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<article 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" xmlns:ali="http://www.niso.org/schemas/ali/1.0/" article-type="research-article" dtd-version="1.2" xml:lang="en"><front><journal-meta><journal-id journal-id-type="publisher-id">Russian Journal of Genetics</journal-id><journal-title-group><journal-title xml:lang="en">Russian Journal of Genetics</journal-title><trans-title-group xml:lang="ru"><trans-title>Генетика</trans-title></trans-title-group></journal-title-group><issn publication-format="print">0016-6758</issn><issn publication-format="electronic">3034-5103</issn><publisher><publisher-name xml:lang="en">The Russian Academy of Sciences</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="publisher-id">686162</article-id><article-id pub-id-type="doi">10.31857/S0016675825010016</article-id><article-id pub-id-type="edn">VFMYSM</article-id><article-categories><subj-group subj-group-type="toc-heading"><subject>ОБЗОРНЫЕ И ТЕОРЕТИЧЕСКИЕ СТАТЬИ</subject></subj-group><subj-group subj-group-type="article-type"><subject>Research Article</subject></subj-group></article-categories><title-group><article-title xml:lang="en">The role of long non-coding RNAs in plants</article-title><trans-title-group xml:lang="ru"><trans-title>Роль длинных некодирующих РНК в растениях</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author"><name-alternatives><name xml:lang="en"><surname>Pronozin</surname><given-names>А. Y.</given-names></name><name xml:lang="ru"><surname>Пронозин</surname><given-names>А. Ю.</given-names></name></name-alternatives><address><country country="RU">Russian Federation</country></address><bio xml:lang="en"><p>Kurchatov Genomic Center</p></bio><bio xml:lang="ru"><p>Курчатовский геномный центр</p></bio><email>pronozinartem95@gmail.com</email><xref ref-type="aff" rid="aff1"/><xref ref-type="aff" rid="aff2"/></contrib><contrib contrib-type="author"><name-alternatives><name xml:lang="en"><surname>Afonnikov</surname><given-names>D. A.</given-names></name><name xml:lang="ru"><surname>Афонников</surname><given-names>Д. А.</given-names></name></name-alternatives><address><country country="RU">Russian Federation</country></address><bio xml:lang="en"><p>Kurchatov Genomic Center</p></bio><bio xml:lang="ru"><p>Курчатовский геномный центр</p></bio><email>pronozinartem95@gmail.com</email><xref ref-type="aff" rid="aff1"/><xref ref-type="aff" rid="aff2"/><xref ref-type="aff" rid="aff3"/></contrib></contrib-group><aff-alternatives id="aff1"><aff><institution xml:lang="en">Institute of Cytology and Genetics, Siberian Branch of the Russian Academy of Sciences</institution></aff><aff><institution xml:lang="ru">Федеральный исследовательский центр Институт цитологии и генетики Сибирского отделения Российской академии наук</institution></aff></aff-alternatives><aff-alternatives id="aff2"><aff><institution xml:lang="en">Institute of Cytology and Genetics, Siberian Branch, Russian Academy of Sciences</institution></aff><aff><institution xml:lang="ru">Институт цитологии и генетики Сибирского отделения Российской академии наук</institution></aff></aff-alternatives><aff-alternatives id="aff3"><aff><institution xml:lang="en">Novosibirsk National Research State University</institution></aff><aff><institution xml:lang="ru">Новосибирский национальный исследовательский государственный университет</institution></aff></aff-alternatives><pub-date date-type="pub" iso-8601-date="2025-01-15" publication-format="electronic"><day>15</day><month>01</month><year>2025</year></pub-date><volume>61</volume><issue>1</issue><fpage>3</fpage><lpage>23</lpage><history><date date-type="received" iso-8601-date="2025-06-27"><day>27</day><month>06</month><year>2025</year></date></history><permissions><copyright-statement xml:lang="en">Copyright ©; 2025, Russian Academy of Sciences</copyright-statement><copyright-statement xml:lang="ru">Copyright ©; 2025, Российская академия наук</copyright-statement><copyright-year>2025</copyright-year><copyright-holder xml:lang="en">Russian Academy of Sciences</copyright-holder><copyright-holder xml:lang="ru">Российская академия наук</copyright-holder></permissions><self-uri xlink:href="https://transsyst.ru/0016-6758/article/view/686162">https://transsyst.ru/0016-6758/article/view/686162</self-uri><abstract xml:lang="en"><p>Long non-coding RNAs (lncRNAs) are a class of linear or circular RNA molecules longer than 200 nucleotides without open reading frames. Experimental studies have shown the involvement of lncRNAs in the regulation of resistance to cold, salt, and heat stress, and in fruit, root, and leaf development. However, experimental methods are labor-intensive and costly approaches and cannot yet be used for genome-wide mass studies of lncRNAs. For this purpose, bioinformatic approaches that aim at large-scale recognition of lncRNA sequences in genomes and transcriptomes have been applied. However, despite the growing number of studies devoted to the structural and functional analysis of lncRNAs, this type of molecule remains poorly understood. This is due to the many factors that need to be considered when identifying lncRNAs. The use of pan-genomes and pan-transcriptomes will improve the efficiency of the study and the total number of predicted lncRNAs compared to using the genome of a single species representative. This review focuses on describing the molecular and biological functions of lncRNAs, experimental and bioinformatic methods of identification, patterns of evolution, detection and analysis of lncRNAs at the scale of pan-genomes and pan-transcriptomes.</p></abstract><trans-abstract xml:lang="ru"><p>Длинные некодирующие РНК (днРНК) представляют собой класс линейных или кольцевых молекул РНК длиной более 200 нуклеотидов без открытых рамок считывания. Экспериментальные исследования показали участие днРНК в регуляции устойчивости к холодовому, солевому, тепловому стрессу, в развитии плодов, корня и листьев. Однако экспериментальные методы являются трудоемкими и дорогостоящими подходами и пока еще не могут быть использованы для массового изучения днРНК в масштабах всего генома. Для этих целей применяют биоинформатические подходы, которые направлены на широкомасштабное распознавание последовательностей днРНК в геномах и транскриптомах. Несмотря на растущее число исследований, посвященных структурно-функциональному анализу днРНК, данный тип молекул по-прежнему остается малоизученным. Это связано со множеством факторов, которые нужно учитывать при идентификации днРНК. Применение пан-геномов и пан-транскриптомов позволит повысить эффективность исследования и общее количество предсказанных днРНК по сравнению с использованием генома одного представителя вида. Настоящий обзор посвящен описанию молекулярных и биологических функций днРНК, экспериментальных и биоинформатических методов идентификации, а также закономерностям эволюции, выявлению и анализу днРНК в масштабах пан-геномов и пан-транскриптомов.</p></trans-abstract><kwd-group xml:lang="en"><kwd>long non-coding RNAs</kwd><kwd>microRNAs</kwd><kwd>transcription regulation</kwd><kwd>pan-genome</kwd></kwd-group><kwd-group xml:lang="ru"><kwd>длинные некодирующие РНК</kwd><kwd>микроРНК</kwd><kwd>регуляция транскрипции</kwd><kwd>пан-геном</kwd></kwd-group><funding-group/></article-meta></front><body></body><back><ref-list><ref id="B1"><label>1.</label><mixed-citation>Deng P., Liu S., Nie X., Wu L. Conservation analysis of long non-coding RNAs in plants // Sci. China Life Sci. 2018. V. 61. P. 190–198.</mixed-citation></ref><ref id="B2"><label>2.</label><mixed-citation>Wu H.-J., Wang Z-M., Wang M., Wang X-J. Widespread long noncoding RNAs as endogenous target mimics for microRNAs in plants // Plant Physiol. 2013. V. 161. № 4. P. 1875–1884.</mixed-citation></ref><ref id="B3"><label>3.</label><mixed-citation>Zhu Q.-H., Wang M.-B. Molecular functions of long non-coding RNAs in plants // Genes. 2012. V. 3. № 1. P. 176–190.</mixed-citation></ref><ref id="B4"><label>4.</label><mixed-citation>Назипова Н.Н. Разнообразие некодирующих РНК в геномах эукариот // Матем. биол. и биоинформат. 2021. Т. 16. № 2. С. 256–298.</mixed-citation></ref><ref id="B5"><label>5.</label><mixed-citation>Joshi A., Romanowska J. Recent advances in computational-based approaches in epigenetics studies // Epigenetics Methods. 2020. P. 569–590.</mixed-citation></ref><ref id="B6"><label>6.</label><mixed-citation>Kim E.-D., Sung S. Long noncoding RNA: Unveiling hidden layer of gene regulatory networks // Trends Plant Sci. 2012. V. 17. № 1. P. 16–21.</mixed-citation></ref><ref id="B7"><label>7.</label><mixed-citation>Karlik E., Ari S., Gozukirmizi N. LncRNAs: Genetic and epigenetic effects in plants // Biotechnol. Biotechnol. 2019. V. 33. № 1. P. 429–439. https://doi.org/10.1080/13102818.2019.1581085</mixed-citation></ref><ref id="B8"><label>8.</label><mixed-citation>Tsai M.-C., Manor O., Wan Y. et al. Long noncoding RNA as modular scaffold of histone modification complexes // Science. 2010. V. 329. № 5992. P. 689–693. https://doi.org/10.1126/science.1192002</mixed-citation></ref><ref id="B9"><label>9.</label><mixed-citation>Sousa C., Johansson C., Charonet C. et al. Translational and structural requirements of the early nodulin gene enod40 , a short-open reading frame-containing RNA, for elicitation of a cell-specific growth response in the alfalfa root cortex // Mol. Cell. Biol. 2001. V. 21. № 1. P. 354–366. https://doi.org/10.1128/MCB.21.1.354-366.2001</mixed-citation></ref><ref id="B10"><label>10.</label><mixed-citation>Medvedeva Y.A., Lennartsson A., Ehsani R. et al. EpiFactors: A comprehensive database of human epigenetic factors and complexes // Database. 2015. V. 2015. P. bav067.</mixed-citation></ref><ref id="B11"><label>11.</label><mixed-citation>Frankish A., Diekhans M., Jungreis I. et al. GENCODE 2021 // Nucl. Acids Res. 2021. V. 49. № D1. P. D916–D923.</mixed-citation></ref><ref id="B12"><label>12.</label><mixed-citation>Гордиюк В.В. Длинные некодирующие РНК – камертон в регуляции клеточных процессов // Ukr. Biochem. J. 2014. V. 86. № 2. С. 5–15.</mixed-citation></ref><ref id="B13"><label>13.</label><mixed-citation>Zhao X., Li J., Lian B. et al. Global identification of Arabidopsis lncRNAs reveals the regulation of MAF4 by a natural antisense RNA // Nat. Commun. 2018. V. 9. № 1. P. 5056.</mixed-citation></ref><ref id="B14"><label>14.</label><mixed-citation>Li X., Wu Z., Fu X. et al. lncRNAs: Insights into their function and mechanics in underlying disorders // Mutat. Res. Mutat. Res. 2014. V. 762. P. 1–21.</mixed-citation></ref><ref id="B15"><label>15.</label><mixed-citation>Ahmad P., Bensaoud C., Mekki I. et al. Long non-coding RNAs and their potential roles in the vector–host–pathogen triad // Life. MDPI. 2021. V. 11. № 1. P. 56.</mixed-citation></ref><ref id="B16"><label>16.</label><mixed-citation>De Quattro C., Pè M.E., Bertolini E. Long noncoding RNAs in the model species Brachypodium distachyon // Sci. Rep. 2017. V. 7. № 1. P. 11252.</mixed-citation></ref><ref id="B17"><label>17.</label><mixed-citation>Ma L., Bajic V.B., Zhang Z. On the classification of long non-coding RNAs // RNA Biol. 2013. V. 10. № 6. P. 924–933. https://doi.org/10.4161/rna.24604</mixed-citation></ref><ref id="B18"><label>18.</label><mixed-citation>Chen L., Zhu Q.-H., Kaufmann K. Long non-coding RNAs in plants: Emerging modulators of gene activity in development and stress responses // Planta. 2020. V. 252. № 5. P. 92. https://doi.org/10.1007/s00425-020-03480-5</mixed-citation></ref><ref id="B19"><label>19.</label><mixed-citation>Griffiths-Jones S. miRBase: The microRNA sequence database // MicroRNA Protocols. 2006. V. 342. P. 129–138.</mixed-citation></ref><ref id="B20"><label>20.</label><mixed-citation>Amaral P.P., Mattick J.S. Noncoding RNA in development // Mamm. Genome. 2008. V. 19. P. 454–492.</mixed-citation></ref><ref id="B21"><label>21.</label><mixed-citation>Бейлерли О.А., Гареев И.Ф. Длинные некодирующие РНК: какие перспективы? // Профилактическая медицина. 2020. Т. 23. № 2. С. 124–128.</mixed-citation></ref><ref id="B22"><label>22.</label><mixed-citation>Blythe A.J., Fox A.H., Bond C.S. The ins and outs of lncRNA structure: How, why and what comes next? // Biochim. Biophys. Acta BBA-Gene Regul. Mech. 2016. V. 1859. № 1. P. 46–58.</mixed-citation></ref><ref id="B23"><label>23.</label><mixed-citation>Bryzghalov O., Makałowska I., Szcześniak M.W. lncEvo: Automated identification and conservation study of long noncoding RNAs // BMC Bioinformatics. 2021. V. 22. № 1. P. 59. https://doi.org/10.1186/s12859-021-03991-2</mixed-citation></ref><ref id="B24"><label>24.</label><mixed-citation>Zhao Q., Sun Y., Wang D. et al. LncPipe: A nextflow-based pipeline for identification and analysis of long non-coding RNAs from RNA-Seq data // J. Genet. Genomics. 2018. V. 45. https://doi.org/10.1016/j.jgg.2018.06.005</mixed-citation></ref><ref id="B25"><label>25.</label><mixed-citation>Talyan S., Filipów S., Ignarski M. et al. CALINCA – a novel pipeline for the identification of lncRNAs in podocyte disease // Cells. MDPI. 2021. V. 10. № 3. P. 692.</mixed-citation></ref><ref id="B26"><label>26.</label><mixed-citation>Campalans A., Kondorosi A., Crespi M. Enod40, a short open reading frame – containing mRNA, induces cytoplasmic localization of a nuclear RNA binding protein in Medicago truncatula // Plant Cell. 2004. V. 16. № 4. P. 1047–1059.</mixed-citation></ref><ref id="B27"><label>27.</label><mixed-citation>Unver T., Tombuloglu H. Barley long non-coding RNAs (lncRNA) responsive to excess boron // Genomics. 2020. V. 112. № 2. P. 1947–1955.</mixed-citation></ref><ref id="B28"><label>28.</label><mixed-citation>Khorkova O., Hsiao J., Wahlestedt C. Basic biology and therapeutic implications of lncRNA // Adv. Drug Deliv. Rev. 2015. V. 87. P. 15–24.</mixed-citation></ref><ref id="B29"><label>29.</label><mixed-citation>Duret L., Chureau С., Samain S. et al. The Xist RNA Gene evolved in eutherians by pseudogenization of a protein-coding gene // Science. 2006. V. 312. № 5780. P. 1653–1655. https://doi.org/10.1126/science.1126316</mixed-citation></ref><ref id="B30"><label>30.</label><mixed-citation>Graf J., Kretz M. From structure to function: Route to understanding lncRNA mechanism // BioEssays. 2020. V. 42. № 12. https://doi.org/10.1002/bies.202000027</mixed-citation></ref><ref id="B31"><label>31.</label><mixed-citation>Golicz A.A., Singh M.B., Bhalla P.L. The long intergenic noncoding RNA (LincRNA) landscape of the soybean genome // Plant Physiol. 2018. V. 176. № 3. P. 2133–2147.</mixed-citation></ref><ref id="B32"><label>32.</label><mixed-citation>Cheng F., Wu J., Fang L., Wang X. Syntenic gene analysis between Brassica rapa and other Brassicaceae species // Front. Plant Sci. Frontiers. 2012. V. 3. P. 30895.</mixed-citation></ref><ref id="B33"><label>33.</label><mixed-citation>Huang L., Dong H., Zhou D. et al. Systematic identification of long non-coding RNAs during pollen development and fertilization in Brassica rapa // Plant J. 2018. V. 96. № 1. P. 203–222. https://doi.org/10.1111/tpj.14016</mixed-citation></ref><ref id="B34"><label>34.</label><mixed-citation>Derrien T., Johnson R., Bussotti G. et al. The GENCODE v7 catalog of human long noncoding RNAs: Analysis of their gene structure, evolution, and expression // Genome Res. 2012. V. 22. № 9. P. 1775–1789.</mixed-citation></ref><ref id="B35"><label>35.</label><mixed-citation>Ponting C.P., Oliver P.L., Reik W. Evolution and functions of long noncoding RNAs // Cell. 2009. V. 136. № 4. P. 629–641.</mixed-citation></ref><ref id="B36"><label>36.</label><mixed-citation>Golicz A.A., Bayer P.E., Barker G.C. et al. The pangenome of an agronomically important crop plant Brassica oleracea // Nat. Commun. 2016. V. 7. № 1. P. 13390.</mixed-citation></ref><ref id="B37"><label>37.</label><mixed-citation>Meile L., Croll D., Brunner P.C. et al. A fungal avirulence factor encoded in a highly plastic genomic region triggers partial resistance to septoria tritici blotch // New Phytol. 2018. V. 219. № 3. P. 1048–1061. https://doi.org/10.1111/nph.15180</mixed-citation></ref><ref id="B38"><label>38.</label><mixed-citation>Alcaraz L.D., Moreno-Hagelsieb G., Eguiarte L.E. et al. Understanding the evolutionary relationships and major traits of Bacillus through comparative genomics // BMC Genomics. 2010. V. 11. № 1. P. 332. https://doi.org/10.1186/1471-2164-11-332</mixed-citation></ref><ref id="B39"><label>39.</label><mixed-citation>Rasko D.A., Rosovitz M.J., Myers G.S.A. et al. The pangenome structure of Escherichia coli: Comparative genomic analysis of E. coli commensal and pathogenic isolates // J. Bacteriol. 2008. V. 190. № 20. P. 6881–6893. https://doi.org/10.1128/JB.00619-08</mixed-citation></ref><ref id="B40"><label>40.</label><mixed-citation>Merot-L’anthoene V., Tournebize R., Darracq O. et al. Development and evaluation of a genome – wide Coffee 8.5K SNP array and its application for high-density genetic mapping and for investigating the origin of Coffea arabica L. // Plant Biotechnol. J. 2019. V. 17. № 7. P. 1418–1430. https://doi.org/10.1111/pbi.13066</mixed-citation></ref><ref id="B41"><label>41.</label><mixed-citation>Budak H., Kaya S.B., Cagirici H.B. Long non-coding RNA in plants in the era of reference sequences // Front. Plant Sci. Frontiers. 2020. V. 11. P. 441273.</mixed-citation></ref><ref id="B42"><label>42.</label><mixed-citation>Britto-Kido S. de A., Neto J.R.C.F., Pandolfi V. et al. Natural antisense transcripts in plants: A review and identification in soybean infected with Phakopsora pachyrhizi SuperSAGE Library // Sci. World J. 2013. V. 2013.</mixed-citation></ref><ref id="B43"><label>43.</label><mixed-citation>Kapusta A., Kronenberg Z., Lynch V.J. et al. Transposable elements are major contributors to the origin, diversification, and regulation of vertebrate long noncoding RNAs // PLoS Genet. 2013. V. 9. № 4. P. e1003470.</mixed-citation></ref><ref id="B44"><label>44.</label><mixed-citation>Лукина С.С., Бурденный А.М., Филиппова Е.А. et al. Роль длинных некодирующих РНК и ДНК-метилирования в патогенезе рака яичников // Патол. физиол. и эксперим. терапия. 2022. Т. 66. № 4. С. 143–156.</mixed-citation></ref><ref id="B45"><label>45.</label><mixed-citation>Ulitsky I., Bartel D.P. lincRNAs: Genomics, evolution, and mechanisms // Cell. 2013. V. 154. № 1. P. 26–46.</mixed-citation></ref><ref id="B46"><label>46.</label><mixed-citation>Dinger M.E., Pang K.C., Mercer T.R. et al. Differentiating protein-coding and noncoding RNA: Challenges and ambiguities // PLoS Comput. Biol. 2008. V. 4. № 11. P. e1000176.</mixed-citation></ref><ref id="B47"><label>47.</label><mixed-citation>Xie C., Yuan J., Li H. et al. NONCODEv4: Exploring the world of long non-coding RNA genes // Nucleic Acids Res. 2014. V. 42. № D1. P. D98–D103.</mixed-citation></ref><ref id="B48"><label>48.</label><mixed-citation>Pal D., Rao M.R.S. Long noncoding RNAs in pluripotency of stem cells and cell fate specification // Long Non Coding RNA Biology / Ed. Rao M.R.S. Singapore: Springer, 2017. V. 1008. P. 223–252.</mixed-citation></ref><ref id="B49"><label>49.</label><mixed-citation>Татосян К.А., Зиневич Л.С., Демин Д.Э., Шварц А.М. Функциональные особенности длинных некодирующих РНК, содержащих последовательности мобильных генетических элементов // Мол. биол. 2020. Т. 54. № 5. P. 718–724.</mixed-citation></ref><ref id="B50"><label>50.</label><mixed-citation>Zhao Z., Zang S., Zou W. et al. Long non-coding RNAs: New players in plants // Int. J. Mol. Sci. MDPI. 2022. V. 23. № 16. P. 9301.</mixed-citation></ref><ref id="B51"><label>51.</label><mixed-citation>Kopp F., Mendell J.T. Functional classification and experimental dissection of long noncoding RNAs // Cell. Elsevier. 2018. V. 172. № 3. P. 393–407.</mixed-citation></ref><ref id="B52"><label>52.</label><mixed-citation>Huarte M., Rinn J.L. Large non-coding RNAs: missing links in cancer? // Hum. Mol. Genet. 2010. V. 19. № R2. P. R152–R161.</mixed-citation></ref><ref id="B53"><label>53.</label><mixed-citation>Hung Т., Wang Y., Lin M.F. et al. Extensive and coordinated transcription of noncoding RNAs within cell-cycle promoters // Nat. Genet. 2011. V. 43. № 7. P. 621–629.</mixed-citation></ref><ref id="B54"><label>54.</label><mixed-citation>Loewer S., Cabili M.N., Guttman M. et al. Large intergenic non-coding RNA-RoR modulates reprogramming of human induced pluripotent stem cells // Nat. Genet. 2010. V. 42. № 12. P. 1113–1117.</mixed-citation></ref><ref id="B55"><label>55.</label><mixed-citation>Wang K.C., Chang H.Y. Molecular mechanisms of long noncoding RNAs // Mol. Cell. 2011. V. 43. № 6. P. 904–914.</mixed-citation></ref><ref id="B56"><label>56.</label><mixed-citation>Chen J., Wang H., Yao Y. Experimental study of nonlinear ultrasonic behavior of soil materials during the compaction // Ultrasonics. 2016. V. 69. P. 19–24.</mixed-citation></ref><ref id="B57"><label>57.</label><mixed-citation>Kotake Y., Nakagawa T., Kitagawa K. et al. Long non-coding RNA ANRIL is required for the PRC2 recruitment to and silencing of p15INK4B tumor suppressor gene // Oncogene. 2011. V. 30. № 16. P. 1956–1962.</mixed-citation></ref><ref id="B58"><label>58.</label><mixed-citation>Ye X., Wang S., Zhao X. et al. Role of lncRNAs in cis- and trans-regulatory responses to salt in Populus trichocarpa // Plant J. 2022. V. 110. № 4. P. 978–993. https://doi.org/10.1111/tpj.15714</mixed-citation></ref><ref id="B59"><label>59.</label><mixed-citation>Yang L., Froberg J.E., Lee J.T. Long noncoding RNAs: Fresh perspectives into the RNA world // Trends Biochem. Sci. 2014. V. 39. № 1. P. 35–43.</mixed-citation></ref><ref id="B60"><label>60.</label><mixed-citation>Lam M.T.Y., Cho H., Lesch H.P. et al. Rev-Erbs repress macrophage gene expression by inhibiting enhancer-directed transcription // Nature. 2013. V. 498. № 7455. P. 511–515.</mixed-citation></ref><ref id="B61"><label>61.</label><mixed-citation>Kim T.-K., Hemberg M., Gray J.M. Enhancer RNAs: A class of long noncoding RNAs synthesized at enhancers // Cold Spring Harb. Perspect. Biol. 2015. V. 7. № 1. P. a018622.</mixed-citation></ref><ref id="B62"><label>62.</label><mixed-citation>Kim T.-K., Hemberg M., Gray J.M. et al. Widespread transcription at neuronal activity-regulated enhancers // Nature. 2010. V. 465. № 7295. P. 182–187.</mixed-citation></ref><ref id="B63"><label>63.</label><mixed-citation>Lai F., Orom U.A., Cesaroni M. et al. Activating RNAs associate with Mediator to enhance chromatin architecture and transcription // Nature. 2013. V. 494. № 7438. P. 497–501.</mixed-citation></ref><ref id="B64"><label>64.</label><mixed-citation>Melo C.A., Drost J., Wijchers P.J. et al. eRNAs are required for p53-dependent enhancer activity and gene transcription // Mol. Cell. 2013. V. 49. № 3. P. 524–535.</mixed-citation></ref><ref id="B65"><label>65.</label><mixed-citation>Zhang P., Meng J., Luan Y. et al. Plant miRNA–lncRNA interaction prediction with the ensemble of CNN and IndRNN // Interdiscip. Sci. Comput. Life Sci. 2020. V. 12. P. 82–89.</mixed-citation></ref><ref id="B66"><label>66.</label><mixed-citation>Chen K., Rajewsky N. The evolution of gene regulation by transcription factors and microRNAs // Nat. Rev. Genet. 2007. V. 8. № 2. P. 93–103.</mixed-citation></ref><ref id="B67"><label>67.</label><mixed-citation>Jin Q., Zhao Z., Zhao Q. et al. Long noncoding RNAs: Emerging roles in pulmonary hypertension // Heart Fail. Rev. 2020. V. 25. P. 795–815.</mixed-citation></ref><ref id="B68"><label>68.</label><mixed-citation>Huntzinger E., Izaurralde E. Gene silencing by microRNAs: Contributions of translational repression and mRNA decay // Nat. Rev. Genet. 2011. V. 12. № 2. P. 99–110.</mixed-citation></ref><ref id="B69"><label>69.</label><mixed-citation>Yoon J.-H., Abdelmohsen K., Kim J. et al. Scaffold function of long non-coding RNA HOTAIR in protein ubiquitination // Nat. Commun. 2013. V. 4. № 1. P. 2939.</mixed-citation></ref><ref id="B70"><label>70.</label><mixed-citation>Mukherjee N., Corcoran D.L., Nusbaum J.D. et al. Integrative regulatory mapping indicates that the RNA-binding protein HuR couples pre-mRNA processing and mRNA stability // Mol. Cell. 2011. V. 43. № 3. P. 327–339.</mixed-citation></ref><ref id="B71"><label>71.</label><mixed-citation>Thomson D.W., Dinger M.E. Endogenous microRNA sponges: Evidence and controversy // Nat. Rev. Genet. 2016. V. 17. № 5. P. 272–283.</mixed-citation></ref><ref id="B72"><label>72.</label><mixed-citation>Franco-Zorrilla J.M., Valli A., Todesco M. et al. Target mimicry provides a new mechanism for regulation of microRNA activity // Nat. Genet. 2007. V. 39. № 8. P. 1033–1037.</mixed-citation></ref><ref id="B73"><label>73.</label><mixed-citation>Du Q., Wang K., Zou C. et al. The PILNCR1-miR399 regulatory module is important for low phosphate tolerance in maize // Plant Physiol. 2018. V. 177. № 4. P. 1743–1753.</mixed-citation></ref><ref id="B74"><label>74.</label><mixed-citation>Wang T., Zhao M., Zhang X. et al. Novel phosphate deficiency-responsive long non-coding RNAs in the legume model plant Medicago truncatula // J. Exp. Bot. 2017. V. 68. № 21–22. P. 5937–5948.</mixed-citation></ref><ref id="B75"><label>75.</label><mixed-citation>Faghihi M.A., Zhang M., Huang J. et al. Evidence for natural antisense transcript-mediated inhibition of microRNA function // Genome Biol. 2010. V. 11. № 5. https://doi.org/10.1186/gb-2010-11-5-r56</mixed-citation></ref><ref id="B76"><label>76.</label><mixed-citation>Kimura T., Jiang S., Nishizawa M. et al. Stabilization of human interferon-α1 mRNA by its antisense RNA // Cell. Mol. Life Sci. 2013. V. 70. № 8. P. 1451–1467. https://doi.org/10.1007/s00018-012-1216-x</mixed-citation></ref><ref id="B77"><label>77.</label><mixed-citation>Wang Y., Pang W.J., Wei N. et al. Identification, stability and expression of Sirt1 antisense long non-coding RNA // Gene. 2014. V. 539. № 1. P. 117–124.</mixed-citation></ref><ref id="B78"><label>78.</label><mixed-citation>Cai X., Cullen B.R. The imprinted H19 noncoding RNA is a primary microRNA precursor // RNА. 2007. V. 13. № 3. P. 313–316.</mixed-citation></ref><ref id="B79"><label>79.</label><mixed-citation>Augoff K., McCue B., Plow E.F. et al. miR-31 and its host gene lncRNA LOC554202 are regulated by promoter hypermethylation in triple-negative breast cancer // Mol. Cancer. 2012. V. 11. № 1. https://doi.org/10.1186/1476-4598-11-5</mixed-citation></ref><ref id="B80"><label>80.</label><mixed-citation>Kallen A.N., Zhou X.B., Xu J. et al. The imprinted H19 lncRNA antagonizes let-7 microRNAs // Mol. Cell. 2013. V. 52. № 1. P. 101–112.</mixed-citation></ref><ref id="B81"><label>81.</label><mixed-citation>Amor B.B., Wirth S., Merchan F. et al. Novel long non-protein coding RNAs involved in Arabidopsis differentiation and stress responses // Genome Res. 2009. V. 19. № 1. P. 57–69.</mixed-citation></ref><ref id="B82"><label>82.</label><mixed-citation>Hirsch J., Lefort V., Vankersschaver M. et al. Characterization of 43 non-protein-coding mRNA genes in Arabidopsis, including the MIR162a-derived transcripts // Plant Physiol. 2006. V. 140. № 4. P. 1192–1204.</mixed-citation></ref><ref id="B83"><label>83.</label><mixed-citation>Lamin-Samu A.T., Zhuo S., Ali M., Lu G. Long non-coding RNA transcriptome landscape of anthers at different developmental stages in response to drought stress in tomato // Genomics. 2022. V. 114. № 4. P. 110383.</mixed-citation></ref><ref id="B84"><label>84.</label><mixed-citation>Kryuchkova-Mostacci N., Robinson-Rechavi M. A benchmark of gene expression tissue-specificity metrics // Brief. Bioinform. 2017. V. 18. № 2. P. 205–214.</mixed-citation></ref><ref id="B85"><label>85.</label><mixed-citation>Li L., Eichten S.R., Shimizu R. et al. Genome-wide discovery and characterization of maize long non-coding RNAs // Genome Biol. 2014. V. 15. № 2. P. https://doi.org/10.1186/gb-2014-15-2-r40.</mixed-citation></ref><ref id="B86"><label>86.</label><mixed-citation>Han L., Mu Z., Luo Z. et al. New lncRNA annotation reveals extensive functional divergence of the transcriptome in maize // J. Integr. Plant Biol. 2019. V. 61. № 4. P. 394–405. https://doi.org/10.1111/jipb.12708</mixed-citation></ref><ref id="B87"><label>87.</label><mixed-citation>Subramanian S., Kumar S. Gene expression intensity shapes evolutionary rates of the proteins encoded by the vertebrate genome // Genetics. 2004. V. 168. № 1. P. 373–381.</mixed-citation></ref><ref id="B88"><label>88.</label><mixed-citation>Yanai I., Benjamin H., Shmoish M. et al. Genome-wide midrange transcription profiles reveal expression level relationships in human tissue specification // Bioinformatics. 2005. V. 21. № 5. P. 650–659.</mixed-citation></ref><ref id="B89"><label>89.</label><mixed-citation>Ceriani L., Verme P. The origins of the Gini index: Extracts from Variabilità e Mutabilità (1912) by Corrado Gini // J. Econ. Inequal. 2012. V. 10. P. 421–443.</mixed-citation></ref><ref id="B90"><label>90.</label><mixed-citation>Julien P., Brawand D., Soumillon M. et al. Mechanisms and evolutionary patterns of mammalian and avian dosage compensation // PLoS Biol. 2012. V. 10. № 5. P. e1001328.</mixed-citation></ref><ref id="B91"><label>91.</label><mixed-citation>Xiao S.-J., Zhang C., Zou Q., Ji Z.L. TiSGeD: A database for tissue-specific genes // Bioinformatics. 2010. V. 26. № 9. P. 1273–1275.</mixed-citation></ref><ref id="B92"><label>92.</label><mixed-citation>Yu X., Lin J., Zack D.J., Qian J. Computational analysis of tissue-specific combinatorial gene regulation: Predicting interaction between transcription factors in human tissues // Nucl. Acids Res. 2006. V. 34. № 17. P. 4925–4936.</mixed-citation></ref><ref id="B93"><label>93.</label><mixed-citation>Huang X., Li S.Z., Wang Y. Jensen-Shannon boosting learning for object recognition // 2005 IEEE Computer Society Conference on Computer Vision and Pattern Recognition (CVPR’05). IEEE. 2005. V. 2. P. 144–149.</mixed-citation></ref><ref id="B94"><label>94.</label><mixed-citation>Marquardt S., Raitskin O., Wu Z. et al. Functional consequences of splicing of the antisense transcript COOLAIR on FLC transcription // Mol. Cell. 2014. V. 54. № 1. P. 156–165.</mixed-citation></ref><ref id="B95"><label>95.</label><mixed-citation>Liu X., Hao L., Li D. et al. Long non-coding RNAs and their biological roles in plants // Genomics Proteomics Bioinformatics. 2015. V. 13. № 3. P. 137–147.</mixed-citation></ref><ref id="B96"><label>96.</label><mixed-citation>Ding J., Lu Q., Ouyang Y. et al. A long noncoding RNA regulates photoperiod-sensitive male sterility, an essential component of hybrid rice // Proc. Natl Acad. Sci. USA. 2012. V. 109. № 7. P. 2654–2659. https://doi.org/10.1073/pnas.1121374109</mixed-citation></ref><ref id="B97"><label>97.</label><mixed-citation>Song J.-H., Cao J.-S., Wang C.-G. BcMF11, a novel non-coding RNA gene from Brassica campestris, is required for pollen development and male fertility // Plant Cell Rep. 2013. V. 32. P. 21–30.</mixed-citation></ref><ref id="B98"><label>98.</label><mixed-citation>Kim J., Yi H., Choi G. et al. Functional characterization of phytochrome interacting factor 3 in phytochrome-mediated light signal transduction // Plant Cell. 2003. V. 15. № 10. P. 2399–2407.</mixed-citation></ref><ref id="B99"><label>99.</label><mixed-citation>Wang J., Meng X., Dobrovolskaya O.B. et al. Non-coding RNAs and their roles in stress response in plants: 5 // Genomics Proteomics Bioinformatics. 2017. V. 15. № 5. P. 301–312. https://doi.org/10.1016/j.gpb.2017.01.007</mixed-citation></ref><ref id="B100"><label>100.</label><mixed-citation>Wang A., Hu J., Gao C. et al. Genome-wide analysis of long non-coding RNAs unveils the regulatory roles in the heat tolerance of Chinese cabbage (Brassica rapa ssp. chinensis) // Sci. Rep. 2019. V. 9. № 1. P. 5002</mixed-citation></ref><ref id="B101"><label>101.</label><mixed-citation>Wang P., Dai L., Ai J. et al. Identification and functional prediction of cold-related long non-coding RNA (lncRNA) in grapevine // Sci. Rep. 2019. V. 9. № 1. P. 6638.</mixed-citation></ref><ref id="B102"><label>102.</label><mixed-citation>Chung P.J., Jung H., Jeong D.H. et al. Transcriptome profiling of drought responsive noncoding RNAs and their target genes in rice // BMC Genomics. 2016. V. 17. № 1. P. 563. https://doi.org/10.1186/s12864-016-2997-3</mixed-citation></ref><ref id="B103"><label>103.</label><mixed-citation>Calixto C.P., Tzioutziou N.A., James A.B. et al. Cold-dependent expression and alternative splicing of Arabidopsis long non-coding RNAs // Front. Plant Sci. Frontiers Media SA. 2019. V. 10. P. 235.</mixed-citation></ref><ref id="B104"><label>104.</label><mixed-citation>Jha U.C., Nayyar H., Jha R. et al. Long non-coding RNAs: Emerging players regulating plant abiotic stress response and adaptation // BMC Plant Biol. 2020. V. 20. № 1. P. 466. https://doi.org/10.1186/s12870-020-02595-x</mixed-citation></ref><ref id="B105"><label>105.</label><mixed-citation>Zhang X., Dong J., Deng F. et al. The long non-coding RNA lncRNA973 is involved in cotton response to salt stress // BMC Plant Biol. 2019. V. 19. № 1. P. 459. https://doi.org/10.1186/s12870-019-2088-0</mixed-citation></ref><ref id="B106"><label>106.</label><mixed-citation>Wang X., Fan H., Wang B., Yuan F. Research progress on the roles of lncRNAs in plant development and stress responses // Front. Plant Sci. Frontiers Media SA. 2023. V. 14. P. 1138901.</mixed-citation></ref><ref id="B107"><label>107.</label><mixed-citation>Chen J., Zhong Y., Qi X. LncRNA TCONS_00021861 is functionally associated with drought tolerance in rice (Oryza sativa L.) via competing endogenous RNA regulation // BMC Plant Biol. 2021. V. 21. № 1. P. 410. https://doi.org/10.1186/s12870-021-03195-z</mixed-citation></ref><ref id="B108"><label>108.</label><mixed-citation>Kazemzadeh M., Safaralizadeh R., Orang A.V. LncRNAs: Emerging players in gene regulation and disease pathogenesis // J. Genet. 2015. V. 94. P. 771–784.</mixed-citation></ref><ref id="B109"><label>109.</label><mixed-citation>Zhu Y., Chen L., Hong X. et al. Revealing the novel complexity of plant long non-coding RNA by strand-specific and whole transcriptome sequencing for evolutionarily representative plant species // BMC Genomics. 2022. V. 23. № S4. P. 381. https://doi.org/10.1186/s12864-022-08602-9</mixed-citation></ref><ref id="B110"><label>110.</label><mixed-citation>Ulitsky I. Evolution to the rescue: Using comparative genomics to understand long non-coding RNAs // Nat. Rev. Genet. 2016. V. 17. № 10. P. 601–614.</mixed-citation></ref><ref id="B111"><label>111.</label><mixed-citation>Wang H., Niu Q.W., Wu H.W. et al. Analysis of non-coding transcriptome in rice and maize uncovers roles of conserved lnc RNAs associated with agriculture traits // Plant J. 2015. V. 84. № 2. P. 404–416. https://doi.org/10.1111/tpj.13018</mixed-citation></ref><ref id="B112"><label>112.</label><mixed-citation>Nitsche A., Stadler P.F. Evolutionary clues in lncRNAs // WIREs RNA. 2017. V. 8. № 1. https://doi.org/10.1002/wrna.1376</mixed-citation></ref><ref id="B113"><label>113.</label><mixed-citation>Sang S., Chen W., Zhang D. et al. Data integration and evolutionary analysis of long non-coding RNAs in 25 flowering plants: 3 // BMC Genomics. 2021. V. 22. № 3. P. 739. https://doi.org/10.1186/s12864-021-08047-6</mixed-citation></ref><ref id="B114"><label>114.</label><mixed-citation>Zhang Y.-C., Liao J.Y., Li Z.Y. et al. Genome-wide screening and functional analysis identify a large number of long noncoding RNAs involved in the sexual reproduction of rice // Genome Biol. 2014. V. 15. № 12. P. 512. https://doi.org/10.1186/s13059-014-0512-1</mixed-citation></ref><ref id="B115"><label>115.</label><mixed-citation>Pronozin A.Y., Bragina M.K., Salina E.A. Crop pangenomes // Vavilov. J. Genet. Breed. 2021. V. 25. № 1. P. 57.</mixed-citation></ref><ref id="B116"><label>116.</label><mixed-citation>Vernikos G., Medini D., Riley D.R., Tettelin H. et al. Ten years of pan-genome analyses // Curr. Opin. Microbiol. 2015. V. 23. P. 148–154.</mixed-citation></ref><ref id="B117"><label>117.</label><mixed-citation>Lapierre P., Gogarten J.P. Estimating the size of the bacterial pan-genome // Trends Genet. 2009. V. 25. № 3. P. 107–110.</mixed-citation></ref><ref id="B118"><label>118.</label><mixed-citation>Chekanova J.A., Gregory B.D., Reverdatto S.V. et al. Genome-wide high-resolution mapping of exosome substrates reveals hidden features in the Arabidopsis transcriptome // Cell. 2007. V. 131. № 7. P. 1340–1353.</mixed-citation></ref><ref id="B119"><label>119.</label><mixed-citation>Medina C.A., Samac D.A., Yu L.-X. Pan-transcriptome identifying master genes and regulation network in response to drought and salt stresses in Alfalfa (Medicago sativa L.) // Sci. Rep. 2021. V. 11. № 1. P. 17203.</mixed-citation></ref><ref id="B120"><label>120.</label><mixed-citation>Jin M., Liu H., He C. et al. Maize pan-transcriptome provides novel insights into genome complexity and quantitative trait variation // Sci. Rep. 2016. V. 6. № 1. P. 18936.</mixed-citation></ref><ref id="B121"><label>121.</label><mixed-citation>Chowdhary A., Satagopam V., Schneider R. Long non-coding RNAs: Mechanisms, experimental, and computational approaches in identification, characterization, and their biomarker potential in cancer // Front. Genet. 2021. V. 12. https://doi.org/10.3389/fgene.2021.649619</mixed-citation></ref><ref id="B122"><label>122.</label><mixed-citation>Svergun D.I., Koch M.H. Small-angle scattering studies of biological macromolecules in solution // Rep. Prog. Phys. 2003. V. 66. № 10. P. 1735.</mixed-citation></ref><ref id="B123"><label>123.</label><mixed-citation>Schön P. Atomic force microscopy of RNA: State of the art and recent advancements // Seminars in Cell &amp; Developmental Biology. 2018. V. 73. P. 209–219.</mixed-citation></ref><ref id="B124"><label>124.</label><mixed-citation>Chillón I., Marcia M., Legiewicz M. et al. Native purification and analysis of long RNAs // Methods in Enzymology. 2015. V. 558. P. 3–37.</mixed-citation></ref><ref id="B125"><label>125.</label><mixed-citation>Cheung F., Haas B.J., Goldberg S.M.D. et al. Sequencing Medicago truncatula expressed sequenced tags using 454 Life Sciences technology // BMC Genomics. 2006. V. 7. № 1. https://doi.org/10.1186/1471-2164-7-272</mixed-citation></ref><ref id="B126"><label>126.</label><mixed-citation>Au P.C.K., Zhu Q.-H. Identification of lncRNAs using computational and experimental approaches // Regulatory RNAs / Eds Mallick B., Ghosh Z. Berlin; Heidelberg: Springer, 2012. P. 319–340.</mixed-citation></ref><ref id="B127"><label>127.</label><mixed-citation>Babak T., Blencowe B.J., Hughes T.R. A systematic search for new mammalian noncoding RNAs indicates little conserved intergenic transcription // BMC Genomics. 2005. V. 6. № 1. P. 104. https://doi.org/10.1186/1471-2164-6-104</mixed-citation></ref><ref id="B128"><label>128.</label><mixed-citation>Shiraki T., Kondo S., Katayama S. et al. Cap analysis gene expression for high-throughput analysis of transcriptional starting point and identification of promoter usage // Proc. Natl Acad. Sci. USA. 2003. V. 100. № 26. P. 15776–15781. https://doi.org/10.1073/pnas.2136655100</mixed-citation></ref><ref id="B129"><label>129.</label><mixed-citation>Merino E.J., Wilkinson K.A., Coughlan J.L., Weeks K.M. RNA structure analysis at single nucleotide resolution by Selective 2‘-Hydroxyl Acylation and Primer Extension (SHAPE) // J. Am. Chem. Soc. 2005. V. 127. № 12. P. 4223–4231. https://doi.org/10.1021/ja043822v</mixed-citation></ref><ref id="B130"><label>130.</label><mixed-citation>Kertesz M., Wan Y., Mazor E. et al. Genome-wide measurement of RNA secondary structure in yeast // Nature. 2010. V. 467. № 7311. P. 103–107.</mixed-citation></ref><ref id="B131"><label>131.</label><mixed-citation>Hawkes E.J., Hennelly S.P., Novikova I.V. et al. COOLAIR antisense RNAs form evolutionarily conserved elaborate secondary structures // Cell Rep. 2016. V. 16. № 12. P. 3087–3096.</mixed-citation></ref><ref id="B132"><label>132.</label><mixed-citation>Kim D.N., Thiel B.C., Mrozowich T. et al. Zinc-finger protein CNBP alters the 3-D structure of lncRNA Braveheart in solution // Nat. Commun. 2020. V. 11. № 1. P. 148.</mixed-citation></ref><ref id="B133"><label>133.</label><mixed-citation>Li A., Zhang J., Zhou Z. PLEK: A tool for predicting long non-coding RNAs and messenger RNAs based on an improved k-mer scheme // BMC Bioinformatics. 2014. V. 15. № 1. P. 311. https://doi.org/10.1186/1471-2105-15-311</mixed-citation></ref><ref id="B134"><label>134.</label><mixed-citation>Kang Y.-J., Yang D.C., Kong L. et al. CPC2: A fast and accurate coding potential calculator based on sequence intrinsic features // Nucl. Acids Res. 2017. V. 45. № W1. P. W12–W16.</mixed-citation></ref><ref id="B135"><label>135.</label><mixed-citation>Wang L., Park H.J., Dasari S. et al. CPAT: Coding-Potential Assessment Tool using an alignment-free logistic regression model // Nucl. Acids Res. 2013. V. 41. № 6. P. e74.</mixed-citation></ref><ref id="B136"><label>136.</label><mixed-citation>Da Costa Negri T., Paschoal A.R., Alves W.A.L. Comparison tools for lncRNA identification: Analysis among plants and humans // 2020 IEEE Conference on Computational Intelligence in Bioinformatics and Computational Biology (CIBCB). IEEE. 2020. P. 1–8.</mixed-citation></ref><ref id="B137"><label>137.</label><mixed-citation>Pronozin A.Y., Afonnikov D.A. ICAnnoLncRNA: A snakemake pipeline for a long non-coding-RNA search and annotation in transcriptomic sequences // Genes. MDPI. 2023. V. 14. № 7. P. 1331.</mixed-citation></ref><ref id="B138"><label>138.</label><mixed-citation>Gong Y., Huang H.T., Liang Y. et al. lncRNA-screen: An interactive platform for computationally screening long non-coding RNAs in large genomics datasets // BMC Genomics. 2017. V. 18. № 1. P. 434. https://doi.org/10.1186/s12864-017-3817-0</mixed-citation></ref><ref id="B139"><label>139.</label><mixed-citation>Кит О.И., Кириченко Е.Ю., Кириченко Ю.Г. и др. Длинные некодирующие РНК, ассоциированные с канцерогенезом: биологическое значение и перспективы применения в диагностике // Клин. лаб. диагностика. 2016. Т. 61. № 1. P. 13–16.</mixed-citation></ref><ref id="B140"><label>140.</label><mixed-citation>Gao L., Gonda I., Sun H. et al. The tomato pan-genome uncovers new genes and a rare allele regulating fruit flavor // Nat. Genet. 2019. V. 51. № 6. P. 1044–1051.</mixed-citation></ref></ref-list></back></article>
