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Invert4 2 139 149 Tolstenkov.PM6 Çîîëîãèÿ áåñïîçâîíî÷íûõ, 2007, 4(2): 139149 © ÇÎÎËÎÃÈß ÁÅÑÏÎÇÂÎÍÎ×ÍÛÕ, 2007 Îðãàíèçàöèÿ ìûøå÷íîé ñèñòåìû è ðàñïðåäåëåíèå NO-åðãè÷åñêèõ è ñåðîòîíèíåðãè÷åñêèõ ýëåìåíòîâ ó òðåìàòîä Allocreadium isoporum Looss, 1894 (Allocreadiidae) è Paramphistomum cervi Zeder, 1790 (Paramphistomatidae) Î.Î. Òîëñòåíêîâ1, Í.Á. Òåðåíèíà1, Í.Ì. Øàëàåâà2, Ò.Â. Ãàéâîðîíñêàÿ1 1 Öåíòð ïàðàçèòîëîãèè ôèëèàë Èíñòèòóòà ïðîáëåì ýêîëîãèè è ýâîëþöèè èìåíè À.Í. Ñåâåðöîâà ÐÀÍ, Ëåíèíñêèé ïðîñïåêò 33, 119071 Ìîñêâà, Ðîññèÿ. e-mail: [email protected] 2 Êàôåäðà çîîëîãèè áåñïîçâîíî÷íûõ Áèîëîãè÷åñêîãî ôàêóëüòåòà Ìîñêîâñêîãî Ãîñóäàð- ñòâåííîãî Óíèâåðñèòåòà, Âîðîáüåâû ãîðû, 119991 Ìîñêâà, Ðîññèÿ. ÐÅÇÞÌÅ: Ñ èñïîëüçîâàíèåì TRITC-ìå÷åííîãî ôàëëîèäèíà èçó÷åíà àíàòîìè÷åñêàÿ îðãàíèçàöèÿ ìûøå÷íîé ñèñòåìû ó òðåìàòîä Allocreadium isoporum è Paramphistomum cervi. Èññëåäîâàíî ðàñïðåäåëåíèå NO-åðãè÷åñêèõ è 5-ÍÒ-åðãè÷åñêèõ ýëåìåíòîâ â íåðâíîé ñèñòåìå è ìóñêóëàòóðå. Ñòðóêòóðû, ãåíåðèðóþùèå îêñèä àçîòà, è 5-ÍÒ- åðãè÷åñêèå ýëåìåíòû íàéäåíû â öåíòðàëüíîé è ïåðèôåðè÷åñêîé íåðâíîé ñèñòåìå îáîèõ âèäîâ. Íàëè÷èå NO-åðãè÷åñêèõ ñòðóêòóð è 5-ÍÒ-åðãè÷åñêèõ ýëåìåíòîâ â òîëùå ìóñêóëàòóðû ñâèäåòåëüñòâóåò îá ó÷àñòèè îêñèäà àçîòà è ñåðîòîíèíà â ðåãóëÿ- öèè ñîêðàòèòåëüíîé àêòèâíîñòè òðåìàòîä. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: òðåìàòîäû, Allocreadium isoporum, Paramphistomum cervi, ìóñêóëàòóðà, íåðâíàÿ ñèñòåìà, íåéðîìåäèàòîðû, îêñèä àçîòà, ñåðîòîíèí. The organization of the muscular system and the distribution of NO-ergic and serotoninergic elements of trematodes Allocreadium isoporum Looss, 1894 (Allocreadiidae) and Paramphistomum cervi Zeder, 1790 (Paramphistomatidae) O.O. Tolstenkov1, N.B. Terenina1, N.M. Shalaeva2, T.V. Gaivoronskaja1 1 Center of Parasitology of A.N. Severtsov Institute of Ecology and Evolution, Russian Academy of Science, Leninskiy prospect 33, 119071 Moscow,, Russia. e-mail: [email protected] 2 Department of Invertebrate Zoology, Biological Faculty, Moscow State University, Vorobievy Gori, 119899 Moscow, Russia ABSTRACT: TRITC-conjugated phalloidin staining was used to investigate the anatomical organization of the musculature of Allocreadium isoporum and Paramphistomum cervi. 140 Î.Î. Òîëñòåíêîâ, Í.Á. Òåðåíèíà, Í.Ì. Øàëàåâà, Ò.Â. Ãàéâîðîíñêàÿ The distribution of NO-ergic and 5-HT-ergic elements in the nervous system and the musculature was studied. Structures generating NO and 5-HT-ergic elements were observed in the central and peripheral nervous system of both species. The presence of NO-ergic and 5-HT-ergic nerve elements in the musculature testified the participation of Nitric oxide and serotonin in the regulation of the motility of trematodes. KEY WORDS: Trematoda, Allocreadium isoporum, Paramphistomum cervi, musculature, nervous system, neuromediators, nitric oxide, serotonin. Ââåäåíèå Ìîñêîâñêîé îáëàñòè. Ìàòåðèàë ôèêñèðîâà- ëè â 4%-íîì ïàðàôîðìàëüäåãèäå â 0,01 Ì Ïðèìåíåíèå ãèñòîõèìè÷åñêèõ è èììó- ôîñôàòíîì áóôåðíîì ðàñòâîðå (ðH 7,4) ïðè íîöèòîõèìè÷åñêèõ ìåòîäîâ ïîçâîëèëî âûÿ- 4°Ñ è çàòåì ñîõðàíÿëè â 10%-íîé ñàõàðîçå, âèòü â íåðâíîé ñèñòåìå ãåëüìèíòîâ õîëè- ïðèãîòîâëåííîé íà 0,01 Ì ôîñôàòíîì áóôå- íåðãè÷åñêèå, ñåðîòîíèíåðãè÷åñêèå, ïåïòè- ðå. ×àñòü ìàòåðèàëà áûëà èñïîëüçîâàíà äëÿ äåðãè÷åñêèå ýëåìåíòû (Øèøîâ, 1991; Barton ïðèãîòîâëåíèÿ êðèîñòàòíûõ ñðåçîâ òîëùè- et al., 1993; Biserova et al., 1996; Biserova et íîé 20 ìêì, äðóãóþ ÷àñòü èñïîëüçîâàëè äëÿ al., 2000; Gzubaj, Grabda-Kazubska, 1993; îêðàøèâàíèÿ öåëûõ ïàðàçèòîâ. Marks et al., 1995; Niewiadomska et al., 1996 è Îêðàñêó ìóñêóëàòóðû ïðîâîäèëè ñ èñ- äð.). Îòêðûòèå ôóíêöèè îêñèäà àçîòà êàê ïîëüçîâàíèåì TRITC (òåòðàìåòèëðîäàìèí àòèïè÷íîãî íåéðîíàëüíîãî ìåññåíäæåðà â èçîòèîöèàíàò) ìå÷åííîãî ôàëëîèäèíà (â íåðâíîé ñèñòåìå ìëåêîïèòàþùèõ (Garth- ðàçâåäåíèè 1:200) âî âëàæíîé êàìåðå â òå÷å- waite, 1991; Snyder, 1992) äàëî òîë÷îê ê èñ- íèå 1 ÷ â òåìíîòå ïðè òåìïåðàòóðå 4°C â ñëåäîâàíèþ ýòîãî âåùåñòâà íå òîëüêî ó ìëå- ñîîòâåòñòâèè ñ ìåòîäîì, îïèñàííûì Wahl- êîïèòàþùèõ, íî è ó äðóãèõ æèâîòíûõ, â òîì berg (1998). ÷èñëå è áåñïîçâîíî÷íûõ.  ïîñëåäíèå ãîäû Äëÿ èäåíòèôèêàöèè íåðâíûõ ñòðóêòóð, ïîÿâèëèñü äàííûå î íàëè÷èè ñòðóêòóð, ãåíå- ñîäåðæàùèõ ñèíòàçó îêñèäà àçîòà ôåð- ðèðóþùèõ îêñèä àçîòà, ó ãåëüìèíòîâ (Gus- ìåíò ñèíòåçà îêñèäà àçîòà (NO-ñèíòàçó), tafsson et al., 2001; Terenina, Gustafsson, 2002; èñïîëüçîâàëè ÍÀÄÔÍ (íèêîòèíàìèäàäåíèí- Terenina et al., 2006). äèíóêëåîòèäôîñôàò, âîññòàíîâëåííàÿ ôîð-  íàñòîÿùåé ðàáîòå ïðèâåäåíû ðåçóëü- ìà)-äèàôîðàçíûé (ÍÀÄÔÍ-ä) ãèñòîõèìè- òàòû èçó÷åíèÿ ìûøå÷íîé ñèñòåìû, à òàêæå ÷åñêèé ìåòîä. Ìåòîä îñíîâàí íà îïðåäåëå- äàííûå èììóíîöèòîõèìè÷åñêîãî è ãèñòîõè- íèè ôåðìåíòà ÍÀÄÔÍ-äèàôîðàçû, èçâåñò- ìè÷åñêîãî èññëåäîâàíèÿ ñåðîòîíèíåðãè÷åñ- íîãî êàê ìàðê¸ð NO-ñèíòàçû, ñîãëàñíî ñòàí- êèõ íåðâíûõ ñòðóêòóð è ñòðóêòóð, ãåíåðèðó- äàðòíîé ìåòîäèêå (Hope, Vincent, 1989). þùèõ îêñèä àçîòà (NO-åðãè÷åñêèõ, íèòðî- Ëîêàëèçàöèþ ñåðîòîíèíåðãè÷åñêèõ (5- êñèäåðãè÷åñêèõ), ó äâóõ âèäîâ òðåìàòîä: ÍÒ-åðãè÷åñêèõ) íåðâíûõ ñòðóêòóð îïðåäå- Allocreadium isoporum Looss, 1894 (ñåì. Allo- ëÿëè èììóíîöèòîõèìè÷åñêè â ñîîòâåòñòâèè creadiidae) è Paramphistomum cervi Zeder, ñ ìåòîäîì Coons et al. (1955). Îêðàøèâàíèå 1790 (ñåì. Paramphistomatidae). ïðîâîäèëè íà êðèîñòàòíûõ ñðåçàõ è òîòàëü- íûõ ïðåïàðàòàõ. Ñðåçû èíêóáèðîâàëè â ïåð- âè÷íîé àíòèñûâîðîòêå (Incstar, USA, â ðàç- Ìàòåðèàë è ìåòîäû âåäåíèè 1:500) 48 ÷ ïðè òåìïåðàòóðå 4°C; âî âòîðè÷íîé àíòèñûâîðîòêå (TRITC, DAKO, â Âçðîñëûå ôîðìû A. isoporum áûëè âçÿòû ðàçâåäåíèè 1:30) ïðåïàðàòû èíêóáèðîâàëè â èç êèøå÷íèêà óêëåè (Alburnus alburnus), òå÷åíèå 3 ÷. ïîéìàííîé â âîäîåìàõ Êàðåëèè, P. cervi Äëÿ ñîïîñòàâëåíèÿ ÍÀÄÔÍ-ä ñòðóêòóð, èç ðóáöà äîìàøíåé êîðîâû (Bos taurus) â ñòðóêòóð, ñîäåðæàùèõ ñåðîòîíèí, è ìûøå÷- Îðãàíèçàöèÿ ìûøå÷íîé ñèñòåìû ó òðåìàòîä 141 íûõ âîëîêîí áûëî ïðîâåäåíî òðîéíîå îêðà- öåâûìè âîëîêíàìè. Âåòâè êèøå÷íèêà â ðàé- øèâàíèå ñðåçîâ: ÍÀÄÔÍ-äèàôîðàçíûì ãè- îíå áèôóðêàöèè è äî èçãèáà â êàóäàëüíîì ñòîõèìè÷åñêèì ìåòîäîì, çàòåì èììóíîöè- íàïðàâëåíèè òàêæå èìåþò äâà ìûøå÷íûõ ñëîÿ òîõèìè÷åñêîå îêðàøèâàíèå íà ñåðîòîíèí è ìåíåå ðàçâèòûé íàðóæíûé ïðîäîëüíûé è îêðàñêà ìûøå÷íûõ âîëîêîí ñ èñïîëüçîâà- âíóòðåííèé êîëüöåâîé.  öåëîì, ìóñêóëàòó- íèåì TRITC-ìå÷åííîãî ôàëëîèäèíà. ðà êèøå÷íèêà ñòàíîâèòñÿ ìåíåå âûðàæåííîé Ïðåïàðàòû èññëåäîâàëè ñ ïîìîùüþ êîí- ïî ìåðå óäàëåíèÿ îò ìåñòà áèôóðêàöèè. ôîêàëüíîãî ñêàíèðóþùåãî ëàçåðíîãî ìèê-  ïîëîâîé ñèñòåìå A. isoporum ìûøå÷- ðîñêîïà Leica TCS 4D, ñîåäèíåííîãî ñ ôëó- íûé ñëîé íàèáîëåå ñèëüíî ðàçâèò â îáëàñòè îðåñöåíòíûì ìèêðîñêîïîì Leitz Aristoplan, åå äèñòàëüíûõ îòäåëîâ. Äèñòàëüíûé îòäåë à òàêæå ìèêðîñêîïà Axiostar plus, Zeiss. ìàòêè ìåòðàòåðì, à òàêæå äèñòàëüíûå îòäåëû ìóæñêîé ïîëîâîé ñèñòåìû èìåþò Ðåçóëüòàòû òîëñòóþ ìûøå÷íóþ ñòåíêó, ñîñòîÿùóþ èç õîðîøî ðàçâèòîãî êîëüöåâîãî ñëîÿ è ìåíåå Allocreadium isoporum. Ìóñêóëàòóðà ìîùíîãî ïðîäîëüíîãî ñëîÿ (ðèñ. 1D). Êîëü- ñòåíêè òåëà A. isoporum ïðåäñòàâëåíà òðåìÿ öåâûå è ïðîäîëüíûå ìûøå÷íûå âîëîêíà ñëîÿìè ìûøå÷íûõ âîëîêîí: íàðóæíûì êîëü- îêðóæàþò ïîëîâîå îòâåðñòèå, îòêðûâàþùå- öåâûì, ïðîìåæóòî÷íûì ïðîäîëüíûì, à òàê- åñÿ ìåäèàííî. Îîòèï èìååò óòîëùåííóþ æå ñëîåì âíóòðåííèõ äèàãîíàëüíûõ âîëî- ìûøå÷íóþ ñòåíêó, ñîñòîÿùóþ èç êîëüöå- êîí (ðèñ. 1À). Êîëüöåâûå âîëîêíà îðèåíòè- âûõ è ïðîäîëüíûõ ìûøå÷íûõ âîëîêîí. Ïðî- ðîâàíû â îñíîâíîì ïîä ïðÿìûì óãëîì ê êñèìàëüíûå îòäåëû ïîëîâîé ñèñòåìû èìå- ïðîäîëüíîé îñè òåëà. Îíè ÿâëÿþòñÿ ñàìûìè þò òîíêóþ ìûøå÷íóþ ñòåíêó, ñîñòîÿùóþ òîíêèìè è ìíîãî÷èñëåííûìè, ðàñïîëîæåíû ïðåèìóùåñòâåííî èç êîëüöåâûõ âîëîêîí. ÷àùå äðóãèõ. Ïðîäîëüíûå âîëîêíà èìåþò Ïðè èññëåäîâàíèè NO-åðãè÷åñêèõ ñòðóê- áîëüøóþ òîëùèíó, ÷åì êîëüöåâûå, è ðàñïî- òóð ó A. isoporum îáíàðóæåíî, ÷òî ÍÀÄÔÍ- ëîæåíû ðåæå. Òîëñòûå äèàãîíàëüíûå âîëîê- äèàôîðàçíàÿ ðåàêöèÿ âûÿâëÿåòñÿ ïî áîêàì íà ðàñïîëàãàþòñÿ ïàðàìè è ïåðåñåêàþòñÿ ãëîòêè, íà óðîâíå âåðõíåé åå ÷àñòè, ò.å. â ïîä óãëîì 120150°. îáëàñòè öåðåáðàëüíûõ ãàíãëèåâ (ðèñ. 1E). Ìîùíî ðàçâèòàÿ ìóñêóëàòóðà ðîòîâîé è NO-åðãè÷åñêèå âîëîêíà âèäíû â îòõîäÿùèõ áðþøíîé ïðèñîñîê ïðåäñòàâëåíà òðåìÿ òè- îò öåðåáðàëüíûõ ãàíãëèåâ ïðîäîëüíûõ íå- ïàìè ìûøå÷íûõ âîëîêîí: ðàäèàëüíûìè, ðâíûõ ñòâîëàõ, êîòîðûå ïðîñëåæèâàþòñÿ íà êîëüöåâûìè è ïðîäîëüíûìè. Êîëüöåâûå âî- ñðåçàõ äî óðîâíÿ áðþøíîé ïðèñîñêè.  äè- ëîêíà îáðàçóþò âíåøíèé ìûøå÷íûé ñëîé ñòàëüíîé ÷àñòè òåëà ïàðàçèòà ïîëîæèòåëü- ñòåíêè ïðèñîñêè, ïîä íèìè ðàñïîëàãàåòñÿ íîå îêðàøèâàíèå ïðîäîëüíûõ íåðâíûõ ñòâî- òîíêèé ñëîé ïðîäîëüíûõ âîëîêîí, ðàäèàëü- ëîâ íå âûÿâëåíî. Ñïåöèôè÷åñêè îêðàøåí- íûå âîëîêíà ïðîõîäÿò îò âíóòðåííåé ê âíåø- íûå âîëîêíà îòìå÷åíû ìåæäó ðàäèàëüíûìè íåé ñòåíêå ïðèñîñêè. Îò ñòåíêè òåëà ê áðþø- ìûøå÷íûìè âîëîêíàìè áðþøíîé è ðîòî- íîé ïðèñîñêå ñ ÷åòûðåõ ñòîðîí èäóò ïó÷êè âîé ïðèñîñîê â ñóáòåãóìåíòå. Ïîìèìî ýòî- ìûøå÷íûõ âîëîêîí (ðèñ. 1B). Ìóñêóëèñòàÿ ãî, ñðåäè ìûøå÷íûõ âîëîêîí ðîòîâîé è ãëîòêà èìååò øàðîîáðàçíóþ ôîðìó è âêëþ- áðþøíîé ïðèñîñîê, à òàêæå ãëîòêè îáíàðó- ÷àåò äâà ñëîÿ ìûøå÷íûõ âîëîêîí: õîðîøî æåíû íåñêîëüêî ðàäèàëüíî ðàñïîëîæåííûõ ðàçâèòûé ðàäèàëüíûé ñëîé è êîëüöåâîé ñëîé ñòðóêòóð, èìåþùèõ ïîëîæèòåëüíóþ ðåàê- (ðèñ. 1C). Íà ãðàíèöå ðîòîâîé ïðèñîñêè è öèþ íà ÍÀÄÔÍ-äèàôîðàçó (ðèñ. 1F). Ðåàê- ïðåôàðèíêñà èìååòñÿ ñóæåíèå òèïà ñôèíêòå- öèÿ íàáëþäàëàñü òàêæå â âîëîêíàõ, ðàñïî- ðà, îãðàíè÷åííîå êîëüöåâûìè âîëîêíàìè. ëîæåííûõ âáëèçè áðþøíîé ïðèñîñêè. Ïèùåâîä èìååò ðàçâèòûé ìûøå÷íûé ñëîé, Ñèëüíîå ïîëîæèòåëüíîå îêðàøèâàíèå íà ïðåäñòàâëåííûé õîðîøî âûðàæåííûìè íà- ÍÀÄÔÍ-äèàôîðàçó îáíàðóæåíî â äèñòàëü- ðóæíûìè ïðîäîëüíûìè è âíóòðåííèìè êîëü- íûõ è ïðîêñèìàëüíûõ îòäåëàõ ìóæñêîé è 142 Î.Î. Òîëñòåíêîâ, Í.Á. Òåðåíèíà, Í.Ì. Øàëàåâà, Ò.Â. Ãàéâîðîíñêàÿ Îðãàíèçàöèÿ ìûøå÷íîé ñèñòåìû ó òðåìàòîä 143 æåíñêîé ïîëîâîé ñèñòåìû: â ìûøå÷íîé ñòåí- òóðó è ñîñòîÿò èç òîíêîãî ñëîÿ ïðîäîëüíûõ êå ñåìÿèçâåðãàòåëüíîãî êàíàëà, à òàêæå â è êîëüöåâûõ âîëîêîí, à òàêæå ìíîãî÷èñëåí-
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