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Анатомия И Морфология Anatomy and Morphology 86 Turczaninowia 2013, 16 (3) : 86–95 АНАТОМИЯ И МОРФОЛОГИЯ ANATOMY AND MORPHOLOGY УДК 582.998 (581.471) DOI: http:/dx.doi.org/10.14258/turczaninowia.16.3.15 Э.В. Бойко E.V. Boyko ПАПИЛЛЫ НА ПОВЕРХНОСТИ СЕМЯНОК ВИДОВ ASTERACEAE PAPILLAE ON THE SURFACE OF THE SEEDS OF SPECIES OF ASTERACEAE Аннотация. Методом сканирующей электронной микроскопии (СЭМ) исследована скульптура по- верхности семянок 371 вида из 168 родов, 21 трибы, пяти подсемейств семейства Asteraceae. У 75 видов из 45 родов, относящихся к 8 трибам, а также у Symphyllocarpus exilis (вида с неустановленным таксономическим положением в системе семейства) было выявлено наличие папилл. На поверхности семянок большинства видов трибы Cichorieae находятся папиллы разнообразного строения. Только у видов этой трибы папиллы об- разуют гребни (виды подтрибы Hypochaeridinae) и на поверхности экзокарпия многих видов находятся окру- глые гладкие гранулы эпикутикулярного воска, отсутствующие у видов других триб. На поверхности семянок видов триб Anthemideae, Astereae, Athroismeae, Calenduleae, Gnaphalieae, Heliantheae, Senecioneae папиллы имеются только у некоторых видов. У исследованных нами видов триб Barnadesieae, Mutisieae, Cynareae, Vernonieae, Arctotideae, Inuleae, Helenieae, Coreopsideae, Tageteae, Madieae, Millerieae, Eupatorieae папиллы не обнаружены. Длина, форма папилл, их расположение на поверхности семянки различны у разных видов и могут быть использованы как таксономический признак. Ключевые слова: Asteraceae, семянки, папиллы, морфология, сканирующая электронная микроскопия (СЭМ), систематика. Summary. The sculpture of the surface of cypselae (scanning electron microscopy) of 371 species of 168 gene­ ra, 21 tribes, five subfamilies of the family Asteraceae was investigated. 75 species of 45 genera belonging to 8 tribes and the species Symphyllocarpus exilis with uncertain taxonomic position, papillae were detected. Papillae of diverse structure have been revealed on the surface of the cypselae of the majority of Cichorieae species. Only in species of this tribe papillae form crests (genera of subtribe Hypochaeridinae) and on the surface of exocarp of many species are rounded smooth granules epicuticular wax, missing in species of other tribes. On the cypselae surface of species of the tribes Anthemideae, Astereae, Athroismeae, Calenduleae, Gnaphalieae, Heliantheae, and Senecioneae, papil- lae are present only in some species. Papillae were not found in the investigated species of the tribes Barnadesieae, Mutisieae, Cynareae, Vernonieae, Arctotideae, Inuleae, Helenieae, Coreopsideae, Tageteae, Madieae, Millerieae, Eupatorieae. Length and shape of papillae, their location on cypselae surface vary greatly between species and can be used as taxonomic characters. Key words: Asteraceae, achenes, papillae, morphology, SEM, taxonomy. На поверхности семянок видов семейства ки, не отделенное от последней перегородкой. Asteraceae могут находятся волоски (простые и Многие семянки имеют выпуклые клетки экзо- железисты) и папиллы. Волосок – самостоятель- карпия, при этом высота оболочки увеличивает- ная клетка, образовавшаяся из клетки эпидермы ся постепенно и незначительно. Нередко возвы- и отделенная от неё клеточной стенкой. Волоски шение периклинальной стенки бывает резким и могут быть одноклеточными и многоклеточны- значительным, в результате этого формируется па- ми, простыми или железистыми. Папилла (со- пилла. В семействе Asteraceae папиллы находятся сочек) – это выпячивание эпидермальной клет- на семянках видов из различных триб. Важность Тихоокеанский институт биоорганической химии ДВО РАН (ТИБОХ ДВО РАН), пр-т 100 лет Владивостоку, 159; 690022, Владивосток, Россия; e-mail: [email protected] Pacific Institute of Bioorganic Chemistry, Far East Branch, Russian Academy of Sciences (PIBOC FEB RAS) 159 Pr-t 100 let Vladivostoku; 690022, Vladivostok, Russia Поступило в редакцию 19.08.2013 г. Submitted 19.08.2013 Принято к публикации 12.09.2013 г. Accepted 12.092013 Бойко Э.В. Папиллы на поверхности семянок видов Asteraceae 87 признаков строения трихом и папилл на поверх- coronopifolia L. № 1545. Сахалинская обл., о. Ку- ности семянок для филогении и классификации нашир, пос. Головнино, 28. 08. 1983, Басаргин семейства указывается в ряде работ (Алексан- Д., Баранов В. Tripleurospermum inodorum (L.) дров, Савченко, 1951; Carlquist, 1958; Jeffrey, Sch. Bip. № 2039. Амурская обл., г. Белогорск, Chen, 1984; Mukherjee, Nordenstam, 2012; Napp­ пустырь, 19. 07. 2002, Аистова Е. Ursinia anethoi- Zinn, Eble, 1980; Ramayya, 1962; Schneider, Bold- des (L.) Poir. № 2479. Г. Владивосток, в культуре, rini, 2011). 01. 09. 2012, Бойко Э. Данная работа является продолжением Триба Astereae Cass. – Aster alpinus L. исследования поверхности семянок сложноц- Амурская обл.: № 1322. Пос. Джалинда, 31. 08. ветных (Бойко, 2011, 2012). Цель работы – изу­ 85, Бойко Э.; № 1209. Пос. Игнашино, 02. 09. чение папилл на поверхности семянок с исполь- 1986, Волкова С.; № 2102. Чукотский Н.О., г. зованием сканирующей электронной микроскопии Анадырь, скалистые обнажения берега лимана, и выявление возможности использования получен- 15. 08. 1975. Здоровьева Е., Федореев С.А. A. ных данных для таксономических и филогенети- amellus L. № 1402. Чехословакия, 26. 09. 1958. ческих исследований в семействе Asteraceae. Heteropappus biennis (Ledeb.) Tamamsch. ex Grub. Материал и методы. Материалом для № 1247. Приморский край, Хасанский р­н, пос. исследования послужили зрелые плоды (семянки) Славянка, 03. 10. 1986, Здоровьева Е.; Амурская растений из семейства Asteraceae, собранные обл.: № 827. C. Мухино, 25. 08. 1969, Здоровьева автором и сотрудниками лаборатории хемотак- Е.; № 1342. Шимановский р­н, пос. Калашнико- сономии растений Тихоокеанского института во, устье р. Анго, 24. 09. 1975, Басаргин Д. H. биоорганической химии им. Г.Б. Елякова (ТИ- hispidus (Thunb.) Less. № 830. Г. Владивосток, БОХ ДВО РАН) в различных районах Дальнего Ботанический сад, 20. 09. 1981, Бойко Э.; Амурс­ Востока России. Некоторые образцы получены из кая обл.: № 1243. пос. Буссе, 04. 09. 1986, Бойко отечественных и зарубежных гербариев: Влади- Э.; № 1244. С. Томское, 12. 09. 1986, Бойко Э. востока (VLA), Ташкента (TASH), Новосибирска Olearia lirata (Sims) Hutch. № 2489. Amsterdam, (NS), Благовещенска (БГПУ), Кью (К). В данной 2013. Townsendia annua Beaman № 1401. Colo- работе мы придерживаемся системы, принятой в rado, USA, 07. 07. 1965, Salamun. Turczaninowia сводке под редакцией K. Kubitzki «The Families and fastigiata (Fisch.) DC. № 837. Амурская обл. с. Genera of Vascular Plants» (Kadereit, Jeffrey, 2007), Бузунда, 20. 08. 1969, Здоровьева Е.; № 2181. в которой семейство Asteraceae представлено 5 Приморский кр., Шкотовский р­н, пос. Речица, подсемействами. Нами исследована поверхность 12. 10. 2009, Бойко Э.В. семянок 371 вида из 168 родов и 21 трибы, пред- Триба Athroismeae Panero. – Centipeda ставляющих все пять подсемейств семейства. minima (L.) A. Br. et Aschers. № 1546. Хабаровс­ Изучение поверхности семянок прово- кий кр., с. Бичевая, в пойме р. Хор, 18. 08. 1965, дились на сканирующих электронных микроско- Бойко Э., Петроченкова А. пах LEO 430 и EVO 40 при увеличении от ×30 Триба Calenduleae Cass. – Calendula of- до ×3000. Объекты готовили по стандартной ме- ficinalisL. № 2323. Приморский кр., г. Владивос­ тодике: семянки наклеивали лаком на специаль- ток, 15. 08. 2002, Бойко Э. ные столики определенного типа. Столики с об- Триба Cichorieae Lam. et DC. – Askellia разцами напыляли золотом или смесью платины nana (Richards.) W.A. Weber (=Crepis nana Ri­ и палладия. Для сравнения семянки одного вида chards.) № 1847. Магаданская обл., Теньки́ нский брались из нескольких точек ареала; исследова- р­н, пос. Кулу, 03. 08. 1978. Бойко Э. Chorisis re- лось по 3–6 семянок каждого образца из разных pens (L.) DC. № 1956. Приморский кр., Хасанский частей корзинки. Расположение папилл на по- р­н, бухта Витязь, 30. 08. 1977, Бойко Э. Cicho- верхности неравномерное, поэтому исследова- rium intybus L. № 1278. Приморский кр., Пар- лись участки разных частей зрелых семянок. тизанский р­н, пос. Золотая Долина, 10. 08. 83, Приводим список исследованных видов, Бойко Э. Crepidiastrum denticulatum (Houtt.) J.H. на семянках которых обнаружены папиллы (но- Pak et Kawano (= Paraixeris denticulata (Houtt.) мер образца соответствует номеру в коллекции Nakai) № 1272. Приморский кр., Шкотовский семянок, которая хранится в лаборатории хемо- р­н, пос. Анисимовка, 21. 09. 1984, Бойко Э. C. таксономии растений ТИБОХ ДВО РАН). sonchifolium (Bunge) Pak et Kawano (= Paraixeris serotina (Maxim.) Tzvel.) № 1775. Окр. г. Благове- Триба Anthemideae Cass. – Cota austriaca щенска, 02. 09. 98. Старченко В. Crepis burejensis (Jacq.) Sch. Bip. № 2203. Stuttgart, 2010. Cotula Fr. Schmidt № 931. Хабаровский кр., Баджальс­ 88 Бойко Э.В. Папиллы на поверхности семянок видов Asteraceae кий хребет, 17. 08. 1973, Бойко Э., Шаповал И. ко Э.; № 1976. Хасанский р-н, п-ов Гамова, 01. C. chrysantha (Ledeb.) Turcz. № 582. Магаданс­ 10. 2003, Горовой П. Gnaphalium californicum кая обл., пос. Беринговский, бухта Угольная, 18. DC. № 2228. USA, Santa Barbara, у дороги, 02. 08. 1968, Горовой П. C. coreana (Nakai) Sennikov 07. 2009, Чернышев В. G. mandshuricum Kirp. (=H. coreanum Nakai) № 579. Хабаровский кр., № 2103. Приморский кр., Хасанский р­н, п­ов Баджальский хребет, 15. 08. 1973, Бойко Э. C. Краббе, 26. 07. 2008, Нечаев В. G. sylvaticum gmelinii (L.) Tausch № 2120. Магаданская обл., L. № 1043. Сахалинская обл., пос. Анива, бе- 2–4 км трассы Сусуман­Буркандья, 12. 08. 1980, рег залива, 18.
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