<?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">biosel</journal-id><journal-title-group><journal-title xml:lang="ru">Биотехнология и селекция растений</journal-title><trans-title-group xml:lang="en"><trans-title>Plant Biotechnology and Breeding</trans-title></trans-title-group></journal-title-group><issn pub-type="ppub">2658-6266</issn><issn pub-type="epub">2658-6258</issn><publisher><publisher-name>VIR</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="doi">10.30901/2658-6266-2025-4-o2</article-id><article-id custom-type="elpub" pub-id-type="custom">biosel-288</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>ИЗУЧЕНИЕ ГЕНЕТИЧЕСКИХ РЕСУРСОВ РАСТЕНИЙ С ИСПОЛЬЗОВАНИЕМ МЕТОДОВ МОЛЕКУЛЯРНОЙ ГЕНЕТИКИ</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>STUDY OF PLANT GENETIC RESOURCES USING MOLECULAR GENETICS METHODS</subject></subj-group></article-categories><title-group><article-title>Особенности синтеза картамина в соцветиях сафлора красильного Carthamus tinctorius L.</article-title><trans-title-group xml:lang="en"><trans-title>Features of carthamin synthesis in inflorescences of safflower Carthamus tinctorius L.</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-9574-0356</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Бемова</surname><given-names>В. Д.</given-names></name><name name-style="western" xml:lang="en"><surname>Bemova</surname><given-names>V. D.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Виктория Дмитриевна Бемова, младший научный сотрудник, лаборатория генетики, селекции и биотехнологии декоративных и ягодных культур, ВИР</p><p>190000 Россия, Санкт-Петербург, ул. Большая Морская, 42, 44</p></bio><bio xml:lang="en"><p>Viktoria D. Bemova, Junior Researcher, Laboratory of Genetics, Breeding and Biotechnology of Ornamental and Berry Crops, VIR</p><p>42, 44, Bolshaya Morskaya Street, St. Petersburg, 190000 Russia</p></bio><email xlink:type="simple">viktoria.bemova@yandex.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></aff><aff xml:lang="en"><institution>N.I.  Vavilov All-Russian Institute of Plant Genetic Resources</institution></aff></aff-alternatives><pub-date pub-type="collection"><year>2025</year></pub-date><pub-date pub-type="epub"><day>30</day><month>12</month><year>2025</year></pub-date><volume>8</volume><issue>4</issue><fpage>127</fpage><lpage>135</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Бемова В.Д., 2025</copyright-statement><copyright-year>2025</copyright-year><copyright-holder xml:lang="ru">Бемова В.Д.</copyright-holder><copyright-holder xml:lang="en">Bemova V.D.</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://biosel.elpub.ru/jour/article/view/288">https://biosel.elpub.ru/jour/article/view/288</self-uri><abstract><p>Сафлор Carthamus tinctorius L., относящийся к семейству сложноцветных (Asteraceae), – важная масличная культура, его семена богаты жирными кислотами, в частности олеиновой и линолевой. Сафлор используется также в декоративных целях; на протяжении столетий его активно выращивали во многих странах мира. В последние годы особый интерес вызывают вторичные метаболиты, получаемые из соцветий сафлора, в частности флавоноиды. Флавоноиды сафлора можно разделить на две группы: специальные, представленные хинохалконами, и общие. Многие из этих веществ существенно влияют на окраску соцветий сафлора, которая изменяется в зависимости от стадии цветения (от желтой к оранжевой и красной при увядании). Флавоноиды сафлора активно используются в медицине и в качестве натуральных красителей при изготовлении тканей, косметики, а также в пищевой промышленности. Процесс биосинтеза пигментов в соцветиях до сих пор изучается, остаются неисследованными многие этапы, неизвестны механические аспекты их образования. Особый интерес представляет синтез красного пигмента – картамина, уникального димерного хинохалкона, добываемого только из оранжевых и красных соцветий сафлора. При правильном очищении этот пигмент приобретает металлический золотистый блеск. В исследовании 2021 года из соцветий сафлора были выделены белки картамин-синтазы (CarS), отвечающие за заключительный этап преобразования прекартамина в картамин. Гены CarS (CtPOD1, CtPOD2 и CtPOD3) экспрессируются в тканях сафлора независимо от окраски цветка. Предположительно, прекартамин накапливается в структурах венчика, которые физически отделены от клеточного компартмента, содержащего CarS. В ходе старения клетки соцветия сафлора разрушаются, что позволяет CarS взаимодействовать с прекартамином и образовывать картамин, который адсорбируется клеточной стенкой венчика и тем самым достигается стабилизация красной пигментации. В этом обзоре собраны данные об особенностях синтеза картамина, особенно о последнем этапе – преобразовании прекартамина в картамин и накопление его в соцветиях.</p></abstract><trans-abstract xml:lang="en"><p>Safflower Carthamus tinctorius L., a member of the Asteraceae family, is an important oilseed crop; its seeds are rich in fatty acids, particularly oleic and linoleic. Safflower is also used for ornamental purposes and has been extensively cultivated in many countries for centuries. In recent years, secondary metabolites obtained from safflower inflorescences, particularly flavonoids, have attracted particular interest. Safflower flavonoids can be divided into two groups: specialized flavonoids, represented by quinochalcones, and general ones. Many of these substances significantly influence the color of safflower inflorescences, which changes depending on the flowering stage (from yellow to orange and red at fading). Safflower flavonoids are widely used in medicine and as natural dyes in the manufacture of fabrics, cosmetics, and in the food industry. The process of pigment biosynthesis in inflorescences is still being studied; many stages remain unexplored, and the mechanical aspects of their formation are unknown. Of particular interest is the synthesis of the red pigment, carthamin, a unique dimeric quinochalcone extracted only from orange and red safflower inflorescences. When properly purified, this pigment acquires a metallic golden sheen. A 2021 study used safflower inflorescences for extracting carthamin synthase (CarS) proteins responsible for the final step in converting precarthamin to carthamin. The CarS genes (CtPOD1, CtPOD2, and CtPOD3) are expressed in safflower tissues regardless of flower color. Presumably, precarthamin accumulates in corolla cellular structures that are physically separated from the cellular compartment containing CarS. During floral senescence, cells degrade, allowing CarS to interact with precarthamin and form carthamin, which is adsorbed by the corolla cell wall, thereby stabilizing the red pigmentation. This review summarizes data on the specifics of carthamin synthesis, particularly the final step – the conversion of precarthamin to carthamin and its accumulation in inflorescences.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>картамин-синтаза</kwd><kwd>пигменты</kwd><kwd>флавоноиды</kwd><kwd>хинохалконы</kwd><kwd>CarS</kwd></kwd-group><kwd-group xml:lang="en"><kwd>carthamin synthase</kwd><kwd>pigments</kwd><kwd>flavonoids</kwd><kwd>quinochalcones</kwd><kwd>CarS</kwd></kwd-group><funding-group><funding-statement xml:lang="ru">Работа выполнена в рамках Государственного задания согласно тематическому плану ВИР по проекту № FGEM-2025-0008</funding-statement><funding-statement xml:lang="en">The research was performed within the framework of the State Assignment according to the theme plan of VIR, Project No. FGEM-2025-0008.</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">Azami K., Hayashi T., Kusumi T., Ohmori K., Suzuki K. Total synthesis of carthamin, a traditional natural red pigment. Angewandte Chemie International Edition. 2019;58: 5321-5326. DOI: 10.1002/anie.201900454.</mixed-citation><mixed-citation xml:lang="en">Azami K., Hayashi T., Kusumi T., Ohmori K., Suzuki K. Total synthesis of carthamin, a traditional natural red pigment. Angewandte Chemie International Edition. 2019;58: 5321-5326. DOI: 10.1002/anie.201900454.</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Davies K.M., Jibran R., Zhou Y., Albert N.W., Brummell D.A., Jordan B.R., Bowman J.L., Schwinn K.E. The evolution of flavonoid biosynthesis: a bryophyte perspective. Frontiers in Plant Science. 2020;11:7. DOI: 10.3389/fpls.2020.00007</mixed-citation><mixed-citation xml:lang="en">Davies K.M., Jibran R., Zhou Y., Albert N.W., Brummell D.A., Jordan B.R., Bowman J.L., Schwinn K.E. The evolution of flavonoid biosynthesis: a bryophyte perspective. Frontiers in Plant Science. 2020;11:7. DOI: 10.3389/fpls.2020.00007</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Guo D-D., Liu F., Tu Y-H., He B-X., Gao Y., Guo M-L. Expression patterns of three UGT genes in different chemotype safflower lines and under MeJA stimulus revealed their potential role in flavonoid biosynthesis. PLoS One. 2016;11(7):e0158159. DOI: 10.1371/journal.pone.0158159</mixed-citation><mixed-citation xml:lang="en">Guo D-D., Liu F., Tu Y-H., He B-X., Gao Y., Guo M-L. Expression patterns of three UGT genes in different chemotype safflower lines and under MeJA stimulus revealed their potential role in flavonoid biosynthesis. PLoS One. 2016;11(7):e0158159. DOI: 10.1371/journal.pone.0158159</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Gupta R.K., Singh S.B. Diallel analysis for seed yield, oil content and other economic traits in safflower (Carthamus tinctorius L.). Genetika-Yugoslavia. 1988;20:161-173.</mixed-citation><mixed-citation xml:lang="en">Gupta R.K., Singh S.B. Diallel analysis for seed yield, oil content and other economic traits in safflower (Carthamus tinctorius L.). Genetika-Yugoslavia. 1988;20:161-173.</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Hartman A. Inheritance of corolla color in safflower (Carthamus tinctorius L.). Davis: University of California. 1967;76:2.</mixed-citation><mixed-citation xml:lang="en">Hartman A. Inheritance of corolla color in safflower (Carthamus tinctorius L.). Davis: University of California. 1967;76:2.</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Kametaka T., Perkin A.G. CXXX. – Carthamine. Part I. Journal of the Chemical Society, Transactions. 1910;97:1415-1427. DOI: 10.1039/CT9109701415</mixed-citation><mixed-citation xml:lang="en">Kametaka T., Perkin A.G. CXXX. – Carthamine. Part I. Journal of the Chemical Society, Transactions. 1910;97:1415-1427. DOI: 10.1039/CT9109701415</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Kanehira T., Saito K. Decomposition of carthamin by peroxidases from Carthamus tinctorius. Biochemie und Physiologie der Pflanzen. 1990;186(3):179-187. DOI: 10.1016/S0015-3796(96)80006-0</mixed-citation><mixed-citation xml:lang="en">Kanehira T., Saito K. Decomposition of carthamin by peroxidases from Carthamus tinctorius. Biochemie und Physiologie der Pflanzen. 1990;186(3):179-187. DOI: 10.1016/S0015-3796(96)80006-0</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Knowles P.F. Variability in oleic and linoleic acid contents of safflower oil. Economic Botany. 1965;19:53-62. DOI: 10.1007/BF02971186</mixed-citation><mixed-citation xml:lang="en">Knowles P.F. Variability in oleic and linoleic acid contents of safflower oil. Economic Botany. 1965;19:53-62. DOI: 10.1007/BF02971186</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Kosoto H. The history of Chinese drugs recorded in the ‘‘Japanese Pharmacopoeia. Fifteenth Edition”. Japanese Society of Pharmacognosy. 2007;61:68-78. [in Japanese]</mixed-citation><mixed-citation xml:lang="en">Kosoto H. The history of Chinese drugs recorded in the ‘‘Japanese Pharmacopoeia. Fifteenth Edition”. Japanese Society of Pharmacognosy. 2007;61:68-78. [in Japanese]</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Kotecha A. Inheritance and association of six traits in safflower. Crop Science. 1979;19(4):523-527. DOI: 10.2135/cropsci1979.0011183X001900040022x</mixed-citation><mixed-citation xml:lang="en">Kotecha A. Inheritance and association of six traits in safflower. Crop Science. 1979;19(4):523-527. DOI: 10.2135/cropsci1979.0011183X001900040022x</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Kumazawa T., Amano Y., Haga T., Matsuba S., Sato S., Kawamoto K., Onodera J. Synthesis of model compounds of the precursor of carthamin, a colouring matter of safflower, and their conversion into carthamin-type compounds. Chemistry Letters. 1995;24(8):625-626.</mixed-citation><mixed-citation xml:lang="en">Kumazawa T., Amano Y., Haga T., Matsuba S., Sato S., Kawamoto K., Onodera J. Synthesis of model compounds of the precursor of carthamin, a colouring matter of safflower, and their conversion into carthamin-type compounds. Chemistry Letters. 1995;24(8):625-626.</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Леус Т.В. Типы взаимодействия генов при наследовании окраски цветков у сафлора красильного. Вестник Санкт-Петербургского университета. 2016;3(4):108-116. DOI: 10.21638/11701/spbu03.2016.408</mixed-citation><mixed-citation xml:lang="en">Leus T.V. Interallelic interactions genes’ types in inheritance of corolla colour of safflower. Bulletin of St. Petersburg University. 2016;3(4):108-116. [in Russian]. DOI: 10.21638/11701/spbu03.2016.408</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Liu X.M., Ahmad N., Yang L.Y., Fu T.Y., Kong J., Yao N., Dong Y.Y., Wang N., Li X.W., Wang F.W., Liu X., Liu W.C., Li H.Y. Molecular cloning and functional characterization of chalcone isomerase from Carthamus tinctorius. AMB Express. 2019;9(1):132. DOI: 10.1186/s13568-019-0854-x</mixed-citation><mixed-citation xml:lang="en">Liu X.M., Ahmad N., Yang L.Y., Fu T.Y., Kong J., Yao N., Dong Y.Y., Wang N., Li X.W., Wang F.W., Liu X., Liu W.C., Li H.Y. Molecular cloning and functional characterization of chalcone isomerase from Carthamus tinctorius. AMB Express. 2019;9(1):132. DOI: 10.1186/s13568-019-0854-x</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Liu X.M., Dong Y.Y., Yao N., Zhang Y., Wang N., Cui X.Y., Li X.W., Wang Y.F., Wang F.W., Yang J., Guan L.L., Du L.N., Li H.Y., Li X.K. De novo sequencing and analysis of the safflower transcriptome to discover putative genes associated with safflor yellow in Carthamus tinctorius L. International Journal of Molecular Science. 2015;16(10):25657-25677. DOI: 10.3390/ijms161025657</mixed-citation><mixed-citation xml:lang="en">Liu X.M., Dong Y.Y., Yao N., Zhang Y., Wang N., Cui X.Y., Li X.W., Wang Y.F., Wang F.W., Yang J., Guan L.L., Du L.N., Li H.Y., Li X.K. De novo sequencing and analysis of the safflower transcriptome to discover putative genes associated with safflor yellow in Carthamus tinctorius L. International Journal of Molecular Science. 2015;16(10):25657-25677. DOI: 10.3390/ijms161025657</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Narkhede B.N., Deokar A.B. Inheritance of corolla color in safflower. Journal of the Maharashtra Agricultural Universities 1986;11:278-281.</mixed-citation><mixed-citation xml:lang="en">Narkhede B.N., Deokar A.B. Inheritance of corolla color in safflower. Journal of the Maharashtra Agricultural Universities 1986;11:278-281.</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Narkhede B.N., Deokar A.B. Inheritance of spininess and pericarp types in safflower. Journal of the Maharashtra Agricultural Universities 1990;15:279-281.</mixed-citation><mixed-citation xml:lang="en">Narkhede B.N., Deokar A.B. Inheritance of spininess and pericarp types in safflower. Journal of the Maharashtra Agricultural Universities 1990;15:279-281.</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Pahlavani M.H., Saeidi G., Mirlohi A.F. Inheritance of flower color and spininess in safflower (Carthamus tinctorius L.). Journal of Heredity. 2004;95:265-267. DOI: 10.1093/jhered/esh030.</mixed-citation><mixed-citation xml:lang="en">Pahlavani M.H., Saeidi G., Mirlohi A.F. Inheritance of flower color and spininess in safflower (Carthamus tinctorius L.). Journal of Heredity. 2004;95:265-267. DOI: 10.1093/jhered/esh030.</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Pu Z., Zhang S., Tang Y., Shi X., Tao H., Yan H., Chen J., Yue S., Chen Y., Zhu Z., Zhou G., Su S., Duan J. Study on changes in pigment composition during the blooming period of safflower based on plant metabolomics and semi-quantitative analysis. Journal of Separation Science. 2021;44(22):4082-4091. DOI: 10.1002/jssc.202100439.</mixed-citation><mixed-citation xml:lang="en">Pu Z., Zhang S., Tang Y., Shi X., Tao H., Yan H., Chen J., Yue S., Chen Y., Zhu Z., Zhou G., Su S., Duan J. Study on changes in pigment composition during the blooming period of safflower based on plant metabolomics and semi-quantitative analysis. Journal of Separation Science. 2021;44(22):4082-4091. DOI: 10.1002/jssc.202100439.</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">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] (Рахмангулов Р.С. Применение системы CRISPR/Cas для редактирования генов декоративных культур. Биотехнология и селекция растений. 2022;5(3):33-41). DOI: 10.30901/2658-6266-2022-3-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: 10.30901/2658-6266-2022-3-o1</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</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="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Ren C., Chen C., Dong S., Wang R., Xian B., Liu T., Xi Z., Pei J., Chen J. Integrated metabolomics and transcriptome analysis on flavonoid biosynthesis in flowers of safflower (Carthamus tinctorius L.) during colour-transition. PeerJ. 2022;10:e13591. DOI: 10.7717/peerj.13591</mixed-citation><mixed-citation xml:lang="en">Ren C., Chen C., Dong S., Wang R., Xian B., Liu T., Xi Z., Pei J., Chen J. Integrated metabolomics and transcriptome analysis on flavonoid biosynthesis in flowers of safflower (Carthamus tinctorius L.) during colour-transition. PeerJ. 2022;10:e13591. DOI: 10.7717/peerj.13591</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Ren C.X., Tang X.H., Chen C.P., Chen J., Pei J., Wu Y.Y., Wu Q.H. Cloning and expression analysis of a new chalcone isomerase gene during flowering in safflower. Turkish Journal of Botany. 2019;43(2):143-150. DOI: 10.3906/bot-1809-25</mixed-citation><mixed-citation xml:lang="en">Ren C.X., Tang X.H., Chen C.P., Chen J., Pei J., Wu Y.Y., Wu Q.H. Cloning and expression analysis of a new chalcone isomerase gene during flowering in safflower. Turkish Journal of Botany. 2019;43(2):143-150. DOI: 10.3906/bot-1809-25</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Saito K. Glucose oxidase, a potential contributor towards flower colour modification in the capitula of Carthamus tinctorius L. Biochemie und Physiologie der Pflanzen. 1993;188:405-417.</mixed-citation><mixed-citation xml:lang="en">Saito K. Glucose oxidase, a potential contributor towards flower colour modification in the capitula of Carthamus tinctorius L. Biochemie und Physiologie der Pflanzen. 1993;188:405-417.</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Saito K., Miyakawa K.-I., Murata T., Enomoto Y. Phytohormone-mediated induction of red colour in the flower florets of a cultivar of dyer’s saffron (Carthamus tinctorius). Zeitschrift fur Naturforschung C. 1998;53(9-10):828-832. DOI: 10.1515/znc-1998-9-1008</mixed-citation><mixed-citation xml:lang="en">Saito K., Miyakawa K.-I., Murata T., Enomoto Y. Phytohormone-mediated induction of red colour in the flower florets of a cultivar of dyer’s saffron (Carthamus tinctorius). Zeitschrift fur Naturforschung C. 1998;53(9-10):828-832. DOI: 10.1515/znc-1998-9-1008</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Sato S., Obara H., Kumazawa T., Onodera J., Furuhata K. Synthesis of (+), (−)-Model compounds and absolute configuration of carthamin; a red pigment in the flower petals of safflower. Chemistry Letters. 1996;25:833-834.</mixed-citation><mixed-citation xml:lang="en">Sato S., Obara H., Kumazawa T., Onodera J., Furuhata K. Synthesis of (+), (−)-Model compounds and absolute configuration of carthamin; a red pigment in the flower petals of safflower. Chemistry Letters. 1996;25:833-834.</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Seshadri T.R., Thakur R.S. The coloring matter of the flowers of Carthamus tinctorius. Current Science. 1960;29:54-55.</mixed-citation><mixed-citation xml:lang="en">Seshadri T.R., Thakur R.S. The coloring matter of the flowers of Carthamus tinctorius. Current Science. 1960;29:54-55.</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Shimokoriyama M., Hattori S. On the formation of carthamin in the flowers of Carthamus tinctorius. Archives of Biochemistry and Biophysics. 1955;54:93-101.</mixed-citation><mixed-citation xml:lang="en">Shimokoriyama M., Hattori S. On the formation of carthamin in the flowers of Carthamus tinctorius. Archives of Biochemistry and Biophysics. 1955;54:93-101.</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Singer A.C., Crowley D.E., Thompson I.P. Secondary plant metabolites in phytoremediation and biotransformation. Trends in Biotechnology. 2003;21(3):123-130. DOI: 10.1016/S0167-7799(02)00041-0</mixed-citation><mixed-citation xml:lang="en">Singer A.C., Crowley D.E., Thompson I.P. Secondary plant metabolites in phytoremediation and biotransformation. Trends in Biotechnology. 2003;21(3):123-130. DOI: 10.1016/S0167-7799(02)00041-0</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Tamburini D., Dyer J., David P., Aceto M., Turina V., Borla M., Vandenbeusch M., Gulmini M. Compositional and micro-morphological characterisation of red colourants in archaeological textiles from Pharaonic Egypt. Molecules. 2019;24(20):3761. DOI: 10.3390/molecules24203761</mixed-citation><mixed-citation xml:lang="en">Tamburini D., Dyer J., David P., Aceto M., Turina V., Borla M., Vandenbeusch M., Gulmini M. Compositional and micro-morphological characterisation of red colourants in archaeological textiles from Pharaonic Egypt. Molecules. 2019;24(20):3761. DOI: 10.3390/molecules24203761</mixed-citation></citation-alternatives></ref><ref id="cit30"><label>30</label><citation-alternatives><mixed-citation xml:lang="ru">Tu YH, Liu F., Guo DD., Fan LJ., Zhu ZX., Xue YR., Gao Y., Guo ML. Molecular characterization of flavanone 3-hydroxylase gene and flavonoid accumulation in two chemotyped safflower lines in response to methyl jasmonate stimulation. BMC Plant Biology. 2016;16:132. DOI: 10.1186/s12870-016-0813-5</mixed-citation><mixed-citation xml:lang="en">Tu YH, Liu F., Guo DD., Fan LJ., Zhu ZX., Xue YR., Gao Y., Guo ML. Molecular characterization of flavanone 3-hydroxylase gene and flavonoid accumulation in two chemotyped safflower lines in response to methyl jasmonate stimulation. BMC Plant Biology. 2016;16:132. DOI: 10.1186/s12870-016-0813-5</mixed-citation></citation-alternatives></ref><ref id="cit31"><label>31</label><citation-alternatives><mixed-citation xml:lang="ru">Urage E., Weyessa B. Genetic diversity of Ethiopian safflower collections. In: V. Ranga Rao, M. Ramachandran, (eds) Proceedings Second International Safflower Conference; 1989 Jan. 9-13; Hyderabad, India. Indian Society of Oilseeds Research, Directorate of Oilseeds Research; 1991. p. 175-178.</mixed-citation><mixed-citation xml:lang="en">Urage E., Weyessa B. Genetic diversity of Ethiopian safflower collections. In: V. Ranga Rao, M. Ramachandran, (eds) Proceedings Second International Safflower Conference; 1989 Jan. 9-13; Hyderabad, India. Indian Society of Oilseeds Research, Directorate of Oilseeds Research; 1991. p. 175-178.</mixed-citation></citation-alternatives></ref><ref id="cit32"><label>32</label><citation-alternatives><mixed-citation xml:lang="ru">Вахрушева Т.Е., Иваненко Е.Н. Классификатор вида Carthamus tinctorius L. (сафлор красильный) / под ред. В.А. Корнейчук. Ленинград: ВИР; 1985.</mixed-citation><mixed-citation xml:lang="en">Vakhrusheva T.E., Ivanenko E.N. Descriptor list for the species Carthamus tinctrorius L. (safflower) (Klassifikator vida Carthamus tinctorius L. (saflor krasil'nyi). V.A. Korneichuk (ed.). Leningrad: VIR; 1985. [in Russian]</mixed-citation></citation-alternatives></ref><ref id="cit33"><label>33</label><citation-alternatives><mixed-citation xml:lang="ru">Waki T., Terashita M., Fujita N., Fukuda K., Kato M., Negishi T., Uchida H., Aoki Y., Takahashi S., Nakayama T. Identification of the genes coding for carthamin synthase, peroxidase homologs that catalyze the final enzymatic step of red pigmentation in safflower (Carthamus tinctorius L.). Plant Cell Physiology. 2021;62(10):1528-1541. DOI: 10.1093/pcp/pcab122</mixed-citation><mixed-citation xml:lang="en">Waki T., Terashita M., Fujita N., Fukuda K., Kato M., Negishi T., Uchida H., Aoki Y., Takahashi S., Nakayama T. Identification of the genes coding for carthamin synthase, peroxidase homologs that catalyze the final enzymatic step of red pigmentation in safflower (Carthamus tinctorius L.). Plant Cell Physiology. 2021;62(10):1528-1541. DOI: 10.1093/pcp/pcab122</mixed-citation></citation-alternatives></ref><ref id="cit34"><label>34</label><citation-alternatives><mixed-citation xml:lang="ru">Wang R., Ren C.X., Dong S., Chen C., Xian B., Wu Q.H., Wang J., Pei J., Chen J. Integrated metabolomics and transcriptome analysis of flavonoid biosynthesis in safflower (Carthamus tinctorius L.) with different colors. Frontiers in Plant Science. 2021;12:712038. DOI: 10.3389/fpls.2021.712038</mixed-citation><mixed-citation xml:lang="en">Wang R., Ren C.X., Dong S., Chen C., Xian B., Wu Q.H., Wang J., Pei J., Chen J. Integrated metabolomics and transcriptome analysis of flavonoid biosynthesis in safflower (Carthamus tinctorius L.) with different colors. Frontiers in Plant Science. 2021;12:712038. DOI: 10.3389/fpls.2021.712038</mixed-citation></citation-alternatives></ref><ref id="cit35"><label>35</label><citation-alternatives><mixed-citation xml:lang="ru">Watanabe S. Cultivation, introduction, and historical note of the Benibana (safflower: Carthamus tinctorius L.) in Yamagata Prefecture. Journal of the Yamagata Agriculture and Forestry Society. 1977;34:73-76.</mixed-citation><mixed-citation xml:lang="en">Watanabe S. Cultivation, introduction, and historical note of the Benibana (safflower: Carthamus tinctorius L.) in Yamagata Prefecture. Journal of the Yamagata Agriculture and Forestry Society. 1977;34:73-76.</mixed-citation></citation-alternatives></ref><ref id="cit36"><label>36</label><citation-alternatives><mixed-citation xml:lang="ru">Wen W.W., Alseekh S., Fernie A.R. Conservation and diversification of flavonoid metabolism in the plant kingdom. Current Opinion in Plant Biology. 2020;55:100-108. DOI: 10.1016/j.pbi.2020.04.004</mixed-citation><mixed-citation xml:lang="en">Wen W.W., Alseekh S., Fernie A.R. Conservation and diversification of flavonoid metabolism in the plant kingdom. Current Opinion in Plant Biology. 2020;55:100-108. DOI: 10.1016/j.pbi.2020.04.004</mixed-citation></citation-alternatives></ref><ref id="cit37"><label>37</label><citation-alternatives><mixed-citation xml:lang="ru">Yang W.T., Liu X.M., Wan Q., Yao N., Wang N., Zhang X.M., Jiao Z.D., Li H.Y., Li X.K. Full-length cDNA cloning of flavonol synthase genes of Carthamus tinctorius and construction plant expression vector. Zhongguo Zhong Yao Za Zhi. 2015;40(4):634-638. [in Chinese]</mixed-citation><mixed-citation xml:lang="en">Yang W.T., Liu X.M., Wan Q., Yao N., Wang N., Zhang X.M., Jiao Z.D., Li H.Y., Li X.K. Full-length cDNA cloning of flavonol synthase genes of Carthamus tinctorius and construction plant expression vector. Zhongguo Zhong Yao Za Zhi. 2015;40(4):634-638. [in Chinese]</mixed-citation></citation-alternatives></ref><ref id="cit38"><label>38</label><citation-alternatives><mixed-citation xml:lang="ru">Yue S., Tang Y., Li S., Duan J.A. Chemical and biological properties of quinochalcone C-glycosides from the florets of Carthamus tinctorius. Molecules. 2013;18(12):15220-15254. DOI: 10.3390/molecules181215220</mixed-citation><mixed-citation xml:lang="en">Yue S., Tang Y., Li S., Duan J.A. Chemical and biological properties of quinochalcone C-glycosides from the florets of Carthamus tinctorius. Molecules. 2013;18(12):15220-15254. DOI: 10.3390/molecules181215220</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>
