13
Мембрана клетки
План
Емкость мембраны
Постоянная времени
Емкость в кабеле
Сенситизация и S интернейроны
Вывод
Емкость мембраны
Мембрана клетки не только проводит ионные токи, но и накапливает заряд
на своей внешней или внутренней поверхности. С точки зрения теории
электричества, разделение зарядов на мембране означает, что мембрана
обладает свойствами конденсатора. В общем виде конденсатор состоит из
двух проводящих пластин, отделенных друг от друга изолирующим
материалом; в промышленных конденсаторах проводящие пластины обычно
сделаны из металлической фольги, а изолирующая прослойка между ними — из
пластика. В случае нервной клетки проводниками являются два слоя
жидкости, находящиеся по обе стороны мембраны, а сама мембрана играет
роль изолирующей прослойки. При зарядке конденсатора от батареи на одной
из пластинок накапливается положительный заряд, в то время как на второй
пластинке создается равный по величине запас отрицательного заряда.
Емкость конденсатора (С) определяется количеством заряда (Q), запасаемым
на один вольт потенциала (V), приложенного к пластинам конденсатора: С =
Q/V. С измеряется в кулонах, деленных на вольт, т. е. в фарадах (Ф). Чем
ближе друг к другу расположены пластины, тем более эффективно
конденсатор способен разделять и накапливать заряд. Поскольку толщина
мембраны клетки всего 5 нм, она способна накапливать достаточно большой
заряд. Обычно емкость мембраны нервных клеток составляет 1 мкФ/см2.
Преобразовав выражение, получаем Q = CV. При потенциале покоя —80 мВ,
количество избыточного отрицательного заряда на внутренней стороне
мембраны составит (1 · 10–6) х (80 ·10–3) = 8· 10–8 кулонов, деленных на
см2, что соответствует 5 · 1011 одновалентных ионов (0,8 пмоль) на
квадратный сантиметр мембраны.
Величину тока, протекающего внутрь конденсатора или из него, можно
подсчитать на основе соотношения заряда и напряжения, учитывая, что ток
(i, в амперах) есть скорость изменения заряда во времени, т.е. 1 ампер =
(1 кулон)/(1 с). Поскольку Q = C/V, получим:
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Скорость изменения заряда на конденсаторе прямо пропорциональна величине
тока. Если ток постоянен, то потенциал будет меняться с постоянной
скоростью dV/dt = i/C.
Соотношение тока и напряжения в цепи, содержащей резисторы
(сопротивления) и конденсаторы (емкости), соединенные параллельно.
Прямоугольный скачок тока величиной i, приложенный к резистору (R),
создает скачок напряжения на резисторе величиной V = iR. Если тот же
скачок тока приложить к конденсатору (С), то напряжение на конденсаторе
будет накапливаться со скоростью dV/dt = i/C. Когда эти два элемента,
резистор и конденсатор, соединены параллельно, то весь ток пойдет
сначала на зарядку конденсатора со скоростью i/С; однако, как только на
конденсаторе накопится какой-то заряд, ток потечет и через резистор. По
мере нарастания тока, все большая его часть будет проходить через
сопротивление, потому что скорость зарядки конденсатора будет постепенно
снижаться. В конце концов весь ток будет течь через резистор, создавая
на нем потенциал V = iR, a конденсатор будет полностью заряжен. По
завершении скачка тока заряд из конденсатора постепенно рассеется на
резисторе, а напряжение вернется к нулю. Из экспериментов на аксоне
кальмара можно заключить, что гипотеза, высказанная Бернштейном в 1902
г., была близка к истине: трансмембранный градиент калия является
важным, хотя и не единственным фактором, влияющим на мембранный
потенциал. Чем можно объяснить отклонение экспериментальной кривой от
уравнения Нернста? Оказывается, для этого достаточно снять ограничение с
модели, состоящее в том, что мембрана непроницаема для ионов натрия.
Мембрана реальной клетки действительно обладает натриевой
проницаемостью, которая составляет от 1 до 10 % калиевой.
Для рассмотрения роли натриевой проницаемости обратимся к модели
идеальной клетки и временно исключим из поля зрения перемещение ионов
хлора. Мембранный потенциал равен калиевому равновесному потенциалу,
поэтому перемещение суммарного заряда через мембрану отсутствует, клетка
находится в покое. Если теперь ввести в модель натриевую проницаемость,
то натрий будет стремиться войти в клетку благодаря как своему
концентрационному градиенту, так и мембранному потенциалу. По мере входа
натрия на внутренней поверхности мембраны накапливается положительный
заряд и мембрана деполяризуется. В результате ионы калия выходят из
равновесия и начинают покидать клетку. С увеличением деполяризации
мембраны движущая сила для входа натрия снижается, в то время как
движущая сила для выхода калия возрастает. Процесс продолжается до тех
пор, пока оба ионных потока не уравновесят друг друга. В этот момент
изменение мембранного потенциала прекращается, поскольку какое-либо
накопление заряда отсутствует. Вообще говоря, значение мембранного
потенциала расположено между калиевым и натриевым равновесными
потенциалами и определяется равновесием между калиевым и натриевым
токами, равными по величине и направленными в противоположные стороны.
Ионы хлора также участвуют в этом процессе, однако, как мы убедились
ранее, равновесный потенциал для хлора подстраивается под новое значение
мембранного потенциала за счет изменения внутриклеточной концентрации
этого иона. По мере того как токи катионов постепенно приходят в
равновесие, внутриклеточный уровень хлора возрастает до тех пор, пока
суммарный хлорный ток не станет равным нулю.
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22172a3d3e1f032a343f40191c1606130b04050706050607070605030202030306070707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Рис. 1. Распространение потенциала вдоль аксона омара, регистрируемое с
помощью поверхностного электрода.
Постоянная времени
Нарастание и спад потенциала происходит по экспоненциальной кривой, как
показано на рис. 1В. Фаза роста описывается уравнением: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где t — время от начала импульса. Постоянная времени равна произведению
RC. Это время, за которое потенциал возрастаетдо 63 % (1 – 1/e) своего
максимального значения. Спад напряжения тоже экспоненциален, с той же
постоянной времени. Ток через резистор, iR, должен изменяться во времени
по тому же закону, что и напряжение. Следовательно, на фазе роста ток
начинает расти от нуля до своего максимального значения i. Емкостной
ток, наоборот, начинается с величины i и спадает до нуля по экспоненте с
той же постоянной времени. По завершении импульса, поскольку внешнего
источника тока нет, единственным током на резисторе будет ток,
генерируемый напряжением на конденсаторе. Следовательно, ток на
резисторе равен по величине емкостному току и противоположен ему по
направлению, как показано на рисунке.
Вышеописанную цепь, состоящую из параллельно соединенных резистора и
конденсатора, можно использовать для описания сферической нервной
клетки, аксон и дендриты которой настолько малы, что их вкладом в
электрические свойства клеток можно пренебречь. В эквивалентной цепочке
для аксона или мышечного волокна как емкость мембраны, так и ее
сопротивление распределены по всей длине волокна, как показано на рис.
7.2D. Емкость мембраны на единицу длины ст (измеряемая в мкФ/см) зависит
от удельной емкости на единицу площади Сm (в мкФ/см2) по формуле сm =
2Ст, где — радиус волокна.
Постоянная времени мембраны сферической клетки или волокна не зависит от
размера клетки или волокна. Причина этого в том, что увеличение радиуса
(а следовательно, и площади поверхности мембраны) влечет за собой не
только увеличение емкости, но и соответствующее снижение сопротивления,
так что произведение двух величин не меняется. Поскольку показано, что
величина Ст приблизительно одинакова для всех нервных и мышечных волокон
(1 мкФ/см2), то величина ф является удобным параметром, характеризующим
удельное сопротивление мембраны для данной клетки. Постоянная времени —
это третий параметр, который, наряду с входным сопротивлением и
постоянной длины, характеризует поведение аксона. Диапазон значений
постоянной времени в различных типах нервных и мышечных клеток
составляет от 1 до 20 мс.
Емкость в кабеле
Каково влияние постоянной времени на ток в кабеле? Как и в случае
простой RC-цепочки (рис. 1С), нарастание и спад потенциала в ответ на
скачкообразное изменение тока замедляется благодаря наличию
конденсатора. Ситуация осложняется тем, что ток течет уже не через один
конденсатор, но каждый сегмент цепочки является одним из
резистивно-емкостных элементов, взаимодействующих между собой. Благодаря
этим взаимодействиям временной ход в отдельном сегменте нельзя описать
простой экспонентой, и фазы роста и спада замедляются по мере удаления
от точки инъекции тока (рис. 2). Поскольку скорость нарастания
потенциала зависит от расстояния между отводящим электродом и местом
инъекции тока, постоянную времени спада уже нельзя рассчитать на основе
простого измерения времени 63 %-ного роста потенциала, за исключением
единственной точки вдоль волокна, в которой это расстояние равно
постоянной длины.
Обратимся к рассмотрению движения ионов. Если в аксон инъецирован
положительный ток, внутриклеточные ионы (главным образом, калия) будут
распространяться вдоль волокна. Некоторая часть ионов уйдет на изменение
заряда на емкости мембраны, другая часть будет протекать по мембранному
сопротивлению. Одновременно с этим процессом будет происходить
перемещение отрицательных ионов в противоположном направлении.
Постепенно потенциал на мембране достигнет нового устойчивого значения,
емкости, распределенные по мембране, будут полностью заряжены до нового
уровня потенциала, а через мембрану будет протекать постоянный ионный
ток. Время, необходимое для достижения нового устойчивого состояния,
определяется постоянной времени.
Рис. 2. Мембранный ток во время прохождения потенциала действия.
.picscalex10001000900000318cc00000000f3cb0000000004000000030108000500000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e48795a97a47facad9191919191919191a68e6574ac9e535179514a4851a0d48ea4a684b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Сенситизация и S интернейроны
Механизм того, как отдельный нейрон может играть координирующую роль в
поведении животного, тщательным образом на насекомых и ракообразных, у
которых были описаны отдельные «командные» нейроны, способные
дирижировать оркестром поведенческих реакций всего организма. У пиявки
эту роль играет непарный интернейрон, присутствующий в каждом ганглии и
называемый S клеткой. Этот нейрон получает возбуждающие импульсы от
чувствительных клеток при касании и давлении и в свою очередь возбуждает
L мотонейрон, который способствует сокращению тела (как ранее было
указано). Каждая S клетка связана с S клеткой соседних сегментов
посредством толстого аксона с большой скоростью проведения через
электрический синапс. То, что синапс находится посередине между
сегментами, можно определить путем введения в S клетку пероксидазы
хрена, которая не может распространяться через щелевые контакты.
Последовательная цепь S клеток является ключевым механизмом адаптивных
реакций пиявки. При повторной тактильной стимуляции прикосновением к
определенному сегменту тела сначала возникает рефлекс сокращения. Ответ
становится все слабее и слабее при каждом следующем прикосновении —
данный эффект известен как «привыкание» (habituation). Однако после
более сильного раздражения, когда активируются С (давление) и N (боль)
клетки, ответ на дальнейшую стимуляцию только Т клеток (при касании)
вновь становится заметным. Этот процесс известен как «отвыкание»
(dishabituation) Сходным же образом, если более сильный раздражитель
наносится до процедуры «обучения», это вызывает сенситизацию, т. е.
ответ на стимуляцию при касании будет больше обычного. Активность S
клеток во время «привыкания» не меняется, однако возрастает как при
сенситизации, так и при «отвыкании» В технически сложно выполнимых
экспериментах, когда избирательно перерезается аксон S клетки, либо
клетки убиваются инъекцией проназ, процессы «привыкания» и сокращения не
изменяются, однако полностью исчезает «отвыкание» и сенситизация.
Во второй серии экспериментов аксоны S клеток были повреждены, чтобы
исследовать их регенерацию. Удивительным свойством S клеток является то,
что после повреждения их аксоны следуют по старому пути и заново
образуют электрические соединения с соседним S нейроном с поразительной
точностью. Как и ожидалось, перерыв передачи импульсов по S клеткам
между отдельными сегментами приводил к исчезновению сенситизации, однако
через несколько недель после полной регенерации аксонов и восстановления
прежних связей данное свойство также восстанавливалось.
Эти примеры наглядно демонстрируют, каким образом отдельная нервная
клетка способна принимать участие в таком сложном поведенческом
процессе, как сенситизация. Было установлено, что на самом деле процессы
сенситизации во многом обусловлены серотонином. (Необычная серия
совпадений способствовала тому, что название «S клетка» стало очень
подходящим для данных клеток. Изначально она была так названа, потому
что ее импульс образует долгий импульс (spike) в месте контакта. Затем,
обнаружив, что эти клетки очень тесно связаны друг с другом, полагали,
что они могут быть синцитием. Сейчас же они называются S клетками от
слов «сенситизация» и «серотонин»!) У Aplysia процесс сенситизации (в
котором серотонин тоже играет важную роль) был очень тщательно изучен
как на клеточном, так и на молекулярном уровне).
Вывод
Короткие сигналы распространяются на более короткие расстояния, чем
длительные сигналы. В случае достаточной длительности сигнала, в течение
которого потенциал успевает достигнуть своего максимального значения,
емкость заряжается полностью, и пространственное распределение
потенциала определяется сопротивлениями мембраны и цитоплазмы: Vx =
V0е–x/. Для коротких импульсов, таких как синаптический потенциал, ток
прекращается еще до того, как емкость успевает полностью зарядиться. Это
выражаетсяв уменьшении расстояния, на которое потенциал распространяется
вдоль волокна. Другими словами, эффективная постоянная длины для
коротких сигналов меньше, чем для длительных. Кроме того, форма коротких
сигналов искажается по мере их перемещения по волокну, а их амплитуда
снижается за счет «закругления» пика, достигаемого все позднее и позднее
Литература
1. Пенроуз Р. НОВЫЙ УМ КОРОЛЯ. О компьютерах, мышлении и законах физики.
2. Грегори Р. Л. Разумный глаз.
3. Леках В. А. Ключ к пониманию физиологии.
4. Гамов Г., Ичас М. Мистер Томпкинс внутри самого себя: Приключения в
новой биологии.
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