Строение Семенной Кожуры Видов Asteraceae. I. Трибы Arctotideae, Cardueae, Mutisieae, Vernonieae

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Строение Семенной Кожуры Видов Asteraceae. I. Трибы Arctotideae, Cardueae, Mutisieae, Vernonieae Turczaninowia 21 (4): 44–62 (2018) ISSN 1560–7259 (print edition) DOI: 10.14258/turczaninowia.21.4.6 TURCZANINOWIA http://turczaninowia.asu.ru ISSN 1560–7267 (online edition) УДК 582.998:581.48 Строение семенной кожуры видов Asteraceae. I. Трибы Arctotideae, Cardueae, Mutisieae, Vernonieae Э. В. Бойко, Е. В. Новожилова ФГБУН Тихоокеанский институт биоорганической химии им. Г. Б. Елякова ДВО РАН, пр. 100 лет Владивостоку, 159, г. Владивосток, 690022, Россия. E-mail: [email protected], [email protected] Ключевые слова: анатомия, семенная кожура (теста), семянки, систематика, сканирующая электронная микроскопия (СЭМ). Аннотация. Представлены результаты исследований методами световой и сканирующей микроскопии скульптуры поверхности семян и анатомического строения семенной кожуры 35 видов из 28 родов семейства Asteraceae, относящихся к четырем трибам: Arctotideae (Arctotis, Berkheya, Gazania), Cardueae (Acroptilon, Al- fredia, Ancathia, Berardia, Carthamus, Cirsium, Echinops, Klasea, Lamyropappus, Olgaea, Onopordum, Rhaponti- cum, Serratula, Silybum, Synurus, Syreitschikovia, Takeikadzuchia), Mutisieae (Adenocaulon, Dicoma, Gochnatia, Jungia, Leibnitzia, Trichocline, Trixis) и Vernonieae (Vernonia). Семенная кожура исследованных видов хорошо выражена и сохраняется до полной зрелости семянок. Для характеристики семенной кожуры основное зна- чение имеет строение клеток экзотесты, которые имеют преимущественно один тип строения по всей по- верхности семени. Однако есть виды, у которых клетки экзотесты представлены несколькими типами. Так, у Acroptilon repens по всей поверхности семени клетки с утолщёнными стенками чередуются с тонкостенными. У Dicoma schimperi клетки экзотесты в апикальной и базальной частях семени имеют различное строение. Установлено, что морфологическое разнообразие клеток экзотесты связано с формой, размерами, простран- ственной ориентацией клеток по отношению к оси семени, а также характером утолщений тангентальных и радиальных стенок клеток. Комплекс вышеперечисленных признаков отражает видовую специфику струк- туры семенной кожуры. Выявлены и описаны признаки, которые могут быть использованы в систематике исследованных триб. Признаки строения семенной кожуры, характерные для каждой отдельной трибы, не установлены. Structure of the seed coat of the Asteraceae species. I. Tribes Arctotideae, Cardueae, Mutisieae, Vernonieae E. V. Boyko, E. V. Novozhilova G. B. Elyаkov Pacific Institute of Bioorganic Chemistry, Far East Branch, Russian Academy of Sciences, 159 Pr-t 100-let Vladivostoku, Vladivostok, 690022, Russian Federation Keywords: anatomy, cypselas, scanning electronic microscopy (SEM), seed coat, systematics. Summary. The sculpture of the seeds surface and the anatomic structure of the seed coat of 35 species from 28 genera from 4 tribes of the Asteraceae: Arctotideae (Arctotis, Berkheya, Gazania), Cardueae (Acroptilon, Alfredia, Ancathia, Berardia, Carthamus, Cirsium, Echinops, Klasea, Lamyropappus, Olgaea, Onopordum, Rhaponticum, Ser- ratula, Silybum, Synurus, Syreitschikovia, Takeikadzuchia), Mutisieae (Adenocaulon, Dicoma, Gochnatia, Jungia, Поступило в редакцию 06.04.2018 Submitted 06.04.2018 Принято к публикации 21.11.2018 Accepted 21.11.2018 Turczaninowia 21 (4): 44–62 (2018) 45 Leibnitzia, Trichocline, Trixis) and Vernonieae (Vernonia) are investigated by methods of the light and scanning mi- croscopy. The seed coat of the studied species is well developed and remains until the complete maturity of cypselas. The structure of the cells of an exotesta has a basic meaning for the characteristic of a seed coat. Cells of the exotesta have mainly one type of structure on all surfaces of the seed, however there are types at which cells of the exotesta are presented by several types. For example, Acroptilon repens has thin-walled cells alternating with thick-walled cells on all surface of the seed. The exotesta cells of Dicoma schimperi have different structure in apical and basal parts of the seed. As a result of this research it is established that the morphological differences of exotesta cells are caused by form, size, dimensional orientation of cells in relation to the seed axis and also degree of thickenings of tangential and radial walls of cells. The complex of the revealed features of a seed coat characterizes specifics of taxon at the species level. The features of seed coat structure, the characteristic for each separate tribe, are not established. Введение казал константность признаков экзотесты, кото- рые эволюционно обусловлены, стабильны и не Семянка сложноцветных (Asteraceae) – сухой зависят от внешних условий, в связи с чем об- нераскрывающийся односемянный плод, произ- ладают высокой систематической ценностью. водный от нижней завязи. Плодовая оболочка Он выявил в строении экзотесты признаки, ха- (перикарпий), семенная кожура (теста, спермо- рактерные как для отдельных видов, родов, так и дерма) и эндосперм защищают семязачаток. По- для ряда подтриб. кровы семянок сложноцветных разнообразны по A. M. R. Davies (2010), изучая особенности анатомическому строению. Соотношение тол- строения клеток экзотесты представителей три- щины перикарпия и семенной кожуры у разных бы Mutisieae (Chaetanthera Ruiz et Pav., Oriastrum видов неодинаково. Обычно функция защиты Poepp. et Endl.) с поверхности и на поперечных зародыша от неблагоприятных экологических срезах семянок, выявил четыре типа орнамента факторов выполняется преимущественно одним поверхности: параллельные, параллельно-про- из этих покровов. У видов, произрастающих в долговатые клетки; извилистые, псевдодендрит- засушливых местообитаниях, обычно перикар- ные клетки; округлые параллельно-продолго- пий плотный, состоит из толстостенных клеток. ватые клетки; плотно извилистые мозаичные Перикарпий у видов мезофитных условий чаще клетки, а на поперечных срезах разнообразные представлен тонкостенными клетками. склеренхиматозные утолщения, которые клас- Признаки перикарпия и тесты важны для сифицировал в четыре типа: равномерно утол- таксономии семейства. Исследователи плодов щённые стенки; множественные U-образные сложноцветных уделяют основное внимание ребра в клетках; клеточные стенки равномерно строению перикарпия. У многих представителей U-образно утолщённые; неравномерно утолщён- семейства семенная кожура к моменту созрева- ные стенки клеток. Полученные данные автор ния семянки облитерируется, в связи с чем осо- использовал для ревизии р. Chaetanthera с двумя бенности ее строения не используются в систе- подродами и восстановил р. Oriastrum с двумя матике. Для зрелых семянок некоторых таксонов подродами. теста описана в виде гомогенной пленки (Reese, Данные о скульптуре поверхности семени 1989; Galastri, Oliveira, 2010). Поэтому указания единичны (Robinson, 1981; Tegel, 2002). В насто- на строение семенной кожуры видов семейства ящее время в систематике сложноцветных всё Asteraceae в литературе приводятся в основном большее значение приобретают комплексные при изучении анатомического строения семянок исследования структуры семянок с использова- видов триб Cardueae (Dittrich, 1970, 1977; Häff- нием светового и сканирующего микроскопов, ner, 2000), Cichorieae (Tegel, 2002) и Mutisieae которые позволяют получить более подробные (Grau, 1980; Davies, 2010), у которых она хорошо сведения об особенностях структуры и ультра- выражена и имеет разнообразное строение. скульптуры поверхности семянок для система- В трибе Cardueae особенности строения экзо- тики сем. Asteraceae. тесты имеют первостепенное значение и широко Целью данного исследования явилось изуче- применяются для таксономических построений ние семян (скульптура поверхности и анатоми- (Knorring, Tamamshyan, 1953; Karawya et al., ческое строение) видов базальных триб семей- 1974; Tscherneva, Surukhina, 1979). ства Asteraceae (Arctotideae, Cardueae, Mutisieae, F. Tegel (2002), исследуя семенную кожуру Vernonieae) для пополнения данных о строении 300 видов из 90 родов Lactuceae и 18 видов из семенной кожуры плодов сложноцветных и вы- других триб (Liabeae, Arctoteae, Vernonieae), по- явления признаков, которые можно использовать 46 Бойко Э. В., Новожилова Е. В. Строение семенной кожуры видов Asteraceae в таксономии и филогенетических построениях бритвы. Срезы окрашивали сафранином, с по- семейства. следующим заключением в глицерин-желатин. Фотографии препаратов выполняли с помощью Материал и методы микроскопов Leica 4500 (Leica) и Axio Imager.Z2 Материалом для карпологического исследо- (Carl Zeiss). вания послужили семянки, собранные авторами. Для изучения внешней поверхности семени Также использованы материалы Гербария лабо- семянку раскалывали лезвием бритвы на отдель- ратории хемотаксономии ФГБУН Тихоокеанско- ные элементы. Образцы (поверхность семянки, го института биоорганической химии им. Г. Б. сколы семянок, обратная сторона перикарпия, Елякова ДВО РАН (далее ТИБОХ). Некоторые поверхность и сколы семенной кожуры) напыля- образцы взяты из Гербариев Ботанического ин- ли хромом в вакууме с помощью Quorum tech- ститута им. В. Л. Комарова РАН (LE, г. Санкт- nology Q150T ES. Строение и микроскульптуру Петербург), Главного ботанического сада им. поверхности изучали по препаратам участков Н. В. Цицина РАН (MHA, г. Москва) и получены средней части семянок в трехкратной повтор- по “Delectus seminum” из ботанических садов ности c помощью сканирующего микроскопа мира. В данной работе мы придерживаемся си- EVO 40 XVP (Carl Zeiss) в центре коллективного стемы семейства Asteraceae, принятой в сводке пользования «Национального научного центра “The Families and Genera of Vascular Plants” (Ka- морской
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