Регуляторные гены, опосредующие генетическую стабильность и радиочувствительность дрожжей Saccharomyces cerevisiae тема диссертации и автореферата по ВАК РФ 03.00.01, доктор биологических наук Колтовая, Наталия Алексеевна

• Колтовая, Наталия Алексеевна
• доктор биологических наук
• 2006

Оглавление диссертации доктор биологических наук Колтовая, Наталия Алексеевна

ВВЕДЕНИЕ.

ГЛАВА 1. МИЮХОНДРИАЛЬНЫЙ ГЕНОМ ДРОЖЖЕЙ Saccharomyces cerevisia обзор литературы).

1.1 Структура митохондриальной ДНК.

1.2 Структурная организация митохондрий.

1.3 Функции митохондрий.

1.3.1 Образование энергии.

1.3.1.1 Окислительное фосфорилирование.

1.3.1.2 РеМе-позитивные и реШе-пегативные дрожжи.

1.3 2 Образование активных форм кислорода.

1 3.3 Метаболизм железо-серных белков.

1.3.4 Апоитоз.

1.3.4.1 Программируемая гибель клеток млекопитающих.

1.3.4.2 Программируемая гибель дрожжей.

1.4 Наследование митохондрий.

1.5 Наследование митохондриальнои ДНК.

1.6 Реиликация митохондриального генома.

1.7 Сегрегация митохондриального генома.

1.8 Индукция мутаций petite бромистым этидием.

1.9 Репарация митохондриальной ДНК.

1.10 Нарушения митохондриального генома и его наследования вызывают заболевания человека.

1.11 Выделение мутаций srm, снижающих спонтанную г/го"-мутабильность.Ю

1.11.1 Получение мутаций srml-srm5.

1.11.2 Попарное взаимодействие мутаций srml, srm2, srm3 и srm5.

1 11.3 Влияние мутаций srml-srmS па спонтанную гАо'-мутабильность.

1.11.4 Влияние мутаций srml-srm5 па r/zo'-мутагенез под действием БЭ.

1.11.5 Влияние мутаций srml-srm5 на стабильность хромосом.

1.11.6 Влияние мутаций srml-srm5 на стабильность рекомбинантных нлазмид

1.11.7 Картирование мутации srm5.

ЭКСПЕРИМЕНТАЛЬНАЯ ЧАСТЬ

ГЛАВА 2. МАТЕРИАЛЫ И МЕТОДЫ.

2.1 Линии.

2.2 Плазмиды и библиотека геномной ДНК дрожжей.

2.3 Состав сред и растворов.

2.4 Основные методики.

ГЛАВА 3. ПОЛУЧЕНИЕ И СВОЙСВА МУТАЦИЙ ыт, ОДНОВРЕМЕННО СНИЖАЮЩИХ СПОНТАННУЮ гЬо- -МУТАБИЛЬНОСТЬ И МИ ГОТИЧЕСКУЮ СТАБИЛЬНОСТЬ ХРОМОСОМ.

3.1 Получение мутаций чгт8, эгтП, згт15 и БгтП.

3.2 Попарное взаимодействие мутаций чгт8, ¿>гт12, 5ш/и .мж5.

3.3 Феиотинические особенности отобранных мутантов чгт.

3.3.1 Форма клеток.

3.3.2 Время генерации.

3.3.3 Хронологическое старение.

3.3.4 Характер почкования.

3.4 Влияние мутаций на митохондриальную г/ш'-мутабилыюсть.

3.4.1 Влияние мутаций эгт на спонтанную г/го'-мутабильиость.

3.4.2 Влияние мутаций зпя на г/ш"-мута1 енез под действием бромистого этидия.

3.4.3 Влияние мутаций чгт на индукцию мутаций гИо' под действием УФ-света

3.5 Мутации бгш и точечный митохондриальный мута1 енез.

3.6 Рекомбинация и сегрегация митохондриальных генетических маркеров у мутантов 5гт.

3.7 Влияние мутаций згт на теми возникновения ядерных генных мутаций и митотической рекомбинации.

3.8 Влияние мутаций эгт на митотическую стабильность хромосом.

3 9 Влияние мутаций 5гт на митотическую стабильность рекомбинантных плазмид.

3.10 Влияние нарушений митохондриального генома на стабильность плазмид.

3.11 Обсуждение.

ГЛАВА 4. КАРТИРОВАНИЕ, КЛОНИРОВАНИЕ И СЕКВЕПИРОВАНИЕ ГЕНОВ 5ЯМ.

4.1Ген£Ш5.

4.1.1 Локализация гена ЯЯМЗ.

4.1.2 Влияние различных аллелей сс1с28 на скорость размножения.

4.1.3 Влияние различных аллелей сс!с28 на спонтанный г/го" -мутагенез.

4.1.4 Влияние различных аллелей cdc28 на радиочувствительность.

4.1.5 Определение нуклеотидной последовательности аллеля cdc28-srm.

4.1.6 Молекулярно-динамическое моделирование киназы.

4.1.7 Ген CDC28 (обсуждение).

4.2 Ген SRM8.

4 2.1 Картирование мутации ыт8.

4.2.2 Клонирование и идентификация нуклеотидной последовательности гена SRM8.

4.2.3 Определение нуклеотидной последовательности аллеля srm8/netl-srm.

4.2 4 Ген CFI1/NET1 (обзор литературы).

4.3 Ген SRMI2.

4.3.1 Клонирование и идентификация нуклеотиднои последовательности гена SRM12.

4.3.2 Определение нуклеотидной последовательности аллеля srml 2/hfll-srm.

4.3.3 Ген HFJI (обзор литературы).

4.4 Обсуждение.

ГЛАВА 5. МОДИФИКАЦИЯ ГЕНАМИ SRMЧУВСТВИТЕЛЬНОСТИ К у- ИЗЛУЧЕНИЮ И СНЕСКРОШТ-КОНТРОЛЯ.

5.1 Чувствительность мутантов srm к у-излучению.

5.1.1 Чувствительность гаплоидных клеток мутантов чгт к летальному действию у-излучения и УФ-света.

5.1.2 Чувствительность диплоидных мутантов srm к летальному действию у-излучения.

5.1.3 Взаимодействие мутаций srml, srm2 и srm5.

5.1.4 Влияние элиминации митохондриального генома на радиочувствительность штаммов дикого типа и мутантов srml и srm2.

5.1.5 Пострадиационное восстановление жизнеспособности мутантов srm2 и srm5.

5.1.6 Пострадиационное восстановление жизнеспособности дыхательно-недостаточных мутантов с генотипом SRMt, srm2, srm5.

5.2 Взаимодействие i еиов SRM с генами репарации и checkpoint-контроля.

5.2.1 Мутация cdc28-srm и эпистатические группы мутаций чувствительности к ионизирующему излучению RAD6 и RAD52.

5.2.2 Взаимодействие гена SRM5/CDC28 с checkpoint-генами RAD9, RAD17, RAD24, RAD53.

5.2.3 Взаимодействие между checkpoint-генами RAD9, RADI 7, RAD24.

5.2.4 Взаимодействие гена RAD53 с ¡енами RAD9, RAD 17, RAD24.

5 2.5 Взаимодействие гена SRM8/NET1 с checkpoint-генами RAD9, RAD17, RAD24,

RAD53.

5.2 6 Взаимодействие гена SRM12/HF11 с checkpoint-генами СОС28, RAD9, RAD24, RAD53.

5.3. Участие генов SRM в checkpoint-контроле.

5 3 1 Влияние мутаций srm на остановку клеточного цикла в GO и G1/S под действием УФ-света.

5.3.2 Участие генов SRM8 и SRM12 в остановке деления клеток в фазе S под действием гидроксимочевины.

5.3 3 Ген CDC28 необходим для остановки клеточного цикла в фаю G2 при повреждении ДНК у мутантов cdc9-l и cdc6-l при нспермиссивнои температуре.

5.3.4 Остановка деления клеток в фазе G2 при повреждении ДНК под действием у-излучения.

5.3.5 Влияние checkpoint-генов на мутабильность митохондриального генома 277 5.4 Обсуждение.

Введение диссертации (часть автореферата) на тему «Регуляторные гены, опосредующие генетическую стабильность и радиочувствительность дрожжей Saccharomyces cerevisiae»

Изучение генетического контроля поддержания целостности геномов, как ядерного, так и митохондриального, является фундаментальной проблемой генетики В поддержании наследственного материала участвуют различные процессы, в том числе процессы восстановления от эндогенных и экзогенных повреждений ДНК, часто наблюдается корреляция между генетической стабильностью и радиочувствительностью. Изучение действия ионизирующих излучений на живые организмы и генетического контроля воздействия особенно актуально в связи с последствиями радиационных аварий и использованием радиации в диагностических и терапевтических целях.

Дрожжи сахаромицеты служат удобной моделью для анализа важнейших механизмов, действующих в клетках высших эукариот. Функциональные гомологи генов, ассоциированных с некоторыми заболеваниями человека, и многих генов, необходимых для репарации повреждений ДНК, вызванных ионизирующей радиацией в клетках человека, вначале были охарактеризованы у дрожжей (Resnick, Сох, 2000, Yu et al, 1999, Foury, 1997). Результаты анализа изменений экспрессии в масштабе 1енома (genome-wide expression analysis) в ответ на изменения окружающей среды (DeRisi et al., 1997; Causton et al., 2001) и, в частности, на действие различных повреждающих агентов (Jelinsky et al., 2000), существенно расширяют возможности использования дрожжей в качестве модельного организма.

Систематический анализ делеционных мутантов выявил новые локусы, опосредующие у-чувствительность дрожжей (Bennett et al., 2001; Game et al., 2003) Идентифицированные 1ены разбиваются на группы ответственные за репликацию ДНК, генетическую рекомбинацию, ремоделирование хроматина, сегрегацию хромосом, checkpoint, транскрипцию, убиквитин-опосредованную деградацию белков, образование ядерных пор, поддержание клеточных стенок, активность аппарата Гольджи/вакуоли, цитокинез и активность митохондрий. Однако анализ делеционных мутантов имеет ограничения. Вне поля зрения остаются жизненно важные гены, для выделения которых необходимы иные подходы. Кроме того, при тотальном скринише возникают сложности с отбором генов слабо-чувствительных или снижающих жизнеспособность клеток. Часто именно такие гены оказываются регуляторными и являются многофункциональными и многомишенными, знание энзиматической функции которых недостаточно для понимания их роли в восстановлении клетки. Поэтому остается актуальным целенаправленное выделение и функциональный и генетический анализ отдельных генов.

При рассмотрении вопросов генетической стабильности и чувстви!елыюсти к повреждающим агентам, как правило, рассматривают повреждения ядерной ДНК. Однако в клетке наследственный материал обнаруживается и в органеллах. К настоящему времени накоплен значительный объем сведений относительно механизмов, опосредующих стабильное поддержание хромосомного аппарата клеток Сведения по генетике поддержания наследственных структур ор1анелл, в частности митохондрий, в целом более скромны. Для большинства эукариотических организмов митохондриальная ДНК (мтДНК) жизненно важна В частности, было показано, что делеции, дупликации и точечные мутации мтДНК вызывают заболевания у человека (Grossman, Shoubridge, 1996; Howell, 1999; Larsson, Clayton, 1995; Wallace, 1999; Zeviani et al, 1997). Случаи, в которых мутантная мтДНК обнаруживается в культуре клеток вместе с нормальной немутантнои мтДНК определяют как гетероплазмию. Однако отношение двух митохондриальных гаплотипов часто изменяется в течение жизни индивидуума и может сильно отличаться между различными типами клеток. Обнаружено, что некоторые ядерные гены причастны к накоплению нескольких классов делетированных молекул мтДНК у одного и тою же индивидуума Несмотря на важность в фундаментальных и клинических исследованиях, факторы, ответственные за распределение между нормальными и мутантными молекулами мтДНК в процессе развития индивидуума, относительно слабо изучены. Только один из них в настоящее время клонирован и охарактеризован (Nishino et al, 1999) Он кодирует тимидин фосфорилазу, фермент, участвующий в катаболизме тимидина и возможно необходимый для поддержания мтДНК. Направление исследования роли ядерно1 о генофона активно развивается (Dunbar et al., 1995).

Хотя на настоящий момент времени очень мало известно об ядерных генах, прямо или косвенно контролирующих поддержание мтДНК у высших эукариот, имеются многочисленные данные, касающиеся почкующихся дрожжей S cerevisiae, для которых характерна высокая частота выщепления дегенеративных митохондриальных мутантов petite. Обзор этих данных содержится в главе 1 настоящей работы. У этого вида дрожжей развитие митохондриальной 1енетики и изучение ядерного контроля целостности и передачи мтДНК более доступно по двум причинам: 1) эти дрожжи являются факультативными аэробами, и 2) просты и доступны в смысле классической и молекулярной генетики. Несмотря на статус удобной модели их использование ограничено в связи с наличием свойств дрожжей S cerevisiae, которые отличают их от высших эукариот (одноклеточный организм, факультативный аэроб, не поддерживает стабильно гетероплазмическое состояние и структура их мтДНК значительно отличается от таковой для высших эукариот). Тем не менее, можно полагать, что, по крайней мере, некоторые ядерные факторы, контролирующие целостность и передачу молекул мтДНК консервативны в процессе эволюции. Понимание ядерно-митохондриальнот взаимодействия у дрожжей может облегчить поиск соответствующих ядерных генов у высших эукариот.

Представления о функциональной значимости митохондрий в жизнедеятельности клетки постоянно расширяются. Большие усилия в настоящее время направлены на изучение роли митохондрий в апоптозе, запрограммированной i ибели клеток (Li et al, 1997). Внимание исследователей направлено также на изучение интеграции митохондрий в многочисленные клеточные процессы и динамику поведения в качестве субклеточной органеллы внутри клетки. Начинают вырисовываться механизмы репликации и экспрессии митохондриально1 о генома, но еще остается много вопросов в отношении возникновения мутаций, репарации и сегрегации мтДНК (Contamine, Picard, 2000). Главным достижением последних лет является понимание центральной роли мтДНК в некоторых заболеваниях человека. Наконец, расширяется ассортимент многочисленных антибиотиков с новыми мишенями, традиционно используемых для изучения биохимических механизмов в митохондриях. Таким образом, изучение стабильности митохондриального генома и роли митохондрий и ядерно-митохондриальных генетических взаимодействий в определении радиорезистентности и стабильности генома клеток представляет медицинский и общебиологический интерес.

Работа по идентификации ядерных генов (SRM - spontaneous rho~ mutability), необходимых для высокой мутабильности митохондриального генома, и выяснение роли этих генов в жизнедеятельности дрожжевых клеток была начата около трех десятков лет назад. Исследования выявили наличие механизмов, одновременно оказывающих заметное влияние на митотическую стабильность различных генетических структур, митохондриальных и ядерных, природных и рекомбинантных; и связи этих механизмов с регуляцией прохождения клеточного цикла. В настоящей работе продолжено выделение новых i енов SRM. Для целенаправленного выделения мутаций, влияющих на генетическую стабильность различных наследственных систем, был разработан специальный селективный метод. Помимо изучения непосредственно генетического контроля стабильности наследственных структур, сама генетическая стабильность использовалась в качестве инструмента для выделения генов, контролирующих радиочувствительность клеток дрожжей. Исследования, описанные в диссертации, впервые продемонстрировали тесную связь между изменчивостью митохондриального генома, поддержанием хромосом и радиочувствительностью клеток дрожжей, которая, как представляется, заслуживает специального исследования.

Введение митохондриальною генома в рассмотрение генетической стабильности позволяет усилить селекцию регуляторных генов. Поскольку репарация мтДНК осуществляется в основном специфическими митохондриальными ферментами, то мутации влияющие на стабильность одновременно всех наследственных структур, ядерных и митохондриальных, скорее всею затрагивают регуляторные гены Действительно, нами впервые было показано участие генов, контролирующих локализацию деацетилазы (NET1), стабильность ацетилтрансферазных комплексов (HFI1), а также жизненно-важною гена протеинкиназы CDC28, в поддержании целостности генома и контроле радиочувствительности. Влияние гена HFI1 на у-чувствительность было позднее обнаружено и при изучении делеционных мутантов (Bennett et al., 2001).

Известно, что существенный вклад в стабильность генома и определение радиочувствительности вносит репарация и checkpoint-контроль остановки клеточного деления. Значительный интерес исследователей к данному регуляторному механизму связан, в частности, с широко распространившимися представлениями о тесной связи между нарушениями checkpoint и малигнизацией клеток Несмотря на значительные успехи последних лет в этих исследованиях многие конкретные детали остаются неясными. У дрожжей S cerevisiae генетический контроль механизмов репарации и checkpoint наиболее изучен. Однако эпистатические взаимодействия checkpoint-генов, отражающиеся на у-чувствительности клеток, исследованы недостаточно. В настоящей работе среди генов SRM выделены новые гены, участвующие в этих процессах. Анализ взаимодействия между checkpoint-i енами позволил выявить помимо остановки клеточного цикла их дополнительную роль в определении уровня радиочувствительности и охарактеризовать их принадлежность к путям, определяющим чувствительность клеток дрожжей к повреждающему действию ионизирующей радиации. Знаиие энзиматических активностей, определяемых выделенными генами, позволяет предположить, что обнаруженные генетические и радиобиологические эффекты вызваны модификациями белков, в том числе, определяющих нуклеосомную структуру хромосом или нуклеоидную структуру мтДНК. Полученные данные хорошо вписываются в современные представления о молекулярных механизмах репарации и checkpoint, согласно которым для функционирования репарационных комплексов и комплексов, запускающих checkpoint, необходима предварительная модификация хроматина в области повреждения ДНК. Проверка высказанных предположений актуальна и открывает большие перспективы проводимых исследований.

Заключение диссертации по теме «Радиобиология», Колтовая, Наталия Алексеевна

выводы

1. Разработан метод отбора мутантов со сниженной стабильностью хромосом и rho" -мугабильностью митохондриальною генома С помощью этого метода получены новые ядерные генные мутации srm (spontaneous rho" mutability). Мутации srm снижаю1 часгогу спонтанных миюхондриальных мутаций rho" (в среднем на порядок), митотическую стабильность природных хромосом у дисомиков (снижение до двух порядков) и рекомбинантных минихромосом (в несколько раз)

2 Клонированы и секвенированы 1ены SRM8 и SRM12, показана их аллельиость 1снам регулятора выхода из мигоза и локализации деацетилазы NET1 и компонента гистонацетилтрансферазных комплексов HF11, соответственно. Определена нуклеотидная последовательность мутантных аллелей srm8/netl-srm (+2А), srm!2/hfil-srm [L196P Stop] и srm5/cdc28-srm [G20S]. Мутация srm5/cdc28-srm предсгавчяет собой единичную замену глицина на серии в консервативной G-богатои петле киназы CDC28. Проведегго МД моделирование мутантною белка с использованием киназы человека CDK2 в качестве модельной системы. Показано, что упомянутая аминокислотная замена вьвываег заметные изменения структуры киназы, которые влекут за собой изменения взаимодействия кина}ы с циклинами, субстратами и молекулой АТФ

3 Впервые показапо участие генов CDC28, NET1 и HFI1 в регуляции радиорезистентности клеток Мутации cdc28-srm, netl-srm, hfil-srm снижают резистентность клеток к летальному действию у-излучепия, мутация sгт2 повышает радиорезистентность клеток Мутации srm2, netl-srm, hfil-srm снижают частоту индукции мутаций и рекомбинации. Влияние мутации cdc28-srm на чувствительность клеток дрожжей к у-излучению, а также на спонтанную и индуцированную рекомбинацию позволяет предположить участие гена CDC28 в рекомбипационной репарации Ген CDC28 не относится к RAD6- и /?/Ш52-эпистатическим группам генов

4 Checkpoint-rciibi RAD9, RAD17, RAD24 и RAD53 относятся к одной эпистатической группе генов, опосредующих чувствительность к у-излучению диплоидных штаммов в стационарной фазе клеточною цикла. Мутация гена CDC28 взаимодействует энистатически с мутациями checkpoint-iehob RAD9, RAD 17, RAD24, но аддитивно с мутацией i сна RAD53 в отношении чувствительности к у-излучеиию. У двойных мутантов cdc28-srm rad53 у-чувствительность выше, чем у каждого из соответствующих одиночных мутантов, и выше, чем у двойных мутантов rad9A cdc28-srm и rad9A rad53. Можно предположить существование /¿Л£Н>-независимого ретуляторною механизма, блокируемого при одновременном повреждении киназ

CDC28 и RAD53. Мутация 1ена NET1 взаимодействует эпистатически с мутациями генов RAD9 и RAD53, но аддитивно с мугациями генов RADI7 и RAD24 Мутация гена HF11 также взаимодействует эпистатически с мутацией 1ена RAD9, но аддитивно с мутацией 1ена RAD24. Таким образом, эффекты мутации генов RAD17, RAD24, RAD53, CDC28, NET1, HFII эпистатичны эффекту мутации гена RAD9 Изученные гены контролируют один механизм определения радиочувствителньости, но образуют разветвленную сеть контроля чувствительности к у-излучению

5 Впервые показано участие 1енов CDC28, NET! и HF11 в chcckpoint-конфоле Мутации cdc28-srm, netl-srm, hfil-srm сокращают остановку клеточного цикла при повреждении ДНК и влияют на G0/G1 (cdc28-srm netl-srm), Gl/S (cdc28-srm, netl-srm, hfil-srm), S (netl-srm, hfil-srm) и G2/M (cdc28-srm) checkpoint-контроль По-видимому, 1ены CDC28, NET1 и 11F11/ADA1 участвуют в формировании отклика клетки на повреждения ДНК через регуляцию репарации и остановку клеточного цикла (checkpoint).

6. Мутации в checkpoint-i енах RAD9, RAD17, RAD24, RAD53 вызывают повышение частоты возникновения мугаций rho~ Мутации в генах CDC28, NET1, HFI1, хотя и нарушают checkpoint-контроль, по приводят к снижению частоты возникновения мутаций rho. По-видимому, влияние мутаций генов CDC28, NET!, HF11 на митохондриальную муибильносгь не обусловлено непосредственно нарушением задержки клеточного цикла Анализ показал, что 1ен CDC28 принимает участие в регуляции рекомбинации и сегрегации мтДНК

7. Выявлены ядерно-митохондриальные 1енетические взаимодействия с участием генов SRM, способные модулировать1 а) резистентность клекж дрожжей к повреждающему действию ионизирующей радиации, б) поддержание клетками рскомбинашных диетических структур и в) жизнеспособность клеток с возрастом культур.

8. Представленные данные показывают, что эффекты мутаций генов CDC28, NET1 и HFI1, являющихся пюбальными регуляторами множественных жизненно важных клеточных механизмов, проявляются, в частности, в координированных изменениях генетической стабильности и радиочувствительности клеток, а также в нарушениях рефляции клегочною цикла и, вероятно, опосредованы химической модификацией структурных эпемептов хроматина (в частности, фосфорилированием, апеллированием и деацегилированием белков) Полученные данные позволяют предположить существование общих регуляторных элементов упаковки хромосом и мтДНК

ЗАКЛЮЧЕНИЕ

В настоящей работе разработан метод идентификации генов, влияющих на стабильность митохондриального 1енома и хромосом при митотическом делении клеюк дрожжей S cerevisiae С помощью этой селективной системы была получена колчекция ядерных шшых мутаций srm (spontaneous rho' mutability), влияющих на надежность передачи генетического материала, как митохондриального, так и ядерного (природных хромосом и рекомбинантных структур). Некоторые из них вызывают морфологические изменения, снижение скорости размножения и жизнеспособности клеток. Мутации ыт характеризуются широкими плейотропными проявлениями, что свидетельсгвуег о принадлежности генов к высоким ступеням иерархии. Действительно, три идентифицированных гена SRM5, SRM8 и SRM12 аллельны 1енам CDC28, NEl'l и 11F11, соответственно. Известно, что протеинкиназа CDC28 играет центральную роль в регуляции клеточного цикла; белок NEFI регулирует выход из митоза и локализацию деацетилазы Sir2p; SRMI2/ADAI/HFÜ является компонентом i истонацетилтрансферазных комплексов SAGA, ADA1/GCN5, SLIK и участвует в экспрессии большей части iciiob дрожжей.

Данные, представленные в настоящей работе, показывают, что у дрожжей наблюдается координация стабичыюсти различных генетических структур, но принадлежат ли 1ены SRM к единой системе, контролирующей ¡енетическую стабильность, или механизм их действия различен, несмотря на схожесть фенотипа, пока не известно. Можно предположить, что ре1уляторная роль наиболее и ученных нами генов SRM опосредуется через модификацию нуклеосомной структуры хромосом и нуклеоидной структуры мтДНК. Известно, что субстратами протеинкиназы являются компоненты модифицирующих и ремоделирующих хромосомы комплексов CAF1 (R112p), FACT (Sptlóp), SWI/SNF (Sip2p - Snflp), SAGA (Ngglp/Ada3p), SLIK (Ngglp/Ada3p), a также белки, участвующие в сайленсипге (Net 1 р, Sir4p, ORC). Помимо модифицирования хроматина SAGA-комплекс необходим для сборки ремоделирующих комплексов Эти факторы принимают участие, как в активации транскрипции, так и в саиленсинге. Причем SAGA-комплекс служит не только активатором транскрипции, но и барьером, определяющим область распространения сайленсинга Таким образом, белок Hfilp может оказывать влияние на активацию транскрипции, сайленсиш и активацию ретро1радной системы. Ремоделирование также окашвает влияние на репликацию ДНК. Нарушение правильной упаковки и регуляция упаковки ДНК может влиять на стабильность хромосом. Эти комплексы могут также участвовать в регуляции процессов, протекающих в митохондриях В митохондриях обнаружены белки сайленсинга Sir4p, белки конденсипа

Smc2p (взаимодействует с Smc4p, являющимся субстратом киназы Cdc28p), и белок Spt7p, входящий в состав комтексов SAGA и SLIK. Выделена 1еликаза Rrm3p, локализованная в ядре и митохондриях и облегчающая репликацию в специфических местах (молчащие ARS, области саиленсинга, CEN). Можно предположить, что влияние генов CDC28, HF11, NET1 на стабильность хромосом и мтДНК опосредовано их влиянием на структуру и репликацию ДНК в ядре и митохондриях. Стабильность и копийность мтДНК регулируется несколькими механизмами, в том числе опосредованными ПКА-АЫ2р и Mecl/Rad53-checkpoint Оба пути, по-видимому, зависят от киназы CDC28 Однако высказанные предположения нуждакмся в прямой проверке

Актуальность проводимых исследонаний обусловлена открытием в конце 80-ых юдов у дрожжей S cerevisiae ре1уляторною механизма высокого уровня иерархии, контролирующего прохождение клеточного цикла, репликацию и репарацию, так называемого checkpoint-контроля (Weinert, Hartwell, 1988). При наличии эндогенных ичи экзогенных повреждений ДНК checkpoint-контроль осуществляет остановку клеточного цикла и активацию репарационных процессов. После исправления повреждения клетка возобновляет деление. Последствиями нарушений checkpoint-регуляции клеточного деления в ответ на повреждения ДНК во многих случаях являются снижение точности митотической передачи компонентов наследственного аппарата и повышение чувствительности клеток к летальному действию ДНК-трогшых агентов Значительный интерес исследователей к рассматриваемому peí уляторному механизму связан, в частности, с широко распространившимися представлениями о тесной связи между нарушениями checkpoint-контроля и малигнизациеи клеток. Несмотря на значительные успехи последних лет в этих исследованиях многие конкретные детали остаются неясными.

Некоторые выделенные нами мутации srm сопровождаются не только снижением частоты спонтанных митохондриальных мутаций petite и падением митотической стабильности природных и рекомбинантных ядерных 1енегических струкгур, но и повышением чувствительности к летальному действию у-излучения. Нами впервые показано участие генов CDC28, NET1 и HF11 в определении уровня радиочувствительности и в checkpoint-контроле. В дальнейшем нредаавляе1ся целесообразным детальный анализ peí улягорной роли 1енов SRM и исследование функций этих i енов в chcckpoint-регуляции, репарации и наследовании мтДНК. Изучение механизмов стабилизации митохопдриального генома приобретает все большую важность в свете интенсивных исследований митохондриальных заболеваний человека.

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