invert5_2 173_180 Shepeleva.p65

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Çîîëîãèÿ áåñïîçâîíî÷íûõ, 2008, 5(2): 173–180 © ÇÎÎËÎÃÈß ÁÅÑÏÎÇÂÎÍÎ×ÍÛÕ, 2008
Ñòðîåíèå è îïòè÷åñêèå ñâîéñòâà ãëàçà íàçåìíîãî
áðþõîíîãîãî ìîëëþñêà [email protected]= (Linné, 1758)
(Pulmonata: Stylommatophora) èç Þæíîé Øâåöèè
È.Ï. Øåïåëåâà
Êàôåäðà áèîëîãèè êëåòêè è îðãàíèçìà, Ôàêóëüòåò íàóêè, Óíèâåðñèòåò ã. Ëóíäà, Õåëüãîíàâàãåí, 3, Ëóíä 22362 Øâåöèÿ.
e-mail: [email protected]
ÐÅÇÞÌÅ: Ñ èñïîëüçîâàíèåì ìåòîäîâ ñâåòîâîé ìèêðîñêîïèè èññëåäîâàíû ñòðóêòóðà è îïòè÷åñêèå ñâîéñòâà ãëàçà íàçåìíîãî áðþõîíîãîãî ìîëëþñêà Trichia hispida
(Linné, 1758) èç Þæíîé Øâåöèè. Ãëàçà ïîñòðîåíû ïî òèïó êàìåðíîãî ãëàçà ñ
ôèêñèðîâàííîé îïòèêîé. Îáíàðóæåíû ñóùåñòâåííûå îòëè÷èÿ â ñòðîåíèè è îïòè÷åñêèõ ñâîéñòâàõ ãëàçà T. hispida èç Þæíîé Øâåöèè â ñðàâíåíèè ñ ðàíåå èññëåäîâàííîé T. hispida èç Êàëèíèíãðàäñêîé îáëàñòè. Ãëàç T. hispida èç Þæíîé Øâåöèè
ïðèñïîñîáëåí ê çðåíèþ â äíåâíîå è ñóìåðå÷íîå âðåìÿ, òîãäà êàê ãëàç T. hispida èç
Êàëèíèíãðàäñêîé îáëàñòè ôóíêöèîíèðóåò òîëüêî êàê äåòåêòîð îñâåùåííîñòè.
ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: áðþõîíîãèå ìîëëþñêè, Trichia hispida, ñòðîåíèå ãëàçà, îïòè÷åñêèå ñâîéñòâà.
Structure and optical properties of the eye of terrestrial
gastropod snail [email protected]= (Linné, 1758) (Pulmonata:
Stylommatophora) from South Sweden
I.P. Shepeleva
Department of Cell and Organism Biology, Faculty of Science, University of Lund, Helgonavagen
3, S-22362 Lund, Sweden.
e-mail: [email protected]
ABSTRACT: Structure and optical properties of the eye of terrestrial gastropod snail
Trichia hispida (Linné, 1758) from South Sweden were investigated by light microscopy
methods. The structure of T. hispida eye is typical camera eye with fixed optics. Eye
structure and optical properties differences were revealed between the specimens T. hispida
from South Sweden and the specimens of T. hispida from Kaliningrad District previously
studied. The eye of T. hispida from South Sweden is adapted to the vision during daylight
and twilight hours, whereas the eye of T. hispida from Kaliningrad District functions as an
illumination detector only.
KEY WORDS: gastropod mollusks, Trichia hispida, eye structure, optical properties.
Printed in 2009
174
È.Ï. Øåïåëåâà
Ââåäåíèå
Áðþõîíîãèå ìîëëþñêè çàñåëèëè ìíîãèå
áèîòîïû îêåàíîâ, ìîðåé, ïðåñíûõ âîäîåìîâ è
ñóøè. Ñòðîåíèå èõ îðãàíîâ ÷óâñò⠗ íàãëÿäíûé ïðèìåð ýêñòåíñèâíîé àäàïòèâíîé ðàäèàöèè è ïëàñòè÷íîñòè (Seed, 1983; Audesirk,
Audesirk, 1985). Äàæå â ïðåäåëàõ ñåìåéñòâà è
ðîäà ýòè æèâîòíûå äåìîíñòðèðóþò çíà÷èòåëüíîå ðàçíîîáðàçèå ñòðîåíèÿ ïåðèôåðè÷åñêèõ
îòäåëîâ çðèòåëüíîé ñèñòåìû (Eakin, 1972;
Salvini-Plawen, Mayr, 1977; Hamilton, 1991;
Bobkova et al., 2004; Gal et al., 2004; Øåïåëåâà
2005à,á, 2006à,á,â, 2007). Ñðàâíèòåëüíîå èçó÷åíèå çðèòåëüíîé ñèñòåìû ýêîëîãè÷åñêè è ãåîãðàôè÷åñêè èçîëèðîâàííûõ ãðóïï áðþõîíîãèõ ìîëëþñêîâ ìîæåò ïîêàçàòü ñòåïåíü âëèÿíèÿ ôàêòîðîâ îêðóæàþùåé ñðåäû íà åå ðàçâèòèå â ôèëîãåíåçå, âûÿâèòü ôóíêöèîíàëüíîå çíà÷åíèå ýòîé ñèñòåìû äëÿ æèâîòíûõ è
îïðåäåëèòü âîçìîæíûé äèàïàçîí åå àäàïòèâíûõ èçìåíåíèé (Hamilton, Winter, 1984).
Trichia hispida (Linné, 1758) — ïðåäñòàâèòåëü ëåãî÷íûõ áðþõîíîãèõ ìîëëþñêîâ, øèðîêî ðàñïðîñòðàíåííûé â Ñåâåðíîé Åâðîïå, ïðè÷åì â ðàçíûõ ÷àñòÿõ àðåàëà ýêîëîãèÿ ýòîãî
âèäà ðàçëè÷àåòñÿ. Òàê, â Êàëèíèíãðàäñêîé
îáëàñòè T. hispida ïîñòîÿííî îáèòàåò â çàòåìíåííûõ ìåñòàõ ïîä êàìíÿìè, ïîâàëåííûìè
äåðåâüÿìè èëè îïàâøèìè ëèñòüÿìè è ïèòàåòñÿ ðàçðóøàþùèìèñÿ ðàñòèòåëüíûìè îñòàòêàìè (Øèëåéêî, 1978). Íà þãå Øâåöèè T.
hispida îáèòàåò â ëåñàõ íà äîñòàòî÷íî õîðîøî îñâåùåííûõ ó÷àñòêàõ íà íåâûñîêîé ìåëêîëèñòâåííîé ðàñòèòåëüíîñòè, êîòîðîé è ïèòàåòñÿ (ñîáñòâåííûå íàáëþäåíèÿ).  ðàíåå
îïóáëèêîâàííûõ ðàáîòàõ áûëè èçó÷åíû ñòðîåíèå è îïòè÷åñêèå ñâîéñòâà ãëàçà ó T. hispida,
îáèòàþùåé â Êàëèíèíãðàäñêîé îáëàñòè (Øåïåëåâà, 2002, 2005á; Bobkova et al., 2004; Gal
et al., 2004). Çàäà÷à íàñòîÿùåé ðàáîòû —
èññëåäîâàíèå ñòðîåíèÿ è îïòè÷åñêèõ ñâîéñòâ
ãëàçà T. hispida èç Þæíîé Øâåöèè.
Ìàòåðèàë è ìåòîäû
Èññëåäîâàíèÿ ïðîâîäèëè íà âçðîñëûõ
îñîáÿõ íàçåìíîãî áðþõîíîãîãî ëåãî÷íîãî
ìîëëþñêà Trichia hispida (Linné, 1758) (Pulmonata: Stylommatophora: Hygromiidae: Trichiinae) ñ ðàçìåðîì ðàêîâèíû 5–6´7–8 ìì.
Ìîëëþñêîâ ñîáèðàëè â ïðèãîðîäå ã. Ëóíäà —
Âîìáå (Øâåöèÿ) â èþíå–èþëå 2004 ã. Æèâîòíûõ ñîäåðæàëè â òåððàðèóìàõ ñ ïî÷âîé
ïðè òåìïåðàòóðå 18–20 °Ñ è åñòåñòâåííîì
ñâåòîâîì ðåæèìå.  êà÷åñòâå êîðìà èñïîëüçîâàëè ëèñòüÿ îäóâàí÷èêîâ.
Ïðåïàðàòû èçîëèðîâàííûõ ãëàç è õðóñòàëèêîâ ãîòîâèëè ïîä áèíîêóëÿðíûì ìèêðîñêîïîì Carl Zeiss ñ èñïîëüçîâàíèåì ôèçèîëîãè÷åñêîãî ðàñòâîðà (Takeuchi et al., 1996). Ôîêóñíîå ðàññòîÿíèå õðóñòàëèêîâ îöåíèâàëè ïî ðàçìåðàì ñîçäàâàåìûõ èçîáðàæåíèé (Nilsson et
al., 1988; Seyer, 1992; Øåïåëåâà, 2005á). Ïðèáëèçèòåëüíûé ïîêàçàòåëü ïðåëîìëåíèÿ õðóñòàëèêîâ, êîòîðûé ñîñòàâëÿåò 1,54, îïðåäåëÿëè ïî ôîðìóëå äëÿ ãîìîãåííîãî ñôåðè÷åñêîãî õðóñòàëèêà (Land, 1981). Èçîëèðîâàííûå ãëàçà, õðóñòàëèêè è ñîçäàâàåìûå èìè
èçîáðàæåíèÿ èçó÷àëè ïðè ïîìîùè ñâåòîâîãî ìèêðîñêîïà Zeiss Axiophot (îáúåêòèâ
25´0,80 ÂÈ). Öèôðîâûå èçîáðàæåíèÿ ïîëó÷àëè ïðè ïîìîùè êàìåðû Olympus DP 50.
Äëÿ ãèñòîëîãè÷åñêîãî èçó÷åíèå èñïîëüçîâàëè ãëàçíûå ñòåáåëüêè òåìíîàäàïòèðîâàííûõ ìîëëþñêîâ. Ãëàçíûå ñòåáåëüêè ôèêñèðîâàëè â 2,5%-íîì ðàñòâîðå ãëóòàðàëüäåãèäà íà 0,1 Ì êàêîäèëàòíîì áóôåðå (ðÍ=8,0)
â òå÷åíèå 2 ÷ ïðè 4 °Ñ. Ïîñëå îòìûâàíèÿ îò
ãëóòàðàëüäåãèäà â íåñêîëüêèõ ïîðöèÿõ áóôåðà ïðåïàðàòû ôèêñèðîâàëè â 1%-íîì ðàñòâîðå OsO4 íà 0,1 Ì êàêîäèëàòíîì áóôåðå
(ðÍ=8,0) â òå÷åíèå 1 ÷ ïðè 4 °Ñ. Îáåçâîæèâàíèå ïðîâîäèëè â ñåðèè ýòèëîâîãî ñïèðòà
âîçðàñòàþùåé êîíöåíòðàöèè è ïðîïèòûâàëè ýïîíîì-812. Èç ïîëó÷åííûõ áëîêîâ äåëàëè ñåðèéíûå ïîëóòîíêèå ñðåçû (2 ìêì) ïðè
ïîìîùè óëüòðàìèêðîòîìà V LKB 2088, êîòîðûå îêðàøèâàëè 0,5%-íûì ðàñòâîðîì òîëóèäèíîâîãî ñèíåãî ñ äîáàâëåíèåì 1%-íîãî
êàðáîíàòà íàòðèÿ è ïðîñìàòðèâàëè ïðè ïîìîùè ñâåòîâîãî ìèêðîñêîïà Zeiss Axiophot.
Öèôðîâûå èçîáðàæåíèÿ ïîëó÷àëè ïðè ïîìîùè êàìåðû Olympus DP 50.
Ìîðôîìåòðèþ âûïîëíÿëè íà öèôðîâûõ
èçîáðàæåíèÿõ ïðåïàðàòîâ è ñðåçîâ ïðè ïî-
Ñòðîåíèå è îïòè÷åñêèå ñâîéñòâà ãëàçà Trichia hispida
ìîùè êîìïüþòåðíîé ïðîãðàììû CorelDraw
10. Äëÿ ðàñ÷åòà ïàðàìåòðîâ ãëàçà èñïîëüçîâàëè ñëåäóþùèå ôîðìóëû.
Äëÿ ïëîòíîñòè ðàñïîëîæåíèÿ ôîòîðåöåïòîðîâ (H, ìêì–2) (Gal et al., 2004):
r=
2
3 ´ p2
,
(1)
ãäå p — àíàòîìè÷åñêîå ðàññòîÿíèå ìåæäó
öåíòðàìè ñîñåäíèõ ôîòîðåöåïòîðîâ.
Äëÿ îïòè÷åñêîé ÷óâñòâèòåëüíîñòè ãëàçà ê áåëîìó ñâåòó (SÁ, ìêì2·ñð–1) (Warrant,
Nilsson, 1998):
p
d
kl
,
(2)
SÁ = ( )2 ´ A2 ´ ( )2 ´
4
(2,3 + kl)
f
ãäå À — äèàìåòð àïåðòóðû, d — äèàìåòð
ôîòîðåöåïòîðà, f — ôîêóñíîå ðàññòîÿíèå
îïòè÷åñêîé ñèñòåìû, k — êîýôôèöèåíò ýêñòèíêöèè ôîòîïèãìåíòà ó ðàêîîáðàçíûõ in
situ, ðàâíûé 0,0067 ìêì–1 è l — äëèíà ôîòîðåöåïòîðà, ñîîòâåòñòâóþùàÿ òîëùèíå ìèêðîâèëëÿðíîãî ñëîÿ.
Äëÿ ðàçðåøàåìîãî óãëîâîãî ðàññòîÿíèÿ
ôîòîðåöåïòîðîâ ( Dj, ãðàä) (Land, 1981):
Dj =
p
,
f
(3)
ãäå p — àíàòîìè÷åñêîå ðàññòîÿíèå ìåæäó
öåíòðàìè ñîñåäíèõ ôîòîðåöåïòîðîâ è f —
ôîêóñíîå ðàññòîÿíèå îïòè÷åñêîé ñèñòåìû.
Äëÿ ðàçðåøàþùåé ñïîñîáíîñòè ãëàçà ïðè
ãåêñàãîíàëüíîì ðàñïîëîæåíèè ôîòîðåöåïòîðîâ (R, ðàä–1) (Land, 1981):
R=
1
3 ´ Dj
,
(4)
ãäå Dj — ðàçðåøàåìîå óãëîâîå ðàññòîÿíèå
ôîòîðåöåïòîðîâ.
Äëÿ F-÷èñëà (Land, 1981):
f
(5)
F= ,
A
ãäå f — ôîêóñíîå ðàññòîÿíèå îïòè÷åñêîé
ñèñòåìû è À — äèàìåòð àïåðòóðû.
Äëÿ îòíîñèòåëüíîé àïåðòóðû ãëàçà
(Land, 1981):
A
,
(6)
f
ãäå À — äèàìåòð àïåðòóðû è f — ôîêóñíîå
ðàññòîÿíèå îïòè÷åñêîé ñèñòåìû.
Äëÿ äèàìåòðà äèñêà Ýéðè (DÝ, ìêì)
(Land, 1981), îáðàçóåìîãî íà ñåò÷àòêå òî-
175
÷å÷íûì èñòî÷íèêîì ñâåòà ïðè l=500 íì (çíà÷åíèå, áëèçêîå ê ìàêñèìóìàì ÷óâñòâèòåëüíîñòè ôîòîðåöåïòîðíûõ êëåòîê ó áîëüøèíñòâà èññëåäîâàííûõ âèäîâ áðþõîíîãèõ ìîëëþñêîâ: Æóêîâ, Ãðèáàêèí, 1990; ×åðíîðèçîâ è äð., 1992; Dennis, 1967; Gillary, Wolbarsht 1967; Hughes, 1970; von Berg, Schneider,
1972; Suzuki et al., 1979; Katagiri et al., 1985):
ælö
DÝ = 2,44 ´ f ´ ç ÷ ,
(7)
è Aø
ãäå f — ôîêóñíîå ðàññòîÿíèå îïòè÷åñêîé
ñèñòåìû è À — äèàìåòð àïåðòóðû.
Äëÿ âñåõ ðàñ÷åòîâ èñïîëüçîâàëè ñðåäíèå
çíà÷åíèÿ ïîëó÷åííûõ äàííûõ.  òàáëèöàõ
ýòè çíà÷åíèÿ ïðèâåäåíû ñî ñòàíäàðòíûì
îòêëîíåíèåì (ñì. Ëàêèí, 1990).
Ðåçóëüòàòû
Ãëàçà Trichia hispida ïîñòðîåíû ïî òèïó
êàìåðíîãî ãëàçà ñ ôèêñèðîâàííîé îïòèêîé
(Ðèñ. 1À, 2À). Äèîïòðè÷åñêèé àïïàðàò ãëàçà
ñîñòîèò èç äâóõ ëèíç. Ïåðâàÿ ëèíçà ïðåäñòàâëåíà ïðèëåæàùèìè äðóã ê äðóãó ùóïàëüöåâûì ýïèäåðìèñîì è ðîãîâèöåé. Îíè
îáðàçîâàíû îäíîñëîéíûìè íåïèãìåíòèðîâàííûìè ñòîëá÷àòûìè ýïèòåëèÿìè. Âòîðàÿ
ëèíçà — íåêëåòî÷íûé õðóñòàëèê, êîòîðûé
èìååò ñðàâíèòåëüíî ìÿãêóþ êîíñèñòåíöèþ
è ýëëèïñîèäíóþ ôîðìó ïåðïåíäèêóëÿðíî
îïòè÷åñêîé îñè. Âäîëü êîðîòêîé îñè õðóñòàëèê ðàçäåëåí íà äâå çîíû — öåíòðàëüíóþ è
ïåðèôåðè÷åñêóþ.  öåíòðàëüíîé çîíå èìåþòñÿ íåìíîãî÷èñëåííûå ðàññåÿííûå âåçèêóëû èëè ìíîæåñòâî ïëîòíî ðàñïîëîæåííûõ âåçèêóë ýëëèïñîèäíîé ôîðìû ðàçìåðîì îò ìåíåå 1 äî 2,5´5 ìêì (Ðèñ. 2B). Â
îäíîì è òîì æå õðóñòàëèêå ïðèñóòñòâóþò
êàê òåìíûå, òàê è ñâåòëûå âåçèêóëû. Ïî
âíóòðåííåé ñòðóêòóðå âåçèêóëû ãîìîãåííû.
Ïåðèôåðè÷åñêàÿ çîíà õðóñòàëèêîâ èìååò
ñëîèñòîñòü. Èçîëèðîâàííûå õðóñòàëèêè ôîðìèðóþò äîñòàòî÷íî îò÷åòëèâûå èçîáðàæåíèÿ òåñòîâîãî îáúåêòà (Ðèñ. 1C, D).
Îò ìèêðîâèëëÿðíîãî ñëîÿ ñåò÷àòêè õðóñòàëèê îòäåëåí òîíêèì ñëîåì ïðîçðà÷íîãî
ñòóäåíèñòîãî ñòåêëîâèäíîãî òåëà. Äîìèíèðóþùèìè êëåòêàìè ñåò÷àòêè ÿâëÿþòñÿ ìèê-
176
È.Ï. Øåïåëåâà
Ðèñ. 1. Öèôðîâûå èçîáðàæåíèÿ äëÿ T. hispida èç Þæíîé Øâåöèè.
A — èçîëèðîâàííûé ãëàç; B — èçîëèðîâàííûé õðóñòàëèê; C — òåñòîâûé îáúåêò — ìåäíàÿ ñåòêà äëÿ
ýëåêòðîííîãî ìèêðîñêîïà ñ äëèíîé ñòîðîíû ÿ÷åéêè 88 ìêì; D — èçîáðàæåíèå òåñòîâîãî îáúåêòà, ñôîðìèðîâàííîå õðóñòàëèêîì. pa — îáëàñòü çðà÷êà, na — îáëàñòü íåéðîïèëÿ. Ìàñøòàá: 50 ìêì (À, C) è 10 ìêì (B, D).
Fig. 1. Digital pictures for T. hispida from South Sweden.
A — isolated eye; B — isolated lens; C — test object, a copper grid for electron microscope with square hole of 88-mm
side length; D — the image of the test object formed by the isolated lens. pa — pupil area; na — neuropil area. Scale
bars: 50 mm (À, C) and 10 mm (B, D).
ðîâèëëÿðíûå ôîòîðåöåïòîðû è ïèãìåíòíûå
êëåòêè, êîòîðûå èçîëèðóþò ðåöåïòîðíûå
êëåòêè äðóã îò äðóãà íà óðîâíå êëåòî÷íûõ
òåë (Ðèñ. 2À). Íàñêîëüêî ìîæíî ñóäèòü ïî
ïîëóòîíêèì ñðåçàì, ìèêðîâèëëû îïòè÷åñêè
íå èçîëèðîâàíû äðóã îò äðóãà.
Àíàòîìè÷åñêèå è çðèòåëüíûå ïàðàìåòðû
ãëàçà Trichia hispida èç Þæíîé Øâåöèè,
ðàññ÷èòàííûå ïî ôîðìóëàì (1) – (7), ïðèâåäåíû â Òàáë. 1–3.
Îáñóæäåíèå
Ñóäÿ ïî ðàñ÷åòíûì äàííûì, ïðèâåäåííûì â Òàáë. 1–3, ãëàç T. hispida ñïîñîáåí
ôîðìèðîâàòü îò÷åòëèâîå èçîáðàæåíèå â ïðåäåëàõ ìèêðîâèëëÿðíîãî ñëîÿ ôîòîðåöåïòîðîâ (Gal et al., 2004; Øåïåëåâà, 2006à). Ðàññ÷èòàííàÿ ïîòåíöèàëüíàÿ ðàçðåøàþùàÿ ñïîñîáíîñòü ôîòîðåöåïòîðîâ (4,0 ðàä–1) èäåíòè÷íà ðàçðåøàþùåé ñïîñîáíîñòè ôîòîðåöåïòîðîâ íàçåìíîãî ëåãî÷íîãî ìîëëþñêà
Ñòðîåíèå è îïòè÷åñêèå ñâîéñòâà ãëàçà Trichia hispida
177
Ðèñ. 2. Öèôðîâûå èçîáðàæåíèÿ ïðîäîëüíûõ ïîëóòîíêèõ ñðåçîâ ãëàç T. hispida.
A — èç Þæíîé Øâåöèè; B — èç Êàëèíèíãðàäñêîé îáëàñòè (ïî Bobkova et al., 2004). te — ùóïàëüöåâûé
ýïèäåðìèñ, c — ðîãîâèöà, l — õðóñòàëèê, mv — ìèêðîâèëëÿðíûé ñëîé ñåò÷àòêè, pg — ïèãìåíòíûé ñëîé
ñåò÷àòêè. Ìàñøòàá 50 ìêì (À, Â).
Fig. 2. Digital pictures of longitudinal semithin sections of T. hispida eyes.
A —from South Sweden; B — from Kaliningrad District (after Bobkova et al., 2004). te — tentacular epidermis, c — cornea,
l — lens, mv — microvillar layer of the retina, pg — pigmented layer of the retina. Scale bars: 50 mm (A, B).
Cepaea nemoralis (Linné, 1758) (4,0 ðàä–1)
(Gal et al., 2004). Îïòè÷åñêàÿ ÷óâñòâèòåëüíîñòü ôîòîðåöåïòîðîâ T. hispida — 1,8 ìêì2
·ñð–1 (ñì. Òàáë. 3) ñðàâíèìà ñ ÷óâñòâèòåëüíîñòüþ ôîòîðåöåïòîðîâ ïðåñíîâîäíûõ ëåãî÷íûõ ìîëëþñêîâ, íàñåëÿþùèõ íåãëóáîêèå
âîäîåìû: Planorbarius corneus (Linné,
1758) — 1,4 ìêì2·ñð–1, Lymnaea stagnalis
(Linné, 1758) — 1,8 ìêì2·ñð–1 è Physa fontinalis
(Linné, 1758) — 2,0 ìêì2·ñð–1 (Gal et al.,
2004). Íèçêîå çíà÷åíèå F-÷èñëà ãëàçà T. hispida, ðàâíîå 1,1 (ñì. Òàáë. 3), óêàçûâàåò íà
ïðèñóòñòâèå ñôåðè÷åñêîé àáåððàöèè, â òî
æå âðåìÿ, àáñîëþòíûé äèàìåòð àïåðòóðû,
ðàâíûé 90 ìêì (ñì. Òàáë. 1), ïîçâîëÿåò ïðåíåáðå÷ü õðîìàòè÷åñêîé àáåððàöèåé. Äèôðàêöèÿ ñâåòà íà àïåðòóðå ãëàçà, âåðîÿòíî, íå
îãðàíè÷èâàåò åãî ðàçðåøàþùóþ ñïîñîáíîñòü áëàãîäàðÿ íåáîëüøîìó äèàìåòðó äèñêîâ Ýéðè (1,3 ìêì) ïî ñðàâíåíèþ ñ ìåæðåöåïòîðíûìè ðàññòîÿíèÿìè — 14 ìêì (ñì.
Òàáë. 2). Îäíàêî îñòðîòà çðåíèÿ T. hispida
äîëæíà íåçíà÷èòåëüíî ñíèæàòüñÿ èç-çà
ðàññåèâàíèÿ ñâåòà íà ìèêðîâèëëàõ ñîñåäíèõ
ôîòîðåöåïòîðíûõ êëåòîê, ïîñêîëüêó ìåæäó
íèìè îòñóòñòâóåò ýêðàíèðóþùèé ïèãìåíò.
Äëÿ èäåíòèôèêàöèè îêðóæàþùèõ îáúåêòîâ, ðàçðåøàåìîå óãëîâîå ðàññòîÿíèå ôîòî-
ðåöåïòîðîâ ãëàçà íå äîëæíî ïðåâûøàòü íåñêîëüêî ãðàäóñîâ (Land, Fernald, 1992), à
îïòè÷åñêàÿ ÷óâñòâèòåëüíîñòü ãëàçà äîëæíà
ñîîòâåòñòâîâàòü ñâåòîâûì óñëîâèÿì ìåñò
îáèòàíèÿ (Warrant, McIntyre, 1993). Äíåâíûå æèâîòíûå èìåþò ÷óâñòâèòåëüíîñòü íèæå
åäèíèöû, ñóìåðå÷íûå — îò åäèíèöû äî ñòà
(Land, Nilsson, 2002). Îïòè÷åñêàÿ ÷óâñòâèòåëüíîñòü ãëàçà T. hispida èç Þæíîé Øâåöèè íåñêîëüêî ïðåâûøàåò 1 ìêì2·ñð–1 (ñì.
Òàáë. 3). Ýòî îçíà÷àåò, ÷òî îí, ñêîðåå âñåãî,
ïðèñïîñîáëåí ê çðåíèþ â óñëîâèÿõ îñâåùåííîñòè â äíåâíûå è ñóìåðå÷íûå ÷àñû.
Ðàíåå áûë èññëåäîâàí ãëàç T. hispida èç
Êàëèíèíãðàäñêîé îáëàñòè (Øåïåëåâà, 2002;
Bobkova et al., 2004; Gal et al., 2004; Øåïåëåâà, 2005á). Â ýòîì ðåãèîíå T. hispida îáèòàåò
â óñëîâèÿõ áîëåå íèçêîé îñâåùåííîñòè, è
ñîîòâåòñòâåííî ýòîìó, ãëàç ìîëëþñêà èìååò
÷óâñòâèòåëüíîñòü, ðàâíóþ 4 ìêì2·ñð–1 (ñì.
Òàáë. 3). Ó T. hispida èç Êàëèíèíãðàäñêîé
îáëàñòè ôîêóñ íå ñîâïàäàåò ñ ìèêðîâèëëÿðíûì ñëîåì ñåò÷àòêè (ñì. Òàáë. 3). Ýòî îçíà÷àåò, ÷òî ìîëëþñê èìååò íèçêóþ îñòðîòó
çðåíèÿ, è ãëàç ìîæåò ñëóæèòü òîëüêî äåòåêòîðîì îñâåùåííîñòè (Gal et al., 2004). Õîòÿ
ìîëëþñê èç Êàëèíèíãðàäñêîé îáëàñòè èìååò áîëüøèé ðàçìåð ãëàçíîãî áîêàëà è áîëåå
178
È.Ï. Øåïåëåâà
Òàáëèöà 1. Àíàòîìè÷åñêèå ïàðàìåòðû ãëàç T. hispida.
Table 1. Anatomical parameters of T. hispida eye.
Ïàðàìåòð (ìêì)
Ðàçìåð ãëàçíîãî áîêàëà
Äèàìåòð àïåðòóðû, A
Íàðóæíûéé ðàäèóñ êðèâèçíû ýïèòåëèÿ, r1
Òîëùèíà ýïèòåëèÿ
Âíóòðåííèéé ðàäèóñ êðèâèçíû ýïèòåëèÿ, r2
Íàðóæíûéé ðàäèóñ êðèâèçíû ðîãîâèöû, r1
Òîëùèíà ðîãîâèöû
Âíóòðåííèéé ðàäèóñ êðèâèçíû ðîãîâèöû, r2
é ïîâåðõíîñòüþ
Ðàññòîÿíèå ìåæäó íàðóæíîé
õðóñòàëèêà è âíóòðåííåéé ïîâåðõíîñòüþ
ðîãîâèöû
Ðàçìåð õðóñòàëèêà
Ðàäèóñ êðèâèçíû ïåðåäíåéé è çàäíåéé
ïîâåðõíîñòè õðóñòàëèêà
Ðàññòîÿíèå ìåæäó çàäíåéé ïîâåðõíîñòüþ
õðóñòàëèêà è âåðõíåéé ãðàíèöåéé
ìèêðîâèëëÿðíîãî ñëîÿ
T. hispida1
T. hispida2
130±4,0´190±4,0
142±9,0´220±8,0 (n=7)
90±5,0
96±5,0
7±2,0
89±2,0
89±2,0
8±2,0
81±3,0
92±3,0 (n=7)
137±7,0 (n=7)
9±0,4 (n=7)
128±7,0 (n=7)
128±7,0 (n=7)
13±2,0 (n=7)
49±5,0 (n=7)
2±0,1
–
63±5,0´96±3,0
81±3,0* (n=5)
69±5,0
63±6,0 (n=9)
2±0,1
–
T. hispida1 — ìîëëþñê èç Þæíîé Øâåöèè, T. hispida2 — ìîëëþñê èç Êàëèíèíãðàäñêîé îáëàñòè. Ñðåäíèå
çíà÷åíèÿ ïàðàìåòðîâ ðàññ÷èòàíû íà îñíîâå äàííûõ, ïîëó÷åííûõ îò 12 ãëàç äëÿ T. hispida1. Ïàðàìåòðû äëÿ T.
hispida2 âçÿòû èç Bobkova et al. (2004). * — ðàññòîÿíèå ìåæäó íàðóæíîé è âíóòðåííåé ïîâåðõíîñòüþ õðóñòàëèêà.
T. hispida1 — mollusk from South Sweden, T. hispida2 — mollusk from Kaliningrad District.
Average values of parameters were calculated on the basis of data taken from 12 eyes for T. hispida1. Parameters
for T. hispida2 were taken from Bobkova et al. (2004). * — distance between outer and inner lens surfaces.
Òàáëèöà 2. Õàðàêòåðèñòèêà ôîòîðåöåïòîðíûõ êëåòîê öåíòðàëüíîé îáëàñòè ñåò÷àòêè ãëàç T. hispida.
Table 2. Description of the photoreceptor cells from the central region of T. hispida retina.
Ïàðàìåòð
Äèàìåòð ôîòîðåöåïòîðà, d (ìêì)
Äëèíà ôîòîðåöåïòîðà, l (ìêì)
Ðàññòîÿíèå ìåæäó öåíòðàìè ñîñåäíèõ
ôîòîðåöåïòîðîâ, p (ìêì)
Ïëîòíîñòü ðàñïîëîæåíèÿ
ôîòîðåöåïòîðîâ, r (ìêì-2)
T. hispida1
10±2,0 (n=72)
12±2,0 (n=72)
T. hispida2
14±3,0 (n=25)
15±3,0 (n=25)
14±3,0 (n=72)
15±1,0 (n=25)
0,006
0,005
T. hispida1 — ìîëëþñê èç Þæíîé Øâåöèè, T. hispida2 — ìîëëþñê èç Êàëèíèíãðàäñêîé îáëàñòè. Ïàðàìåòðû
äëÿ T. hispida2 âçÿòû èç Bobkova et al. (2004) è Gal et al. (2004).
T. hispida1 — mollusk from South Sweden, T. hispida2 — mollusk from Kaliningrad District.
Parameters for T. hispida2 were taken from Bobkova et al. (2004) and Gal et al. (2004).
äëèííîå ôîêóñíîå ðàññòîÿíèå, îäíàêî ìåíüøàÿ ïëîòíîñòü ðàñïîëîæåíèÿ ôîòîðåöåïòîðîâ îçíà÷àåò ìåíüøóþ ðàçðåøàþùóþ ñïîñîáíîñòü. Áîëüøèé ðàçìåð ðåöåïòîðîâ äàåò
áîëåå âûñîêîå çíà÷åíèå îïòè÷åñêîé ÷óâñòâè-
òåëüíîñòè ãëàçà, îäíàêî ó ìîëëþñêà èç Êàëèíèíãðàäñêîé îáëàñòè îíà óâåëè÷åíà â
óùåðá îñòðîòå çðåíèÿ (Òàáë. 1–3). Ãëàç T.
hispida èç Êàëèíèíãðàäñêîé îáëàñòè (â îòëè÷èå îò T. hispida èç Þæíîé Øâåöèè) õàðàê-
Ñòðîåíèå è îïòè÷åñêèå ñâîéñòâà ãëàçà Trichia hispida
Òàáëèöà 3. Ðàññ÷èòàííûå çðèòåëüíûå
ïàðàìåòðû ãëàç T. hispida.
Table 3. Visual parameters of T. hispida eye calculated.
Ïàðàìåòð
fõ (ìêì)
fð+ý (ìêì)
fîáù (ìêì)
SÁ (ìêì2·ñð-1)
D? j (ãðàä)
R (ðàä-1)
F-÷èñëî
A/f
DÝ (ìêì)
T. hispida1
T. hispida2
118±0,1 (n=12)
381
96
1,8
8,4
4,0
1,1
0,9
1,3
122±19 (n=9)
–
113
,
4.0
13
2.5
,
1,2
0,8
1,5
T. hispida1 — ìîëëþñê èç Þæíîé Øâåöèè, T.
hispida2 — ìîëëþñê èç Êàëèíèíãðàäñêîé îáëàñòè. fõ —
ôîêóñíîå ðàññòîÿíèå õðóñòàëèêà, fð+ý — ôîêóñíîå
ðàññòîÿíèå ëèíçû, îáðàçîâàííîé ðîãîâèöåé è
ýïèòåëèåì, fîáù — ôîêóñíîå ðàññòîÿíèå îïòè÷åñêîé
ñèñòåìû ãëàçà, SÁ — îïòè÷åñêàÿ ÷óâñòâèòåëüíîñòü
ôîòîðåöåïòîðîâ â öåíòðàëüíîé ñåò÷àòêå, Dj —
ðàçðåøàåìîå óãëîâîå ðàññòîÿíèå ôîòîðåöåïòîðîâ â
öåíòðàëüíîé ñåò÷àòêå, R — ðàçðåøàþùàÿ ñïîñîáíîñòü
ôîòîðåöåïòîðîâ â öåíòðàëüíîé ñåò÷àòêå, A/f —
îòíîñèòåëüíàÿ àïåðòóðà ãëàçà, DÝ — äèàìåòð äèñêà
Ýéðè. Ïàðàìåòðû äëÿ T. hispida2 âçÿòû èç Gal et al.
(2004).
T. hispida1 — mollusk from South Sweden, T. hispida2 — mollusk from Kaliningrad District.
fl is the focal length of the lens, fc+e is the focal length
of the lens formed by cornea and epithelium, fcom is the
focal length of the optical system, Sw is the optical sensitivity of photoreceptors in the central retina, Dj is the
resolving angular distance of photoreceptors in the central
retina, R is the resolving ability of photoreceptors in the
central retina, DA is diameter of the Airy disc. Parameters
for T. hispida2 were taken from Gal et al. (2004).
òåðèçóåòñÿ ìåíüøèìè ðàçìåðàìè õðóñòàëèêà ïî ñðàâíåíèþ ñ ðàçìåðîì ãëàçíîãî áîêàëà, à ôîòîðåöåïòîðíûå êëåòêè èìåþò àïèêàëüíûå îòðîñòêè, õîðîøî çàìåòíûå íà ïîëóòîíêèõ ñðåçàõ (Ðèñ. 2B).
 íàñòîÿùåå âðåìÿ ìû âîçäåðæèâàåìñÿ
îò ñóæäåíèé î òîì, íàñêîëüêî âûÿâëåííûå
ðàçëè÷èÿ ìîãóò îòðàæàòü òàêñîíîìè÷åñêóþ
äèôôåðåíöèðîâêó ìåæäó ïîïóëÿöèÿìè T.
hispida èç Þæíîé Øâåöèè è Êàëèíèíãðàäñêîé îáëàñòè. Îòìåòèì òîëüêî, ÷òî ñòðóêòóðíûå ðàçëè÷èÿ ãëàç è èõ îïòè÷åñêèõ ñâîéñòâ
îáíàðóæèâàþòñÿ è ó äðóãèõ áëèçêîðîäñòâåííûõ áðþõîíîãèõ ìîëëþñêîâ. Ïåðèôåðè÷åñêèé îòäåë çðèòåëüíîé ñèñòåìû ëåãî÷íûõ
179
óëèòîê Cepaea hortensis (Müller,
ü
1774), îáèòàþùåé â çàðîñëÿõ êðàïèâû íà ïîëÿõ (Øåïåëåâà, 2006â), è C. nemoralis, æèâóùåé íà
êóñòàðíèêàõ è ñòâîëàõ äåðåâüåâ â ñàäàõ (Øåïåëåâà, 2002, 2005á; Bobkova et al., 2004; Gal
et al., 2004), ðàçëè÷àåòñÿ ôîðìîé ãëàçíîãî
áîêàëà è ïîëîæåíèåì õðóñòàëèêà îòíîñèòåëüíî îïòè÷åñêîé îñè. Ïðè ýòîì C. hortensis
ìîæåò îïðåäåëÿòü ëèøü èçìåíåíèÿ óðîâíÿ
îñâåùåííîñòè, à C. nemoralis, âåðîÿòíî, ìîæåò ðàçëè÷àòü îáúåêòû îêðóæàþùåé åå ñðåäû. Ìîðñêèå ïåðåäíåæàáåðíûå ìîëëþñêè
Littorina littorea (Linné, 1758) è L. irrorata
(Say, 1822) àêòèâíû äíåì, íî â ðàçíûõ óñëîâèÿõ: L. littorea, ãëàâíûì îáðàçîì, â âîäíîé,
à L. irrorata — â âîçäóøíîé ñðåäå. Ãëàçà ýòèõ
âèäîâ ïðèñïîñîáëåíû ê çðåíèþ â ñîîòâåòñòâóþùèõ ñðåäàõ, ÷òî ñêàçûâàåòñÿ íà îðèåíòàöèè ýëëèïñîèäíûõ õðóñòàëèêîâ îòíîñèòåëüíî îïòè÷åñêîé îñè (Newell, 1965; Hamilton et al., 1983; Seyer, 1992).
Áëàãîäàðíîñòè
Àâòîð áëàãîäàðèò ïðîôåññîðà Ý.Äæ. Âîððàíòà çà ïðåäîñòàâëåííóþ âîçìîæíîñòü ïðîâåäåíèÿ èññëåäîâàíèé íà êàôåäðå áèîëîãèè
êëåòêè è îðãàíèçìà óíèâåðñèòåòà ã. Ëóíäà
(Øâåöèÿ) è Ì. Ñîðåíññîíà çà óêàçàíèå ìåñò
îáèòàíèÿ ìîëëþñêîâ è îïðåäåëåíèå âèäà.
Ëèòåðàòóðà
Æóêîâ Â.Â., Ãðèáàêèí Ô.Ã. 1990. Ñïåêòðàëüíàÿ ÷óâñòâèòåëüíîñòü ãëàçà ìîëëþñêîâ Lymnaea stagnalis
è Planorbarius corneus â óëüòðàôèîëåòîâîé è âèäèìîé îáëàñòè ñïåêòðà // Ñåíñîðíûå ñèñòåìû. Ò.4.
¹ 4. Ñ.341–350.
Ëàêèí Ã.Ô. 1990. Áèîìåòðèÿ. Ì.: Âûñøàÿ øêîëà. 351 ñ.
×åðíîðèçîâ À.Ì., Øåõòåð Å.Ä., Àðàêåëîâ Ã.Ã., Çèìà÷åâ Ì.Ì. 1992. Çðåíèå âèíîãðàäíîé óëèòêè: ñïåêòðàëüíàÿ ÷óâñòâèòåëüíîñòü òåìíî-àäàïòèðîâàííîãî ãëàçà // Æóðíàë âûñøåé íåðâíîé äåÿòåëüíîñòè.
Ò.42. Ñ.1150–1155.
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