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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">vir-nw</journal-id><journal-title-group><journal-title xml:lang="ru">Труды по прикладной ботанике, генетике и селекции</journal-title><trans-title-group xml:lang="en"><trans-title>Proceedings on applied botany, genetics and breeding</trans-title></trans-title-group></journal-title-group><issn pub-type="ppub">2227-8834</issn><issn pub-type="epub">2619-0982</issn><publisher><publisher-name>N.I. Vavilov All-Russian Institute of Plant Genetic Resources</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="doi">10.30901/2227-8834-2025-1-242-256</article-id><article-id custom-type="elpub" pub-id-type="custom">vir-nw-2266</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>ОБЗОРЫ</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>SURVEYS</subject></subj-group></article-categories><title-group><article-title>Ген f3’5’h (d) розовой окраски цветка льна – представитель семейства CYP75 класса генов CYP450: от функции класса до функции гена</article-title><trans-title-group xml:lang="en"><trans-title>The f3’5’h (d) gene for pink coloring of flax flowers is a member of the CYP75 family within the CYP450 gene class: from class function to gene function</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-2333-1980</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Слободкина</surname><given-names>А. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Slobodkina</surname><given-names>A. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Анастасия Александровна Слободкина, младший научный сотрудник</p><p>190000, Санкт-Петербург, ул. Б. Морская, 42, 44</p></bio><bio xml:lang="en"><p>Anastasia А. Slobodkina, Associate Researcher</p><p>42, 44 Bolshaya Morskaya Street, St. Petersburg 190000</p></bio><email xlink:type="simple">a.slobodkina@vir.nw.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/0000-0002-5098-4904</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Павлов</surname><given-names>А. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Pavlov</surname><given-names>A. V.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Андрей Валерьевич Павлов, кандидат сельскохозяйственных наук, старший научный сотрудник</p><p>190000, Санкт-Петербург, ул. Б. Морская, 42, 44</p></bio><bio xml:lang="en"><p>Andrey V. Pavlov, Cand. Sci. (Agriculture), Senior Researcher</p><p>42, 44 Bolshaya Morskaya Street, St. Petersburg 190000</p></bio><email xlink:type="simple">avpavlov77@yandex.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/0000-0002-8328-9684</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Пороховинова</surname><given-names>Е. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Porokhovinova</surname><given-names>E. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Елизавета Александровна Пороховинова, доктор биологических наук, ведущий научный сотрудник</p><p>190000, Санкт-Петербург, ул. Б. Морская, 42, 44</p></bio><bio xml:lang="en"><p>Elizaveta A. Porokhovinova, Dr. Sci. (Biology), Leading Researcher</p><p>42, 44 Bolshaya Morskaya Street, St. Petersburg 190000</p></bio><email xlink:type="simple">e.porohovinova@mail.ru</email><xref ref-type="aff" rid="aff-1"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru"><institution>Федеральный исследовательский центр Всероссийский институт генетических ресурсов растений имени Н.И. Вавилова</institution><country>Россия</country></aff><aff xml:lang="en"><institution>N.I. Vavilov All-Russian AllRussian Institute of Plant Genetic Resources</institution><country>Russian Federation</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2025</year></pub-date><pub-date pub-type="epub"><day>13</day><month>04</month><year>2025</year></pub-date><volume>186</volume><issue>1</issue><fpage>242</fpage><lpage>256</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Слободкина А.А., Павлов А.В., Пороховинова Е.А., 2025</copyright-statement><copyright-year>2025</copyright-year><copyright-holder xml:lang="ru">Слободкина А.А., Павлов А.В., Пороховинова Е.А.</copyright-holder><copyright-holder xml:lang="en">Slobodkina A.A., Pavlov A.V., Porokhovinova E.A.</copyright-holder><license xml:lang="ru" license-type="creative-commons-attribution" xlink:href="https://creativecommons.org/licenses/by/4.0/" xlink:type="simple"><license-p>Данная работа распространяется под лицензией Creative Commons Attribution 4.0.</license-p></license><license xml:lang="en" 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://elpub.vir.nw.ru/jour/article/view/2266">https://elpub.vir.nw.ru/jour/article/view/2266</self-uri><abstract><p>В публикации охарактеризованы флавоноиды – пигменты растений, их роль, классификация, биосинтез. Основное внимание уделено ферментам класса цитохром P450, которые помимо участия в биосинтезе флавоноидов играют большую роль во множестве других процессов, таких как биосинтез полимеров, метаболизм гормонов, защита от неблагоприятных факторов. Представлены различные сведения о гене F3’5’H, кодирующем флавоноид-3’,5’-гидроксилазу, освещаются ключевые достижения и основные проблемы его изучения. Говорится о роли данного гена в биосинтезе антоцианов, показана принадлежность его продукта к семейству ферментов CYP75 клана CYP71 цитохромов P450, показаны активные центры фермента.</p><p>Лен обычно имеет голубой венчик и дельфинидин в качестве основного пигмента его окраски. В данной публикации охарактеризован ген d (f3’5’h), в рецессивном состоянии обуславливающий розовую окраску цветка и пеларгонидин в качестве основного пигмента. Показано, что F3’5’H льна относится к ферментам CYP75, но в отличие от большинства других принадлежит к подсемейству CYP75B. Показана возможность множественного аллелизма F3’5’H, приводящего к разным фенотипам, что подтверждает уже известный на формально-генетическом уровне множественный аллелизм гена d у льна. На данный момент лен – единственный вид, у которого проявление F3’5’H-активности шло в ущерб F3’H – возможно, потому что лен филогенетически далек от других древних культурных растений и эволюция этого гена связана с селекцией на желтосемянность.</p></abstract><trans-abstract xml:lang="en"><p>The publication characterizes flavonoids – plant pigments, their role, classification, and biosynthesis. Attention is mainly paid to the enzymes of the cytochrome P450 class, which, in addition to their participation in flavonoid biosynthesis, play a major role in many other processes, such as polymer biosynthesis, hormone metabolism, and defense against unfavorable factors. Various information about the flavonoid 3’,5’-hydroxylase (F3’5’H) gene is presented, highlighting key advances and major problems in its study. The role of this gene in anthocyanin biosynthesis is discussed, its product is shown to belong to the CYP75 enzyme family from the CYP71 clan of cytochromes P450, and the active centers of the enzyme are displayed.</p><p>Flax usually has a blue corolla, and delphinidin as the main pigment in its coloration. In this publication, we characterized the d gene (f3’5’h), which in recessive condition causes pink flower coloration and pelargonidin as the main pigment. It was shown that F3’5’H in flax belongs to CYP75 enzymes, but unlike most others is attributed to the CYP75B subfamily rather than CYP75A. The possibility of multiple allelism of F3’5’H leading to different phenotypes was demonstrated, which confirms the multiple allelism of the d gene in flax, already known at the formal genetic level. At present, flax is the only species wherein the emergence of F3’5’H activity went to the detriment of F3’H, probably because flax is phylogenetically distant from other ancient cultivated plants and its evolution is associated with selection for yellow seed color.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>Linum usitatissimum L.</kwd><kwd>флавоноиды</kwd><kwd>биосинтез антоцианов</kwd><kwd>декоративный лен</kwd><kwd>flavonoid 3’</kwd><kwd>5’-hydroxylase (F3’5’H)</kwd><kwd>цитохром P450</kwd><kwd>CYP75</kwd></kwd-group><kwd-group xml:lang="en"><kwd>Linum usitatissimum L.</kwd><kwd>flavonoids</kwd><kwd>anthocyanin biosynthesis</kwd><kwd>ornamental flax</kwd><kwd>flavonoid 3’</kwd><kwd>5’-hydroxylase (F3’5’H)</kwd><kwd>cytochrome P450</kwd><kwd>CYP75</kwd></kwd-group><funding-group><funding-statement xml:lang="ru">Работа выполнена в рамках государственного задания ВИР согласно тематическому плану НИР (№ FGEM-2025-0008 «Разработка подходов ускоренной селекции для улучшения хозяйственно ценных признаков декоративных и ягодных культур»).</funding-statement><funding-statement xml:lang="en">The work was carried out within the framework of the state task assigned to VIR according to the theme plan of research (No. FGEM-2025-0008 “Development of accelerated breeding approaches to the improvement of valuable agronomic characters in ornamental and berry crops”).</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">Babu P.R., Rao K.V., Reddy V.D. Structural organization and classification of cytochrome P450 genes in flax (Linum usitatissimum L.). Gene. 2013;513(1):156-162. DOI: 10.1016/j.gene.2012.10.040</mixed-citation><mixed-citation xml:lang="en">Babu P.R., Rao K.V., Reddy V.D. Structural organization and classification of cytochrome P450 genes in flax (Linum usitatissimum L.). Gene. 2013;513(1):156-162. DOI: 10.1016/j.gene.2012.10.040</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Bak S., Beisson F., Bishop G., Hamberger B., Höfer R., Paquette S. et al. Cytochromes P450. The Arabidopsis Book. 2011;9:e0144. DOI: 10.1199/tab.0144</mixed-citation><mixed-citation xml:lang="en">Bak S., Beisson F., Bishop G., Hamberger B., Höfer R., Paquette S. et al. Cytochromes P450. The Arabidopsis Book. 2011;9:e0144. DOI: 10.1199/tab.0144</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Berková V., Berka M., Griga M., Kopecká R., Prokopová M., Luklová M. et al. Molecular mechanisms underlying flax (Linum usitatissimum L.) tolerance to cadmium: a case study of proteome and metabolome of four different flax genotypes. Plants. 2022;11(21):2931. DOI: 10.3390/plants11212931</mixed-citation><mixed-citation xml:lang="en">Berková V., Berka M., Griga M., Kopecká R., Prokopová M., Luklová M. et al. Molecular mechanisms underlying flax (Linum usitatissimum L.) tolerance to cadmium: a case study of proteome and metabolome of four different flax genotypes. Plants. 2022;11(21):2931. DOI: 10.3390/plants11212931</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Бриттон Г. Биохимия природных пигментов. Москва: Мир; 1986.</mixed-citation><mixed-citation xml:lang="en">Britton G. The biochemistry of natural pigments. Moscow: Mir; 1986. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Dubois J., Harborne J., Bablom B., Plonka F. The inheritance of flower colors and anthocyanins in flax (Linum usitatissimum L.). Annales de l’amélioration de plantes. 1979;29(3):267-276.</mixed-citation><mixed-citation xml:lang="en">Dubois J., Harborne J., Bablom B., Plonka F. The inheritance of flower colors and anthocyanins in flax (Linum usitatissimum L.). Annales de l’amélioration de plantes. 1979;29(3):267-276.</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Dubois J.A., Harborne J.B. Anthocyanin inheritance in petals of flax, Linum usitatissimum. Phytochemistry. 1975;14:24912494.</mixed-citation><mixed-citation xml:lang="en">Dubois J.A., Harborne J.B. Anthocyanin inheritance in petals of flax, Linum usitatissimum. Phytochemistry. 1975;14:2491-2494.</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Эллади Е.В. Linum usitatissimum L. Vav. consp. nov. – Лён. В кн.: Культурная Флора СССР. Т. 6. Прядильные /под ред. Е.В. Вульфа. Москва; Ленинград; 1940. С.109-208.</mixed-citation><mixed-citation xml:lang="en">Elladi E.V. Linum usitatissimum L. Vav. consp. nov. – Flax (Len). In: E.V. Vulf (ed.). Flora of Cultivated Plants. Vol. 6. Fiber Crops (Pryadilnye). Moscow; Leningrad; 1940. р.109-208. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">FAOSTAT. Food and Agriculture Organization of the United Nations. Food and Agriculture Data: [website]. Available from: https://www.fao.org/faostat/en/#data/QCL [accessed Oct. 23, 2024].</mixed-citation><mixed-citation xml:lang="en">FAOSTAT. Food and Agriculture Organization of the United Nations. Food and Agriculture Data: [website]. Available from: https://www.fao.org/faostat/en/#data/QCL [accessed Oct. 23, 2024].</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Fujiwara Y., Nishiyama S., Onoue N., Matsuzaki R., Yonemori K., Tao R. Candidate gene analysis for ASTRINGENCY controlling fruit astringency in Diospyros kaki based on mRNAand small RNA-sequencing analyses. Acta Horticulturae. 2022;1338:269-276. DOI: 10.17660/ActaHortic.2022.1338.39</mixed-citation><mixed-citation xml:lang="en">Fujiwara Y., Nishiyama S., Onoue N., Matsuzaki R., Yonemori K., Tao R. Candidate gene analysis for ASTRINGENCY controlling fruit astringency in Diospyros kaki based on mRNAand small RNA-sequencing analyses. Acta Horticulturae. 2022;1338:269-276. DOI: 10.17660/ActaHortic.2022.1338.39</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Holton T.A., Cornish E.C. Genetics and biochemistry of anthocyanin biosynthesis. The Plant Cell. 1995;7(7):1071-1083. DOI: 10.1105/tpc.7.7.1071</mixed-citation><mixed-citation xml:lang="en">Holton T.A., Cornish E.C. Genetics and biochemistry of anthocyanin biosynthesis. The Plant Cell. 1995;7(7):1071-1083. DOI: 10.1105/tpc.7.7.1071</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Irmisch S., Ruebsam H., Jancsik S., Man Saint Yuen M., Madilao L.L., Bohlmann J. Flavonol biosynthesis genes and their use in engineering the plant antidiabetic metabolite montbretin A. Plant Physiology. 2019;180(3):1277-1290. DOI: 10.1104/pp.19.00254</mixed-citation><mixed-citation xml:lang="en">Irmisch S., Ruebsam H., Jancsik S., Man Saint Yuen M., Madilao L.L., Bohlmann J. Flavonol biosynthesis genes and their use in engineering the plant antidiabetic metabolite montbretin A. Plant Physiology. 2019;180(3):1277-1290. DOI: 10.1104/pp.19.00254</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Kitamura S. Transport of flavonoids: from cytosolic synthesis to vacuolar accumulation. In: E. Grotewold (ed.). The Science of Flavonoids. New York, NY: Springer; 2008. p.123-146.</mixed-citation><mixed-citation xml:lang="en">Kitamura S. Transport of flavonoids: from cytosolic synthesis to vacuolar accumulation. In: E. Grotewold (ed.). The Science of Flavonoids. New York, NY: Springer; 2008. p.123-146.</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Кольман Я., Рём К.Г. Система цитохрома Р450. В кн.: Я. Кольман, К.Г. Рём. Наглядная биохимия. Москва: Мир; 2004. С.310-311. URL: https://www.chem.msu.ru/rus/teaching/kolman/310.htm [дата обращения: 01.09.2024].</mixed-citation><mixed-citation xml:lang="en">Koolman J., Roehm K.H. Cytochrome P450 system. In: J. Koolman, K.H. Roehm. Taschenatlas der Biochemie. Moscow: Mir; 2004. p.310-311. [in Russian]. URL: https://www.chem.msu.ru/rus/teaching/kolman/310.htm [дата обращения: 01.09.2024].</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Lam P.Y., Liu H., Lo C. Completion of tricin biosynthesis pathway in rice: cytochrome P450 75b4 is a unique chrysoeriol 5’-hydroxylase. Plant Physiology. 2015;168(4):1527-1536. DOI: 10.1104/pp.15.00566</mixed-citation><mixed-citation xml:lang="en">Lam P.Y., Liu H., Lo C. Completion of tricin biosynthesis pathway in rice: cytochrome P450 75b4 is a unique chrysoeriol 5’-hydroxylase. Plant Physiology. 2015;168(4):1527-1536. DOI: 10.1104/pp.15.00566</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Liang C.Y., Rengasamy K.P., Huang L.M., Hsu C.C., Jeng M.F., Chen W.H. et al. Assessment of violet-blue color formation in Phalaenopsis orchids. BMC Plant Biology. 2020;20(1):212. DOI: 10.1186/s12870-020-02402-7</mixed-citation><mixed-citation xml:lang="en">Liang C.Y., Rengasamy K.P., Huang L.M., Hsu C.C., Jeng M.F., Chen W.H. et al. Assessment of violet-blue color formation in Phalaenopsis orchids. BMC Plant Biology. 2020;20(1):212. DOI: 10.1186/s12870-020-02402-7</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">McClean P., Lee R., Howe K., Osborne C., Grimwood J., Levy S. et al. The common bean V gene encodes Flavonoid 3′5′ Hydroxylase: a major mutational target for flavonoid diversity in angiosperm. Frontiers in Plant Science. 2022;13:869582. DOI: 10.3389/fpls.2022.869582</mixed-citation><mixed-citation xml:lang="en">McClean P., Lee R., Howe K., Osborne C., Grimwood J., Levy S. et al. The common bean V gene encodes Flavonoid 3′5′ Hydroxylase: a major mutational target for flavonoid diversity in angiosperm. Frontiers in Plant Science. 2022;13:869582. DOI: 10.3389/fpls.2022.869582</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Minerdi D., Savoi S., Sabbatini P. Role of cytochrome P450 enzyme in plant microorganisms’ communication: a focus on grapevine. International Journal of Molecular Sciences. 2023;24(5):4695. DOI: 10.3390/ijms24054695</mixed-citation><mixed-citation xml:lang="en">Minerdi D., Savoi S., Sabbatini P. Role of cytochrome P450 enzyme in plant microorganisms’ communication: a focus on grapevine. International Journal of Molecular Sciences. 2023;24(5):4695. DOI: 10.3390/ijms24054695</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Миневич И.Э. Научное обоснование и разработка научно-практических основ технологий глубокой переработки семян льна с получением ингредиентов для создания продуктов здорового питания: дис. ... докт. техн. наук. Тверь: ФНЦ лубяных культур; 2022.</mixed-citation><mixed-citation xml:lang="en">Миневич И.Э. Научное обоснование и разработка научно-практических основ технологий глубокой переработки семян льна с получением ингредиентов для создания продуктов здорового питания: дис. ... докт. техн. наук. Тверь: ФНЦ лубяных культур; 2022.</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">NCBI. National Center for Biotechnology Information: [website]. Available from: https://www.ncbi.nlm.nih.gov [accessed June 20, 2024].</mixed-citation><mixed-citation xml:lang="en">NCBI. National Center for Biotechnology Information: [website]. Available from: https://www.ncbi.nlm.nih.gov [accessed June 20, 2024].</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Nelson D.R. Cytochrome P450 diversity in the tree of life. Biochimica et Biophysica Acta. Proteins and Proteomics. 2018;1866(1):141-154. DOI: 10.1016/j.bbapap.2017.05.003</mixed-citation><mixed-citation xml:lang="en">Nelson D.R. Cytochrome P450 diversity in the tree of life. Biochimica et Biophysica Acta. Proteins and Proteomics. 2018;1866(1):141-154. DOI: 10.1016/j.bbapap.2017.05.003</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Olsen K.M., Hehn A., Jugdé H., Slimestad R., Larbat R., Bourgaud F. et al. Identification and characterisation of CYP75A31, a new flavonoid 3’5’-hydroxylase, isolated from Solanum lycopersicum. BMC Plant Biology. 2010;10:21. DOI: 10.1186/1471-2229-10-21</mixed-citation><mixed-citation xml:lang="en">Olsen K.M., Hehn A., Jugdé H., Slimestad R., Larbat R., Bourgaud F. et al. Identification and characterisation of CYP75A31, a new flavonoid 3’5’-hydroxylase, isolated from Solanum lycopersicum. BMC Plant Biology. 2010;10:21. DOI: 10.1186/1471-2229-10-21</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Phytozome 13. The Plant Genomics Resource: [website]. Available from: https://phytozome-next.jgi.doe.gov [accessed June 20, 2024].</mixed-citation><mixed-citation xml:lang="en">Phytozome 13. The Plant Genomics Resource: [website]. Available from: https://phytozome-next.jgi.doe.gov [accessed June 20, 2024].</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Пороховинова Е.А. Создание и изучение генетической коллекции льна: Linum usitatissimum L.: дис. ... канд. биол. наук. Санкт-Петербург: ВИР; 2002.</mixed-citation><mixed-citation xml:lang="en">Porokhovinova E.A. Building up and studying the genetic collection of flax: Linum usitatissimum L. (Sozdaniye i izucheniye geneticheskoy kollektsii lna: Linum usitatissimum L.) [dissertation]. St. Petersburg: VIR; 2002. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Пороховинова Е.А. Генетическая коллекция льна (Linum usitatissimum L.): создание, анализ и перспективы использования дис. ... докт. биол. наук. Санкт-Петербург: ВИР; 2019.</mixed-citation><mixed-citation xml:lang="en">Porokhovinova E.A. Genetic collection of flax (Linum usitatissimum L.): establishment, analysis, and prospects of utilization (Geneticheskaya kollektsiya lna (Linum usitatissimum L.): sozdaniye, analiz i perspectivy ispolzovanya) [dissertation]. St. Petersburg: VIR; 2019. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Пороховинова Е.А., Кутузова С.Н., Павлов А.В., Бузовкина И.С., Брач Н.Б. Разнообразие морфологических признаков льна в генетической коллекции ВИР как результат его доместикации. Экологическая генетика. 2018;16(4):33-50. DOI: 10.17816/ecogen16433-50</mixed-citation><mixed-citation xml:lang="en">Porokhovinova E.A., Kutuzova S.N., Pavlov A.V., Buzovkina I.S., Brutch N.B. Diversity of flax morphological characters in VIR genetic collection as a result of crop domestication. Ecological Genetics. 2018;16(4):33-50. [in Russian]. DOI: 10.17816/ecogen16433-50</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Пороховинова Е.А., Шеленга Т.В., Матвеева Т.В., Павлов А.В., Григорьева Е.А., Брач Н.Б. Полиморфизм генов, контролирующих низкое содержание линоленовой кислоты, у линий генетической коллекции льна ВИР. Экологическая генетика. 2019;17(2):519. DOI: 10.17816/ecogen1725-19</mixed-citation><mixed-citation xml:lang="en">Porokhovinova E.A., Shelenga T.V., Matveeva T.V., Pavlov A.V., Grigorieva E.A., Brutch N.B. Polymorphism of genes controlling low level of linolenic acid in lines from VIR flax genetic collection. Ecological Genetics. 2019;17(2):5-19. [in Russian] . DOI: 10.17816/ecogen1725-19</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Pourcel L., Routaboul J.M., Cheynier V., Lepiniec L., Debeaujon I. Flavonoid oxidation in plants: from biochemical properties to physiological functions. Trends in Plant Science. 2007;12(1):29-36. DOI: 10.1016/j.tplants.2006.11.006</mixed-citation><mixed-citation xml:lang="en">Pourcel L., Routaboul J.M., Cheynier V., Lepiniec L., Debeaujon I. Flavonoid oxidation in plants: from biochemical properties to physiological functions. Trends in Plant Science. 2007;12(1):29-36. DOI: 10.1016/j.tplants.2006.11.006</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Рахмангулов Р.С. Применение системы CRISPR/Cas для редактирования генов декоративных культур. Биотехнология и селекция растений. 2022;5(3):33-41. DOI: 10.30901/2658-62662022-3-o1</mixed-citation><mixed-citation xml:lang="en">Rakhmangulov R.S. Application of the CRISPR/Cas system for gene editing in ornamental crops. Plant Biotechnology and Breeding. 2022;5(3):33-41. [in Russian]. DOI: 10.30901/2658-62662022-3-o1</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Рахмангулов Р.С., Барабанов И.В., Ерастенкова М.В., Иванов А.А., Коваленко Т.В., Межина К.М. и др. Новые направления в генетике, селекции, биотехнологии декоративных и ягодных культур в ВИР им. Н.И. Вавилова. Биотехнология и селекция растений. 2022;5(4):65-78. DOI: 10.30901/2658-6266-2022-4-o3</mixed-citation><mixed-citation xml:lang="en">Rakhmangulov R.S., Barabanov I.V., Erastenkova M.V., Ivanov A.A., Kovalenko T.V., Mezhina K.M. et al. The new directions in genetics, breeding and biotechnology of ornamental and berry crops in the N.I. Vavilov Institute of Plant Genetic Resources (VIR). Plant Biotechnology and Breeding. 2022;5(4):65-78. [in Russian]. DOI: 10.30901/2658-6266-2022-4-o3</mixed-citation></citation-alternatives></ref><ref id="cit30"><label>30</label><citation-alternatives><mixed-citation xml:lang="ru">Рахмангулов Р.С., Тихонова Н.Г. Селекция декоративных растений в России. Биотехнология и селекция растений. 2021;4(4):40-54. DOI: 10.30901/2658-6266-2021-4-o4</mixed-citation><mixed-citation xml:lang="en">Rakhmangulov R.S., Tikhonova N.G. Breeding of ornamental plants in Russia. Plant Biotechnology and Breeding. 2021;4(4):40-54. [in Russian]. DOI: 10.30901/2658-6266-2021-4-o4</mixed-citation></citation-alternatives></ref><ref id="cit31"><label>31</label><citation-alternatives><mixed-citation xml:lang="ru">Rupasinghe S., Baudry J., Schuler M.A. Common active site architecture and binding strategy of four phenylpropanoid P450s from Arabidopsis thaliana as revealed by molecular modeling. Protein Engineering. 2003;16(10):721-731. DOI: 10.1093/protein/gzg094</mixed-citation><mixed-citation xml:lang="en">Rupasinghe S., Baudry J., Schuler M.A. Common active site architecture and binding strategy of four phenylpropanoid P450s from Arabidopsis thaliana as revealed by molecular modeling. Protein Engineering. 2003;16(10):721-731. DOI: 10.1093/protein/gzg094</mixed-citation></citation-alternatives></ref><ref id="cit32"><label>32</label><citation-alternatives><mixed-citation xml:lang="ru">Sato M., Kawabe T., Hosokawa M., Tatsuzawa F., Doi M. Tissue culture-induced flower-color changes in saintpaulia caused by excision of the transposon inserted in the flavonoid 3’, 5’ hydroxylase (F3’5’H) promoter. Plant Cell Reports. 2011;30(5):929-939. DOI: 10.1007/s00299-011-1016-z</mixed-citation><mixed-citation xml:lang="en">Sato M., Kawabe T., Hosokawa M., Tatsuzawa F., Doi M. Tissue culture-induced flower-color changes in saintpaulia caused by excision of the transposon inserted in the flavonoid 3’, 5’ hydroxylase (F3’5’H) promoter. Plant Cell Reports. 2011;30(5):929-939. DOI: 10.1007/s00299-011-1016-z</mixed-citation></citation-alternatives></ref><ref id="cit33"><label>33</label><citation-alternatives><mixed-citation xml:lang="ru">Seitz C., Ameres S., Forkmann G. Identification of the molecular basis for the functional difference between flavonoid 3’-hydroxylase and flavonoid 3’5’-hydroxylase. FEBS Letters. 2007;581(18):3429-3434. DOI: 10.1016/j.febslet.2007.06.045</mixed-citation><mixed-citation xml:lang="en">Seitz C., Ameres S., Forkmann G. Identification of the molecular basis for the functional difference between flavonoid 3’-hydroxylase and flavonoid 3’5’-hydroxylase. FEBS Letters. 2007;581(18):3429-3434. DOI: 10.1016/j.febslet.2007.06.045</mixed-citation></citation-alternatives></ref><ref id="cit34"><label>34</label><citation-alternatives><mixed-citation xml:lang="ru">Seitz C., Ameres S., Schlangen K., Forkmann G., Halbwirth H. Multiple evolution of flavonoid 3’,5’-hydroxylase. Planta. 2015;242(3):561-573. DOI: 10.1007/s00425-015-2293-5</mixed-citation><mixed-citation xml:lang="en">Seitz C., Ameres S., Schlangen K., Forkmann G., Halbwirth H. Multiple evolution of flavonoid 3’,5’-hydroxylase. Planta. 2015;242(3):561-573. DOI: 10.1007/s00425-015-2293-5</mixed-citation></citation-alternatives></ref><ref id="cit35"><label>35</label><citation-alternatives><mixed-citation xml:lang="ru">Шарова Е.И. Антиоксиданты растений. Санкт-Петербург: Санкт-Петербургский государственный университет; 2016.</mixed-citation><mixed-citation xml:lang="en">Sharova E.I. Plant antioxidants. St. Petersburg: St. Petersburg State University; 2016. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit36"><label>36</label><citation-alternatives><mixed-citation xml:lang="ru">Государственная комиссия Российской Федерации по испытанию и охране селекционных достижений: [сайт]. URL: https://gossortrf.ru [дата обращения: 07.08.2024].</mixed-citation><mixed-citation xml:lang="en">State Commission of the Russian Federation for Testing and Protecting Selection Achievements: [website]. [in Russian] (Государственная комиссия Российской Федерации по испытанию и охране селекционных достижений: [сайт]). URL: https://gossortrf.ru [дата обращения: 07.08.2024].</mixed-citation></citation-alternatives></ref><ref id="cit37"><label>37</label><citation-alternatives><mixed-citation xml:lang="ru">Стрыгина К.В., Хлесткина Е.К. Синтез антоцианов у картофеля (Solanum tuberosum L.): генетические маркеры для направленного отбора (обзор). Сельскохозяйственная биология. 2017;52(1):37-49. DOI: 10.15389/agrobiology.2017.1.37rus</mixed-citation><mixed-citation xml:lang="en">Strygina K.V., Khlestkina E.K. Anthocyanins synthesis in potato (Solanum tuberosum L.): genetic markers for smart breeding (review). Agricultural Biology. 2017;52(1):37-49. [in Russian]. DOI: 10.15389/agrobiology.2017.1.37rus</mixed-citation></citation-alternatives></ref><ref id="cit38"><label>38</label><citation-alternatives><mixed-citation xml:lang="ru">Sudarshan G. P., Kulkarni M., Akhov L., Ashe P., Shaterian H., Cloutier S. et al. QTL mapping and molecular characterization of the classical D locus controlling seed and flower color in Linum usitatissimum (flax). Scientific Reports. 2017;7(1):15751. DOI: 10.1038/s41598-017-11565-7</mixed-citation><mixed-citation xml:lang="en">Sudarshan G. P., Kulkarni M., Akhov L., Ashe P., Shaterian H., Cloutier S. et al. QTL mapping and molecular characterization of the classical D locus controlling seed and flower color in Linum usitatissimum (flax). Scientific Reports. 2017;7(1):15751. DOI: 10.1038/s41598-017-11565-7</mixed-citation></citation-alternatives></ref><ref id="cit39"><label>39</label><citation-alternatives><mixed-citation xml:lang="ru">Takahashi R., Dubouzet J.G., Matsumura H., Yasuda K., Iwashina T. A new allele of flower color gene W1 encoding flavonoid 3’5’-hydroxylase is responsible for light purple flowers in wild soybean Glycine soja. BMC Plant Biology. 2010;10:155. DOI: 10.1186/1471-2229-10-155</mixed-citation><mixed-citation xml:lang="en">Takahashi R., Dubouzet J.G., Matsumura H., Yasuda K., Iwashina T. A new allele of flower color gene W1 encoding flavonoid 3’5’-hydroxylase is responsible for light purple flowers in wild soybean Glycine soja. BMC Plant Biology. 2010;10:155. DOI: 10.1186/1471-2229-10-155</mixed-citation></citation-alternatives></ref><ref id="cit40"><label>40</label><citation-alternatives><mixed-citation xml:lang="ru">Tanaka Y., Yonekura K., Fukuchi-Mizutani M., Fukui Y., Fujiwara H., Ashikari T. et al. Molecular and biochemical characterization of three anthocyanin synthetic enzymes from Gentiana triflora. Plant and Cell Physiology. 1996;37(5):711-716. DOI: 10.1093/oxfordjournals.pcp.a029004</mixed-citation><mixed-citation xml:lang="en">Tanaka Y., Yonekura K., Fukuchi-Mizutani M., Fukui Y., Fujiwara H., Ashikari T. et al. Molecular and biochemical characterization of three anthocyanin synthetic enzymes from Gentiana triflora. Plant and Cell Physiology. 1996;37(5):711-716. DOI: 10.1093/oxfordjournals.pcp.a029004</mixed-citation></citation-alternatives></ref><ref id="cit41"><label>41</label><citation-alternatives><mixed-citation xml:lang="ru">Wang Y.S., Xu Y.J., Gao L.P., Yu O., Wang X.Z., He X.J. et al. Functional analysis of flavonoid 3’,5’-hydroxylase from tea plant (Camellia sinensis): critical role in the accumulation of catechins. BMC Plant Biology. 2014;14:347. DOI: 10.1186/s12870-014-0347-7</mixed-citation><mixed-citation xml:lang="en">Wang Y.S., Xu Y.J., Gao L.P., Yu O., Wang X.Z., He X.J. et al. Functional analysis of flavonoid 3’,5’-hydroxylase from tea plant (Camellia sinensis): critical role in the accumulation of catechins. BMC Plant Biology. 2014;14:347. DOI: 10.1186/s12870-014-0347-7</mixed-citation></citation-alternatives></ref><ref id="cit42"><label>42</label><citation-alternatives><mixed-citation xml:lang="ru">Williams C.A., Grayer R.J. Anthocyanins and other flavonoids. Natural Product Reports. 2004;21(4):539-573. DOI: 10.1039/B311404J</mixed-citation><mixed-citation xml:lang="en">Williams C.A., Grayer R.J. Anthocyanins and other flavonoids. Natural Product Reports. 2004;21(4):539-573. DOI: 10.1039/B311404J</mixed-citation></citation-alternatives></ref><ref id="cit43"><label>43</label><citation-alternatives><mixed-citation xml:lang="ru">Winkel B.S.J. The biosynthesis of flavonoids. In: E. Grotewold (ed.). The Science of Flavonoids. New York, NY: Springer; 2008. p.71-95. DOI: 10.1007/978-0-387-28822-2_3</mixed-citation><mixed-citation xml:lang="en">Winkel B.S.J. The biosynthesis of flavonoids. In: E. Grotewold (ed.). The Science of Flavonoids. New York, NY: Springer; 2008. p.71-95. DOI: 10.1007/978-0-387-28822-2_3</mixed-citation></citation-alternatives></ref><ref id="cit44"><label>44</label><citation-alternatives><mixed-citation xml:lang="ru">Xu J., Wang X., Guo W. The cytochrome P450 superfamily: key players in plant development and defense. Journal of Integrative Agriculture. 2015;14(9):1673-1686. DOI: 10.1016/S2095-3119(14)60980-1</mixed-citation><mixed-citation xml:lang="en">Xu J., Wang X., Guo W. The cytochrome P450 superfamily: key players in plant development and defense. Journal of Integrative Agriculture. 2015;14(9):1673-1686. DOI: 10.1016/S2095-3119(14)60980-1</mixed-citation></citation-alternatives></ref><ref id="cit45"><label>45</label><citation-alternatives><mixed-citation xml:lang="ru">Yoshida K., Mori M., Kondo T. Blue flower color development by anthocyanins: from chemical structure to cell physiology. Natural Product Reports. 2009;26(7):884-915. DOI: 10.1039/B800165K</mixed-citation><mixed-citation xml:lang="en">Yoshida K., Mori M., Kondo T. Blue flower color development by anthocyanins: from chemical structure to cell physiology. Natural Product Reports. 2009;26(7):884-915. DOI: 10.1039/B800165K</mixed-citation></citation-alternatives></ref><ref id="cit46"><label>46</label><citation-alternatives><mixed-citation xml:lang="ru">Zabala G., Vodkin L.O. A rearrangement resulting in small tandem repeats in the F3’5’H gene of white flower genotypes is associated with the soybean W1 locus. Crop Science. 2007;47(S2):113-124. DOI: 10.2135/cropsci2006.12.0838tpg</mixed-citation><mixed-citation xml:lang="en">Zabala G., Vodkin L.O. A rearrangement resulting in small tandem repeats in the F3’5’H gene of white flower genotypes is associated with the soybean W1 locus. Crop Science. 2007;47(S2):113-124. DOI: 10.2135/cropsci2006.12.0838tpg</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>
