Множественность управляемых трансмиттерами процессов в период делений дробления

Выводы.

1. Раннее развитие в период делений дробления находится под постоянным контролем трансмиттеров, последовательно или одновременно управляющих рядом ключевых процессов: запуском деления, определением места формирования борозды, завершением ее формирования, адгезией бластомеров и межбластомерными взаимодействиями.

2. Методом микроинъекции на зародышах X. laevis продемонстрирована прямо, а фармакологическими экспериментами, связыванием меченных лигандов и методом электропробоя мембраны на зародышах морских ежей подтверждена внутриклеточная локализация трансмиттерных рецепторов, участвующих в запуске делений дробления.

3. В клетках ранних зародышей одновременно присутствуют и являются функционально активными рецепторы по крайней мере к двум трансмиттерам. Внутриклеточные адренои холинорецепторы ранних зародышей Х.1аеу1з проявляют выраженную специфичность.

4. Трансмиттеры через активацию соответствующих внутриклеточных рецепторов способны влиять на внутриклеточный уровень цАМФ. и Са2+ у ранних зародышей.

5. Функция внутриклеточных трансмиттерных рецепторов реализуется при участии вторичных мессенджеров: циклических нуклеотидов, ионов кальция и фосфоинозитидов, однако именно трансмиттер определяет положение и момент формирования борозды, т. е. является более «адресным» мессенджером.

6. Циклический АМФ непосредственно или через изменение концентрации Са2+1 способен влиять на состояние цитоскелета и сократительных белков ранних зародышей.

7. Внутриклеточные 5-НТ-рецепторы ранних зародышей морских ежей участвуют в процессе адгезии бластомеров морских ежей после деления, являющемся предпосылкой для последующей реализации межбластомерной сигнализации.

8. Методами ри1: сЬ-с1итр, флуоресцентных Са2±зондов, связывания меченных радиолигандов и фармакологическим методом показано присутствие в плазматических мембранах ранних зародышей морских ежей 5-НТи холинорецепторов. Активность этих рецепторов связана с регуляцией уровня Са2+1, причем в отношении эффектов цитотоксических тандемов типа «ФМА+никотин» 5-НТ и АХ проявляют функциональный антагонизм.

9. Мембранные 5-НТ-рецепторы участвуют в процессе завершения формирования борозды дробления у ранних зародышей морских ежей. Процесс завершения борозды дробления сопряжен также с изменениями уровня Са2+1.

10. Феномен сборки-разборки борозды дробления предоставляет возможность псевдостационарной модели развивающегося зародыша, перспективной для исследования эффектов веществ, влияющих на уровень Са2+1.

11. На «микромерной модели» и на интактных зародышах морских ежей показано существование значимых межклеточных взаимодействий в период делений дробления.

12. Мембранные 5-НТи, вероятно, АХрецепторы участвуют в процессе межбластомерных взаимодействий у ранних зародышей морских ежей.

13. Влияние трансмиттеров на межбластомерные взаимодействия реализуется через систему вторичных мессенджеров, в том числе циклическими нуклеотидами, ионами кальция и, вероятно, фосфоинозитидами.

14. Сформулирована гипотеза «протосинапса» — двухсторонней структуры, в которой обе клетки являются как источниками, так и акцепторами сигнальных молекул, идентичных трансмиттерам, на основе которой формируется асимметрия раннего зародыша и который может быть генетическим предшественником дефинитивного синапса, с одной стороны, и основой формирования многоклеточности.

15. Рассмотрены существующие и предложена модифицированная гипотеза эволюции трансмиттерных веществ и соответствующих рецепторов. гипотетическая схема происхождения трансмиттеров.

1 — поток аминокислотпредшественников трансмиттеров- 2 -порог срабатывания трансмиттерсинтезирующего фермента- 3 трансмиттерсинтезирующий фермент- 4 — белковый синтез- 5 -новосинтезированный трансмиттер- 6 — трансмиттерная рецептивная структура- 7 — цветочек.

Автор благодарит своего учителя заведующего Лабораторией эмбриофизиологии ИБР РАН профессора Геннадия Алексеевича Бузникова за науку, доброжелательное сотрудничество в совместных опытых, представленную свободу научного исследования, терпение и взыскательность в дискуссиях. Большая благодарность также сотрудникам Лаборатории за дружеское отношение, советы и поддержку. Большую ценность для автора представляет сотрудничество с коллегами по экспериментам — соавторами опубликованных материалов — в первую очередь акад. Чайлахяну и к.б.н. Григорьеву. Доктору химических наук В. В. Безуглову и коллективу возглавляемой им лаборатории ИБХ РАН автор благодарен за предоставление специально разработанных искусственно функционализированных жирных кислот и сотрудничество в проведении экспериментов. Данная работа не могла бы быть выполнена, если бы не многочисленные коллеги, принимавшие автора на морских станциях России и зарубежьяособая благодарность сотрудникам ДВО РАН докторам наук Сове, Васьковскому, Хотимченко и Жадану, кандидатам биологических наук Латышеву, Деридовичу, Креймеру и Ващенко. Автор благодарен д-ру Майклу Уитекеру (Университет Ньюкасла, Великобритания) и Королевскому Обществу за представленную возможность исследований уровня внутриклеточного кальция в клетках зародышей морских ежей, д-р д-р Элизабетте Тости, Мартину Уайлдингу и Брайену Дэйлу (Зоологическая станция «Антон Дорн» (Неаполь, Италия) за большую методическую помощь в организации опытов по ри1с1>с1итрпрофессору Давиде Файсу и Нино Олива из Университета Палермо (Италия) и академику САНУ Ракичу и сотрудникам Института биологии моря (Котор, СФРЮ) за. помощь в организации исследований. Автор признателен заведующему аквариальной ИБР Льву Гудкову за большую помощь в организации работы на зародышах шпорцевой лягушки. Искренняя признательность академику Т. М. Турпаеву за большую поддержку в организации исследований в кабинете электрофизиологии ИБР РАН и ценное обсущение. Нельзя не поблагодарить чл.-корр. РАН Л. А. Пирузяна, благодаря которому научная карьера автора не прервалась в самом начале. Автор всегда встречал самое доброжелательное отношение руководства Института биологии развития РАН в том числе академика Хрущова и доктора биологических наук Нечаевой, за что искренне благодарен. Последние по очереди, но не последние по глубине благодарности автор приносит своей семье, которая всегда относилась к его работе с пониманием и уважением. А. Б. Шмуклер оказал очень большую помощь в распечатке этой работы.

Заключение

.

От принципиального утверждения о роли трансмиттеров в эмбриогенезе и пионерских опытов, ее доказывающих, исследования, в том числе представленные в данной работе, прошли этап расширения и углубления знаний по этой проблеме и достигли этапа формирования целостных концепций. Универсальность распространения трансмиттерных систем в животном царстве, функциональная активность в ходе всего онтогенеза и консервативность каскадов передачи сигнала позволяют отнести их к фундаментальным регуляторным механизмам. Те черты, которые, согласно первоначальным представлениям, считались специфическими для эмбриональных трансмиттерных систем, в ходе исследований обнаружились в аналогичных механизмах клеток взрослого организма, а стандартные трансмиттерные механизмы дефинитивных клеток обнаруживаются на эмбриональных стадиях. Это подчеркивает генетическое единство трансмиттерной системы в ходе всего онтогенеза.

Функции множества эмбриональных трансмиттеров реализуются на уровне одной клетки с участием сложной системы вторичных мессенджеров. Предполагается, что это отражает эволюционное происхождение трансмиттерных систем как датчиков метаболических процессов. В дефинитивных клетках происходит специализация как трансмиттеров и вторичных мессенджеров, так и их функций. При всем разнообразии функций так или иначе они сводятся к пространственно-временной организации адресного сигнала. Как и в клетках взрослых организмов, трансмиттеры в донервных этапах развития являются «вершиной пирамиды» передачи клеточных сигналов, реализующихся через каскад вторичных мессенджеров.

Изученные в данной работе трансмиттерные механизмы, включающие как внутриклеточные, так и мембранные рецепторы, контролируют последовательно или одновременно практически все существенные события делений дробления, в том числе собственно цитокинез и межбластомерные взаимодействия. Наряду с открытой Г. А. Бузниковым способностью трансмиттеров внутриклеточно регулировать запуск делений дробления, уже на стадии двухклеточного зародыша, вероятно, возникает протосинаптический механизм трансмиттерной межклеточной передачи информации, обладающий значительным сходством с классической синаптической передачей, эволюционным предшественником которой он и является. К этим двум функциям в той или иной степени можно свести и все последующие проявления участия трансмиттеров в индивидуальном развитии, включая дефинитивные функции — синаптическую и управления пролиферацией. Можно предположить, что эти функции сами по себе и лежат в основе возникновения трансмиттерного управления клеточными процессами.

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