От палочки до колбочки: функциональные переходы в эволюции фоторецепторов позвоночных

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Аннотация

Сетчатка позвоночных содержит два типа фоторецепторов: палочки и колбочки, являющиеся рецепторами ночного и дневного зрения соответственно. Они имеют ряд морфологических и биохимических различий, определяющих их функциональную роль. Обнаружение у ряда позвоночных рецепторов промежуточного типа стало основой для теории трансмутаций, постулирующей, что как палочки, так и колбочки в процессе адаптации зрительной системы к различным условиям обитания способны менять свои функциональные роли и превращаться в противоположный своему тип. Можно выделить несколько уровней проявления изменений в физиологии фоторецептора при функциональном переходе: морфологический (общее строение клетки), молекулярно-биохимический (экспрессия специфичных изоформ белков светочувствительного сигнального каскада) и электрофизиологический (чувствительность и кинетика ответа на свет). Фоторецепторы с подтвержденным переходным типом обнаруживаются у наиболее примитивных позвоночных, а также у групп, претерпевших смещение условий обитания в сторону крайне низкой или высокой освещенности. В последние два десятилетия значительно продвинулось понимание молекулярных механизмов функционального перехода от типичных палочек к колбочкам, и наоборот. Вместе с тем ряд аспектов остается малоизученным, в первую очередь из-за того, что многие животные, обладающие переходными рецепторами, далеки от стандартных модельных объектов современной биологии. В рамках данного обзора рассматриваются примеры переходных фоторецепторов в различных таксонах, описывается история их изучения и современные исследования, проливающие свет на молекулярные особенности, лежащие в основе их нестандартной физиологии.

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А. Ю. Ротов

Институт эволюционной физиологии и биохимии им. И.М. Сеченова РАН; Институт физиологии им. Л.А. Орбели НАН РА

Автор, ответственный за переписку.
Email: rotovau@gmail.com
Россия, Санкт-Петербург; Армения, Ереван

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2. Рис. 1. Общая морфология типичных палочек и колбочек. Ключевые различия включают в себя форму и ультраструктуру дисков наружного сегмента, наличие/отсутствие масляной капли (характерна для колбочек лопастеперых и ганоидных рыб, амфибий, ящериц, птиц, однопроходных и сумчатых млекопитающих, см. Toomey, Corbo, 2017) и форму синаптического окончания. Свет распространяется по направлению от синаптического окончания к наружному сегменту.

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3. Рис. 2. Упрощенная схема каскада фототрансдукции в палочках позвоночных. В случае колбочек цГМФ-управляемые каналы также локализованы в мембранных дисках (представляющих собой складки плазматической мембраны). Сплошные стрелки обозначают процессы активации каскада, пунктирные – процессы выключения. Обозначение “Ca” соответствует кальций-чувствительным белкам, регулирующим активность опсинкиназы, гуанилатциклазы и цГМФ-управляемых каналов. Условные обозначения: Р – зрительный пигмент (родопсин), Рф – фосфорилированная форма; Т – трансдуцин; ФДЭ – фосфодиэстераза; ГЦ – гуанилатциклаза; ОпК – опсинкиназа; Арр – аррестин. RGS9/Gβ5 – комплекс, ускоряющий инактивацию трансдуцина.

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