Молекулярно-генетическая организация междисков политенных хромосом дрозофилы на уровне ДНК и междиск-ассоциированных белков тема диссертации и автореферата по ВАК РФ 03.00.15, кандидат биологических наук Горчаков, Андрей Александрович

• Горчаков, Андрей Александрович
• кандидат биологических наук
• 2004

Оглавление диссертации кандидат биологических наук Горчаков, Андрей Александрович

ВВЕДЕНИЕ.

Глава 1. ОБЗОР ЛИТЕРАТУРЫ.

1.1. Организация политенных хромосом (общие сведения).

1.2. Морфологическая организация междисков.

1.3. Молекулярная организация междисков.

1.3.1. Содержание и конформационные особенности междисковой ДНК.

1.3.2. Транскрипционная активность междисков.

1.3.3. Анализ последовательностей нуклеотидов ДНК междисков.

1.3.4. Ассоциированные с междисками белки.

1.4. Метод Р-трансформации.

1.4.1. Явление гибридного дисгенеза.

1.4.2. Строение Р-элемента.

1.4.3. Транспозаза Р-элемента.

1.4.4. Механизм и регуляция транспозиции. Поддержание Р-цитотипа.

1.4.5. Выбор нового сайта.

1.4.6. Р-элемент-опосредованный трансгенез D. melanogaster.

1.4.7. Получение перестроек в районе инсерции транспозонов с использованием Р-элемент опосредованной рекомбинации у самцов.

1.4.8. Направленный мутагенез у дрозофилы.

1.4.9. Трансформация как подход к клонированию ДНК междисков.

Введение диссертации (часть автореферата) на тему «Молекулярно-генетическая организация междисков политенных хромосом дрозофилы на уровне ДНК и междиск-ассоциированных белков»

В настоящее время взаимосвязь структурной организации и генетической активности различных районов интерфазных хромосом несомненна. Однако многие детали этой взаимосвязи до сих пор не ясны. Изучение данной проблемы - одна из приоритетных задач современной биологии, свидетельством чего является большое количество работ и разнообразие выдвигаемых гипотез, посвященных этой теме.

Одной из наиболее удобных моделей для исследования интерфазных хромосом эукариот являются политенные хромосомы D. melanogaster. Эти хромосомы содержат до 1000 хроматид, лежащих вплотную друг к другу, каждая из которых соответствует интерфазной хромосоме (обзор Hill et al., 1987). Гомологичные хромосомы конъюгируюг, поэтому политенные хромосомы представлены в ядре гаплоидным числом. Упорядоченное расположение материала хромосом ведет к образованию поперечной исчерченности, представляющей собой чередование компактных (темных в световом микроскопе) хромомеров, или дисков, и менее плотных, и потому светлых, межхромомеров, или междисков. Именно эти особенности организации политенных хромосом определяют преимущество данной модели перед другими - специфический рисунок дисков и большая степень политении позволяют проводить точную привязку исследуемых последовательностей ДНК к определенным районам, с высоким разрешением картировав сайты связывания многих белковых факторов и проводить тонкий структурно-функциональный анализ хроматина.

Актуальность проблемы

К 60-ым годам XX века сложилось представление о хромомерах, как единственных носителях генов в хромосомах, поэтому подавляющее число работ было посвящено исследованию функциональной нагрузки именно хромомеров. Данные, указывающие на возможную постоянную транскрипцию материала междисков, - включение междисковыми районами 3Н-уридина, локализация в них РНК-полимеразы II, ДНК:РНК-гибридов и РНП-часгиц (см. раздел 1.3.2), - были получены лишь к 80-ым годам. Систематический молекулярный анализ организации междисков, включающий исследования конкретных последовательностей ДНК, картируемых в междисках, упаковки ДНК в них и ассоциированных белковых факторов был начат относительно недавно, и был обязан развитию двух методов - Рэлемент-опосредованной трансформации генома дрозофилы (см. раздел 1.4) и электронно-микроскопического (ЭМ) картирования инсерций транспозонов на политенных хромосомах (см. раздел 1.4.9). В частности, в результате ЭМ анализа политенных хромосом было обнаружено, что встройка транспозона в междисковый район может вызывать формирование нового тонкого диска, чго открывает уникальную возможность решения проблемы точного соотнесения молекулярных и цитологических данных в этих районах. Зная молекулярную организацию транспозона, можно клонировагь и определить последовательности ДНК, прилежащие к встройке, что позволяет проводить прямые эксперименты по изучению генетических и структурных особенностей междисковых районов.

Целью данной работы является комплексное изучение молекулярно-генетической организации междисковых районов политенных хромосом D. melanogaster, а также молекулярно-генетическое описание белкового комплекса Z4/CHRIZ, ассоциированного с междисками политенных хромосом дрозофилы. В связи с эгим были поставлены следующие задачи:

1. Определить молекулярно-генетическую организацию междискового материала: а) клонировать и определить последовательность нуклеотидов ДНК из междисковых районов ЗА4/А6 и 60Е8/Е10; б) провести сравнительный анализ последовательностей нуклеотидов междисковых ДНК; в) исследовать транскрипционную активность ДНК из района междиска 60Е8/Е10.

2. Синтезировать и проверить активность in vivo нового «донорного» транспозона дрозофилы для работы с междисковой ДНК в ненативном геномном окружении.

3. Провести комплексный молекулярно-генетический анализ гена Chriz, продукт которого предположительно ассоциирован с междисками: а) определить паттерн экспрессии гена Chriz\ б) определить паттерн хромосомной локализации комплекса Z4/CHRIZ в диплоидных и политенных клетках; в) получить и охарактеризовать мутантные аллели Chriz; г) идентифицировать эволюционно-консервативные домены белка CHRIZ.

Научная новизна. В данной работе впервые проведено клонирование и определение нуклеогидных последовательностей ДНК междисков ЗА4/А6 и 60Е8/Е10. Проведен сравнительный анализ ДНК исследованных междисков, последовахельноегей ДНК междисков, описанных ранее, и последовательное!ей ДНК дисков. Впервые на молекулярном уровне показана транскрипционная активность ДНК междиска. Впервые получены и протестированы три новых вектора - ве1аоры pFRT и pFRTV, разработанные для изучения междисков и вектор рМН, созданный для регулируемой экспрессии химерных белков в сис1еме Е. coli. Проведен детальный молекуляржыенетический анализ гена Chriz, белковый продукт которого образует комплекс с белком Z4 и специфически связывается с междисками политенных хромосом D. melanogaster.

Практическая ценность. В работе показана высокая технологичность подхода, предложенного ранее для клонирования ДНК междисков. Проведены первые прямые эксперименты по изучению транскрипционных свойств ДНК междисков. Впервые исследованы некоторые белковые составляющие хроматина междисков на молекулярном уровне. Результаты этих экспериментов имеют существенное значение для дальнейшего анализа структурно-функциональной организации интерфазных хромосом.

Апробация работы. Основные результаты данного исследования докладывались на ХШ и XIV Международных симпозиумах по структуре и функции клеючного ядра (Санкт-Петербург, 1999, 2002), 14th International Chromosome Conference (Wuerzburg, Germany, 2001), 28th International Meeting of The Federation of European Biochemical Societies (Istanbul, Turkey, 2002), 18th European Drosophila Research Conference / Eurofly (Gottingen, Germany, 2003), 45th Annual Drosophila Research Conference (Washington DC, USA, 2004), III Съезд ВОГиС (Москва, 2004).

Вклад автора. Основная часть экспериментов была выполнена авюром самостоятельно. Работа по клонированию и трансформации кДНК Chriz была проведена автором в лаборатории проф. X. Заумвебера при техническом содействии докт. X. Эггерта и г-жи Р. Гинапп (Институт Биологии, Берлин, Германия). Электронно-микроскопический анализ районов инсерций гранспозонов был полностью проведен д.б.н. В.Ф. Семешиным (ИЦиГ СО РАН). Анализ транскрипционной активности междиска 60Е8/Е10 методом Нозерн-блог гибридизации, а также начальные эксперименты по секвенированию ДНК междисков проводились совместно и в соавюрсгве с к.б.н. С. Л. Демаковым. Идея создания Р-транспозона pFRT принадлежит к.б.н. Ю. Б. Шварцу (Женевский Университет, Швейцария), трансформация этого вектора в геном дрозофилы была проведена им в лаборатории проф. С.В. Разина (ИБГ РАН, Москва). Часть работы по клонированию междисковых последовагельносгей проводилась автором в лаборатории проф. П. Слонимского (Центр Молекулярной Генетики CNRS, Жиф-сюр-Ивегт, Франция).

Обьем работы. Диссертация состоит из введения, обзора литературы, описания материалов и методов, результатов, обсуждения, а также выводов и списка цитируемой литературы, включающего 416 ссылок. Работа изложена на 164 страницах машинописною текста, содержит 10 таблиц и 32 рисунка.

Заключение диссертации по теме «Генетика», Горчаков, Андрей Александрович

Выводы

1. Последовательности ДНК изученных междисков уникальны в геноме и соответствуют либо регуляторным 5'-нетранскрибируемым областям генов, либо кодирующим областям активно экспрессирующихся генов «домашнего хозяйства».

2. Показано, что транспозон дрозофилы pFRT функционален и может быть использован для картирования последовательностей ДНК, способных формировать междиск в ненативном геномном окружении.

3. Установлено, что комплекс белков Z4/CHRIZ локализуется в большинстве междисковых районов политенных хромосом.

A.Chriz является жизненно-важным геном дрозофилы. Chriz экспрессируется на всех стадиях развития, а белок CHRIZ обнаруживается во всех исследованных клеточных типах, органах и тканях. Изменение экспрессии Chriz приводит к гибели клеток.

5. Установлено, что в диплоидных тканях комплекс Z4/CHRIZ ассоциирован исключительно с интерфазным хроматином. На стадии прометафазы белок CHRIZ колокализуется с микротрубочками веретена деления.

6. Показано, что полученные мутантные аллели Chriz являются слабыми супрессорами эффекта положения мозаичного типа.

7. В структуре CHRIZ обнаруживается несколько эволюционно консервативных доменов, включая хромодомен.

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