Кинетический механизм реакции синтеза АТР, катализируемой F1-F0-АТРазой субмитохондриальных частиц тема диссертации и автореферата по ВАК РФ 03.00.04, кандидат биологических наук Галкин, Михаил Александрович

• Галкин, Михаил Александрович
• кандидат биологических наук
• 1999

Оглавление диссертации кандидат биологических наук Галкин, Михаил Александрович

ОГЛАВЛЕНИЕ

Оф.

ВВЕДЕНИЕ

ПРИНЯТЫЕ СОКРАЩЕНИЯ

лголп mrmnA'nmT.t с V^UOV^r JlJTXi з rJoi

ГЛАВА 1. ПРЕПАРАТЫ АТРаз FjF0 ТИПА

ГЛАВА 2. СТРУКТУРА Н+- АТР - СИНТЕТАЗЫ

2.1. Структура фактора Fj

2.2. Структура фактора F0

ГЛАВА 3. ВЗАИМОДЕЙСТВИЕ АТРАЗЫ СО

СВОИМИ ЕСТЕСТВЕННЫМИ ЛИГАНДАМИ

3.1. Связывание нуклеотидов с факторам Fj

3.2. Функции центров прочного связывания нуклеотидов

3.3. Функционально-важные аминокислотные остатки, входящие в состав нуклеощцсвязывающих центров

3.4. Связывание фосфата с фактором Fj

3.5. Взаимодействие FjFq-АТРэзы с ионами металлов

ГЛАВА 4. РЕАКЦИИ, КАТАЛИЗИРУЕМЫЕ F^-АТРазой

4.1. Реакция синтеза АТР

4.2. Реакция гидролиза АТР

ГЛАВА 5. СОВРЕМЕННЫЕ ПРЕДСТАВЛЕНИЯ О МЕХАНИЗМЕ

ФУНКЦИОНИРОВАНИЯ МИТОХОНДРИАЛЬНОЙ Н+-АТР- СИНТЕТАЗЫ.

5.1. Гипотезы сопряжения

5.2. Структурно-функциональные основы сопряжения

ГЛАВА б. ПРОБЛЕМА ОБРАТИМОСТИ АТРАЗНОЙ РЕАКЦИИ

6.1. Эффекторы FjFq-АТРэзы

Grp.

МЕТОДЫ ИССЛЕДОВАНИЯ

ГЛАВА 1. ПРЕПАРАТИВНЫЕ МЕТОДЫ

ГЛАВА 2. АНАЛИТИЧЕСКИЕ МЕТОДЫ

РЕЗУЛЬТАТЫ ИССЛЕДОВАНИЯ

ГЛАВА I. КИНЕТИЧЕСКИЙ АНАЛИЗ РЕАКЦИИ СИНТЕЗА АТР

ГЛАВА П. "СКРЫТАЯ" Д£н+ -ЗАВИСИМАЯ АКТИВАЦИЯ (Mg2+)ADP-

БЛОКИРОВАННОЙ АТРазы

ГЛАВА Ш. ИЗУЧЕНИЕ МЕХАНИЗМА "СКРЫТОЙ" Дцн+ -ЗАВИСИМОЙ

АКТИВАЦИИ (Мй2^)АОР-БЛОКИРОВАННОЙ АТРазы

ОБСУЖДЕНИЕ РЕЗУЛЬТАТОВ

СПИСОК ЛИТЕРАТУРЫ

Введение диссертации (часть автореферата) на тему «Кинетический механизм реакции синтеза АТР, катализируемой F1-F0-АТРазой субмитохондриальных частиц»

ВВЕДЕНИЕ

Синтез ATP в митохондриях осуществляется сложным протеолигоэдньш комплексом - FjFq-АТРнзой. Для протекания реакции синтеза АТР необходим трансмембранный градиент протонов (Дщ+) . В ходе A[Ijj+- зависимого синтеза АТР осуществляется сопряжение векторного переноса протонов и скалярной реакции образования фосфоэфирной связи. Механизм этого сопряжения не установлен, несмотря на большое количество экспериментальных данных. Большинство авторов, изучая гидролиз или синтез АТР, явно или неявно отождествляют механизмы прямой и обратной реакции. Однако целый ряд экспериментальных данных, касающихся взаимодействия АТРазы с нуклеотидами и другими естественными лигаддами, указывает на то, что механизмы синтеза и гидролиза АТР могут различаться. Прежде всего это вытекает из существования односторонних ингибиторов АТРазы (тормозящих только либо АТРазную, либо АТР синтетазную активность). Одним из таких ингибиторов является ADP. Преинкубация АТРазы с ADP в низкой концентрации в присутствии ионов магния приводит к гистерезисному поведению фермента в АГРазной реакции. При этом блокируется гидролиз АТР, однако скорость синтеза АТР не меняется. Подробное изучение параметров (М$2+)АОР-зависимого торможения АГРазной активности привело к возникновению гипотезы о различии путей синтеза и гидролиза АТР, катализируемых митохоцдриальной АТРазой. Согласно этой гипотезе, фермент может находиться в двух функционально различных состояниях: "синтетазном" и "щдролазном", а переход из одной формы в другую контролируется нуклеотидами - ADP и АТР. В связи с этим становится очевидным, что нельзя исходя из данных о гидролизе АТР судить о механизме синтстазной реакции. Следовательно, необходим обстоятельный кинетический анализ реакции синтеза

АТР} как таковой. Недавно в нашей лаборатории проведен кинетический анализ решщии ГИДролиза AT?, катализируемой препаратом суомигохондриальиьис частц (AS-СМЧ). Поэтому целесообразно провести исследование кинетического механизма АТР синтстазной реакции на этом же типе препарата фермента, что и было осуществлено в данной работе.

Содержание данной работы состоит из : 1) обзора литературных данных; 2) экспериментальной часта, включающей методы и результаты наших исследований; 3) обсуждения полученных данных; 4) выводов, следующих из нашей рабош.

В обзоре литературы будет описана структура Н^-АТР-синтазы, а также приведены данные, касающиеся: а) взаимодействия АТРазы с субстратами и продуктами реакции; б) параметров реакций синтеза и гидролиза АТР; в) участия ионов металлов в катализе и регулировании активности АТРазы; г) современных представлений о механизме синтеза АТР; д) проблемы обратимости АТРазной реакции.

В экспериментальной части будут изложены данные, полученные нами в ходе кинетического анализа АТР синтетазной реакции. Нами определены параметры реакции синтеза АТР для препарата AS-СМЧ из сердца быка, сопряженных субстехиометрическими количествами олигомшщна. Нами установлено, что помимо субстратов - комплекса Mj^ADP и моноаниона неорганического фосфата - для протекания реакции синтеза необходим ион Сродство субстратов и Mg2+ к ферменту не зависит от величины Apjj-h

Также в ходе исследования установлено, что (Mg^^)ADP-блокированная АТРаза под действием ДДц+ переходит в форму фермента, остающуюся неактивной в реакции гидролиза, но способную мгновенно активироваться при снятии ДЦЦ + •

ПРИНЯТЫЕ СОКРАЩЕНИЯ

СМЧ - субмитохондриальныс частацы;

А-СМЧ, AS-СМЧ - различные типы субмигохондриальных частиц; ОЧА - ашихзмицин-чувстщггслъная АТРаза;

Fj, MFj, CFj, EFj, TFj - АТРазы (сопрягающие факторы) митохондрий, хлорогшастов, Е. соН, термофильной бактерии PS3.

Fq - мембранный компонент - АТР-синтазы;

Ajljj+ - трансмембранная разность электрохимических потенциалов протонов;

АТР - аденозинтрифосфорная кислота;

ADP - аденозиндифосфорная кислота;

NTP ( ГГР, GTP, СТР, UTP) - нуклсозидтрифосфаты;

Pj - неорганический фосфат;

NADH, NAD+ - шжешшамидадеыиндинуклеотид ( восстановленная и окисленная формы);

ФЕП - фосфознолпируват; ПК - пируваткиназа; ЛДГ - лактатдегидрогеназа; ЭДТА - этилендиаминтстраацетат;

HEPES - К-2-гидроксиэтилпиперазин-К-этансульфоновая кислота; Трис - трис- (гидроксимстил)аминомстан;

FCCP - карбоншшианид-3-фторфенилгидразон (разобщитель);

Nbf-Q - 4-хлор-7-шпробенэофуразан;

DCCD-NjN'-дихщклогсксилкарбодиимид;

FSBA - 5-р-флуоросульфонилбснзоил-адснозин;

FSBI - 5-р-флуоросульфонилбензоил-инозин.

ОБЗОР ЛИТЕРАТУРЫ

Заключение диссертации по теме «Биохимия», Галкин, Михаил Александрович

выводы

1) Измерены начальные скорости реакции синтеза АТР, катализируемой FjFq - АТРазой AS-СМЧ в широком диапазоне концентраций свободных ADP, неорганического фосфата, ионов Mg2^" и их комплексов.

2) Зависимости начальных скоростей от концентраций лигандов -субстратов хорошо соответствуют модели, в которой истинными субстратами служат комплекс Mg:ADP и моноанион неорганического фосфата, а в качестве активатора синтеза АТР служит ион Mg2* (Kjjj для иона Mg2~*~ составляет 25 мкМ ( рН 8,0; ионная сила - 0,1 мМ; 30°С). Ион магния является формально бесконкурентным активатором реакции синтеза АТР по отношению к комплексу MgiADP и конкурентным активатором по отношению к фосфату.

3) Установлено, что при синтезе АТР реакция протекает с неупорядоченным образованием тройного комплекса фермент 'Mg:ADP • фосфат.

4) Сродство субстратов синтеза и иона к ферменту не зависит от величины Apjj+.

5) Показано, что (Mg2+)ADP- блокированная АТРаза под действием Ajljj+ преобразуется таким образом, что способна мгновенно активироваться при снятии Apfj+. Свободный ADP тормозит AjTjj+ - зависимую активацию с 20 мкМ.

6) Предложена модель для реакции синтеза АТР, включающая Ард+ -зависимую активацию как начальную стадию каталитического цикла синтеза.

Автор хотел бы сердечно поблагодарить АНДРЕЯ ДМИТРИЕВИЧА ВИНОГРАДОВА за отеческое руководство, АНТОНА ВЛАДИМИРОВИЧА СЫРОЕШКИНА - за всестороннюю помощь и поддержку в работе, а также сотрудников лаборатории и кафедры биохимии - за теплое отношение и поддержку.

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