5. Фоторецептор бактерий

Описанная выше система регуляции, основанная на использовании уровня цикло-ди-ГМФ, является достаточно тонкой и позволяет успешно регулировать многие процессы в бактериальных клетках. Компоненты этой системы входят в состав значительного количества мультиферментных комплексов. Одним из самых причудливых примеров таких комплексов является так называемый бактериофитохром, который выполняет роль фоторецептора. Эта структура была впервые описана у Rhodobacter phaeroides под названием BphG1 [78].

Бактериофитохром (рис. 30) функционирует в качестве рецептора к свету в области «красный\далний красный» и регулирует подвижность и синтез пигментов у R. phaeroides. BphG1 состоит из двух основных модулей: PAS-GAF-PHY фотосенсорного и GGDEF-EAL терминального. Было показано, что BphG1может функционировать в качестве фотозависимой дигуанилатциклазы, а так же в качестве фотонезависимой фосфодиэстеразы [78]. Переключение между двумя разными типами активности осуществляется путём расщепления С-терминального EAL домена. Считается, что димеризация EAL домена предупреждает демеризацию домена GGDEF, что делает последний функционально неактивным. Расщепление EAL домена приводит к тому, что GGDEF домен подвергается конформационным изменениям, которые активируются красным светом и приводят к проявлению дигуанилатциклазной активности. Однако не у всех бактерий терминальным является GGDEF-EAL домен. У некоторых видов С-терминальный домен фоторецептора представлен гистидин киназой [78].

Рис. 30 Структура и принципы функционирования бактериофитохрома с GGDEF-EAL С-терминальным доменом [78].

Интересным является то, что в качестве кофермента в фоторецепторе бактерий используется билливирдин IX. Данное соединение является продуктом метаболизма гемина, который играет огромную роль в жизнедеятельности микроорганизмов. Билливирдин IX образуется путём расшепления молекулы гемина по мезоположениям с помощью гемоксигеназ. Большенство этих ферментов способно разрывать молекулу гемина по α-мезо-положению с образованием билливирдин IX α. Однако, у P. aeruginosa описана гемоксигеназа PigA, которая, разрывая макроцикл гемина в других положениях приводит к образованию билливирдин IX β и билливирдин IX δ [81]. Причины таких различий, а так же, как различные биливердины влияют на функционирование фоторецептора, окончательно не выяснены. Так же окончательно не ясны функции самого бактериофитохрома, не смотря на то, что, эти соединения как с GGDEF-EAL так и с киназным С-терминальным доменом обнаружено у значительного количества бактерий, а так же у некоторых грибов.

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