Особенности структуры, экспрессии и взаимодействия с патогенами белка табака Nt-4/1 тема диссертации и автореферата по ВАК РФ 03.01.03, кандидат биологических наук Макарова, Светлана Сергеевна

• Макарова, Светлана Сергеевна
• кандидат биологических наук
• 2013

Оглавление диссертации кандидат биологических наук Макарова, Светлана Сергеевна

ОГЛАВЛЕНИЕ

СПИСОК СОКРАЩЕНИЙ

ВВЕДЕНИЕ

ОБЗОР ЛИТЕРАТУРЫ

1 Транспорт вирусов по растению

1.1 Транспортные белки вирусов растений

1.2 Внутриклеточный транспорт

1.2.1 Роль микротрубочек во внутриклеточном транспорте

1.2.2 Роль микрофиламентов во внутриклеточном транспорте

1.2.3 Везикулярный транспорт

1.3. Межклеточный транспорт

1.3.1 Строение ПД

1.3.2 Транспорт через ПД

1.4 Дальний транспорт вирусов

1.4.1 Строение проводящей системы растений

1.4.2 Дальний транспорт вирусов по флоэме

1.4.2.1 Вирусы, для дальнего транспорта которых не нужен белок оболочки

1.4.2.2 Вирусы, для дальнего транспорта которых необходим белок оболочки40

1.4.2.3 Транспортные системы поти- и клостеровирусов

1.4.2.4 Флоэмно-ограниченные вирусы

2 Белки растений - участники вирусной и вироидной инфекции

2.1 Белки растений, необходимые для транспорта вирусов

2.2 Белки растений, негативно регулирующие транспорт вируса по растению

2.3 Белки растений, участвующие в жизненном цикле вироидов

2.4 4/1 белок растений

МАТЕРИАЛЫ И МЕТОДЫ

РЕЗУЛЬТАТЫ И ОБСУЖДЕНИЕ

1.0 ИССЛЕДОВАНИЕ ЭКСПРЕССИИ ГЕНА Ш-4/1

2.0 СТРУКТУРНЫЕ ОСОБЕННОСТИ БЕЛКА N1-4/1

2.1 Исследование вторичной структуры белка N1-4/1

2.2 Ограниченный протеолиз белка N1-4/1 в клетках Е соИ

2.3 Анализ третичной структуры белка N1-4/1 методом дифференциальной

сканирующей калориметрии

9

3.0 ОЛИГОМЕРИЗАЦИЯ

3.1 Изучение способности к олигомеризации белка Nt-4/l и его делеционных мутантов методом динамического лазерного светорассеяния

3.2 Визуализация мультимерных комплексов, образуемых белком Nt-4/l, с помощью электронной микроскопии

4.0 ИССЛЕДОВАНИЕ БИОХИМИЧЕСКИХ СВОЙСТВ БЕЛКА Nt-4/l

4.1 Анализ РНК-связывающей активности белка Nt-4/l

4.2 in vitro взаимодействие белка AVirPl с РНК ВВКК

4.3 Картирование РНК-связывающего сайта белка Nt-4/1

4.4 Анализ РНК-связывающей активности белка Nt-4/l и его мутантов методом Норт-Вестерн гибридизации

4.5 Определение размера рибонуклеопротеидных комплексов методом динамического лазерного светорассеяния

5.0 ПРЕДСКАЗАНИЕ ТРЕТИЧНОЙ СТРУКТУРЫ БЕЛКА 4/1

ЗАКЛЮЧЕНИЕ

ВЫВОДЫ

СПИСОК ЛИТЕРАТУРЫ

СПИСОК СОКРАЩЕНИЙ

Названия вирусов

ВТМ - вирус табачной мозаики (Tobacco mosaic virus, TMV)

ВНПД - вирус некротической пятнистости дыни {Melon necrotic spot virus, MNSV)

BMT - вирус морщинистости турнепса (Turnip crinkle virus, TCV)

ВПГ - вирус пятнистости гвоздики (Carnation mottled virus, CMV)

ВШМЯ - вирус штриховатой мозаики ячменя (Barley stripe mosaic virus, BSMV)

BCBK - вирус скрученности верхушек картофеля (Potato mop-top virus, PMTV)

ВЖС - вирус желтухи свеклы (Beetyellow virus, BYV)

ВГТ - вирус гравировки табака (Tobacco etch virus, TEV)

ПЛВМ - полулатентный вирус мятлика (Poa semilatent virus, PSLV)

BKB - вирус короткоузлия винограда (Grapevine janleaf virus, GFV)

ВМЦК - вирус мозаики цветной капусты (Cauliflower mosaic virus, CaMV)

ВМБ - вирус мозаики бамбука (Bamboo mosaic virus, BaMV)

BOM - вирус огуречной мозаики (Cucumber mosaic virus, CMV)

ВОЖТ - вирус осветления жилок турнепса (Turnip vein clearing virus, TVCV)

ВМКГ - вирус мозаики коровьего горошка (Cowpea mosaic virus, CPMV)

XBK - Х-вирус картофеля (Potato virus X. PVX)

BMB - вирус мозаики вигны (Сом'реа mosaic virus, CMV)

BMA - вирус мозаики арахиса (Groundnut rosette virus, GRV)

ВПГТ - вирус пятнистой гнилостности томатов (Tomato spotted wilt virus, TSWV)

BHMKK - вирус некротической мозаики красного клевера (Red clover necrotic mosaic virus,

RCNMV)

ВШМП - вирус штриховатой мозаики пшеницы (Wheat streak mosaic virus, WSMV)

BC-J1K - вирус скручивания листьев картофеля (Potato leaf roll virus, PLRV)

ВЗЖС - вирус западной желтухи свеклы (Beet western yellow virus, BWYV)

BMP - вирус мозаики рапса (Oil-seed rape mosaic virus, ORMV)

BMoT - вирус мозаики турнепса (Turnip mosaic virus, TuMV)

ВПГТ - вирус пятнистой гнилостности томатов (Tomato spotted wilt virus, TSWV)

Названия вироидов

ВКХ - вироид карликовости хмеля (Hop stunt viroid, HSVd)

ВВКК - вироид веретеновидности клубней картофеля (.Potato spindle tuber viroid, PSTVd) ВСПА - вироид солнечной пятнистости авокадо (.Avocado sunblotch viroid, ASVd) ВЭЦ - вироид экзокортиса цитрусовых (Citrus exocortis viroid, CEVd) ВЛМП - вироид латентной мозаики персика (Peach latent mosaic viroid, PLMVd)

Другие сокращения

ГТД - плазмодесма

ТБ-транспортный белок

ВРК - вирусный репликативный комплекс

ТК - транспортный комплекс

БО - белок оболочки

вРНП-комплекс - вирусный рибонуклеопротеидный комплекс

вРНК - вирусная рибонуклеиновая кислота

оцРНК - одноцепочечная РНК

ЭПР - эндоплазматический ретикулум

АГ - аппарат Гольджи

МТ - микротрубочки

ПМ - плазматическая мембрана

Пл - пластиды

М - митохондрии

В - вакуоль

Я - ядро

КС - клеточная стенка Хл - хлоропласты СЭ - ситовидный элемент КС - клетка-спутница ПМЭ - пектинметилэстераза

ППС ПД- предельная пропускная способность плазмодесм ДБГ ТБ - двойной блок генов транспортных белков ТБГ ТБ - тройной блок генов транспортных белков

TGBpl - первый белок тройного блока генов

TGBp2 - второй белок тройного блока генов

TGBp3 - третий белок тройного блока генов

ORF - open reading frame, открытая рамка трансляции

GFP - green fluorescent protein, зеленый флуоресцентный белок

mRFP - monomeric red fluorescent protein, мономерный красный флуоресцентный белок

NCAP - non-cell-autonomous-protein, неклеточный автономный белок

МРВ2С - movement protein binding 2С, белок 2С, связывающий транспортный белок

БиФК - бимолекулярная флуоресцентная комплементация

KJICM - конфокальная лазерная сканирующая микроскопия

ДЛС - динамическое лазерное светорассеяние

КД - круговой дихроизм

ДСК - дифференциальная сканирующая калориметрия ПААГ - полиакриламидный гель

Введение диссертации (часть автореферата) на тему «Особенности структуры, экспрессии и взаимодействия с патогенами белка табака Nt-4/1»

ВВЕДЕНИЕ

Распространение вирусов по растению является сложным многостадийным процессом, состоящим из внутриклеточного транспорта к местам репликации, самой репликации и образования транспортного комплекса, с последующим его распространением по всему организму хозяина. Для осуществления транспорта вирусы кодируют специфические транспортные белки, которые обеспечивают межклеточный транспорт, а некоторые - и дальний. После достижения вирусным транспортным комплексом проводящей системы растения, происходит загрузка флоэмы и системное распространение с током ассимилятов в незараженные ткани, что в итоге приводит к заражению всего организма.

Поскольку генетический аппарат вирусов относительно небольшой и кодирует ограниченное число белков, становится очевидным, что для осуществления некоторых этапов своего жизненного цикла вирусам приходится использовать не только белки, кодируемые своим геномом, но и рекрутировать специфические факторы растения-хозяина. Особенно это очевидно в отношении вироидов - патогенов, представляющих собой кольцевую одноцепочечную молекулу РНК, не кодирующую белков.

Помимо широко известных взаимодействий компонентов вирусов и вироидов с белками аппарата транскрипции и трансляции клетки, а также транспортных комплексов вирусов с элементами цитоскелета, появляются данные о вовлеченности новых, неизученных белков растения-хозяина в жизненный цикл патогенов. На первый взгляд не всегда понятна роль таких взаимодействий для вируса или вироида. И даже не всегда известна функция таких белков для самого растения. Так обнаружено, что подавление экспрессии гена VirPl в протопластах N.tabacum и N.benthamiana приводило к нарушению репликации вироида веретеновидности клубней картофеля (ВВКК). Также получены данные, свидетельствующие в пользу того, что белок VirPl необходим для системного распространения ВВКК и вироида экзокортиса цитрусовых (Kalantidis et al., 2007). Известно, что ключевую роль в осуществлении дальнего транспорта умбравирусов и потивирусов играет взаимодействие ORF3 и VPg белков, соответственно, с ядрышковым белком фибрилларином. Снижение уровня экспрессии фибрилларина являлось причиной значительного подавления аккумуляции А вируса картофеля в системных листьях, но не приводило к полной отмене дальнего транспорта, как в случае с умбравирусной инфекцией (Kim et al., 2007а; Rajamaki and Valkonen, 2009).

Таким образом, более детальное исследование факторов растения-хозяина, вовлеченных в осуществление определенных стадий жизненного цикла патогенов, позволит приблизиться к пониманию устойчивости некоторых видов и сортов сельскохозяйственных культур к патогенам и, в перспективе, поможет создать новые сорта резистентные к инфекционным агентам.

В двугибридной дрожжевой системе при скрининге кДНК библиотеки Arabidopsis thaliana против транспортного белка (NSm) вируса пятнистой гнилостности томатов (ВПГТ) был обнаружен миозин- и кинезин-подобный белок с неизвестной функцией, названный At-4/1 (von Bargen et о/.,2001). В экспериментах по совместной экспрессии в' клетках N. benthamiana At-4/l слитого в одной рамке трансляции с GFP и слитого с YFP белка ТБГЗ полулатентного вируса мятлика было показано, что оба белка локализуются в одних и тех же структурах. В нашей лаборатории была клонирована полная кодирующая последовательность белка ортолога At-4/l из растений Nicotiana tabacum (Nt-4/l). С помощью конфокальной лазерной сканирующей микроскопии было определено, что белок локализуется в подвижных везикулярных структурах в цитоплазме и на периферии клетки рядом с ПД. В данной работе продолжаются исследования белка Nt-4/l - вероятного участника жизненного цикла патогенов растений.

Заключение диссертации по теме «Молекулярная биология», Макарова, Светлана Сергеевна

ВЫВОДЫ

1. Промотор Nt-4/l активен в тканях проводящей системы, преимущественно в жилках молодых листьев.

2. Альфа-спиральный белок Nt-4/l состоит из трех структурных доменов.

3. Экспериментально доказано, что белок "Nt-4/1 способен олигомеризоваться in vitro, формируя протяженные фибриллярные структуры.

4. Белок Nt-4/l обладает РНК-связывающей активностью, предпочтительно связывая несовершенные дуплексы РНК.

5. РНК-связывающая активность определяется участком белка, расположенном в его С-терминальном структурном домене.

ЗАКЛЮЧЕНИЕ

Поскольку вирусы являются облигатными паразитами растений, а кодирующая способность их генетического материала значительно ограничена, давно не вызывает сомнения факт эксплуатации вирусами клеточных структур и механизмов для обеспечения своей жизнедеятельности. Помимо трансляционного аппарата клетки вирусы растений используют систему цитоскелета, обеспечивающую направленную доставку по клетке (Boyko et al., 2000, 2007; Heinlein et al, 1995). При межклеточном транспорте вирусные транспортные белки изменяют предельную пропускную способность канала ПД для прохождения генома в соседнюю незараженную клетку. Для этого вирус взаимодействует с целым спектром белков растения-хозяина, такими как пектинметилэстераза, Р-1,3-глюканаза, шапероны (Chen et al., 2000; Dorokhov et al, 1999; Iglesias and Meins, 2000). Для обеспечения последующего системного распространения вирусному транспортному комплексу необходимо проникнуть в проводящую систему растения. Для этого он преодолевает дополнительные барьеры, отграничивающие разные типы клеток в составе проводящего пучка. В этом ему помогает взаимодействие с рядом дополнительных факторов растения-хозяина (Ueki and Citovsky, 2007).

Таким образом, становится очевидным, что в жизненном цикле вируса задействованы разнообразные факторы организма-хозяина, которые обеспечивают успешное инфицирование целого растения. Нарушение взаимодействия между хозяйским белком и белком вируса, лежит в основе устойчивости определенных видов и сортов растений к той или иной инфекции. Поэтому поиск новых белков - партнеров и исследование транспорта вирусов по растению приближает нас к понимаю механизма устойчивости многих дикорастущих предков культурных растений и способствует выведению промышленно ценных сельскохозяйственных сортов растений устойчивых к патогенам.

До сих пор не известна роль белка At-4/l в транспорте вирусов по растению, однако аминокислотная последовательность этого белка выявляет сходство с моторными белками растительной клетки: миозинами и кинезинами (von Bargen et al., 2001).

Исходя из накопленных на данный момент сведений, мы предполагаем, что белок 4/1 вовлечен в осуществление некоторых этапов распространения РНК-содержащих патогенов по растению.

Программа COFACTOR, показала, что по третичной структуре белок 4/1 маршанции похож на РНК-связывающий белок дрожжей She2p. В клетке дрожжей She2p обеспечивает направленную доставку ASH1 мРНК в две новообразованные гаплоидные споры, для дальнейшего определения их «пола» (Niessing et al., 2004; Muller et al., 2009). За такое специфическое узнавание и транспорт отвечает определенная пространственная структура в ASH1 мРНК (zip-code), представляющая собой двуцепочечный участок, разделенный одноцепочечными выпетливаниями, которую и распознает She2p белок (Chartrand et al., 1999; Olivier et al., 2005). Причем направленная доставка рибонуклеопротеидного комплекса She2p-ASH1 осуществляется по тяжам актиновых микрофиламентов при участии миозиновых моторов V класса (Takizawa et al., 1997).

По аналогии с She2p мы предположили, что белок 4/1 также взаимодействует с РНК, содержащей двуцепочечные участки. В данной работе in vitro методом задержки рибонуклеопротеидных комплексов в агарозном геле было установлено, что белок Nt-4/l in vitro образует комплексы с РНК, причем с большей эффективностью с РНК, содержащей одноцепочечные выпетливания и шпильки, в частности продемонстрировано, что Nt-4/l эффективно образует комплексы с РНК вироида веретеновидности клубней картофеля.

Ранее в нашей лаборатории с помощью KJ1CM было показано, что белок Nt-4/l, слитый в одной рамке трансляции с GFP, локализуется в виде везикулярных структур способных перемещаться через ПД в соседнюю клетку. Более того было обнаружено, что

109 при некоторых условиях локализация наблюдается исключительно в жилках инфильтрированных листьев (неопубликованные данные). Также нами было показано, что белок 4/1 слитый в одной рамке трансляции с GFP совместно локализуется с актиновой сетью, а при ее разрушении обработкой клеток цитохалазином В, движение 4/1 содержащих везикул прекращается. Полученные данные наводят на мысль о возможности участия белка Nt-4/l в распространение РНК-содержащих патогенов по растению.

В пользу этого предположения свидетельствуют данные, полученные с использованием трансгенных растений, экспрессирующих ген Р-глюкуронидазы под контролем промотора Nt-4/1. С помощью гистохимического окрашивания мы установили, что промотор работает во флоэмных клетках и паренхиме проводящих пучков стеблей и листьев. В молодых проростках с хорошо развитыми семядолями и двумя первыми листьями гистохимическое окрашивание наблюдалось только в жилках семядолей. Однако по мере развития растения, окрашивание пропадало в старых органах и появлялось в жилках молодых листьев, черешках, гипокотиле и, в последнюю очередь, в стебле. Вероятно, не одинаковый во времени уровень экспрессии гена Nt-4/l может влиять на течение инфекции.

В настоящее время ведутся исследования системного распространения вироида веретеновидности клубней картофеля на растениях, гиперэкспрессирующих белок Nt-4/l, и на растениях, в которых работа данного гена подавлена вирус-индуцированным сайленсингом. Результаты, полученные по завершении этих экспериментов, прольют свет на роль белка Nt-4/l в распространении РНК-содержащих патогенов по растению.

Список литературы диссертационного исследования кандидат биологических наук Макарова, Светлана Сергеевна, 2013 год

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