<?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">10.30901/2658-6266-2020-1-o1</article-id><article-id custom-type="elpub" pub-id-type="custom">biosel-76</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>REVIEW ARTICLES</subject></subj-group></article-categories><title-group><article-title>Генная инженерия как способ получения декоративных растений с изменённой окраской цветков</article-title><trans-title-group xml:lang="en"><trans-title>Genetic engineering as a way to obtain ornamental plants with a changed flower color</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-4861-4805</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Санникова</surname><given-names>В. Ю.</given-names></name><name name-style="western" xml:lang="en"><surname>Sannikova</surname><given-names>V. Yu.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Биологический факультет</p><p>199034, г. Санкт-Петербург, Университетская наб., д. 7–9</p></bio><bio xml:lang="en"><p>Faculty of Biology</p><p>7/9, University Emb., St. Petersburg 199034</p></bio><email xlink:type="simple">st049681@student.spbu.ru</email><xref ref-type="aff" rid="aff-1"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru"><institution>Санкт-Петербургский государственный университет</institution></aff><aff xml:lang="en"><institution>St. Petersburg State University</institution></aff></aff-alternatives><pub-date pub-type="collection"><year>2020</year></pub-date><pub-date pub-type="epub"><day>10</day><month>08</month><year>2020</year></pub-date><volume>3</volume><issue>1</issue><fpage>40</fpage><lpage>45</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Санникова В.Ю., 2020</copyright-statement><copyright-year>2020</copyright-year><copyright-holder xml:lang="ru">Санникова В.Ю.</copyright-holder><copyright-holder xml:lang="en">Sannikova V.Y.</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/76">https://biosel.elpub.ru/jour/article/view/76</self-uri><abstract><p>Важным направлением в цветоводстве является получение новых сортов декоративных растений, среди которых наибольшим спросом пользуются растения с необычной окраской цветков. Ранее для их получения успешно применялись традиционные программы по разведению и селекции. Однако в настоящий момент генная инженерия способна предложить альтернативный путь создания новых форм и сортов. Антоцианы, относящиеся к флавоноидам, беталаины и каротиноиды являются основными типами пигментов, которые синтезируются в растении и отвечают за окраску лепестков цветка. Модификация путей биосинтеза пигментов с помощью методов генной инженерии позволяет добиться результатов, которые не могут быть получены при помощи традиционной селекции. В данном обзоре литературы представлены основные достижения применения методов генной инженерии в цветоводстве путём модификации окраски цветков. Существует несколько основных направлений в работе с генами биосинтеза пигментов. Среди них чаще всего используется стратегия по подавлению экспрессии генов для предотвращения синтеза пигмента или, наоборот, для устранения факторов, препятствующих развитию окраски. Нередко используется метод введения в геном растений дополнительных гетерологичных генов, недостающих в пути биосинтеза пигментов. Также для модификации окраски прибегают к геномному редактированию посредством технологии CRISPR/Cas, но данный метод в отношении декоративных растений стал использоваться относительно недавно. Несмотря на быстрое развитие биотехнологий, существуют препятствия для распространения генномодифицированных растений на мировом рынке. Преодоление ряда проблем сможет сделать производство трансгенных декоративных растений экономически более выгодным и привлекательным, чем выведение новых сортов исключительно с помощью традиционных методов селекции.</p></abstract><trans-abstract xml:lang="en"><p>An important trend in the field of floriculture is the creation of new varieties of ornamental plants, among which varieties with unusual color are most in demand. To this end, traditional breeding and selection programs have been successfully applied for many years. However, currently genetic engineering is able to offer an alternative way to obtain new forms and varieties. Anthocyanins belonging to flavonoids, betalains and carotenoids are the main types of pigments that are synthesized in the plant and are responsible for the color of flower petals. The modification of pigment biosynthesis pathways using genetic engineering techniques can produce results that cannot be obtained by traditional breeding. This review presents the main advances in the application of genetic engineering techniques in floriculture using the example of flower color modification. There are several main areas of work with the genes of pigment biosynthesis. Among them, the strategy of suppressing gene expression is used most often. Expression of certain genes is suppressed to prevent pigment synthesis, or vice versa, to eliminate factors that hinder color development. The method of additional heterologous genes insertion to plants lacking them in the pathway of pigment biosynthesis is often used. Genomic editing, in particular by using the CRISPR/Cas system, is also used for color modification, but the application of this method to ornamental plants is a relatively recent innovation. Despite the rapid development of biotechnology, there are obstacles to the distribution of genetically modified plants on the world market. By addressing a number of problems, the production of transgenic ornamental plants may become economically more cost-effective and attractive than the development of new varieties exclusively through traditional breeding methods.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>декоративные растения</kwd><kwd>генная инженерия</kwd><kwd>окраска цветка</kwd><kwd>агробактериальная трансформация</kwd><kwd>CRISPR/Cas</kwd><kwd>геномное редактирование</kwd></kwd-group><kwd-group xml:lang="en"><kwd>ornamental plants</kwd><kwd>genetic engineering</kwd><kwd>flower color</kwd><kwd>Agro-bacterium-mediated transformation</kwd><kwd>CRISPR/Cas</kwd><kwd>genome editing</kwd></kwd-group><funding-group><funding-statement xml:lang="ru">Обзор подготовлен в рамках магистерской программы «Молекулярная биология и агробиотехнология растений» биологического факультета СПбГУ. Особую благодарность автор выражает д.б.н. Матвеевой Татьяне Валерьевне за предложенную тему обзора и ценные замечания.</funding-statement><funding-statement xml:lang="en">The review was prepared as part of the master degree program in Molecular Biology and Plant Agrobiotechnology of the Faculty of Biology of St. Petersburg State University. The author expresses special gratitude to Dr. Biol. Sci. Tatyana V. Matveeva for the proposed topic of the review and valuable comments.</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">Aida R., Kishimoto S., Tanaka Y., Shibata M. Modification of flower color in torenia (Torenia fournieri Lind.) by genetic transformation. Plant Science. 2000a;151(1):33-42. DOI: 10.1016/S0168-9452(99)00239-3</mixed-citation><mixed-citation xml:lang="en">Aida R., Kishimoto S., Tanaka Y., Shibata M. Modification of flower color in torenia (Torenia fournieri Lind.) by genetic transformation. Plant Science. 2000a;151(1):33-42. DOI: 10.1016/S0168-9452(99)00239-3</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Aida R., Yoshida K., Kondo T., Kishimoto S., Shibata M. Copigmentation gives bluer flowers on transgenic torenia plants with the antisense dihydroflavonol-4-reductase gene. Plant Science. 2000b;160(1):49- 56. DOI: 10.1016/S0168-9452(00)00364-2</mixed-citation><mixed-citation xml:lang="en">Aida R., Yoshida K., Kondo T., Kishimoto S., Shibata M. Copigmentation gives bluer flowers on transgenic torenia plants with the antisense dihydroflavonol-4-reductase gene. Plant Science. 2000b;160(1):49- 56. DOI: 10.1016/S0168-9452(00)00364-2</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Albert N. W., Davies K. M., Lewis D. H., Zhang H., Montefiori M., Brendolise C., Boase M. R., Ngo H., Jameson P. E., Schwinn K. E. A conserved network of transcriptional activators and repressors regulates anthocyanin pigmentation in eudicots. The Plant Cell. 2014;26(3):962-80. DOI: 10.1105/tpc</mixed-citation><mixed-citation xml:lang="en">Albert N. W., Davies K. M., Lewis D. H., Zhang H., Montefiori M., Brendolise C., Boase M. R., Ngo H., Jameson P. E., Schwinn K. E. A conserved network of transcriptional activators and repressors regulates anthocyanin pigmentation in eudicots. The Plant Cell. 2014;26(3):962-80. DOI: 10.1105/tpc</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Boutigny A. L., Dohin N., Pornin D., Rolland M. Overview and detectability of the genetic modifications in ornamental plants. Horticulture Research. 2020;7:11. DOI: 10.1038/s41438-019-0232-5</mixed-citation><mixed-citation xml:lang="en">Boutigny A. L., Dohin N., Pornin D., Rolland M. Overview and detectability of the genetic modifications in ornamental plants. Horticulture Research. 2020;7:11. DOI: 10.1038/s41438-019-0232-5</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Bradley J., Davies K., Dolores S., Bloor S., Lewis D. The maize Lc regulatory gene up regulates the flavonoid biosynthetic pathway of Petunia. The Plant Journal. 2002;13(3):381-392. DOI: 10.1046/j.1365-313X.1998.00031.x</mixed-citation><mixed-citation xml:lang="en">Bradley J., Davies K., Dolores S., Bloor S., Lewis D. The maize Lc regulatory gene up regulates the flavonoid biosynthetic pathway of Petunia. The Plant Journal. 2002;13(3):381-392. DOI: 10.1046/j.1365-313X.1998.00031.x</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Brand M.H. Ornamental Plant Transformation. Journal of Crop Improvement. 2006;17(1):27-50. DOI: 10.1300/J411v17n01_02</mixed-citation><mixed-citation xml:lang="en">Brand M.H. Ornamental Plant Transformation. Journal of Crop Improvement. 2006;17(1):27-50. DOI: 10.1300/J411v17n01_02</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Brugliera F., Tao G.Q., Tems U., Kalc G., Mouradova E., Price K., Stevenson K., Nakamura N., Stacey I., Katsumoto Y., Tanaka Y., Mason J. G. Violet/blue chrysanthemums--metabolic engineering of the anthocyanin biosynthetic pathway results in novel petal colors. Plant and Cell Physiology. 2013;54(10):1696-1710. DOI: 10.1093/pcp/pct110</mixed-citation><mixed-citation xml:lang="en">Brugliera F., Tao G.Q., Tems U., Kalc G., Mouradova E., Price K., Stevenson K., Nakamura N., Stacey I., Katsumoto Y., Tanaka Y., Mason J. G. Violet/blue chrysanthemums--metabolic engineering of the anthocyanin biosynthetic pathway results in novel petal colors. Plant and Cell Physiology. 2013;54(10):1696-1710. DOI: 10.1093/pcp/pct110</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Chandler S.F., Brugliera F. Genetic modification in floriculture. Biotechnology Letters. 2011;33(2):207-214. DOI: 10.1007/s10529-010-0424-4</mixed-citation><mixed-citation xml:lang="en">Chandler S.F., Brugliera F. Genetic modification in floriculture. Biotechnology Letters. 2011;33(2):207-214. DOI: 10.1007/s10529-010-0424-4</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Chandler S.F., Sanchez C. Genetic modification; the development of transgenic ornamental plant varieties. Plant Biotechnology Journal. 2012;10(8):891-903. DOI: 10.1111/j.1467-7652.2012.00693.x</mixed-citation><mixed-citation xml:lang="en">Chandler S.F., Sanchez C. Genetic modification; the development of transgenic ornamental plant varieties. Plant Biotechnology Journal. 2012;10(8):891-903. DOI: 10.1111/j.1467-7652.2012.00693.x</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Chandler S.F., Tanaka Y. Genetic Modification in Floriculture. Critical Reviews in Plant Sciences. 2007;26(4):169-197. DOI: 10.1080/07352680701429381</mixed-citation><mixed-citation xml:lang="en">Chandler S.F., Tanaka Y. Genetic Modification in Floriculture. Critical Reviews in Plant Sciences. 2007;26(4):169-197. DOI: 10.1080/07352680701429381</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Courtney-Gutterson N., Napoli C., Lemieux C., Morgan A., Firoozabady E., Robinson K.E. Modification of flower color in florist's chrysanthemum: production of a white-flowering variety through molecular genetics. Biotechnology. 1994;12(3):268-271. DOI: 10.1038/nbt0394-268</mixed-citation><mixed-citation xml:lang="en">Courtney-Gutterson N., Napoli C., Lemieux C., Morgan A., Firoozabady E., Robinson K.E. Modification of flower color in florist's chrysanthemum: production of a white-flowering variety through molecular genetics. Biotechnology. 1994;12(3):268-271. DOI: 10.1038/nbt0394-268</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Delgado-Vargas F., Jimenez A.R., Paredes-Lopez O. Natural pigments: carotenoids, anthocyanins, and betalains--characteristics, biosynthesis, processing, and stability. Critical Reviews in Food Science and Nutrition. 2000;40(3):173-289. DOI: 10.1080/10408690091189257</mixed-citation><mixed-citation xml:lang="en">Delgado-Vargas F., Jimenez A.R., Paredes-Lopez O. Natural pigments: carotenoids, anthocyanins, and betalains--characteristics, biosynthesis, processing, and stability. Critical Reviews in Food Science and Nutrition. 2000;40(3):173-289. DOI: 10.1080/10408690091189257</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Dobres M. Barriers to Genetically Engineered Ornamentals: An Industry Perspective. In: J.A.T. da Silva (ed.). Floriculture, Ornamental and Plant Biotechnology: Advances and Topical Issues, Volume V. London: Global Science Books; 2008. p.1-14.</mixed-citation><mixed-citation xml:lang="en">Dobres M. Barriers to Genetically Engineered Ornamentals: An Industry Perspective. In: J.A.T. da Silva (ed.). Floriculture, Ornamental and Plant Biotechnology: Advances and Topical Issues, Volume V. London: Global Science Books; 2008. p.1-14.</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Dunwell J. Transgenic Crops: The Next Generation, or an Example of 2020 Vision. Annals of Botany. 1999;84(3):269-277. DOI: 10.1006/anbo.1999.0934</mixed-citation><mixed-citation xml:lang="en">Dunwell J. Transgenic Crops: The Next Generation, or an Example of 2020 Vision. Annals of Botany. 1999;84(3):269-277. DOI: 10.1006/anbo.1999.0934</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Elomaa P., Honkanen J., Puska R., Seppänen P., Helariutta Y., Mehto M., Kotilainen M., Nevalainen L., Teeri T.H. Agrobacterium-Mediated Transfer of Antisense Chalcone Synthase cDNA to Gerbera hybrida Inhibits Flower Pigmentation. Biotechnology. 1993;11:508–511. DOI: 10.1038/nbt0493-508</mixed-citation><mixed-citation xml:lang="en">Elomaa P., Honkanen J., Puska R., Seppänen P., Helariutta Y., Mehto M., Kotilainen M., Nevalainen L., Teeri T.H. Agrobacterium-Mediated Transfer of Antisense Chalcone Synthase cDNA to Gerbera hybrida Inhibits Flower Pigmentation. Biotechnology. 1993;11:508–511. DOI: 10.1038/nbt0493-508</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Katsumoto Y., Fukuchi-Mizutani M., Fukui Y., Brugliera F., Holton T.A., Karan M., Nakamura N., Yonekura-Sakakibara K., Togami J., Pigeaire A., Tao G.Q., Nehra N.S., Lu C.Y., Dyson B.K., Tsuda S., Ashikari T., Kusumi T., Mason J.G., Tanaka Y. Engineering of the Rose Flavonoid Biosynthetic Pathway Successfully Generated Blue-Hued Flowers Accumulating. Plant and Cell Physiology. 2007;48(11):1589-1600. DOI: 10.1093/pcp/pcm131</mixed-citation><mixed-citation xml:lang="en">Katsumoto Y., Fukuchi-Mizutani M., Fukui Y., Brugliera F., Holton T.A., Karan M., Nakamura N., Yonekura-Sakakibara K., Togami J., Pigeaire A., Tao G.Q., Nehra N.S., Lu C.Y., Dyson B.K., Tsuda S., Ashikari T., Kusumi T., Mason J.G., Tanaka Y. Engineering of the Rose Flavonoid Biosynthetic Pathway Successfully Generated Blue-Hued Flowers Accumulating. Plant and Cell Physiology. 2007;48(11):1589-1600. DOI: 10.1093/pcp/pcm131</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Khoo H.E., Azlan A., Tang S.T., Lim S.M. Anthocyanidins and anthocyanins: colored pigments as food, pharmaceutical ingredients, and the potential health benefits. Food and Nutrition Research. 2017;61(1):1361779. DOI: 10.1080/16546628.2017.1361779</mixed-citation><mixed-citation xml:lang="en">Khoo H.E., Azlan A., Tang S.T., Lim S.M. Anthocyanidins and anthocyanins: colored pigments as food, pharmaceutical ingredients, and the potential health benefits. Food and Nutrition Research. 2017;61(1):1361779. DOI: 10.1080/16546628.2017.1361779</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Kishi-Kaboshi M., Aida R., Sasaki K. Genome engineering in ornamental plants: Current status and future prospects. Plant Physiology and Biochemistry. 2018;131:47-52. DOI: 10.1016/j.plaphy.2018.03.015</mixed-citation><mixed-citation xml:lang="en">Kishi-Kaboshi M., Aida R., Sasaki K. Genome engineering in ornamental plants: Current status and future prospects. Plant Physiology and Biochemistry. 2018;131:47-52. DOI: 10.1016/j.plaphy.2018.03.015</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Krol A., Lenting P., Veenstra J., Meer I., Koes R., Gerats A., Mol J., Stuitje A. An anti-sense chalcone synthase gene in transgenic plants inhibits flower pigmentation. Nature. 1998;333:866–869. DOI: 10.1038/333866a0</mixed-citation><mixed-citation xml:lang="en">Krol A., Lenting P., Veenstra J., Meer I., Koes R., Gerats A., Mol J., Stuitje A. An anti-sense chalcone synthase gene in transgenic plants inhibits flower pigmentation. Nature. 1998;333:866–869. DOI: 10.1038/333866a0</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Kuligowska K., Lutken H., Muller R. Towards development of new ornamental plants: status and progress in wide hybridization. Planta. 2016;244(1):1-17. DOI: 10.1007/s00425-016-2493-7</mixed-citation><mixed-citation xml:lang="en">Kuligowska K., Lutken H., Muller R. Towards development of new ornamental plants: status and progress in wide hybridization. Planta. 2016;244(1):1-17. DOI: 10.1007/s00425-016-2493-7</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Lin-Wang K., Bolitho K., Grafton K., Kortstee A., Karunairetnam S., McGhie T.K., Espley R.V., Hellens R.P., Allan A.C. An R2R3 MYB transcription factor associated with regulation of the anthocyanin biosynthetic pathway in Rosaceae. BMC Plant Biology. 2010;10:50. DOI: 10.1186/1471-2229-10-50</mixed-citation><mixed-citation xml:lang="en">Lin-Wang K., Bolitho K., Grafton K., Kortstee A., Karunairetnam S., McGhie T.K., Espley R.V., Hellens R.P., Allan A.C. An R2R3 MYB transcription factor associated with regulation of the anthocyanin biosynthetic pathway in Rosaceae. BMC Plant Biology. 2010;10:50. DOI: 10.1186/1471-2229-10-50</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Meyer P., Heidmann I., Forkmann G., Saedler H. A new petunia flower colour generated by transformation of a mutant with a maize gene. Nature. 1987;330(6149):677-678. DOI: 10.1038/330677a0</mixed-citation><mixed-citation xml:lang="en">Meyer P., Heidmann I., Forkmann G., Saedler H. A new petunia flower colour generated by transformation of a mutant with a maize gene. Nature. 1987;330(6149):677-678. DOI: 10.1038/330677a0</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Morita Y., Saitoh M., Hoshino A., Nitasaka E., Iida S. Isolation of cDNAs for R2R3-MYB, bHLH and WDR Transcriptional Regulators and Identification of c and ca Mutations Conferring White Flowers in the Japanese Morning Glory. Plant and Cell Physiology. 2006;47(4):457-470. DOI: 10.1093/pcp/pcj012</mixed-citation><mixed-citation xml:lang="en">Morita Y., Saitoh M., Hoshino A., Nitasaka E., Iida S. Isolation of cDNAs for R2R3-MYB, bHLH and WDR Transcriptional Regulators and Identification of c and ca Mutations Conferring White Flowers in the Japanese Morning Glory. Plant and Cell Physiology. 2006;47(4):457-470. DOI: 10.1093/pcp/pcj012</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Morita Y., Takagi K., Fukuchi-Mizutani M., Ishiguro K., Tanaka Y., Nitasaka E., Nakayama M., Saito N., Kagami T., Hoshino A., Iida S. A chalcone isomerase-like protein enhances flavonoid production and flower pigmentation. The Plant Journal. 2014;78(2):294-304. DOI: 10.1111/tpj.12469</mixed-citation><mixed-citation xml:lang="en">Morita Y., Takagi K., Fukuchi-Mizutani M., Ishiguro K., Tanaka Y., Nitasaka E., Nakayama M., Saito N., Kagami T., Hoshino A., Iida S. A chalcone isomerase-like protein enhances flavonoid production and flower pigmentation. The Plant Journal. 2014;78(2):294-304. DOI: 10.1111/tpj.12469</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Nakamura N., Fukuchi-Mizutani M., Miyazaki K., Suzuki K., Tanaka Y. RNAi suppression of the anthocyanidin synthase gene in Torenia hybrida yields white flowers with higher frequency and better stability than antisense and sense suppression. Plant Biotechnology. 2006;23(1):13-17. DOI: 10.5511/plantbiotechnology.23.13</mixed-citation><mixed-citation xml:lang="en">Nakamura N., Fukuchi-Mizutani M., Miyazaki K., Suzuki K., Tanaka Y. RNAi suppression of the anthocyanidin synthase gene in Torenia hybrida yields white flowers with higher frequency and better stability than antisense and sense suppression. Plant Biotechnology. 2006;23(1):13-17. DOI: 10.5511/plantbiotechnology.23.13</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Nakatsuka T., Mishiba K., Abe Y., Kubota A., Kakizaki Y., Yamamura S., Nishihara M. Flower color modification of gentian plants by RNAi-mediated gene silencing. Plant Biotechnology. 2008a;25(1):61-68. DOI: 10.5511/plantbiotechnology.25.61</mixed-citation><mixed-citation xml:lang="en">Nakatsuka T., Mishiba K., Abe Y., Kubota A., Kakizaki Y., Yamamura S., Nishihara M. Flower color modification of gentian plants by RNAi-mediated gene silencing. Plant Biotechnology. 2008a;25(1):61-68. DOI: 10.5511/plantbiotechnology.25.61</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Nakatsuka T., Haruta K.S., Pitaksutheepong C., Abe Y., Kakizaki Y., Yamamoto K., Shimada N., Yamamura S., Nishihara M. Identification and characterization of R2R3-MYB and bHLH transcription factors regulating anthocyanin biosynthesis in gentian flowers. Plant and Cell Physiology. 2008b;49(12):1818-1829. DOI: 10.1093/pcp/pcn163</mixed-citation><mixed-citation xml:lang="en">Nakatsuka T., Haruta K.S., Pitaksutheepong C., Abe Y., Kakizaki Y., Yamamoto K., Shimada N., Yamamura S., Nishihara M. Identification and characterization of R2R3-MYB and bHLH transcription factors regulating anthocyanin biosynthesis in gentian flowers. Plant and Cell Physiology. 2008b;49(12):1818-1829. DOI: 10.1093/pcp/pcn163</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Nishihara M., Higuchi A., Watanabe A., Tasaki K. Application of the CRISPR/Cas9 system for modification of flower color in Torenia fournieri. BMC Plant Biology. 2018;18(1):331. DOI: 10.1186/s12870-018-1539-3</mixed-citation><mixed-citation xml:lang="en">Nishihara M., Higuchi A., Watanabe A., Tasaki K. Application of the CRISPR/Cas9 system for modification of flower color in Torenia fournieri. BMC Plant Biology. 2018;18(1):331. DOI: 10.1186/s12870-018-1539-3</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Nishihara M., Nakatsuka T. Genetic engineering of novel flower colors in floricultural plants: recent advances via transgenic approaches. Methods in Molecular Biology. 2010;589:325-347. DOI: 10.1007/978-1-60327-114-1_29</mixed-citation><mixed-citation xml:lang="en">Nishihara M., Nakatsuka T. Genetic engineering of novel flower colors in floricultural plants: recent advances via transgenic approaches. Methods in Molecular Biology. 2010;589:325-347. DOI: 10.1007/978-1-60327-114-1_29</mixed-citation></citation-alternatives></ref><ref id="cit30"><label>30</label><citation-alternatives><mixed-citation xml:lang="ru">Nishihara M., Nakatsuka T. Genetic engineering of flavonoid pigments to modify flower color in floricultural plants. Biotechnology Letters. 2011;33(3):433-441. DOI: 10.1007/s10529-010-0461-z</mixed-citation><mixed-citation xml:lang="en">Nishihara M., Nakatsuka T. Genetic engineering of flavonoid pigments to modify flower color in floricultural plants. Biotechnology Letters. 2011;33(3):433-441. DOI: 10.1007/s10529-010-0461-z</mixed-citation></citation-alternatives></ref><ref id="cit31"><label>31</label><citation-alternatives><mixed-citation xml:lang="ru">Noda N. Recent advances in the research and development of blue flowers. Breeding Science. 2018;68:79-87. DOI: 10.1270/jsbbs.17132</mixed-citation><mixed-citation xml:lang="en">Noda N. Recent advances in the research and development of blue flowers. Breeding Science. 2018;68:79-87. DOI: 10.1270/jsbbs.17132</mixed-citation></citation-alternatives></ref><ref id="cit32"><label>32</label><citation-alternatives><mixed-citation xml:lang="ru">Noda N., Yoshioka S., Kishimoto S., Nakayama M., Douzono M., Tanaka Y., Aida R. Generation of blue chrysanthemums by anthocyanin B-ring hydroxylation and glucosylation and its coloration mechanism. Science Advances. 2017;3(7):e1602785. DOI: 10.1126/sciadv.1602785.</mixed-citation><mixed-citation xml:lang="en">Noda N., Yoshioka S., Kishimoto S., Nakayama M., Douzono M., Tanaka Y., Aida R. Generation of blue chrysanthemums by anthocyanin B-ring hydroxylation and glucosylation and its coloration mechanism. Science Advances. 2017;3(7):e1602785. DOI: 10.1126/sciadv.1602785.</mixed-citation></citation-alternatives></ref><ref id="cit33"><label>33</label><citation-alternatives><mixed-citation xml:lang="ru">Ohmiya A., Kishimoto S., Aida R., Yoshioka S., Sumitomo K. Carotenoid cleavage dioxygenase (CmCCD4a) contributes to white color formation in chrysanthemum petals. Plant Physiology. 2016;142(3):1193-1201. DOI: 10.1104/pp.106.087130</mixed-citation><mixed-citation xml:lang="en">Ohmiya A., Kishimoto S., Aida R., Yoshioka S., Sumitomo K. Carotenoid cleavage dioxygenase (CmCCD4a) contributes to white color formation in chrysanthemum petals. Plant Physiology. 2016;142(3):1193-1201. DOI: 10.1104/pp.106.087130</mixed-citation></citation-alternatives></ref><ref id="cit34"><label>34</label><citation-alternatives><mixed-citation xml:lang="ru">Polturak G., Grossman N., Vela-Corcia D., Dong Y., Nudel A., Pliner M., Levy M., Rogachev I., Aharoni A. Engineered gray mold resistance, antioxidant capacity, and pigmentation in betalain-producing crops and ornamentals. PNAS. 2017;114(34):9062-9067. DOI: 10.1073/pnas.1707176114</mixed-citation><mixed-citation xml:lang="en">Polturak G., Grossman N., Vela-Corcia D., Dong Y., Nudel A., Pliner M., Levy M., Rogachev I., Aharoni A. Engineered gray mold resistance, antioxidant capacity, and pigmentation in betalain-producing crops and ornamentals. PNAS. 2017;114(34):9062-9067. DOI: 10.1073/pnas.1707176114</mixed-citation></citation-alternatives></ref><ref id="cit35"><label>35</label><citation-alternatives><mixed-citation xml:lang="ru">Qi Y., Lou Q., Quan Y., Liu Y., Wang Y. Flower-specific expression of the Phalaenopsis flavonoid 3’, 5’-hydoxylase modifies flower color pigmentation in Petunia and Lilium. Plant Cell, Tissue and Organ Culture. 2013;115:263–273 DOI: 10.1007/s11240-013-0359-2</mixed-citation><mixed-citation xml:lang="en">Qi Y., Lou Q., Quan Y., Liu Y., Wang Y. Flower-specific expression of the Phalaenopsis flavonoid 3’, 5’-hydoxylase modifies flower color pigmentation in Petunia and Lilium. Plant Cell, Tissue and Organ Culture. 2013;115:263–273 DOI: 10.1007/s11240-013-0359-2</mixed-citation></citation-alternatives></ref><ref id="cit36"><label>36</label><citation-alternatives><mixed-citation xml:lang="ru">Quattrocchio F., Baudry A., Lepiniec L., Grotewold E. The Regulation of Flavonoid Biosynthesis. In: Grotewold E. (eds) The Science of Flavonoids. New York: Springer; 2006. p. 97-122. DOI: 10.1007/978-0-387-28822-2_4</mixed-citation><mixed-citation xml:lang="en">Quattrocchio F., Baudry A., Lepiniec L., Grotewold E. The Regulation of Flavonoid Biosynthesis. In: Grotewold E. (eds) The Science of Flavonoids. New York: Springer; 2006. p. 97-122. DOI: 10.1007/978-0-387-28822-2_4</mixed-citation></citation-alternatives></ref><ref id="cit37"><label>37</label><citation-alternatives><mixed-citation xml:lang="ru">Rodriguez-Amaya D.B. Update on natural food pigments - A mini-review on carotenoids, anthocyanins, and betalains. Food Research International. 2019;124:200-205. DOI: 10.1016/j.foodres.2018.05.028</mixed-citation><mixed-citation xml:lang="en">Rodriguez-Amaya D.B. Update on natural food pigments - A mini-review on carotenoids, anthocyanins, and betalains. Food Research International. 2019;124:200-205. DOI: 10.1016/j.foodres.2018.05.028</mixed-citation></citation-alternatives></ref><ref id="cit38"><label>38</label><citation-alternatives><mixed-citation xml:lang="ru">Scarano A., Chieppa M., Santino A. Looking at Flavonoid Biodiversity in Horticultural Crops: A Colored Mine with Nutritional Benefits. Plants (Basel). 2018;7(4):98. DOI: 10.3390/plants7040098</mixed-citation><mixed-citation xml:lang="en">Scarano A., Chieppa M., Santino A. Looking at Flavonoid Biodiversity in Horticultural Crops: A Colored Mine with Nutritional Benefits. Plants (Basel). 2018;7(4):98. DOI: 10.3390/plants7040098</mixed-citation></citation-alternatives></ref><ref id="cit39"><label>39</label><citation-alternatives><mixed-citation xml:lang="ru">Schwinn K.E., Boase M.R., Bradley J.M., Lewis D.H., Deroles S.C., Martin C.R., Davies K.M. MYB and bHLH transcription factor transgenes increase anthocyanin pigmentation in petunia and lisianthus plants, and the petunia phenotypes are strongly enhanced under field conditions. Frontiers in Plant Science. 2014;5:603. DOI: 10.3389/fpls.2014.00603</mixed-citation><mixed-citation xml:lang="en">Schwinn K.E., Boase M.R., Bradley J.M., Lewis D.H., Deroles S.C., Martin C.R., Davies K.M. MYB and bHLH transcription factor transgenes increase anthocyanin pigmentation in petunia and lisianthus plants, and the petunia phenotypes are strongly enhanced under field conditions. Frontiers in Plant Science. 2014;5:603. DOI: 10.3389/fpls.2014.00603</mixed-citation></citation-alternatives></ref><ref id="cit40"><label>40</label><citation-alternatives><mixed-citation xml:lang="ru">Schwinn K., Venail J., Shang Y., Mackay S., Alm V., Butelli E., Oyama R., Bailey P., Davies K., Martin C. A small family of MYB-regulatory genes controls floral pigmentation intensity and patterning in the genus Antirrhinum. The Plant Cell. 2006;18(4):831-851. DOI: 10.1105/tpc.105.039255</mixed-citation><mixed-citation xml:lang="en">Schwinn K., Venail J., Shang Y., Mackay S., Alm V., Butelli E., Oyama R., Bailey P., Davies K., Martin C. A small family of MYB-regulatory genes controls floral pigmentation intensity and patterning in the genus Antirrhinum. The Plant Cell. 2006;18(4):831-851. DOI: 10.1105/tpc.105.039255</mixed-citation></citation-alternatives></ref><ref id="cit41"><label>41</label><citation-alternatives><mixed-citation xml:lang="ru">Shibata M. Importance of genetic transformation in ornamental plant breeding. Plant Biotechnology. 2008;25(1):3-8. DOI: 10.5511/plantbiotechnology.25.3</mixed-citation><mixed-citation xml:lang="en">Shibata M. Importance of genetic transformation in ornamental plant breeding. Plant Biotechnology. 2008;25(1):3-8. DOI: 10.5511/plantbiotechnology.25.3</mixed-citation></citation-alternatives></ref><ref id="cit42"><label>42</label><citation-alternatives><mixed-citation xml:lang="ru">Suzuki S., Nishihara M., Nakatsuka T., Misawa N., Ogiwara I., Yamamura S. Flower color alteration in Lotus japonicus by modification of the carotenoid biosynthetic pathway. Plant Cell Reports. 2007;26(7):951959. DOI: 10.1007/s00299-006-0302-7</mixed-citation><mixed-citation xml:lang="en">Suzuki S., Nishihara M., Nakatsuka T., Misawa N., Ogiwara I., Yamamura S. Flower color alteration in Lotus japonicus by modification of the carotenoid biosynthetic pathway. Plant Cell Reports. 2007;26(7):951959. DOI: 10.1007/s00299-006-0302-7</mixed-citation></citation-alternatives></ref><ref id="cit43"><label>43</label><citation-alternatives><mixed-citation xml:lang="ru">Stafford H.A. Anthocyanins and betalains: evolution of the mutually exclusive pathways. Plant Science. 1994;101(2):91-98. DOI: 10.1016/0168-9452(94)90244-5</mixed-citation><mixed-citation xml:lang="en">Stafford H.A. Anthocyanins and betalains: evolution of the mutually exclusive pathways. Plant Science. 1994;101(2):91-98. DOI: 10.1016/0168-9452(94)90244-5</mixed-citation></citation-alternatives></ref><ref id="cit44"><label>44</label><citation-alternatives><mixed-citation xml:lang="ru">Tanaka Y., Brugliera F. Flower color and cytochromes P450. Philosophical Transactions of the Royal Society B: Biological Sciences. 2013;368(1612):20120432. DOI: 10.1098/rstb.2012.0432</mixed-citation><mixed-citation xml:lang="en">Tanaka Y., Brugliera F. Flower color and cytochromes P450. Philosophical Transactions of the Royal Society B: Biological Sciences. 2013;368(1612):20120432. DOI: 10.1098/rstb.2012.0432</mixed-citation></citation-alternatives></ref><ref id="cit45"><label>45</label><citation-alternatives><mixed-citation xml:lang="ru">Watanabe K., Oda-Yamamizo C., Sage-Ono K., Ohmiya A., Ono M. Alteration of flower color in Ipomoea nil through CRISPR/Cas9-mediated mutagenesis of carotenoid cleavage dioxygenase 4. Transgenic Research. 2018;27(1):25-38. DOI: 10.1007/s11248-017-0051-0</mixed-citation><mixed-citation xml:lang="en">Watanabe K., Oda-Yamamizo C., Sage-Ono K., Ohmiya A., Ono M. Alteration of flower color in Ipomoea nil through CRISPR/Cas9-mediated mutagenesis of carotenoid cleavage dioxygenase 4. Transgenic Research. 2018;27(1):25-38. DOI: 10.1007/s11248-017-0051-0</mixed-citation></citation-alternatives></ref><ref id="cit46"><label>46</label><citation-alternatives><mixed-citation xml:lang="ru">Yamagishi M., Shimoyamada Y., Nakatsuka T., Masuda K. Two R2R3-MYB genes, homologs of Petunia AN2, regulate anthocyanin biosyntheses in flower tepals, tepal spots and leaves of asiatic hybrid lily. Plant and Cell Physiology. 2010;51(3):463-474. DOI: 10.1093/pcp/pcq011</mixed-citation><mixed-citation xml:lang="en">Yamagishi M., Shimoyamada Y., Nakatsuka T., Masuda K. Two R2R3-MYB genes, homologs of Petunia AN2, regulate anthocyanin biosyntheses in flower tepals, tepal spots and leaves of asiatic hybrid lily. Plant and Cell Physiology. 2010;51(3):463-474. DOI: 10.1093/pcp/pcq011</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>
