Бесплатный автореферат и диссертация по биологии на тему

Реконструкция трансляционных инициирующих комплексов на эукариотических мРНК со структурированными 5'-нетранслируемыми областями

ВАК РФ 03.00.03, Молекулярная биология

Содержание диссертации, кандидата биологических наук, Дмитриев, Сергей Евгеньевич

Список сокращений.

Введение.

Обзор литературы

Глава 1. Механизмы инициации трансляции у зукариот.

1.1 Особенности эукариотических мРНК.

1.2 Сканирующая модель инициации трансляции.

1.3 Влияние вторичной структуры мРНК на инициацию трансляции.

1.4 Факторы инициации трансляции.

1.4.1 Факторы, ассоциированные с 4OS субчастицей. eIF2. eIF3. elFl. elFIA.

1.4.2 Факторы, участвующие в привлечении мРНК в инициаторный комплекс. eIF4F и его компоненты. eIF4A. eIF4B. eIF4H.

Прочие белки, участвующие в преодолении вторичных структур мРНК.

1.4.3 Факторы, участвующие в присоединении 60S субчастицы. eIF5. eIF5B

Прочие белки, участвующие в инициации на стадии 80S.

1. 5 Современная модель и перспективы изучения инициации трансляции.

Глава 2. Метод реконструкции инициаторных комплексов из очищенных компонентов и техника тоу-принтинга.

Результаты и обсуждение

1. Наличие в 5'-НТО мРНК даже незначительного числа спаренных оснований приводит к неэффективности сборки 48S при использовании стандартного протокола.

2. Фактор инициации eIF4B критичен для реконструкции 48S комплексов с мРНК, имеющими в 5'-НТО незначительные вторичные структуры.

3. Фактор инициации eIF4H не может заменить eIF4B в системе реконструкции 48S комплексов с мРНК, имеющими в 5'-НТО незначительные вторичные структуры.

4. Эффективная реконструкция 48S комплексов с мРНК, содержащими полный трипартитный лидер и 5'-НТО мРНК р-актина.

5. Незначительные вариации вторичной структуры 5'-НТО изменяют требования мРНК к концентрации фактора инициации eIF2.

6. Изучение перехода 4 8S инициаторного комплекса в 8 0S комплекс методом тоу-принтинга.

Материалы и методы

1. Реактивы и материалы.

2. Плазмидные конструкции.

3. Генноинженерные манипуляции.

4. Получение РНК-транскриптов.

5. Трансляция в лизате ретикулоцитов кролика.

6. Сборка инициаторных комплексов в лизате ретикулоцитов кролика.

7. Зондирование вторичной структуры мРНК.

8. Реконструкция инициаторных комплексов из очищенных компонентов

8.1. Выделение рибосомных субчастиц и факторов инициации.

8.2. Аминоацилирование инициаторной тРНК.

8.3. Собственно сборка инициаторных комплексов.

8.4. Тоу-принтинг.

8.5. Анализ комплексов методом центрифугирования в градиенте сахарозы.

9. Количественная обработка' результатов.

Выводы.

Благодарности.

Введение Диссертация по биологии, на тему "Реконструкция трансляционных инициирующих комплексов на эукариотических мРНК со структурированными 5'-нетранслируемыми областями"

Синтез белковой цепи у эукариот является сложным многостадийным процессом, требующим согласованной работы значительного числа клеточных компонентов. В большинстве случаев его регуляция осуществляется на уровне инициации, и отчасти по этой причине данный этап трансляции привлекает наиболее пристальное внимание исследователей. Сканирующая модель М. Козак, объясняющая в общих чертах принцип инициации трансляции на кеп-зависимых мРНК и считающаяся на сегодняшний день общепринятой, была предложена уже почти четверть века назад, однако её биохимический аспект до сих пор не представляет собой целостную картину. Каждый из известных на данный момент белковых факторов, принимающих участие в этом сложном процессе, по отдельности неплохо изучен как со структурной, так и с функциональной стороны, однако как работает весь ансамбль факторов в целом, какова последовательность событий при инициации трансляции, до сих пор остаётся до конца не выясненным.

Безусловно, одним из наиболее перспективных методов в изучении инициации трансляции является реконструкция инициирующих комплексов из очищенных компонентов. Его сочетание с техникой тоу-принта, применяемой для визуализации комплексов, позволяет убедиться не только в их наличии, но и в правильности положения рибосомы на стартовом AUG-кодоне. Данный подход совершенно незаменим при исследовании трансляционного аппарата млекопитающих, поскольку в этом случае использование генетических методов крайне затруднено. Метод блестяще проявил себя при исследовании внутренней инициации на мРНК некоторых вирусов. Однако его применение к изучению кеп-зависимой инициации до сих пор привело к успешной реконструкции 48S и 80S инициирующих комплексов лишь на мРНК р-глобина и на искусственной матрице с (САА)П-лидером. Все попытки добиться сборки на других кеп-зависимых мРНК как в нашей, так и в других лабораториях заканчивались неудачей, несмотря на то, что эти мРНК (в частности, мРНК р-актина, поздняя аденовирусная мРНК, мРНК белка теплового шока Hsp70 и ряд других) прекрасно транслируются в клеточном лизате.

В ходе данной работы было показано, что сложности в реконструкции инициирующих комплексов на таких мРНК определяются наличием в их 5'-НТО вторичных структур, причём даже незначительное количество спаренных GC-оснований приводит к снижению выхода 488-комплекса. Было проведено исследование с использованием модельных конструкций, показавшее, что наличие в 5'-лидерах мРНК незначительных вторичных структур резко повышает требования к концентрации фактора инициации elF4B, который, будучи очищен до гомогенного состояния, становится очень нестабильным и быстро теряет активность даже при хранении в глубокой заморозке, чем и объяснялись предыдущие неудачи в реконструкции. У таких матриц также наблюдались повышенные требования к факторам elF4F, elF4A и elF2. С учётом этих данных начальная методика реконструкции 488-комплексов из очищенных компонентов была модифицирована, в результате чего впервые удалось добиться сборки 48S на кеп-зависимых мРНК со структурированными 5'-НТО и продемонстрировать правильность положения 40S субчастицы на AUG-кодоне методом тоу-принта.

Другая часть работы была посвящена применению метода тоу-принта для анализа различий между 48S и 80S инициирующими комплексами. Было обнаружено, что паттерны тоу-принтов от 48S и 80S комплексов как в лизате ретикулоцитов кролика, так и в системе реконструкции из очищенных компонентов различаются, что позволило изучить накопление комплексов в лизате под действием разных антибиотиков, обнаружить паузу в переходе 48S в 80S и проанализировать распределение комплексов на трёх AUG-кодонах вируса энцефаломиокардита.

Обзор литературы

Заключение Диссертация по теме "Молекулярная биология", Дмитриев, Сергей Евгеньевич

Выводы.

1. Разработан протокол реконструкции 48S инициирующих комплексов с мРНК, содержащими в 5-НТО G-богатые вторичные структуры.

2. Обнаружено, что даже незначительное спаривание оснований в 5'-НТО мРНК приводит к повышенным требованиям к концентрации и качеству фактора инициации elF4B.

3. Фактор инициации elF4H не в состоянии заменить elF4B в системе реконструкции 48S инициаторных комплексов из очищенных компонентов.

4. Для инициации трансляции по механизму шунтирования на мРНК, содержащих трипартитный лидер поздних аденовирусных мРНК, недостаточно набора канонических факторов инициации.

5. мРНК, имеющие спаренные основания в 5-НТО, проявляют повышенные требования к концентрации elF2.

6. Обнаружена разница в картине тоу-принтов от 48S от 80S инициаторных комплексов.

7. Методом тоу-принтинга продемонстрирована пауза в процессе инициации трансляции на стадии перехода 48S комплекса в 80S.

8. Обнаружен феномен аберрантного распределения 48S комплексов по близко расположенным AUG кодонам мРНК EMCV при невозможности гидролиза GTP/GMP-PNP, связанного с elF2.

Благодарности.

Выражаю искреннюю признательность своему научному руководителю Ивану Николаевичу Шатскому за организацию работы, обучение методам и всестороннюю поддержку; благодарю Илью Теренина за существенную помощь в проведении исследования; Андрея Писарева, Марию Рубцову и Дмитрия Андреева - сотрудникам нашей лаборатории - за обсуждение результатов и ценные советы; Дунаевского Я.Е. - за помощь в выделении факторов инициации; Овчинникова Л.П. - за предоставление препарата белка р50; Народицкого Б.С. -за ДНК, содержащую трипартитный лидер аденовируса; Надеждина Е.В. - за препарат кДНК из мозжечка человека.

Работа выполнена при поддержке Российского фонда фундаментальных исследований (96-04-55009), РФФИ MAC (01-04-06503) и Федеральной целевой программы "Интеграция" (2000-1.5-88).

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