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Светоизолирующий Аппарат Камерного Глаза Наземного Брюхоногого Моллюска Arion Rufus (Heterobranchia, Stylommatophora)

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Светоизолирующий Аппарат Камерного Глаза Наземного Брюхоногого Моллюска Arion Rufus (Heterobranchia, Stylommatophora) Ruthenica, 2021, vol. 31, No. 1: 39-49. © Ruthenica, 2021 Published online January 2, 2021. http: ruthenica.net Светоизолирующий аппарат камерного глаза наземного брюхоногого моллюска Arion rufus (Heterobranchia, Stylommatophora) И.П. ШЕПЕЛЕВА Федеральное государственное бюджетное учреждение науки Институт физиологии им. И.П. Павлова РАН, наб. Макарова, 6, Санкт-Петербург, 199034, РОССИЯ. Международный университет Бремена, Кампус, 1, Бремен, 28759, ГЕРМАНИЯ. Университет Лунда, Хельгонаваген, 3, Лунд, 22362, ШВЕЦИЯ. E-mail: [email protected] РЕЗЮМЕ. При помощи методов световой и электронной микроскопии определен клеточный состав, изучена морфология компонентов, а также оценены экранирующая функция и эволюционные из- менения светоизолирующего аппарата камерного глаза наземного брюхоногого легочного моллюска Arion rufus. По всем рассмотренным характеристикам светоизолирующий аппарат A. rufus является типичным для наземных легочных моллюсков. Эволюционные изменения светоизолирующего аппа- рата, обусловленные сменой моллюсками среды обитания, несущественны и затрагивают клеточный состав, морфологию пигментных клеток и изоляцию светочувствительных частей фоторецепторных клеток друг от друга. Light-insulating apparatus of the camera-like рованных пигментных клетках, расположенных eye of the terrestrial gastropod mollusk Arion rufus по соседству с фоторецепторными клетками, у (Heterobranchia, Stylommatophora) других присутствуют и в самих фоторецептор- ных клетках [Bharti et al., 2006; Land, 1981; Land, I.P. SHEPELEVA Nilsson, 2012]. Пигментные клетки на протяже- Pavlov Institute of Physiology, Russian Academy of Sciences, нии всей эволюции глаз животных помогают Makarova emb. 6, St. Petersburg, 199034, RUSSIA. выполнять те функции, к которым приспособлен International University Bremen, Campus Ring 1, Bremen, каждый тип глаз [D’Alba, Shawkey, 2019]. На- 28759, GERMANY. Lund University, Helgonavägen 3, пример, оба упомянутых выше типа глаз – пиг- Lund, 22362, SWEDEN. E-mail: [email protected] ментный бокал и камерный глаз встречаются у ABSTRACT. The cellular composition, the morphol- брюхоногих моллюсков – единственной группы ogy of the components, and the screening function and беспозвоночных с представителями в водной и evolutionary changes of the light-insulating apparatus наземной среде и доминирующим камерным гла- of the camera-like eye of the terrestrial gastropod pul- зом. У брюхоногих моллюсков пигментный бокал monate mollusk Arion rufus were studied and evalu- состоит из трех компонентов: фиксированного ated using light and electron microscopy. According to all the considered characteristics, the light-insulating зрачка большого размера; двухслойной инвер- apparatus of A. rufus is typical for terrestrial pulmo- тированной или неинвертированной сетчатки, nate mollusks. Evolutionary transformations of the которая может содержать до сотни фоторецеп- light-insulating apparatus, caused by the change in the торных клеток, распределенных по внутренней mollusks’ environment, are insignificant and affect the поверхности чаши из пигментных клеток, и cellular composition, the morphology of pigmented зрительного нерва. При помощи пигментных cells and the isolation of the light-sensitive parts of бокалов животные определяют местоположение photoreceptor cells from each other. источника света благодаря тому, что пигментные клетки создают экран, который обеспечивает Введение проникновение света в глаз только через зрачок и одновременно из разных направлений окружаю- В каждом существующем типе глаз – от пиг- щего пространства, а также выполняют формоо- ментных бокалов беспозвоночных до камерных бразующую, опорную, защитную и трофическую глаз позвоночных – обычно содержатся темные функции, функции синтеза и хранения экрани- экранирующие пигменты. У одних животных эти рующих пигментов [Beklemishev, 1952; Eakin, пигменты сосредоточены только в специализи- 1972; Land, 1981; Bobkova, 1998; Land, Nilsson, 40 И.П. Шепелева 2012]. Камерный глаз состоит из большего числа компонентов: оболочки, образованной рогови- цей и глазной капсулой; фиксированного зрачка меньшего размера; хрусталика; стекловидного тела, окружающего хрусталик; четырехслойной неинвертированной сетчатки, которая может насчитывать до нескольких десятков тысяч фоторецепторных клеток, расположенных между пигментными клетками, и зрительного нерва [Land, 1981; Messenger, 1981; Land, Nilsson, 2012; Shepeleva, 2013a, b, 2018a, b, 2019]. При помо- щи камерных глаз некоторые виды брюхоногих моллюсков различают предметы их естествен- ной среды обитания и геометрические фигуры, которые визуально напоминают эти предметы и, следовательно, используют зрительную ин- формацию для ориентации в окружающей среде [Messenger, 1981; Warrant, 2006; Shepeleva, 2011a, 2019]. Более сложное строение и функции камер- ных глаз по сравнению с таковыми пигментных бокалов указывают на больший набор функций у пигментных клеток. Дополнительно к перечис- ленным выше функциям в пигментных бокалах в РИС. 1. Схематичное изображение камерного глаза камерных глазах пигментные клетки участвуют Arion rufus. Сокращения: c – роговица; l – хрусталик, в формировании двух компонентов – хрусталика окруженный слоем стекловидного тела; mvl – микро- виллярный слой сетчатки; on – зрительный нерв; pgl и стекловидного тела [Eakin, 1972; Shepeleva, – пигментный слой сетчатки; spl – соматический и 2011b]. Эти неклеточные компоненты образуются плексиформный слои сетчатки; 1 – центральная об- из веществ, которые синтезируются в пигментных ласть зрительной части сетчатки; 2 – периферическая клетках, а потом секретируются и транспортиру- область зрительной части сетчатки; 3 – незрительная часть сетчатки. ются в полость глаз при помощи их отростков, как, например, у морских Caenogastropoda Tritia FIG. 1. Schematic drawing of the camera-like eye of Arion rufus. Abbreviations: c – cornea; l – lens, surrounded by obsoleta (Say, 1822) [Gibson, 1984] и Aporrhais a layer of the vitreous body; mvl – microvillar layer of pespelecani (Linnaeus, 1758) [Blumer, 1996], а так- the retina; on – optic nerve; pgl – pigmented layer of the же у наземных легочных моллюсков Helix pomatia retina; spl – somatic and plexiform layer of the retina; 1 – central region of the optic part of the retina; 2 – peripheral (Linnaeus, 1758) [Röhlich, Török, 1963], Cornu region of the optic part of the retina; 3 – non-optic part aspersum (Müller, 1774) [Eakin, Brandenburger, of the retina. 1967] и Limacus flavus (Linnaeus, 1758) [Kataoka, 1977]. Кроме того, в сетчатке камерных глаз пигментные клетки могут выполнять экраниру- ющую функцию, направленную на сохранении потенциальной разрешающей способности. Эта этих клеток необходимы для создания наиболее функция реализуется при помощи отростков полной характеристики светоизолирующего ап- пигментных клеток, заполненных гранулами парата и тем самым – камерных глаз брюхоногих экранирующего пигмента, которые окружают моллюсков. светочувствительную часть каждой фоторецеп- Цель работы – изучить светоизолирующий торной клетки и поглощают рассеянный тканями аппарат камерного глаза наземного брюхоногого глаза свет, предотвращая его распространение легочного моллюска Arion rufus (Linnaeus, 1758) между светочувствительными частями соседних (Arionidae). Экспериментальные задачи – опре- фоторецепторных клеток [Eakin, 1972; Shepeleva, делить клеточный состав светоизолирующего 2011b, 2017, 2018c; Dontsov, Ostrovsky, 2019]. аппарата; охарактеризовать компоненты светои- Таким образом, данные литературы о камерных золирующего аппарата; оценить экранирующую глазах брюхоногих моллюсков показывают, что функцию светоизолирующего аппарата; оценить основные функции специализированных пиг- эволюционные изменения светоизолирующего ментных клеток, которые в совокупности обра- аппарата, обусловленные сменой среды обитания. зуют светоизолирующий аппарат, известны. В то Для работы выбран A. rufus потому, что для этого же время морфологические свойства специали- вида моллюска автором впервые были получены зированных пигментных клеток описаны лишь данные об ультраструктуре сетчатки глаза, кото- поверхностно. Между тем, знания о морфологии рые позволяют дать необходимую характеристи- Светоизолирующий аппарат глаза Arion rufus 41 Табл. 1. Параметры пигментных клеток и пигментного слоя сетчатки камерного глаза Arion rufus. Table 1. Parameters of pigmented cells and pigmented layer of the retina of the camera-like eye of Arion rufus. Отношение параметров Зрительная часть Незри- в центральной в перифериче- в центральной Параметры тельная области и не- ской области и перифе- и перифериче- централь- часть зрительной незрительной рическая ской области ная область части части область Диаметр апикальной 2,6±0,2 2,9±0,2 4,1±0,2 поверхности тела 0,9 0,6 0,7 (n=15) (n=15) (n=10) клетки, (мкм) 7,0±0,8 6,0±0,6 5,2±0,4 Число отростков 1,2 1,4 1,2 (n=15) (n=15) (n=10) Диаметр отростков, 0,1±0,05 0,1±0,05 0,2±0,07 1,0 0,5 0,5 (мкм) (n=20) (n=28) (n=12) Длина отростков, 1,3±0,07 1,3±0,07 1,2±0,07 1,0 1,1 1,1 (мкм) (n=20) (n=28) (n=12) Число раздвоенных от- 2,0±0,5 1,5±0,8 1,1±0,3 1,3 2,0 1,5 ростков (n=15) (n=12) (n=8) Плотность расположе- 2,2±0,9 2,7±0,8 4,3±1,4 ния гранул экранирую- 0,8 0,5 0,6 (n=45) (n=45) (n=21) щего пигмента, (мкм-2) Толщина пигментного 73±3,3 63±2,7 38±4,0 1,2 1,9 1,7 слоя, (мкм) (n=10) (n=10) (n=10) n – число исследованных глаз (для толщины пигментного слоя), клеток (для диаметра апикальной поверх- ности тела клетки, числа отростков, числа раздвоенных отростков), отростков (для диаметра и длины отростков) или значений плотности расположения
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