Â. Ò. Õìóð÷èê1, Ñ. Ì. Áëèíîâ2
1 Èíñòèòóò ýêîëîãèè è ãåíåòèêè ìèêðîîðãàíèçìîâ ÓðÎ ÐÀÍ,
2 Ïåðìñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò, Ïåðìü, Ðîññèÿ
ïî ñêëîíó. Íèæíÿÿ ÷àñòü îòëîæåíèé, ìîùíîñòüþ äî 50 ñì, ïðåäñòàâëåíà ÿðîçèòîì, âåðõíÿÿ — òåìíî-áóðû-ìè ïîðèñòûòåìíî-áóðû-ìè êîðàòåìíî-áóðû-ìè ãèäðîêñèäîâ æåëåçà [4].
 ðåçóëüòàòå èññëåäîâàíèé èçó÷åíû ãåîëîãî-ãèä-ðîãåîëîãè÷åñêèå óñëîâèÿ ïîëÿ øàõòû “Áåëûé ñïîé”, ãèäðîäèíàìè÷åñêèé, òåìïåðàòóðíûé è ãèäðîõèìè÷åñ-êèé ðåæèìû ñàìîèçëèâà, âûïîëíåíî èññëåäîâàíèå ìîðôîëîãè÷åñêèõ, ìèíåðàëîãè÷åñêèõ è ãåîõèìè÷åñ-êèõ îñîáåííîñòåé æåëåçèñòûõ ïëîòèí, äàíà õàðàêòå-ðèñòèêà óñëîâèé èõ ôîðìèðîâàíèÿ è ðàçâèòèÿ, âûÿâ-ëåíà ðîëü ãåîõèìè÷åñêèõ áàðüåðîâ â êîíöåíòðàöèè õèìè÷åñêèõ ýëåìåíòîâ è ó÷àñòèå â ýòèõ ïðîöåññàõ ìèêðîîðãàíèçìîâ è ðàñòèòåëüíîñòè. Óñòàíîâëåíî, ÷òî ôîðìèðîâàíèå îòëîæåíèé ïðîèñõîäèò íà ãåîõèìè÷åñ-êèõ áàðüåðàõ, ñðåäè êîòîðûõ îñíîâíóþ ðîëü èãðàþò áèîãåîõèìè÷åñêèå (ðàáîòà ìèêðîîðãàíèçìîâ, âîäîðîñ-ëåé è äð.) è êèñëîðîäíûé. Îñíîâíûìè ôàêòîðàìè ôîð-ìèðîâàíèÿ îòëîæåíèé ÿâëÿþòñÿ îñîáåííîñòè õèìè÷åñ-êîãî ñîñòàâà øàõòíûõ âîä, õàðàêòåð ðåëüåôà, ìèêðî-áèîëîãè÷åñêèå îñîáåííîñòè è íàëè÷èå è âèäîâîé ñî-ñòàâ âîäîðîñëåé.
Ëèòåðàòóðà
1. Áàëàøîâà Â. Â. Ìèêîïëàçìû è æåëåçîáàêòåðèè. Ì.:
Íàóêà, 1974. 65 ñ.
2. Baker B. J., Banfield J. F. Microbial communities in acid mine drainage // FEMS Microbiol. Ecol., 2003. ¹ 44. Ð. 139—
152.
3. Edwards K. J., Gihring T. M., Banfield J. F. Seasonal variations in microbial populations and environmental conditions in an extreme acid mine drainage environment // Appl. Environ.
Microbiol., 1999. ¹ 65. Ð. 3627—3632.
4. Ïîòàïîâ Ñ. Ñ., Áëèíîâ Ñ. Ì. Ãåîýêîëîãè÷åñêàÿ ñèòóà-öèÿ â Êèçåëîâñêîì óãîëüíîì áàññåéíå íà îñíîâå èçó÷åíèÿ òåõíîãåííûõ ìèíåðàëèçàöèé // Óðàëüñêèé ìèíåðàëîãè÷åñ-êèé ñáîðíèê, 2002. ¹ 12. Ñ. 204—219.
Îäíèìè èç èíòåðåñíåéøèõ ñêåëåòíûõ ýëåìåíòîâ ïàëåîçîéñêèõ êîðàëëîâ ðóãîç ÿâëÿþòñÿ òàê íàçûâàå-ìûå “ ñåïòàëüíûå êîíóñû” èëè, áîëåå ïðàâèëüíî, “ñòå-ðåîïëàçìàòè÷åñêèå êîðêè”–ïåðèîäè÷åñêè îòëàãàâøè-åñÿ áàçàëüíîé ïîâåðõíîñòüþ ïîëèïà èëè åå ÷àñòüþ ñëîè ñòåðåîïëàçìû (ñêëåðåíõèìû).  íàñòîÿùåå âðå-ìÿ ñîñòàâ êîðîê, êàê è ñêåëåòà èñêîïàåìûõ ïàëåîçîéñ-êèõ êîðàëëîâ â öåëîì, êàëüöèòîâûé, õîòÿ íå èñêëþ÷å-íî, ÷òî ïåðâè÷íûé èõ ñîñòàâ áûë àðàãîíèòîâûì. Ñòå-ðåîïëàçìàòè÷åñêèå êîðêè íàèáîëåå âûðàçèòåëüíî ðàç-âèòû ó äåâîíñêèõ ðóãîç-öèñòèôèëëèä. Ìîðôîëîãèþ èõ ñòåðåîïëàçìàòè÷åñêèõ êîðîê èññëåäîâàëè Ð. Âåäåêèíä, Ê. Ã. Âîéíîâñêèé-Êðèãåð, Å. Ä. Ñîøêèíà, Ð. Áèðåíõåéä è àâòîð ñòàòüè [1—6].
Ó ïðåäñòàâèòåëåé ðàçíûõ âèäîâ, ðîäîâ è ñå-ìåéñòâ ðóãîç ñòåðåîïëàçìàòè÷åñêèå êîðêè ðàçâèòû ïî-ðàçíîìó: ó îäíèõ îíè áîëåå èëè ìåíåå ðàâíîìåð-íî ïåðåñåêàþò âñþ ïîëîñòü êîðàëëèòà, ó äðóãèõ ðàç-âèòû ëîêàëüíî. Íà ýòîì îñíîâàíèè ìíîé áûëà ïðåä-ëîæåíà êëàññèôèêàöèÿ êîðîê ïî ñòåïåíè èõ ðàçâè-òèÿ è ïðèóðî÷åííîñòè [5, 6]. Âñå
ñòåðåîïëàçìàòè-÷åñêèå êîðêè áûëè ðàçäåëåíû íà äâà îñíîâíûõ òèïà:
ïîëíûå (ðèñ. 1, à), è íåïîëíûå (ðèñ. 1, b, ñ). Ïîëíûå êîðêè ñåêðåòèðîâàëèñü âñåé áàçàëüíîé ýêòîäåðìîé ïîëèïîâ. Ñðåäè íåïîëíûõ êîðîê, â ñâîþ î÷åðåäü, âûäåëåíû äåñöåíäåíòíûå (ðèñ. 1, b), è àñöåíäåíò-íûå (ðèñ. 1, ñ). Ïåðâûå ÿâëÿþòñÿ ðåçóëüòàòîì ñîîò-âåòñòâóþùèõ ïðîöåññîâ â ïðîêñèìàëüíîé ÷àñòè ýê-òîäåðìû, à àñöåíäåíòíûå–ïðîèçâîäíûå äèñòàëüíîé
÷àñòè ýêòîäåðìû.
 íàñòîÿùåå âðåìÿ â îñíîâíîì ðåøåí âîïðîñ î ïóòÿõ ïîñòóïëåíèÿ êàëüöèÿ è óãëåêèñëîòû, íåîáõî-äèìûõ äëÿ ïîñòðîåíèÿ ñêåëåòà ñîâðåìåííûìè êîðàë-ëàìè. Íàèáîëåå äåòàëüíî ñõåìà êàëüöèôèêàöèîííî-ãî îáìåíà ó êîðàëëîâ ïðåäñòàâëåíà Ê. Êðîññëåíäîì è Ä. Áàðíåññîì (ðèñ. 2). Ïðåäïîëàãàåòñÿ, ÷òî ïîäîá-íàÿ æå ñõåìà ïîñòóïëåíèÿ êàëüöèÿ äëÿ ñòðîèòåëüñòâà ñêåëåòà ñóùåñòâîâàëà è ó èñêîïàåìûõ êîðàëëîâ.
ÌÎÐÔÎËÎÃÈß È ÃÅÍÅÇÈÑ ÑÒÅÐÅÎÏËÀÇÌÀÒÈ×ÅÑÊÈÕ ÊÎÐÎÊ Ó ÏÀËÅÎÇÎÉÑÊÈÕ ÊÎÐÀËËΠÐÓÃÎÇ
Â. Ñ. Öûãàíêî
Èíñòèòóò ãåîëîãèè Êîìè ÍÖ Óðî ÐÀÍ, Ñûêòûâêàð, Ðîññèÿ
Âîçâðàùàÿñü ê îáúåêòó íàøåãî ðàññìîòðåíèÿ, ñëå-äóåò îòìåòèòü, ÷òî äëÿ âûÿñíåíèÿ òàêñîíîìè÷åñêîãî çíà÷åíèÿ ñòåðåîïëàçìàòè÷åñêèõ êîðîê âàæíûì ïðåä-ñòàâëÿåòñÿ èõ ãåíåçèñ. Ïîñëåäíèé òðàêòóåòñÿ èññëåäî-âàòåëÿìè ïî-ðàçíîìó. Îäíè èññëåäîâàòåëè, òàêèå êàê Ò. Ìà [7] è Í. ß. Ñïàññêèé [8], ñâÿçûâàþò îáðàçîâàíèå êîðîê ñ ñåçîííîñòüþ: â îñåííåå–çèìíèé ïåðèîä ïðîèñ-õîäèò çàìåäëåíèå ðîñòà ñêåëåòà êîðàëëà è âìåñòî ïó-çûð÷àòîé òêàíè øëî îáðàçîâàíèå ïëîòíîé òêàíè êàëü-öèòîâîé êîðêè. Ê. Ã. Âîéíîâñêèé-Êðèãåð [2] âûñêàçàë ìûñëü î ñâÿçè ãåíåçèñà êîðîê ñ ïåðèîäè÷íîñòüþ íàêîï-ëåíèÿ èëà. Ìíîé áûëî ñäåëàíî ïðåäïîëîæåíèå î ñâÿçè ñòåðåîïëàçìàòè÷åñêèõ êîðîê ñ ãàìåòîãåíåçîì â ïåðèîä ïîëîâîãî ðàçìíîæåíèÿ ïîëèïîâ [5, 6].
Ñîãëàñíî ìíîãî÷èñëåííûì íàáëþäåíèÿì èññëåäî-âàòåëåé íàä äðóãèìè ãðóïïàìè îðãàíèçìîâ, ïåðèîä ïîëîâîãî ðàçìíîæåíèÿ õàðàêòåðèçóåòñÿ ó íèõ ñóùå-ñòâåííûìè èçìåíåíèÿìè â ìåòàáîëèçìå è
ôèçèîëîãè-÷åñêèõ ïðîöåññàõ. Ãàìåòîãåíåç è ñëåäóþùèå çà íèì ïîëîâîå ðàçìíîæåíèå òðåáóþò áîëåå èíòåíñèâíûõ ïðîöåññîâ ðîñòà è ðàçìíîæåíèÿ êëåòîê, ÷òî îáû÷íî ñâÿçàíî ñ óñèëåíèåì îáìåíà âåùåñòâ â îðãàíèçìå.
Ë. Ãåéëüáðóí [9] ïîäìåòèë, ÷òî ñêîðîñòü âõîæäåíèÿ â êëåòêè îðãàíèçìîâ èîíîâ Ñà+2 ïîâûøàåòñÿ ïðè ïåðå-õîäå êëåòîê â àêòèâíîå ñîñòîÿíèå (âîçáóæäåíèå, îï-ëîäîòâîðåíèå è ò. ï.). Ïðîöåññû æå ãàìåòîãåíåçà ñî-ïðÿæåíû ñ èíòåíñèâíûì ðàçìíîæåíèåì êëåòîê è èõ ïîñëåäóþùèìè ñëîæíûìè ïðåâðàùåíèÿìè. Ñ òàêèìè ïðîöåññàìè, ïðîèñõîäèâøèìè â îðãàíèçìàõ ïàëåîçîé-ñêèõ ïîëèïîâ, è ñâÿçûâàåòñÿ îáðàçîâàíèå ðàññìàòðè-âàåìûõ ñòåðåîïëàçìàòè÷åñêèõ êîðîê. Ê ñîæàëåíèþ, Ðèñ. 1. Òèïû ñòåðåîïëàçìàòè÷åñêèõ êîðîê.
à — ïîëíûå; â, ñ — íåïîëíûå: â — äåñöåíäåíòíûå, ñ — àñöåí-äåíòíûå
Fig. 1. Typen of stereoplasmatic crust.
a — complete, b, c — incomplete: b — descendent, c — ascendent
Ðèñ. 2. Ñõåìà êàëüöèôèêàöèîííîãî îáìåíà ó êîðàëëîâ (ïî: Crossland and Barnes, 1974)
Fig. 2. Scheme of calcificacion exchange from corals (by: Crossland and Barnes, 1974)
ýêñïåðèìåíò íàä ïàëåîçîéñêèìè êîðàëëàìè äëÿ ïîä-òâåðæäåíèÿ âûñêàçàííîé òî÷êè çðåíèÿ ïî îáúåêòèâ-íûì ïðè÷èíàì íåâîçìîæåí. Ïðàêòè÷åñêè íåò òàêæå èññëåäîâàíèé â îòíîøåíèè ñâÿçè ìåæäó ðàçìíîæåíè-åì è ñêîðîñòüþ ñêåëåòîãåíåçèñà è ó ñîâððàçìíîæåíè-åìåííûõ êî-ðàëëîâûõ ïîëèïîâ. Åäèíñòâåííîå íàáëþäåíèå â ýòîì ïëàíå áûëî ñäåëàíî â 1935 ã. Ì. À. Êðåìïôîì [10], êîòîðûé îòìåòèë ñâÿçü ïåðèîäè÷íîñòè â îáðàçîâàíèè ñêåëåòà ó ñîâðåìåííûõ êîðàëëîâ ðîäà Favia ñ ïåðèî-äàìè ïîëîâîé àêòèâíîñòè îðãàíèçìà. Ïðè ýòîì îí ëèøü êîíñòàòèðîâàë, ÷òî îáà ÿâëåíèÿ ñâÿçàíû ìåæäó ñîáîé ïðîñòûì öèòîëîãè÷åñêèì ìåõàíèçìîì. Êîñâåí-íûì ïîäòâåðæäåíèåì âûñêàçàííîé çäåñü òî÷êè çðåíèÿ ìîãóò ñëóæèòü íàáëþäåíèÿ íàä ãàìåòîãåíåçîì ó íåêî-òîðûõ ïðåäñòàâèòåëåé ñîâðåìåííûõ áåñïîçâîíî÷íûõ.
 ÷àñòíîñòè, íàáëþäåíèÿ Áå [11] íàä ãàìåòîãåííîé
êàëüöèôèêàöèåé ó ïëàíêòîííûõ ôîðàìèíèôåð Gloobigerinoides sacculifer (Brady) ïîêàçàëè óâåëè÷å-íèå âåñà ïîñòãàìåòîãåííîé ðàêîâèíû â ñðåäíåì íà 28 % ïî ñðàâíåíèþ ñ ïðåäãàìåòîãåííîé (ðèñ. 3).
Åñëè ñëåäîâàòü àíàëîãèÿì, òî íåëüçÿ èñêëþ÷èòü ñóùåñòâîâàíèå ïîäîáíûõ ïðîöåññîâ è ó èñêîïàåìûõ êîðàëëîâ ðóãîç. Òîò èëè èíîé òèï êîðîê áûë ñâÿçàí â ýòîì ñëó÷àå ñ óñèëåíèåì ñåêðåöèè êàëüöèòà ó÷àñòêà-ìè òåëà ïîëèïà, ãäå ïðîèñõîäèëà ãåíåðàöèÿ ãàìåò. Îò-ñþäà è ðàçëè÷íûå òèïû ñòåðåîïëàçìàòè÷åñêèõ êîðîê.
Ëèòåðàòóðà
1. Wedekind R. 1924. Das Mitteldevon der Eifel. Eine biostrat. Studie. T. 1. Die Tetracorallen des unteren Mitteldevon //
Schrift. Ges. Beford. Nat. Mardurg, Bd. 14(3).
2. Vojnovskij-Kriger C. 1930. Zur Kenntnis des inneren Baues der Korallen Rugosa (eine Kritik der Theorie der Septalkegel) // Èçâ. ÀÍ ÑÑÑÐ, VII ñåð., îòä. ôèç.-ìàò. íàóê,
¹ 4. Ñ. 311—330.
3. Ñîøêèíà Å. Ä. Äåâîíñêèå êîðàëëû Rugosa Óðàëà // Òð.
Ïàëåîíòîëîãè÷åñêîãî èí-òà ÀÍ ÑÑÑÐ, 1949. Ò. 15. Âûï. 4.
Ñ. 3—160
4. Birencheide R. Die “Cystimorpha” (Rugosa) aus dem Eifeler Devon // Abh. Senckenberg nature Ges., 1964. T. 507.
S. 1—120.
5. Öûãàíêî Â. Ñ. “Ñåïòàëüíûå êîíóñû” ÷åòûðåõëó÷åâûõ êîðàëëîâ è èõ ôóíêöèîíàëüíîå çíà÷åíèå // Ïàëåîíòîë. æóðí., 1972. ¹ 4. Ñ. 31—43.
6. Öûãàíêî Â. Ñ. Äåâîíñêèå êîðàëëû ðóãîçû ñåâåðà Óðà-ëà. Ë.: Íàóêà, 1981. 220 ñ.
7. Ma T. Y. H. On the seasonal growth in palaeozoic tetracorals and the climate during the Devonian period //
Palaeontol. Sinica, ser. B, 1937.V. 11. P. 3—97.
8. Ñïàññêèé Í. ß. Ïàëåîýêîëîãèÿ ÷åòûðåõëó÷åâûõ êîðàë-ëîâ // Ïàëåîíòîë. æóðí., 1967. ¹ 2. Ñ. 7—14.
9. Ãåéëüáðóí Ë. Äèíàìèêà æèâîé ïðîòîïëàçìû. Èçä-âî èíîñòð. ëèò., 1957. Ñ. 9—347.
10. Krempf M. A. Inscription maregraphique des cycles de retrogradation des noeuds de la Lune par certains coraux constructeurs de recifs // Compt. rend. Acad. Sci.. Paris, 1935.
T. 198. ¹19. P. 1708—1710.
11. Be A. W. H. Gametogenic calcification in a spinose planctonic foraminifer Glomigerinoides sacculifer (Brady) // Mar.
Micropaleontology, 1980. V. 5. N 3. P. 283—310.
Ðèñ. 3. Ñõåìàòè÷åñêîå ñå÷åíèå ñòåíêè è ïîðû Globigerinoides sacculifer, äåìîíñòðèðóþùåå ïîñëåäîâàòåëüíûé ðîñò ñòàäèé îíòîãåíåòè÷åñêîé êàëüöèôèêàöèè èçîáðàæåííîãî
îðãàíè-÷åñêîãî ñëîÿ. Ãàìåòîãåííàÿ êàëüöèôèêàöèÿ íà÷èíàåòñÿ â ìåñòå, ãäå çàêàí÷èâàåòñÿ øèï (ïî: Be, 1980) Fig. 3. Schematic drawing of wall and pore structure G. sacculifer, showing successive growth stages of ontogenetic calcification delineated by organic layers. Gametogenic calcification starts from
the point where spine is resorberi (Be, 1980)
One of the most interesting skeletal elements of paleo-zoic corals of rugose are so called “septal cones” and/or
“stereoplasmatic crusts”, layers of stereoplasm (scleren-chyma) periodically deposited by basal surface of a polyp or its part. “Septal cones” are most expressively devel-oped with the Devonian rugose-cystifillids. Their morphol-ogy is studied in detail R. Wedekind, K. H. Vojnovskij-Kriger, E. D. Soshkina, R. Birencheide et author this let-ter [1-6]. Nevertheless, while investigating the considered formations the author came to conclusion about absence of precise connection between “cones” and septal organ of rugoses and about appropriateness of using in the given case the term “stereoplasmatic crust” [5, 6]. At that the
“crust” classification was suggested, which was based on the completeness of its development and confinement to this or that area of supposed basal surface of a polyp. In particular, two basic types were determined — complete, isolated by the whole basal surface of a polyp, and incom-plete, divided into descendent “crusts” —a result of vital function of proximal part of ectoderm, and ascendant ones — derivatives of distal part of ectoderm (see the prev.
article in Russian, fig. 1).
To determine a taxonomic value of “crusts” the estab-lishment of their genesis is important, which is treated by researchers differently. One of them suppose seasonal cau-sality of their occurrence connected with slowdown of growth of corals in autumn-winter period forming dense calcite crusts instead of a bubbly tissue [7, 8], the others assume their connection with periodicity of silt accumula-tion [2]. Earlier the author supposed connecaccumula-tion of stereo-plasmatic crusts with gametogenesis during polyp sexual reproduction [5, 6]. In the given report I insists on the point of view stated earlier about the genesis of stereoplasmatic crusts proceeding from the following preconditions:
1. The classification of stereoplasmatic crusts given above does not allow to connect their formation with sea-sonality, as in this case we would always deal with com-plete crusts, which are more or less uniform by thickness and secreted by the whole surface of ectoderm.
2. A question about ways of calcium and carbon car-bonic acid transportation needed for skeleton building among modern corals is now mainly solved [9, 10]. The most detailed pattern of calcium metabolism of corals was presented by C. J. Crossland and D. J. Barnes (see the prev.
article in Russian, fig. 2). It is supposed that similar pat-tern of carbonic acid supply for skeleton building existed also among fossil corals.
3. At present data are available about strengthening calcite secretion during gametogenesis of some
represent-atives of modern invertebrates. In particular, observing gametogenous calcification of planktonic foraminifers Globigerinoides sacculifer (Brady) showed increase in weight of postgametogenous shell in 28 % in average in comparison to pregametogenous one [9]. If to follow anal-ogies, it is impossible to exclude existence of similar proc-esses of fossil corals of rugose too. This or that crust type was connected in this case with the intenficacion of cal-cite secretion at those areas of the polyp body where there was a generation of gametes (see the prev. article in Rus-sian, fig. 3).
4. The validity of the given concept is indirectly proved by the use of morphology of stereoplasmatic crusts as tax-onomic sign of rugoses [1, 3—6] et al.. In case of seasonal causality of crusts their taxonomic value bring to nought.
References
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