</italic>
genes were rarely regulated by miRNAs such as in <italic>Poaceae</italic>
and <italic>Brassicaceae</italic>
genomes. For example, some miRNAs also have a high duplication rate such as miR482/miR2118 in tandem duplication in genomes [<xref rid="B60-ijms-20-00335" ref-type="bibr">60</xref>
,<xref rid="B61-ijms-20-00335" ref-type="bibr">61</xref>
,<xref rid="B62-ijms-20-00335" ref-type="bibr">62</xref>
], which may enhance the expression dosage. </p>
<p>Newly emerged miRNAs were periodically derived from duplicated/redundant <italic>NBS-LRRs</italic>
from different gene families. And most of these new birth miRNAs target these <italic>NBS-LRR</italic>
gene regions of conserved, encoded protein motif, which follow in the convergent evolution model (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
B). The miRNAs may drive the rapid diploidization of these <italic>NBS-LRR</italic>
genes in polyploid plants. These <italic>NBS-LRR</italic>
associated miRNAs had a rapid diversity. The nucleotide diversity of the target site region in the wobble position of the codons drives the diversification of miRNAs. These characters of high duplication rate and rapid diversity were similar to their target genes. The co-evolutionary model between <italic>NBS-LRRs</italic>
and miRNAs in plants makes the plants balance the costs and benefits of disease resistance [<xref rid="B18-ijms-20-00335" ref-type="bibr">18</xref>
].</p>
</sec>
</sec>
<sec id="sec6-ijms-20-00335"><title>6. The Strategies of Defense Pathogens in plants</title>
<sec id="sec6dot1-ijms-20-00335"><title>6.1. The First Strategy: Utilize the Disease Resistance Genes by a Molecular Switch </title>
<p>Up to now, a number of genes were exemplified to be involved in plant immunity defense. By over-expressing such defense genes can dramatically enhance disease resistance in plants, while is often associated with significant penalties to fitness and make the resulting products undesirable. Thus, it is difficult in agricultural applications. Recently, it has been developed a strategy to utilize these disease defense genes from the angle of plant genes or their regulators [<xref rid="B83-ijms-20-00335" ref-type="bibr">83</xref>
]. The strategy is to introduce immunity-inducible promoter and other two pathogen-responsive upstream open reading frames of the <italic>TBF1</italic>
gene. It is called uORFsTBF1, which is a key immune regulator and its translation is transiently and rapidly induced upon pathogen challenge (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
C, uORF). It has been demonstrated that inclusion of the <italic>uORFsTBF1</italic>
-mediated translational control over the production of AtNPR1 in rice and an auto-activated immunity receptor snc1-1 in <italic>Arabidopsis</italic>
did not reduce the plant fitness in the laboratory or in the field [<xref rid="B83-ijms-20-00335" ref-type="bibr">83</xref>
]. This strategy using a molecular switch enables us to engineer more broad-spectrum disease resistance genes with minimal adverse effects on plant growth and development in the agriculture application. </p>
</sec>
<sec id="sec6dot2-ijms-20-00335"><title>6.2. The Second Strategy: Host-Induced Gene Silencing (HIGS)</title>
<p>Transgene-derived artificial sRNAs in plants can induce the target gene silencing in certain interacting insects [<xref rid="B84-ijms-20-00335" ref-type="bibr">84</xref>
,<xref rid="B85-ijms-20-00335" ref-type="bibr">85</xref>
], nematodes [<xref rid="B86-ijms-20-00335" ref-type="bibr">86</xref>
], fungi [<xref rid="B87-ijms-20-00335" ref-type="bibr">87</xref>
,<xref rid="B88-ijms-20-00335" ref-type="bibr">88</xref>
,<xref rid="B89-ijms-20-00335" ref-type="bibr">89</xref>
,<xref rid="B90-ijms-20-00335" ref-type="bibr">90</xref>
], oomycetes [<xref rid="B91-ijms-20-00335" ref-type="bibr">91</xref>
,<xref rid="B92-ijms-20-00335" ref-type="bibr">92</xref>
], and even plants–plants [<xref rid="B141-ijms-20-00335" ref-type="bibr">141</xref>
]. The phenomenon was called host-induced gene silencing (HIGS). The artificial sRNAs can travel from host plants to pathogens or pests and then function in trans (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
C, HIGS). It had been well used in many plants in the decades. By plant RNAi suppressing a bollworm <italic>P450 monooxygenase</italic>
gene of cotton impaired larval tolerance of gossypol [<xref rid="B85-ijms-20-00335" ref-type="bibr">85</xref>
]. In transgenic plants, by RNAi silencing of a conserved and essential root-knot nematode parasitism gene engineered broad root-knot resistance [<xref rid="B86-ijms-20-00335" ref-type="bibr">86</xref>
]. HIGS of nematode fitness and reproductive genes decreases fecundity of <italic>Heterodera glycines Ichinohe</italic>
. Double-stranded RNA complementary to <italic>cytochrome P450 lanosterol C14 alpha-demethylase-encoding</italic>
genes of <italic>Fusarium</italic>
in <italic>Arabidopsis</italic>
and barley contributes to strong resistance to <italic>Fusarium</italic>
species [<xref rid="B90-ijms-20-00335" ref-type="bibr">90</xref>
]. HIGS to the <italic>MAPKK</italic>
gene <italic>PsFUZ7</italic>
in wheat enhance stable resistance to wheat stripe rust [<xref rid="B159-ijms-20-00335" ref-type="bibr">159</xref>
]. HIGS of an important pathogenicity factor <italic>PsCPK1</italic>
in <italic>Puccinia striiformis f. sp. tritici</italic>
conferred resistance of wheat to stripe rust [<xref rid="B160-ijms-20-00335" ref-type="bibr">160</xref>
]. By transgene-mediated cross-kingdome RNAi mechanism, HIGS by transgene is a good and effect strategy to improve the crop disease resistance in a broad spectrum.</p>
</sec>
<sec id="sec6dot3-ijms-20-00335"><title>6.3. The Third Strategy: Spray-Induced Gene Silencing (SIGS) </title>
<p>The pathogens and pests are capable to take up the double RNAs or small RNAs from the plants or the environments [<xref rid="B93-ijms-20-00335" ref-type="bibr">93</xref>
]. Based on this and according to the mechanism of cross-kingdom/organism RNA interference, researchers have developed a strategy to control crop disease. It is spray-induced gene silencing (SIGS) that spraying dsRNAs and sRNAs on plant surfaces can target pathogen genes to repression pathogen virulence (<xref ref-type="fig" rid="ijms-20-00335-f002">Figure 2</xref>
C, SIGS). For modern crop protection strategies, it is a natural blueprint. Evidences suggest that nematodes [<xref rid="B94-ijms-20-00335" ref-type="bibr">94</xref>
], insects [<xref rid="B84-ijms-20-00335" ref-type="bibr">84</xref>
] and fungi [<xref rid="B95-ijms-20-00335" ref-type="bibr">95</xref>
] could uptake up the environmental dsRNA or sRNAs. Directly spraying the dsRNAs that target the fungal <italic>cytochrome P450 lanosterol C-14alpha-demethylases</italic>
of fungal gene can suppress fungal growth [<xref rid="B95-ijms-20-00335" ref-type="bibr">95</xref>
]. On barley leaves, spraying <italic>CYP51</italic>
-targeting dsRNA at a concentration range of 1–20 ng/mL, inhibited growth of <italic>Fusarium</italic>
species [<xref rid="B3-ijms-20-00335" ref-type="bibr">3</xref>
]. It has been demonstrated that spraying naked sRNAs and dsRNA on plants was successful to protect fruits and vegetables against pathogens. However, pesticide effect of the naked sRNAs and dsRNAs can only last 5–8 days. Mitter, et al. developed a method to load dsRNAs on designer, non-toxic, degradable, layered double hydroxide (LDH) clay nanosheets. This LDH made the dsRNA does not be wash off, and can be sustained released for 30 days [<xref rid="B96-ijms-20-00335" ref-type="bibr">96</xref>
]. This SIGS broadly application of new strategy may contribute to reduced use of chemical pesticides and lightening of selective pressure for resistant pathogens. The new-generation of RNA-based fungicides and pesticides are powerful, eco-friendly, which can be easily adapted to control multiple plant diseases simultaneously. </p>
</sec>
</sec>
<sec id="sec7-ijms-20-00335"><title>7. Conclusions</title>
<p>Plants deployed PTI, ETI, and CKRI innate immune systems to arm race with different pathogen stresses. Pathogens developed more advanced effectors to defeat plant defense immunity. A number of genes have been exemplified to play important role between the host-pathogen interactions in plants. These signaling genes will be helpful to improve plant disease resistance against various pathogens. The sustainable and broaden spectrum resistance genes and their regulators such as miRNAs will be applied in developing crop varieties by introducing the molecular switch. From the cross-kingdom angle, the HIGS can also be used to crop breeding by transgenic approach, which can also confer the broaden spectrum resistance to hosts. The SIGS can also make plants yield the broaden spectrum resistance by spraying the designed dsRNAs/sRNA. Further function studies in plants will dissect more and more defense genes and hopefully unravel the intricate defense regulation network. More and more molecular technologies will be invented and adapted to help develop the eco-friendly disease-resistance cultivars.</p>
</sec>
</body>
<back><ack><title>Acknowledgments</title>
<p>This work was mainly supported by the Young Elite Scientists Sponsorship Program By CAST (2016QNRC001), the National Natural Science Foundation of China (31501312, 31601301 from SAAS), and the Natural Science Foundation of Shandong Province (ZR2014CM006 from SAAS) for the design of the study. Collection of data was supported by the Agricultural Science and Technology Innovation Project of Shandong Academy of Agricultural Sciences (CXGC2016C09 from SAAS), the Youth Talent Program of Shandong Academy of Agricultural Sciences from SAAS. Data analysis, organization, and interpretation were supported by the Ministry of Agriculture of China (2018ZX08009-10B from SAAS), the Youth Foundation of Shandong Academy of Agricultural Sciences (2016YQN01 from SAAS), the Ministry of Science and Technology of China (2016YFD0100500 from SAAS) and Key R&D Programme of Shandong Province (2017GNC10113 from SAAS), the National Science and Technology Major Project of Breeding New Varieties of Genetically Modified Organisms (2018ZX0800910B from SAAS), and Projects of ENCODE of Tobacco Genome (110201601033(JY-07)).</p>
</ack>
<notes><title>Author Contributions</title>
<p>S.Z. and R.Z. conceived and designed the project. F.Z., R.Z., S.Z., S.W., and P.C. downloaded the data and analyzed the data. R.Z., F.Z., and G.L. prepared and drafted the manuscript. S.Z. and P.C. revised the manuscript. All the authors read and approved the final manuscript.</p>
</notes>
<notes notes-type="COI-statement"><title>Conflicts of Interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as potential conflicts of interest.</p>
</notes>
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<floats-group><fig id="ijms-20-00335-f001" orientation="portrait" position="float"><label>Figure 1</label>
<caption><p>Categories of the genes/regulators in the three defense layers in plants. The data was downloaded from PRGdb database and the recent publications [<xref rid="B20-ijms-20-00335" ref-type="bibr">20</xref>
,<xref rid="B21-ijms-20-00335" ref-type="bibr">21</xref>
,<xref rid="B22-ijms-20-00335" ref-type="bibr">22</xref>
,<xref rid="B23-ijms-20-00335" ref-type="bibr">23</xref>
,<xref rid="B24-ijms-20-00335" ref-type="bibr">24</xref>
,<xref rid="B25-ijms-20-00335" ref-type="bibr">25</xref>
,<xref rid="B26-ijms-20-00335" ref-type="bibr">26</xref>
,<xref rid="B27-ijms-20-00335" ref-type="bibr">27</xref>
,<xref rid="B28-ijms-20-00335" ref-type="bibr">28</xref>
,<xref rid="B29-ijms-20-00335" ref-type="bibr">29</xref>
,<xref rid="B30-ijms-20-00335" ref-type="bibr">30</xref>
,<xref rid="B31-ijms-20-00335" ref-type="bibr">31</xref>
,<xref rid="B32-ijms-20-00335" ref-type="bibr">32</xref>
,<xref rid="B33-ijms-20-00335" ref-type="bibr">33</xref>
,<xref rid="B34-ijms-20-00335" ref-type="bibr">34</xref>
,<xref rid="B35-ijms-20-00335" ref-type="bibr">35</xref>
,<xref rid="B36-ijms-20-00335" ref-type="bibr">36</xref>
,<xref rid="B37-ijms-20-00335" ref-type="bibr">37</xref>
,<xref rid="B38-ijms-20-00335" ref-type="bibr">38</xref>
,<xref rid="B39-ijms-20-00335" ref-type="bibr">39</xref>
,<xref rid="B40-ijms-20-00335" ref-type="bibr">40</xref>
,<xref rid="B41-ijms-20-00335" ref-type="bibr">41</xref>
,<xref rid="B42-ijms-20-00335" ref-type="bibr">42</xref>
,<xref rid="B43-ijms-20-00335" ref-type="bibr">43</xref>
,<xref rid="B44-ijms-20-00335" ref-type="bibr">44</xref>
,<xref rid="B45-ijms-20-00335" ref-type="bibr">45</xref>
,<xref rid="B46-ijms-20-00335" ref-type="bibr">46</xref>
,<xref rid="B47-ijms-20-00335" ref-type="bibr">47</xref>
,<xref rid="B48-ijms-20-00335" ref-type="bibr">48</xref>
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,<xref rid="B60-ijms-20-00335" ref-type="bibr">60</xref>
,<xref rid="B61-ijms-20-00335" ref-type="bibr">61</xref>
,<xref rid="B62-ijms-20-00335" ref-type="bibr">62</xref>
]. PTI: pathogen-associated molecular patterns (PAMP) triggered immunity; ETI: effector-triggered immunity; CRKI: cross-kingdom RNA interference.</p>
</caption>
<graphic xlink:href="ijms-20-00335-g001"></graphic>
</fig>
<fig id="ijms-20-00335-f002" orientation="portrait" position="float"><label>Figure 2</label>
<caption><p>The interaction mechanisms of plants-pathogens from three interacted and miRNA regulation layers. (<bold>A</bold>
) The three defensive layers in plants including the PTI, ETI, and cross-kingdom RNA interference (CKRI), and the three infection layers in pathogens including pattern recognition receptors (PRR), effector and CKRI. (<bold>B</bold>
) The evolution of <italic>NBS-LRR</italic>
genes and their regulator miRNAs. (<bold>C</bold>
) The three strategies of defense to biotic stresses including uORF [<xref rid="B83-ijms-20-00335" ref-type="bibr">83</xref>
], host-induced gene silencing (HIGS) [<xref rid="B84-ijms-20-00335" ref-type="bibr">84</xref>
,<xref rid="B85-ijms-20-00335" ref-type="bibr">85</xref>
,<xref rid="B86-ijms-20-00335" ref-type="bibr">86</xref>
,<xref rid="B87-ijms-20-00335" ref-type="bibr">87</xref>
,<xref rid="B88-ijms-20-00335" ref-type="bibr">88</xref>
,<xref rid="B89-ijms-20-00335" ref-type="bibr">89</xref>
,<xref rid="B90-ijms-20-00335" ref-type="bibr">90</xref>
,<xref rid="B91-ijms-20-00335" ref-type="bibr">91</xref>
,<xref rid="B92-ijms-20-00335" ref-type="bibr">92</xref>
] and spray-induced gene silencing (SIGS) [<xref rid="B3-ijms-20-00335" ref-type="bibr">3</xref>
,<xref rid="B93-ijms-20-00335" ref-type="bibr">93</xref>
,<xref rid="B94-ijms-20-00335" ref-type="bibr">94</xref>
,<xref rid="B95-ijms-20-00335" ref-type="bibr">95</xref>
,<xref rid="B96-ijms-20-00335" ref-type="bibr">96</xref>
] in plants.</p>
</caption>
<graphic xlink:href="ijms-20-00335-g002"></graphic>
</fig>
<table-wrap id="ijms-20-00335-t001" orientation="portrait" position="float"><object-id pub-id-type="pii">ijms-20-00335-t001_Table 1</object-id>
<label>Table 1</label>
<caption><p>Disease resistance genes and their regulator miRNAs in plants [<xref rid="B98-ijms-20-00335" ref-type="bibr">98</xref>
]. Mbp, million base pair; Nb, number; R-gene, disease resistance genes.</p>
</caption>
<table frame="hsides" rules="groups"><thead><tr><th rowspan="2" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" colspan="1">Species</th>
<th rowspan="2" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" colspan="1">Nb Chr.</th>
<th rowspan="2" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" colspan="1">Size (Mbp)</th>
<th rowspan="2" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" colspan="1">Nb Gene</th>
<th colspan="4" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1">R-Genes</th>
</tr>
<tr><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nb R-Genes</th>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">(%) <sup>1</sup>
</th>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nb MiRNA Targets</th>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">(%) <sup>2</sup>
</th>
</tr>
</thead>
<tbody><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><bold>Monocots</bold>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1"><italic>Oryza sativa</italic>
(rice)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">12</td>
<td align="center" valign="middle" rowspan="1" colspan="1">372</td>
<td align="center" valign="middle" rowspan="1" colspan="1">41,046</td>
<td align="center" valign="middle" rowspan="1" colspan="1">1196</td>
<td align="center" valign="middle" rowspan="1" colspan="1">2.91</td>
<td align="center" valign="middle" rowspan="1" colspan="1">144</td>
<td align="center" valign="middle" rowspan="1" colspan="1">12.04</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1"><italic>Sorghum bicolor</italic>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">10</td>
<td align="center" valign="middle" rowspan="1" colspan="1">659</td>
<td align="center" valign="middle" rowspan="1" colspan="1">34,008</td>
<td align="center" valign="middle" rowspan="1" colspan="1">625</td>
<td align="center" valign="middle" rowspan="1" colspan="1">1.84</td>
<td align="center" valign="middle" rowspan="1" colspan="1">109</td>
<td align="center" valign="middle" rowspan="1" colspan="1">17.44</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1"><italic>Zea mays</italic>
(maize)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">10</td>
<td align="center" valign="middle" rowspan="1" colspan="1">2365</td>
<td align="center" valign="middle" rowspan="1" colspan="1">32,540</td>
<td align="center" valign="middle" rowspan="1" colspan="1">673</td>
<td align="center" valign="middle" rowspan="1" colspan="1">2.07</td>
<td align="center" valign="middle" rowspan="1" colspan="1">0</td>
<td align="center" valign="middle" rowspan="1" colspan="1">0</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Brachypodium distachyon</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">5</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">271</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">25,504</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">537</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2.11</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">149</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">27.75</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><bold>Eudicots</bold>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1"><italic>Arabidopsis thaliana</italic>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">5</td>
<td align="center" valign="middle" rowspan="1" colspan="1">119</td>
<td align="center" valign="middle" rowspan="1" colspan="1">33,198</td>
<td align="center" valign="middle" rowspan="1" colspan="1">503</td>
<td align="center" valign="middle" rowspan="1" colspan="1">1.52</td>
<td align="center" valign="middle" rowspan="1" colspan="1">81</td>
<td align="center" valign="middle" rowspan="1" colspan="1">16.1</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1"><italic>Populus trichocarpa</italic>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">19</td>
<td align="center" valign="middle" rowspan="1" colspan="1">294</td>
<td align="center" valign="middle" rowspan="1" colspan="1">30,260</td>
<td align="center" valign="middle" rowspan="1" colspan="1">446</td>
<td align="center" valign="middle" rowspan="1" colspan="1">1.47</td>
<td align="center" valign="middle" rowspan="1" colspan="1">382</td>
<td align="center" valign="middle" rowspan="1" colspan="1">85.65</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">Carica papaya</td>
<td align="center" valign="middle" rowspan="1" colspan="1">9</td>
<td align="center" valign="middle" rowspan="1" colspan="1">234</td>
<td align="center" valign="middle" rowspan="1" colspan="1">19,205</td>
<td align="center" valign="middle" rowspan="1" colspan="1">228</td>
<td align="center" valign="middle" rowspan="1" colspan="1">1.19</td>
<td align="center" valign="middle" rowspan="1" colspan="1">0</td>
<td align="center" valign="middle" rowspan="1" colspan="1">0</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1"><italic>Glycine max</italic>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">20</td>
<td align="center" valign="middle" rowspan="1" colspan="1">949</td>
<td align="center" valign="middle" rowspan="1" colspan="1">46,164</td>
<td align="center" valign="middle" rowspan="1" colspan="1">1171</td>
<td align="center" valign="middle" rowspan="1" colspan="1">2.54</td>
<td align="center" valign="middle" rowspan="1" colspan="1">290</td>
<td align="center" valign="middle" rowspan="1" colspan="1">24.77</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Malusx</italic>
<italic>domestica</italic>
(apple)</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">17</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">742</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">58,979</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2052</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3.48</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">256</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">12.48</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn><p><sup>1</sup>
the percentage of the R-genes from the total coding genes; <sup>2</sup>
percentage of the miRNA target genes from the R-genes.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="ijms-20-00335-t002" orientation="portrait" position="float"><object-id pub-id-type="pii">ijms-20-00335-t002_Table 2</object-id>
<label>Table 2</label>
<caption><p>The validated disease resistance genes and their pathogens in plants. The data were derived from PRGDB database.</p>
</caption>
<table frame="hsides" rules="groups"><thead><tr><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Plant Species</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Disease</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Pathogens</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Avirus Genes</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Types of Pathogens</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Genes</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Types</th>
<th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">GenBank Locus</th>
</tr>
</thead>
<tbody><tr><td rowspan="6" align="center" valign="middle" style="border-bottom:solid thin" colspan="1"><italic>Arabidopsis thaliana</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">White rust of crucifers</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Albugo candida</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Oomycetes</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">RAC1</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AY522496</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Cucumber Mosaic Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Cucumber mosaic virus</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">RCY1</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AB087829</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bacterial Blight</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Pseudomonas syringae/Xanthomonas oryzae</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">avrRpm1; avrRpt2; avrPphB; N; avrRps4</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bacterium</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">RPM1; Rps2; RPS5; SSI4; Rps4</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL; CNL; CNL; TNL; TNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">NM_111584; NM_118742; NM_101094; AY179750; NM_123893</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Downy mildew of cucurbits</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Pseudoperonospora cubensis</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Oomycetes</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">RPP13/RPP8; RPP1/RPP4; RPP5</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL/CNL; TNL/TNL; TNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">NM_114520/NM_123713; NM_114316/NM_117790; NM_117798</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bacterial wilt of potato</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ralstonia solanacearum</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bacterium</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">RRS1</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">NM_001085246</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Turnip crinkle virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Turnip crinkle virus</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">HRT</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">NM_128190</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Aegilops tauschii</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Cereal cyst nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Heterodera avenae</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Cre1</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AY124651</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Capsicum chacoense</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bacterial spot of tomato</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Xanthomonas campestris</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AvrBs2</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bacterium</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bs2</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AF202179</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Capsicum chinense</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Pepper mild mottle virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Pepper mild mottle virus</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">L3</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">BAJ33559</td>
</tr>
<tr><td rowspan="3" align="center" valign="middle" style="border-bottom:solid thin" colspan="1"><italic>Cucumis melo</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fusarium Wilt</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Fusarium oxysporum</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">FOM-2</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DQ287965</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Melon aphid disease</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Aphis gossypii</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">insect</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">VAT</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AGH33848</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">zucchini yellow mosaic virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>zucchini yellow mosaic virus</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">FOM1</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AIU36098</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Glycine max</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Soybean mosaic virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>soybean mosaic virus</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">KR1</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AF327903</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Helianthus annuus</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Downy mildew of sunflower</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Plasmopara halstedii</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Oomycetes</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Pl8</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AY490793</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Hordeum vulgare</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Powdery mildew (barley)</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Blumeria graminis</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLA10/MLA1/MLA13</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AY266445; GU245961; AF523683</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Lactuca sativa</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Downy mildew of lettuce</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Bremia lactucae</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Avr3</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Oomycetes</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Dm3 (RGC2B)</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AH007213</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Linum usitatissimum</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Flax rust</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Melampsora lini</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">P2/L6/M; L,L1-L11; P,P1-4</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TNL; TNL; TNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AF310960/U27081/U73916; AAD25974/AAK28806</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Nicotiana glutinosa</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tobacco Mosaic Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Tobacco mosaic virus</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">N</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">U15605</td>
</tr>
<tr><td rowspan="3" align="center" valign="middle" style="border-bottom:solid thin" colspan="1"><italic>Oryza sativa</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Rice blast disease</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Magnaporthe grisea</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Avr-Pita</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Pi-ta/PIB</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AY196754</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bacterial Blight</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Pseudomonas syringae/Xanthomonas oryzae</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bacterium</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">XA1</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AB002266</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Rice blast disease</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Magnaporthe oryzae</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">RGA5</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">EU883792</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Oryza sativa Indica Group</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Rice blast disease</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Magnaporthe grisea</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Pi36/Pi9/Pi2</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DQ900896/DQ285630/DQ352453</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Oryza sativa Japonica Group</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Rice blast disease</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Magnaporthe grisea</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Piz-t/Pikm1-TS/Pikm2-TS/Pid3/Pi5-1/Pi5-2/Pit/Pikp-2</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DQ352040/AB462324/AB462325/FJ773286/EU869185/EU869186/AB379816/HM035360</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Rice blast disease</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Magnaporthe oryzae</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Pia; Pi37; Rpr1</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL; CNL; CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AB604626; DQ923494; AC119670</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Solanum acaule</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Latent mosaic of potato/Beet cyst nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Potato virus X/Heterodera schachtii</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus/Nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Rx2</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AJ249448</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Solanum bulbocastanum</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Late Blight of tomato</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Phytophthora infestans</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Oomycete</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Rpi-blb1/Rpi-blb2; RB</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL; CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AY336128; DQ122125; AY426259</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Solanum demissum</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Late Blight of tomato</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Phytophthora infestans</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Oomycete</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">R1</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AF447489</td>
</tr>
<tr><td rowspan="5" align="center" valign="middle" style="border-bottom:solid thin" colspan="1"><italic>Solanum lycopersicum</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bacterial spot of tomato</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Xanthomonas campestris</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Hax4/AvrBs4</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bacterium</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bs4</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AY438027</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Yellow potato cyst nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Yellow potato cyst nematode</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Hero</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AJ457052</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">root-knot nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Meloidogyne incognita</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Mi1.2</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AF039682</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tomato Spotted Wilt</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Tomato spotted wilt virus</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Sw-5</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AY007366</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tobacco Mosaic Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Tobacco mosaic virus</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MP</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tm-2a/Tm-2</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F536201/AF536200</td>
</tr>
<tr><td rowspan="2" align="center" valign="middle" style="border-bottom:solid thin" colspan="1"><italic>Solanum pimpinellifolium</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bacterial Speck of tomato</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Pseudomonas syringae</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AvrPto/AvrPtoB</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Bacterium</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Prf</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AF220602</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Late blight</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Phytophthora infestans</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Oomycete</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Ph-3</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">KJ563933</td>
</tr>
<tr><td rowspan="3" align="center" valign="middle" style="border-bottom:solid thin" colspan="1"><italic>Solanum tuberosum</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Yellow potato cyst nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Globodera</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Gpa2</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AF195939</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Late Blight of potato</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Phytophthora infestans</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Gro1.4</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AY196151</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Latent mosaic of potato/Beet cyst nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Potato virus X/Heterodera schachtii</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Rx</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AJ011801</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Solanum tuberosum subsp andigena</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Potato virus Y</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Potato virus Y</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">RY-1</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AJ300266</td>
</tr>
<tr><td rowspan="5" align="center" valign="middle" style="border-bottom:solid thin" colspan="1"><italic>Triticum aestivum</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Brown wheat rust of potato</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Puccinia triticina</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Lr10/Lr21/Lr1</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AY270157/FJ876280/EF439840</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">powdery mildew</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Blumeria graminis f. sp. Tritici</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Pm3</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AY325736</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">stem rust</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Puccinia graminis f. sp. Tritici</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Sr33</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">KF031303</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nematode disease</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Heterodera avenae</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Nematode</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Cre3</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AF052641</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Yellow rust</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Puccinia striiformis Westend. f.sp. Tritici</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Yr10</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AF149114</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Triticum monococcum subsp. Monococcum</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">stem rust</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Puccinia graminis f. sp. Tritici</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Sr35</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AGP75918</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Zea mays</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Common rust of maize</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Puccinia sorghi</italic>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungal</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Rp1-D</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CNL</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">AF107293</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="ijms-20-00335-t003" orientation="portrait" position="float"><object-id pub-id-type="pii">ijms-20-00335-t003_Table 3</object-id>
<label>Table 3</label>
<caption><p>List of regulators involved in the immunity response to pathogens in plants.</p>
</caption>
<table frame="hsides" rules="groups"><thead><tr><th rowspan="2" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" colspan="1">Plant miRNAs</th>
<th rowspan="2" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" colspan="1">Immunity Response</th>
<th rowspan="2" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" colspan="1">Targets in Plants or Pathogens</th>
<th rowspan="2" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" colspan="1">Positive (+)/Negative (−) Regulator</th>
<th colspan="2" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1">Pathogens</th>
</tr>
<tr><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Classification</th>
<th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Pathogen/Plant</th>
</tr>
</thead>
<tbody><tr><td align="center" valign="middle" rowspan="1" colspan="1">miR393</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PTI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">F-box auxin receptors</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Positive</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Bacteria</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Pseudomonas syringae</italic>
/<italic>Arabidopsis</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR160a</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PTI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">auxin response factors16</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Positive</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Bacteria</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Pseudomonas syringae</italic>
/<italic>Arabidopsis</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR319</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PTI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">TCP21</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Rice ragged stunt virus</italic>
(RRSV)/RICE</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR773</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PTI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">METHYLTRANSFERASE 2</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Bacteria; Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Pseudomonas syringae</italic>
/<italic>Arabidopsis</italic>
; <italic>Plectosphaerella cucumerina/Arabidopsis</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR169</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PTI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">NFYA</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Bacteria; Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Magnaporthe oryzae</italic>
/RICE</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR396</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PTI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">GRF</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Plectosphaerella cucumerina</italic>
/<italic>Arabidopsis</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR156</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PTI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">MdWRKYN1</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Alternaria alternaria</italic>
/APPLE</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR395</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PTI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">MdWRKY26</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Alternaria alternaria</italic>
/APPLE</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR5272</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PTI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">MKK6</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Fusarium oxysporum</italic>
/COTTON</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">MIR398</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PTI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">CSD2</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Bacteria</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Pseudomonas syringae</italic>
/<italic>Arabidopsis</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR164</td>
<td align="center" valign="middle" rowspan="1" colspan="1">PTI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">NAC</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Magnaporthe oryzae</italic>
/RICE</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR393*</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">MEMB12 (Membrin 12)</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Positive</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Bacteria</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Pseudomonas syringae</italic>
/<italic>Arabidopsis</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR444</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">MADS</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Positive</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Rice stripe virus</italic>
(RSV)/RICE</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR171</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">OsSCL6-Iia/b/c</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Positive</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Rice stripe virus</italic>
(RSV)/RICE</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR863-3p</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ARLPK1&ARLPK2</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Positive</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Bacteria</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Pseudomonas syringae</italic>
/<italic>Arabidopsis</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR863-3p</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">SERRATE</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Bacteria</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Pseudomonas syringae</italic>
/<italic>Arabidopsis</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">MIR9863</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>NBS-LRR</italic>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Blumeria graminis</italic>
/Barley</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">MIR482</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>NBS-LRR</italic>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Fusarium oxysporum</italic>
/Tomato</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">MIR5300</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>NBS-LRR</italic>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Fusarium oxysporum</italic>
/Tomato</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR1510</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>NBS-LRR</italic>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Phytophthora sojae</italic>
/Soybean</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR6019</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>NBS-LRR</italic>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Tobacco mosaic virus</italic>
/<italic>Tobacco</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR6020</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>NBS-LRR</italic>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Tobacco mosaic virus</italic>
/<italic>Tobacco</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR1885</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>NBS-LRR</italic>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Virus</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Turnip mosaic virus</italic>
/<italic>Brassica</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR472</td>
<td align="center" valign="middle" rowspan="1" colspan="1">ETI</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>NBS-LRR</italic>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Negative</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Bacteria</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Pseudomonas syringae</italic>
/<italic>Arabidopsis</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR166</td>
<td align="center" valign="middle" rowspan="1" colspan="1">CKRI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Clp-1 <sup>1</sup>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Positive</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Verticillium dahliae/Cotton</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">miR159</td>
<td align="center" valign="middle" rowspan="1" colspan="1">CKRI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">HiC-15 <sup>1</sup>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Positive</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Verticillium dahliae/Cotton</italic>
</td>
</tr>
<tr><td align="center" valign="middle" rowspan="1" colspan="1">TAS1c-siR483</td>
<td align="center" valign="middle" rowspan="1" colspan="1">CKRI</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Bc-Vps51&Bc-DCTN1 <sup>1</sup>
</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Positive</td>
<td align="center" valign="middle" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" rowspan="1" colspan="1"><italic>Botrytis cinerea /Arabidopsis</italic>
</td>
</tr>
<tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TAS2-siR453</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">CKRI</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">BC1T_08464 <sup>1</sup>
</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Positive</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Fungul</td>
<td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Botrytis cinerea /Arabidopsis</italic>
</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn><p>CKRI: Cross-kingdom RNA interference; <sup>1</sup>
Target gene from pathogen.</p>
</fn>
</table-wrap-foot>
</table-wrap>
</floats-group>
</pmc>
</record>
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