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<article article-type="research-article" dtd-version="1.3" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xml:lang="ru"><front><journal-meta><journal-id journal-id-type="publisher-id">vavilov</journal-id><journal-title-group><journal-title xml:lang="ru">Вавиловский журнал генетики и селекции</journal-title><trans-title-group xml:lang="en"><trans-title>Vavilov Journal of Genetics and Breeding</trans-title></trans-title-group></journal-title-group><issn pub-type="epub">2500-3259</issn><publisher><publisher-name>Institute of Cytology and Genetics of Siberian Branch of the RAS</publisher-name></publisher></journal-meta><article-meta><article-id custom-type="elpub" pub-id-type="custom">vavilov-28</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="ru"><subject>Статьи</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>Articles</subject></subj-group></article-categories><title-group><article-title>МОДЕЛЬ ГЕННОЙ СЕТИ РЕГУЛЯЦИИ ВРЕМЕНИ ЦВЕТЕНИЯ У ОЗИМОЙ ПШЕНИЦЫ И ЯЧМЕНЯ</article-title><trans-title-group xml:lang="en"><trans-title>A GENE REGULATORY NETWORK MODEL FOR VERNALIZATION AND SEASONAL FLOWERING RESPONSE IN WINTER WHEAT AND BARLEY</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Степаненко</surname><given-names>И. Л.</given-names></name><name name-style="western" xml:lang="en"><surname>Stepanenko</surname><given-names>I. L.</given-names></name></name-alternatives><email xlink:type="simple">stepan@bionet.nsc.ru</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Смирнова</surname><given-names>О. Г.</given-names></name><name name-style="western" xml:lang="en"><surname>Smirnova</surname><given-names>O. G.</given-names></name></name-alternatives><email xlink:type="simple">stepan@bionet.nsc.ru</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Титов</surname><given-names>И. И.</given-names></name><name name-style="western" xml:lang="en"><surname>Titov</surname><given-names>I. I.</given-names></name></name-alternatives><email xlink:type="simple">stepan@bionet.nsc.ru</email><xref ref-type="aff" rid="aff-1"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru">Федеральное государственное бюджетное научное учреждение «Федеральный исследовательский центр Институт цитологии и генетики Сибирского отделения Российской академии наук», Новосибирск, Россия<country>Россия</country></aff><aff xml:lang="en">Institute of Cytology and Genetics, SB RAS, Novosibirsk, Russia<country>Russian Federation</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2012</year></pub-date><pub-date pub-type="epub"><day>10</day><month>12</month><year>2014</year></pub-date><volume>16</volume><issue>1</issue><fpage>99</fpage><lpage>106</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Степаненко И.Л., Смирнова О.Г., Титов И.И., 2014</copyright-statement><copyright-year>2014</copyright-year><copyright-holder xml:lang="ru">Степаненко И.Л., Смирнова О.Г., Титов И.И.</copyright-holder><copyright-holder xml:lang="en">Stepanenko I.L., Smirnova O.G., Titov I.I.</copyright-holder><license license-type="creative-commons-attribution" xlink:href="https://creativecommons.org/licenses/by/4.0/" xlink:type="simple"><license-p>This work is licensed under a Creative Commons Attribution 4.0 License.</license-p></license></permissions><self-uri xlink:href="https://vavilov.elpub.ru/jour/article/view/28">https://vavilov.elpub.ru/jour/article/view/28</self-uri><abstract><p>Переход от вегетативной к репродуктивной стадии развития растений зависит от фотопериода и яровизации. Предложена простая логическая модель генной сети, регулирующей время цветения озимой пшеницы. У озимых сортов злаков чувствительность к яровизации контролируется генами VRN1, VRN2 и VRN3. После яровизации продукт гена VRN1 подавляет ген VRN2, кодирующий репрессор цветения. В результате уровень экспрессии VRN3 возрастает и стимулирует дальнейшее увеличение уровня транскрипции гена VRN1. Гены образуют петлю с положительной обратной связью,что усиливает транскрипцию VRN1 до уровня, необходимого для инициации цветения. В условиях длинного дня экспрессию VRN3 усиливают продукты генов PPD1 и CO2, определяющие чувствительность к фотопериоду. Сезонные изменения длины дня через фоторецепторы передаются циркадным часам, которые модулируют время цветения. Данные о генах ячменя и пшеницы, контролирующих чувствительность к яровизации и фотопериоду, интегрированы в генную сеть. С помощью синхронной булевой модели воспроизведена динамика генной сети. Расширенная модель генной сети может быть использована для проверки согласованности экспериментальных данных и выдвижения новых гипотез о взаимодействии генов.</p></abstract><trans-abstract xml:lang="en"><p>The transition from vegetative to reproductive development in wheat is regulated by seasonal cues, including vernalization and photoperiod. Here we present a simple logical model of the wheat development gene network. Vernalization accelerates flowering in winter cereals. It is regulated mainly by the vernalization genes VRN1, VRN2, and VRN3. After vernalization, VRN1 downregulates the VRN2 flowering repressor, thereby increasing the VRN3 level. The expression of VRN3 promotes further increases in the VRN1 transcription level, generating a positive feedback loop, which enhances VRN1 transcription to a threshold level required to initiate flowering. The products of the PPD1 and CO2 photoperiod genes increase VRN3 expression under long daylight conditions. Seasonal changes in day length are perceived by plant photoreceptors and transmitted to the circadian clock to modulate flowering time. Here we integrate data on vernalization and photoperiod genes in a gene network. Using a synchronous Boolean model, we have simulated the network dynamics. This model can be useful to test the coherence of experimental data and to hypothesize gene interactions that remain to be discovered.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>пшеница</kwd><kwd>ячмень</kwd><kwd>VRN1</kwd><kwd>VRN2</kwd><kwd>VRN3</kwd><kwd>яровизация</kwd><kwd>фотопериод</kwd><kwd>длина дня</kwd><kwd>циркадный ритм</kwd><kwd>цветение</kwd><kwd>генная сеть</kwd><kwd>синхронная булева модель</kwd></kwd-group><kwd-group xml:lang="en"><kwd>wheat</kwd><kwd>barley</kwd><kwd>VRN1</kwd><kwd>VRN2</kwd><kwd>VRN3</kwd><kwd>vernalization</kwd><kwd>photoperiod</kwd><kwd>day length</kwd><kwd>circadian rhythm</kwd><kwd>flowering</kwd><kwd>gene network</kwd><kwd>synchronous Boolean model</kwd></kwd-group><funding-group xml:lang="ru"><funding-statement>Минобрнауки, СО РАН</funding-statement></funding-group></article-meta></front><back><ref-list><title>References</title><ref id="cit1"><label>1</label><citation-alternatives><mixed-citation xml:lang="ru">Ananko E.A., Podkolodny N.L., Stepanenko I.L. et al. GeneNet in 2005 // Nucl. Acids Res. 2005. V. 33. D425–427.</mixed-citation><mixed-citation xml:lang="en">Ananko E.A., Podkolodny N.L., Stepanenko I.L. et al. GeneNet in 2005 // Nucl. Acids Res. 2005. V. 33. D425–427.</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Casao M.C., Igartua E., Karsai I. et al. Expression analysis of vernalization and day-length response genes in barley (Hordeum vulgare L.) indicates that VRNH2 is a repressor of PPDH2 (HvFT3) under long days // J. Exp. Bot. 2011. V. 62. P. 1939–1949.</mixed-citation><mixed-citation xml:lang="en">Casao M.C., Igartua E., Karsai I. et al. Expression analysis of vernalization and day-length response genes in barley (Hordeum vulgare L.) indicates that VRNH2 is a repressor of PPDH2 (HvFT3) under long days // J. Exp. Bot. 2011. V. 62. P. 1939–1949.</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Corbesier L., Vincent C., Jang S.H. et al. FT protein movement contributes to long-distance signaling in fl oral induction of Arabidopsis // Sci. 2007. V. 316. P. 1030–1033.</mixed-citation><mixed-citation xml:lang="en">Corbesier L., Vincent C., Jang S.H. et al. FT protein movement contributes to long-distance signaling in fl oral induction of Arabidopsis // Sci. 2007. V. 316. P. 1030–1033.</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Davidich M., Bornholdt S. Boolean network model predicts cell cycle sequence of fi ssion yeast // PLoS ONE. 2008. V. 3. P. e1672.</mixed-citation><mixed-citation xml:lang="en">Davidich M., Bornholdt S. Boolean network model predicts cell cycle sequence of fi ssion yeast // PLoS ONE. 2008. V. 3. P. e1672.</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Distelfeld A., Li C., Dubcovsky J. Regulation of fl owering in temperate cereals // Curr. Opin. Plant Biol. 2009. V. 12. P. 178–184.</mixed-citation><mixed-citation xml:lang="en">Distelfeld A., Li C., Dubcovsky J. Regulation of fl owering in temperate cereals // Curr. Opin. Plant Biol. 2009. V. 12. P. 178–184.</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Distelfeld A., Dubcovsky J. Characterization of the maintained vegetative phase deletions from diploid wheat and their effect on VRN2 and FT transcript levels // Mol. Genet. Genomics. 2010. V. 283. P. 223–232.</mixed-citation><mixed-citation xml:lang="en">Distelfeld A., Dubcovsky J. Characterization of the maintained vegetative phase deletions from diploid wheat and their effect on VRN2 and FT transcript levels // Mol. Genet. Genomics. 2010. V. 283. P. 223–232.</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Greenup A., Peacock W.J., Dennis E.S., Trevaskis B. The molecular biology of seasonal fl owering-responses in Arabidopsis and the cereals // Ann. Bot. 2009. V. 103. P. 1165–1172.</mixed-citation><mixed-citation xml:lang="en">Greenup A., Peacock W.J., Dennis E.S., Trevaskis B. The molecular biology of seasonal fl owering-responses in Arabidopsis and the cereals // Ann. Bot. 2009. V. 103. P. 1165–1172.</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Greenup A.G., Sasani S., Oliver S.N. et al. ODDSOC2 is a MADS box fl oral repressor that is down-regulated by vernalization in temperate cereals // Plant Physiol. 2010. V. 153. P. 1062–1073.</mixed-citation><mixed-citation xml:lang="en">Greenup A.G., Sasani S., Oliver S.N. et al. ODDSOC2 is a MADS box fl oral repressor that is down-regulated by vernalization in temperate cereals // Plant Physiol. 2010. V. 153. P. 1062–1073.</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Greenup A.G., Sasani S., Oliver S.N. et al. Transcriptome analysis of the vernalization response in barley (Hordeum vulgare) seedlings // PLoS One. 2011. V. 6. P. e17900.</mixed-citation><mixed-citation xml:lang="en">Greenup A.G., Sasani S., Oliver S.N. et al. Transcriptome analysis of the vernalization response in barley (Hordeum vulgare) seedlings // PLoS One. 2011. V. 6. P. e17900.</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Hemming M.N., Peacock W.J., Dennis, E.S., Trevaskis B. Low-temperature and daylength cues are integrated to regulate FLOWERING LOCUS T in barley // Plant Physiol. 2008. V. 147. P. 355–366.</mixed-citation><mixed-citation xml:lang="en">Hemming M.N., Peacock W.J., Dennis, E.S., Trevaskis B. Low-temperature and daylength cues are integrated to regulate FLOWERING LOCUS T in barley // Plant Physiol. 2008. V. 147. P. 355–366.</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Higgins J.A., Bailey P.C., Laurie D.A. Comparative genomics of flowering time pathways using Brachypodium distachyon as a model for the temperate grasses // PLoS One. 2010. V. 19. P. e10065.</mixed-citation><mixed-citation xml:lang="en">Higgins J.A., Bailey P.C., Laurie D.A. Comparative genomics of flowering time pathways using Brachypodium distachyon as a model for the temperate grasses // PLoS One. 2010. V. 19. P. e10065.</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Imaizumi T. Arabidopsis circadian clock and photoperiodism: Time to think about location // Curr. Opin. Plant Biol. 2010. V. 13. P. 83–89.</mixed-citation><mixed-citation xml:lang="en">Imaizumi T. Arabidopsis circadian clock and photoperiodism: Time to think about location // Curr. Opin. Plant Biol. 2010. V. 13. P. 83–89.</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Kauffman S., Peterson C., Samuelsson B. et al. Genetic networks with canalyzing Boolean rules are always stable // Proc. Natl Acad. Sci. USA. 2004. V. 101. P. 17102–17107.</mixed-citation><mixed-citation xml:lang="en">Kauffman S., Peterson C., Samuelsson B. et al. Genetic networks with canalyzing Boolean rules are always stable // Proc. Natl Acad. Sci. USA. 2004. V. 101. P. 17102–17107.</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Kane N.A., Agharbaoui Z., Diallo A.O. et al. TaVRT2 represses transcription of the wheat vernalization gene TaVRN1 // Plant J. 2007. V. 51. P. 670–680.</mixed-citation><mixed-citation xml:lang="en">Kane N.A., Agharbaoui Z., Diallo A.O. et al. TaVRT2 represses transcription of the wheat vernalization gene TaVRN1 // Plant J. 2007. V. 51. P. 670–680.</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Li C., Distelfeld A., Comis A., Dubcovsky J. Wheat fl owering repressor VRN2 and promoter CO2 compete for interactions with NUCLEAR FACTOR-Y complexes // Plant J. 2011. V. 67. P. 763–773.</mixed-citation><mixed-citation xml:lang="en">Li C., Distelfeld A., Comis A., Dubcovsky J. Wheat fl owering repressor VRN2 and promoter CO2 compete for interactions with NUCLEAR FACTOR-Y complexes // Plant J. 2011. V. 67. P. 763–773.</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Li C., Dubcovsky J. Wheat FT protein regulates VRN1 transcription through interactions with FDL2 // Plant J. 2008. V. 55. P. 543–554.</mixed-citation><mixed-citation xml:lang="en">Li C., Dubcovsky J. Wheat FT protein regulates VRN1 transcription through interactions with FDL2 // Plant J. 2008. V. 55. P. 543–554.</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Mendoza L., Thieffry D., Alvarez-Buylla E. Genetic control of fl ower morphogenesis in Arabidopsis thaliana: а logical analysis // Bioinformatics. 1999. V. 15. P. 593–606.</mixed-citation><mixed-citation xml:lang="en">Mendoza L., Thieffry D., Alvarez-Buylla E. Genetic control of fl ower morphogenesis in Arabidopsis thaliana: а logical analysis // Bioinformatics. 1999. V. 15. P. 593–606.</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Oliver S.N., Finnegan E.J., Dennis E.S. et al. Vernalizationinduced fl owering in cereals is associated with changes in histone methylation at the VERNALIZATION1 gene // Proc. Natl Acad. Sci. USA. 2009. V. 106. P. 8386–8391.</mixed-citation><mixed-citation xml:lang="en">Oliver S.N., Finnegan E.J., Dennis E.S. et al. Vernalizationinduced fl owering in cereals is associated with changes in histone methylation at the VERNALIZATION1 gene // Proc. Natl Acad. Sci. USA. 2009. V. 106. P. 8386–8391.</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Papin J.A., Hunter T., Palsson B.O., Subramaniam S. Reconstruction of cellular signalling networks and analysis of their properties // Nat. Rev. Mol. Cell Biol. 2005. V. 6. P. 99–111.</mixed-citation><mixed-citation xml:lang="en">Papin J.A., Hunter T., Palsson B.O., Subramaniam S. Reconstruction of cellular signalling networks and analysis of their properties // Nat. Rev. Mol. Cell Biol. 2005. V. 6. P. 99–111.</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Sawa M., Nusinow D.A., Kay S.A., Imaizumi T. FKF1 and GIGANTEa complex formation is required for daylength measurement in Arabidopsis // Sci. 2007. V. 318. P. 261–265.</mixed-citation><mixed-citation xml:lang="en">Sawa M., Nusinow D.A., Kay S.A., Imaizumi T. FKF1 and GIGANTEa complex formation is required for daylength measurement in Arabidopsis // Sci. 2007. V. 318. P. 261–265.</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Shimada S., Ogawa T., Kitagawa S. et al. A genetic network of fl owering-time genes in wheat leaves, in which an APETALA1/FRUITFULL-like gene, VRN1, is upstream of FLOWERING LOCUS T // Plant J. 2009. V. 58. P. 668–681.</mixed-citation><mixed-citation xml:lang="en">Shimada S., Ogawa T., Kitagawa S. et al. A genetic network of fl owering-time genes in wheat leaves, in which an APETALA1/FRUITFULL-like gene, VRN1, is upstream of FLOWERING LOCUS T // Plant J. 2009. V. 58. P. 668–681.</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Smirnova O.G., Stepanenko I.L., Shumny V.K. The role of the COP1, SPA, and PIF proteins in plant photomorphogenesis // Biol. Bull. Rev. 2011. V. 1. Nо 4. P. 314–324.</mixed-citation><mixed-citation xml:lang="en">Smirnova O.G., Stepanenko I.L., Shumny V.K. The role of the COP1, SPA, and PIF proteins in plant photomorphogenesis // Biol. Bull. Rev. 2011. V. 1. Nо 4. P. 314–324.</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Song Y.H., Ito S., Imaizumi T. Similarities in the circadian clock and photoperiodism in plants // Curr. Opin. Plant Biol. 2010. V. 13. P. 594–603.</mixed-citation><mixed-citation xml:lang="en">Song Y.H., Ito S., Imaizumi T. Similarities in the circadian clock and photoperiodism in plants // Curr. Opin. Plant Biol. 2010. V. 13. P. 594–603.</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Stephenson T.J., McIntyre C.L., Collet C. et al. Genome-wide identifi cation and expression analysis of the NF-Y family of transcription factors in Triticum aestivum // Plant Mol. Biol. 2007. V. 65 P. 77–92.</mixed-citation><mixed-citation xml:lang="en">Stephenson T.J., McIntyre C.L., Collet C. et al. Genome-wide identifi cation and expression analysis of the NF-Y family of transcription factors in Triticum aestivum // Plant Mol. Biol. 2007. V. 65 P. 77–92.</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Tiwari S.B., Shen Y., Chang H.C. et al. The fl owering time regulator CONSTANS is recruited to the FLOWERING LOCUS T promoter via a unique cis-element // New Phytol. 2010. V. 187. P. 57–66.</mixed-citation><mixed-citation xml:lang="en">Tiwari S.B., Shen Y., Chang H.C. et al. The fl owering time regulator CONSTANS is recruited to the FLOWERING LOCUS T promoter via a unique cis-element // New Phytol. 2010. V. 187. P. 57–66.</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Trevaskis B., Hemming M.N., Dennis E.S. et al. The molecular basis of vernalization-induced fl owering in cereals // Trends Plant Sci. 2007. V. 12. P. 352–357.</mixed-citation><mixed-citation xml:lang="en">Trevaskis B., Hemming M.N., Dennis E.S. et al. The molecular basis of vernalization-induced fl owering in cereals // Trends Plant Sci. 2007. V. 12. P. 352–357.</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Trevaskis B., Hemming M.N., Peacock W.J., Dennis E.S. HvVRN2 responds to daylength, whereas HvVRN1 is regulated by vernalization and developmental status // Plant Physiol. 2006. V. 140. P. 1397–1405.</mixed-citation><mixed-citation xml:lang="en">Trevaskis B., Hemming M.N., Peacock W.J., Dennis E.S. HvVRN2 responds to daylength, whereas HvVRN1 is regulated by vernalization and developmental status // Plant Physiol. 2006. V. 140. P. 1397–1405.</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Turner A., Beales J., Faure S. et al. The pseudo-response regulator Ppd-H1 provides adaptation to photoperiod in barley // Sci. 2005. V. 310. P. 1031–1034.</mixed-citation><mixed-citation xml:lang="en">Turner A., Beales J., Faure S. et al. The pseudo-response regulator Ppd-H1 provides adaptation to photoperiod in barley // Sci. 2005. V. 310. P. 1031–1034.</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Yan L., Fu D., Li C., Blechl A. et al. The wheat and barley vernalization gene VRN3 is an orthologue of FT // Proc. Natl Acad. Sci. USA. 2006. V. 103. P. 19581–19586.</mixed-citation><mixed-citation xml:lang="en">Yan L., Fu D., Li C., Blechl A. et al. The wheat and barley vernalization gene VRN3 is an orthologue of FT // Proc. Natl Acad. Sci. USA. 2006. V. 103. P. 19581–19586.</mixed-citation></citation-alternatives></ref><ref id="cit30"><label>30</label><citation-alternatives><mixed-citation xml:lang="ru">Yan L., Loukoianov A., Blechl A. et al. The wheat VRN2 gene is a fl owering repressor down-regulated by vernalization // Sci. 2004. V. 303. Nо 5664. P. 1640–1644.</mixed-citation><mixed-citation xml:lang="en">Yan L., Loukoianov A., Blechl A. et al. The wheat VRN2 gene is a fl owering repressor down-regulated by vernalization // Sci. 2004. V. 303. Nо 5664. P. 1640–1644.</mixed-citation></citation-alternatives></ref><ref id="cit31"><label>31</label><citation-alternatives><mixed-citation xml:lang="ru">Yan L., Loukoianov A., Tranquilli G. et al. Positional cloning of wheat vernalization gene VRN1 // Proc. Natl Acad. Sci. USA. 2003. V. 100. P. 6263–6268.</mixed-citation><mixed-citation xml:lang="en">Yan L., Loukoianov A., Tranquilli G. et al. Positional cloning of wheat vernalization gene VRN1 // Proc. Natl Acad. Sci. USA. 2003. V. 100. P. 6263–6268.</mixed-citation></citation-alternatives></ref><ref id="cit32"><label>32</label><citation-alternatives><mixed-citation xml:lang="ru">Yu J.W., Rubio V., Lee N.Y. et al. COP1 and ELF3 control circadian function and photoperiodic flowering by regulating GI stability // Mol. Cell. 2008. V. 32. P. 617–630.</mixed-citation><mixed-citation xml:lang="en">Yu J.W., Rubio V., Lee N.Y. et al. COP1 and ELF3 control circadian function and photoperiodic flowering by regulating GI stability // Mol. Cell. 2008. V. 32. P. 617–630.</mixed-citation></citation-alternatives></ref><ref id="cit33"><label>33</label><citation-alternatives><mixed-citation xml:lang="ru">Zhao X.Y., Liu M.S., Li J.R. et al. The wheat TaGI1, involved in photoperiodic fl owering, encodes an Arabidopsis GI ortholog // Plant Mol. Biol. 2005. V. 58. P. 53–64.</mixed-citation><mixed-citation xml:lang="en">Zhao X.Y., Liu M.S., Li J.R. et al. The wheat TaGI1, involved in photoperiodic fl owering, encodes an Arabidopsis GI ortholog // Plant Mol. Biol. 2005. V. 58. P. 53–64.</mixed-citation></citation-alternatives></ref></ref-list><fn-group><fn fn-type="conflict"><p>The authors declare that there are no conflicts of interest present.</p></fn></fn-group></back></article>
