Факторы морозоустойчивости пшеницы – белки-ингибиторы рекристаллизации льда

Для озимой пшеницы зимостойкость – один из комплексных признаков, определяющих успешное возделывание этой культуры, а отвечающие за нее гены признаны высоко значимыми для селекционных работ. Накопление белков, препятствующих рекристаллизации льда (ice recrystallization inhibition proteins, IRIP), коррелирует с выживаемостью озимой пшеницы, что указывает на важность учета этого признака при получении более морозоустойчивых сортов. Значение IRIP для выживаемости определяется их способностью встраиваться в растущие кристаллы льда, что ограничивает формирование крупных ледяных конгломератов в тканях озимых растений. IRIP пшеницы, накапливающиеся преимущественно в апопласте листьев и узлов кущения в период холодовой акклимации, характеризуются типичной двойственной структурной организацией, определяющей как проявление IRI-активности, так и антипатогенные свойства. Молекула IRIP пшеницы содержит на С-конце консервативный несколько раз повторяющийся фрагмент NxVx(x)G, формирующий ответственную за связывание с поверхностью льда β-спираль; на N-конце расположены типичная для активируемых патогенами киназ LRR-последовательность, а также направляющий сигнальный пептид. Геном пшеницы содержит до 11 IRI-генов. Промотор генов TaIRI содержит типичные основные цисактивирующие элементы и некоторые элементы, реагирующие на абиотический стресс и гормоны. Изоформы ответственных за защиту от патогенов белков (pathogenesis related proteins, PRP), накапливающихся у озимой пшеницы в период холодовой акклимации, также обладают IRI-активностью. Экспрессия генов IRIP и PRP положительно коррелирует с холодоустойчивостью растений озимой пшеницы. Регуляция генов IRIP и генов PRP, активируемых холодом, согласно современным данным, АБК-независимая, но зависит от присутствия жасмоновой кислоты и от некоторых протеомных факторов транскрипции. В обзоре приведены примеры практического использования изолированных IRIP озимой пшеницы. Вопрос о факторах регуляции активности генов IRIP и активируемых холодом PRP является наименее разработанным на сегодняшний день. Связь этих белков с зимостойкостью пшеницы указывает на перспективность их дальнейшего изучения.

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