Адаптивное значение и происхождение генов биосинтеза флавоноидов в зерновке культурных злаков

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Большинство культурных злаков – кукуруза, рис, пшеница, ячмень, овес и рожь представлены многочисленными сортами без антоциановой окраски или со слабой окраской вегетативных органов и/или зерновок. Интенсивной окраской растений и/или зерна обладают редкие местные расы и дикорастущие родственные виды. Окраска зерновок связана с биосинтезом окрашенных флавоноидов в материнских (перикарп и теста) и гибридных (алейрон) тканях зерновки и контролируется доминантными аллелями регуляторных генов, кодирующих консервативные транскрипционные факторы семейств MYB, bHLH-MYC и WD40, составляющих белковый регуляторный комплекс MBW. Исследованиями последних лет доказано участие неокрашенных и окрашенных флавоноидов в реакции растений на биотические и абиотические стрессовые факторы, установлена их функциональная значимость в составе пищевых продуктов из цельного зерна. Однако многие вопросы по адаптивности и полезности антоцианов остаются без ответа или даже не ставятся. В частности, не ясны причины, по которым в ходе одомашнивания и селекции зерновых злаков не получили широкого распространения доминантные аллели регуляторных генов окраски перикарпа, несмотря на то, что именно этим генам уделяется особое внимание в связи с оздоровительным эффектом зернового питания. Настоящая статья посвящена рассмотрению сходства и специфики генетического контроля биосинтеза флавоноидов в зерновке у трех родственных культурных злаков – пшеницы, ячменя и ржи, их биологической роли в ходе развития зерновки и прорастания семян.

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А. Н. Буланов

Санкт-Петербургский государственный университет; Институт общей генетики им. Н. И. Вавилова Российской академии наук

Автор, ответственный за переписку.
Email: an.bulanov20002014@gmail.com
Россия, Санкт-Петербург, 199034; Москва, 119991

А. В. Войлоков

Институт общей генетики им. Н. И. Вавилова Российской академии наук

Email: av_voylokov@mail.ru
Россия, Москва, 119991

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2. Рис. 1. Схема пути биосинтеза флавоноидов у растений. Классы неокрашенных флавоноидов выделены оттенками серого цвета, окрашенные флавоноиды (флобафены, проантоцианидины, антоцианидины и антоцианы) выделены цветами. Путь биосинтеза флавоноидов у растений начинается, как и у всех фенольных соединений, с последовательных превращений фенилаланина, катализируемых фенилаланинаммиаклиазой (PAL, phenylalanine ammonia-lyase), 4-гидроксилазой коричной кислоты (C4H, cinnamic acid 4-hydroxylase) и 4-кумарат-КоА-лигазой (4CL, 4-Coumarate-CoA ligase). Халконсинтаза (CHS, chalcone synthase) опосредует синтез нарингенин-халкона – предшественника большинства флавоноидов, а совместное действие халконсинтазы и халконредуктазы (CHR, chalcone reductase) обусловливает синтез халкона-предшественника изофлавоноидов изоликвиринтигенина. Халконы дают начало всем классам флавоноидов, что обусловливается работой следующих ферментов: халконизомеразы (CHI, chalcone isomerase), флавонсинтаз 1 и 2 (flavone synthase 1 and 2), изофлавонсинтазы (IFS, isoflavone synthase), флаванон-3-гидроксилазы (F3H, flavanone-3-hydroxylase), флавоноид-3′- и –3′5′-гидроксилаз (F3′H / F3′5′H, flavonoid-3′- / 3′5′-hydroxylase), флавонолсинтазы (FLS, flavonol synthase), дигидрофлавонолредуктазы (DFR, dihydroflavonol reductase), лейкоантоцианидинредуктазы (LAR, leucoanthocyanidin reductase) и антоцианидинредуктазы/ лейкоантоцианидиндиоксигеназы (ANR, anthocyanidin reductase / LDOX, leucoanthocyanidin dioxygenase). Антоцианы синтезируются посредством модификации антоцианидинов посредством метилтрансфераз (MT, methyltransferase), ацилтрансфераз (AT, acyltransferase) и гликозилтрансфераз (GT, glycosyltransferase). Флаваноны и дигидрофлавонолы могут иметь разное количество гидроксильных групп, что опосредуется работой флавоноид-3′- и -3′5′-гидроксилаз и впоследствии приводит к синтезу различных флобафенов, флавонов, флавонолов, проантоцианидинов и антоцианидинов. Так, цианидин характеризуется наличием гидроксильной группы в 3′-положении, а дельфинидин – и в 3′-, и в 5′-положении. Экспрессия различных структурных генов биосинтеза флавоноидов напрямую активируется посредством связывания с их промоторами MBW-комплексов, состоящих из двух димеризованных молекул транскрипционного фактора bHLH-MYC, связанных каждая с одной молекулой транскрипционного фактора R2R3-MYB. Такой комплекс стабилизируется белком WD40.

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