ЛЕКЦИЯ
ДНК-СОДЕРЖАЩИЕ ВИРУСЫ И ФАГИ
РНК-СОДЕРЖАЩИЕ ВИРУСЫ
От лат. «virus» – яд
Вирусы — это внеклеточная форма жизни, обладающая собственным геномом и
способная к воспроизведению только в клетках живых организмов.
Вирион (или вирусная частица) состоит из одной или нескольких молекул
ДНК или РНК, заключенных в белковую оболочку (капсид), иногда содержащую
также липидные и углеводные компоненты
Диаметр вирусных частиц (их называют также вирионами) равен 20-300 нм.
Т.е. они намного меньше, чем мельчайшие из прокариотических клеток. Так
как размеры белков и некоторых ам.к. находятся в диапазоне 2-50 нм, то
вирусную частицу можно было бы считать просто комплексом макромолекул.
Вследствие их малых размеров и неспособности к самовоспроизведению
вирусы часто относят к разряду «неживого».
Говорят «Вирус – это промежуточная форма жизни, или нежизни», т.к. вне
клетки хозяина он превращается в кристалл.
Говорят: в. это переход от химии к живому
Вирусы являются внутриклеточными паразитами на генетическом уровне и
используют для своего размножения белок-синтезирующий аппарат
клетки-хозяина.
Жизненный цикл вируса начинается
1. с проникновения внутрь клетки.
2. Для этого он связывается со специфическими рецепторами на ее
поверхности и
а) либо вводит свою нуклеиновую кислоту внутрь клетки, оставляя белки
вириона на ее поверхности,
б) либо проникает целиком в результате эндоцитоза. В последнем случае
после проникновения вируса внутрь клетки следует его «раздевание» —
освобождение геномных нуклеиновых кислот от белков оболочки.
3. В результате этой процедуры вирусный геном становится доступным для
ферментных систем клетки, обеспечивающих экспрессию генов вируса.
4. Именно после проникновения вирусной геномной нуклеиновой кислоты в
клетку заключенная в ней генетическая информация расшифровывается
генетическими системами хозяина и используется для синтеза компонентов
вирусных частиц.
По сравнению с геномами других организмов вирусный геном относительно
мал и кодирует лишь ограниченное число белков, в основном белки капсида
и один или несколько белков, участвующих в репликации и экспрессии
вирусного генома. Необходимые метаболиты и энергия поставляются
хозяйской клеткой.
ДНК-СОДЕРЖАЩИЕ ВИРУСЫ
ДНК-содержащие вирусы несут в качестве генетического материала либо
одно-, либо двухцепочечную ДНК, которая может быть как линейной, так и
кольцевой. В ДНК закодирована информация о всех белках вируса. Вирусы
классифицируют в зависимости от того, одно или двухцепочечная у них ДНК,
и про- или эукариотической является клетка-хозяин. Вирусы заражающие
бактерии называются бактериофагами.
ДНК-содержащие вирусы
1 — вирусы оспы; 2 — вирусы герпеса; 3 – аденовирусы; 4 — паповавирусы;
5 — гепаднавирусы; 6 — парвовирусы;
Первая группа — вирусы с двуцепочечной ДНК,
– репликация осуществляется по схеме: ДНК —»РНК -> ДНК.
– они получили название ретроидные вирусы.
– представителями этой группы вирусов являются вирус гепатита В и вирус
мозаики цветной капусты.
1. Репликация ДНК-генома этих вирусов осуществляется при посредстве
промежуточных молекул РНК: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. Молекулы РНК образуются в результате транскрипции вирусных ДНК в
клеточном ядре хозяйским ферментом ДНК-зависимой РНК-полимеразой.
3. Транскрибируется только одна из нитей вирусной ДНК.
4. Синтез ДНК на РНК-матрице происходит в результате реакции,
катализируемой обратной транскриптазой; сначала синтезируется (-) нить
ДНК,
5. а затем на вновь синтезированной (-) нити ДНК тот же фермент строит
(+) нить.
В целом общая схема репликации генома ретроидных вирусов поразительно
похожа на схему репликации генома ретровирусов. По-видимому, данное
сходство имеет под собой и эволюционную основу, так как первичная
структура обратных транскриптаз этих вирусов выявляет определенное
сходство между собой.
Вторая группа — вирусы с двуцепочечной ДНК,
– репликация осуществляется по схеме ДНК -> ДНК.
– с генома этих вирусов в зараженной клетке ДНК-зависимая РНК-полимераза
транскрибирует молекулы мРНК (т.е. (+) РНК),
– мРНК (т.е. (+) РНК) принимает участие в синтезе вирусных белков,
– размножение вирусного генома осуществляет фермент ДНК-зависимая
ДНК-полимераза: (±) днк ? (+)РНК
В одних случаях производством как мРНК, так и ДНК занимаются клеточные
ферменты; в других случаях вирусы используют собственные ферменты.
Бывает, что те и другие ферменты обслуживают процесс репликации и
транскрипции. К этой группе относятся вирусы герпеса, оспы и др.
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a000000000000008100840000000000040000002701ffff030000000000- вирус
герпеса
Оспа – естественный враг СПИДа (нет оспы – есть СПИД). О СПИДе есть
информация в Ветхом завете. В нашем геноме есть генетические метки
прежних пандемий СПИДа
Вирусные заболевания периодичны: ОСПА ? ПРОКАЗА?ЧУМА ? †
Третья группа — вирусы с одноцепочечной ДНК, либо с негативной, либо с
позитивной полярностью.
– Попав в клетку, вирусный геном сначала превращается в двуцепочечную
форму,
– это превращение обеспечивает клеточная ДНК-зависимая ДНК-полимераза:
(+) ДНК
или (±) ДНК ? (+) РНК
(-) ДНК
Транскрипция и репликация на последующих этапах происходит так же, как и
для вирусов, с (±) ДНК-геномом.
Структура вируса: это молекула ДНК в белковой оболочке, называемой
капсидом. Однако есть много разных вариантов строения вирусов: от просто
покрытой белком ДНК (например, бактериофаг Pf1) до сложных
макромолекулярных комплексов, окруженных мембранными структурами,
например, вирус оспы. Если у вируса есть мембрана‚ говорят, что он в
оболочке, а если мембраны нет, то вирус называют «раздетым». Различают
четыре основных вида капсидов: спиральные, икосаэдрические, сложные без
оболочки, сложные с оболочкой.(см. презентация)
Спиральные капсиды
– обычно встречаются у нитевидных вирусов.
– образуются путем самосборки асимметричных белковых субъединиц
(капсомеров), объединяющихся в трубчатую структуру со спиральной
симметрией (например у Pf1).
– Субъединицы в большинстве случаев гомогенны, так, что поверхность
вириона состоит из множества копий одного и того же белка, хотя под
наружним капсидом могут находиться и другие белки.
– ДНК в таких вирусах либо вытянута, либо может быть туго скручена в
комплексе со специальными связывающими белками.
Икосаэдрические капсиды
– свойственны большинству сферических ДНК-содержащих вирусов
– икосаэдр – это многогранник с двадцатью треугольными гранями, имеющий
кубическую симметрию и приблизительно сферическую форму.
– Вершины треугольников соединяясь образуют двенадцать вершин икосаэдра;
– в местах соединения располагаются обычно пентамерные белковые
структуры – пентоны; там же могут находится участки, на которых
формируются белковые нити, нередко ассоциированные с вершинами (
например у ф Х174 прозрачка 1).
– Грани икосаэдра заполнены другими белковыми субъединицами,
сгруппированными обычно в гексамерные структуры – гексоны (апример, у
аденовируса прозрачка 1).
– Количество субъединиц, необходиимое для заполнения граней,
определяется размерами вириона в целом, и разные икосаэдрические вирусы
содержат поэтому разное число гексонов – обычно при неизменном числе
пентонов.
– ДНК обычно плотно свернута внутри капсида;
– иногда она связана с белками или полипептидами, способными
стабилизировать ее структуру.
Сложные капсиды без оболочки
– типичны для бактериофагов:
– они состоят из частей с разными типами симметрии.
– У бактериофага Т2, например, ДНК находится в икосаэдрической головке,
а для «узнавания» бактерии и введения в нее ДНК служат трубчатые и
фибриллярные структуры (в узнавании участвует также лизоцим,
расположенный на дистальном конце хвостового отростка).
Сложные капсиды с оболочкой
– есть только у вирусов эукариотических клеток.
– В них ДНК-белковые комплексы окружены одним или несколькими белковыми
слоями и наружней мембраной, почти все белковые компоненты которой
являются вирусными по своему происхождению, а липидные структуры –
клеточными.
Инфицирование – процесс, посредством которого вирус внедряется в
клетку-хозяина и «настраивает» ее метаболический аппарат на
воспроизведение вирионов.
– зараженные вирусом клетки либо остаются живыми (тогда говорят, что
вирус невирулентен),
– либо подвергаются лизису, приводящему к высвобождению вирусных частиц.
– Неизменным итогом заражения клеток ДНК-содержащими бактериофагами
является лизис.
– ДНК-содержащие вирусы животных вызывают лизис редко; однако клетки
могут погибнуть из-за возникших при заражении хромосомных повреждений,
вследствии иммунологической реакцииорганизма или просто в результате
нарушения вирусом нормальных клеточных функций.
Размножение вируса – четко очерченный цикл, приводящий в конечном счете,
после синтеза новых молекул вирусных белков и большого числа копий
вирусной ДНК, к формированию зрелых вирусных частиц. У вирусов бактерий
весь цикл может завершаться менее чем за час, тогда как у многих вирусов
животных он занимает не один день.
Адсорбция вируса на клетке-хозяине – первый этап инфицирования. Она
происходит на специфических рецепторных участках (белковых или липидных)
клеточной поверхности, которые узнаются особыми выступающими частями
вириона и к которым он прочно прикрепляется. У вирусов без оболочки
такими частями могут быть белковые отростки (например, у аденовируса и
бактериофага Т2), а у вирусов с оболочкой это, как правило белки,
погруженные в вирусную мембрану. В процессе адсорбции осуществляются, в
частности, такие белок-белковые взаимодействия, результатом которых
является инициация стадии проникновения ДНК в клетку.
Проникновение вирусной ДНК в клетку-хозяина у разных вирусов происходит
по-разному. ДНК многих бактериофагов, например, бактериофага Т2:
белковый стержень сплющивается, подобно телескопической конструкции, и
ДНК «впрыскивается» в бактерию. Из вирусов животных ДНК обычно переходит
в клетку в результате как бы слияния наружнего слоя вириона с клеточной
мембраной. В отличие от ДНК большинства бактериофагов ДНК вирусов
животных всегда входят в клетку вместе с непосредственно прилегающими к
ней белками; последующее освобождение ДНК от этих белков осуществляется
с помощью ферментов.
Транскрипция и репликация генетического материала вируса осуществляется
обычно с участием ферментов клетки-хозяина. Сначала вирусная ДНК
копируется РНК-полимеразами клетки-хозяина, в результате чего образуется
мРНК, которая затем транслируется. На некоторых молекулах вирусной ДНК
синтезируются также ее ДНК-копии – с помощью либо клеточной, либо
кодируемой вирусом ДНК-полимеразы. Эти ДНК-копии используются в
последствии при сборке вирусных частиц. В некоторых случаях, например, у
бактериофага Т4 первые же новосинтезированнные молекулы вирусной мРНК
транслируются с образованием специальных белков, модифицирующих
полимеразы клетки-хозяина таким образом, что те прекращают транскрипцию
клеточных генов, не теряя пари этом способности транскрибировать
вирусные. В какой части клетки протекают процессы транскрипции и
репликации вирусной ДНК эукариотических вирусов: в ядре или цитоплазме?
У одних транскрипция и репликация происходит в ядре клетки-хозяина (
например, у вируса герпеса), а у других – в цитоплазме, например, у
поксивирусов.
Трансляция вирусной мРНК на рибосомах клетки-хозяина приводит к
образованию вирусных белков. Некоторые из этих белков используются в
последствии для построения капсидов, другие связываются с вирусной ДНК,
стабилизируя ее (у многих вирусов животных), третьи хотя и не войдут
никогда в состав зрелых вирионов, участвуют в процессе их сборки в
качестве ферментов (например, у бактериофага Т2)
Сборка вируса из его компонентов в клетке-хозяине может происходить
спонтанно (самосборка), но может зависеть и от участия вспомогательных
белков. Вирусная ДНК обычно покрывается слоем белка – капсидом. Капсид,
в свою очередь, может заключаться в мембранную структуру, получаемую
вирионом обычно от клетки-хозяина: покидая клетку путем отпочковывания
от нее, вирусная частица оказывается окруженной плазматической
мембраной.
ДНК-содержащие опухолеродные вирусы разделяются на следующие 5 классов.
1. Полиомавирусы – обезьяний вирус SV40, вирус полиомы мышей и вирусы
человека ВК и JC.
2. Папилломавирусы – 16 вирусов папилломы человека и множество
папилломовирусов животных.
3. Аденовирусы – 37 вирусов человека, множество аденовирусов животных
(например, 24 вируса обезьян и 9 вирусов крупного рогатого скота).
4. Герповирусы – вирусы простого герпеса человека, цитомегаловирус
человека, вирус Эпштейна–Барр и онкогенные вирусы приматов, лошадей,
кур, кроликов, лягушек.
5. Вирусы, подобные вирусу гепатита В, – вирус гепатита В человека,
гепатита североамериканского сурка, гепатита земляных белок и гепатита
уток.
РНК-СОДЕРЖАЩИЕ ВИРУСЫ
РНК-содержащие вирусы
1 — парамиксовирусы; 2 — вирусы гриппа; 3 — коронавирусы; 4 —
аренавирусы; 5 — ретровирусы; 6 — реовирусы; 7 — пикорнавирусы; 8 —
капицивирусы; 9 — рабдовирусы; 10 — тогавирусы, флавивирусы; 11 —
буньявирусы
РНК-содержащие вирусы (РНК-» РНК). не имеют ДНК; генетическая информация
закодирована в РНК. РНК может быть одно и двуцепочечной, а клетка-хозяин
– про, или эукариотической. Только вирусы с одноцепочечной РНК заражают
бактерии, а эукариотические вирусы могут быть как одно – так и
двухцепочечными.
Геномы почти всех известных РНК-содержащих вирусов — это линейные
молекулы, их удобно разделить на 3 группы.
Первая группа — это однонитевые геномы положительной полярности, т.е. с
нуклеотидной последовательностью, соответствующей таковой у мРНК.
– Такие геномы обозначают как (+) РНК.
– Вирусные (+) РНК-геномы кодируют несколько белков, среди которых
РНК-зависимая РНК-полимераза (репликаза), способная синтезировать
молекулы РНК без участия ДНК.
– С помощью этого фермента синтезируются сначала (-) нити РНК фага,
– затем при наличии особого белка, называемого «хозяйским фактором»,
репликаза осуществляет синтез (+) нити РНК.
– на заключительной стадии из накопившихся вирусных белков и (+) РНК
формируются вирионы.
– Упрощенная схема этого процесса такова:
(+) РНК (-) РНК
– Однонитевый (+) РНК-геном характерен для
а) фага Q?,
б) вирусов табачной мозаики,
Вирус табачной мозаики – пример + одноцепочечного вируса растенийвирус
не имеет оболочки, спиральный, содержит 2130 идентичных молекул белка
капрсида и одну цепь РНК. РНК располагается в спиральном желобке,
обрапзованном белковыми субъединицами, и удерживается многочисленными
слабыми связями.
Инфекционный процесс, протекающий по схеме (прозрачка 2 внизу), состоит
в проникновении вируса в растительную клетку с последующей быстрой
утратой им капсида. Затем в результате трансляции непосредственно
+одноцепочечной вирусной РНК рибосомами клетки-хозяина образуются
несколько белков, часть которых необходима для репликации вирусного
генома.
Репликация осуществляется РНК-репликазой, продуцирующей копии РНК для
новых вирионов. Синтез белка капсида происходит только после того как
инфицировавшая клетку РНК подвергается некоторой модификации, делающей
возможным присоединение рибосом клетки к тому участку РНК, которым
кодируется этот белок. Сборка вириона начинается с образования дисков из
белка капсида. Два таких белковых диска, располагаясь концентрически,
образую похожую на бисквит структуру, которая после связывания с ней РНК
приобретает форму спирали. Последующее присоединение молекул белка
продолжается до тех пор, пока РНК не будет покрыта полностью. В своей
окончательной форме вирион представляет собой цилиндр длиной 300 нм.
3) полиомиелита,
4) клещевого энцефалита.
Вторая группа — это однонитевые геномы с негативной полярностью, т.е.
(-) РНК-геномы.
Поскольку (-) РНК не может выполнять функции мРНК, для образования
«своих» мРНК вирус внедряет в клетку не только геном, но и фермент,
умеющий снимать с этого генома комплементарные копии по схеме:
(-) РНК (+) РНК
– Этот вирусный фермент (РНК-зависимая РНК-полимераза, синтезированная в
предыдущем цикле размножения) упакован в вирионе в удобной для доставки
в клетку форме.
– Инфекционный процесс начинается с того, что вирусный фермент копирует
вирусный геном, образуя (+) РНК, которая выступает в качестве матрицы
для синтеза вирусных белков, в том числе РНК-зависимой РНК-полимеразы,
которая входит в состав образующихся вирионов
– К вирусам с негативным РНК-геномом относятся
а) вирусы гриппа,
б) кори,
в) бешенства,
г) желтой карликовости картофеля и др.
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– Вирусная частица содержит две молекулы геномной одноцепочечной (+)
РНК.
– В вирусном геноме закодирован необычный фермент (обратная
транскриптаза, или ревертаза), который обладает свойствами как
РНК-зависимой, так и ДНК-зависимой ДНК-полимеразы.
Только в 1970 г. американские ученые Г. Темин и Мицутани и независимо от
них Д.Балтимор разрешили эту загадку. Они доказали возможность передачи
генетической информации от РНК к ДНК. Это открытие перевернуло
центральную догму молекулярной биологии о том, что генетическая
информация может переноситься только в направлении ДНК–РНК–белок. Пять
лет понадобилось Г. Темину для обнаружения фермента, осуществляющего
перенос информации от РНК к ДНК, – РНК-зависимой ДНК-полимеразы. Этот
фермент получил название обратной транскриптазы. Г. Темину удалось не
только получить фрагменты ДНК, комплементарные заданной цепи РНК, но и
доказать что ДНК-копии могут встраиваться в ДНК клеток и передаваться
потомству.
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fffffffffffffffffffff3fffffffffffffffffffffffffffffffe0fffffffffffffffff
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fffff3fffffffffffffffffffffffffffffffe0ffffffffffffffffffffffffffffff3ff
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0000000f300fb0000000000040000002701ffff030000000000
– Этот фермент попадает в заражаемую клетку вместе с вирусной РНК и
обеспечивает синтез ее ДНК-копии сначала в одноцепочечной форме [(-)
ДНК], а затем в двуцепочечной [(±) ДНК]:
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0010100ffffffffffffffffffffffffffffffffffffffffffffffffffffffffffffffff8
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0010101ffffffffffffffffffffffffffffffffffffffffffffffffffffffffffffffff8
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00101000c00000040092900aa00000000000000e701010100000000040000002701ffff0
30000000000
– Вирусный геном в форме нормального дуплекса ДНК (так называемая
провирусная ДНК) встраивается в хромосому клетки хозяина.
– В результате двуцепочечная ДНК вируса представляет собой в сущности
дополнительный набор генов клетки, который реплицируется вместе с ДНК
хозяина при делении.
– Для образования новых ретровирусных частиц провирусные гены (гены
вируса в хромосомах хозяина) транскрибируются в ядре клетки
транскрипционным аппаратом хозяина в (+) РНК-транскрипты.
– Одни из них становятся геномом нового «потомства» ретровирусов, а
другие подвергаются процессингу в мРНК и используются для трансляции
белков, необходимых для сборки вирусных частиц
В эту группу входят
а) вирус иммунодефицита человека (ВИЧ)
– информация о СПИДе есть в Ветхом Завете
– в нашем ганоме есть генетические метки прежних пандемий СПИДа
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