Активность ацетилхолинэстеразы

Первой нашей задачей было сопоставить количество ацетилхолина в синаптических везикулах с его общим содержанием в нервных окончаниях. Если обработать нервные окончания кислотой, что инактивирует ферменты, то разрушаются как наружная мембрана нервных окончаний, так и внутренние мембраны синаптических везикул. При этом ацетилхолинэстераза на поверхности окончаний теряет активность и ацетилхолин полностью сохраняется. Таким образом, кислотная обработка позволяет определить общее количество ацетилхолина в нервном окончании.

Если же препарат нервных окончаний подвергнуть быстрому замораживанию и последующему оттаиванию или определенному механическому воздействию, то разрушается только наружная мембрана синаптосом. Активность ацетилхолинэстеразы при этом не подавляется, и фермент гидролизует весь ацетилхолин, находящийся в цитоплазме вне везикул. Ацетилхолин, содержащийся в везикулах, не распадается, потому что от действия фермента он защищен мембранами везикул. Теперь количество содержащегося в везикулах ацетилхолина может быть определено после кислотной обработки, разрушающей мембраны везикул и инактивирующей ацетилхолинэстеразу.

В 1968 г. подтвердились данные о том, что в синаптических везикулах запасается большое количество ацетилхолина. Мы обнаружили, что в везикулах электрического органа, как и в везикулах из мозга, сосредоточена примерно половина всего ацетилхолина нервных окончаний. Казалось бы, это свидетельствует в пользу везикулярной гипотезы. Напомним, однако, что баланс ацетилхолина определяется, по всей вероятности, в цитоплазме окончаний, поскольку синтезирующий его фермент, холинаце- тилтрансфераза, находится в цитоплазме.

На данной стадии исследований стало ясно, какие эксперименты следует поставить, чтобы подтвердить везикулярную гипотезу. Если она верна, то стимуляция нервных окончаний должна приводить к истощению везикулярного ацетилхолина, прежде чем произойдет его высвобождение из других отделов клетки. Для проверки этого предположения очень подходил такой объект, как концентрированная масса нервных окончаний, которую предоставлял электрический орган Torpedo.

 

 

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