<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Publishing DTD v1.3 20210610//EN" "JATS-journalpublishing1-3.dtd">
<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 pub-id-type="doi">10.18699/vjgb-25-128</article-id><article-id custom-type="elpub" pub-id-type="custom">vavilov-4921</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>УСТОЙЧИВОСТЬ РАСТЕНИЙ К СТРЕССОВЫМ ФАКТОРАМ</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>RESISTANCE OF PLANTS TO STRESS FACTORS</subject></subj-group></article-categories><title-group><article-title>Дифференциальный профиль экспрессии генов транскрипционных факторов подсемейства DREB2 в динамике солевого стресса и послестрессового восстановления растений томата</article-title><trans-title-group xml:lang="en"><trans-title>Differential expression profile of DREB2 subfamily transcription factor genes in the dynamics of salt stress and post-stress recovery in tomato plants</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-3668-7601</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Филюшин</surname><given-names>М. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Filyushin</surname><given-names>M. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Москва</p></bio><bio xml:lang="en"><p>Moscow</p></bio><email xlink:type="simple">michel7753@mail.ru</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-4692-3727</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Щенникова</surname><given-names>А. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Shchennikova</surname><given-names>A. V.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Москва</p></bio><bio xml:lang="en"><p>Moscow</p></bio><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-6091-0765</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Кочиева</surname><given-names>Е. З.</given-names></name><name name-style="western" xml:lang="en"><surname>Kochieva</surname><given-names>E. Z.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Москва</p></bio><bio xml:lang="en"><p>Moscow</p></bio><xref ref-type="aff" rid="aff-1"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru">Федеральный исследовательский центр «Фундаментальные основы биотехнологии» Российской академии наук<country>Россия</country></aff><aff xml:lang="en">Institute of Bioengineering, Research Center of Biotechnology of the Russian Academy of Sciences<country>Russian Federation</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2025</year></pub-date><pub-date pub-type="epub"><day>11</day><month>01</month><year>2026</year></pub-date><volume>29</volume><issue>8</issue><fpage>1213</fpage><lpage>1220</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Филюшин М.А., Щенникова А.В., Кочиева Е.З., 2026</copyright-statement><copyright-year>2026</copyright-year><copyright-holder xml:lang="ru">Филюшин М.А., Щенникова А.В., Кочиева Е.З.</copyright-holder><copyright-holder xml:lang="en">Filyushin M.A., Shchennikova A.V., Kochieva E.Z.</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/4921">https://vavilov.elpub.ru/jour/article/view/4921</self-uri><abstract><p>В ответ на стрессовое воздействие в геноме растения происходят эпигенетические модификации, вместе формирующие стрессовую память, которая может наследоваться и повышает эффективность защитной реакции растения на повторные стрессовые события. Гены, чья экспрессия становится мишенью эпигенетических модификаций, служат биомаркерами стрессовой памяти. Их характерными признаками считаются профиль экспрессии, различающийся между ответами на первичное и повторное стрессовые события, а также длительное удерживание изменений после отмены стресса. Томат (Solanum lycopersicum L.) – важная овощная культура, урожайность которой снижается при засолении почв. К генам, индуцируемым солевым стрессом, относятся гены транскрипционных факторов подсемейства DREB2 (DEHYDRATION-RESPONSIVE ELEMENT-BINDING PROTEIN 2). В настоящей работе проведена оценка генов SlDREB2 томата как возможных генов-маркеров памяти о солевом стрессе. Для этого в листьях растений двух сортов, Гном и Отрадный, с разной степенью солеустойчивости была определена экспрессия генов SlDREB16, 20, 22, 24, 43, 44 и 46 в ответ на 24-ч воздействие NaCl и в динамике продолжительного (14 дней) послестрессового периода восстановления. Выявлены значительные генотип специфичные колебания уровней транскриптов генов как в контроле, так и в подвергнутых стрессу растениях. Показано, что в процессе длительной фазы памяти экспрессия генов возвращается к контрольным показателям либо временно (SlDREB24, 44 и 46 у среднеустойчивого сорта Гном через 7 дней; через 14 дней экспрессия снова меняется), либо медленно (SlDREB16 и 43 у высокоустойчивого сорта Отрадный через 14 дней восстановления). Было определено, что только два гена, SlDREB22 и 46, имеют сходный между сортами паттерн колебаний экспрессии в динамике стресса и восстановления, а ген SlDREB20 не экспрессируется ни в контроле, ни в опыте. Полученные данные позволили предположить, что гены подсемейства SlDREB2, кроме SlDREB20, участвуют в ответе S. lycopersicum на солевой стресс генотип-специфичным образом и могут служить маркерами стрессовой памяти, сцепленными с эпигенетической регуляцией адаптации томата к солевому стрессу. Гены SlDREB16, 28, 43 и 44 могут вносить вклад в определение различий в механизме регуляции ответа растений на солевой стресс между солеустойчивыми генотипами S. lycopersicum. Полученные результаты могут стать основой для дальнейших исследований роли генов SlDREB2 в эпигенетической регуляции адаптации растений томата к солевому стрессу, что может быть использовано в селекции солеустойчивых сортов.</p></abstract><trans-abstract xml:lang="en"><p>In response to stress, epigenetic modifications occur in the plant genome, which together form a stress memory that can be inherited and increases the efficiency of the plant's defense response to repeated stress events. Genes whose expression becomes the target of epigenetic modifications serve as biomarkers of stress memory. Their characteristic features are considered to be an expression profile that differs between responses to primary and repeated stress events, as well as long-term retention of changes after the stress is canceled. Tomato (Solanum lycopersicum L.) is an important vegetable crop whose yield decreases with soil salinity. Genes induced by salt stress include genes encoding transcription factors of the DREB2 (DEHYDRATION-RESPONSIVE ELEMENT-BINDING PROTEIN 2) subfamily. In this work, we evaluated the SlDREB2 genes of tomato as possible marker genes of salt stress memory. The expression of the genes SlDREB16, 20, 22, 24, 43, 44 and 46 was determined in the leaves of two plant varieties (Gnom, Otradnyi) with different degrees of salt tolerance in response to 24 h of NaCl exposure and in the dynamics of a long-term (14 days) post-stress recovery period. Significant genotype-specific fluctuations in the levels of gene transcripts were revealed both in the control and in the stressed plants. It was shown that during the long-term memory phase, gene expression returns to the control values either temporarily (SlDREB24, 44 and 46 in the moderately resistant Gnom variety after 7 days; after 14 days, the expression changed again) or slowly (SlDREB16 and 43 in the highly resistant Otradnyi variety after 14 days of recovery). Only two genes (SlDREB22 and 46) showed a similar between varieties pattern of expression fluctuations in the dynamics of stress and recovery, and the SlDREB20 gene was not expressed in either the control or the experiment. The data obtained suggest that the SlDREB2 subfamily genes (except SlDREB20) are involved in the response of S. lycopersicum to salt stress in a genotype-specific manner and can serve as markers of stress memory linked to the epigenetic regulation of tomato adaptation to salt stress. The SlDREB16, 28, 43 and 44 genes may contribute to the determination of differences in the mechanism of regulation of plant response to salt stress between salt-tolerant genotypes of S. lycopersicum. The obtained results can form the basis for further studies of the role of SlDREB2 genes in the epigenetic regulation of tomato plant adaptation to salt stress, which can be used in breeding salt-tolerant varieties.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>томат</kwd><kwd>Solanum lycopersicum L.</kwd><kwd>солевой стресс</kwd><kwd>стрессовая память</kwd><kwd>фаза памяти</kwd><kwd>транскрипционные  факторы SlDREB2</kwd><kwd>экспрессия генов</kwd><kwd>гены-кандидаты стрессовой памяти</kwd></kwd-group><kwd-group xml:lang="en"><kwd>tomato</kwd><kwd>Solanum lycopersicum L.</kwd><kwd>salt stress</kwd><kwd>stress memory</kwd><kwd>memory phase</kwd><kwd>SlDREB2 transcription factors</kwd><kwd>gene expression</kwd><kwd>potential stress memory genes</kwd></kwd-group><funding-group xml:lang="ru"><funding-statement>The study was supported by the RSF grant No. 24-16-00043 (analysis of gene expression) and the Ministry of  Science and Higher Education of the Russian Federation (plant material)</funding-statement></funding-group><funding-group xml:lang="en"><funding-statement>The study was supported by the RSF grant No. 24-16-00043 (analysis of gene expression) and the Ministry of  Science and Higher Education of the Russian Federation (plant material).</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">Aina O., Bakare O.O., Fadaka A.O., Keyster M., Klein A. Plant bio markers as early detection tools in stress management in food crops: A review. Planta. 2024;259(3):60. doi 10.1007/s00425-024-04333-1</mixed-citation><mixed-citation xml:lang="en">Aina O., Bakare O.O., Fadaka A.O., Keyster M., Klein A. Plant bio markers as early detection tools in stress management in food crops: A review. Planta. 2024;259(3):60. doi 10.1007/s00425-024-04333-1</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Akbudak M.A., Filiz E., Kontbay K. DREB2 (dehydration-responsive element-binding protein 2) type transcription factor in sorghum (Sorghum bicolor): genome-wide identification, characterization and expression profiles under cadmium and salt stresses. 3 Biotech. 2018;8(10):426. doi 10.1007/s13205-018-1454-1</mixed-citation><mixed-citation xml:lang="en">Akbudak M.A., Filiz E., Kontbay K. DREB2 (dehydration-responsive element-binding protein 2) type transcription factor in sorghum (Sorghum bicolor): genome-wide identification, characterization and expression profiles under cadmium and salt stresses. 3 Biotech. 2018;8(10):426. doi 10.1007/s13205-018-1454-1</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Anisimova O.K., Shchennikova A.V., Kochieva E.Z., Filyushin M.A. Identification of chalcone synthase genes from garlic (Allium sati vum L.) and their expression levels in response to stress factors. Acta Nat. 2025;17(2):4-14. doi 10.32607/actanaturae.27639</mixed-citation><mixed-citation xml:lang="en">Anisimova O.K., Shchennikova A.V., Kochieva E.Z., Filyushin M.A. Identification of chalcone synthase genes from garlic (Allium sati vum L.) and their expression levels in response to stress factors. Acta Nat. 2025;17(2):4-14. doi 10.32607/actanaturae.27639</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Bai Y., Sunarti S., Kissoudis C., Visser R.G.F., van der Linden C.G. The role of tomato WRKY genes in plant responses to combined abio tic and biotic stresses. Front Plant Sci. 2018;9:801. doi 10.3389/fpls.2018.00801</mixed-citation><mixed-citation xml:lang="en">Bai Y., Sunarti S., Kissoudis C., Visser R.G.F., van der Linden C.G. The role of tomato WRKY genes in plant responses to combined abio tic and biotic stresses. Front Plant Sci. 2018;9:801. doi 10.3389/fpls.2018.00801</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Baillo E.H., Kimotho R.N., Zhang Z., Xu P. Transcription factors asso ciated with abiotic and biotic stress tolerance and their potential for crops improvement. Genes (Basel). 2019;10(10):771. doi 10.3390/genes10100771</mixed-citation><mixed-citation xml:lang="en">Baillo E.H., Kimotho R.N., Zhang Z., Xu P. Transcription factors asso ciated with abiotic and biotic stress tolerance and their potential for crops improvement. Genes (Basel). 2019;10(10):771. doi 10.3390/genes10100771</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Bäurle I., Trindade I. Chromatin regulation of somatic abiotic stress memory. J Exp Bot. 2020;71(17):5269-5279. doi 10.1093/jxb/eraa098</mixed-citation><mixed-citation xml:lang="en">Bäurle I., Trindade I. Chromatin regulation of somatic abiotic stress memory. J Exp Bot. 2020;71(17):5269-5279. doi 10.1093/jxb/eraa098</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Borsani O., Cuartero J., Fernández J.A., Valpuesta V., Botella M.A. Identification of two loci in tomato reveals distinct mechanisms for salt tolerance. Plant Cell. 2001;13(4):873-887. doi 10.1105/tpc.13.4.873</mixed-citation><mixed-citation xml:lang="en">Borsani O., Cuartero J., Fernández J.A., Valpuesta V., Botella M.A. Identification of two loci in tomato reveals distinct mechanisms for salt tolerance. Plant Cell. 2001;13(4):873-887. doi 10.1105/tpc.13.4.873</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Calone R., Mircea D.M., González-Orenga S., Boscaiu M., Zuzunaga Rosas J., Barbanti L., Vicente O. Effect of recurrent salt and drought stress treatments on the endangered halophyte Limonium anguste bracteatum Erben. Plants (Basel). 2023;12(1):191. doi 10.3390/plants12010191</mixed-citation><mixed-citation xml:lang="en">Calone R., Mircea D.M., González-Orenga S., Boscaiu M., Zuzunaga Rosas J., Barbanti L., Vicente O. Effect of recurrent salt and drought stress treatments on the endangered halophyte Limonium anguste bracteatum Erben. Plants (Basel). 2023;12(1):191. doi 10.3390/plants12010191</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Chen Y., Li C., Yi J., Yang Y., Lei C., Gong M. Transcriptome response to drought, rehydration and re-dehydration in potato. Int J Mol Sci. 2019;21(1):159. doi 10.3390/ijms21010159</mixed-citation><mixed-citation xml:lang="en">Chen Y., Li C., Yi J., Yang Y., Lei C., Gong M. Transcriptome response to drought, rehydration and re-dehydration in potato. Int J Mol Sci. 2019;21(1):159. doi 10.3390/ijms21010159</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Ding Y., Liu N., Virlouvet L., Riethoven J.J., Fromm M., Avramova Z. Four distinct types of dehydration stress memory genes in Arabi dopsis thaliana. BMC Plant Biol. 2013;13:229. doi 10.1186/14712229-13-229</mixed-citation><mixed-citation xml:lang="en">Ding Y., Liu N., Virlouvet L., Riethoven J.J., Fromm M., Avramova Z. Four distinct types of dehydration stress memory genes in Arabi dopsis thaliana. BMC Plant Biol. 2013;13:229. doi 10.1186/14712229-13-229</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Ding Y., Virlouvet L., Liu N., Riethoven J.J., Fromm M., Avramova Z. Dehydration stress memory genes of Zea mays; comparison with Arabidopsis thaliana. BMC Plant Biol. 2014;14:141. doi 10.1186/ 1471-2229-14-141</mixed-citation><mixed-citation xml:lang="en">Ding Y., Virlouvet L., Liu N., Riethoven J.J., Fromm M., Avramova Z. Dehydration stress memory genes of Zea mays; comparison with Arabidopsis thaliana. BMC Plant Biol. 2014;14:141. doi 10.1186/ 1471-2229-14-141</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Dubouzet J.G., Sakuma Y., Ito Y., Kasuga M., Dubouzet E.G., Miura S., Seki M., Shinozaki K., Yamaguchi-Shinozaki K. OsDREB genes in rice, Oryza sativa L., encode transcription activators that function in drought-, high-salt- and cold-responsive gene expression. Plant J. 2003;33(4):751-763. doi 10.1046/j.1365-313x.2003.01661.x</mixed-citation><mixed-citation xml:lang="en">Dubouzet J.G., Sakuma Y., Ito Y., Kasuga M., Dubouzet E.G., Miura S., Seki M., Shinozaki K., Yamaguchi-Shinozaki K. OsDREB genes in rice, Oryza sativa L., encode transcription activators that function in drought-, high-salt- and cold-responsive gene expression. Plant J. 2003;33(4):751-763. doi 10.1046/j.1365-313x.2003.01661.x</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Efremov G.I., Slugina M.A., Shchennikova A.V., Kochieva E.Z. Dif ferential regulation of phytoene synthase PSY1 during fruit caro tenogenesis in cultivated and wild tomato species (Solanum section Lycopersicon). Plants (Basel). 2020;9(9):1169 doi 10.3390/plants9091169</mixed-citation><mixed-citation xml:lang="en">Efremov G.I., Slugina M.A., Shchennikova A.V., Kochieva E.Z. Dif ferential regulation of phytoene synthase PSY1 during fruit caro tenogenesis in cultivated and wild tomato species (Solanum section Lycopersicon). Plants (Basel). 2020;9(9):1169 doi 10.3390/plants9091169</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Filyushin M.A., Shagdarova B.T., Shchennikova A.V., Il’ina A.V., Kochieva E.Z., Varlamov V.P. Pretreatment with chitosan prevents Fusarium infection and induces the expression of chitinases and β-1,3-glucanases in garlic (Allium sativum L.). Horticulturae. 2022; 8(5):383. doi 10.3390/horticulturae8050383</mixed-citation><mixed-citation xml:lang="en">Filyushin M.A., Shagdarova B.T., Shchennikova A.V., Il’ina A.V., Kochieva E.Z., Varlamov V.P. Pretreatment with chitosan prevents Fusarium infection and induces the expression of chitinases and β-1,3-glucanases in garlic (Allium sativum L.). Horticulturae. 2022; 8(5):383. doi 10.3390/horticulturae8050383</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Filyushin M.A., Anisimova O.K., Shchennikova A.V., Kochieva E.Z. DREB1 and DREB2 genes in garlic (Allium sativum L.): Genome wide identification, characterization, and stress response. Plants (Basel). 2023;12(13):2538. doi 10.3390/plants12132538</mixed-citation><mixed-citation xml:lang="en">Filyushin M.A., Anisimova O.K., Shchennikova A.V., Kochieva E.Z. DREB1 and DREB2 genes in garlic (Allium sativum L.): Genome wide identification, characterization, and stress response. Plants (Basel). 2023;12(13):2538. doi 10.3390/plants12132538</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Friedrich T., Faivre L., Bäurle I., Schubert D. Chromatin-based mecha nisms of temperature memory in plants. Plant Cell Environ. 2019; 42(3):762-770. doi 10.1111/pce.13373</mixed-citation><mixed-citation xml:lang="en">Friedrich T., Faivre L., Bäurle I., Schubert D. Chromatin-based mecha nisms of temperature memory in plants. Plant Cell Environ. 2019; 42(3):762-770. doi 10.1111/pce.13373</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Gallusci P., Agius D.R., Moschou P.N., Dobránszki J., Kaiserli E., Mar tinelli F. Deep inside the epigenetic memories of stressed plants. Trends Plant Sci. 2023;28(2):142-153. doi 10.1016/j.tplants.2022.09.004 Guo M., Wang X.S., Guo H.D., Bai S.Y., Khan A., Wang X.M., Gao Y.M., Li J.S. Tomato salt tolerance mechanisms and their po tential applications for fighting salinity: A review. Front Plant Sci. 2022;13:949541. doi 10.3389/fpls.2022.949541</mixed-citation><mixed-citation xml:lang="en">Gallusci P., Agius D.R., Moschou P.N., Dobránszki J., Kaiserli E., Mar tinelli F. Deep inside the epigenetic memories of stressed plants. Trends Plant Sci. 2023;28(2):142-153. doi 10.1016/j.tplants.2022.09.004 Guo M., Wang X.S., Guo H.D., Bai S.Y., Khan A., Wang X.M., Gao Y.M., Li J.S. Tomato salt tolerance mechanisms and their po tential applications for fighting salinity: A review. Front Plant Sci. 2022;13:949541. doi 10.3389/fpls.2022.949541</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Hassan S., Berk K., Aronsson H. Evolution and identification of DREB transcription factors in the wheat genome: modeling, docking and simulation of DREB proteins associated with salt stress. J Biomol Struct Dyn. 2022;40(16):7191-7204. doi 10.1080/07391102.2021.1894980</mixed-citation><mixed-citation xml:lang="en">Hassan S., Berk K., Aronsson H. Evolution and identification of DREB transcription factors in the wheat genome: modeling, docking and simulation of DREB proteins associated with salt stress. J Biomol Struct Dyn. 2022;40(16):7191-7204. doi 10.1080/07391102.2021.1894980</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Ismail A.M., Horie T. Genomics, physiology, and molecular breeding approaches for improving salt tolerance. Annu Rev Plant Biol. 2017; 68:405-434. doi 10.1146/annurev-arplant-042916-040936 Jacques C., Salon C., Barnard R.L., Vernoud V., Prudent M. Drought stress memory at the plant cycle level: A review. Plants (Basel). 2021;10(9):1873. doi 10.3390/plants10091873</mixed-citation><mixed-citation xml:lang="en">Ismail A.M., Horie T. Genomics, physiology, and molecular breeding approaches for improving salt tolerance. Annu Rev Plant Biol. 2017; 68:405-434. doi 10.1146/annurev-arplant-042916-040936 Jacques C., Salon C., Barnard R.L., Vernoud V., Prudent M. Drought stress memory at the plant cycle level: A review. Plants (Basel). 2021;10(9):1873. doi 10.3390/plants10091873</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Kudo M., Kidokoro S., Yoshida T., Mizoi J., Todaka D., Fernie A.R., Shinozaki K., Yamaguchi-Shinozaki K. Double overexpression of DREB and PIF transcription factors improves drought stress toler ance and cell elongation in transgenic plants. Plant Biotechnol J. 2017;15(4):458-471. doi 10.1111/pbi.12644</mixed-citation><mixed-citation xml:lang="en">Kudo M., Kidokoro S., Yoshida T., Mizoi J., Todaka D., Fernie A.R., Shinozaki K., Yamaguchi-Shinozaki K. Double overexpression of DREB and PIF transcription factors improves drought stress toler ance and cell elongation in transgenic plants. Plant Biotechnol J. 2017;15(4):458-471. doi 10.1111/pbi.12644</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Maqsood H., Munir F., Amir R., Gul A. Genome-wide identification, comprehensive characterization of transcription factors, cis-regu latory elements, protein homology, and protein interaction network of DREB gene family in Solanum lycopersicum. Front Plant Sci. 2022;13:1031679. doi10.3389/fpls.2022.1031679</mixed-citation><mixed-citation xml:lang="en">Maqsood H., Munir F., Amir R., Gul A. Genome-wide identification, comprehensive characterization of transcription factors, cis-regu latory elements, protein homology, and protein interaction network of DREB gene family in Solanum lycopersicum. Front Plant Sci. 2022;13:1031679. doi10.3389/fpls.2022.1031679</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Matsukura S., Mizoi J., Yoshida T., Todaka D., Ito Y., Maruyama K., Shinozaki K., Yamaguchi-Shinozaki K. Comprehensive analysis of rice DREB2-type genes that encode transcription factors involved in the expression of abiotic stress-responsive genes. Mol Genet Genomics. 2010;283(2):185-196. doi 10.1007/s00438-009-0506-y</mixed-citation><mixed-citation xml:lang="en">Matsukura S., Mizoi J., Yoshida T., Todaka D., Ito Y., Maruyama K., Shinozaki K., Yamaguchi-Shinozaki K. Comprehensive analysis of rice DREB2-type genes that encode transcription factors involved in the expression of abiotic stress-responsive genes. Mol Genet Genomics. 2010;283(2):185-196. doi 10.1007/s00438-009-0506-y</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Shi Y., Ding Y., Yang S. Molecular regulation of CBF signaling in cold acclimation. Trends Plant Sci. 2018;23(7):623-637. doi 10.1016/j.tplants.2018.04.002</mixed-citation><mixed-citation xml:lang="en">Shi Y., Ding Y., Yang S. Molecular regulation of CBF signaling in cold acclimation. Trends Plant Sci. 2018;23(7):623-637. doi 10.1016/j.tplants.2018.04.002</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Sun N., Sun X., Zhou J., Zhou X., Gao Z., Zhu X., Xu X., Liu Y., Li D., Zhan R., Wang L., Zhang H. Genome-wide characterization of pep per DREB family members and biological function of CaDREB32 in response to salt and osmotic stresses. Plant Physiol Biochem. 2025;222:109736. doi 10.1016/j.plaphy.2025.109736</mixed-citation><mixed-citation xml:lang="en">Sun N., Sun X., Zhou J., Zhou X., Gao Z., Zhu X., Xu X., Liu Y., Li D., Zhan R., Wang L., Zhang H. Genome-wide characterization of pep per DREB family members and biological function of CaDREB32 in response to salt and osmotic stresses. Plant Physiol Biochem. 2025;222:109736. doi 10.1016/j.plaphy.2025.109736</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Villagómez-Aranda A.L., Feregrino-Pérez A.A., García-Ortega L.F., González-Chavira M.M., Torres-Pacheco I., Guevara-González R.G. Activating stress memory: eustressors as potential tools for plant breeding. Plant Cell Rep. 2022;41(7):1481-1498. doi 10.1007/s00299-022-02858-x</mixed-citation><mixed-citation xml:lang="en">Villagómez-Aranda A.L., Feregrino-Pérez A.A., García-Ortega L.F., González-Chavira M.M., Torres-Pacheco I., Guevara-González R.G. Activating stress memory: eustressors as potential tools for plant breeding. Plant Cell Rep. 2022;41(7):1481-1498. doi 10.1007/s00299-022-02858-x</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Virlouvet L., Avenson T.J., Du Q., Zhang C., Liu N., Fromm M., Avramova Z., Russo S.E. Dehydration stress memory: gene networks linked to physiological responses during repeated stresses of Zea mays. Front Plant Sci. 2018;9:1058. doi 10.3389/fpls.2018.01058</mixed-citation><mixed-citation xml:lang="en">Virlouvet L., Avenson T.J., Du Q., Zhang C., Liu N., Fromm M., Avramova Z., Russo S.E. Dehydration stress memory: gene networks linked to physiological responses during repeated stresses of Zea mays. Front Plant Sci. 2018;9:1058. doi 10.3389/fpls.2018.01058</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Zhu Z., Dai Y., Yu G., Zhang X., Chen Q., Kou X., Mehareb E.M., Raza G., Zhang B., Wang B., Wang K., Han J. Dynamic physiological and transcriptomic changes reveal memory effects of salt stress in maize. BMC Genomics. 2023;24(1):726. doi 10.1186/s12864023-09845-w</mixed-citation><mixed-citation xml:lang="en">Zhu Z., Dai Y., Yu G., Zhang X., Chen Q., Kou X., Mehareb E.M., Raza G., Zhang B., Wang B., Wang K., Han J. Dynamic physiological and transcriptomic changes reveal memory effects of salt stress in maize. BMC Genomics. 2023;24(1):726. doi 10.1186/s12864023-09845-w</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Zuo D.D., Ahammed G.J., Guo D.L. Plant transcriptional memory and associated mechanism of abiotic stress tolerance. Plant Physiol Bio chem. 2023;201:107917. doi 10.1016/j.plaphy.2023.107917</mixed-citation><mixed-citation xml:lang="en">Zuo D.D., Ahammed G.J., Guo D.L. Plant transcriptional memory and associated mechanism of abiotic stress tolerance. Plant Physiol Bio chem. 2023;201:107917. doi 10.1016/j.plaphy.2023.107917</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>
