Вместе с рисом искусственному отбору подвергся и мимикрирующий под него сорняк

Рис. 1. Крестьяне эпохи династии Сун (980–1279 годы н. э.) пропалывают рис. Сорняки, которые по виду больше других походят рис, остаются на поле, дают семена и высеваются со следующим урожаем. Именно так и появилась рисоподобная форма ежовника обыкновенного, самого распространенного сорняка рисовых полей. Рисунок с сайта www.readworks.org

Международная группа ботаников и генетиков разобралась с тем, как появился сорняк, похожий внешне на обычный рис. Это растение ежовник, у которого есть разные фенотипы: и рисоподобный прямостоячий, и исходный «приземленный». Сделав полногеномный анализ 328 экземпляров ежовника, ученые выявили действие отбора, который привел к формированию сходных с рисом морфологических признаков. Похожий на рис фенотип сформировался около тысячи лет назад, когда в бассейне реки Янцзы активизировалось рисоводство, заместив культуру пшеницы. Исследователям удалось показать, что в этом регионе в ходе селекции риса и эволюции сопутствующего ему ежовника параллельно менялись одни и те же гены, контролирующие одни и те же морфологические признаки. Последнее особенно важно для понимания сущности явления конвергенции.

Искусственный отбор и доместикация растений и животных — важная черта хозяйственной деятельности человека, прошедшего рубеж бронзового века. Если рассматривать одомашненные виды растений и животных, то они, с эволюционной точки зрения, исключительно успешны: они достигли высочайшей численности и распространились гораздо шире своих диких предков. Очевидно, что «быть как они» выгодно, по крайней мере, в краткосрочной перспективе.

По пути подражания успешным окультуренным видам пошли многие сорняки возделываемых растений. Они приобрели признаки своих полезных полевых сожителей, став похожими на них, и таким образом, получили все их преимущества. Это явление известно как вавиловская мимикрия: благодаря ей из обширного круга различных подражателей из-за внешнего сходства выделяются пары «культурное растение — сорняк». Механизм такой мимикрии предположительно связан с искусственным отбором, а не с «естественными» попытками сорняков оптимальнее соответствовать условиям среды. При всей логичности такой гипотезы есть не так уж много хорошо разобранных примеров исследования селекции на уровне фенотипов и генотипов сорняков и их культурных моделей, да еще с историко-демографической базой.

Такой пример, показательный во всех отношениях, опубликовали специалисты из Китая и США (представляющие Чжэцзянский университет, Государственный институт рисоводства (оба расположены в Ханчжоу, Китай), Лабораторию генетики растений Института генетики и биологии развития (Пекин, Китай), биологическое отделение Вашингтонского университета (Сент-Луис, США) и Виргинский университет (Шарлоттсвиль, США)). Они работали с рисом (Oryza sativa) — главной сельскохозяйственной культурой Китая — и его сорняком ежовником обыкновенным (Echinochloa crus-galli). Эти два растения — типичная пара вавиловской мимикрии (рис. 2).

Вместе с рисом искусственному отбору подвергся и мимикрирующий под него сорняк
Вместе с рисом искусственному отбору подвергся и мимикрирующий под него сорняк

Рис. 2. Ежовник (справа) маскируется под культурный рис (слева). Если посадки риса пропалывать вручную (а именно так и происходило во все времена), то не всякий земледелец с легкостью отличит полезное растение от его сожителя, оставив сорняк расти. Рисунок из обсуждаемой статьи в Nature Ecology & Evolution

Данная пара удобна для исследования помимо прочего тем, что ежовник очень изменчив, у него может проявиться и рисоподобный фенотип, и множество других, на рис совсем не похожих (рис. 3). Ученые выделили четыре основных фенотипических признака, отличающие растения-подражатели от других фенотипов ежовника. Это угол наклона стебля к земле (у подражателей стебель близок к пряморастущему, у других — наклонен к земле), цвет основания стеблей (зеленый или красноватый), искривленность междоузлий (прямые или угловатые), собранность листьев (собранные или расставленные).

Вместе с рисом искусственному отбору подвергся и мимикрирующий под него сорняк
Вместе с рисом искусственному отбору подвергся и мимикрирующий под него сорняк

Рис. 3. Фенотипы ежовника: слева — рисоподобный, справа — «приземленный». Они различаются по четырем признакам: наклон стеблей к земле, сгруппированность листьев, кривизна стебельков в междоузлиях, цвет основания стеблей. Фото из обсуждаемой статьи в Nature Ecology & Evolution

Используя градации для этих признаков, можно собрать растения и количественно оценить степень схожести (в работе используется индекс мимикрии, равный сумме баллов каждого признака: чем меньше балл, тем меньше мимикрия по признаку, и, следовательно, индекс тем меньше, чем меньше сходства по всем признакам) риса и его сорняка.

Растения собирали по всему сельскохозяйственному региону бассейна Янцзы, всего в 328 местах: как на рисовых полях, так и в отдалении от них. Семена растений из каждой точки (всего по 8 семян) проращивали в теплице, а затем высаживали на опытных чеках. Уровень сходства оценивали у трехнедельных проростков. Чтобы убедиться в устойчивости фенотипа, оценивали и фенотип потомков на следующий год, и реакцию выборочных растений на разные световые режимы. Нужно подчеркнуть, что и повторные высаживания, и выращивание при разных продолжительностях светового дня показали стабильность фенотипов, что подразумевает генетическую базу данного фенотипического сдвига. У каждого растения был прочитан полный геном (написать это быстро, а сама работа в действительности гигантская!).

Вместе с рисом искусственному отбору подвергся и мимикрирующий под него сорняк

Рис. 4. Филогенетическое дерево ежовника, построенное по однонуклеотидным заменам после анализа 328 экземпляров. Цветами обозначены кластеры, включающие рисоподобный фенотип (MIC), исходный фенотип (NMC) и смешанный (MIX). Рисунок из обсуждаемой статьи в Nature Ecology & Evolution

Очевидно, что растения с рисоподобным обликом имели высокий индекс мимикрии, близкий к 8 или больше, а у растений с исходной внешностью индекс мимикрии обычно не превышал 2. Анализ генотипов всех растений позволили определить у них порядка 9 млн однонуклеотидных замен. Построенное по ним дерево показало четкое разделение на три кластера (рис. 4). И эти кластеры совпали с тремя фенотипическими группами: с высоким индексом мимикрии, с низким индексом мимикрии и с промежуточным показателем. Это означает, что рисоподобный фенотип имеет монофилетическое происхождение: если бы рисоподобный ежовник возникал много раз независимо, то в трех кластерах представители фенотипов были бы перемешаны.

У подражателей оказалось урезанное разнообразие однонуклеотидных замен по сравнению с «приземленными» родичами: все варианты замен, найденные у подражателей, имеются у «приземленного» фенотипа, но он сам имеет и такие, которых нет у подражателей. Судя по расчетам, сделанным несколькими методами, рисоподобный фенотип берет начало от «приземленного» (а значит, исходного), пройдя через бутылочное горлышко. Все это происходило около тысячи лет назад. Это период династии Сун — как раз тогда в бассейне реки Янцзы население быстро разрослось и стало очень многочисленным, а пшеница — основная зерновая культура — была замещена рисом.

Распределение мутаций (замен, вставок и делеций) у растений-подражателей и у растений, имеющих исходный фенотип, позволило выявить семь участков генома размером больше 10 тысяч пар нуклеотидов, где различия в два раза выше, чем в среднем по геному. Такая ситуация напоминает «острова дивергенции» — участки генома, куда нацелены силы естественного отбора. В таких крупных участках с пониженным разнообразием и повышенным уровнем различий удалось выявить четкий сигнал действия отбора. В них в общей сложности сосредоточено 1986 генов с 8373 несинонимичными (то есть потенциально важных для фенотипа) заменами.

Чтобы добавить конкретики этим цифрам, ученые рассмотрели гены, функции которых связаны с архитектурой растения. В участках дивергенции среди выявленных 1986 генов-кандидатов количество таких генов (всего 87) выше среднего по геному. Например, ген LAZY1 контролирует угол роста стебля за счет распределения в стебле ауксина. Участок генома с этим геном у рисоподобного фенотипа сильно отличается от своего исходного аналога, разнообразие у подражателей в этом участке резко понижено, и при этом большая часть вариаций (70%) приходится на один гаплотип, весьма малочисленный (15%) среди растений с исходным фенотипом. Важно, что и у самого культурного риса под действие селекции попал именно этот ген. Так что на примере риса и сопутствующего сорняка ежовника мы видим параллельное изменение одного и того же гена под действием одного и того же усилия селекционеров-рисоводов.

Говоря о мимикрии, нельзя не вспомнить о феномене конвергентной эволюции, когда в самых разных группах организмов появляются внешне сходных формы или формы со сходными сложными признаками. Считается, что конвергентные признаки формируются в силу сходных требований среды и соответственно сходных функциональных решений у разных видов, но наборы генов, вовлеченных в формирование таких признаков, должны быть разными. Сейчас появляется все больше данных, что и гены, участвующие в этом процессе, могут быть сходными. Мимикрия в чем-то похожа на конвергенцию — это формирование сходного облика у двух эволюционно далеких групп. Пример с геном распределения ауксина LAZY1 добавляет доводов в пользу гипотезы, что сходные признаки у филогенетически далеких видов могут формироваться за счет одних и тех же генов.

Источник: elementy.ru

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