ЗАКЛЮЧЕНИЕ. Гладкомышечные клетки, которые находятся в разных органах и тканях могут значительно различаться между собой и являются наиболее разнообразным типом мышц

Гладкомышечные клетки, которые находятся в разных органах и тканях могут значительно различаться между собой и являются наиболее разнообразным типом мышц. Кроме деления на мультиунитарные и унитарные, гладкие мышцы различаются по характеру нервных и гуморальных влияний, могут иметь разную электрическую активность и характеристики сокращения.

Даже среди гладких мышц одного типа ткани могут наблюдаться значительные функциональные различия. Например, сократительные ответы васкулярных гладкомышечных клеток стенок двух артериол, которые кровоснабжают разные органы, могут отличаться друг от друга. Различия могут наблюдаться даже между гладкомышечными клетками одной артерии в двух достаточно удаленных точках [26, 122, 336, 372].

Фенотип гладкой мышцы может вариировать в соответствии с изменяющимися условиями. Например, миометрий матки образуют гладкомышечные клетки, которые претерпевают значительную трансформацию во время беременности с целью подготовки матки к родам. Кроме гипертрофии в них увеличивается степень межклеточного взаимодействия благодаря увеличению количества нексусов. Клетки также претерпевают изменения в отношении экспрессии изоформ сократительных белков. При изменении экспрессии ионных каналов и рецепторов усиливается ритмическая электрическая активность, которая координируется распространяющимся потенциалом действия и повышением концентрации ионов Са²⁺. Несмотря на то, что эти ритмические, скоординированные сокращения возникают спонтанно, они находятся под строгим контролем со стороны гормона окситоцина, уровень которого повышается до, во время и непосредственно после родов.

Значительные различия в функционировании гладких мышц, которые могут наблюдаться даже на протяжении жизни одной клетки, возможно, отражают различия в структуре белков. Показано, что в отличие от скелетной мышцы, гладкомышечные клетки демонстрируют большую вариабельность изоформ сократительных и регуляторных белков. Эта вариабельность является результатом как мультипликации генов, так и альтернативного сплайсинга. Данное разнообразие играет огромную роль в механизмах функционирования гладкой мышцы. Однако тонкие взаимоотношения между структурой и функциями этих изоформ протеинов до конца еще не выяснены.

Механизмы возбуждения и сокращения гладкомышечных клеток по многим параметрам значительно отличаются от механизмов возбуждения и сокращения скелетной мышцы:

- Гладкая мышца кроме потенциала действия может генерировать медленную градуальную деполяризацию и спонтанную активность. Потенциал действия сопровождается фазным сократительным ответом, медленная деполяризация – тоническим сокращением.

- От уровня деполяризации зависит количество открытых кальциевых каналов и, соответственно, уровень внутриклеточного кальция. В свою очередь, количество цитозольного кальция определяет силу сокращения гладкомышечной клетки. Таким образом, чем выше деполяризация мембраны, тем сильнее будет сокращаться гладкомышечная клетка.

- Наряду с электромеханическим в гладкой мышце существует фармакомеханический тип сопряжения процессов возбуждения и сокращения.

- Сократительный ответ гладкой мышцы может возникнуть и без изменения потенциала на мембране.

- На мембрану гладкой мышцы одновременно могут влиять множество факторов, иногда противоположных – например, симпатические и парасимпатические влияния. Таким образом, конечный результат зависит от соотношения интенсивности возбуждающих и тормозных влияний.

- Для инициации сокращения в гладкой мышце существует два основных источника Са²⁺: 1) из саркоплазматического ретикулума, и 2) из внеклеточной среды (через потенциал-зависимые и рецептор-управляемые ионные каналы).

- В процессе освобождения ионов Са²⁺ из саркоплазматического ретикулума активное участие принимают вторичные посредники (например, инозитол-1, 4, 5-трифосфат).

- Концентрация ионов Са²⁺ в саркоплазме гладкомышечной клетки может уменьшаться или увеличиваться в зависимости от градуальных деполяризационных или гиперполяризационных изменений мембранного потенциала, от которых зависит количество открытых кальциевых каналов.

- Наличие в плазматической мембране гладкой мышцы депо-управляемых Са²⁺ каналов способствует тонкой регуляции внутриклеточной концентрации ионов Са²⁺.

- В отличие от скелетной мышцы Са²⁺ в саркоплазме гладкой мышцы связывается с кальмодулином. Комплекс Са²⁺-кальмодулин активирует киназу легких цепей миозина, которая фосфорилирует миозин и инициирует запуск цикла образования поперечных мостиков.

- В скелетной мышце при возникновении одиночного потенциала действия освобождается количество ионов Са²⁺, достаточное для вовлечения в процесс всех поперечных мостиков. В гладкой мышце в результате физиологической стимуляции активируется только часть поперечных мостиков, т.е. существует резервное количество мостиков. Это дает возможность постепенного увеличения силы сокращения гладкой мышцы в зависимости от увеличения внутриклеточной концентрации ионов Са²⁺.

- Сокращение гладкой мышцы, которое определяется как состояние latch state является энергетически экономичным. Это энергосберегающее состояние является чрезвычайно важным для гладкой мышцы, которой приходится поддерживать сокращение в течение длительного времени.

- Скорость гидролиза АТФ миозином гладких мышц относительно скелетных мышц ниже, поэтому гладкая мышца укорачивается более медленно, чем скелетная.

- Процесс расслабления гладкой мышцы инициирует фермент фосфатаза, которая дефосфорилирует легкую цепь миозина. Одновременно с этим происходит ресеквестрация ионов Са²⁺ в саркоплазматический ретикулум.

Эффективной работе и регуляции сокращений гладкой мышцы способствуют разнообразные факторов. К этим факторам относятся:

- нейротрансмиттеры вегетативных нейронов,

- гормональные влияния,

- спонтанная электрическая активность,

- местные факторы (паракринные вещества, концентрации ионов, осмотическое давление, кислотность),

- растяжение гладкой мышцы.

В гладкомышечных клетках представлено большое количество рецепторов к нейротрансмиттерам и гормонам. Гладкомышечные клетки широко различаются в отношении типов рецепторов, располагающихся на поверхности мембраны. Причем, стимуляция рецепторов может приводить как к сокращению, так и к расслаблению гладкой мышцы. Большое количество биологически активных веществ могут действовать через разнообразные подтипы рецепторов, которые могут функционировать благодаря различным механизмам. Например, в то время как некоторые рецепторы могут представлять из себя лиганд-активируемые ионные каналы, другие действуют через G-белок. Гетеротример G-белка может действовать как непосредственно, так и посредством активации системы внутриклеточных посредников, таких как ц-АМФ, ц-ГМФ, ионы Са²⁺, инозитол-1, 4, 5-трифосфат и диацилглицерол [356]. Основные отличительные особенности строения и функционирования гладких мышц в сравнении со скелетными и сердечными мышцами даны в таблице 1.

Перечень нейротрансмиттеров, гормонов и местных факторов, регулирующих функции гладких мышц, огромен. Среди основных необходимо отметить адреналин, норадреналин, серотонин, ангиотензин, вазопрессин, нейропептид Y, оксид азота, эндотелии и кислород. Интересно отметить, что одни и те же стимулы в различных гладких мышцах могут приводить к противоположным результатам. Например, при уменьшении напряжения кислорода гладкомышечные клетки артерий большого круга кровообращения расслабляются, в то время как гладкомышечные клетки легочной артерии сокращаются.

И в заключение хотелось бы отметить, что разнообразие, пластичность и уникальные свойства гладкой мышцы дают возможность формировать мышечную стенку внутренних органов, выполняющих самые разные функции, что позволяет своеобразно реагировать на сигналы и тонко регулировать функционирование организма. Расширение наших представлений о механизмах, лежащих в основе функционирования гладких мышц, позволит обнаружить новые пути для разработки лекарственных препаратов и создать более селективные и эффективные вазоактивные, антиспастические и др. средства.

Таблица 1. Основные сравнительные характеристики скелетной, сердечной и гладкой мышцы.

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