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«Антипролиферативное действие карнозина и его производных на опухолевые клетки нейрального происхождения ...»

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ФЕДЕРАЛЬНОЕ ГОСУДАРСТВЕННОЕ БЮДЖЕТНОЕ УЧРЕЖДЕНИЕ «НАУЧНЫЙ ЦЕНТР НЕВРОЛОГИИ» РОССИЙСКОЙ АКАДЕМИИ МЕДИЦИНСКИХ НАУК

На правах рукописи

РЫБАКОВА ЮЛИЯ СЕРГЕЕВНА

Антипролиферативное действие карнозина и его производных на опухолевые клетки нейрального происхождения

03.01.04 – биохимия

Диссертация на соискание ученой степени кандидата биологических наук

Научный руководитель:

доктор биологических наук

Федорова Татьяна Николаевна

МОСКВА 2013

СОДЕРЖАНИЕ

TOC \h \z \t "Глава,1,Раздел,2,Подраздел,3" ГЛАВА 1. ОБЗОР ЛИТЕРАТУРЫ. PAGEREF _Toc368852406 \h 11РАЗДЕЛ 1.1. Редокс-статус клетки определяется соотношением внутриклеточных про- и антиоксидантов. PAGEREF _Toc368852407 \h 111.1.1. Активные формы кислорода и их значение в жизни клетки. PAGEREF _Toc368852408 \h 111.1.2. Основные представители антиоксидантной системы клетки и их роль в поддержании редокс-статуса. PAGEREF _Toc368852409 \h 13РАЗДЕЛ 1.2. Редокс-регуляция клеточной пролиферации. PAGEREF _Toc368852410 \h 171.2.1. Клеточный цикл и его регуляция. PAGEREF _Toc368852411 \h 181.2.2. Редокс-регуляция клеточного цикла. PAGEREF _Toc368852412 \h 26РАЗДЕЛ 1.3. Современное состояние знаний о противоопухолевом эффекте короткоцепочечных пептидов. PAGEREF _Toc368852413 \h 361.



3.1. Карнозин и его производные. Общая характеристика. PAGEREF _Toc368852414 \h 361.3.2. Антиоксидантные свойства карнозина и его производных. PAGEREF _Toc368852415 \h 391.3.3. Эффект карнозина и его производных на пролиферацию нормальных и опухолевых клеток. PAGEREF _Toc368852416 \h 421.3.4. Пинеалон. Общая характеристика и свойства. PAGEREF _Toc368852417 \h 45ГЛАВА 2. МЕТОДЫ ИССЛЕДОВАНИЯ. PAGEREF _Toc368852418 \h 472.1. Культуры клеток. PAGEREF _Toc368852419 \h 472.2. Исследуемые соединения. PAGEREF _Toc368852420 \h 482.3. Исследование клеточной пролиферации. PAGEREF _Toc368852421 \h 482.3.1. Время удвоения популяции. PAGEREF _Toc368852422 \h 482.3.2. Эффективность посева и доля выживших клеток. PAGEREF _Toc368852423 \h 492.4. Проточная цитометрия. PAGEREF _Toc368852424 \h 492.4.1. Измерение уровня АФК. PAGEREF _Toc368852425 \h 502.4.2. Определение доли мертвых клеток. PAGEREF _Toc368852426 \h 512.4.3. Анализ клеточного цикла. PAGEREF _Toc368852427 \h 512.4.4. Двухпараметрический анализ клеточного цикла. PAGEREF _Toc368852428 \h 532.5. Иммуноблоттинг. PAGEREF _Toc368852429 \h 532.6. Определение активности MnСОД. PAGEREF _Toc368852430 \h 542.7. Определение активности каталазы. PAGEREF _Toc368852431 \h 542.8. Полимеразная цепная реакция (ПЦР) в режиме реального времени. PAGEREF _Toc368852432 \h 552.9. Статистическая обработка результатов. PAGEREF _Toc368852433 \h 55ГЛАВА 3. РЕЗУЛЬТАТЫ ИССЛЕДОВАНИЯ. PAGEREF _Toc368852434 \h 56РАЗДЕЛ 3.1. Изучение характера действия карнозина на пролиферацию клеток. PAGEREF _Toc368852435 \h 563.1.1. Карнозин снижает количество клеток РС-12 в культуре. PAGEREF _Toc368852436 \h 563.1.2. Карнозин снижает уровень АФК в клетках культуры РС-12. PAGEREF _Toc368852437 \h 583.1.3. Карнозин и его производные индуцируют накопление клеток РС-12 в S и G2/М фазах клеточного цикла. PAGEREF _Toc368852438 \h 593.1.4. Изучение воздействия синтетического трипептида пинеалона на клеточный цикл. PAGEREF _Toc368852439 \h 63РАЗДЕЛ 3.2. Изучение механизма регуляции клеточного цикла под действием карнозина.




PAGEREF _Toc368852440 \h 663.2.1. Карнозин избирательно ингибирует пролиферацию клеток глиобластомы. PAGEREF _Toc368852441 \h 663.2.2. Карнозин снижает уровень АФК и индуцирует экспрессию MnСОД. PAGEREF _Toc368852442 \h 703.2.3. Карнозин индуцирует G2 блок и усиливает экспрессию циклина В1. PAGEREF _Toc368852443 \h 74РАЗДЕЛ 3.3. Изучение функциональной нагрузки структурных элементов в молекуле карнозина при антипролиферативном эффекте. PAGEREF _Toc368852444 \h 78РАЗДЕЛ 3.4. Возможности применения карнозина в радиотерапии. PAGEREF _Toc368852445 \h 82ГЛАВА 4. ОБСУЖДЕНИЕ РЕЗУЛЬТАТОВ PAGEREF _Toc368852446 \h 83ВЫВОДЫ PAGEREF _Toc368852447 \h 96СПИСОК ЛИТЕРАТУРЫ PAGEREF _Toc368852448 \h 97

ВВЕДЕНИЕ

Карнозин представляет собой природный дипептид, состоящий из двух аминокислот – -аланина и L-гистидина. В организме человека карнозин представлен во многих тканях (мышцы, мозг, печень, ткани глаза, желудок, почки, легочная ткань) достигая наивысшей концентрации в скелетных мышцах (~ 20 мM) и в обонятельных луковицах головного мозга (~ 2.5 мM) PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5NYW5uaW9uPC9BdXRob3I+PFllYXI+MTk5MjwvWWVhcj48

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RW5kTm90ZT5=

ADDIN EN.CITE.DATA [1, 2]. Карнозин синтезируется с помощью фермента карнозин-синтетазы, которая катализирует образование пептидной связи между -аланином и L-гистидином используя энергию АФТ и ионы Mg2+ с использованием энергии АТФ PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5LYWx5YW5rYXI8L0F1dGhvcj48WWVhcj4xOTU5PC9ZZWFy

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ADDIN EN.CITE.DATA [8, 9].

Карнозин демонстрирует многообразие функций, приведем некоторые из них. Во время интенсивной мышечной работы карнозин выполняет функцию протонного буфера, связывающего избыток протонов, образующихся вместе с молочной кислотой, и препятствующего развитию ацидоза ADDIN EN.CITE <EndNote><Cite><Author>Северин С. Е.</Author><Year>1953</Year><RecNum>200</RecNum><DisplayText>[10]</DisplayText><record><rec-number>200</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">200</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author><style face="normal" font="default" charset="204" size="100%">Северин С.

Е., Кирзон, М.В., Кафтанова, Т.М.</style></author></authors></contributors><titles><title><style face="normal" font="default" charset="204" size="100%">Влияние карнозина и ансерина на работу изолированной мышцы лягушки.</style></title><secondary-title><style face="normal" font="default" charset="204" size="100%">Доклад. АН СССР</style></secondary-title></titles><periodical><full-title>Доклад. АН СССР</full-title></periodical><pages><style face="normal" font="default" charset="204" size="100%">691-694</style></pages><volume><style face="normal" font="default" charset="204" size="100%">91</style></volume><dates><year><style face="normal" font="default" charset="204" size="100%">1953</style></year></dates><urls></urls></record></Cite></EndNote>[10]. Карнозин также способен образовывать комплексы с металлами переменной валентности (Cu2+, Zn2+, Fe2+ и др.) PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5WaW9sYTwvQXV0aG9yPjxZZWFyPjE5Nzk8L1llYXI+PFJl

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ADDIN EN.CITE.DATA [11-13]. Антиоксидантные свойства карнозина заключаются в его способности образовывать комплексы с двумя высоко реакционноспособными окислителями – гидроксильным радикалом и супероксидным анионом, снижая их реактивность и предотвращая дальнейшее распространение окислительного повреждения PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5QYXZsb3Y8L0F1dGhvcj48WWVhcj4xOTkwPC9ZZWFyPjxS

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ADDIN EN.CITE.DATA [14-17]. Кроме того, в нескольких независимых исследованиях было продемонстрировано эффективное подавление накопления продуктов перекисного окисления липидов (малоновый диальдегид, пероксильные радикалы, ненасыщенные альдегиды, 4-гидроксиноненаль) в присутствии карнозина, что снижало уровень окислительного повреждения макромолекул, способствуя сохранению их структуры и функций PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5EdXBpbjwvQXV0aG9yPjxZZWFyPjE5ODQ8L1llYXI+PFJl

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ADDIN EN.CITE.DATA [15, 18, 19]. Антипролиферативный эффект карнозина на опухолевые клетки был впервые описан Nagai и Suda в 1986 году ADDIN EN.CITE <EndNote><Cite><Author>Nagai</Author><Year>1986</Year><RecNum>88</RecNum><DisplayText>[20]</DisplayText><record><rec-number>88</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">88</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Nagai, K.</author><author>Suda, T.</author></authors></contributors><titles><title>[Antineoplastic effects of carnosine and beta-alanine--physiological considerations of its antineoplastic effects]</title><secondary-title>Nihon Seirigaku Zasshi</secondary-title><alt-title>Nihon seirigaku zasshi. Journal of the Physiological Society of Japan</alt-title></titles><periodical><full-title>Nihon Seirigaku Zasshi</full-title><abbr-1>Nihon seirigaku zasshi. Journal of the Physiological Society of Japan</abbr-1></periodical><alt-periodical><full-title>Nihon Seirigaku Zasshi</full-title><abbr-1>Nihon seirigaku zasshi. Journal of the Physiological Society of Japan</abbr-1></alt-periodical><pages>741-7</pages><volume>48</volume><number>11</number><edition>1986/01/01</edition><keywords><keyword>Alanine/ pharmacology</keyword><keyword>Animals</keyword><keyword>Antineoplastic Agents</keyword><keyword>Carnosine/administration &amp; dosage/ pharmacology/therapeutic use</keyword><keyword>Dipeptides/ pharmacology</keyword><keyword>Male</keyword><keyword>Mice</keyword><keyword>Mitomycin</keyword><keyword>Mitomycins/administration &amp; dosage</keyword><keyword>Picibanil/administration &amp; dosage</keyword><keyword>Sarcoma 180/drug therapy</keyword><keyword>beta-Alanine/administration &amp; dosage/ pharmacology/therapeutic use</keyword></keywords><dates><year>1986</year></dates><isbn>0031-9341 (Print)&#xD;0031-9341 (Linking)</isbn><accession-num>3102721</accession-num><urls></urls><remote-database-provider>NLM</remote-database-provider><language>jpn</language></record></Cite></EndNote>[20]. В их работе подкожные инъекции карнозина мышам линии ddY, с предварительно вживленными клетками саркомы, способствовали снижению интенсивности пролиферации опухолевых клеток и увеличению выживаемости животных. Позже в независимом исследовании Renner et al. на мышах с ксенографтами HER2/neu NIH3T3 фибробластов показали, что ежедневные внутрибрюшинные инъекции карнозина существенно подавляют пролиферацию злокачественных клеток и опухолевый рост PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5SZW5uZXI8L0F1dGhvcj48WWVhcj4yMDEwPC9ZZWFyPjxS

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ADDIN EN.CITE.DATA [21]. Holliday и McFarland продемонстрировали избирательность действия карнозина на опухолевые клетки. Добавление карнозина к смеси клеток карциномы (HeLa) и нормальных человеческих фибробластов (MRC-5) приводило к выборочному уничтожению HeLa клеток и удлиняло продолжительность жизни фибробластов PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5NY0ZhcmxhbmQ8L0F1dGhvcj48WWVhcj4xOTk0PC9ZZWFy

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ADDIN EN.CITE.DATA [22, 23]. Обработка карнозином нормальных эмбриональных стволовых клеток и злокачественных клеток эмбриональной терато-карциномы также приводило к избирательной гибели последних. Полученные данные указывали на то, что ни степень дифференцировки, ни скорость пролиферации клеток не являются определяющими факторами избирательного действия карнозина на опухолевые клетки. Несмотря на наличие большого количества экспериментальных данных об антипролиферативном действии карнозина на опухолевые клетки, механизм противоопухолевого эффекта остается не выясненным.

На сегодняшний день в литературе накоплено множество данных о том, что интенсивность пролиферации зависит от соотношения про- и антиоксидантов внутри клетки PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5CdXJoYW5zPC9BdXRob3I+PFllYXI+MjAwOTwvWWVhcj48

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ADDIN EN.CITE.DATA [24, 25]. Показано, что быстро пролиферирующие клетки характеризуются более высоким содержанием прооксидантов и пониженной активностью антиоксидантной системы по сравнению с клетками, делящимися медленно или находящимися в состоянии покоя PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Hb3N3YW1pPC9BdXRob3I+PFllYXI+MjAwMDwvWWVhcj48

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