Morfologiczne przystosowania naczyn krwionosnych i limfatycznych wiezadla szerokiego macicy owcy do lokalnych regulacji czynnosci narzadu rozrodczego

• Autorzy: Zezula-Szpyra A
• Języki publikacji: PL

Abstrakty

PL

W pracy przedstawiono, opracowaną na poziomie LM, TEM i SEM, morfologię oraz wzajemne stosunki topograficzne naczyń krwionośnych i limfatycznych więzadła szerokiego macicy owcy i przeanalizowano ich przypuszczalne znaczenie w mechanizmie przeciwprądowego przenikania hormonów i innych czynników regulujących. W doświadczalnej części pracy starano się wykazać ewentualny lokalny wpływ hormonów jajnikowych na niektóre przystosowania czynnościowe naczyń związane z dopływem krwi do macicy i jajnika. Badania przeprowadzono na dorosłych owcach w okresie aktywności płciowej, sezonowego anestrus, a także w 7-8 tygodni po owariektomii. Owce cykliczne posłużyły do badań morfologicznych, w których wykorzystano: preparaty naczyniowe po wypełnieniu środkiem kontrastującym i mikrofilem, merkoksowe odlewy korozyjne naczyń i preparaty tkankowe analizowane w skaningowym mikroskopie elektronowym (SEM), preparaty histologiczne obserwowane w mikroskopie świetlnym, a także ultrastrukturę naczyń limfatycznych ocenianą w transmisyjnym mikroskopie elektronowym (ТЕМ). W materiale pobranym od owiec w cyklu rujowym oraz owariektomizowanych, które otrzymywały lub nie benzoesan estradiolu określano: zachowanie się komórek śródbłonka tętnic macicznej i krezkowej na podstawie preparatów tkankowych analizowanych w SEM, natomiast w badaniach histochemicznych określano aktywność NADPH-diaforazy (markera syntazy tlenku azotu) w śródbłonku naczyń krwionośnych i limfatycznych z obszaru więzadła szerokiego macicy. Makroskopowe obserwacje naczyń krwionośnych wykazały, że mimo istniejących różnic osobniczych, gałązki - jajowodowa i maciczna tętnicy jajnikowej mają kręty przebieg, chociaż nie tak intensywny, jak jej gałązka jajnikowa. Gałązki - jajowodowa i maciczna tętnicy jajnikowej układały się obok lub oplatały odpowiadające im naczynia żylne. Nie stwierdzono jednak, aby te naczynia tętnicze, zaopatrujące jajowód i przednią część rogu macicy, układały się w bliskim sąsiedztwie żył jajnikowych odprowadzających krew żylną bogatą w hormony jajnikowe. Zaobserwowano także po raz pierwszy, że gałązka jajnikowa tętnicy jajnikowej, przeważnie prosta tuż przed jajnikiem, w jego części rdzeniowej ulega bardzo intensywnej spiralizacji. Podobnie spiralny przebieg wykazywały także mniejsze rozgałęzienia odchodzące od spętlonej tętnicy, a kierujące się ku części korowej jajnika. Tętnica maciczna i jej gałązki w obrębie krezki macicznej wykazywały prosty przebieg. Naczynia żylne odprowadzające krew zarówno z macicy, jak i z jajnika były bardzo rozbudowa­ne i miały dobrze wykształcone zastawki. Mimo ich bliskiego kontaktu z naczyniami tętniczymi zaopatrującymi jajnik czy macicę nie zaobserwowano przypadków redukowania grubości ścian sąsiadujących ze sobą naczyń. Zaobserwowano, że odjajnikowe naczynia limfatyczne schodzące wzdłuż krezki jajnikowej, chociaż niezbyt liczne, rozdzielały się i układały nie tylko w pobliżu gałązki jajnikowej, ale także towarzyszyły gałązce jajowodowej i macicznej tętnicy jajnikowej. Niektóre z odjajnikowych naczyń limfatycznych, zwykle małe i cienkościenne, wchodziły pomię­dzy pętle spiralnej gałązki jajnikowej tętnicy jajnikowej. Odmaciczne naczynia limfatyczne tworzyły natomiast wyraźnie dwa pasma - jedno mniej rozbudowane, kierujące się ku tętnicy macicznej i drugie, prowadzące limfę z przyjajowodowej i środkowej części rogu macicy, które na wysokości jajnika wchodziło na teren krezki jajnikowej. Przeważnie gubościenne naczynia tego drugiego pasma układały się, od strony brzusznej krezki jajnikowej, pomiędzy jej warstwą mięśniową a naczyniami krwionośnymi. Ułożone wzdłuż krezki jajnikowej odmaciczne naczynia limfatyczne stopniowo łączyły się między sobą, a następnie z odjajnikowymi naczyniami limfatycznymi i tworzyły dwa lub trzy naczynia zbiorcze. Przed osiągnięciem przyaortowego węzła chłonnego naczynia limfatyczne układały się obok tętnicy jajnikowej i żyły maciczno-jajnikowej. Analiza odlewów korozyjnych naczyń w SEM wykazała, że większe naczynia tętnicze i żylne, a także odjajnikowe i odmaciczne naczynia limfatyczne otoczone były mikronaczyniowymi (naczynia naczyń) sieciami. Występowały jednak znaczne różnice dotyczące zarówno rozmieszczenia, jak i zagęszczenia tych sieci. W początkowym przebiegu tętnicy macicznej i jajnikowej występowały mniej rozbudowane sieci naczyń niż na ich mniejszych, już bardziej cienkościen­nych rozgałęzieniach. Najbardziej rozbudowane, wielowarstwowe sieci vasa vasorum obserwo­wano na krętej gałązce jajnikowej i macicznej tętnicy jajnikowej, a także na żyle jajnikowej. Gałązka jajowodowa tętnicy jajnikowej i gałązki odchodzące od tętnicy macicznej otaczały zwykle jednowarstwowe mikronaczyniowe sieci. Zaobserwowano, że powstające z vasa vasorum żyłki układały się w zewnętrznej warstwie przydanki lub poza przydanką naczyń tętniczych czy żylnych. Dobrze rozbudowane, chociaż zwykle jednowarstwowe sieci vasa vasorum otaczały także naczynia limfatyczne. Analiza w SEM wykazała tętniczo-żylny charakter mikronaczyniowych sieci otaczających naczynia limfatyczne. Uzyskane wyniki badań wskazują na istnienie u owcy kilku innych - niż to sugerowano dotychczas - morfologicznych przystosowań naczyń krwionośnych i limfatycznych do przeciwprądowego przenikania hormonów, a przypuszczalnie i innych związków regulujących w obsza­rze więzadła szerokiego macicy. Przeciwprądowemu przenikaniu i lokalnemu zaopatrzeniu macicy i jajowodu w hormony steroidowe wydaje się sprzyjać bliskie sąsiedztwo gałązek macicznej i jajowodowej z odjajnikowymi naczyniami limfatycznymi prowadzącymi bogatą w hormony jajnikowe limfę (Daniel i in. 1963). Przypuszczalnie odmaciczne naczynia limfatyczne przebiegające od strony brzusznej wzdłuż krezki jajnikowej są dla jajnika nie mniej obfitym źródłem PGF 2α niż towarzysząca tętnicy jajnikowej żyła maciczno-jajnikowa czy odprowadza­jące maciczno-jajnikową limfę, naczynia limfatyczne. Wydaje się także, że obserwowane przez innych morfologów (Del Campo, Ginther, 1973a, Lee, O'Shea 1976) i rozpatrywane jako ważne drogi dla przeciwprądowego przenikania różnych substancji, sploty żylne na gałązce jajnikowej tętnicy jajnikowej uczestniczą tylko pośrednio w mechanizmie przeciwprądowego przenikania u owcy. Obecność mikronaczyniowych sieci na naczyniach krwionośnych i limfatycznych i ich zróżnicowanie pod względem obfitości i rozmieszczenia wskazuje, że jest to główne i znaczące morfologiczne przystosowanie naczyń krwionośnych, służące przeciwprądowemu przenikaniu biologicznie aktywnych substancji w obszarze więzadła szerokiego macicy u owcy. W doświadczalnej części pracy wykazano, że oprócz przystosowań morfologicznych naczyń krwionośnych i limfatycznych występują także pojawiające się przejściowo przystosowania czynnościowe. Wskazują na to obserwowane zmiany kształtu i ułożenia komórek śródbłonka w tętnicy macicznej w zależności od fazy cyklu rujowego, a także u owiec owariektomizowanych, otrzymujących benzoesan estradiolu. Stwierdzono także, że wstrzykiwanie benzoesanu estradiolu owariektomizowanym owcom powodowało istotnie wyższy wzrost aktywności NADPH-diaforazy (markera syntazy tlenku azotu) w komórkach śródbłonka naczyń tętniczych i żylnych, a także w naczyniach limfatycznych. Zaobserwowane w cyklu rujowym i po podaniu estradiolu zmiany morfologii komórek śródbłonka oraz wzrost aktywności NADPH-diaforazy pozwalają przypuszczać, że istnieje powiązanie pomiędzy podwyższoną lokalnie, dzięki mechanizmowi przeciwprądowego przenikania, koncentracją hormonów jajnikowych a regulowaniem dopływu krwi do macicy i jajnika.

EN

The paper presents the morphology and mutual topographic relations between blood and lymph vessels of the broad ligament of ewe's uterus, examined employing LM, ТЕМ and SEM, and their presumable significance for the mechanism of counter-current transfer of hormones and other regulation agents. The experimental part of the paper constitutes an attempt to show a potential, local effect of ovarian hormones on some functional adjustments of vessels, connected with blood inflow to the uterus and ovary. The research was conducted on adult ewes during the reproductive period and seasonal anestrous, as well as 7-8 weeks after ovariectomy. The cycling ewes were used for morphological examinations, employing: vessel specimens filled with contrast medium and Microfil, corrosion casts of vessels and tissue specimens analyzed in a scanning electron microscope (SEM), histological specimens observed in a light microscope, and ultrastructure of lymph vessels studied by means of a transmission electron microscope (ТЕМ). The experimental material collected from ewes in the oestrus cycle and ovariectomized ones (treated with estradiol benzoate or not) enabled to determine: the response of endothelial cells of uterine and mesentric arteries - on the basis of tissue specimens analyzed in a SEM; and the activity of NADPH- diaphorase (marker of nitric oxide synthase) in the endothelium of blood and lymph vessels from the area of broad ligament - in the course of histochemical examinations. Macroscopic observations of blood vessels indicate that despite certain individual differen­ces, the oviductal branch and the uterine branch of the ovarian artery are characterized by tortuosity, which is less intensive in their case than in that of the ovarian branch. The oviductal branch and the uterine branch of the ovarian artery were situated side by side, or surrounded venous vessels. However, the arterial vessels supplying the oviduct and the anterior part of the uterine horn were not found to be located in a close neighbourhood of ovarian veins, carrying venous blood rich in ovarian hormones. It was also observed, for the first time, that the ovarian branch of the ovarian artery, most often straight in front of the ovary, becomes spiral in its medulla. Similar spiralization was also discovered in smaller ramifications emanating from the looped artery and heading for the cortical part of the ovary. The uterine artery and its small branches were straight within the mesometrium. Venous vessels, transporting blood both from the uterus and ovary, turned out to be very well developed. So were the valves. Despite their close contact with arterial vessels, supplying the ovary or the uterus, no cases of wall thickness reduction were observed in neighbouring vessels. It was found that the ovarian lymphatics situated along the mesovarium - although not numerous - split up and arrange not only in the vicinity of the ovarian branch, but also accompany the oviductal and uterine branches of the ovarian artery. Some of ovarian lympha­tics, usually small and thin-walled, get in between the loops of the spiral ovarian branch of the ovarian artery. Uterine lymphatics, on the other hand, formed two lines: one - less developed - headed for the uterine artery; the other transported lymph from the central and paraoviductal parts of the uterine horn and, at the ovary level, entered the mesovarium area. The vessels of the other line, most of them thick-walled, were situated on the ventral side of the mesovarium, between its muscular layer and blood vessels. Lymph vessels emanating from the uterus and located along the mesovarium used to join gradually with one another and then with lymph vessels emanating from the ovary, to form two or three collecting vessels. Before reaching the paraortal lymph node, lymph vessels were arranged next to the ovarian artery and utero- ovarian vein. An analysis of vessel corrosion casts performed by means of a SEM proved that bigger arterial and venous vessels, as well as lymph nodes emanating from the ovary and uterus, were surrounded by networks of microvessels (vasa vasorum). However, there were significant differences in both distribution and density of those networks. Less developed ones were present in the anterior parts of the uterine and ovarian arteries, rather than on their smaller, thin-walled ramifications. The most developed, multilayer vasa vasorum networks were obse­rved on tortuous ovarian and uterine branches of the ovarian artery, and the ovarian vein. The oviductal branch of the ovarian artery, as well as the branches emanating from the uterine artery, were in most cases surrounded by single-layer networks of microvessels. It was found that venules originating from vasa vasorum were situated in the external layer of the adventitia of arterial and venous vessels, or outside it. Well-developed, although mostly single- layer vasa vasorum networks surrounded also lymph vessels. An analysis in a SEM confirmed the arteriovenous character of microvessel networks surrounding lymph vessels. The research results indicate the existence in ewes of several - different from the ones that have been described so far - morphological adjustments of blood and lymph vessels to the counter-current transfer of hormones and, presumably, some other regulation agents, in the area of the uterus broad ligament. A close neighbourhood of the uterine and oviductal branches, with lymph vessels emanating from the ovary and transporting lymph rich in ovarian hormo­nes, seems conducive to the counter-current transfer and local supply of the uterus and oviduct with steroid hormones (Daniel et al. 1963). It may be suggested that lymph vessels emanating from the uterus and located on the ventral side of the mesovarium are the source of PGF 2α for the ovary, equally abundant as the utero-ovarian vein, accompanying the ovarian artery, or lymph vessels carrying utero-ovarian lymph. Moreover, it seems that venous plexuses on the ovarian branch of the ovarian artery - observed by other morphologists (Del Campo, Ginther 1973a; Lee, O'Shea 1976) and considered important routes of counter-current transfer of different substances - only indirectly participate in this mechanism in ewes. The presence of microvessel networks on blood and lymph vessels, as well as the differences between them concerning their density and distribution, indicate that this it the main and crucial morphologi­cal adjustment of blood vessels, designed for the counter-current transfer of biologically active substances in the area of the broad ligament of ewe's uterus. The experimental part of the paper shows that apart from morphological adjustments of blood and lymph vessels, there exist certain functional adjustments, which appear transiently. It finds confirmation in the changes in the shape and arrangement of endothelial cells that were observed in the uterine artery. The changes depend on the phase of the oestrus cycle and were also noted in ovariectomized ewes treated with estradiol benzoate. Moreover, it was found that injections of estradiol benzoate administered to ovariectomized ewes resulted in a significantly higher increase in the activity of NADPH-diaphorase (marker of nitric oxide synthase) in endothelial cells of arterial and venous vessels, as well as lymph vessels. The changes in the morphology of endothelial cells, observed during the oestrus cycle and after estradiol admini­stration, as well as an increase in the activity of NADPH-diaphorase, suggest that there is a relationship between locally higher (due to the mechanism of counter-current transfer) concen­tration of ovarian hormones and regulation of blood inflow to the uterus and ovary.

Słowa kluczowe

PL

morfologia zwierzat; anatomia topograficzna; adaptacja morfologiczna; owce; mikroskopia elektronowa; badania mikroskopowe; mikroskop elektronowy transmisyjny; mikroskop elektronowy skaningowy; naczynia krwionosne; naczynia limfatyczne; wiezadlo szerokie macicy; ultrastruktura; naczyniowe odlewy korozyjne

EN

animal morphology; topographical anatomy; morphological adaptation; sheep; electron microscopy; microscopy research; transmission electron microscope; scanning electron microscope; blood vessel; lymph vessel; broad ligament; uterus; ultrastructure; vascular corrosion cast

• Wydawca: -
• Czasopismo: Acta Academiae Agriculturae ac Technicae Olstenensis. Veterinaria
• Rocznik: 1998
• Tom: 25,Suppl.A
• Opis fizyczny: s.1-105,rys.,fot.,bibliogr.

Twórcy

autor

Zezula-Szpyra A

Bibliografia

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