Внутриклеточный транспорт белков ТБГ2 и ТБГ3 вирусов растений тема диссертации и автореферата по ВАК РФ 03.00.06, кандидат биологических наук Щепетильников, Михаил Вячеславович

• Щепетильников, Михаил Вячеславович
• кандидат биологических наук
• 2006

Оглавление диссертации кандидат биологических наук Щепетильников, Михаил Вячеславович

СПИСОК СОКРАЩЕНИЙ

1. ВВЕДЕНИЕ

2. ОБЗОР ЛИТЕРАТУРЫ 9 2.1. Внутриклеточный везикулярный транспорт в клетках растений

2.1.1. Роль малых клеточных ГТФаз семейства Arf во внутриклеточном 10 транспорте

2.1.1.1. Малые клеточные ГТФазы подсемейства Arf

2.1.1.2. Структура малых клеточных ГТФаз Arf

2.1.1.3. Факторы обмена ГТФ (GEF)

2.1.1.4. Активаторные белки ГТФаз Arf (GAP)

2.1.1.5. Заякоривание ГТФаз Arf на мембране

2.1.1.6. Формирование COPI-везикул с участием малых клеточных ГТФаз 14 Afr

2.1.1.7. Клатрин транспортные везикулы

2.1.2. Малые клеточные ГТФазы Sar подсемейства

2.1.2.1. Структура белков Sar

2.1.2.2. Участие ГТФазы Sar в сборке COPII-окаймленных везикул

2.1.3. Белки SNARE

2.1.3.1. Классификация SNARE

2.1.3.2. Структурные особенности N-концевого домена белков SNARE

2.1.3.3. Структура SNARE комплекса

2.1.3.4. Белки SNARE растений

2.1.3.5. Механизм действия белков SNARE в везикулярном транспорте

2.1.3.6. Механизм разборки цис-SNARE комплекса

2.1.3.7. Функции и регуляция белков SNARE

2.1.3.8. Факторы, регулирующие активность SNARE

2.1.3.9. Внутриклеточная локализация белков SNARE

2.1.3.10. Сигналы внутриклеточной локализации

2.1.3.11. Взаимодействие с другими регуляторными белками

2.1.4. Белковые tethering факторы

2.1.4.1. Coiled-coil tethering факторы

2.1.4.2. Мульти-субъединичные tethering факторы

2.1.4.3. Механизм действия tethering факторов. Модель моста

2.1.4.4. Внутриклеточная локализация tethering факторов

2.1.5. Малые клеточные ГТФазы семейства Rab

2.1.5.1. Механизм действия ГТФаз Rab семейства

2.1.5.2. Структурные особенности регуляторных факторов белков Rab

2.1.5.3. Заякоревание на мембране

2.1.5.4. Внутриклеточная локализация ГТФаз Rab

2.1.5.5. Взаимодействие Rab с эффекторными белками

2.1.5.6. Транспортная функция ГТФаз Rab

2.1.6. Внутриклеточный везикулярный транспорт в высших растениях

2.1.6.1. Экзоцитозный путь

2.1.6.1.1. Транспорт между ЭПР и АГ

2.1.6.1.2. Антероградный COPII-зависимый транспорт

2.1.6.1.3. Модели везикулярного транспорта между структурами ЭПР и АГ

2.1.6.1.4. Ретроградный COPI-зависимый транспорт

2.1.6.1.5. Аппарат Гольджи

2.1.6.1.6. Сеть транс-Гольджи

2.1.6.1.7. Секреторный транспорт в вакуоли

2.1.6.1.8. Секреторный транспорт к плазматической мембране

2.1.6.2. Эндоцитозный путь 52 2.1.6.2.1. Роль сети транс-Гольджи и эндосом в эндоцитозе 54 2.2. Внутриклеточный транспорт вирусов, кодирующих тройной блок транспортных генов (ТБГ), в растениях

2.2.1 Вирусы, кодирующие тройной блок транспортных генов (ТБГ)

2.2.2 Структура генома ТБГ-содержащих вирусов

2.2.3. Структура доменов и сравнение последовательностей белков, 59 кодируемых тройным блоком транспортных генов

2.2.4. Два класса ТБГ-транспортных модулей

2.2.5. Особенности экспрессии тройного блока транспортных генов

2.2.6. Структура белков ТБГ

2.2.7. Активности ТБГ белков in vivo и их функции в вирусном 64 транспорте

2.2.8. Структура и транспортные функции РНП ТБГ-содержащих 65 вирусов

2.2.9. Роль белка оболочки

2.2.10. Дальний транспорт

2.2.11. Внутриклеточная локализация белков тройного блока 69 транспортных генов

2.2.12. Сигналы внутриклеточной локализации белков тройного блока 71 трапспортных генов

2.2.13. Ко-локализация белков ТБГ2 и ТБГЗ в клетке

2.2.14. Ко-локализация ТБГ1 и ТБГ2/ТБГЗ в клетках

2.2.15. Модель транспорта

Введение диссертации (часть автореферата) на тему «Внутриклеточный транспорт белков ТБГ2 и ТБГ3 вирусов растений»

Транспортные белкн вирусов способны транспортироваться к местам своейспецифической локализации и обеспечивают при этом перепое вирусного геномавнутри клетки, являясь, таким образом, удобной модельной системой для изучениямолекулярных механизмов внутриклеточного транспорта в клетках растений. Наосновании особенностей организации генома припято вьщелять группу вирусов стройным блоком транспортных генов (ТБГ), который содержит три гена, кодирующихбелки ТБГ1, ТБГ2 и ТБГЗ. Функциональное взаимодействие всех трех белковнеобходимо для транспорта вируса в зараженном растении.Объектом настоящей работы являлись белки ТБГ2 и ТБГЗ ТБГ-содержащихвирусов растений. Ранее была показана специфическая локализация белка ТБГ2 вмембранах эндоплазматического ретикулума, а также способность ТБГЗтранспортироваться на периферию клетки и формировать мембранные структуры вобласти плазмодесмы, но сигналы, нанравляющие их внутриклеточный транспорт, пебыли картированы. Для специфической внутриклеточной локализации транспортныхбелков вирусов растений, принадлежащих различным систематическим группам,необходимы структуры цитоскелета в клетках. Однако, возможный вклад актиновыхфиламентов и микротрубочек в процесс транспорта белков ТБГ2 и ТБГЗ не был до сихпор исследован. Кроме того, отсутствовали сведения об участии классическогосекреторного пути и отдельных внутриклеточных мембранных компартментов, как,например, аппарата Гольджи, в трапспорте этих белков.В настоящей работе изучались механизмы внутриклеточного транспорта белковТБГ2 и ТБГЗ вирусов растений. В ходе работы решались следующие основные задачи:изучение последовательностей белка ТБГ2, отвечающих за его специфическуюлокализацию в клетках растений, картирование районов белка ТБГЗ, необходимых дляего транспорта па периферию клеток растепий, изучение роли СОРП-везикул втранспорте белка ТБГЗ из ЭПР, определение способности белка ТБГЗ использоватьклассический секреторный путь для специфической локализации па периферии клетки,изучение влияния элементов цитоскелета па внутриклеточпый транспорт белка ТБГЗ.

Заключение диссертации по теме «Вирусология», Щепетильников, Михаил Вячеславович

1. Картированы носледовательности белков ТБГ2 и ТБГЗ, определяющие их

специфическую внутриклеточную локализацию. 2. Показано, что транспорт белка ТБЗ является COPI- и СОРП-независимым. 3. Показано, что внутриклеточный транснорт белка ТБГЗ происходит независимо

от структур цитоскелета клетки. 4. Установлено, что белок ТБГЗ транспортируется на нериферию клетки, минуя

классический секреторный путь.

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