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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Positional cloning and characterization reveal the molecular basis for soybean maturity locus <italic>E1</italic>
that regulates photoperiodic flowering</title>
<author><name sortKey="Xia, Zhengjun" sort="Xia, Zhengjun" uniqKey="Xia Z" first="Zhengjun" last="Xia">Zhengjun Xia</name>
<affiliation><nlm:aff id="aff1">Northeast Institute of Geography and Agroecology,<institution>Chinese Academy of Sciences</institution>
, Harbin 150081,<country>China</country>
;</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="aff2">Soybean Applied Genomics Research Unit,<institution>National Institute of Agrobiological Sciences</institution>
, Tsukuba 305-8602,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Watanabe, Satoshi" sort="Watanabe, Satoshi" uniqKey="Watanabe S" first="Satoshi" last="Watanabe">Satoshi Watanabe</name>
<affiliation><nlm:aff id="aff2">Soybean Applied Genomics Research Unit,<institution>National Institute of Agrobiological Sciences</institution>
, Tsukuba 305-8602,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Yamada, Tetsuya" sort="Yamada, Tetsuya" uniqKey="Yamada T" first="Tetsuya" last="Yamada">Tetsuya Yamada</name>
<affiliation><nlm:aff id="aff3">Graduate School of Agriculture,<institution>Hokkaido University</institution>
, Sapporo 060-8589,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Tsubokura, Yasutaka" sort="Tsubokura, Yasutaka" uniqKey="Tsubokura Y" first="Yasutaka" last="Tsubokura">Yasutaka Tsubokura</name>
<affiliation><nlm:aff id="aff2">Soybean Applied Genomics Research Unit,<institution>National Institute of Agrobiological Sciences</institution>
, Tsukuba 305-8602,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Nakashima, Hiroko" sort="Nakashima, Hiroko" uniqKey="Nakashima H" first="Hiroko" last="Nakashima">Hiroko Nakashima</name>
<affiliation><nlm:aff id="aff4">Faculty of Agriculture,<institution>Saga University</institution>
, Saga 840-8502,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Zhai, Hong" sort="Zhai, Hong" uniqKey="Zhai H" first="Hong" last="Zhai">Hong Zhai</name>
<affiliation><nlm:aff id="aff1">Northeast Institute of Geography and Agroecology,<institution>Chinese Academy of Sciences</institution>
, Harbin 150081,<country>China</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Anai, Toyoaki" sort="Anai, Toyoaki" uniqKey="Anai T" first="Toyoaki" last="Anai">Toyoaki Anai</name>
<affiliation><nlm:aff id="aff4">Faculty of Agriculture,<institution>Saga University</institution>
, Saga 840-8502,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Sato, Shusei" sort="Sato, Shusei" uniqKey="Sato S" first="Shusei" last="Sato">Shusei Sato</name>
<affiliation><nlm:aff wicri:cut="; and" id="aff5">Department of Plant Genome Research,<institution>Kazusa DNA Research Institute</institution>
, Kisarazu 292-0812,<country>Japan</country>
</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Yamazaki, Toshimasa" sort="Yamazaki, Toshimasa" uniqKey="Yamazaki T" first="Toshimasa" last="Yamazaki">Toshimasa Yamazaki</name>
<affiliation><nlm:aff id="aff6">Biomolecular Research Unit,<institution>National Institute of Agrobiological Sciences</institution>
, Tsukuba 305-8602,<country>Japan</country>
</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Lu, Shixiang" sort="Lu, Shixiang" uniqKey="Lu S" first="Shixiang" last="Lü">Shixiang Lü</name>
<affiliation><nlm:aff id="aff1">Northeast Institute of Geography and Agroecology,<institution>Chinese Academy of Sciences</institution>
, Harbin 150081,<country>China</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Wu, Hongyan" sort="Wu, Hongyan" uniqKey="Wu H" first="Hongyan" last="Wu">Hongyan Wu</name>
<affiliation><nlm:aff id="aff1">Northeast Institute of Geography and Agroecology,<institution>Chinese Academy of Sciences</institution>
, Harbin 150081,<country>China</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Tabata, Satoshi" sort="Tabata, Satoshi" uniqKey="Tabata S" first="Satoshi" last="Tabata">Satoshi Tabata</name>
<affiliation><nlm:aff wicri:cut="; and" id="aff5">Department of Plant Genome Research,<institution>Kazusa DNA Research Institute</institution>
, Kisarazu 292-0812,<country>Japan</country>
</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Harada, Kyuya" sort="Harada, Kyuya" uniqKey="Harada K" first="Kyuya" last="Harada">Kyuya Harada</name>
<affiliation><nlm:aff id="aff2">Soybean Applied Genomics Research Unit,<institution>National Institute of Agrobiological Sciences</institution>
, Tsukuba 305-8602,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
</titleStmt>
<publicationStmt><idno type="wicri:source">PMC</idno>
<idno type="pmid">22619331</idno>
<idno type="pmc">3420212</idno>
<idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3420212</idno>
<idno type="RBID">PMC:3420212</idno>
<idno type="doi">10.1073/pnas.1117982109</idno>
<date when="2012">2012</date>
<idno type="wicri:Area/Pmc/Corpus">000D37</idno>
</publicationStmt>
<sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Positional cloning and characterization reveal the molecular basis for soybean maturity locus <italic>E1</italic>
that regulates photoperiodic flowering</title>
<author><name sortKey="Xia, Zhengjun" sort="Xia, Zhengjun" uniqKey="Xia Z" first="Zhengjun" last="Xia">Zhengjun Xia</name>
<affiliation><nlm:aff id="aff1">Northeast Institute of Geography and Agroecology,<institution>Chinese Academy of Sciences</institution>
, Harbin 150081,<country>China</country>
;</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="aff2">Soybean Applied Genomics Research Unit,<institution>National Institute of Agrobiological Sciences</institution>
, Tsukuba 305-8602,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Watanabe, Satoshi" sort="Watanabe, Satoshi" uniqKey="Watanabe S" first="Satoshi" last="Watanabe">Satoshi Watanabe</name>
<affiliation><nlm:aff id="aff2">Soybean Applied Genomics Research Unit,<institution>National Institute of Agrobiological Sciences</institution>
, Tsukuba 305-8602,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Yamada, Tetsuya" sort="Yamada, Tetsuya" uniqKey="Yamada T" first="Tetsuya" last="Yamada">Tetsuya Yamada</name>
<affiliation><nlm:aff id="aff3">Graduate School of Agriculture,<institution>Hokkaido University</institution>
, Sapporo 060-8589,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Tsubokura, Yasutaka" sort="Tsubokura, Yasutaka" uniqKey="Tsubokura Y" first="Yasutaka" last="Tsubokura">Yasutaka Tsubokura</name>
<affiliation><nlm:aff id="aff2">Soybean Applied Genomics Research Unit,<institution>National Institute of Agrobiological Sciences</institution>
, Tsukuba 305-8602,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Nakashima, Hiroko" sort="Nakashima, Hiroko" uniqKey="Nakashima H" first="Hiroko" last="Nakashima">Hiroko Nakashima</name>
<affiliation><nlm:aff id="aff4">Faculty of Agriculture,<institution>Saga University</institution>
, Saga 840-8502,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Zhai, Hong" sort="Zhai, Hong" uniqKey="Zhai H" first="Hong" last="Zhai">Hong Zhai</name>
<affiliation><nlm:aff id="aff1">Northeast Institute of Geography and Agroecology,<institution>Chinese Academy of Sciences</institution>
, Harbin 150081,<country>China</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Anai, Toyoaki" sort="Anai, Toyoaki" uniqKey="Anai T" first="Toyoaki" last="Anai">Toyoaki Anai</name>
<affiliation><nlm:aff id="aff4">Faculty of Agriculture,<institution>Saga University</institution>
, Saga 840-8502,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Sato, Shusei" sort="Sato, Shusei" uniqKey="Sato S" first="Shusei" last="Sato">Shusei Sato</name>
<affiliation><nlm:aff wicri:cut="; and" id="aff5">Department of Plant Genome Research,<institution>Kazusa DNA Research Institute</institution>
, Kisarazu 292-0812,<country>Japan</country>
</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Yamazaki, Toshimasa" sort="Yamazaki, Toshimasa" uniqKey="Yamazaki T" first="Toshimasa" last="Yamazaki">Toshimasa Yamazaki</name>
<affiliation><nlm:aff id="aff6">Biomolecular Research Unit,<institution>National Institute of Agrobiological Sciences</institution>
, Tsukuba 305-8602,<country>Japan</country>
</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Lu, Shixiang" sort="Lu, Shixiang" uniqKey="Lu S" first="Shixiang" last="Lü">Shixiang Lü</name>
<affiliation><nlm:aff id="aff1">Northeast Institute of Geography and Agroecology,<institution>Chinese Academy of Sciences</institution>
, Harbin 150081,<country>China</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Wu, Hongyan" sort="Wu, Hongyan" uniqKey="Wu H" first="Hongyan" last="Wu">Hongyan Wu</name>
<affiliation><nlm:aff id="aff1">Northeast Institute of Geography and Agroecology,<institution>Chinese Academy of Sciences</institution>
, Harbin 150081,<country>China</country>
;</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Tabata, Satoshi" sort="Tabata, Satoshi" uniqKey="Tabata S" first="Satoshi" last="Tabata">Satoshi Tabata</name>
<affiliation><nlm:aff wicri:cut="; and" id="aff5">Department of Plant Genome Research,<institution>Kazusa DNA Research Institute</institution>
, Kisarazu 292-0812,<country>Japan</country>
</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Harada, Kyuya" sort="Harada, Kyuya" uniqKey="Harada K" first="Kyuya" last="Harada">Kyuya Harada</name>
<affiliation><nlm:aff id="aff2">Soybean Applied Genomics Research Unit,<institution>National Institute of Agrobiological Sciences</institution>
, Tsukuba 305-8602,<country>Japan</country>
;</nlm:aff>
</affiliation>
</author>
</analytic>
<series><title level="j">Proceedings of the National Academy of Sciences of the United States of America</title>
<idno type="ISSN">0027-8424</idno>
<idno type="eISSN">1091-6490</idno>
<imprint><date when="2012">2012</date>
</imprint>
</series>
</biblStruct>
</sourceDesc>
</fileDesc>
<profileDesc><textClass></textClass>
</profileDesc>
</teiHeader>
<front><div type="abstract" xml:lang="en"><p>The complex and coordinated regulation of flowering has high ecological and agricultural significance. The maturity locus <italic>E1</italic>
has a large impact on flowering time in soybean, but the molecular basis for the <italic>E1</italic>
locus is largely unknown. Through positional cloning, we delimited the <italic>E1</italic>
locus to a 17.4-kb region containing an intron-free gene (<italic>E1</italic>
). The E1 protein contains a putative bipartite nuclear localization signal and a region distantly related to B3 domain. In the recessive allele, a nonsynonymous substitution occurred in the putative nuclear localization signal, leading to the loss of localization specificity of the E1 protein and earlier flowering. The early-flowering phenotype was consistently observed in three ethylmethanesulfonate-induced mutants and two natural mutations that harbored a premature stop codon or a deletion of the entire <italic>E1</italic>
gene. <italic>E1</italic>
expression was significantly suppressed under short-day conditions and showed a bimodal diurnal pattern under long-day conditions, suggesting its response to photoperiod and its dominant effect induced by long day length. When a functional <italic>E1</italic>
gene was transformed into the early-flowering cultivar Kariyutaka with low <italic>E1</italic>
expression, transgenic plants carrying exogenous <italic>E1</italic>
displayed late flowering. Furthermore, the transcript abundance of <italic>E1</italic>
was negatively correlated with that of <italic>GmFT2a</italic>
and <italic>GmFT5a</italic>
, homologues of <italic>FLOWERING LOCUS T</italic>
that promote flowering. These findings demonstrated the key role of <italic>E1</italic>
in repressing flowering and delaying maturity in soybean. The molecular identification of the maturity locus <italic>E1</italic>
will contribute to our understanding of the molecular mechanisms by which a short-day plant regulates flowering time and maturity.</p>
</div>
</front>
</TEI>
<pmc article-type="research-article"><pmc-comment>The publisher of this article does not allow downloading of the full text in XML form.</pmc-comment>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Proc Natl Acad Sci U S A</journal-id>
<journal-id journal-id-type="iso-abbrev">Proc. Natl. Acad. Sci. U.S.A</journal-id>
<journal-id journal-id-type="hwp">pnas</journal-id>
<journal-id journal-id-type="pmc">pnas</journal-id>
<journal-id journal-id-type="publisher-id">PNAS</journal-id>
<journal-title-group><journal-title>Proceedings of the National Academy of Sciences of the United States of America</journal-title>
</journal-title-group>
<issn pub-type="ppub">0027-8424</issn>
<issn pub-type="epub">1091-6490</issn>
<publisher><publisher-name>National Academy of Sciences</publisher-name>
</publisher>
</journal-meta>
<article-meta><article-id pub-id-type="pmid">22619331</article-id>
<article-id pub-id-type="pmc">3420212</article-id>
<article-id pub-id-type="publisher-id">201117982</article-id>
<article-id pub-id-type="doi">10.1073/pnas.1117982109</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>PNAS Plus</subject>
</subj-group>
<subj-group subj-group-type="heading"><subject>Biological Sciences</subject>
<subj-group><subject>Genetics</subject>
</subj-group>
</subj-group>
<series-title>PNAS Plus</series-title>
</article-categories>
<title-group><article-title>Positional cloning and characterization reveal the molecular basis for soybean maturity locus <italic>E1</italic>
that regulates photoperiodic flowering</article-title>
<alt-title alt-title-type="short">Molecular identification of the soybean <italic>E1</italic>
locus</alt-title>
</title-group>
<contrib-group><contrib contrib-type="author"><name><surname>Xia</surname>
<given-names>Zhengjun</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>a</sup>
</xref>
<xref ref-type="aff" rid="aff2"><sup>b</sup>
</xref>
<xref ref-type="corresp" rid="cor1"><sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Watanabe</surname>
<given-names>Satoshi</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>b</sup>
</xref>
<xref ref-type="corresp" rid="cor1"><sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Yamada</surname>
<given-names>Tetsuya</given-names>
</name>
<xref ref-type="aff" rid="aff3"><sup>c</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Tsubokura</surname>
<given-names>Yasutaka</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>b</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Nakashima</surname>
<given-names>Hiroko</given-names>
</name>
<xref ref-type="aff" rid="aff4"><sup>d</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Zhai</surname>
<given-names>Hong</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>a</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Anai</surname>
<given-names>Toyoaki</given-names>
</name>
<xref ref-type="aff" rid="aff4"><sup>d</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Sato</surname>
<given-names>Shusei</given-names>
</name>
<xref ref-type="aff" rid="aff5"><sup>e</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Yamazaki</surname>
<given-names>Toshimasa</given-names>
</name>
<xref ref-type="aff" rid="aff6"><sup>f</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Lü</surname>
<given-names>Shixiang</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>a</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Wu</surname>
<given-names>Hongyan</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>a</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Tabata</surname>
<given-names>Satoshi</given-names>
</name>
<xref ref-type="aff" rid="aff5"><sup>e</sup>
</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Harada</surname>
<given-names>Kyuya</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>b</sup>
</xref>
<xref ref-type="corresp" rid="cor1"><sup>1</sup>
</xref>
</contrib>
<aff id="aff1"><sup>a</sup>
Northeast Institute of Geography and Agroecology,<institution>Chinese Academy of Sciences</institution>
, Harbin 150081,<country>China</country>
;</aff>
<aff id="aff2"><sup>b</sup>
Soybean Applied Genomics Research Unit,<institution>National Institute of Agrobiological Sciences</institution>
, Tsukuba 305-8602,<country>Japan</country>
;</aff>
<aff id="aff3"><sup>c</sup>
Graduate School of Agriculture,<institution>Hokkaido University</institution>
, Sapporo 060-8589,<country>Japan</country>
;</aff>
<aff id="aff4"><sup>d</sup>
Faculty of Agriculture,<institution>Saga University</institution>
, Saga 840-8502,<country>Japan</country>
;</aff>
<aff id="aff5"><sup>e</sup>
Department of Plant Genome Research,<institution>Kazusa DNA Research Institute</institution>
, Kisarazu 292-0812,<country>Japan</country>
; and</aff>
<aff id="aff6"><sup>f</sup>
Biomolecular Research Unit,<institution>National Institute of Agrobiological Sciences</institution>
, Tsukuba 305-8602,<country>Japan</country>
</aff>
</contrib-group>
<author-notes><corresp id="cor1"><sup>1</sup>
To whom correspondence may be addressed. E-mail: <email>xiazhj@neigaehrb.ac.cn</email>
, <email>nabemame@affrc.go.jp</email>
, or <email>haradaq@nifty.com</email>
.</corresp>
<fn fn-type="edited-by"><p>Edited by Detlef Weigel, Max Planck Institute for Developmental Biology, Tübingen, Germany, and approved April 5, 2012 (received for review November 10, 2011)</p>
</fn>
<fn fn-type="con"><p>Author contributions: Z.X. and K.H. designed research; Z.X., S.W., T. Yamada, Y.T., H.N., H.Z., T.A., S.S., S.L., and H.W. performed research; T. Yamazaki and S.T. analyzed data; and Z.X., S.W., and K.H. wrote the paper.</p>
</fn>
</author-notes>
<pub-date pub-type="ppub"><day>7</day>
<month>8</month>
<year>2012</year>
</pub-date>
<pub-date pub-type="epub"><day>22</day>
<month>5</month>
<year>2012</year>
</pub-date>
<pub-date pub-type="pmc-release"><day>22</day>
<month>5</month>
<year>2012</year>
</pub-date>
<pmc-comment> PMC Release delay is 0 months and 0 days and was based on the
. </pmc-comment>
<volume>109</volume>
<issue>32</issue>
<fpage>E2155</fpage>
<lpage>E2164</lpage>
<page-range>E2155–E2164</page-range>
<permissions><license license-type="open-access"><license-p>Freely available online through the PNAS open access option.</license-p>
</license>
</permissions>
<self-uri content-type="pdf" xlink:type="simple" xlink:href="pnas.201117982.pdf"></self-uri>
<self-uri content-type="author-summary-pdf" xlink:type="simple" xlink:href="pnas.201117982_summary.pdf"></self-uri>
<abstract id="d34e291"><p>The complex and coordinated regulation of flowering has high ecological and agricultural significance. The maturity locus <italic>E1</italic>
has a large impact on flowering time in soybean, but the molecular basis for the <italic>E1</italic>
locus is largely unknown. Through positional cloning, we delimited the <italic>E1</italic>
locus to a 17.4-kb region containing an intron-free gene (<italic>E1</italic>
). The E1 protein contains a putative bipartite nuclear localization signal and a region distantly related to B3 domain. In the recessive allele, a nonsynonymous substitution occurred in the putative nuclear localization signal, leading to the loss of localization specificity of the E1 protein and earlier flowering. The early-flowering phenotype was consistently observed in three ethylmethanesulfonate-induced mutants and two natural mutations that harbored a premature stop codon or a deletion of the entire <italic>E1</italic>
gene. <italic>E1</italic>
expression was significantly suppressed under short-day conditions and showed a bimodal diurnal pattern under long-day conditions, suggesting its response to photoperiod and its dominant effect induced by long day length. When a functional <italic>E1</italic>
gene was transformed into the early-flowering cultivar Kariyutaka with low <italic>E1</italic>
expression, transgenic plants carrying exogenous <italic>E1</italic>
displayed late flowering. Furthermore, the transcript abundance of <italic>E1</italic>
was negatively correlated with that of <italic>GmFT2a</italic>
and <italic>GmFT5a</italic>
, homologues of <italic>FLOWERING LOCUS T</italic>
that promote flowering. These findings demonstrated the key role of <italic>E1</italic>
in repressing flowering and delaying maturity in soybean. The molecular identification of the maturity locus <italic>E1</italic>
will contribute to our understanding of the molecular mechanisms by which a short-day plant regulates flowering time and maturity.</p>
</abstract>
<kwd-group><kwd>photoperiodism</kwd>
<kwd>quantitative trait locus</kwd>
<kwd>photoperiodic insensibility</kwd>
</kwd-group>
<custom-meta-group><custom-meta><meta-name>author-summary</meta-name>
<meta-value><xref ref-type="other" rid="author-summary"></xref>
</meta-value>
</custom-meta>
</custom-meta-group>
</article-meta>
</front>
</pmc>
</record>
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