Гибридизация однодомных и двудомных растений увеличивает разнообразие половых фенотипов

Рис. 1. Стрелолист широколистный (Sagittaria latifolia). Фото с сайта www.esd.ornl.gov

В последнее время накапливается все больше сведений в поддержку так называемой «сетчатой эволюции». Речь идет о происхождении новых форм путем гибридизации двух других. Первый шаг этого процесса — появление гибридных популяций с какими-то особыми свойствами. Часто это причудливая смесь черт родительских форм — неважно, принадлежат ли они к одному виду или к разным. Некоторые виды растений распадаются на однодомные и двудомные популяции. В однодомных на каждом растении есть и мужские, и женские органы, а в двудомных мужские и женские органы находятся на разных растениях. На примере стрелолиста канадские ученые показали, что гибридизация исходно однодомных и двудомных популяций может вести к формированию смешанных популяций, в которых в разных соотношениях присутствуют гермафродиты (однодомные) и мужские и/или женские растения. Это новый механизм формирования разнообразия половых фенотипов в популяциях растений.

Многие годы ученые игнорировали значения для популяций животных такого явления, как межвидовая гибридизация. Но в последние десятилетия стали накапливаться данные о важной роли гибридизации в некоторых случаях. Главное значение ее в том, что приток генов от другого вида изменяет и разнообразит генофонд популяции данного вида. А это, в свою очередь, может повлиять на его судьбу, приспособленность к среде обитания и прочие параметры. О некоторых известных случаях можно прочесть на «Элементах»: см., к примеру, про дарвиновых вьюрков (Разнообразие дарвиновых вьюрков сокращается из-за межвидовой гибридизации, «Элементы», 24.03.2014); про партеногенетические виды, возникшие в результате гибридизации (Партеногенетические самки рыбки моллинезии более агрессивны, «Элементы», 27.08.2015); про влияние гибридизации на полиморфизм (Где какой цвет в моде, «Элементы», 17.08.2015).

Недавно канадские ученые выяснили, как и к чему еще может приводить межвидовая гибридизация — на этот раз у высших растений.

Известно, что большинство представителей этой группы являются однодомными. Это значит, что каждое растение обладает и пестиками (женские половые органы), и тычинками (мужские половые органы) и является таким образом гермафродитом. Но около 6–7% всех видов — двудомные, у которых пестики и тычинки находятся на разных (мужских и женских) растениях. А иногда встречаются и интереснейшие ситуации, переходные между однодомностью и двудомностью. Например, в одной популяции могут одновременно присутствовать растения мужские, женские и гермафродиты.

Откуда взялось такое разнообразие систем размножения? Обычно считают, что в эволюции растений чаще встречался переход от однодомности к двудомности, нежели наоборот, а переходная ситуация является неустойчивой. Она возникает, когда женские растения попадают из двудомной популяции в однодомную. Они «помогают» здесь однодомным гермафродитам избегать инбридинга (проще говоря — самоопыления). Постепенно у гермафродитов женские половые органы перестают функционировать, и они становятся мужскими растениями, то есть популяция становится двудомной. Но в целом такие смешанные популяции растений изучены еще недостаточно, поэтому тут может быть много открытий. С одной из таких ситуаций и разобрались ученые, написавшие обсуждаемую статью.

Они изучали стрелолист широколистный (Sagittaria latifolia) из семейства Частуховые (Alismataceae); рис. 1. Это многолетнее водное растение широко распространено в Северной Америке и встречается в самых разных типах водоемов. На северо-востоке Северной Америки около 2/3 популяций являются либо однодомными, либо двудомными (рис. 2).

Гибридизация однодомных и двудомных растений увеличивает разнообразие половых фенотипов

Рис. 2. Географическое распространение однодомных (количество популяций n = 25, серые кружки), двудомных (n = 27, белые квадраты), смешанных (гермафродиты и мужские и женские фенотипы, n = 13, черные треугольники), андро-двудомных (мужские фенотипы и гермафродиты, n=6, черные шестиугольники) и гино-двудомных (женские фенотипы и гермафродиты, n = 1, ромб). Всего 72 популяции. Рисунок из обсуждаемой статьи в Evolution

В остальных доля гермафродитов (то есть однодомных растений) варьирует от 11 до 79%, а остальные — это мужские и/или женские растения (рис. 3).

Гибридизация однодомных и двудомных растений увеличивает разнообразие половых фенотипов

Рис. 3. Разнообразие фенотипического состава 116 популяций (включая те 72, что изображены на рис. 2) стрелолиста широколистного. По оси Y — доля разных фенотипов. Внизу все популяции условно классифицированы на три группы: к смешанным здесь относятся как собственно смешанные, так и андро-двудомные и гино-двудомные (см. рис. 2). Рисунок из статьи S. B. Yakimowski, S. C. H. Barret, 2014. Variation and evolution of sex ration at the northern range limit of a sexual polymorphic plant

У стрелолиста широколистного есть еще одна интересная особенность. Половой фенотип растения определяется всего двумя локусами. Первый — это локус «женской» стерильности, доминантный аллель которого (SuF) подавляет развитие женских половых органов. А в локусе «мужской» стерильности всё наоборот: доминантный аллель SM необходим для развития мужских половых органов. Таким образом, в однодомных популяциях все растения имеют генотип SufSM/SufSM. В двудомных популяциях генотип женского растения SufSm/SufSm, а мужские фенотипы имеют одну из двух комбинаций: SuFSM/SufSm или SufSM/SufSm. Выше большая надстрочная буква обозначает доминантный аллель данного гена (например, SM), а маленькая — рецессивный (например, Sm). Схожая система определения пола известна и у некоторых других растений (см. Пол цветка определяется взаимодействием трех генов, «Элементы», 10.11.2015).

Разница между однодомными и двудомными популяциями не сводится только к различиям в комбинациях аллелей. Интересно, что они различаются и экологией — требованиям к местообитанию, временем цветения, массой семян и др. (рис. 4).

Гибридизация однодомных и двудомных растений увеличивает разнообразие половых фенотипов

Рис. 4. Диаграмма рассеяния, показывающая различия между однодомными (n = 16, белые кружки) и двудомными (n = 11, черные квадраты) популяциями стрелолиста широколистного. PC1 и PC2 — соответственно первая и вторая компоненты, полученные анализом главных компонент. Эти компоненты объясняют 29% и 23% изменчивости массива данных из многих переменных, характеризующих особенности жизненного цикла. Среди них высота растения, число листьев на одном растении, день цветения, общая масса семян одного растения, масса одного семени и др. Таким образом, диаграмма характеризует различия между однодомными и двудомными популяциями сразу по многим экологическим параметрам. Рисунок из статьи M. E. Dorken, S. C. H. Barrett, 2003. Life-history differentiation and the maintenance of monoecy and dioecy in  Sagittaria lafifolia (Alismataceae)

Итак, что же выяснилось в данном исследовании? Первоначальной задачей было проанализировать хлоропластную ДНК (хлДНК), то есть ДНК, содержащуюся в хлоропластах. Всего было изучено 66 популяций. Оказалось, что здесь можно выделить две группы гаплотипов — D и M. При этом двудомные популяции (n = 22) включали растения почти исключительно с гаплотипом D, а большинство однодомных (22 из 25) — с гаплотипом M. Популяции смешанного состава (в которых наряду с гермафродитами присутствовали мужские и/или женские растения) по составу гаплотипов были более разнообразны. В большинстве случае они состояли из растений обоих гаплотипов в тех или иных пропорциях (рис. 5).

Гибридизация однодомных и двудомных растений увеличивает разнообразие половых фенотипов

Рис. 5. Частоты гаплотипов D (черный) и М (серый) в двудомных (А), смешанных (В) и однодомных (С) популяциях стрелолиста. Сокращения латинскими буквами указывают на отдельные популяции. Рисунок из обсуждаемой статьи в Evolution

Эти данные навели ученых на мысль о возможности гибридного происхождения смешанных популяций. Для проверки этого предположения они дополнительно изучили набор микросателлитов. Микросателлиты — это короткие повторы последовательностей из нескольких пар оснований. Они характеризуются значительной индивидуальной и популяционной специфичностью и потому часто используются, например, для определения родства или выявления гибридных особей. В последнем случае в специальных программах можно рассчитать гибридные индексы, характеризующие вероятность гибридного происхождения данной особи.

Оказалось, что набор используемых микросателлитов позволяет легко разделять двудомные и однодомные популяции стрелолиста (рис. 6, А и B). А вот в смешанных популяциях различного состава присутствовало много особей гибридного происхождения (рис. 6, С). Таким образом, данный анализ подтвердил, что в смешанных популяциях имеется много гибридов между растениями из однодомных и двудомных популяций. При этом такие гибриды (не обязательно первого поколения) могут быть как гермафродитами, так и женскими или мужскими особями.

Гибридизация однодомных и двудомных растений увеличивает разнообразие половых фенотипов

Рис. 6. Гибридные индексы (по оси Y) особей из однодомных (A), двудомных (B) и смешанных (C) популяций стрелолиста. Номера особей даны по оси X. Значение индекса, близкое к 1,0, указывает на принадлежность к однодомным популяциям. Близкое к 0,0 — к двудомным. А значение 0,5 характеризует гибрида. Рисунок из обсуждаемой статьи в Evolution

Значит, происхождение смешанных популяций стрелолиста во многих случаях обусловлено гибридизацией растений из исходно двудомных и однодомных популяций. Итог такой гибридизации — увеличение разнообразия половых фенотипов в данной гибридной популяции. Пока не ясно, как такое увеличение может отразиться (и отразится ли) на эволюционном успехе. Однако такой необычный итог гибридизации представляет интерес и сам по себе.

Источник: elementy.ru

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