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Structural Diversity of a Novel LTR Retrotransposon, RTPOSON, in the Genus Oryza

Identifieur interne : 001E84 ( Ncbi/Merge ); précédent : 001E83; suivant : 001E85

Structural 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 :

RBID : PMC:4718150

Abstract

Retrotransposons with long terminal repeats (LTRs) are the most abundant transposable elements in plant genomes. A novel LTR retrotransposon named RTPOSON primarily occurs in the genus Oryza and in several species of the Poaceae family. RTPOSON has been identified in the Ty1-copia group of retrotransposons because two of its open reading frames encode an uncharacterized protein and UBN2_2 and zinc knuckle, respectively. More than 700 RTPOSONs were identified in Oryza genomes; 127 RTPOSONs with LTRs and gag-pol elements were classified into three subgroups. The subgroup RTPOSON_sub3 had the smallest DNA size and 97% (32/33) of RTPOSON elements from Oryza punctata are classified in this group. The insertion time of these RTPOSONs 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 RTPOSONs, with different nested transposable elements and gene fragments, were identified by comparing the genomes of ssp. japonica cv. Nipponbare and ssp. indica cv. 93–11. A part of intact RTPOSON elements was evolved independently after the divergence of indica and japonica. In addition, intact RTPOSONs and homologous fragments were preferentially retained or integrated in genic regions. This novel LTR retrotransposon, RTPOSON, might have an impact on genome evolution, genic innovation, and genetic variation.


Url:
DOI: 10.4137/EBO.S35158
PubMed: 26819544
PubMed Central: 4718150

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PMC:4718150

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<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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<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>
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</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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