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Ультраструктура Сперматозоидов Четырех Видов Двустворчатых Моллюсков – Представителей Семейств Cardiidae И Astartidae Из Японского Моря С.А

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Ультраструктура Сперматозоидов Четырех Видов Двустворчатых Моллюсков – Представителей Семейств Cardiidae И Astartidae Из Японского Моря С.А Бюллетень Дальневосточного The Bulletin of the Russian малакологического общества Far East Malacological Society 2012, вып. 15/16, с. 176–182 2012, vol. 15/16, pp. 176–182 Ультраструктура сперматозоидов четырех видов двустворчатых моллюсков – представителей семейств Cardiidae и Astartidae из Японского моря С.А. Тюрин1, А.Л. Дроздов1,2 1Институт биологии моря им. А.В. Жирмунского ДВО РАН, Владивосток 690059, Россия 2Дальневосточный федеральный университет, Владивосток, 690950, Россия e-mail: [email protected] Изучена ультраструктура спермиев четырех видов двустворчатых моллюсков из семейств Cardiidae (Serripes groenlandicus, Clinocardium californiense, Clinocardium ciliatum) и Astartidae (Astarte borealis). Показано, что описанные спермии представляют собой классические акваспер- мии и состоят из головки, средней части и хвоста. Форма головки варьирует от конической изо- гнутой (сем. Cardiidae) до стержневидной (сем. Astartidae). Средняя часть имеет сходное строение и представлена четырьмя митохондриями, которые окружают перпендикулярно расположенные центриоли, от дистальной центриоли берет начало аксонема. Коническая изогнутая форма головки спермиев у представителей сем. Cardiidae подтверждает ранее опубликованные данные. Сравни- тельный анализ строения спермиев двустворчатых моллюсков свидетельствует о специфичности формы сперматозоидов для семейств. Ключевые слова: двустворчатые моллюски, Cardiidae, Astartidae, строение спермиев, Япон- ское море. Spermatozoa ultrastructure of four bivalve species of the families Cardiidae and Astartidae from the Sea of Japan S.A. Tyurin1, A.L. Drozdov1,2 1A.V. Zhirmunsky Institute of Marine Biology, Far East Branch, Russian Academy of Sciences, Vladivostok 690059, Russia 2Far Eastern Federal University, Vladivostok 690950, Russia e-mail: [email protected] Sperm ultrastructure of three bivalve mollusks of the family Cardiidae (Serripes groenlandicus, Clino- cardium californiense, Clinocardium ciliatum), and one species of the family Astartidae (Astarte borealis) from the Sea of Japan was examined. It is shown that studied spermatozoa are of «classical» aquasperm type, they consist of head, mid-piece and tale. Spermatozoa of the cardiid species are conical and slightly curved while ones of the astartid are rod-like. Mid-peace of all spermatozoa has similar structure: it com- prises four spherical mitochondria, surrounding two perpendicularly located centrioles, the distal centriole gives rise to axoneme. The described conical and curved shape of sperm head of the cardiid species is in agreement with previous studies. Comparative analysis of sperm morphology corroborates the idea that shape of sperm is specific for bivalve families. Key words: bivalve mollusks, Cardiidae, Astartidae, spermatozoa structure, Sea of Japan. 176 Мужские гаметы – сперматозои- десятков из нескольких тысяч извест- ды – неоднократно использовалось как ных науке видов. для разделения видов-двойников, так и В свете значимости знания строения для уточнения таксономического стату- спермиев для таксономии и филогении, са более высокого порядка [Аксенова, необходимо изучить ультраструктуру 1978; Дроздов, Иванков, 2000; Тюрин, гамет у гораздо большего количества Дроздов, 2003; Eckelbarger, Grassle, видов, чем это пока сделано. В настоя- 1987; Ferraguti et al., 1994; Hodgson, щей работе мы изучили ультраструкту- 1995; Pashchenko, Drozdov, 1998]. Од- ру спермиев 4 видов двустворчатых мол- нако необходимо отметить, что уль- люсков из семейств Cardiidae и Astartidae траструктура спермиев двустворчатых и сравнили их строение со спермиями моллюсков изучена лишь у нескольких некоторых ранее изученных видов. Материал и методика Исследованы сперматозоиды сле- мывали фильтрованной морской водой дующих видов моллюсков, принад- и дофиксировали в 4% растворе OsO4 в лежащих к подклассу Heterodonta, от- течении 30 мин. Фиксированный мате- ряду Veneroida, надсем. Cardioidea и риал обезвоживали в спиртах возрас- сем. Cardiidae: Clinocardium (Ciliatocar- тающей концентрации и заключали в dium) ciliatum (Fabricius, 1790), Clino- смесь эпон-аралдит. Предварительный cardium (Keenocardium) californiense просмотр материала осуществляли на (Deshayes, 1839), Serripes groenlandi- полутонких срезах, окрашенных мети- cus (Mohr, 1796), и надсем. Astartoidea леновым синим. Ультратонкие срезы и сем. Astartidae: Astarte borealis (Schu- получали на микротоме Ultracut фирмы macher, 1817). Классификация дана по Reichert. Для исследования выбирались статье: Lutaenko [2005]. только серебристые и золотистые срезы. Для ультраструктурного исследо- Контрастировали уранилацетатом и вания сперматозоидов кусочки семен- цитратом свинца. Просмотр и фото- ников фиксировали 2.5% раствором графирование срезов осуществляли в глутаральдегида на морской воде, про- электронном микроскопе JEOL-100S. Результаты Описанные спермии двустворчатых Serripes groenlandicus. Головка моллюсков представляют собой ти- спермиев коническая, слегка изогнутая пичные хвостатые спермии, адаптиро- (рис. 1А, 2A). Длина 4, ширина от 0.5– ванные для передвижения в воде. Они 1 мкм. Акросома коническая (рис. 1B). состоят из головки, средней части и Высота 0.8 мкм. Периакросомный ма- хвоста, представленного жгутиком. Го- териал содержит лишь электронносвет- ловка включает в себя акросому и ядро, лый гранулярный компонент, который средняя часть – митохондрии и цент- равномерно распределен по всему про- риолярный аппарат. странству под акросомным пузырьком. 177 Рис. 1. Срезы сперматозоидов. Serripes groenlandicus: A – продольный срез, B – акросома, C – мито- хондрии и центриоли; Сlinocardium californiense: D – продольный срез, E – акросома, F – митохон- дрии и центриоли; Clinocardium ciliatum: G – продольный срез, H – акросома, I – митохондрии и центриоли; Astarte borealis: J – продольный срез, K – митохондрии и центриоли. Условные обозна- чения: av – акросомная гранула, ce – центриоли, fl – жгутик, m – митохондрии, n – ядро. Масштаб A, C, D, F, G, H, I, J, K – 1 мкм; B, E – 0.5 мкм. Fig. 1. Sections of the spermatozoa. Serripes groenlandicus: A – longitudal section, B – acrosome, C – mitochondria and centrioles; Сlinocardium californiense: D – longitudal section, E – acrosome, F – mito- chondria and centrioles; Clinocardium ciliatum: G – longitudal section, H – acrosome, I – mitochondria and centrioles; Astarte borealis: J – longitudal section, K – mitochondria and centrioles. Abbreviations: av – acrosomal vesicle, ce – centrioles, fl – flagellum, n – nucleus, m – mitochondrion. Scale bar for A, C, D, F, G, H, I, J, K – 1 µm; for B, E – 0.5 µm. 178 Рис. 2. Схемы строения сперматозоидов: А – Serripes groenlandicus, B – Сlinocardium californiense, C – Clinocardium ciliatum, D – Astarte borealis. Обозначения см. в подписи к рис. 1. Масштаб 1 мкм. Fig. 2. Schemes of sperm: А – Serripes groenlandicus, B – Сlinocardium californiense, C – Clinocardium ciliatum, D – Astarte borealis. Abbreviations: see caption to Fig. 1. Scale bar 1 µm. В средней части сперматозоида лежат Длина 7, ширина от 1 в широкой части четыре сферические митохондрии диа- до 0.5 мкм в узкой. Акросома коническая метром 0.6–0.7 мкм (рис. 1C). Внутри (рис. 1E). Высота 1.5 мкм. Периакросом- кольца митохондрий лежат 2 взаимно ный материал содержит лишь электрон- перпендикулярные центриоли. Прок- носветлый гранулярный компонент, симальная центриоль соединяется с который равномерно распределен по небольшим углубление дистальной всему пространству под акросомным поверхности ядра жгутиковым кореш- пузырьком. В средней части сперматозо- ком. Дистальная центриоль окружена ида лежат четыре сферические митохон- сателлитным комплексом, которым она дрии диаметром 0.6–0.7 мкм (рис. 1F). прикреплена к мембране. От дисталь- Clinocardium ciliatum. Головка ной центриоли берет начало хвостовой спермиев вытянутая, коническая, изо- жгутик. гнутая (рис. 1G, 2C). Длина 8 мкм, ши- Clinocardium californiense. Головка рина от 0.8 до 0.4 мкм. Акросома вы- спермиев этого вида вытянутая, кони- тянутая, остро коническая (рис. 1H). ческая, слегка изогнутая (рис. 1D, 2B). Высота 3 мкм, периакросомный мате- 179 риал содержит лишь электронносвет- головку (рис. 1J, 2D), длиной 12, шириной лый гранулярный компонент. В сред- до 1.2 мкм. Акросома небольшая, остро ней части сперматозоида лежат четыре вытянутая, высотой до 1.7 мкм. Пери- сферические митохондрии диаметром акросомный материал имеет фибрил- 0.6–0.7 мкм (рис. 1I) и центриоли, рас- лярный компонент. В средней части положенные перпендикулярно. спермия лежат 4 митохондрии и цен- Astarte borealis. Спермии данного триоли (рис. 1K). Митохондрии сфери- вида имеют вытянутую, стержневидную ческие, диаметр 0.8–0.9 мкм. Обсуждение Моллюски – один из наиболее раз- Семейство Cardiidae – одно из нообразных и специализированных самых многочисленных семейств дву- типов животных, тем не менее, вопрос створчатых моллюсков мелководных, о происхождении сестринских групп преимущественно тепловодных морей, моллюсков не решен [Winnepenninckx имеющих широкое распространение в et al., 1996; Giribet et al., 2006]. Нет Мировом океане. Оно включает около устоявшихся представлений и о вза- 200 видов, которые очень разноо- имоотношении таксонов внутри типа бразны, в связи с чем семейство иногда моллюсков, и об их филогении. Всего разделяют на несколько самостоятель- в мире описано примерно 8 тыс. видов ных семейств. В одном только Арало- двустворчатых моллюсков, в Японском Понто-Каспийском регионе насчиты- море обитает около 400–500 видов. вается 43 современных вида кардиид, Спермии моллюсков демонстрируют 9 из которых принадлежат подсем. очень
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