Structural Diversity of a Novel LTR Retrotransposon, RTPOSON, in the Genus Oryza
Identifieur interne : 001E84 ( Ncbi/Merge ); précédent : 001E83; suivant : 001E85Structural Diversity of a Novel LTR Retrotransposon, RTPOSON, in the Genus Oryza
Auteurs : Yu-Chia Hsu [Taïwan] ; Chang-Sheng Wang [Taïwan] ; Yann-Rong Lin [Taïwan] ; Yong-Pei Wu [Taïwan]Source :
- Evolutionary Bioinformatics Online [ 1176-9343 ] ; 2016.
Abstract
Retrotransposons with long terminal repeats (LTRs) are the most abundant transposable elements in plant genomes. A novel LTR retrotransposon named
Url:
DOI: 10.4137/EBO.S35158
PubMed: 26819544
PubMed Central: 4718150
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<author><name sortKey="Hsu, Yu Chia" sort="Hsu, Yu Chia" uniqKey="Hsu Y" first="Yu-Chia" last="Hsu">Yu-Chia Hsu</name>
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<author><name sortKey="Wang, Chang Sheng" sort="Wang, Chang Sheng" uniqKey="Wang C" first="Chang-Sheng" last="Wang">Chang-Sheng Wang</name>
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<author><name sortKey="Lin, Yann Rong" sort="Lin, Yann Rong" uniqKey="Lin Y" first="Yann-Rong" last="Lin">Yann-Rong Lin</name>
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<author><name sortKey="Wu, Yong Pei" sort="Wu, Yong Pei" uniqKey="Wu Y" first="Yong-Pei" last="Wu">Yong-Pei Wu</name>
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<sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Structural Diversity of a Novel LTR Retrotransposon, <italic>RTPOSON</italic>
, in the Genus <italic>Oryza</italic>
</title>
<author><name sortKey="Hsu, Yu Chia" sort="Hsu, Yu Chia" uniqKey="Hsu Y" first="Yu-Chia" last="Hsu">Yu-Chia Hsu</name>
<affiliation wicri:level="1"><nlm:aff id="af1-ebo-12-2016-029">Department of Agronomy, Chiayi Agricultural Experiment Station, Taiwan Agricultural Research Institute, Chiayi, Taiwan.</nlm:aff>
<country xml:lang="fr" wicri:curation="lc">Taïwan</country>
<wicri:regionArea>Department of Agronomy, Chiayi Agricultural Experiment Station, Taiwan Agricultural Research Institute, Chiayi</wicri:regionArea>
<wicri:noRegion>Chiayi</wicri:noRegion>
</affiliation>
</author>
<author><name sortKey="Wang, Chang Sheng" sort="Wang, Chang Sheng" uniqKey="Wang C" first="Chang-Sheng" last="Wang">Chang-Sheng Wang</name>
<affiliation wicri:level="1"><nlm:aff id="af2-ebo-12-2016-029">Department of Agronomy, National Chung Hsing University, Taichung, Taiwan.</nlm:aff>
<country xml:lang="fr" wicri:curation="lc">Taïwan</country>
<wicri:regionArea>Department of Agronomy, National Chung Hsing University, Taichung</wicri:regionArea>
<wicri:noRegion>Taichung</wicri:noRegion>
</affiliation>
</author>
<author><name sortKey="Lin, Yann Rong" sort="Lin, Yann Rong" uniqKey="Lin Y" first="Yann-Rong" last="Lin">Yann-Rong Lin</name>
<affiliation wicri:level="1"><nlm:aff id="af3-ebo-12-2016-029">Department of Agronomy, National Taiwan University, Taipei, Taiwan.</nlm:aff>
<country xml:lang="fr" wicri:curation="lc">Taïwan</country>
<wicri:regionArea>Department of Agronomy, National Taiwan University, Taipei</wicri:regionArea>
<wicri:noRegion>Taipei</wicri:noRegion>
</affiliation>
</author>
<author><name sortKey="Wu, Yong Pei" sort="Wu, Yong Pei" uniqKey="Wu Y" first="Yong-Pei" last="Wu">Yong-Pei Wu</name>
<affiliation wicri:level="1"><nlm:aff id="af1-ebo-12-2016-029">Department of Agronomy, Chiayi Agricultural Experiment Station, Taiwan Agricultural Research Institute, Chiayi, Taiwan.</nlm:aff>
<country xml:lang="fr" wicri:curation="lc">Taïwan</country>
<wicri:regionArea>Department of Agronomy, Chiayi Agricultural Experiment Station, Taiwan Agricultural Research Institute, Chiayi</wicri:regionArea>
<wicri:noRegion>Chiayi</wicri:noRegion>
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<series><title level="j">Evolutionary Bioinformatics Online</title>
<idno type="eISSN">1176-9343</idno>
<imprint><date when="2016">2016</date>
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<front><div type="abstract" xml:lang="en"><p>Retrotransposons with long terminal repeats (LTRs) are the most abundant transposable elements in plant genomes. A novel LTR retrotransposon named <italic>RTPOSON</italic>
primarily occurs in the genus <italic>Oryza</italic>
and in several species of the Poaceae family. <italic>RTPOSON</italic>
has been identified in the <italic>Ty1-copia</italic>
group of retrotransposons because two of its open reading frames encode an uncharacterized protein and UBN2_2 and zinc knuckle, respectively. More than 700 <italic>RTPOSONs</italic>
were identified in <italic>Oryza</italic>
genomes; 127 <italic>RTPOSONs</italic>
with LTRs and <italic>gag-pol</italic>
elements were classified into three subgroups. The subgroup <italic>RTPOSON_sub3</italic>
had the smallest DNA size and 97% (32/33) of <italic>RTPOSON</italic>
elements from <italic>Oryza punctata</italic>
are classified in this group. The insertion time of these <italic>RTPOSONs</italic>
varied and their proliferation occurred within the last 8 Mya, with two bursting periods within the last 1.5–5.0 Mya. A total of 37 different orthologous insertions of <italic>RTPOSONs</italic>
, with different nested transposable elements and gene fragments, were identified by comparing the genomes of ssp. <italic>japonica</italic>
cv. Nipponbare and ssp. <italic>indica</italic>
cv. 93–11. A part of intact <italic>RTPOSON</italic>
elements was evolved independently after the divergence of <italic>indica</italic>
and <italic>japonica</italic>
. In addition, intact <italic>RTPOSONs</italic>
and homologous fragments were preferentially retained or integrated in genic regions. This novel LTR retrotransposon, <italic>RTPOSON</italic>
, might have an impact on genome evolution, genic innovation, and genetic variation.</p>
</div>
</front>
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</author>
<author><name sortKey="Estill, Jc" uniqKey="Estill J">JC Estill</name>
</author>
<author><name sortKey="Leebens Mack, J" uniqKey="Leebens Mack J">J Leebens-Mack</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Zhang, Lb" uniqKey="Zhang L">LB Zhang</name>
</author>
<author><name sortKey="Ge, S" uniqKey="Ge S">S Ge</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Huang, X" uniqKey="Huang X">X Huang</name>
</author>
<author><name sortKey="Lu, G" uniqKey="Lu G">G Lu</name>
</author>
<author><name sortKey="Zhao, Q" uniqKey="Zhao Q">Q Zhao</name>
</author>
</analytic>
</biblStruct>
<biblStruct><analytic><author><name sortKey="Kashkush, K" uniqKey="Kashkush K">K Kashkush</name>
</author>
<author><name sortKey="Feldman, M" uniqKey="Feldman M">M Feldman</name>
</author>
<author><name sortKey="Levy, Aa" uniqKey="Levy A">AA Levy</name>
</author>
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</biblStruct>
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</author>
<author><name sortKey="Sarazin, A" uniqKey="Sarazin A">A Sarazin</name>
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</biblStruct>
</listBibl>
</div1>
</back>
</TEI>
<pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Evol Bioinform Online</journal-id>
<journal-id journal-id-type="iso-abbrev">Evol. Bioinform. Online</journal-id>
<journal-id journal-id-type="publisher-id">101256319</journal-id>
<journal-title-group><journal-title>Evolutionary Bioinformatics Online</journal-title>
</journal-title-group>
<issn pub-type="epub">1176-9343</issn>
<publisher><publisher-name>Libertas Academica</publisher-name>
</publisher>
</journal-meta>
<article-meta><article-id pub-id-type="pmid">26819544</article-id>
<article-id pub-id-type="pmc">4718150</article-id>
<article-id pub-id-type="doi">10.4137/EBO.S35158</article-id>
<article-id pub-id-type="publisher-id">ebo-12-2016-029</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Original Research</subject>
</subj-group>
</article-categories>
<title-group><article-title>Structural Diversity of a Novel LTR Retrotransposon, <italic>RTPOSON</italic>
, in the Genus <italic>Oryza</italic>
</article-title>
</title-group>
<contrib-group><contrib contrib-type="author"><name><surname>Hsu</surname>
<given-names>Yu-Chia</given-names>
</name>
<xref ref-type="aff" rid="af1-ebo-12-2016-029">1</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Wang</surname>
<given-names>Chang-Sheng</given-names>
</name>
<xref ref-type="aff" rid="af2-ebo-12-2016-029">2</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Lin</surname>
<given-names>Yann-Rong</given-names>
</name>
<xref ref-type="aff" rid="af3-ebo-12-2016-029">3</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Wu</surname>
<given-names>Yong-Pei</given-names>
</name>
<xref ref-type="aff" rid="af1-ebo-12-2016-029">1</xref>
<xref ref-type="corresp" rid="c1-ebo-12-2016-029"></xref>
</contrib>
</contrib-group>
<aff id="af1-ebo-12-2016-029"><label>1</label>
Department of Agronomy, Chiayi Agricultural Experiment Station, Taiwan Agricultural Research Institute, Chiayi, Taiwan.</aff>
<aff id="af2-ebo-12-2016-029"><label>2</label>
Department of Agronomy, National Chung Hsing University, Taichung, Taiwan.</aff>
<aff id="af3-ebo-12-2016-029"><label>3</label>
Department of Agronomy, National Taiwan University, Taipei, Taiwan.</aff>
<author-notes><corresp id="c1-ebo-12-2016-029">CORRESPONDENCE: <email>wuypei@dns.caes.gov.tw</email>
</corresp>
</author-notes>
<pub-date pub-type="collection"><year>2016</year>
</pub-date>
<pub-date pub-type="epub"><day>18</day>
<month>1</month>
<year>2016</year>
</pub-date>
<volume>12</volume>
<fpage>29</fpage>
<lpage>40</lpage>
<history><date date-type="received"><day>21</day>
<month>9</month>
<year>2015</year>
</date>
<date date-type="rev-recd"><day>08</day>
<month>11</month>
<year>2015</year>
</date>
<date date-type="accepted"><day>16</day>
<month>11</month>
<year>2015</year>
</date>
</history>
<permissions><copyright-statement>© 2016 the author(s), publisher and licensee Libertas Academica Ltd.</copyright-statement>
<copyright-year>2016</copyright-year>
<license license-type="open-access"><license-p>This is an open-access article distributed under the terms of the Creative Commons CC-BY-NC 3.0 License.</license-p>
</license>
</permissions>
<abstract><p>Retrotransposons with long terminal repeats (LTRs) are the most abundant transposable elements in plant genomes. A novel LTR retrotransposon named <italic>RTPOSON</italic>
primarily occurs in the genus <italic>Oryza</italic>
and in several species of the Poaceae family. <italic>RTPOSON</italic>
has been identified in the <italic>Ty1-copia</italic>
group of retrotransposons because two of its open reading frames encode an uncharacterized protein and UBN2_2 and zinc knuckle, respectively. More than 700 <italic>RTPOSONs</italic>
were identified in <italic>Oryza</italic>
genomes; 127 <italic>RTPOSONs</italic>
with LTRs and <italic>gag-pol</italic>
elements were classified into three subgroups. The subgroup <italic>RTPOSON_sub3</italic>
had the smallest DNA size and 97% (32/33) of <italic>RTPOSON</italic>
elements from <italic>Oryza punctata</italic>
are classified in this group. The insertion time of these <italic>RTPOSONs</italic>
varied and their proliferation occurred within the last 8 Mya, with two bursting periods within the last 1.5–5.0 Mya. A total of 37 different orthologous insertions of <italic>RTPOSONs</italic>
, with different nested transposable elements and gene fragments, were identified by comparing the genomes of ssp. <italic>japonica</italic>
cv. Nipponbare and ssp. <italic>indica</italic>
cv. 93–11. A part of intact <italic>RTPOSON</italic>
elements was evolved independently after the divergence of <italic>indica</italic>
and <italic>japonica</italic>
. In addition, intact <italic>RTPOSONs</italic>
and homologous fragments were preferentially retained or integrated in genic regions. This novel LTR retrotransposon, <italic>RTPOSON</italic>
, might have an impact on genome evolution, genic innovation, and genetic variation.</p>
</abstract>
<kwd-group><kwd>LTR retrotransposon</kwd>
<kwd><italic>RTPOSON</italic>
</kwd>
<kwd>genome</kwd>
<kwd><italic>Oryza</italic>
</kwd>
<kwd>evolution</kwd>
<kwd>genetic variation</kwd>
</kwd-group>
</article-meta>
</front>
<floats-group><fig id="f1-ebo-12-2016-029" position="float"><label>Figure 1</label>
<caption><p>The gene structure of the novel retrotransposon, <italic>RTPOSON</italic>
. (<bold>A</bold>
) Structural annotation for one of the intact <italic>RTPOSON</italic>
elements from <italic>O. sativa</italic>
ssp. <italic>indica</italic>
, 93-11. The 5′- and 3′-LTRs are shown in the light blue arrows; TSD indicates the 5-bp target site duplication; PBS indicates the primer-binding site; PPT indicates the PPT; <italic>gag-pol</italic>
is able to transcribe a polyprotein that is provided for the transposition activity of retrotransposon. The gene, <italic>gag-pol</italic>
, contains two ORFs, of which ORF1 contains the uncharacterized protein and ORF2 contains two domains, UBN2_2 (pfam144227) for <italic>gag</italic>
and zinc knuckle (pfam00098). (<bold>B</bold>
) The sequence alignment of two LTRs of intact <italic>RTPOSON</italic>
elements randomly selected from 93-11. The identical nucleotides are indicated by gray shading. (<bold>C</bold>
) Homology matrix between the <italic>RTPOSON</italic>
element and the tobacco Ty1-<italic>copia Tnt1</italic>
(X13777) internal coding region. The approximate regions corresponding to the <italic>gag</italic>
, zinc knuckle, and PR domains of tobacco Ty1-<italic>copia</italic>
encoded by <italic>pol</italic>
are indicated below the dotted lines. The second long match region is similar to zinc knuckle of <italic>Tnt1</italic>
.</p>
</caption>
<graphic xlink:href="ebo-12-2016-029f1"></graphic>
</fig>
<fig id="f2-ebo-12-2016-029" position="float"><label>Figure 2</label>
<caption><p>Phylogenetic tree and schematic representation for different groups of <italic>RTPOSON</italic>
retrotransposons from the genus <italic>Oryza</italic>
. (<bold>A</bold>
) The neighbor-joining tree was constructed based on <italic>gag</italic>
sequences. The <italic>gag</italic>
sequences were aligned by using MUSCLE,<xref rid="b55-ebo-12-2016-029" ref-type="bibr">55</xref>
and then the tree was constructed by using MEGA version 6.0. Bootstrap values were calculated for 1,000 replicates. The scale bar indicates nucleotide sequence divergence. (<bold>B</bold>
) Organization and structural features of the three groups of <italic>RTPOSON</italic>
retrotransposons. The representative elements were randomly selected from each group. The light and dark blue arrows represent LTRs and <italic>gag-pol</italic>
regions, respectively. The gray areas represent the conserved regions between the different groups.</p>
</caption>
<graphic xlink:href="ebo-12-2016-029f2"></graphic>
</fig>
<fig id="f3-ebo-12-2016-029" position="float"><label>Figure 3</label>
<caption><p>The estimated insertion times of <italic>RTPOSON</italic>
elements in <italic>Oryza</italic>
rice. (<bold>A</bold>
) The 127 intact elements from <italic>Oryza</italic>
were used to estimate the insertion times of <italic>RTPOSON</italic>
. Kimura distances were converted to millions of years ago using the substitution rate of 1.3 × 10<sup>−8</sup>
(<bold>B</bold>
) Insertion times of intact <italic>RTPOSON</italic>
from eight <italic>Oryza</italic>
species.</p>
</caption>
<graphic xlink:href="ebo-12-2016-029f3"></graphic>
</fig>
<fig id="f4-ebo-12-2016-029" position="float"><label>Figure 4</label>
<caption><p>The orthologous regions of <italic>RTPOSON</italic>
elements between Nipponbare and 93-11. The differential locations of <italic>RTPOSON</italic>
elements between Nipponbare and 93-11 were aligned and annotated by using the Vector NTI 7.0 program. The order shown is based on the number of TE insertion types, such as Ty1-<italic>copia</italic>
, Ty3-<italic>gypsy</italic>
, and <italic>En-Spm</italic>
. (<bold>A</bold>
–<bold>F</bold>
) Different types of TEs were inserted within <italic>RTPOSON</italic>
in Nipponbare and 93-11. (<bold>G</bold>
) An additional copy of <italic>RTPOSON</italic>
within its structure in 93-11. (<bold>H</bold>
–<bold>K</bold>
) The TEs or functional genes were inserted in the structure of <italic>RTPOSON</italic>
in Nipponbare. (<bold>L</bold>
) The truncated gene structure of <italic>RTPOSON</italic>
in 93-11. (<bold>M</bold>
) The intact <italic>RTPOSON</italic>
was inserted in Nipponbare. A scale and a key for the domains represented in the schematic representation are shown in the bottom right-hand corner. Abbreviations and color coding of domains: LTR, long terminal repeat (light blue); <italic>gag-pol</italic>
, gag and polyprotein (dark blue); genes are represented by red arrows; TEs are indicated by gray arrows; homolog flanking sequences between Nipponbare and 93-11 are represented by red and blue bars, respectively; <italic>HYP</italic>
, hypothetical proteins.</p>
</caption>
<graphic xlink:href="ebo-12-2016-029f4"></graphic>
</fig>
<fig id="f5-ebo-12-2016-029" position="float"><label>Figure 5</label>
<caption><p>The orthologous insertion and <italic>RTPOSON</italic>
elements across the genus <italic>Oryza</italic>
. <italic>RTPOSON</italic>
elements are present in (<bold>A</bold>
) STE kinase (LOC_ Os02g44642), (<bold>B</bold>
) disease-resistant RGH2B protein (LOC_Os10g22484), (<bold>C</bold>
) disease-resistant RPP13-like protein 1 (LOC_Os10g04090), (<bold>D</bold>
) glycosyl transferase (LOC_Os11g03160). The LTRs of <italic>RTPOSON</italic>
share high similarity of UTR with (<bold>E</bold>
) EARLY flowering protein (LOC_Os08g27870) and (<bold>F</bold>
) dehydrogenase (LOC_Os10g41170), in each <italic>Oryza</italic>
species. The white boxes and arrows represent the 5′- and 3′ UTR of the orthologous genes. The black boxes indicate the exons of genes. The light and dark blue arrows represent LTRs and <italic>gag-pol</italic>
of <italic>RTPOSON</italic>
, respectively. Gray areas represent conserved regions between <italic>RTPOSON</italic>
and genes. The number in gray areas is sequence identity (%). The UTR position was defined based on RAP-DB (<ext-link ext-link-type="uri" xlink:href="http://rapdb.dna.affrc.go.jp/">http://rapdb.dna.affrc.go.jp/</ext-link>
).</p>
<p><bold>Abbreviation:</bold>
UTR, untranslated region.</p>
</caption>
<graphic xlink:href="ebo-12-2016-029f5"></graphic>
</fig>
<table-wrap id="t1-ebo-12-2016-029" position="float"><label>Table 1</label>
<caption><p>Distribution of <italic>RTPOSON</italic>
in 13 <italic>Oryza</italic>
species.</p>
</caption>
<table frame="box" rules="rows"><thead><tr><th rowspan="2" valign="top" align="left" colspan="1">SPECIES</th>
<th rowspan="2" valign="top" align="left" colspan="1">GENOME</th>
<th rowspan="2" valign="top" align="left" colspan="1">GENOME SIZE (Mb)<xref ref-type="table-fn" rid="tfn1-ebo-12-2016-029">1</xref>
</th>
<th colspan="4" valign="top" align="left" rowspan="1">COPY NUMBER</th>
</tr>
<tr><th valign="top" align="left" rowspan="1" colspan="1">INTACT</th>
<th valign="top" align="left" rowspan="1" colspan="1">FRAGMENT</th>
<th valign="top" align="left" rowspan="1" colspan="1">SOLO-LTR</th>
<th valign="top" align="left" rowspan="1" colspan="1">TOTAL</th>
</tr>
</thead>
<tbody><tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. sativa</italic>
ssp. <italic>japonica</italic>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">AA</td>
<td valign="top" align="left" rowspan="1" colspan="1">389</td>
<td valign="top" align="left" rowspan="1" colspan="1">15</td>
<td valign="top" align="left" rowspan="1" colspan="1">37</td>
<td valign="top" align="left" rowspan="1" colspan="1">41</td>
<td valign="top" align="left" rowspan="1" colspan="1">93</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. sativa</italic>
ssp. <italic>indica</italic>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">AA</td>
<td valign="top" align="left" rowspan="1" colspan="1">400</td>
<td valign="top" align="left" rowspan="1" colspan="1">13</td>
<td valign="top" align="left" rowspan="1" colspan="1">40</td>
<td valign="top" align="left" rowspan="1" colspan="1">32</td>
<td valign="top" align="left" rowspan="1" colspan="1">85</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. nivara</italic>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">AA</td>
<td valign="top" align="left" rowspan="1" colspan="1">448</td>
<td valign="top" align="left" rowspan="1" colspan="1">14</td>
<td valign="top" align="left" rowspan="1" colspan="1">26</td>
<td valign="top" align="left" rowspan="1" colspan="1">30</td>
<td valign="top" align="left" rowspan="1" colspan="1">70</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. rufipogon</italic>
<xref ref-type="table-fn" rid="tfn2-ebo-12-2016-029">2</xref>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">AA</td>
<td valign="top" align="left" rowspan="1" colspan="1">445</td>
<td valign="top" align="left" rowspan="1" colspan="1">1</td>
<td valign="top" align="left" rowspan="1" colspan="1">4</td>
<td valign="top" align="left" rowspan="1" colspan="1">2</td>
<td valign="top" align="left" rowspan="1" colspan="1">7</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. glaberrima</italic>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">AA</td>
<td valign="top" align="left" rowspan="1" colspan="1">354</td>
<td valign="top" align="left" rowspan="1" colspan="1">16</td>
<td valign="top" align="left" rowspan="1" colspan="1">18</td>
<td valign="top" align="left" rowspan="1" colspan="1">33</td>
<td valign="top" align="left" rowspan="1" colspan="1">67</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. barthii</italic>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">AA</td>
<td valign="top" align="left" rowspan="1" colspan="1">411</td>
<td valign="top" align="left" rowspan="1" colspan="1">18</td>
<td valign="top" align="left" rowspan="1" colspan="1">28</td>
<td valign="top" align="left" rowspan="1" colspan="1">26</td>
<td valign="top" align="left" rowspan="1" colspan="1">72</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. meridionalis</italic>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">AA</td>
<td valign="top" align="left" rowspan="1" colspan="1">435</td>
<td valign="top" align="left" rowspan="1" colspan="1">5</td>
<td valign="top" align="left" rowspan="1" colspan="1">27</td>
<td valign="top" align="left" rowspan="1" colspan="1">41</td>
<td valign="top" align="left" rowspan="1" colspan="1">73</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. glumaepatula</italic>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">AA</td>
<td valign="top" align="left" rowspan="1" colspan="1">464</td>
<td valign="top" align="left" rowspan="1" colspan="1">10</td>
<td valign="top" align="left" rowspan="1" colspan="1">32</td>
<td valign="top" align="left" rowspan="1" colspan="1">41</td>
<td valign="top" align="left" rowspan="1" colspan="1">83</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. punctata</italic>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">BB</td>
<td valign="top" align="left" rowspan="1" colspan="1">423</td>
<td valign="top" align="left" rowspan="1" colspan="1">33</td>
<td valign="top" align="left" rowspan="1" colspan="1">25</td>
<td valign="top" align="left" rowspan="1" colspan="1">18</td>
<td valign="top" align="left" rowspan="1" colspan="1">76</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. minuta</italic>
<xref ref-type="table-fn" rid="tfn2-ebo-12-2016-029">2</xref>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">BBCC</td>
<td valign="top" align="left" rowspan="1" colspan="1">1,124</td>
<td valign="top" align="left" rowspan="1" colspan="1">1</td>
<td valign="top" align="left" rowspan="1" colspan="1">2</td>
<td valign="top" align="left" rowspan="1" colspan="1">7</td>
<td valign="top" align="left" rowspan="1" colspan="1">10</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. officinalis</italic>
<xref ref-type="table-fn" rid="tfn2-ebo-12-2016-029">2</xref>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">CC</td>
<td valign="top" align="left" rowspan="1" colspan="1">653</td>
<td valign="top" align="left" rowspan="1" colspan="1">1</td>
<td valign="top" align="left" rowspan="1" colspan="1">11</td>
<td valign="top" align="left" rowspan="1" colspan="1">13</td>
<td valign="top" align="left" rowspan="1" colspan="1">25</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. brachyantha</italic>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">FF</td>
<td valign="top" align="left" rowspan="1" colspan="1">260</td>
<td valign="top" align="left" rowspan="1" colspan="1">0</td>
<td valign="top" align="left" rowspan="1" colspan="1">6</td>
<td valign="top" align="left" rowspan="1" colspan="1">13</td>
<td valign="top" align="left" rowspan="1" colspan="1">19</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"><italic>O. granulata</italic>
<xref ref-type="table-fn" rid="tfn2-ebo-12-2016-029">2</xref>
</td>
<td valign="top" align="left" rowspan="1" colspan="1">GG</td>
<td valign="top" align="left" rowspan="1" colspan="1">862</td>
<td valign="top" align="left" rowspan="1" colspan="1">0</td>
<td valign="top" align="left" rowspan="1" colspan="1">6</td>
<td valign="top" align="left" rowspan="1" colspan="1">19</td>
<td valign="top" align="left" rowspan="1" colspan="1">25</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1">Total</td>
<td valign="top" align="left" rowspan="1" colspan="1"></td>
<td valign="top" align="left" rowspan="1" colspan="1"></td>
<td valign="top" align="left" rowspan="1" colspan="1">127</td>
<td valign="top" align="left" rowspan="1" colspan="1">262</td>
<td valign="top" align="left" rowspan="1" colspan="1">316</td>
<td valign="top" align="left" rowspan="1" colspan="1">705</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn><p><bold>Notes:</bold>
</p>
</fn>
<fn id="tfn1-ebo-12-2016-029"><label>1</label>
<p>Reference from the study by Jacquemin et al.<xref rid="b18-ebo-12-2016-029" ref-type="bibr">18</xref>
</p>
</fn>
<fn id="tfn2-ebo-12-2016-029"><label>2</label>
<p>A limited number of sequences were searched and collected from rice chromosome 3 in Gramene website.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="t2-ebo-12-2016-029" position="float"><label>Table 2</label>
<caption><p>Insertion sites of intact <italic>RTPOSON</italic>
in <italic>japonica</italic>
rice genome.</p>
</caption>
<table frame="box" rules="rows"><thead><tr><th valign="top" align="left" rowspan="1" colspan="1">INSERTION SITE</th>
<th valign="top" align="left" rowspan="1" colspan="1">NO. OF ELEMENT</th>
</tr>
</thead>
<tbody><tr><td valign="top" align="left" rowspan="1" colspan="1">Intergenic region</td>
<td valign="top" align="left" rowspan="1" colspan="1">64</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1">Gene</td>
<td valign="top" align="left" rowspan="1" colspan="1">11</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"> intron</td>
<td valign="top" align="left" rowspan="1" colspan="1">9</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"> exon</td>
<td valign="top" align="left" rowspan="1" colspan="1">0</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1"> 5′- or 3′-UTR</td>
<td valign="top" align="left" rowspan="1" colspan="1">2</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1">Putative gene</td>
<td valign="top" align="left" rowspan="1" colspan="1">9</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1">Within 1 kb flaking gene</td>
<td valign="top" align="left" rowspan="1" colspan="1">24</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1">Within other types of TEs</td>
<td valign="top" align="left" rowspan="1" colspan="1">19</td>
</tr>
<tr><td valign="top" align="left" rowspan="1" colspan="1">Total</td>
<td valign="top" align="left" rowspan="1" colspan="1">127</td>
</tr>
</tbody>
</table>
</table-wrap>
</floats-group>
</pmc>
<affiliations><list><country><li>Taïwan</li>
</country>
</list>
<tree><country name="Taïwan"><noRegion><name sortKey="Hsu, Yu Chia" sort="Hsu, Yu Chia" uniqKey="Hsu Y" first="Yu-Chia" last="Hsu">Yu-Chia Hsu</name>
</noRegion>
<name sortKey="Lin, Yann Rong" sort="Lin, Yann Rong" uniqKey="Lin Y" first="Yann-Rong" last="Lin">Yann-Rong Lin</name>
<name sortKey="Wang, Chang Sheng" sort="Wang, Chang Sheng" uniqKey="Wang C" first="Chang-Sheng" last="Wang">Chang-Sheng Wang</name>
<name sortKey="Wu, Yong Pei" sort="Wu, Yong Pei" uniqKey="Wu Y" first="Yong-Pei" last="Wu">Yong-Pei Wu</name>
</country>
</tree>
</affiliations>
</record>
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