Развитие этмоидного отдела хрящевого черепа в эмбриогенезе ужа обыкновенного, Natrix natrix (Ophidia, Colubridae)
Описано развитие этмоидного отдела хрящевого черепа Natrix natrix Linnaeus, 1758 на разных стадиях эмбриогенеза. Впервые для змей описана независимая от париетотектального хряща закладка planum antorbitale и ее самостоятельный очаг охрящевения, а также установлено наличие processus maxillaris anteri...
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irk-123456789-1093792016-11-28T03:02:28Z Развитие этмоидного отдела хрящевого черепа в эмбриогенезе ужа обыкновенного, Natrix natrix (Ophidia, Colubridae) Шевердюкова, А.В. Морфология Описано развитие этмоидного отдела хрящевого черепа Natrix natrix Linnaeus, 1758 на разных стадиях эмбриогенеза. Впервые для змей описана независимая от париетотектального хряща закладка planum antorbitale и ее самостоятельный очаг охрящевения, а также установлено наличие processus maxillaris anterior. Гипохоанальный хрящ формируется как независимая структура. The development of the ethmoidal region of Natrix natrix`s Linnaeus, 1758 chondrocranium at different stages of development is described. For the first time an independent appearance of the planum antorbitale and its independent centre of chondrification are described in snakes. Also for the first time for snakes the presence of processus maxillaris anterior was revealed. The hypochoanal cartilage is shown to develop as an independent structure. 2012 Article Развитие этмоидного отдела хрящевого черепа в эмбриогенезе ужа обыкновенного, Natrix natrix (Ophidia, Colubridae) / А.В. Шевердюкова // Вестник зоологии. — 2012. — Т. 46, № 3. — С. 247-257. — Бібліогр.: 14 назв. — рос. 0084-5604 http://dspace.nbuv.gov.ua/handle/123456789/109379 [591.3+591.471.4]:598.115.31 ru Вестник зоологии Інститут зоології ім. І.І. Шмальгаузена НАН України |
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Морфология Морфология Шевердюкова, А.В. Развитие этмоидного отдела хрящевого черепа в эмбриогенезе ужа обыкновенного, Natrix natrix (Ophidia, Colubridae) Вестник зоологии |
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Описано развитие этмоидного отдела хрящевого черепа Natrix natrix Linnaeus, 1758 на разных стадиях эмбриогенеза. Впервые для змей описана независимая от париетотектального хряща закладка planum antorbitale и ее самостоятельный очаг охрящевения, а также установлено наличие processus maxillaris anterior. Гипохоанальный хрящ формируется как независимая структура. |
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Article |
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Шевердюкова, А.В. |
| author_facet |
Шевердюкова, А.В. |
| author_sort |
Шевердюкова, А.В. |
| title |
Развитие этмоидного отдела хрящевого черепа в эмбриогенезе ужа обыкновенного, Natrix natrix (Ophidia, Colubridae) |
| title_short |
Развитие этмоидного отдела хрящевого черепа в эмбриогенезе ужа обыкновенного, Natrix natrix (Ophidia, Colubridae) |
| title_full |
Развитие этмоидного отдела хрящевого черепа в эмбриогенезе ужа обыкновенного, Natrix natrix (Ophidia, Colubridae) |
| title_fullStr |
Развитие этмоидного отдела хрящевого черепа в эмбриогенезе ужа обыкновенного, Natrix natrix (Ophidia, Colubridae) |
| title_full_unstemmed |
Развитие этмоидного отдела хрящевого черепа в эмбриогенезе ужа обыкновенного, Natrix natrix (Ophidia, Colubridae) |
| title_sort |
развитие этмоидного отдела хрящевого черепа в эмбриогенезе ужа обыкновенного, natrix natrix (ophidia, colubridae) |
| publisher |
Інститут зоології ім. І.І. Шмальгаузена НАН України |
| publishDate |
2012 |
| topic_facet |
Морфология |
| url |
http://dspace.nbuv.gov.ua/handle/123456789/109379 |
| citation_txt |
Развитие этмоидного отдела хрящевого черепа в эмбриогенезе ужа обыкновенного, Natrix natrix (Ophidia, Colubridae) / А.В. Шевердюкова // Вестник зоологии. — 2012. — Т. 46, № 3. — С. 247-257. — Бібліогр.: 14 назв. — рос. |
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Вестник зоологии |
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ÓÄÊ [591.3+591.471.4]:598.115.31
ÐÀÇÂÈÒÈÅ ÝÒÌÎÈÄÍÎÃÎ ÎÒÄÅËÀ ÕÐßÙÅÂÎÃÎ ×ÅÐÅÏÀ
 ÝÌÁÐÈÎÃÅÍÅÇÅ ÓÆÀ ÎÁÛÊÍÎÂÅÍÍÎÃÎ,
NATRIX NATRIX (OPHIDIA, COLUBRIDAE)
À. Â. Øåâåðäþêîâà
Èíñòèòóò çîîëîãèè èì. È. È. Øìàëüãàóçåíà ÍÀÍ Óêðàèíû,
óë. Á. Õìåëüíèöêîãî, 15, Êèåâ, 01601 Óêðàèíà
E-mail: hstramontana@gmail.com
Ïîëó÷åíî 12 èþëÿ 2011
Ïðèíÿòî 10 íîÿáðÿ 2011
Ðàçâèòèå ýòìîèäíîãî îòäåëà õðÿùåâîãî ÷åðåïà â ýìáðèîãåíåçå óæà îáûêíîâåííîãî, Natrix natrix
(Ophidia, Colubridae). Øåâåðäþêîâà À. Â. – Îïèñàíî ðàçâèòèå ýòìîèäíîãî îòäåëà õðÿùåâîãî
÷åðåïà Natrix natrix Linnaeus, 1758 íà ðàçíûõ ñòàäèÿõ ýìáðèîãåíåçà. Âïåðâûå äëÿ çìåé îïèñàíà
íåçàâèñèìàÿ îò ïàðèåòîòåêòàëüíîãî õðÿùà çàêëàäêà planum antorbitale è åå ñàìîñòîÿòåëüíûé
î÷àã îõðÿùåâåíèÿ, à òàêæå óñòàíîâëåíî íàëè÷èå processus maxillaris anterior. Ãèïîõîàíàëüíûé
õðÿù ôîðìèðóåòñÿ êàê íåçàâèñèìàÿ ñòðóêòóðà.
Êëþ÷åâûå ñ ëîâ à: ýìáðèîíàëüíîå ðàçâèòèå, õðÿùåâîé ÷åðåï, ýòìîèäíûé îòäåë, Natrix natrix.
Development of the Ethmoidal Region of the Chondrocranium in the Grass Snake, Natrix natrix (Ophidia,
Colubridae), Embryogenesis. Sheverdyukova H. V. – The development of the ethmoidal region of
Natrix natrix`s Linnaeus, 1758 chondrocranium at different stages of development is described. For the
first time an independent appearance of the planum antorbitale and its independent centre of chondri-
fication are described in snakes. Also for the first time for snakes the presence of processus maxillaris
anterior was revealed. The hypochoanal cartilage is shown to develop as an independent structure.
Ke y wo r d s: embryonic development, chondrocranium, ethmoidal region, Natrix natrix.
Ââåäåíèå
Öåíòðàëüíûì ýëåìåíòîì ýòìîèäíîãî îòäåëà õðÿùåâîãî ÷åðåïà çìåé ÿâëÿåòñÿ íîñîâàÿ ïåðåãî-
ðîäêà, ðàçäåëÿþùàÿ åãî íà äâå ñèììåòðè÷íûå íîñîâûå êàïñóëû. Ïðîèçâîäíûì íîñîâîé ïåðåãîðîä-
êè ñ÷èòàåòñÿ ïàðèåòîòåêòàëüíûé õðÿù, êîòîðûé ôîðìèðóåò êðûøó íîñîâîé êàïñóëû.  ëèòåðàòóðå
îðàëüíóþ ÷àñòü ïàðèåòîòåêòàëüíîãî õðÿùà íàçûâàþò cupola anterior, öåíòðàëüíóþ – paries lateralis
nasi, àáîðàëüíóþ – planum antorbitale (lamina orbitonasalis). Paries lateralis nasi, èçãèáàÿñü âåíòðàëüíî,
îáðàçóåò íîñîâóþ ðàêîâèíó. Äíî íîñîâîé êàïñóëû ïðåäñòàâëåíî lamina transversalis anterior, õðÿùîì
îðãàíà ßêîáñîíà, ýêòîõîàíàëüíûì è ãèïîõîàíàëüíûì («îòðîñòîê Êå») õðÿùàìè (Bäckström, 1931;
Pringle, 1954; Haluska, Alberch, 1983; Deiques, 2004).
Ñâåäåíèÿ î ñòðîåíèè è ðàçâèòèè ýòìîèäíîãî îòäåëà õðÿùåâîãî ÷åðåïà Natrix natrix íåìíîãî÷èñ-
ëåííû.  îòäåëüíûõ ðàáîòàõ ïðåäñòàâëåíî îïèñàíèå ñòðîåíèÿ õðÿùåâîãî ÷åðåïà íà îäíîé èç ñòàäèé
(Born, 1883; Gaupp, 1906, öèò. ïî: Kamal, Hammouda, 1965 à); â äðóãèõ – íà íåñêîëüêèõ îòäåëüíûõ
ñòàäèÿõ ýìáðèîíàëüíîãî ðàçâèòèÿ (Parker, 1878; Bäckström, 1931; ×åêàíîâñêàÿ, 1936).
Îòíîñèòåëüíî ñïîñîáà çàêëàäêè è ôîðìèðîâàíèÿ îòäåëüíûõ ñòðóêòóð ýòìîèäíîãî îòäåëà ó ðàç-
ëè÷íûõ àâòîðîâ âñòðå÷àþòñÿ ðàçíîãëàñèÿ. Â. Ê. Ïàðêåð è Ã. Áîðí ñ÷èòàëè ýêòîõîàíàëüíûé õðÿù
ñàìîñòîÿòåëüíîé ñòðóêòóðîé. Ê. Áåêñòðîì îáíàðóæèë ó N. natrix íåçàâèñèìóþ çàêëàäó íîñîâîé ïåðå-
ãîðîäêè è åå ñàìîñòîÿòåëüíûé î÷àã îõðÿùåâåíèÿ è â îòëè÷èå îò íèõ ñ÷èòàë ýêòîõîàíàëüíûé õðÿù
àáîðàëüíûì ïðîäîëæåíèåì õðÿùà îðãàíà ßêîáñîíà, íå èìåþùèì ñàìîñòîÿòåëüíîãî î÷àãà îõðÿùåâå-
íèÿ. Ïðîòèâîðå÷èâû ëèòåðàòóðíûå äàííûå è îòíîñèòåëüíî êîíòàêòà planum antorbitale è àáîðàëüíîé
÷àñòè íîñîâîé ïåðåãîðîäêè.  ÷àñòíîñòè Ê. Áåêñòðîì îòðèöàë ñëèÿíèå ýòèõ ñòðóêòóð ó N. natrix; à íà
ðåêîíñòðóêöèÿõ ÷åðåïà, ñäåëàííûõ Î. Â. ×åêàíîâñêîé, âèäíî, ÷òî ýòè ñòðóêòóðû èìåþò øèðîêîå
ñîåäèíåíèå åùå íà ðàííèõ ñòàäèÿõ ðàçâèòèÿ.
Vestnik zoologii, 46(3): 247—257, 2012 Ìîðôîëîãèÿ
Ñ öåëüþ âûÿñíåíèÿ ñïîðíûõ âîïðîñîâ ìû ïðîñëåäèëè äèíàìèêó ðàçâèòèÿ ñòðóêòóð ýòìîèäíî-
ãî îòäåëà õðÿùåâîãî ÷åðåïà N. natrix íà ðàçíûõ ñòàäèÿõ ýìáðèîãåíåçà.
Ìàòåðèàë è ìåòîäû
Ìàòåðèàë ñîáðàí â èþíå–àâãóñòå 2010 ã. â îêð. ñ. Âàêàëîâùèíà Ñóìñêîé îáë. Áåðåìåííûå
ñàìêè N. natrix áûëè îòëîâëåíû â åñòåñòâåííûõ óñëîâèÿõ è ïîìåùåíû â òåððàðèóìû. Ïîñëå îòêëàä-
êè ÿèö ñàìîê âûïóñêàëè â ìåñòà îòëîâà. ßéöà èíêóáèðîâàëè â óâëàæíåííîì âåðìèêóëèòå ïðè òåì-
ïåðàòóðå 27—30 ÎÑ. Èíêóáàöèÿ äëèëàñü 30 ñóò (± 1 ñóò).
Ýìáðèîíàëüíûé ìàòåðèàë ôèêñèðîâàëè â 4 %-íîì ðàñòâîðå ôîðìàëèíà. Áûëè èçãîòîâëåíû
ñåðèéíûå ãèñòîëîãè÷åñêèå ñðåçû òîëùèíîé 5—7 ìêì, êîòîðûå îêðàøèâàëè àëüöèàíîâûì ñèíèì è
ãåìàòîêñèëèí-ýîçèíîì. Äëÿ âûÿâëåíèÿ îòëîæåíèé ñîëåé êàëüöèÿ â êîñòíîì ìàòðèêñå èñïîëüçîâàëè
ìåòîä Êîññà (Ðîìåéñ, 1954).
Ñòàäèè ðàçâèòèÿ ýìáðèîíîâ îïðåäåëÿëè ïî òàáëèöå íîðìàëüíîãî ðàçâèòèÿ, ðàçðàáîòàííîé äëÿ
Thamnophis sirtalis (Colubridae) (Zehr, 1962). Âñåãî èññëåäîâàíî 27 ýìáðèîíîâ 27—37 ñòàäèé íîðìàëü-
íîãî ðàçâèòèÿ. Èññëåäîâàëè íåñêîëüêî ýìáðèîíîâ îäíîé ñòàäèè. Ñòàäèè, íà êîòîðûõ ðàçâèòèå çàìåò-
íî ïðîäâèíóëîñü, íî íå äîñòèãëî ñëåäóþùåé ñòàäèè, îáîçíà÷åíû çíàêîì «+».
Ìèêðîôîòîãðàôèè ãèñòîëîãè÷åñêèõ ïðåïàðàòîâ ñäåëàíû ñ ïîìîùüþ ìèêðîñêîïà Zeiss Axio
Imager M1 è ïðîãðàììíîãî îáåñïå÷åíèÿ Zeiss Axio Vision v. 4.63 â öåíòðå êîëëåêòèâíîãî ïîëüçîâà-
íèÿ óíèêàëüíûì îáîðóäîâàíèåì Èíñòèòóòà çîîëîãèè èì. È. È. Øìàëüãàóçåíà ÍÀÍ Óêðàèíû.
Òîòàëüíûå ïðåïàðàòû èçãîòàâëåíû ïî ìåòîäèêå Å. Â. Ñèìîíñà è Ä. Ð. Âàí Õîðíà (1971).
Ðåçóëüòàòû
Ñòàäèÿ 2 7
Íà ýòîé ñòàäèè ïîÿâëÿþòñÿ ïåðâûå çàêëàäêè ñòðóêòóð ýòìîèäíîãî îòäåëà
õðÿùåâîãî ÷åðåïà. Ê ýòîìó ìîìåíòó ñôîðìèðîâàíû îáîíÿòåëüíûå ìåøêè è
õîðîøî ðàçâèòû îðãàíû ßêîáñîíà.  îáùåé ìåçåíõèìíîé ìàññå ðàçëè÷èìî ñêîï-
ëåíèå ìåçåíõèìíûõ êëåòîê, ðàçäåëÿþùåå îòäåë íà äâå ñèììåòðè÷íûå ïîëîâèíû.
Ýòî ñêîïëåíèå ÿâëÿåòñÿ îðàëüíûì ïðîäîëæåíèåì ïåðåäíèõ ìåçåíõèìíûõ êîíöîâ
÷åðåïíûõ òðàáåêóë è ïðåäñòàâëÿåò ñîáîé çàêëàäêó áóäóùåé íîñîâîé ïåðåãîðîä-
êè. Îáîíÿòåëüíûå ìåøêè è îðãàíû ßêîáñîíà ðàñïîëîæåíû ëàòåðàëüíî îò íåå.
Îðàëüíî ìåçåíõèìíàÿ çàêëàäêà íîñîâîé ïåðåãîðîäêè ðàçäâàèâàåòñÿ, îáðàçóÿ
çàêëàäêè ñupola anterior, êîòîðûå îãðàíè÷èâàþò îáîíÿòåëüíûå ìåøêè ìåäèàëü-
248 À. Â. Øåâåðäþêîâà
Ðèñ. 1. Ñðåç ãîëîâû ýìáðèîíà N. natrix â ïîïåðå÷íîé ïëîñêîñòè: à – 27-ÿ ñòàäèÿ ðàçâèòèÿ (äàëåå ñò.
ð.): 1 – ïåðåäíèé ìîçã; 2 – îðãàí ßêîáñîíà; 3 – îáîíÿòåëüíûé íåðâ; 4 – ãëàç; 5 – ìåçåíõèìíàÿ
çàêëàäêà cupola anterior; 6 – ìåçåíõèìíàÿ çàêëàäêà õðÿùà îðãàíà ßêîáñîíà; b – 28+ ñò. ð.: 1 – îáî-
íÿòåëüíûé ìåøîê; 2 – ìåçåíõèìíàÿ çàêëàäêà cupola anterior.
Fig. 1. Cross-section of N. natrix embryo head: a – the 27 stage of development: 1 – brain; 2 – Jacobson’s
organ; 3 – olfactory nerve; 4 – eye; 5 – primordium of cupola anterior; 6 – primordium of cartilage of
Jacobson’s organ; b – the 28+ stage of development: 1 – cavum nasi; 2 – primordium of cupola anterior.
íî (ðèñ. 1, à). Âåíòðàëüíî îò îðãàíà ßêîáñîíà ðàñïîëîæåíî ñêîïëåíèå ìåçåíõè-
ìû, êîòîðîå ïðåäñòàâëÿåò ñîáîé çàêëàäêó õðÿùà îðãàíà ßêîáñîíà (ðèñ. 1, à).
Ìåçåíõèìíàÿ çàêëàäêà planum antorbitale òÿíåòñÿ îò ïåðåäíèõ ìåçåíõèìíûõ
êîíöîâ òðàáåêóë â ëàòåðàëüíîì íàïðàâëåíèè íàä îðãàíàìè ßêîáñîíà (ðèñ. 2, à).
Ñ ò à äèÿ 2 8
Îò ëàòåðàëüíûõ êðàåâ ìåçåíõèìíûõ çàêëàäîê cupola anterior â àáîðàëüíîì
íàïðàâëåíèè òÿíóòñÿ ìåçåíõèìíûå òÿæè, ïðåäñòàâëÿþùèå ñîáîé çàêëàäêè paries
lateralis nasi. Îíè ñëóæàò ëàòåðàëüíîé ãðàíèöåé áóäóùèõ fenestra olfactoria advec-
hens, ÷åðåç êîòîðûå ïðîõîäÿò îáîíÿòåëüíûå íåðâû.
249Ðàçâèòèå ýòìîèäíîãî îòäåëà õðÿùåâîãî ÷åðåïà â ýìáðèîãåíåçå óæà îáûêíîâåííîãî…
Ðèñ. 2. Ñðåç ãîëîâû ýìáðèîíà N. natrix â ïîïåðå÷íîé ïëîñêîñòè: à – 27-ÿ ñò. ð.: 1 – ìåçåíõèìíàÿ
çàêëàäêà planum antorbitale; 2 – îðãàí ßêîáñîíà; 3 – ïåðåäíèé ìîçã; 4 – òðàáåêóëû; 5 – ãëàç; á –
28+ ñò. ð.: 1 – ïåðåäíèé ìîçã; 2 – ìåçåíõèìíàÿ çàêëàäêà planum antorbitale; 3 – òðàáåêóëÿðíàÿ ïëà-
ñòèíêà; 4 – îðãàí ßêîáñîíà; 5 – õîàíà; 6 – ãëàç; 7 – ðîòîâàÿ ïîëîñòü; c – 29-ÿ ñò. ð.: 1 – ìîçã;
2 – òðàáåêóëÿðíàÿ ïëàñòèíêà; 3 – ìåçåíõèìíàÿ çàêëàäêà planum antorbitale; 4 – õîàíà; 5 – ãëàç; 6 –
ðîòîâàÿ ïîëîñòü; 7 – ÿçûê; 8 – Ìåêêåëåâ õðÿù.
Fig. 2. Cross-section of N. natrix embryo head: à – the 27 stage of development: 1 – primordium of planum
antorbitale; 2 – Jacobson’s organ; 3 – brain; 4 – trabecula; 5 – eye; á – the 28+ stage of development:
1 – brain; 2 – primordium of planum antorbitale; 3 – trabecula communis; 4 – Jacobson’s organ; 5 –
choana; 6 – eye; 7 – stomodeum; c – at the 29 stage of development: 1 – brain; 2 – trabecula commu-
nis; 3 – primordium of planum antorbitale; 4 – choana; 5 – eye; 6 – stomodeum; 7 – tongue; 8 –
Meckel`s cartilage.
Ñòàäèÿ 2 8+
Îñòàâàÿñü ìåçåíõèìíûìè, çàêëàäêè ñupola anterior óäëèíèëèñü â îðàëüíîì
íàïðàâëåíèè. Èõ ïåðåäíèå êðàÿ îêðóæàþò îáîíÿòåëüíûå ìåøêè ìåäèàëüíî, äîð-
ñàëüíî è äîðñîëàòåðàëüíî (ðèñ. 1, b).
Ïðîèñõîäèò ñëèÿíèå ïåðåäíèõ êîíöîâ òðàáåêóë, âñëåäñòâèå ÷åãî ôîðìèðó-
åòñÿ õðÿùåâàÿ òðàáåêóëÿðíàÿ ïëàñòèíêà (ðèñ. 2, b). Ïðîöåññ îõðÿùåâåíèÿ ðàñ-
ïðîñòðàíÿåòñÿ â îðàëüíîì íàïðàâëåíèè îò òðàáåêóëÿðíîé ïëàñòèíêè äî ñåðåäè-
íû íîñîâîé ïåðåãîðîäêè.
Ìåçåíõèìíûå çàêëàäêè paries lateralis nasi èçîãíóòû S-îáðàçíî: âåíòðàëüíî,
à çàòåì ìåäèàëüíî, òàê, ÷òî çàõîäÿò â èçãèá îáîíÿòåëüíûõ ìåøêîâ, îáðàçóÿ
çàêëàäêó íîñîâîé ðàêîâèíû (ðèñ. 3). Çàêëàäêè õðÿùà îðãàíà ßêîáñîíà è íîñî-
âîé ðàêîâèíû ñîåäèíåíû ëàòåðàëüíî ïîëîñêîé ìåçåíõèìíîé òêàíè.
Ïðîèñõîäèò ñëèÿíèå àáîðàëüíîãî êðàÿ ìåçåíõèìíûõ çàêëàäîê paries lateralis
nasi è planum antorbitale. Ìåçåíõèìíûå çàêëàäêè planum antorbitale óäëèíåíû â
ëàòåðàëüíîì íàïðàâëåíèè, èõ ëàòåðàëüíûå êðàÿ äóãîîáðàçíî èçîãíóòû âåíòðî-
ëàòåðàëüíî (ðèñ. 2, b).
Ñ ò à äèÿ 2 9
Îõðÿùåâåíèå íîñîâîé ïåðåãîðîäêè ïðîäîëæàåòñÿ â îðàëüíîì íàïðàâëåíèè.
Ñupola anterior óâåëè÷èëèñü è îêðóæàþò îáîíÿòåëüíûå ìåøêè ìåäèàëüíî,
âåíòðîìåäèàëüíî, äîðñàëüíî è äîðñîëàòåðàëüíî. Ïðîöåññ îõðÿùåâåíèÿ ðàñïðî-
ñòðàíèëñÿ îò íîñîâîé ïåðåãîðîäêè íà ìåäèàëüíûå ó÷àñòêè cupola anterior.
 ìåñòå, ãäå ïåðåäíèé êðàé íîñîâîé ïåðåãîðîäêè ïåðåõîäèò â cupola anterior,
âèäíû çàêëàäêè lamina transversalis anterior â âèäå êîðîòêèõ ìåçåíõèìíûõ òÿæåé,
íàïðàâëåííûõ àáîðàëüíî.
250 À. Â. Øåâåðäþêîâà
Ðèñ. 3. Ñðåç ãîëîâû ýìáðèîíà N. natrix â ïîïåðå÷íîé ïëîñêîñòè, 28+ ñò. ð.: 1 – ïåðåäíèé ìîçã; 2 –
îáîíÿòåëüíûé íåðâ; 3 – íîñîâàÿ ïåðåãîðîäêà; 4 – îðãàí ßêîáñîíà; 5 – îáîíÿòåëüíûé ìåøîê; 6 –
ìåçåíõèìíàÿ çàêëàäêà õðÿùà îðãàíà ßêîáñîíà; 7 – ìåçåíõèìíàÿ çàêëàäêà paries lateralis nasi; 8 –
ìåçåíõèìíàÿ çàêëàäêà íîñîâîé ðàêîâèíû; 9 – ìåçåíõèìíîå ñîåäèíåíèå ìåæäó ìåçåíõèìíûìè
çàêëàäêàìè õðÿùà îðãàíà ßêîáñîíà è íîñîâîé ðàêîâèíû.
Fig. 3. Cross-section of N. natrix embryo head at the 28+ stage of development: 1 – brain; 2 – olfactory
nerve; 3 – nasal septum; 4 – cartilage of Jacobson’s organ; 5 – cavum nasi; 6 – primordium of cartilage
of Jacobson’s organ; 7 – paries lateralis nasi; 8 – primordium of nasal concha; 9 – mesenchymal connec-
tion between primordium of cartilage of Jacobson’s organ and nasal concha.
Èçîãíóâøèåñÿ âåíòðàëüíî íà ïðåäûäóùåé ñòàäèè ëàòåðàëüíûå êðàÿ ìåçåí-
õèìíûõ çàêëàäîê planum antorbitale óäëèíèëèñü, äîñòèãíóâ óðîâíÿ íèæíåãî êðàÿ
ãëàç (ðèñ. 2, c).
Ñ ò à äèÿ 29 +
Íîñîâàÿ ïåðåãîðîäêà ïîëíîñòüþ îõðÿùåâåëà. Íà ïîñëåäóþùèõ ñòàäèÿõ ðàç-
âèòèÿ ïðîèñõîäèò ëèøü åå ðîñò. Â îðàëüíîàáîðàëüíîì íàïðàâëåíèè îò cupola
anterior íà÷èíàþò îõðÿùåâåâàòü ìåçåíõèìíûå çàêëàäêè paries lateralis nasi, íîñî-
âàÿ ðàêîâèíà è lamina transversalis anterior; íåçàâèñèìûå î÷àãè îõðÿùåâåíèÿ
ïîÿâèëèñü â çàêëàäêå õðÿùà îðãàíà ßêîáñîíà è planum antorbitale (ðèñ. 4).
Ñ ò à äèÿ 3 0
Îõðÿùåâåíèå â çàêëàäêàõ ñupola anterior ðàñïðîñòðàíÿåòñÿ ëàòåðàëüíî,
ìåçåíõèìíûìè îñòàëèñü òîëüêî èõ ëàòåðàëüíûå êðàÿ. Ïåðåäíåëàòåðàëüíûé êðàé
251Ðàçâèòèå ýòìîèäíîãî îòäåëà õðÿùåâîãî ÷åðåïà â ýìáðèîãåíåçå óæà îáûêíîâåííîãî…
Ðèñ. 5. Ñðåç ãîëîâû ýìáðèîíà N. natrix â ïîïåðå÷íîé ïëîñêîñòè, 33-ÿ ñò. ð.: 1 – íîñîâàÿ ïåðåãîðîä-
êà; 2 – cupola anterior; 3 – îáîíÿòåëüíûé ìåøîê; 4 – îðãàí ßêîáñîíà; 5 – âîìåðîíàçàëüíàÿ êàìå-
ðà; 6 – õðÿù ßêîáñîíîâà îðãàíà; 7 – íîñîâàÿ êîñòü; 8 – âåðõíå÷åëþñòíàÿ êîñòü; 9 – ñîøíèê; 10 –
ñåïòîìàêñèëëà.
Fig. 5. Cross-section of N. natrix embryo head at the 33 stage of the development: 1 – nasal septum; 2 –
cupola anterior; 3 – cavum nasi; 4 – Jacobson‘s organ; 5 – concha of vomero-nasal organ; 6 – cartilage
of Jacobson‘s organ; 7 – nasal bone; 8 – maxilla; 9 – vomer; 10 – septomaxilla.
Ðèñ. 4. Ñðåç ãîëîâû ýìáðèîíà N. natrix â ïîïåðå÷íîé ïëîñêîñòè, 29+ ñò. ð.: 1 – îðãàí ßêîáñîíà; 2 –
íîñîâàÿ ïåðåãîðîäêà; 3 – îáîíÿòåëüíûé íåðâ; 4 – î÷àã îõðÿùåâåíèÿ â çàêëàäêå õðÿùà îðãàíà
ßêîáñîíà; 5 – î÷àã îõðÿùåâåíèÿ â çàêëàäêå planum antorbitale; 6 – õîàíà; 7 – ãëàç.
Fig. 4. Cross-section of N. natrix embryo head at the 29+ stage of development: 1 – Jacobson‘s organ; 2 –
nasal septum; 3 – olfactory nerve; 4 – centre of chondrification in the primordium of cartilage of Jacobson‘s
organ; 5 – centre of chondrification in the primordium of planum antorbitale; 6 – choana; 7 – eye.
ñupola anterior ïðåäñòàâëÿåò ñîáîé processus alaris superior, îãðàíè÷èâàþùèé
íîçäðþ äîðñàëüíî.
Âåíòðîàáîðàëüíî îò íîñîâîé ðàêîâèíû ïîÿâëÿåòñÿ õðÿùåâîé îòðîñòîê,
êîòîðûé ñëèâàåòñÿ ñ îðàëüíûì êðàåì planum antorbitale (ðèñ. 8).
Lamina transversalis anterior çíà÷èòåëüíî óäëèíèëèñü è îõðÿùåâåëè. Ðàññòîÿ -
íèå ìåæäó àáîðàëüíûìè êîíöàìè lamina transversalis anterior è ïîëíîñòüþ îõðÿ-
ùåâåâøèì íà ýòîé ñòàäèè õðÿùåì îðãàíà ßêîáñîíà î÷åíü íåçíà÷èòåëüíîå, íî
ýòè ñòðóêòóðû íå ñëèâàþòñÿ.
Ìåäèàëüíûé ó÷àñòîê õðÿùà îðãàíà ßêîáñîíà – âîìåðîíàçàëüíàÿ êàìåðà –
îãðàíè÷èâàåò îðãàí ßêîáñîíà âåíòðàëüíî, à õðÿù îðãàíà ßêîáñîíà – âåíòðîëà-
òåðàëüíî (ðèñ. 5).
Âåíòðîëàòåðàëüíî îò îðãàíà ßêîáñîíà ðàñïîëàãàåòñÿ ñàìîñòîÿòåëüíàÿ
ìåçåíõèìíàÿ çàêëàäêà ãèïîõîàíàëüíîãî õðÿùà. Îíà ïðåäñòàâëÿåò ñîáîé òÿæ,
îðàëüíûé êîíåö êîòîðîãî äîñòèãàåò óðîâíÿ õðÿùà îðãàíà ßêîáñîíà, à àáîðàëü-
íûé – óðîâíÿ çàäíåé ñòåíêè íîñîâîé êàïñóëû. Íà ýòîé æå ñòàäèè â ñðåäèííîé
÷àñòè ýòîé ñòðóêòóðû âèäåí î÷àã îõðÿùåâåíèÿ.
Çàêëàäêè planum antorbitale îõðÿùåâåëè íåðàâíîìåðíî: ó÷àñòêè, ãðàíè÷àùèå
ñ òðàáåêóëÿðíîé ïëàñòèíêîé, îñòàëèñü ìåçåíõèìíûìè. Íà ýòîé æå ñòàäèè îò
îðàëüíîâåíòðàëüíîé ïîâåðõíîñòè planum antorbitale â îðàëüíîì íàïðàâëåíèè
òÿíåòñÿ îõðÿùåâåâàþùèé îòðîñòîê (ðèñ. 8). Îñíîâûâàÿñü íà åãî ïðîèñõîæäåíèè,
ìû ñ÷èòàåì, ÷òî ýòî processus maxillaris anterior.
Ñ ò à äèÿ 3 1
Ñupola anterior óâåëè÷èëèñü â âåíòðîàáîðàëüíîì íàïðàâëåíèè, îáðàçîâàâ
ïàðíûå êóïîëîîáðàçíûå ñòðóêòóðû.  ïîïåðå÷íîì ñå÷åíèè ýòè ñòðóêòóðû âûãëÿ-
äÿò êàê îêðóæíîñòè, ñîåäèíåííûå âåíòðîìåäèàëüíî äóãîîáðàçíî èçîãíóòîé
íîñîâîé ïåðåãîðîäêîé (ðèñ. 6, à). Ñ âåíòðîëàòåðàëüíîé ñòîðîíû ñupola anterior
ïîÿâëÿåòñÿ ìåçåíõèìíàÿ çàêëàäêà processus alaris inferior.
Àáîðàëüíî îò õðÿùà îðãàíà ßêîáñîíà ïîÿâëÿåòñÿ ìåçåíõèìíûé òÿæ, ïðåä-
ñòàâëÿþùèé ñîáîé çàêëàäêó ýêòîõîàíàëüíîãî õðÿùà.
Ìåçåíõèìíàÿ ñâÿçêà, êîòîðàÿ ïîÿâèëàñü íà ñòàäèè 28+, ìåæäó õðÿùîì îðãà-
íà ßêîáñîíà è paries lateralis nasi, îõðÿùåâåëà.
Ðrocessus maxillaris anterior, êîòîðûé ïîÿâèëñÿ íà ïðåäûäóùåé ñòàäèè, ïîë-
íîñòüþ îõðÿùåâåë.
Ñ ò à äèÿ 3 1+
Ïðîöåññ îõðÿùåâåíèÿ ðàñïðîñòðàíÿåòñÿ îò cupola anterior íà ðrocessus alaris
inferior.
Ïðîèñõîäèò ñëèÿíèå lamina transversalis anterior è õðÿùà îðãàíà ßêîáñîíà
(ðèñ. 8). Ïðîöåññ îõðÿùåâåíèÿ ðàñïðîñòðàíèëñÿ ñî ñòîðîíû õðÿùà îðãàíà
ßêîáñîíà íà çàêëàäêó ýêòîõîàíàëüíîãî õðÿùà.
Ñò à äèÿ 3 2
Ãèïîõîàíàëüíûé õðÿù ïîëíîñòüþ ñôîðìèðîâàí. Äðóãèõ âèäèìûõ èçìåíå-
íèé â ýòìîèäíîì îòäåëå íà ýòîé ñòàäèè íå îáíàðóæåíî.
Ñò à äèÿ 3 3
Âåíòðîëàòåðàëüíûé êðàé èçãèáà paries lateralis nasi óòîëùàåòñÿ, îáðàçóÿ îòðî-
ñòîê íàïðàâëåííûé âåíòðàëüíî (ðèñ. 6, c).
252 À. Â. Øåâåðäþêîâà
Ðrocessus alaris inferior ïîëíîñòüþ îõðÿùåâåë. Ðrocessus alaris inferior è proces-
sus alaris superior íå ñîåäèíÿþòñÿ, òàêèì îáðàçîì, íîçäðÿ èçíà÷àëüíî ïðåäñòàâ-
ëÿåò ñîáîé âûðåçêó, îáðàùåííóþ ëàòåðàëüíî (ðèñ. 6, b).
Ñ ò à äèÿ 3 4
Ýêòî- è ãèïîõîàíàëüíûé õðÿùè ñâîèìè àáîðàëüíûìè êîíöàìè ñëèëèñü,
îáðàçîâàâ ýêòîõîàíàëüíóþ ïëàñòèíêó (ðèñ. 9, à, b).
Ñ ò à äèÿ 3 5
Âñëåäñòâèå ïîñòåïåííîãî ñáëèæåíèÿ planum antorbitale è òðàáåêóëÿðíîé
ïëàñòèíêè â õîäå ýìáðèîãåíåçà íà 35-é ñòàäèè ýòè ñòðóêòóðû ñëèâàþòñÿ (ðèñ. 7).
253Ðàçâèòèå ýòìîèäíîãî îòäåëà õðÿùåâîãî ÷åðåïà â ýìáðèîãåíåçå óæà îáûêíîâåííîãî…
Ðèñ. 6. Ñðåçû ãîëîâû ýìáðèîíà N. natrix â ïîïåðå÷íîé ïëîñêîñòè 33-ÿ ñò. ð. (ïðåäñòàâëåíû ñðåçû
ðàçíûõ óðîâíåé): a – 1 – íîñîâàÿ ïåðåãîðîäêà; 2 – ÿéöîâûé çóá; 3 – cupola anterior; 4 – ïðåäìàê-
ñèëëà; b – 1 – íîñîâàÿ ïåðåãîðîäêà; 2 – îáîíÿòåëüíûé ìåøîê; 3 – ïàðèåòîòåêòàëüíûé õðÿù; 4 –
íîñîâàÿ êîñòü; 5 – processus alaris inferior; 6 – ñåïòîìàêñèëëà; 7 – lamina transversalis anterior; 8 –
çàêëàäêà âåðõíå÷åëþñòíîé êîñòè; ñ – 1 – íîñîâàÿ ïåðåãîðîäêà; 2 – îðãàí ßêîáñîíà; 3 – îáîíÿòåëü-
íûé ìåøîê; 4 – fenestra olfactoria advehens; 5 – ïàðèåòîòåêòàëüíûé õðÿù; 6 – ïðåäëîáíàÿ êîñòü; 7 –
íîñîâàÿ ðàêîâèíà; 8 – fenestra latiralis nasi; 9 – processus lateralis nasi; 10 – ñîøíèê; 11 – ýêòîõîà-
íàëüíûé õðÿù; 12 – ãèïîõîàíàëíûé õðÿù; 13 – ñåïòîìàêñèëëà.
Fig. 6. Cross-section of N. natrix embryo head at the 33 stage of development at the different levels: a – 1 –
nasal septum; 2 – egg tooth; 3 – cupola anterior; 4 – premaxilla; b – 1 – nasal septum; 2 – cavum nasi;
3 – parietotectal cartilage; 4 – nasal bone; 5 – processus alaris inferior; 6 – septomaxilla; 7 – lamina
transversalis anterior; 8 – maxilla; ñ – 1 – nasal septum; 2 – Jacobson‘s organ; 3 – cavum nasi; 4 – fen-
estra olfactoria advehens; 5 – parietotectal cartilage; 6 – prefrontal; 7 – nasal concha; 8 – fenestra lati-
ralis nasi; 9 – processus lateralis nasi; 10 – vomer; 11 – ectochoanal cartilage; 12 – hypochoanal carti-
lage; 13 – septomaxilla.
Òàêèì îáðàçîì, fenestra olfactoria advechens ïðåäñòàâëÿåò ñîáîé îòâåðñòèå, îãðà-
íè÷åííîå õðÿùåâûìè ñòðóêòóðàìè: îðàëüíî – cupola anterior, ëàòåðàëüíî – pari-
es lateralis nasi, àáîðàëüíî – planum antorbitale è çàìûêàþùååñÿ íà óðîâíå òðàáå-
êóëÿðíîé ïëàñòèíêè.
Ñò à äèÿ 3 6
Processus alaris inferior ñìåùåí ëàòåðàëüíåå processus alaris superior. Òàêèì
îáðàçîì, íîçäðÿ íà ýòîé ñòàäèè îáðàùåíà äîðñîëàòåðàëüíî.
Íà ïîñëåäíåé, 37-é ñòàäèè ðàçâèòèÿ, âèäèìûõ èçìåíåíèé â ýòìîèäíîì îòäå-
ëå õðÿùåâîãî ÷åðåïà N. natrix íå îáíàðóæåíî.
254 À. Â. Øåâåðäþêîâà
Ðèñ. 8. Òîòàëüíûé ïðåïàðàò ýòìîèäíîãî îòäåëà ýìáðèîíà N. natrix 35-ÿ ñò. ð.; âèä ñ ëàòåðàëüíîé ñòî-
ðîíû: 1 – cupola anterior; 2 – lamina transversalis anterior; 3 – õðÿù îðãàíà ßêîáñîíà; 4 – ãèïîõîà-
íàëüíûé õðÿù; 5 – õðÿùåâîå ñîåäèíåíèå íîñîâîé ðàêîâèíû è õðÿùà îðãàíà ßêîáñîíà; 6 – íîñîâàÿ
ðàêîâèíà è å¸ îòðîñòîê; 7 – planum antorbitale; 8 – processus maxillaris anterior; 9 – òðàáåêóëà; 10 –
ïðåäìàêñèëëà; 11 – ñåïòîìàêñèëëà; 12 – íîñîâàÿ êîñòü.
Fig. 8. Cleared and stained ethmoidal region of N. natrix embryo at the 35 stage of development from the
lateral view: 1 – cupola anterior; 2 – lamina transversalis anterior; 3 – cartilage of Jacobson‘s organ; 4 –
hypochoanal cartilage; 5 – cartilaginous connection between nasal concha and cartilage of Jacobson‘s organ;
6 – nasal concha and its process; 7 – planum antorbitale; 8 – processus maxillaris anterior; 9 – trabecu-
la; 10 – premaxilla; 11 – septomaxilla; 12 – nasal bone.
Ðèñ. 7. Ñðåç ãîëîâû ýìáðèîíà N. natrix â ïîïåðå÷íîé ïëîñêîñòè íà óðîâíå òðàáåêóëÿðíîé ïëàñòèí-
êè 35-ÿ ñò. ð.: 1 – òðàáåêóëÿðíàÿ ïëàñòèíêà; 2 – planum antorbitale; 3 – ñîøíèê; 4 – ëîáíàÿ êîñòü.
Fig. 7. Cross-section of N. natrix embryo head at the 35 stage of development at the level of trabecula com-
munis: 1 – trabecula communis; 2 – planum antorbitale; 3 – vomer; 4 – frontale.
Îáñóæäåíèå
Ïîëàãàåì, ÷òî ñðàâíåíèå ïîëó÷åííûõ íàìè ðåçóëüòàòîâ ñ ëèòåðàòóðíûìè
äàííûìè è èõ îáñóæäåíèå áóäåò áîëåå íàãëÿäíûì, åñëè âåñòè åãî ïî îòäåëüíûì
ñòðóêòóðàì ýòìîèäíîãî îòäåëà.
Îòíîñèòåëüíî ïðîèñõîæäåíèÿ íîñîâîé ïåðåãîðîäêè ó N. natrix íàøè äàííûå
ñîâïàäàþò ñ äàííûìè Ý. Ãàóïïà (Gaupp, 1906) è Î. Â. ×åêàíîâñêîé
(×åêàíîâñêàÿ, 1936). Ìû îáíàðóæèëè, ÷òî íà 27 ñòàäèè ðàçâèòèÿ, êîãäà òðàáåêó-
ëÿðíàÿ ïëàñòèíêà åùå íå ñôîðìèðîâàíà, ìåçåíõèìíàÿ çàêëàäêà íîñîâîé ïåðåãî-
ðîäêè âûãëÿäèò êàê îðàëüíîå ïðîäîëæåíèå ïåðåäíèõ êîíöîâ òðàáåêóë. Íàì íå
óäàëîñü óâèäåòü ñàìîñòîÿòåëüíîé çàêëàäêè ýòîé ñòðóêòóðû, êàê îïèñûâàë
Ê. Áåêñòðîì (Bäckström, 1931). Íà ñòàäèè 28+ ïîñëå îáðàçîâàíèÿ òðàáåêóëÿðíîé
ïëàñòèíêè, îò íåå íà÷èíàåòñÿ îõðÿùåâåíèå íîñîâîé ïåðåãîðîäêè. Òàêèì îáðà-
çîì, íàøè äàííûå ïîäòâåðæäàþò ìíåíèå áîëüøèíñòâà àâòîðîâ, ÷òî íîñîâàÿ
ïåðåãîðîäêà ÿâëÿåòñÿ îðàëüíûì ïðîäîëæåíèåì ñëèòûõ ïåðåäíèõ êîíöîâ òðàáå-
êóë è íå èìååò ñàìîñòîÿòåëüíîãî î÷àãà îõðÿùåâåíèÿ (Kamal, Hammouda, 1965 a,
b, c; El-Toubi et al., 1970; El-Toubi et al., 1973).
Íîñîâ àÿ ð àêîâèíà. Íà 30-é ñòàäèè âåíòðîàáîðàëüíî îò íîñîâîé ðàêî-
âèíû îòõîäèò õðÿùåâîé îòðîñòîê, êîòîðûé ñëèâàåòñÿ ñ îðàëüíûì êðàåì planum
antorbitale. Êàê âèäíî èç ãðàôè÷åñêèõ ðåêîíñòðóêöèé ÷åðåïà, òàêîé îòðîñòîê áûë
255Ðàçâèòèå ýòìîèäíîãî îòäåëà õðÿùåâîãî ÷åðåïà â ýìáðèîãåíåçå óæà îáûêíîâåííîãî…
Ðèñ. 9. Òîòàëüíûé ïðåïàðàò ýòìîèäíîãî îòäåëà N. natrix: a – ëåâàÿ íîñîâàÿ êàïñóëà ýìáðèîíà 33-é
ñò. ð.; âèä ñ âåíòðàëüíîé ñòîðîíû: 1 – cupola anterior; 2 – processus alaris inferior; 3 – õðÿù îðãàíà
ßêîáñîíà; 4 – ýêòîõîàíàëüíûé õðÿù; 5 – ãèïîõîàíàëüíûé õðÿù; 6 – ïðåäìàêñèëëà; 7 – ñîøíèê;
8 – ñåïòîìàêñèëëà; b – Ëåâàÿ íîñîâà êàïñóëà ýìáðèîíà 34-é ñòàäèè ðàçâèòèÿ; âèä ñ äîðñàëüíîé ñòî-
ðîíû (äíî íîñîâîé êàïñóëû îòâåäåíî â ñòîðîíó): 1 – cupola anterior; 2 – processus alaris inferior; 3 –
paries lateralis nasi; 4 – õðÿù îðãàíà ßêîáñîíà; 5 – ýêòîõîàíàëüíûé õðÿù; 6 – ãèïîõîàíàëüíûé õðÿù;
7 – ýêòîõîàíàëüíàÿ ïëàñòèíêà; 8 – íîñîâàÿ ðàêîâèíà; 9 – planum antorbitale; 10 – fenestra olfacto-
ria advehens; 11 – íîñîâàÿ êîñòü; 12 – ïðåäìàêñèëëà.
Fig. 9. Cleared and stained ethmoidal region (left nasal capsule) of N. natrix embryo: a – at the 33 stage of
development from the ventral view: 1 – cupola anterior; 2 – processus alaris inferior; 3 – cartilage of
Jacobson‘s organ; 4 – ectochoanal cartilage; 5 – hypochoanal cartilage; 6 – premaxilla; 7 – vomer; 8 –
septomaxilla; b – at the 34 stage of development from the dorsal view (solum nasi is turned over to the side):
1 – cupola anterior; 2 – processus alaris inferior; 3 – paries lateralis nasi; 4 – cartilage of Jacobson‘s organ;
5 – ectochoanal cartilage; 6 – hypochoanal cartilage; 7 – ectochoanal plate; 8 – nasal concha; 9 – planum
antorbitale; 10 – fenestra olfactoria advehens; 11 – nasal bone; 12 – premaxilla.
îáíàðóæåí ðàíüøå ó N. natrix (Bäckström, 1931) è íåêîòîðûõ äðóãèõ çìåé (Pringle,
1954; Kamal, Hammouda, 1965 à; El-Toubi et al., 1973).
Íà 33-é ñòàäèè âåíòðîëàòåðàëüíûé êðàé èçãèáà paries lateralis nasi óòîëùàåò-
ñÿ, îáðàçóÿ îòðîñòîê, îïèñàííûé Ê. Áåêñòðîìîì (Bäckström, 1931) êàê processus
lateralis nasi. Òàêîé æå îòðîñòîê îïèñàí ó Cerastes vipera (Viperidae) (Kamal,
Hammouda, 1965 ñ) è íàçâàí ïàðàíàçàëüíûì îòðîñòêîì (paranasal cartilage), òàê
êàê èññëåäîâàòåëè ñ÷èòàëè åãî ãîìîëîãîì ïàðàíàçàëüíîãî õðÿùà ÿùåðèö.
Zona a nnu l a r i s. Ïîä òåðìèíîì «zona annularis» Ý. Ãàóïï (1906) ïîäðàçó-
ìåâàë ïîëîñòü íîñîâîé êàïñóëû, îêðóæåííóþ õðÿùåâûìè ñòðóêòóðàìè.
Íàøè èññëåäîâàíèÿ ïîäòâåðæäàþò äàííûå Ê. Áåêñòðîìà îòíîñèòåëüíî
ñóùåñòâîâàíèÿ zona annularis ó N. natrix. Íà ñòàäèè 31+ âèäíî, ÷òî ïîëîñòü êàï-
ñóëû îêðóæåíà ñî âñåõ ñòîðîí õðÿùåâûìè ñòðóêòóðàìè: äîðñàëüíî – paries late-
ralis nasi, ëàòåðàëüíî – õðÿùåâûì ñîåäèíåíèåì ìåæäó ìåæäó paries lateralis nasi
è õðÿùîì îðãàíà ßêîáñîíà, âåíòðàëüíî – õðÿùîì îðãàíà ßêîáñîíà è lamina
transversalis anterior, ìåäèàëüíî – íîñîâîé ïåðåãîðîäêîé.
Õðÿù îð ã àí à ßêîáñîíà è ý ê òî õî àíàë üíûé õ ðÿù. Íàøè äàííûå
ïîäòâåðæäàþò ðåçóëüòàòû Ê. Áåêñòðîìà (Bäckström, 1931): ýêòîõîàíàëüíûé õðÿù
ó N. natrix, êàê è ó áîëüøèíñòâà èçó÷åííûõ çìåé, ÿâëÿåòñÿ àáîðàëüíûì ïðîäîë-
æåíèåì õðÿùà îðãàíà ßêîáñîíà (ïîÿâëÿåòñÿ íà 31-é ñòàäèè) è íå èìååò îòäåëü-
íîãî î÷àãà «îõðÿùåâåíèÿ». Ýòî îïðîâåðãàåò äàííûå Â. Ê. Ïàðêåðà è (Parker,
1878) è Ã. Áîðíà (Born, 1883) îòíîñèòåëüíî ñàìîñòîÿòåëüíîñòè çàêëàäêè ýòîé
ñòðóêòóðû.
Ãèïîõî àíàë üíûé õ ðÿù. Ìû îáíàðóæèëè ñàìîñòîÿòåëüíóþ çàêëàäêó
ýòîé ñòðóêòóðû íà 30-é ñòàäèè. Íà ýòîé æå ñòàäèè â ñðåäèííîé åå ÷àñòè çàìåòåí
î÷àã îõðÿùåâåíèÿ. Ñëèÿíèå õðÿùåâûõ àáîðàëüíûõ êîíöîâ ýêòî- è ãèïîõîàíàëü-
íûõ õðÿùåé ñ îáðàçîâàíèåì ýêòîõîàíàëüíîé ïëàñòèíêè ïðîèñõîäèò íà 34-é ñòà-
äèè.
Òàêèì îáðàçîì, íàøè ðåçóëüòàòû íå ïîäòâåðæäàþò ëèòåðàòóðíûõ äàííûõ
îòíîñèòåëüíî îäíîãî î÷àãà îõðÿùåâåíèÿ ýêòîõîàíàëüíîãî, ãèïîõîàíàëüíîãî õðÿ-
ùåé è ýêòîõîàíàëüíîé ïëàñòèíêè (Kamal, Hammouda, 1965 a, b, c; El-Toubi et al.,
1970; El-Toubi et al., 1973; Deiques, 2004).
Ð l a num an t o r b i t a l e (Lamina orbitonasalis). Ïî ëèòåðàòóðíûì äàííûì,
planum antorbitale ó çìåé ÿâëÿåòñÿ àáîðàëüíûì ó÷àñòêîì ïàðèåòîòåêòàëüíîãî
õðÿùà è íå èìååò îòäåëüíîãî î÷àãà îõðÿùåâåíèÿ (Bäckström 1931; Kamal,
Hammouda, 1965 a, b, c; Kamal et al., 1970; El-Toubi et al., 1970; El-Toubi et al.,
1973). Ïî íàøèì äàííûì, planum antorbitale ó N. natrix çàêëàäûâàåòñÿ îòäåëüíî
îò ïàðèåòîòåêòàëüíîãî õðÿùà íà 27-é ñòàäèè, ñëèâàåòñÿ ñ íèì åùå íà ñòàäèè
ìåçåíõèìíîé çàêëàäêè (ñòàäèÿ 28+) è èìååò ñàìîñòîÿòåëüíûé î÷àã îõðÿùåâåíèÿ
(ñòàäèÿ 29+).  ëèòåðàòóðå åäèíñòâåííûé ñëó÷àé îáíàðóæåíèÿ ñàìîñòîÿòåëüíî-
ãî î÷àãà îõðÿùåâåíèÿ àáîðàëüíîãî ó÷àñòêà ïàðèåòîòåêòàëüíîãî õðÿùà ó çìåé áûë
îïèñàí ïî Malpolon monspessulana (Colubridae) (El-Toubi et al., 1973). Ïî óòâåð-
æäåíèþ ýòèõ àâòîðîâ óêàçàííûé ó÷àñòîê íå ÿâëÿåòñÿ planum antorbitale.
Ñ÷èòàåòñÿ, ÷òî ðlanum antorbitale çìåé ëèøåíà processus maxillaris anterior
(Parker, 1878; Bäckström, 1931; Pringle, 1954; El-Toubi et al., 1970; El-Toubi et al.,
1973; Haluska, Alberch, 1983; Deiques, 2004). Èñõîäÿ èç òîïîãðàôèè è ïðîèñõîæ-
äåíèÿ, ñ÷èòàåì, ÷òî îòðîñòîê, êîòîðûé ïîÿâëÿåòñÿ íà 30-é ñòàäèè îò îðàëüíî-
âåíòðàëüíîé ïîâåðõíîñòè planum antorbitale è òÿíåòñÿ â îðàëüíîì íàïðàâëåíèè,
ïðåäñòàâëÿåò ñîáîé processus maxillaris anterior. Ýòîò îòðîñòîê î÷åíü ïîõîæ íà
òàêîâîé, îïèñàííûé Ä. À. Ïðèíãëîì ó Dasypeltis scaber (Colubridae) (Pringle,
1954). Îäíàêî Ä. À. Ïðèíãë íå îáíàðóæèë ñâÿçè ýòîãî îòðîñòêà ñ planum antor-
bitale è íàçâàë åãî «posterior process of concha nasalå». Ýòîò îòðîñòîê íàçâàí èì
ïàðàíàçàëüíûì. Èç ðåêîíñòðóêöèé, ñäåëàííûõ Ê. Áåêñòðîìîì (Bäckström, 1931)
256 À. Â. Øåâåðäþêîâà
è Ì. Ð. Åë-Òîóáè ñ ñîàâòîðàìè (El-Toubi et al., 1973), âèäíî, ÷òî processus maxil-
laris anterior, âåðîÿòíî, åñòü è ó îïèñàííûõ èìè îáúåêòîâ. Ñóäÿ ïî âñåìó, îíè
îáíàðóæèëè ñâÿçü «posterior process of concha nasalis» ñ planum antorbitale, íî íå
èäåíòèôèöèðîâàëè ïîñëåäíþþ êàê òàêîâóþ. Ñòðóêòóðó, îò êîòîðîé îòõîäèò pro-
cessus maxillaris anterior îíè îïðåäåëèëè êàê «concha» (Bäckström, 1931) è «poste-
rior process of concha nasalis» (El-Toubi et al., 1973). Òàêèå ðàçíîãëàñèÿ, âåðîÿòíî,
ìîæíî îáúÿñíèòü çíà÷èòåëüíîé òîíêîñòüþ õðÿùåâûõ ñòðóêòóð àáîðàëüíîé ÷àñòè
ýòìîèäíîãî îòäåëà (ðèñ. 8).
Ëèòåðàòóðíûå äàííûå îòíîñèòåëüíî êîíòàêòà planum antorbitale è àáîðàëü-
íîé ÷àñòè íîñîâîé ïåðåãîðîäêè ïðîòèâîðå÷èâû. Ê. Áåêñòðîì (Bäckström, 1931)
îòðèöàë ñëèÿíèå ýòèõ ñòðóêòóð ó N. natrix; íà ðåêîíñòðóêöèÿõ, ñäåëàííûõ
Î. Â. ×åêàíîâñêîé (1936), âèäíî ñîåäèíåíèå ýòèõ ñòðóêòóð åùå íà ðàííèõ ñòà-
äèÿõ ðàçâèòèÿ. Íà íàøèõ ïðåïàðàòàõ âèäíî, ÷òî planum antorbitale ñëèâàåòñÿ ñ
òðàáåêóëÿðíîé ïëàñòèíêîé (ðèñ. 2, a, b; 7). Èç-çà íåðàâíîìåðíîãî ïðîöåññà
îõðÿùåâåíèÿ çàêëàäêè äîðñàëüíîãî ó÷àñòêà planum antorbitale ñáëèæåíèå åå ñ
òðàáåêóëÿðíîé ïëàñòèíêîé ïðîèñõîäèò ïîñòåïåííî, à èõ ñëèÿíèå – äîñòàòî÷íî
ïîçäíî – íà 35-é ñòàäèè ðàçâèòèÿ. Äàííûå îòíîñèòåëüíî ñëèÿíèÿ ýòèõ ñòðóêòóð
ó äðóãèõ èññëåäîâàííûõ çìåé òàêæå ïðîòèâîðå÷èâû. Ïðè÷èíîé ïðîòèâîðå÷èé,
î÷åâèäíî, ÿâëÿåòñÿ òî, ÷òî àâòîðàìè áûëè èññëåäîâàíû ýìáðèîíû ðàçíûõ ñòàäèé
ðàçâèòèÿ.
Ðîìåéñ Á. Ìèêðîñêîïè÷åñêàÿ òåõíèêà. – Ì. : Èçä-âî èíîñòð. ëèò., 1953. – 720 ñ.
×åêàíîâñêàÿ Î. Â. Ðàçâèòèå ÷åðåïà óæà (Tropidonotus natrix) // Àðõèâ àíàòîìèè, ãèñòîëîãèè è
ýìáðèîëîãèè. – 1936. – 15, âûï. 3. – Ñ. 3—33.
Bäckström K. Rekonstruktionsbilder zur Ontogenie des Koopfskelettes von Tropidonotus natrix // Acta Zool.
(Stock.). – 1931. – 12. – P. 83—144.
Deiques C. H. On the ontogeny of the ethmoidal region of Bothrops jararaca and Crotolus durissus terrificus
(Serpentes, Viperidae), and its phylogenetic significance // Acta Biologica Leopoldensia. – 2004. –
26, N 2. – P. 285—298.
El-Toubi M. R., Kamal A. M., Mokhtar F. M. The chondrocranium of late embryos of Egyptian Cobra, Naja
naje // Anat. Anz. – 1970. – 127. – P. 233—289.
El-Toubi M. R., Kamal A. M., Zaher M. M. The development of the chondrocranium of the snake, Malpolon
monspessulana. II. The fully formed stage // Acta anat. – 1973. – 85. – P. 593—619.
Haluska F., Alberch P. The cranial development of Elaphe obsolete (Ophidia, Colubridae) // J. Morph. –
1983. – 178. – P. 37—55.
Kamal A. M., Hammouda H. G. The development of the skull of Psammophis sibilans. II. The fully formed
chondrocranium // J. Morph. – 1965 a. – 116. – P. 247—296.
Kamal A. M., Hammouda H. G. The chondrocranium of the snake Eryx jaculus // Acta Zool. – 1965 b. –
46. – P. 176—208.
Kamal A. M., Hammouda H. G. Observations on the chondrocranium of the snake, Cerastes vipera // Morph.
Jb. – 1965 c. – 107. – P. 58—98.
Parker W. K. On the structure and development of the skull in the common snake (Tropidonotus natrix) //
Phil. Trans. Royal Soc. London. – 1878. – 169. – P. 385—417.
Pringle J. A. The cranial development of certain South African snakes and the relationship of these groups //
Proc. Zool. Soc. London – 1954. – 123. – P. 813—865.
Simons E. V., Van Horn J. R. A new procedure for whole-mount alcian blue staining of the cartilaginous
skeleton of chicken embryos, adapted to the clearing procedure in potassium hydroxide // Acta
Morphol. Neerl. – Scand. – 1971. – 8. – P. 281—292.
Zehr D. R. Stages in the normal development of the common garter Snake Thamnophis sirtalis sirtalis //
Copeia. – 1962. – 2. – P. 322—329.
257Ðàçâèòèå ýòìîèäíîãî îòäåëà õðÿùåâîãî ÷åðåïà â ýìáðèîãåíåçå óæà îáûêíîâåííîãî…
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/ColorImageDict <<
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>>
/JPEG2000ColorACSImageDict <<
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>>
/JPEG2000ColorImageDict <<
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>>
/AntiAliasGrayImages false
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/GrayImageDownsampleType /Bicubic
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/GrayImageDepth -1
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/GrayImageFilter /DCTEncode
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/GrayACSImageDict <<
/QFactor 0.15
/HSamples [1 1 1 1] /VSamples [1 1 1 1]
>>
/GrayImageDict <<
/QFactor 0.15
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>>
/JPEG2000GrayACSImageDict <<
/TileWidth 256
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>>
/JPEG2000GrayImageDict <<
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>>
/AntiAliasMonoImages false
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/MonoImageFilter /CCITTFaxEncode
/MonoImageDict <<
/K -1
>>
/AllowPSXObjects false
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/None
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0.00000
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0.00000
]
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0.00000
0.00000
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/NLD (Gebruik deze instellingen om Adobe PDF-documenten te maken die zijn geoptimaliseerd voor prepress-afdrukken van hoge kwaliteit. De gemaakte PDF-documenten kunnen worden geopend met Acrobat en Adobe Reader 5.0 en hoger.)
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/ENU (Use these settings to create Adobe PDF documents best suited for high-quality prepress printing. Created PDF documents can be opened with Acrobat and Adobe Reader 5.0 and later.)
>>
/Namespace [
(Adobe)
(Common)
(1.0)
]
/OtherNamespaces [
<<
/AsReaderSpreads false
/CropImagesToFrames true
/ErrorControl /WarnAndContinue
/FlattenerIgnoreSpreadOverrides false
/IncludeGuidesGrids false
/IncludeNonPrinting false
/IncludeSlug false
/Namespace [
(Adobe)
(InDesign)
(4.0)
]
/OmitPlacedBitmaps false
/OmitPlacedEPS false
/OmitPlacedPDF false
/SimulateOverprint /Legacy
>>
<<
/AddBleedMarks false
/AddColorBars false
/AddCropMarks false
/AddPageInfo false
/AddRegMarks false
/ConvertColors /ConvertToCMYK
/DestinationProfileName ()
/DestinationProfileSelector /DocumentCMYK
/Downsample16BitImages true
/FlattenerPreset <<
/PresetSelector /MediumResolution
>>
/FormElements false
/GenerateStructure false
/IncludeBookmarks false
/IncludeHyperlinks false
/IncludeInteractive false
/IncludeLayers false
/IncludeProfiles false
/MultimediaHandling /UseObjectSettings
/Namespace [
(Adobe)
(CreativeSuite)
(2.0)
]
/PDFXOutputIntentProfileSelector /DocumentCMYK
/PreserveEditing true
/UntaggedCMYKHandling /LeaveUntagged
/UntaggedRGBHandling /UseDocumentProfile
/UseDocumentBleed false
>>
]
>> setdistillerparams
<<
/HWResolution [2400 2400]
/PageSize [612.000 792.000]
>> setpagedevice
|