References

vavilov

Вавиловский журнал генетики и селекции

Vavilov Journal of Genetics and Breeding

2500-3259

Institute of Cytology and Genetics of Siberian Branch of the RAS

10.18699/VJ19.492

vavilov-2018

Research Article

Генетика растений

Plant genetics

Накопление витамина С в сочных плодах: биосинтез и рециркуляция, гены и ферменты

Vitamin C in fleshy fruits: biosynthesis, recycling, genes, and enzymes

Тяпкина

Д. Ю.

Tyapkina

D. Y.

https://orcid.org/0000-0002-6091-0765

Кочиева

Е. З.

Kochieva

E. Z.

Слугина

М. А.

Slugina

M. A.

mashinmail@mail.ru

Институт биоинженерии Федерального исследовательского центра «Фундаментальные основы биотехнологии» Российской академии наукРоссияInstitute of Bioengineering, Research Center of Biotechnology, RASRussian Federation

Институт биоинженерии Федерального исследовательского центра «Фундаментальные основы биотехнологии» Российской академии наук; Московский государственный университет им. М.В. ЛомоносоваРоссияInstitute of Bioengineering, Research Center of Biotechnology, RAS; Lomonosov Moscow State UniversityRussian Federation

2019

14052019

233270280

2019

Тяпкина Д.Ю., Кочиева Е.З., Слугина М.А.

Tyapkina D.Y., Kochieva E.Z., Slugina M.A.

This work is licensed under a Creative Commons Attribution 4.0 License.

https://vavilov.elpub.ru/jour/article/view/2018

L-аскорбиновая кислота (витамин C) – вторичный метаболит растений, выполняющий множество разнообразных функций как в растительных тканях, так и в организме человека. Основным источником витамина С в питании человека служат растения, и прежде всего плоды цитрусовых, шиповника, перца, смородины, томата, клубники, папайи, киви. Однако, несмотря на то что L-аскорбиновая кислота – важное биологически активное вещество, путь ее биосинтеза в растительной клетке был описан лишь в 2007 г. на примере модельного растения Arabidopsis thaliana. В настоящем обзоре рассмотрены известные на сегодняшний день пути биосинтеза L-аскорбиновой кислоты в тканях растений. Это L-галактозный, L-гулозный, галактуроновый и мио-инозитоловый пути. Наиболее изучен из них L-галактозный путь (путь Смирнова–Уилера), для которого определены все ферменты, катализирующие последовательную цепь реакций. Для других путей известна лишь предположительная последовательность метаболитов, при этом многие ферменты, катализирующие их превращение, еще не выявлены. Выделены ключевые гены, которые участвуют в биосинтезе и накоплении аскорбиновой кислоты в сочных плодах. Среди них ферменты L-галактозного пути (ГДФ-маннозофосфорилаза (GMP, VTC1), ГДФ-D-маннозо-3’5’-эпимераза (GME), ГДФ-L-галактозофосфорилаза (GGP, VTC2/VTC5), L-галактозо-1-фосфатфосфатаза (GPP/VTC4), L-галактозо-1-дегидрогеназа (GalDH) и L-галактоно1,4-лактондегидрогеназа (GalLDH)); ферменты D-галактуронового пути (NADPH-зависимая D-галактуронатредуктаза (GalUR)) и ферменты рециркуляции АК (дегидроаскорбатредуктаза (DHAR1) и монодегидро аскорбатредуктаза (MDHAR)). До сих пор нет однозначного описания всех путей биосинтеза и накопления L-аскорбиновой кислоты в плодах. В настоящее время нельзя однозначно утверждать, что какой-то из четырех известных путей биосинтеза аскорбиновой кислоты является преобладающим в плодах растений. Так, в плодах персика и киви основным является L-галактозный путь, тогда как в плодах винограда и клубники – по всей видимости, D-галактуроновый. В то же время у ряда растений, например цитрусовых или томата, по мере созревания плодов может происходить смена различных путей биосинтеза. Отмечается, что уровни накопления аскорбиновой кислоты зависят не только от биосинтеза, но и от скорости ее окисления и рециркуляции.

L-ascorbic acid (vitamin C) is a plant secondary metabolite that has a variety of functions both in plant tissues and in the human body. Plants are the main source of vitamin C in human nutrition, especially citrus, rose hip, tomato, strawberry, pepper, papaya, kiwi, and currant fruits. However, in spite of the biological significance of L-ascorbic acid, the pathways of its biosynthesis in plants were fully understood only in 2007 by the example of a model plant Arabidopsis thaliana. In the present review, the main biosynthetic pathways of vitamin C are described: the L-galactose pathway, L-gulose pathway, galacturonic and myo-inositol pathway. To date, the best studied is the L-galactose pathway (Smyrnoff–Wheeler pathway). Only for this pathway all the enzymes and the entire cascade of reactions have been described. For other pathways, only hypothetical metabolites are proposed and not all the catalyzing enzymes have been identified. The key genes participating in ascorbic acid biosynthesis and accumulation in fleshy fruits are highlighted. Among them are L-galactose pathway proteins (GDP-mannose phosphorylase (GMP, VTC1), GDP-D-mannose epimerase (GME), GDP-L-galactose phosphorylase (GGP, VTC2/VTC5), L-galactose-1-phosphate phosphatase (GPP/VTC4), L-galactose-1-dehydrogenase (GalDH), and L-galactono1,4-lactone dehydrogenase (GalLDH)); D-galacturonic pathway enzymes (NADPH-dependent D-galacturonate reductase (GalUR)); and proteins, controlling the recycling of ascorbic acid (dehydroascorbate reductase (DHAR1) and monodehydroascorbate reductase (MDHAR)). Until now, there is no clear and unequivocal evidence for the existence of one predominant pathway of vitamin C biosynthesis in fleshy fruits. For example, the L-galactose pathway is predominant in peach and kiwi fruits, whereas the D-galacturonic pathway seems to be the most essential in grape and strawberry fruits. However, in some plants, such as citrus and tomato fruits, there is a switch between different pathways during ripening. It is noted that the final ascorbic acid content in fruits depends not only on biosynthesis but also on the rate of its oxidation and recirculation.

L-аскорбиновая кислотавитамин Сплодыметаболизмгены биосинтеза аскорбиновой кислоты

L-ascorbic acidvitamin Cfruitsmetabolismthe key genes of ascorbic acid biosynthesis

This work was partially supported by grants from the Russian Foundation for Basic Research (#18-316-00033) and the Ministry of Science and Higher Education of the Russian Federation.

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The authors declare that there are no conflicts of interest present.