Представители разных семейств и подотрядов палочников, у которых были изучены пектиназы. По часовой стрелке, начиная с левого верхнего угла: калимантанский палочник Aretaon asperrimus (фото с сайта biolib.cz), калифорнийский палочник Timema cristinae (фото с сайта sciencebrainwaves.com), вьетнамский палочник Ramulus artemis (фото с сайта skorpione.de), кордильерский палочник Peruphasma schultei (фото с сайта insekten-kleinanzeigen.net), розовокрылый мускусный палочник Sipyloidea sipylus (фото с сайта wikimedia.org)

Представители отряда палочников имеют в своем биохимическом арсенале пектиназы — ферменты, расщепляющие пектин. Сравнивая гены пектиназ у палочников и двух других групп животных (жуков-листоедов и нематод), а также у различных бактерий, международная группа ученых выявила источник этих генов. Палочники, как выяснилось, получили гены пектиназ от своих бактериальных соседей — гамма-протеобактерий, обитающих в их кишечнике и на их кормовых растениях. Новое биохимическое свойство оказалось для растительноядных палочников чрезвычайно полезным: оно резко ускорило видообразование, так что в итоге сформировалась обширная группа, насчитывающая около 3000 видов. Это исключительно наглядный пример формирования таксона высокого ранга — отряда — после обретения нового признака, в данном случае биохимического и к тому же позаимствованного у бактерий за счет горизонтального переноса генов.

Новое убедительное доказательство горизонтального переноса генов (ГПГ) от прокариот к эукариотам представили ученые из Института химической экологии имени Макса Планка в Йене (Германия) со своими коллегами из Института зоологии и эволюционной биологии Йенского университета, Института зоологии и антропологии Гёттингенского университета, Университета Ниццы (Франция) и Национального китайского банка генов в Шэньчжэне. Сам факт ГПГ от бактерий к эукариотам уже не слишком удивляет: к настоящему времени найдено множество нагляднейших примеров этого явления (Бактериальные гены помогают нематодам паразитировать на растениях, «Элементы», 18.10.2010; Животные обмениваются генами с паразитическими бактериями, «Элементы», 05.09.2007; Горизонтальный обмен генами заменяет коловраткам половое размножение, «Элементы», 07.06.2008). Новый пример, помимо надежного доказательства факта горизонтального переноса от бактерий к эукариотам, интересен двумя дополнительными аспектами. Во-первых, это случай множественного независимого ГПГ со сходной функцией; во-вторых, для этого случая хорошо обоснована идея о взрывной эволюции, которая привела к формированию обширного таксона высокого ранга. Иными словами, мы видим пример становления большого таксона после приобретения полезного бактериального гена.

Речь идет о весьма разнообразном отряде палочников Phasmatodea, объединяющем около 3000 видов. Современные систематики подразделяют их на несколько подотрядов и семейств, самый примитивный из которых Timematodea. Он включает один род Timema с 20–30 видами. Палочники, а именно предки Timema, появились примерно 125 млн лет назад, однако интенсивное видообразование в этой группе, судя по молекулярным данным, происходило в начале кайнозоя, 60 млн лет назад. Тогда к Timema добавились все остальные семейства палочников. Период их активного видообразования как раз совпадает с бурной радиацией цветковых растений.

В этом совпадении нет ничего удивительного, ведь палочники предпочитают строгую растительную диету. Они сосут соки растений, и для эффективного потребления этой пищи им необходимо уметь расщеплять вещества прочных растительных тканей — целлюлозу и пектины. Обычно у насекомых этим занимаются их микросимбионты, живущие в пищеварительном тракте. Но не всегда. Так, доказано, что у некоторых групп насекомых имеются собственные пектиназы (см. Pectinase), способные расщеплять пектин не хуже бактериальных аналогов. К таким счастливым обладателям пектиназ относятся представители жуков-листоедов, долгоносиков и палочников, а кроме них собственной пектиназной базой обзавелись некоторые нематоды, живущие в корнях растений. Откуда пектиназы у этих групп?

На этот счет предлагаются две очевидных гипотезы. Первая — пектиназные аналоги имелись у предков линяющих (не будем забывать, что пектиназы есть у нематод и неродственных отрядов насекомых); в ходе эволюции у всех линий членистоногих, предпочитающих белковый рацион, пектиназы исчезли, оставшись только у растительноядных специалистов. Вторая гипотеза — ГПГ, за счет которого отдаленные группы животных получили пектиназные гены независимо друг от друга. Источником генов пектиназ могли быть различные группы бактерий. Эти гипотезы можно проверить. Если верна первая из них, то все пектиназы, имеющиеся у нематод и насекомых, будут сходны. Их происхождение можно будет вывести из какой-то одной предковой линии. Если же верна вторая гипотеза, то в происхождении наборов пектиназ у трех групп будет мало общего: вероятно, у них реконструируются два или три изначальных корня.

Ученые выбрали из базы генетической информации GenBank все данные по нуклеотидным последовательностям пектиназ насекомых, добавили собственные данные, полученные по транскриптомам шести представителей палочников, четыре из которых изучили наиболее тщательно. Получилось около 90 вариантов пектиназ. Затем их сравнили между собой и с пектиназами различных линий бактерий. Результаты сравнения показали, что пектиназы нематод, жуков-листоедов и палочников группируются по отдельности, и каждому кластеру соответствуют свои специфические бактериальные пектиназы. Нематодные пектиназы близки к ферментам гамма- и бета-протеобактерий, пектиназы жуков — к пектиназам бактерий из типа Bacteroidetes, а палочников — гамма-протеобактерий, не родственных линиям носителей нематодных пектиназ.

Филогения генов пектиназ у насекомых, нематод и бактерий. Палочники — зеленый кластер, листоеды — оранжевый кластер, нематоды — красный кластер. Видно, что каждая группа животных близка к своей бактериальной линии (они показаны синим). Рисунок из обсуждаемой статьи в Scientific Reports

Это безоговорочное свидетельство независимого приобретения генов пектиназ от бактерий. Значит, сценарий изначального присутствия пектиназ у общих предков линяющих животных не подтверждается. Насекомые и нематоды позаимствовали гены пектиназ, когда в них возникла необходимость и, скорее всего, волею счастливой случайности. Правообладателями этих генов были или бактерии-симбионты, или свободноживущие бактерии, населяющие кормовые растения.

Пектиназы были найдены у четырех из пяти крупных подгрупп палочников. Активность пектиназ в каждой из этих четырех подгрупп имеет свои особенности, формируя четкие кластеры. Иными словами, по активности пектиназ можно точно указать, к какому семейству относится исследуемый представитель палочников, или, наоборот, для каждого представителя палочников можно предсказать набор его пектиназ. Но самое любопытное в этой картине не присутствие, а отсутствие пектиназ у пятой подгруппы палочников. Этой подгруппой является Timematidae, примитивные палочники.

Филогения родственных палочникам групп насекомых и наличие у них пектиназ (зеленая область). У палочников Timema пектиназ нет. Шкала внизу слева указывает на время разделения ветвей (в млн лет назад). Рисунок из обсуждаемой статьи в Scientific Reports

Ученые с полным основанием полагают, что пектиназные гены попали к палочникам после становления этого семейства, но до начала интенсивного видообразования в этой группе. Timematidae существовали в течение 60 млн лет, пользуясь услугами бактериальных сожителей, расщепляющих пектин. Затем, после получения от них гена/генов пектиназ в личное пользование, началось бурное видообразование палочников. В итоге очень быстро сложилось сотни новых видов. Эти виды объединяются в отдельный отряд. Но неизвестно, как поступили бы систематики, если бы из всех палочников им были бы известны только Timema. Вероятно, присоединили бы к отряду родственных эмбий или тараканосверчков.

Но, так или иначе, перед нами целый разнообразный отряд, который сформировался не в последнюю очередь в результате приобретения полезного биохимического признака. Он дал им возможность лучше справляться с обильной, но трудно перевариваемой пищей. Данный признак не сложился постепенно в ходе эволюции, а был получен от бактериальных соседей с помощью ГПГ. Это событие — удачный горизонтальный перенос генов — случилось в правильное время (радиация цветковых растений) и у правильной группы (предрасположенной к растительному рациону).

Источник: elementy.ru

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