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134.Журнал Сибирского федерального университета. Сер. Биология №2 2011

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Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Журнал Сибирского федерального университета
2011
Journal of Siberian Federal University
4 (2)
Биология
Biology
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??.-?. ???, ?-? ???.-???.????
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??.-?. ???, ?-? ???.-???. ????
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?. ?. ????
CONTENTS / СОДЕРЖАНИЕ
Л.В. Волкова
Conioselinum tataricum Hoffm. (Apiaceae) , ??% ???%2,???*,?
C%?,?,, " ??!??"/. ???=. q=?=,!?*%?% *! ?=
? 107 ?
И.А. Кайгородова
c?3K%*%"%??= -=3?= %?,?%.?2 (Annelida, Oligochaeta) "
!=L%?? ?,?!%2?!?=???%?% ,?2%??,*= K3.2/ t!%?,.=, %?.
a=L*=? (q,K,!?, p%??, )
? 117 ?
Editorial Advisory Board
Chairman:
Eugene A. Vaganov
Members:
Josef J. Gitelzon
Vasily F. Shabanov
Andrey G. Degermendzhy
Valery L. Mironov
Gennady L. Pashkov
Vladimir V. Shaidurov
Vladimir V. Zuev
Editorial Board:
Editor-in-Chief:
Mikhail I. Gladyshev
Founding Editor:
Vladimir I. Kolmakov
Managing Editor:
Olga F. Alexandrova
В.М. Сафронов, Е.С. Захаров, А.П. Захаров
q%K%?? " ?=C=??/. %2!%?=. b?!.% ??*%?% .!?K2=
? 133 ?
В.В. Салмин, А.Б. Салмина,А.А. Фурсов,
О.В. Фролова, Д.И. Лалетин, М.А. Фурсов,
Г.В. Юдин, Н.А. Малиновская,
Л.Д. Зыкова, А.С. Проворов
h?C%???%"=?,? -?3%!?????2?%L ?C?*2!%?*%C,, ??
,???,???*%?% C%"!?????, ?,%*=!?=
%???*,
? 142 ?
Natalia O. Zhila, Tatiana G. Volova,
Elena D. Nikolaeva and Daria A. Syrvacheva
Microbial Synthesis and Characterization of
(3-Hydroxybutyrate-co-4-Hydroxybutyrate) Copolymers
Poly
? 158 ?
Executive Editor for Biology:
Nadezhda N.Sushchik
???????? ?.?. ?????? ????????? ?.?. ??????????
???????????? ??????? ?.?. ?????????
????????? ? ?????? 17.06.2011 ?. ?????? 84x108/16. ???. ???. ?. 8,45.
??.-???. ?. 7,95. ?????? ???. ?????? ????????. ????? 1000 ???. ????? 5235.
?????????? ? ?? ???. 660041 ??????????, ??. ?????????, 82?.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Editorial board for Biology:
Sergey I. Bartsev
Alexander Y. Bolsunovsky
Tatiana G. Volova
Eugene S. Vysotski
Nikolai A. Gaevsky
Egor S. Zadereev
Valentina A. Kratasyuk
Elena N. Muratova
J. Woodland Hastings
Frank D. Salisbury
Malcolm K. Hughes
Ernst-Detlef Schulze
Akira Osawa
Takayoshi Koike
Marc D?Alarcao
Mikhail G. Karpinsky
Liliana Zalizniak
А.А. Егоршина, Р.М. Хайруллин, М.А. Лукьянцев,
З.М. Курамшина, Ю.В. Смирнова
t%?-=2-?%K,?,?3??= =*2,"?%?2? .??%-,2?/. ?2=??%"
Bacillus subtilis , ,. "?, ?,? ?= ?2?C??? ?,*%!,?=?,, *%!??L
C???,?/
? 172 ?
И.В. Свидерская, В.Г. Суховольский,
Е.Ю. Радостева, А.В.Кирдянов
l%?????=
%???*= %C2,?=???%?% ?%%2?%???,
????3
2%??,?%L *??2%??%L ?2??*, , !=???!%? ?????= 3 2!=.?,?
."%L?/.
? 183 ?
М.Ю. Салтыков, С.И. Барцев, Ю.П. Ланкин
g=",?,?%?2? 3?2%L?,"%?2, ?%????L ?=?*?32/. .*%?,?2?? %2
?,??= ",?%"
????????????? ? ??????????? ???
?? ? ??77-28-725 ?? 29.06.2007 ?.
????? ???????? ? «???????? ??????? ????????????? ??????? ???????? ? ???????, ? ??????? ??????
???? ???????????? ???????? ??????? ?????????? ??????????? ??
????????? ?????? ??????? ??????? ?
????????? ????» (???????? 2010 ?.)
? 197 ?
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2011 4) 107-116
~~~
??? 581.522+581.412(571.1)
Conioselinum tataricum Hoffm. (Apiaceae)
? ??? ???????????? ???????
? ???????? ????? ??????????? ?????
?.?. ???????*
??????????? ????????? ???????????? ??? ?? ???,
?????? 630090, ???????????, ??. ???????????????, 101 1
Received 3.06.2011, received in revised form 10.06.2011, accepted 17.06.2011
??????? ????????? 11 ????????????? (??) Conioselinum tataricum Hoffm. (???. Apiaceae)
? ????????????? ??????????? ??????????? ????? (?? ???????? ??????). ?? ????
??????????????? ????????? ? ??????????? ????. ????????? ?? ??????????? ?? ??????
????????????? ?????. C. tataricum ???????????? ? ??????????? ??? ???????????? ???????.
??????????? ???????? ???? ? ????????? ??? ?? ??????????????? ? ????????? ?????????
C. tataricum ? ????????????? ? ????????????? ??????????? ???????.
???????? ?????: ?????????????, ?????????? ?????????, ?????????, ?????????.
????????
Conioselinum tataricum Hoffm. ? ??????????? ??????????? ?????????? ?? ????????? ????????? (Apiaceae). ????????????? ?
?????? ???? ?? ???? ??????? ??????, ? ???????? ? ????????? ??????, ? ?????? ???????
??????? ????. ? ????? ??????? ???????????
?? ??????? ??????? ????. ?????? ??????????? ? ??????????? ? ??????? ?????, ? ???????, ?????????? ? ?????? ?????. ??? ???????? ???????? ?????????. ???????????? ?
??????????? ????????????, ????????? ?
????????-???????? ?????, ?? ?? ???????,
???????, ?? ?????? ?????, ?? ??????? ?????-
*
1
???, ?????????? ???????, ???????? ?????;
????? ?????? ?????? ? ??????? (????? ???????? ??????, 1935; ????? ????, 1951; ?????
??????, 1996).
??? ??????? ? ??? ???????????? ??????? ???? (?????????? ???????, ?????, ??????, ?????????? ????????), ??? ?????? ????
????????? ??? ??????, ????????, ??????????? ? ?????? (??????? ????? ??????????
???????, 1998; ??????? ????? ??????????
????????, 2006). C. tataricum ????? ??????
? ?????? ?????? ? ???????? ????? ? ????????????? ? ???????? ????????, ?????????? ?
????????-????????? ???????????, ? ???????-
Corresponding author E-mail address: wolklv@mail.ru
© Siberian Federal University. All rights reserved
# 107 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. Conioselinum tataricum Hoffm. (Apiaceae) ? ??? ???????????? ??????? ? ???????? ??????
???? ??????? ???????????? ??? ????????????? ???????? (??????? ????? ??????????? ???????, 2004).
?? ?????????? ???????? ??????
C. tataricum ????????????? ??????????? ?
???????? ????? ??????? ?????????. ???? ???????????? ????????????? ???????? ?????
???? ????????? ? ???????? ????? ??????????? ????? (?? ???????? ??????, ????????????? ???????).
?????????? ???? ? ??? ????????? ??????
????????, ????? ???????? ????? ?????????
?????? ???????, ?????????? ? ???????????
???????????? ???????-????????? ???????.
???????? ? ???????-????????, ??? ??? ?????????? ????????, ???? ???????? ??????? ?????????????? ? ?????? ????? ?????. ???? ??
????????????? ???????????? ???????? ?????
?????????? ??? ?????? ? ?????? ???????? ?
??? ????????? ?????, ???????????????? ??
???????. ???????? ???? ??? ?????? 1,5 ? ??-
?? ??????? ???????? ???????? ????? ??????????? ????? ???? ????????, ??? ????????
???? ????? ?????? ???????????? ???? ? ???????? ??????????? (????????? ? ??., 1991),
??????? ???????? ????????? ??????????????
????? ???????????? ????? ???????? ??? ???????? ???????????? ???????????????? ????????? ????? ??? ???????????? ??????????
???????????? ?????????.
??????????? ?????????? ? ????????
C. tataricum ? ???????? ????? ??????? ??????? ????? (???????,1993, 2000?, 2001?, 2004,
2009). ? ????????? ?????? ???????????? ?????????? ???????? ????????? ????????????? C. tataricum ?? ???????.
???? ?????? ? ?? ????????? ?????????
?????????? ?? ???????? ???? ? ????????? ???
????????????? ?????????? ???????????? ??????? C. tataricum ? ???????? ????? ??????????? ?????.
???????? ??????? ????????????. ?????? ?????????? ? ?????????? ???????? ????? ?????????? ????????????? ???????? Ranunculaceae,
Apiaceae, Asteraceae, ??????? ???????????? ? ?????????? ? ????????? ?????????? ? ???????????? (Aconitum septentrionale
Koelle, Aegopodium podagraria L., Bupleurum
longifolium L. subsp. aureum (Fisch. ex Hoffm.),
Crepis sibirica L., Cirsium helenioides (L.) Hill.
? ??.).
??? ??????? ???????-???????? ? ???????? ?????, ? ???????? ?? ?????? ???????,
????????????? ????? ?? ???? ???????? ????????? ?????? ???????? ????????, ???????????? ?????? ?? ??????? ????????? ?? 2 ?
? ??????, ????? ???? ????????? ? ?????????????. C. tataricum (Apiaceae) ? ???? ?? ?????????? ????? ????????? ????? ?????????????
???????????. ?.?. ????????? ??? ??????? ?
???????-???????????? ?????? ????????? ???????????? (?????????, ?????????, 1993).
????????? ? ??????
???????????? ????????? ? ????????
????? ??????????? ????? (54,6-54,7є ?.?.,
84,8-85є ?.?.). ???????? ???????? ? ???????
???? ????????????? ??????? 1990-1992 ??.
???????? ??? ?????? ????????? ????????????? C. tataricum ??? ?????? ??? ? ????????, ??? ? ? ??????????? ????????????
??????????? ????????? ????????: ? ???????? ?????, ? ???????-????????? ? ?????????
???? ?????????? ????????????? (?????????????? ?? ????????), ?? ?????? ????????????? ????? ????????????? ?????????????
? ?? ?????? ????????????-?????? ?????,
?????????????? ?????????????? ??????????? (???????????). ????? ???????????
11 ?????????????. ??? ??????? ????????
? ???????-???????? ????? ?????????????
???? ?? ????????? ? ????? ? ??? ???????????
??? ?????? ?????????? ????????????? (????? 1 ????? ?? 10 ? 2).
# 108 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. Conioselinum tataricum Hoffm. (Apiaceae) ? ??? ???????????? ??????? ? ???????? ??????
?????? ????????? (???????????), ????????? ????????????????? ??????? ? ?????????????
?????????
?????????????
C. tataricum ????????? ?? ??????????? ???????? (????????????? ?, 1976), ? ??????
?????????? ????? ???????????? ?????????
???????? ????????. ???? ???????????, ???
? ?????? ??????????????? ? ??????? ???????????????? ???????????????? ?????????
??????????? ??????????????? ??????????
????????? ? ?????? ?????????????? ??????
(???????, 2001?). ??? ? ????? ???? ? ???????????? ????? ?????? ???? ??????????
????? ?????????? ???????? ? ?????????
????????? ????? ?????????????? ???????, ? ????? ?? ?????????????? ? ??????????. ??????????? ???????????? ?????????
???????? ??????????? ????? ????? ??????
?????? ???????????????? ????????? ???
???????? ??????? ????????? ??????? ????????????? C. tataricum (???????, 2009). ?
???????????? ? ???????????? ????? ???????????? ???? ?????? ?????????? ?? ?????????? ?????????? ? ??? ????? ? ? ????????
???? ? ?? ?????? ???????? ????. ???????
???????????? ??? ?????? ?????????????
?????????? ?? 10 ?? 30 ? 2, ? ??????????? ??
???????? ???????.
???????? ????????????? ? ?????? ????? ???? ??: ?.?.?????????, 1995.
??????????
?????? ?? ??????????? ?????????????
????????????? ????????? ? ???????????
????, ?.?. ???????? ? ???????????????? ???????????????. ?? ???? ??????? ??????????? ?
?? ??????? ?????? ??? ??????????, ??? ? ?????????? ?????? ?????????? (???????). ????????????? ? 5 (?????? ????????????? ???)
??????? ??????? ? ??????????? ? ? ?? ???????
???????????? ?????? ?????? ? ??????????
?????.
??????? ????? ?????????? ????????????? ?????????????. ? ?? ??????? ??????????? ?? ???? ?? ???? ???????????????? ?????,
?? ?????? ??????? (????????????? ?? 2,
3, 6, 8, 9, 10). ????????????? ?? 1, 4, 11, ?
????? ? 7, ? ??????? ??????????? ????????? ?????, ??????? ??????? ? ????????????.
?????????? ??????? ??? ???????? ?????????
?????? ??? ?????? ?????????????? ????????????? ?????? ?? ??????????? ?? ????????
(????????????? ?, 1976).
? ??????? ?????????????? ????????????? ?????????? ?????? ?????? ????? ???????????????? ?????? ??? ??????? ? ??????????????? (?? 2, 9) ??? ???????????? (?? 3,
6, 8, 10). ??????? ???????? ?????????????????
??????? ???? ????????????? ?????????????,
?????????????, ? ?????????? ? ??????????????? ??? ???????????? ??????? ???????. ???
???????????? ????????????? ?????????? ?????????? ?????? ? ??????? ?????? ????????
??????????, ?.?. ??????? ?????????????????
??????? ????? ????????????? ?????????????, ????- ??? ?????????????. ???? ?? ?????????? ?????????? ?? ??????????????? ??????? ???????, ?????? ? ?? ???????????? (??
1, 7, 11) ??? ???????????????? ??????? (? 4).
??? ??????? ???????? ????????? ?? ?????????? ?? ?????? ???????????? ?????? (?? 7,
11). ???????????? ????????????? ?????????? ?? ?????????????? ?????????? ????????
???????????????? ?????? (???????).
?????????????? ?? ??????? ????????????? ? ????????????? ? ??????????????
(???????????????) ????????? ????? ????????????? ? ???????? ?????????????, ???????????? ??????????? ??????? ???? ? ?????
(?????????????..., 1976) ??? ????????? ???????? ? ????????? ????????? «???? ?????????????» (??????, 1975).
? ??????????? ?????????? ????????????? C. tataricum ?????????? ????????????
# 109 #
2,0
?????? ????
????????????-??????
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10
11
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pl
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14,3
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0,0
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j
21,2
0,0
60,0
14,3
10,9
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0,0
20,6
19,0
0,0
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im
1,5
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0,0
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19,0
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20,0
v
6,1
38,2
10,0
28,5
21,8
27,6
0,0
10,3
28,6
0,0
6,0
g1
18,2
47,1
20,0
21,4
26,1
37,9
0,0
4,4
28,6
33,3
16,0
g2
??????? ?????????? ?????, %
3,0
8,8
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s
????????????, ss - ???????????? , s ? ?????????.
??????? ?????????? ?????????: pl ? ??????, j - ??????????, im - ??????????, v - ????????????, g1 ? ??????? ????????????, g2 ? ?????? ????????????, g3 ? ??????
3,3
3,5
4,7
????????????? ???????????????
8
4,6
???????-?????????????
7
3,9
1,0
6
5
2,6
????????????? ????
????????????-????????
????????????? ???????????????
0,9
???????-????????? ???, ????????????-????????
????????? ???, ???????????-?????????
0,2
2,5
?????????,
??/?2
3
?????? ??????
???????? ???, ???????-?????????????
???????? ???, ???????-?????????????
?????????
4
2
1
???
???????. ????????? ? ?????????? ????????? ????????????? Conioselinum tataricum ? ??????????? ????????? ???????? ??????????? ?????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. Conioselinum tataricum Hoffm. (Apiaceae) ? ??? ???????????? ??????? ? ???????? ??????
?????. ????? ????????????? ??????? ? ?????? (????????????????) (???????, ?? 3, 6,
8, 10) ??? ?????????? ? ?????? (?? 1, 2, 7).
??????? ?????????????, ? ??????? ???????
?????????? ??????????????? ?????, ???????? ? ????????? ???? ? ?? ?????? ????? (??
4, 9, 11).
???????? ?? ???? ???????? ??, ??? ????????????? ???????? ?????????????????
??????? ??????????? ? ??????????? ? ????????? ?????????? ???????-????????????
???????? ?, ????????, ? ?????????? ???????????, ?.?. ? ??????? ???????-????????????
????????, ??????????? ?????????????, ????????? ?? ??????????? ????????????????
?????. ?? ???? ??????? ?? ????????? ????????? ?????????? ?? ?????????????? ????????? ?????????? ???????, ??? ???????????,
??? ????????????????? ??????? C. tataricum
??? ???????????? ??????? ??????????????.
???????????? ????????? ????????????? ?????????????? ?? ?????? ????????????? ?????, 3,9 ? 4,7 ??????/?2, ????????? ????
??? ?? ?????? ????? ? ??????????? ???? ???
??????? ?????? ??????? (???????). ??? ??,
??? ? ???????????????? ??????, ?????????
????????????? ???? ? ???????? ??????????
??????????? ???????????.
??????????
C. tataricum ? ?????????????????? ????????, ?? ??????? ?????????????????? ??????? ????????????? ???????????. ???????????? ??????????? ? ????? ????????????
????????? ???????? ?????? ???????????
????? ? ?????? ? ???????? ??????? ???????????, ?????????????? ?? ?????? ????????????? ????? ? ?????? ??????????? ????????
(???????, 2001?). ???????? ?????? ??????????? ???? ? ??????? ? ???????? ?????.
???????? ???????????????? ?????????
????????? ????????????? ??????????????? ?
?????????????? ????????? ??????????? ????
? ???????????? ???????? ????????. ???? ??
?????? ????????????? ????????? ????????????? C. tataricum ? ??????? ? ???????????
????????? ????????. ?? ?????? ????????????? C. tataricum ????????? ? ????????????
?????????: ????????? ? ????? ???? ? ??????
???????, ???????? ???????????????? ?????
?????????? ? ????? ???????? ? ???????. ? ????? ? ???????? ??????? ?????????? ????????
?????? ??????? ???????? ????????? ??????
? ??????????? ???????? ???????. ????????
?????????????? ???????? ? ?????? ????? ?
?????????????? ???????????, ???????????
????????? ? ????????? ???? ? ???????????
?? ????????????? ??????? ????????????? ??????? (???????, 1993).
???????? ?? ????????? ?????????????
???? ????????? ? ??????? ???? ?????????????
???????. ?? ?????????? ????? ??????? ? ??????????? ??????? ?? ???? ????????????????
?? ?????? ??????? C. tataricum, ?? ? ??????
?????? 4-6 ???.1
????????, ???????? ????? ?????????????? ????????? «???? ?????????????» ?
?????????????? C. tataricum ?????????? ??
????? 3 - 5 ???.
????? ????????????, ??? ???????????? ????????????? C. tataricum ??????????? ? ???????? ????? ???????????
????????????? «???? ?????????????», ? ?
???????????? ?????????????, ?????????????? ?? ???????, ???????? ????? ?????????? ???????? ????? ??????????
«???????». ????????? ? ????????????,
? ?????????????? ????????????? ????
??????????? ? ?????????? ???????????,
????????????? ????????????? «???? ?????????????» ? C. tataricum ?? ??????? ??
??????? ?????????????. ???????????????
?????????? ????????????? C. tataricum,
??-????????, ???????????? ? ?????? ???-
# 111 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. Conioselinum tataricum Hoffm. (Apiaceae) ? ??? ???????????? ??????? ? ???????? ??????
???? ????????????? ???????? ? ?????????????????? ????????????? ???????, ??
??????? ??????? ???????? ??????????????
???????? ? ?????? ????? ? ?????????????.
? ???? ? ??????????????? ?????? ??????
???????? ?????????????? ????????????
?????? ?????????? ?? ???? ?????????? ????? ? ????????? 2-3-?? ????????.
???????????? ?????????????, ? ??????? ???? ???????????????? ??????, ????????
?? ??????? ? ?????????? ???????? ???????
???? ?????, ?.?. ???????? ???????????. ????????????? ???? ????????????? ? ??????
????????????-???????? ??? ?? ?????????
?????? ? ???????-???????? ????. ? ??? ?????????? ????? ????????? ???????? ??????? ????? C. tataricum. ????????? ?????????
??????, ????????? ? ???????????? ????? ?
???????? ????, ???????????? ??? ?????????
??????????? C. tataricum ? ??????? ????? ????? ??????????.
??????, ??? ????????????????, ?????????????, ? ??????? ?????????? ???????????? ?????, ????????? ???????? ???????????
(??????????????, 1976).
?????? ????????????? C. tataricum ???????? ?? ???? ???? ????? ???????????. ???
??????????????? ? ??????? ?????????????
??? ???? ??????? ???????? ??? ? ??????, ???
? ? ??????? ??????? ?????????????? ????.
?? ??????????? ???????? ?????????????
??????????????? ?????? ????? ??????????
????????????? ???????? ? ???????. ??????
? ??????? ???? ????????????? ?????? ????
?????????, ???????????????? ??????. ???
????????? ??????? ?????? ?????????????
????? ?? ?????? ??????????? ??????????
?????? ?????????????, ? ???? ???????? ????????????? ???? ? ???????????. ????????
???????? ???????????? ???????? ????????????? ???? ?????? ?????????????? ??? ??????????? ? ???????.
???????????? ?????? ?????????????
C. tataricum ???????????? ????????? ?????
???????? ????????, ??????????????? ??????????????? ? ???????????? ???????. ? ???????????? ??????????? ????????? ????????
??????? ????????? C. tataricum ??????????
?????????????????. ??? ???? ?? ??????????
????????????????, ? ?????????, ?????? ???????? ?????? ? ????????????????? ?????????
?????????? ????????? ? ???????? ? ?????
????????????????? ?????????? ??????????? ???????? ??????????? ??? ? ??????, ???
? ? ??????? ??????? (???????, 2000?, 2001?).
???????????? ???????????????? ??????? ?????????? 14-18 ???, ????????????? ? 10-12 ???,
????????????????? ? 5-6 ??? ??? ????? ????????????????? ?????????? ? 30-35 ???.
???????????????? ????????? ?????????
?????????????? C. tataricum ???????? ?????
????????????? ??????????? ????, ??? ??????????????? ???????? ?????? ???????????,
?????????????? ????????? ??????????? ????
? ??????? ?????????? ????????????? ?????
????????-???????????? ????????, ????????? ????? ???????? ??????.
???? ???????????????? ????????? ? ???
??????? ? ??????? ???? ???????? ????????????? ???????????, ???????? ????, ?? ????????? (???????????) ????????????? ????????
??????????? ???????????????? ???? ? ????????????? ? ?????????? ????????, ??? ????????? ?????????? «????????? ?????????» ????
(?????????????...,1988). «????????? ?????????» ????? ???????????????, ? ?????????
?.?. ????????? (1975), ??? ???????? ?????????, ????????????? ?? ???? ?????????????
????????????, ??????????? ???? ?????????? ???????????? ??????? ? ??????.
? ???????? ????? ????????????? ??????????? ??????? ????????? ?????????????
????????, ??? ??????? ?? ????????? ???????? ? ???????????? ? ?????????? ? ????????
# 112 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. Conioselinum tataricum Hoffm. (Apiaceae) ? ??? ???????????? ??????? ? ???????? ??????
???????????. ???????? ???????????? ??????, C. tataricum ?? ?????????? ????? ??????
????? ?????? ?????? ? ???????????? ??????????? (????? ????, 1951 ? ??. ?????????), ???
????? ???????? ? ??? ?????? ?????????????????????.
? ???????? ???????? ????? ???????
???????? ??????? «????????? ?????????»
C. tataricum ?? ?????? ????????????? ????? ? ????? ??????????? ???????????? ???
???? ????????? ?????????????, ??????????? ?????????? ?????? ????????????? ??????????? ????, ????????? ? ???????????????. ??????? ?????? ????????????? ?????
???????????? ????? ??? ??????? ???? ? ??????? ?? ????? ??????????.
?????? ????????????? ???? ? ?????? ???????????? ??????????? ?? ???????, ??????????? ? ?????????? ????????????? ???????
?????????. ??????????? ???????? ????????????? ????????????? ???????????? ???
????????????? ????????? ? ???????? ??????????????: ?????? ?????, ??????? ?????????
??????????????? ???? ????? ?? ??????????
????? ?????????????? ??????, ?????????? ??????????? ????? ? ?????? ?????? (????????...,
1991). ??????????? ??????????? ?????? ????????????? ?????, ?????????? ?? ?? ??????
??????? ????????, ? ????????? ??????? ?????? ?, ??? ?????????, ?????? ???????????????
??????? ?????????? ????? (??????????,
1988). ??????????????? ???????? ?????????? ?? ?????? ????????????? ????? ???????
? ???????? ??????? ?????????? ??????????? ? ??????????? (????????, 1983). ???
???????????-??????????? ???? ??? ???? ??
????????????? ????????.
?????? ?????? ????? ????????????
??? ?????????? ??????. ?????????? ??????????? ???????? ? ?????????? ????????? ?
??????????? ????????? ??????????? ?????
? ?????????????? ??????? ??????. ??????-
??????? ???? ???????? ?? ??????????. ??
????????? ?????? ?????????? ?????? ????????????? ????? ??????????? ????????????- ? ???????????-?????? ???? (??????????,
1988). ?? ???? ????????? ?????? ??????????
??????????? ????????????? ?????? ? ????????????? ?????, ?? ?????????? ?????????? ???? ?????? ? ?????????, ?????????
??????????????? ??????? ?????????? ?????.
?????????? ???????? ??????????? ?????
?????? ????? ???? ????? ?? ?????? ???????? ????????? ????????????? C. tataricum
? ?????? ???????????. ?? ????????? ??????????? ???????? ????? ??? ????????? ???????? ?? ??????????.
??? ???????? ?????????? C. tataricum
???????????? ????? ??????????? ????????
????, ??? ??????????? ?????? ?????????
???????? ??????? (??????????, ??????????) ? ???????? ?????? (????????????, ?????????) ??????, ??? ??????????? ?? ????
????????????? ??????????? (???????,
2001?, 2004, 2009). ????????????????? ????????? ?????? ???? ????? ?????????? ?? ????? 5-6 ??????, ? ???????? ???? ?????????
????? ??????? ???????? ???????, ? ? ?????
???? ?? ????????? ??????? ???????? ???
?? ?????????????? ? ??????????. ?????????
????????????? ??????? ????????????? ????????? ??????? ??????? ?????????????? ???
????????? (?????????, 1991), ???????????
?????? ????????? ???????? ??????????????????? ????? ?????? C. tataricum ?????
????????????? ??? ??????? ?? ????????????
?????? ??? ????????????? ? ?????? ????????? ??????????.
??????? ? ????????? ????? C. tataricum,
???????? ??????? ?????????? ? ????? ??????? ?????????, ? ???? ???????? ????????????????????. ?????? ????????, ???????? ?????
????????, ???????????? ? ???????????? ?????, ??????? ??????? ?????????????? ????-
# 113 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. Conioselinum tataricum Hoffm. (Apiaceae) ? ??? ???????????? ??????? ? ???????? ??????
?? ? ??????????? ????????? ?????, ?????????? ???? ? ???????? ?? ????? ??????????????
??????.
???????? ???????? ???? ???????, ???
???????, ???????????, ???????????, ?????????? ???????????? ??? ????????????? ???????????? ?????????, ? ??? ??????? ??????
???????? ????????? ?????. ?????? ???????
C. tataricum ??????????? ?????????? ? ????????? ????????????? ?????? ? ????????????.
? ???????? ????? ???????-???????? ? ???????? ????? C. tataricum ?????????? ?????, ?????????? ???????. ???????????? ?????????
????????????? C. tataricum ? ?????? ???????
???????????? ??? ?????????? ????????? ??????, ??? ? ???????????????? ??? ???? ??????????????????? ?????????, ???????????
???????????? ?????????.
???????? ????????? ?????????????
???? ??? ?????????? ???????? ???????????
? ?????? ??????? ? ?? ?????? ????? ????????????? C. tataricum ??? ????????????? ????????.
«????????? ?????????» C. tataricum ?
???????? ????? ???????? ?? ???????? ????????????? ?????????????, ????????? ?????? ???????? ?????? ? ??????????? ???????
????????? ???????? ???????????????????
????? ??????. ????????? ????? ????????
?????? ???????????? ????????????? ????????????? ????????????? ? ???????? ????????????? ????????? ??????????? ????????.
??????????? ?????? ????????? ????? ?????? ? ????????????? ???????????? ??????
???????? ? ???????? ????????????? ???????
? ????????? ???? ?? ??????? ??????????.
???????????????? ?????????? ????????????
?????????? ???? ????????? «????????? ?????????» ???? ?????????? ?? ??????? ?????????????: ??? ? ??????, ??? ? ? ???????, ??? ?
????????, ??? ? ? ??????????? ???????????
?????????????? ????.
? ???????? ????? ??????????? ?????
?????? ???????? ???????????? ?????? ??
?????????????? ????????? ? ????????????
????????. ??????????? ??????????? ???????? ????????? ???? ???????????? ?????
??????????? «??????» ? «????????» ??????
(??????????, ?????????, 1987; ?????????,
1990; ????????? ? ??., 1991).
?? ??????? ????? ????????? ?????????????? ????????? ???????????? (Bupleurum
longifolium L., Aconitum septentrionale Koell.
? ??.) ???? ???????? ??????? ????????? ????? ???????????? ? ???????????? ???????? ???????? ????? (???????, 1991, 2000?,
2000?, 2001?). ????????? ????? ?????????????? ?? ?????? ???????????????? ?????????? ??????, ??? ??????? ????????????,
??? ????????? ????? ????????? ??????? ???????????? ???? ? ????????? ??????????
???????? ?????????????. ? C. tataricum ?????? ????????? ???, ???????????? ???? ???
? ??????????? ????????????? ? ???????????. «????????? ?????????» ???? ? ????????
????????????? ??????????? ????????????
??????????.
?? ???? ????????? C. tataricum ?? ??????? ???????? ? ?????????????? ??????????????????? ?????? ????????? ????????????. ??????????????? ???? ? ???????? ?????
??????? ??????????? ???????? ????????????? ????????.
??????????
? ????????????? ??????????? ???????? ???????? ????? ??????????? ?????
C. tataricum ????????? ????????????? ??????????????? ??????????? ????, ???????????? ??? ???????????????, ????????? ????????????, ????????? ?????????. ? ????????
?????????? ??????????? ???????? ?????????????? ???? ?????????? ??????????. ?????????????? ??? ???? ??????? ??????? ???????
# 114 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. Conioselinum tataricum Hoffm. (Apiaceae) ? ??? ???????????? ??????? ? ???????? ??????
??????? ??????????????????? ??????? ? ?????????? ??????? ?????????? ???????????.
«????????? ?????????» C. tataricum
???????? ?? ???????? ??????????? ????????, ????????? ?????? ???????? ??????,
??????????? ??????? ????????? ????????
??????????????????? ????? ?????? ? ???????????? ?????????? ??? ????, ????? ?????????? ??????? ???????????? ???? ???? ?
??????? ????????????? ???????????. ? ???1
????????? ??????????? ????????? ????????
??????? C. tataricum ???????????? ? ????????
???????????, ??? ???????????? ???????.
??? ?? ???????? ??????????? ???????????? ????????? ? ???????????????? ????????? ????????????? ? ?? ??????????? ? ????????????, ??????????? ???????? ????????
?????. ??? ?? ????????? ???????? C. tataricum
? ???????? ?????????????? ??????????????????? ?????? ?????????? ?????????
????????????.
??????????? ?????????????? ????????? ????? ?????? ? ???????????? ??????? ????????? ???????? ?????
????????? ?????????? ?????????? ??????? ???????? ?????? ?? ??????????? ????????????? ????????? (???????,
2001?).
?????? ??????????
??????? ?.?. (1991) ????????? ????? ????????? ?????????? ?? ???????. ?: ?????????
????? ????? ???????? ? ???????????? ????????????? ???????. ???????????, ?. 36-53.
??????? ?.?. (1993) ???????????? ????????? ? ???????? ????? ???????. ???. ????. ????.
4: 65-69.
??????? ?.?. (2000?) ??????????? ???????? ????????? ? ?????????????? ????? ??? ???????? ??????. ???. ??????. 12: 1-12.
??????? ?.?. (2000?) ????????????? Aconitum septentrionale Koell. ? ???????? ????? ??????????? ?????. ????? ? ?????????????? ?????. 5, ???. 1, ?. 47-52.
??????? ?.?. (2001?) ????????? ? ??????????? ???????? Conioselinum tataricum (Apiaceae).
???. ????. 8: 85-93.
??????? ?.?. (2001?) Aconitum septentrionale Koell. ? ???????? ????? ??????????? ?????.
??????. ???????. 37(3): 34-40.
??????? ?.?. (2004) ????????? Conioselinum tataricum Hoffm. ?: ???????????????? ?????
????????????? ????????. ??????-???, ?.106-110.
??????? ?.?. (2009) ??????????? ????????? ???????? Conioselinum tataricum Hoffm. (Apiaceae) ? ????? ? ????????? ??????????? ??????? ?????????????. ???????????? ??? ?????????
??????. 1: 77-83.
??????? ????? ?????????? ??????? (1998). ????????? ?????????. ?.: ?????, ??????? ???????????, 421 ?.
??????? ????? ??????????? ??????? (2004). ??? 2. ???????? ? ?????. ???. ???????? ?.?.,
??????? ?.?. ???????, 360 ?.
??????? ????? ?????????? ???????? (2006). ?????: ??????????? ???????????? (?????), 456 ?.
????????? ?.?., ???????? ?.?., ????????? ?.?. ??. (1991) ??????? ????????? ?? ????????????? ????????? ????? ? ???????? ????? ???????. ???????????.1: 66-69.
????????? ?.?. ??., ????????? ?.?. (1993) ????? ??????????? ?????. ???????????: ???????? ???? ?? ???, 59 ?.
# 115 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. Conioselinum tataricum Hoffm. (Apiaceae) ? ??? ???????????? ??????? ? ???????? ??????
???????? ?.?. (1975) ???????? ???????????? ????????? ? ????? ????????? ?????????
????? ????? ????????. ???. ????. ???. ????. 80(2): 5-17.
???????? ?.?. (1984) ???????????. ?.: ???-?? ???, 319 ?.
?????????? ?.?., ????????? ?.?. (1987) ????????????? ? ????????????? ???????? ????????????? ???????? ????????? ???????. ?: ??????????????? ???????????? ? ???????? ?
??????? ??????. ???????????, ?. 104-111.
?????????? ?.?. (1988) ???????????? ???????? ? ??????? ??????????? ???????. ???????????: ?????, 159 ?.
?????? ?.?. (1975) ?????????? ?????? ????????????????? ??? ??????? ??????? ? ?????????????? ???????? ?????????. ????. ?????. 2: 7-34.
????? ???????? ??????, ???.8 (1935). ?????, ?. 2061-2062.
????? ????, ??? 18 (1951). ?.- ?.: ???-?? ?? ????, ?. 2-3.
????? ??????, ??? 10 (1996). ???????????: ?????, ???.???????????? ????? ???, ?. 73174.
????????????? ???????? (???????? ??????? ? ?????????) (1976). ???. ?????? ?. ?. ?.:
?????, 217 ?.
????????????? ???????? (?????? ????????????? ????????) (1988). ???. ?.?. ???????????,
?.?. ??????, ?. ?. ???????, ???????? ?.?. ? ??. ?.: ?????, 182 ?.
????????? ?. ?. (1995) ?????????? ???????? ?????? ? ???????????? ??????????. ???.:
??? ? ?????, 990 ?.
???????? ????????? ???????? ????? ??????? (1991). ???. ?.?.?????????, ?.? ???????????. ???????????: ?????, 72 ?.
Conioselinum Tataricum Hoffm (Apiaceae)
and its Coenotic Positions
in Subnemoral Forests of the Salair Ridge
Ludmila V. Volkova
Central Siberian Botanical Garden, SB RAN
101 Zolotodolinskaja str., Novosibirsk, 630090 Russia
The structure of 11 coenopopulations (CP) of Conioselinum tataricum Hoffm (Apiaceae) in tallherbaceosus phytocenosis of Salair ridge (south of West Siberia) was studied. The CP are generally
of normal type. CP density is the highest in forest-tall-herbaceosus meadows. C. tataricum is the
coenotical patient in plant communities. The biological and CP-stucture peculiarities show the low
growth adaptation of C. tataricum in tall-herbaceosus phytocenosis of Salair.
Keywords: coenopopulations, age-stucture, density, adaptation.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2011 4) 117-132
~~~
??? 574.587
????????????? ????? ???????? (Annelida, Oligochaeta)
? ?????? ???????????????? ?????????
????? ???????, ??. ?????? (??????, ??????)
?.?. ???????????*
??????????????? ???????? ?? ???
?????? 664033, ?. ???????, ??. ????-?????????, 3, ?/? 278 1
Received 3.06.2011, received in revised form 10.06.2011, accepted 17.06.2011
??????????? ????? ?????????????? ?????? (Oligochaeta: Annelida) ? ??????, ????????????
? ???????????? ??????????? ? ???? ??????? ?????????? ???????????????? ?????????,
?????????????? ? ????? ??????? (???????? ??????). ? ?????? ?????????? ???????? ??
?????????? ?????????? ?????????????? ??????, ????????? ? ???? ??????. ????????
????????????? ??? ??????????? ???? ??????? ??????? ?????????????? ?????????? ????????
????????. ???????????????? ????????? ??????????? ????????? ???????? ??????? ?
?????????????? ????????????? ???????? ?? ???????? ???????? Thioploca ? ???? ??????.
???????? ?????: ???????? ??????, ??????????????? ????, ?????????????? ?????, ???????
??????, ???????????? ?????.
????????
? ????? ??????? (???????? ??????) ? ??????? ????????????? ????????? ?????????
«???» ????? 2009 ?. ????????? ????????????
??? ? ????? ?? ??????? 408?461 ? ? ??????
???????????? ????????? ??????????????????? ?????????? ??????????? ?????????, ??????? ????????? ?? ??????? 420 ? (55?31? N,
109?46? E) (???????, 1979, 1984; Crane et al.,
1991). ???????? ????? ????? ????????? ??-??
???????????? ????????? ??? ?????? ??????
????????? ??????? (Shanks, Callender, 1992),
???????, ??? ????????, ???????? ??????? ???????????? ?????????-????????? ????????-
*
1
???. ? ??????????? ??????, ??? ??????????
??? ? 80-? ??. ?? ????? ?????? ????????????? ?????????? ?? ???????, ?????????? ?
??????? ????????? ????????? ?????????
«??????» (?????? ????????? ??????????????)
? ?????????????? ??????????? ????????? ?
???????????? (???????????? ??????? ??????), ?????????? ??????? ????????????? ??????????, ?????????????? ???????? ???????? ??????????? ??? ????????? ???????????
(?????????, ??????), ??? ? ??????????????
?????????? (?????, ???????, ????????, ?????????, ???????? ? ??.) (???????? ? ??., 1991;
?????? ? ??., 1998). ????????, ??? ??????????
Corresponding author E-mail address: irina@lin.irk.ru
© Siberian Federal University. All rights reserved
# 117 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ????????????? ????? ???????? (Annelida, Oligochaeta) ? ?????? ???????????????? ??????????
???????? ??? ???????????????? ?????????
????????? ? ?????? ????????? ??????????
????? ???????? ???????????? ????????, ?????????? ?? ?????? ?????????? ??????, ?????????? ???????????? ? ??????? ??????????????? ?????????? ???????????, ??? ?????????
?? ??????????? ?????? ????????? ??????????? ????????????? ??????????? (???????? ?
??., 1991), ???????? ??????????? ???????
????????? ?????????? ???????? ????????????
???? Thioploca (????????? ? ??., 1994). ????????????? ???????????? ????????? ?? ??????????? ????????????, ??? ????????? ?????
????? ?????? ?????? ???? ? ?????????????
? ???????????? ??????????? ??????????????? ????????? ????? ??????, ????? ???????? ??? ??????????????? ?????????????
???????? ???? ?????????? ???????? ?????
??? ????? ????????????? ???????? ?? ?????????? ???? 20-??????????????? ???????
????? (Grachev et al., 1995).
???? ?????? ?????? ? ???????????? ???????? ???????????? ????????, ???????????? ?????? ? ????????? ??????????? ?????
???????, ???????????? ? ?????? ??????
?????????? ??????.
????????? ? ??????
????????????? ???????? ??? ??????
???????????? ???????????????? ????????? ?? ??? ?.?. ?????????? ? ?.?. ???????? ?? ????? ?????????? ??? «???-1» ?
«???-2» ? ??????? 2009 ?. ? ?????? ??????
???????????? ????????? (????? ???????,
???????? ??????) ? ?????? ?? 408 ?? 461 ?
(???. 1, ????. 1). ?????????????? ?????
?????????? ?? ???? ? ???????????? ? ??????????? ????????????????? ???????? ???
????????????? ??? ??????, ??? ? ????? ??????????????? ?????? (???????????, 1962;
????????, 2004). ??? ??????????? ??????? ?????????????? ??????????????? ??????????????? ???????, ????????????? ?
?????????????? ?????????? ????????. ???
????????? ??????????????, ???????? ?
????????? ???????????????? ????????? ?
???????? ? ??????.
??????????
?? ?????? ?.?. ??????????? (1987),
??????????? ??????? ?????????????? ???????? ??????????? ????? ?????? ? ????? ?
??????????????? ?? ?????? ?????????????
???. 1. ?????-????? ?????? ????????????. ???????? ?????????? ??????? ? ????? ???????
# 118 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ????????????? ????? ???????? (Annelida, Oligochaeta) ? ?????? ???????????????? ??????????
??????? 1. ?????? ??????? ?????? ???? ? ????????? ????????? ? ???????
?????
???????
????, ????
???????, ?
09106
11.08.2009 ??? «???-1»
457
?????, ??, ?????
09107
11.08.2009 ??? «???-1»
461
????????????? ???, ?????? ??
09111
14.08.2009 ??? «???-2»
441
????????????? ???, ?????????? ??, ?????
09112
14.08.2009 ??? «???-2»
441-448
09113
14.08.2009 ??? «???-2»
461
09119
15.08.2009 ??? «???-2»
439
09120
15.08.2009 ??? «???-2»
408
?????????, ?????? ??? ???? ? Rhyacodriloides
abyssalis ?ekanovskaja, 1975, Lamprodrilus
wagneri Michaelsen, 1901 ? Pseudorhynchelmis
olchonensis (Burow, 1932) ? ??????? ??? ????? ???????. ???? ?????? ?????, ?????????
? ?????? ?. ???????, ??????????? ????????
(????. 2). ??????????????????? ????????
????? ?????????????? ?????? ??? ???????????? ????????? ????? ??????? ???????
?? ???????? Lumbriculidae (???? ???), Propappidae (???? ???), Naididae sensu Ersйus et al.,
2008 (????? 19 ?????). ????????? ?????????
???????? ????, ????????????? ???? ????????????? Naidinae (???? ????? ??????????????? ???????? ???????), Tubificinae (12 ?????)
? Rhyacodrilinae (6 ?????).
??????????????? ?????
NAIDIDAE: Naidinae
1. Naidinae gen. sp.
????????? ????????? ???????????????
???????? ??????? ???? ??????? ????? ????
??????? ? ????????. ????? ???????? ??? ?????? ??????? ???? (????? 5,5 ??, ??????????
0,3 ? 0,4 ??) ? ??????????? ??? ?????????????? ???????????? Naidinae ?????????? ??????? ???????? ????????? (???. 2). ????????????? ?????? ????? ???? ???????? ??????????
??????????? ????????: ??????????? ???????
??? ?????????
????????????? ???
??
????????????? ???, ?? ?? ???????????
??????
?????????? ??, ??????, ?????
?? 30 ?? 50 ??? ?????? ??????????? ?? 1 ? 2
? ?????, ??????? ??????? ?????????? ? IX,
??????? ? ? XV ????????.
NAIDIDAE: Tubificinae
2. Baikalodrilus paradoxus
(Snimschikova, 1984)
??????? ???????. ?????? ?.?. ??????????? (1984) ?? ???????????? ??????? ?
?????? ?. ???? (???????? ??????), ? ???????
10 ?. ????????????? ? ???????? ? ???????
???????. ?????? ?? ???????????, ???????
?? ???????? 5 ? 250 ? (????????, 2004). ???
??????? ??????????? ??? ??????????? ????
???????. ???? ????????? ? ?????????? ? ??????? 441 ? 448 ? 457 ? (????. 2).
3. Baikalodrilus inflatus (Michaelsen,
1901)
??????? ???????. ???????? ???????????? ?????? (Michaelsen, 1905; ??????????,
1987; ????????, 2004) ??? ???????? ?????????? ? ????????????? ?????????????? ????
????? (40?80 ?). ? ????? ?????? ??????????
???? ?????????????? ????? ? ?????, ??????
??????????????? ? ?????????????? ???? ??
??????? 441 ? 448 ?.
4. Baikalodrilus sp.
?????? ? ????? ? 09106. ??????? ?
Baikalodrilus, ??? ??? ????? ????????????
# 119 #
Baikalodrilus inflatus (Michaelsen, 1901)
Baikalodrilus sp.
Isochaetides adenodicystis Semernoy, 1982
Isochaetides baicalensis (Michaelsen, 1901)
Isochaetides compactus Semernoy, 2004
Isochaetides peniacerus Semernoy, 2004
Isochaetides sp.
Tasserkidrilus infundibuliferus (Isossimov, 1972)
Lamadrilus excavatus (Hrab?, 1982)
Lamadrilus proprius Semernoy, 2004*
Limnodrilus sp.
3
4
5
6
7
8
9
10
11
12
13
Rhyacodrilus isossimovi ?ekanovskaja, 1975
Rhyacodrilus korjakovi Semernoy, 2004
Rhyacodrilus propus Semernoy, 2004*
Rhyacodrilus vasalatus Semernoy, 2004
14
15
16
17
NAIDIDAE: RHYACODRILINAE
Baikalodrilus paradoxus (Snimschikova, 1984)
NAIDIDAE: TUBIFICINAE
2
1
2
1
NAIDIDAE Benham, 1890 sensu
Ersйus et al., 2008
Naidinae gen. sp.
??????
?
?/?
+
+
+
+
+
+
+
3
0-20
+
+
+
+
+
+
+
+
4
+
+
+
+
+
+
+
+
+
5
20-50 50-250
+
+
+
+
+
+
+
+
6
250????.
+
???????? ??????
+
+
+
+
+
+
7
0-20
+
+
+
+
+
+
+
8
+
+
+
+
+
+
+
+
+
9
20-50 50-250
???????, ?
??????? ??????
+
+
+
+
10
250????.
????????????? ?? ???????
+
+
+
+
11
0-20
+
+
+
+
12
+
+
+
+
13
20-50 50-250
????? ??????
+
+
+
+
14
250????.
??????? 2. ??????????? ?????? ????? ????????, ????????? ? ?????? ?????? ???????????? ??????????????? ?????????? ? ????? ??????? (???????? ??????),
? ????????? ?????? ?? ?? ??????????????? ?? ??????? (?? ?????????, 2004).
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Lamprodrilus achaetus Isossimov, 1962
LUMBRICULIDAE Vejdovsky, 1884
+
+
+
3
* - ????, ??????? ???????????? ? ???????? ????????? ????? ??????.
21
Propappus glandulosus Michaelsen, 1905
PROPAPPIDAE Coates, 1986
Pararhyacodrilus confusus Semernoy, 2004
20
Rhyacodriloides abyssalis ?ekanovskaja, 1975
19
2
18
1
??????????? ????. 2
+
+
+
4
+
+
+
5
+
+
+
6
+
+
+
7
+
+
+
8
+
+
+
9
+
+
10
+
+
+
11
+
+
+
12
+
+
+
13
+
+
14
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ????????????? ????? ???????? (Annelida, Oligochaeta) ? ?????? ???????????????? ??????????
???. 2. Naididae sp. ? ??????? ??? ???????? ????? ????
????? ???? ??????? ?? ????????? ???? ???????????? ?????? (???. 3). ? ??????? ??
?????? ????? ?????? ???? ?? ????? ????????
???????.
5. Isochaetides adenodicystis Semernoy,
1982
???????. ???????? ?????? ???????????????? ??? ????? ????????? ???????. ??????? ?? ????????????? ???????? ?? ???????
?? 2 ?? 360 ?, ???????????? ??????? ??????
? ????????, ???????? ??????? ?????????
(??????????, 1987). ???? ????????? ?? ??????? 408 ? ? ????? ? 09120.
6. Isochaetides baicalensis (Michaelsen,
1901)
???????. ????????? ? ????????? ???????. ???????? ?? ??????, ???????? ? ????????? ??????? (??????????, 1987). ?.?. ???????? (2004) ??????????, ??? ???? ??? ?????
??????????? ? ????????? ??????????????.
???? ???? ??? ??????? ??????????? ? ??????? ????? 250 ?.
7. Isochaetides compactus Semernoy,
2004
??????? ???????. ????????, ????? ?????? ? ???????? ??? ????? ????. ????? ?? ???
??????? ?? ??????? ???? 250 ?. ? ????? ??????? ?????? ? ??????? ?????????? ?? ??????? 408 ? (????. 2).
8. Isochaetides peniacerus Semernoy,
2004
???????. ?????? ?.?. ???????? (2004).
??????? ????????????? ? ????? ???, ???????
250 ? (???????? ??????). ? ????? ????? ??????????? ? ??????? 408 ? (????. 2).
9. Isochaetides sp.
????? ??????, ?? 5,5 ?? ? ????? ? 0,7 ??
? ???????????; ????? ?? 4 ? 9 ???????????
??????? ? ?????; ?????????? ???????, ????????, ? ???????? ???????? ???????, ??????
????????? ???????????????? ??????????
?????????????? ????? (???. 4). ???? ??????????? ???? ?????, ?????????? ? ????? ?
09120, ????? ?? ???? ???????? ? ???????.
# 122 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
???. 3. Baikalodrilus sp. ? ????????? ???????? ?? ??????????? ????, ?????????? ???????? ??????? ??
?????????????? ? ??????????? ??????
???. 4. Isochaetides sp. ? ?????????? ?? ???????????????? ??????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ????????????? ????? ???????? (Annelida, Oligochaeta) ? ?????? ???????????????? ??????????
NAIDIDAE: Rhyacodrilinae
10. Tasserkidrilus
infundibuliferus
(Isossimov, 1972)
??????? ???????. ?????????? ??? ???
??????? ? ???? Limnodrilus, ?.?. ????????
(2004) ??????? ??? ? ??? Tasserkidrilus. ??? ????????? ? ?????????????, ??????? ? ??????????? ? ???????? ?? ???????????? ??????. ?
????? ????????? ??????? ? ?????? ? 09106,
09107, 09119 (????. 1, 2).
11. Lamadrilus excavatus (Hrab?, 1982)
??????????? ???????. ?.?. ?????????? (1987) ????????? ????? ????????????? ?
?????? ?????? ????????? ??????? ?? ??????? 5 ? 198 ? Tubifex bazikalovae crassiseptus
(Semernoj, 1982), ??????? ???????????? ????
????????? ? Lamadrilus excavatus. ?? ?????
??????, ??? ??????? ?? ????????? ????? ?
?????? ?????????? ??????????? ????????? ??
??????? 457 ?.
12. Lamadrilus proprius Semernoy, 2004
??????? ???????. ??? ?????? ?.?. ???????? (2004) ?? ????????? ????? ? ?????
????, ? ??????? 150 ?. ? ????? ?????????
???????. ?????? ???????????????? ?
??????? ???. ??????? ?????? ?.?. ??????????? (1975) ?? ??????????? ???? ? 512?1610 ?,
? ?? ?? ????? ??? ????????? ?? ? ?? ??????????? ? ?? 100 ?? 330 ?. ? ???????? ???????
???????? ?? ???????? ?? 2 ?? 500 ? (??????????, 1987). ? ????? ?????? ? ???? ?? ????? ????????????? ????? (????. 2). ????????
???????? ????????? ?????? ?? ?????????????
???? (??????? ?????? ? 09112).
15. Rhyacodrilus korjakovi Semernoy,
2004
??????? ???????. ?????? ?.?. ???????? (2004) ?? ?????? ???? ? ??????? 22 ?,
?? ?? ?????? ? ???????? ??????? ?? ???????
250 ?. ? ?????????? ?? ????? ??????? ??????? ???? ? ????? ? 09113 (????. 1).
16. Rhyacodrilus propus Semernoy, 2004
??????? ???????. ???????? (2004) ????????? ? ?????? ???? ??? ?? ?????? ????, ?
????????????? ????? ???? ?????????? ??
?????????? ? ???????????? ??????????????
???? (? 09111 ? 09119, ????. 1), ?? ??????? 439
? 441 ?.
13. Limnodrilus sp.
? ???? ??????, ?????????? ? ??????
????????????? ????????? ?? ??????? 408,
439 ? 461 ? (????. 2), ???? ?????????? ????? ? ???????????? ??? ?????????????? ????
Limnodrilus ??????? ???????? ????????????? ????????: ??????? ????????????????
?????????? ???????? ? ???????? ?????????????? ?????? (???. 5). ??? ?? ?????, ??
???? ?? ???? ????? ????? ????, ????????? ?
????????? ????? ??? ???????, ?? ????????????? ??????????????? ?????????????? ???????????? ???? ?????. ????? ????, ?? ???
??? ???????????? ????????? ???????????
??????????? ????????.
??????? 70 ? 140 ?. ? ????? ????????? ?????????? ??? ???????? R. propus ??? ????????? ? ????? ? 09120 (????. 1).
17. Rhyacodrilus vasalatus Semernoy,
2004
???????. ?????????? ?????? ?? ??????????? ???? ????????? ???????. ????????
(2004) ??????? ??? ?? ???????? 210 ? 350 ?.
???? ??????? ??????????? ? ????? ?????????
??????? (????. 2).
18. Rhyacodriloides abyssalis
?ekanovskaja, 1975
???????. ????????????? ??? ?????? ??
???????? (???????????, 1975). ? ??????????
??????? ?????? ???????? ?? ???????? ?? 10 ?
? ?????? (????????, 2004). ???????? ?????
????????????? ??? ? ????? ?????? (????. 2).
???????? ??????????? ? ???????? ? 09107,
09111 ? 09112 (???. 6), ???, ??? ? ??????? ???-
14. Rhyacodrilus isossimovi ?ekanovskaja,
1975
# 124 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
???. 5. Limnodrilus sp. ???????? ???????? ?????????: 1 ? ?????????? ????????, 2 ? ?????????
?????????????? ??????, 3 ? ??????????
???. 6. Propappus glandulosus: ? ? ???????? ????? ????: ?? ???????? ??????? ??????? ??????????, ???????
???; ? ? ?????????? ????? ??????? IV ????????, SG ? ?????????? ??????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ????????????? ????? ???????? (Annelida, Oligochaeta) ? ?????? ???????????????? ??????????
?????? ???????????? ????????????? ??????????.
19. Pararhyacodrilus confusus Semernoy,
2004
???????. ?????? ?? ???????? ??????? ?
?????? 1969 ?. ?? ??????? 345 ? 420 ?, ??????
?.?. ???????? (2004). ? ?????? ?????????
????????????? ????? ??????? ? ???? ??????
(????. 2).
??????? ?????????? ? ????? ????. ????????
????? ???????? ?????????? ?? ???????? ??????? ?? ????? ????????????? ???? ? ??????????? ??? ?????????????? ???? ??????????
?????? (???. 6).
LUMBRICULIDAE
21. Lamprodrilus achaetus Isossimov, 1962
???????. ? ???????? ??????? ???? ??
???????? ????? ????????????? ????? ???????????. ??????? ??????????? ?? ???????
2 ? 500 ? (??????????, 1987). ?? ???????????, ??? ?? ???? ? ? ????? ??????? (????. 2).
PROPAPPIDAE Coates, 1986
20. Propappus glandulosus Michaelsen,
1905 (?)
???, ????????????? ????????? ?? ?????????? ???? ???????, ???????????? ???
?????? ? ???????????????? ???????? ?????
?? ???????? ?? 120 ? (??????????, 1987),
?????????????? ??????????????
????????? ????????
? ????????????? ???? ?????? ?????????
????????? ???????? ? ??????? ??????????
2837 ???/?2 ??? ??????? ???????? 2,68 ?/?2
(???. 7). ???????????? ?????????? ????????-
? ????? ? ????? ?????? ? ?????? (???????????, 1962). ???? ????????? ???? ?????????
?? ??????? 461 ? ? ????? ? 09107 (????. 1),
6000
7,00
6,00
5000
5,00
4000
4,00
3000
3,00
2000
2,00
1000
1,00
0
0,00
09106
09107
09111
09112
09113
09119
09120
??????? ?????? ????
???????????, ???./??.?
????????, ?/??.?
???. 7. ?????????????? ?????????? ???????? ???????? ? ?????? ????????? ?????????? ??????????
?????????, ?????????????? ? ????? ??????? (???????? ??????). ?????????? ???????? ?? ???????
?????? ????, ??? ???????????? ????????????? ????????? ? ???????? ?????????
# 126 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ????????????? ????? ???????? (Annelida, Oligochaeta) ? ?????? ???????????????? ??????????
??? ???????? ????????? 4777 (????? ? 09119)
? 4624 ???/?2 (????? ? 09106). ????? ??????? ?????????? ???????? ? 5,79 ?/?2 (?????
? 09106). ??????????? ???????? ??? ????????? ?????????, ??? ? ???????? ???????? ????????????? ? ????? ? 09107 ? ??????????
350 ???/?2 ? 0,19 ?/?2 ??????????????.
??????? ???????? ???????? ?? ??????????
? ????????????? ???????????? ????????????
?????? ? ? ??????? ?????????? 2593 ???/?2 (???????? ? 4777 ???/?2, ??????? ? 350 ???/?2)
? 2,03 ?/?2 (???????? ? 4,06 ?/?2, ??????? ?
0,19 ?/?2). ??????? ?????????? ?????? ? ???????? ?????????????? ?????? ?? ???????
?????? ????????? ???? ? 3162 ???/?2, 3,55 ?/?2.
???????????? ???????? ??????????? ? ???????? ??????, ????????? ? ???????? ?????????, ? 4624 ???/?2 ? 5,79 ?/?2, ??????????? ?
1146 ???/?2 ? 1,15 ?/?2.
????????????? ?????? ???????????????? ??????? ????????? ???????? ? ??????
?????? ???????????? ??????? ???????, ???
?? ???? ?????? ???????? ?????????? ????
????????? Naididae sensu Ersйus et al., 2008
(??????? ???????, ?????? Tubificidae) ? ??
83 ?? 100 % (???. 8). ????????????? ??????
???????? ????????? ?????? ? ?????? ? 09106
(Lumbriculidae: L. achaetus ? 8 %) ? ? 09107
(Propappidae: P. glandulosus ? 17 %). ??? ????
????????? Rhyacodriloides abyssalis, Rhyacodrilus isossimovi ? Tasserkidrilus infundibuliferus ???????? ????? ??????????? ? ?????? ?????? ? ?????? ????????????? ????? (???. 8).
?????? ?????????????? ????????? ?????????? ?????? ? ?????? ? 09106 (8 ?????)
? ? 09120 (9 ?????), ??? ?????? ?????????
?????????? ????????? ????? (????. 1, ???. 8).
??????????????? ?????? ?????????? ???????? ? ? 09106 ?????? ???????????? ??? ??????? ??????????? ???????? (???. 7). ?????
???????????? ???????????? (L. achaetus),
??????? (Naidinae gen. sp.), ????????????
(P. confuses, R. isossimovi) ? ?????????? (4
????), ??? ?? ?????, ???????? ???????????
???? ???????????? Tubificinae (59 % ?? ?????
???????????). ???????? ????????? ??????
???????? ????????????? ???? Baikalodrilus
(37 % ?? ????? ???????????). ? ??????? ?
09120 ???? ?????????? ????????? ??????
???????? ??????? (6 ?????) ? ??????????????
??????????? ???????? ?????? (81 % ?? ?????
???????????), ???? 61 % ????? ???????????
?????????? ????? ????? ???? Isochaetides ?? 5
????? (I.adenodicystis, I. baicalensis, I. compactus, I. peniacerus ? Isochaetides sp.). ? 09113 ?
??? ???? ???????, ?????? ? ???????? ??????,
??? ????????? ?????????????? ???????????
(???. 7) ????????????? ???????? ??????? ?
??????? ????????? ?????????? ???????? ?
????????????? ???????????, ??????? ???????, Rhyacodrilus isossimovi (37 % ?? ?????
???????????) (???. 8).
? ??????, ?????? ??????????????? ? ????????????? ??????????? ??? ?????????????
????? ? ???????? ???, ???????? ?????? ????? ?????????????? ??????: Rhyacodriloides
abyssalis, Rhyacodrilus isossimovi, Pararhyacodrilus confuses, Baikalodrilus paradoxus, Baikalodrilus inflatus, Tasserkidrilus infundibuliferus,
Lamadrilus proprius, Limnodrilus sp. ? Propappus glandulosus (???. 8). ?? ???? ???????? ???????? ?????????? ????????????, ???? ?
????? ? 09119 ???????????? ?????????????
??????????. ?? ????? ????????????? ????????? ? ????? ? 09119 ?????? ??????? ????????? ??? ???? ?????????. Rhyacodriloides
abyssalis ? ???????? ???????? ??? ? ?????? ?
???????????? ?????????? ?? Thioploca. ????
??? ????????? ?????????? ? Rhyacodrilus
isossimovi (? ? 09112) ??? ? Tasserkidrilus infundibuliferus (? ? 09107). ?????????? 360
???/?2, Rh. abyssalis ?????????? 82 % ?? ?????
??????????? ???????? ? ????? ? 09111. ???????????? ??????????????? ????????????
# 127 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ????????????? ????? ???????? (Annelida, Oligochaeta) ? ?????? ???????????????? ??????????
???. 8. ??????????? ???????? ??????? ????????? ????????, ????????? ? ?????? ????? ???????.
?????????????? ? ???????????? ??????????????? ?????? ?????????? ??????? ?????????
????????, ????????? ?? ????????? ? ?????????, ??????????? ? ????? ? 09112 (5 ?????)
(???. 8). ?????????? ????? R. isossimovi (51 %
?? ????? ???????????), Rh. abyssalis ????????
?????????????? ????? (35 % ?? ????? ???????????), ????????? ??? ????????? ????????? 520 ???/?2.
??????????
?????? ?????????????, ????????? ??????? ?????????????????? ?????, ???????? ??
???????????? ????????????? ??????????
??????? ??????? ?????????????? ????????
??????? ????????? ?????? ?????????????
?????????? ????????? (???????? ? ??., 1991).
??????? ? ????????? ??????? ? ??. (1998)
???????????, ??? ???????????? ??????? ?
???? ?????? ???????? ?????????, ?????????-
??? 69% ??????????? ? 61% ???????? ???????????????. ? ????????????? ???? ??????
??? ?????????? ????? ????? ???? ??????????? ? ????? ??? ???????? ????????? ???????
(??????????, 1987). ?????? ?????????????? ????? ???????????? ?? ????????? ????????????, ?? ??? ????????? ????????????
??????? ?????????????? ??????????? ?? ???????? ? ??????????? ?????? (???. 7), ???????
?????????? ?? 350 ???/?2 ?? 4777 ???/?2 ?? ??????????? ? ?? 0,19 ?/?2 ?? 5,79 ?/?2 ?? ????????.
??????????? ???????? ????????????? ??? ?
?????? ? ???????? ?????? ? ??????? ????????????? «??????» ? 09107, ??? ? ? ??????,
?????????? ? ?????? ??? ????????????? ???????? ? 09113. ????? ??? ?? ?? ???????????
??????????? ?? ?????? ???????? ? ?????? ?
????????????? ?????????? ??????????????
# 128 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ????????????? ????? ???????? (Annelida, Oligochaeta) ? ?????? ???????????????? ??????????
?????????????: ? 09106 (????????? ?????)
? 09119 (????????????? ????????????? ???
(???. 7). ? ??????? ??????? ???????? ???????? ?? ?????????? ? ????????????? ???????????? ????????? ????????? ???? ??????? ??
??????? ??????. ????? ???????, ??????????
????? ??????? ???????? ? ????? ?????????
?? ???????????. ???????? ?? ??????????????
??????, ????? ??????? ?????, ??? ????????????? ?????????? ???? ?? ???? ?? ???????? ??????????????? ??????????? ????? ???
???????? ?, ??? ?? ?????, ? ?????? ????????????? ????????? ??????????? ?????????
???????? ??? ????????????? ???? ???????
?????????? ?????????????? ??????.
????????? ???????? ??????? ?????? ????????? ???????? ? ?????? ?????? ????? ?.?.
?????????? (1987) ????????? 30 ????? ????????, ????????? ?? ???????????? ???????? ? ??? ??? ???? ????? ????????? ???????;
??????????? ?? ??? ????????? ? ?????????
Lumbriculidae. ??????? ???? ?????? ??? ???????? ??????? ?????? ??? ????? ??????
(????????, 2004). ????? ???????, ?? ?????????? ??????? ??????????? ?????? ?????
????????????? ???? ???? ???????? ????????? ??????? ??????? 40 ????? ????????, ??
??????? ????? ???????? ?????????? ????????????. ?? ????????? ??????? ????????? ??????? ? ???? ????????????????? ??????????? ?????????-?????????? ????????? ??
??????? 21 ??? ???????? (????. 2); ??????????? ??????????? ?? ??? ????????? ? ????????? Tubificidae (???????? ? ?????? Naididae ?
????????? ?????), ?????? ??????????????? ?
???????, ? ???? ???? ??? ???????????. ?????? ????? ???????????????, ? ????? ???????, ? ????????? ? ???????????? ????? ????
?????? ?????????????? ???????? ? ????
?????????? ??????, ? ? ?????? ? ? ??????????????? ????????????? ???????? ??? ?????? ????????? Lumbriculidae, ??????????, ???
????????, ?? ????????????? ????????????
?????????? (????????, 1962).
??????????????? ????? ??????????????? ?????? ?????????????? ????????????
?????????, ????? ????????, ??? ??????? ???
????? (90 %) ?????????? ?? ???? ??????????
??????????? ?????, ?? ????????? ? ???? ? ????? ??????? ???????, ????? ??? Isochaetides
baicalensis ? Propappus glandulosus; ??? ????
?????????? ????? ? ????? ??? ?????????????? ? ???????? ??????? (???????????, 1962;
????????, 2004). ????? ?????, ????????? ?
????. 2, ????? ???????????? ??????????????? ? ?????????? ???? ??? ????? ?????????
???????. ??? ???? Lamadrilus proprius ? Rhyacodrilus propus, ????????? ??????? ?? ???????? ??????? (????????, 2004), ??????? ?????????? ? ???????? ????????? ?????. ???? ?????
(Baikalodrilus inflatus, Isochaetides adenodicystis, Rhyacodrilus isossimovi, Rhyacodriloides
abyssalis ? Lamprodrilus achaetus) ????????
??????????? ? ??????? ??????????????
???????????????? ?? ???????. ????????
????????????? ????????? ???? ??? ???? ??
?????????? ???????: Tasserkidrilus infundibuliferus, Rhyacodrilus vasalatus ? Pararhyacodrilus confusus. ?????? ????? ?????? ?? ?????????? ??? ??????????? ????. Isochaetides
adenodicystis ? Rhyacodrilus vasalatus ???? ?
????????????? ? ???????????? ????????,
??? ?? ????? ??????? ?????????? ?? ???????
????? 360 ?. ????? ????, ?? ???????????? ????????? ??????? ???????????? ??? ???????
?????? ????? ?????????????? ??????? ?????????????? (Naidinae gen. sp., Baikalodrilus
sp., Isochaetides sp., Limnodrilus sp.), ???????
??????? ??????????????? ????????, ??? ???,
?????? ????????, ??? ????? ??? ????? ????.
??? ?? ??? ? Naidinae gen. sp. ? Limnodrilus sp. ? ????????? ? ???????, ??????????
?????? ??????????? ??????? ? ?????? ?????
????????????? ?????? 20 ?.
# 129 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ????????????? ????? ???????? (Annelida, Oligochaeta) ? ?????? ???????????????? ??????????
????? ???????, ??????? ? ????? ???????
??????????? ????????? ???????, ??-????????,
??????? ?????, ??????? ? ???????? ?? ?????
???????? ???????? ???????? ?????????, ???, ?
???? ???????, ?????????? ???? ????, ????????????? ? ????? ??? ???????????? ??????. ?
?????????? ? ?????????? ?????? ???????????? ?????????? ?????????? ????? ??????????
????, ? ??????? ???????????? 11 ????????
??? ????????, ??????? ??????????, ? 10 ?????????????? ?????.
? ???? ?????? ?? ?????????? ????????
?? ?????? ?? ??????, ????????? ????????
??????? ?????? ???????????? ?????????
??????, ?? ???????? ??????? ?? ??????????
???????? ?? ???????????? Thioploca.
?????????? ????????????? ??????????,
??? ???????? ???????? ? ????????????
????????? ? ?????? ????? ???????? ??????
???????? ????????????? ???????, ?????????????? ??????????, ? ?????????? ????????
?????????????? ??????? ???????? ?????????
???, ??????????? ?????? ?????? ?????????
??????? (???????? ? ??., 1991). ?????????, ???
??????? ???? ? ??????? ???????? ??????
????????. ?????????, ????????? ????? ?????
???? ??????????????? ???????, ??????????
???????????? ???????? ?? ?????? ????????
????????????? ??????????, ?? ? ???????????? ????? ????? ??????????????? (??????,
1958; ???????????, 1962).
??? ??????????? ???? «????????????»
????????????? ?????????? ? ???????? ???????? ??? ???????? ????????????? ?????? ???????????????? ??????? ????????? ???????? ?
?????? ?????? ???????????? ???????. ???????, ??? ?? ???? ?????? ??????????? ????,
????????????? ????????????? Tubificinae
? Rhyacodrilinae, ???????? ?? 83 ?? 100 %
???????? ???????????? ????????? ????????
(???. 8). ????????????? ???????? Lumbriculidae (L. achaetus) ? Propappidae (P. glandulo-
sus) ??????? ?????? ? ???? ??????: ? 09106
? ? 09107. ???????? ????? ?????????????
??? Rhyacodriloides abyssalis ???????? ?????????? ?? ??????????? (???. 8). Rhyacodrilus
isossimovi ? Tasserkidrilus infundibuliferus ?
???????? ????, ?? ????? ??????????? ? ?????
?????, ????????, ?????????? ?? ??????. ??????? ?????????? ??????????, ????? ???????,
??? ? ?????? ? ??????? ??????? ???????????
?????? ????????????? ? ??????????? ?? ???????? ?????? ?????????? ?????? (????. 1,
???. 8). ????? ?????????? ????? ??? ???? ??
?????? ???????????? ?????? ??? ????? ??????
?? ??????????? Baikalodrilus inflatus, Propappus glandulosus ? Lamadrilus proprius.
??? ?????????? ????, ? ??????, ?????? ?
????????????? ??????????, ???????? ????
9 ????? ?? ????? ?????????? ??????? ?????????????? ?????? (????. 1, ???. 8). ? ??????????? ?? ???? ???? ????????? ???????????? ? ??????? ????????????? R. abyssalis
? ?????? ?? ????? ????????????? ?????????
? ????? ? 09119 ?????????? ????????????? ?????????? ??? ???????? ???????? T. infundibuliferus.
????????? ??? ??????????? ???? ??????????? ? ????? ? 09107 ????????? ???
???????????? ?????????? P. glandulosus, ????????????? 17 % ?? ????? ??????????? ?????????? ????????, ?????????? ? ???? ?????.
L. proprius, ??????????? ????? ??? ?????????
???????????????? ???? ?????? ???? (??????? ??????), ??????? ??????????? ??? ????????? ???????. ??? ?????? ?????? ? ?????? ?
??????? ??????????? ??????? ????????, ???
??? ?????? ?????????? 9 ? 10 % (???. 8).
??????????
??????????? ??????????? ?????? ????????????? ????? ???????? ????? ???????
?? ?????? ???????????? ? ????????? ??????
???????? 23 ????. ? ???? ?????? ?? ?????-
# 130 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ????????????? ????? ???????? (Annelida, Oligochaeta) ? ?????? ???????????????? ??????????
??? ???????? ?? ?????????? ??????????
?????????????? ??????, ????????? ?? 17
????? ???????? ? 4 ???? ???????????????
???????? ???????, ????????? ?? ????????,
??????????? ? ?????? ????????????? ??????????? ?????????, ?????????? ?????????
??????? ????? ?????? ?????. ????????????
???? ???? ??????????? 11 ?????, ???????,
? ???? ???????, ????????? ? ???? ?????????? Oligochaeta, ?? ??? ? Naididae sensu
Ersйus et al. (2008), ??????? ??? ????????
???????????? Naidinae, Tubificinae ? Rhyacodrilinae ? 19 ?????, Propappidae ? 1 ??? ?
1 ??? ?? ????????? Lumbriculidae. ??????
???????????????? ????????? ???? ????????
????? ?????????. ???? ??? Propappidae ??
??????????? ???????? ?? ?????????????
????????, ?? ??? ?????????????? ????????
????????? Lumbriculidae ???????????? ????????????? ????? ????????? ??????????????? ???????? ?????.
? ?????? ??????????? ????????? ???????? ????????????? ??? ??????????? ????
??????? ??????? ?????????????? ?????????? ???????? ????????? ????????. ????????,
??? ????????? ????????? ? ????? ???????? ?
??????????? ?????? ????????? ??????????
?????????? ? 2-3 ????. ??? ?? ?????, ???????????? ???????? ??????? ????????? ????????
? ?????????? ?? ???????? ???????? ?? ??????? ?? ???????? ???????? Thioploca.
?????????????
???????
???????????????
?.?.?.
?.?. ?????????? (??? ?? ???) ?? ??????????????? ????????.
?????? ????????? ??? ?????????? ????????? ???? (?????? ? 09-04-00781,
11-04-90452_??? ? 11-04-00194).
?????? ??????????
?????? ?.?., ???????? ?.?., ????????? ?.?., ?????? ?.?. (1993) ???? ????????????? ???????? ? ??????? ????????????? ???????? ? ????? ??????? (??. ??????) ? ???????? ??????????? ????????? ??????. ???????? ???, ???. ????. 6: 903?908.
??????? ?.?. (1979) ???????? ????? ????? ??????? ????? ??????. ??????? ?? ???? 245(6):
658.
??????? ?.?. (1984) ????????? ???????????? ????????? ?? ??????? ??????? ??????????
???????????????????. ???????? ?? ????. ?????? ????? 1: 104.
???????? ?.?. (1962) ?????????????? ????? ????????? Lumbriculidae. ?: ??????? ?.?.
(???.) ?????????????? ????? ? ???????? ????? ??????. ?.-?.: ???-?? ?? ????, ?. 3 ? 127.
?????? ?.?., ???????? ?.?., ???????? ?.?. (1998) ?????? ????????? ? ???????????? ?
?????????? ????????????? ???????? ???????. ?: «???????? ?????????? ???????????????»,
????????? ??????????? «???????? ????????», ???????, 28?30 ???????? 1997, ???????????:
?????, ?. 61?63.
???????? ?.?., ??????? ?.?., ????? ?.?., ??????? ?.?., ???????? ?.?. (1991) ????? ? ??????? ???????. ??????????, ?????????? ?? ????????????? ???????????. ???????? ?? ????.
???. ????. 5: 766?772.
????????? ?.?., ????? ?.?., ???????? ?.?., ??????? ?.?., ??????? ?.?., ????????? ?.?.
(1994) ??????? ???????? ? ????????? ??????? ? ?????????? ????????????? ???????? ? ????????? ????? ????? ??????. ????????????? 63 (2): 344?341.
# 131 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ????????????? ????? ???????? (Annelida, Oligochaeta) ? ?????? ???????????????? ??????????
?????? ?.?. (1958) ?????????????? ? ????????? ?????????????? ??????. ???????????????: ???-?? ?? ????, 171 ?.
???????? ?.?. (2004) ????????? ????? ??????. ???????????: ?????, 528 ?.
?????????? ?.?. (1984) ????? ???? ???? Peloscolex (Oligochaeta, Tubificida) ?? ?????????
???????. ?: ??????? ?.?. (???.) ????? ? ????? ???????. ???????????: ?????, ?. 10?26.
?????????? ?.?. (1987) ????????? ????????? ???????. ???????????: ?????, 105 ?.
??????????? ?.?. (1962) ?????? ?????????????? ????? ????? ????. ?.-?.: ???-?? ??
????, 412 ?.
??????????? ?.?. (1975) ????? ?????????? (Oligochaeta, Tubificidae) ?? ???????? ?????
??????. ?: ??????? ?.?. (???.) ????? ? ????? ???????. ???????????: ?????, ?. 112?130.
Crane K., Hecher B., Golubev V. (1991) Hydrothermal vents in Lake Baikal. Nature 350: 281.
Ersйus C., Wetzel M.J., Gustavson L. (2008) ICZN rules ? a farewell to Tubificidae (Annelida,
Clitellata). Zootaxa 1744: 66?68.
Grachev M.A., Fialkov V.A., Nakamura T., Ohta T., Kawai T. (1995) Extant fauna of ancient carbon. Nature 374: 123?124.
Michaelsen W. (1905) Die Oligochaeten des Baikalsees. Wissenschaftliche Ergebnisse einer zoologischen Expedition nach dem Baikalsee in den Jahren 1900-1902. ? R. Friedlander und Sohn, KiewBerlin, Lief. 1 ? S. 1? 69.
Shanks W.C., Callender E. (1992) Thermal springs in Lake Baikal. Geology 20: 495?497.
Deep-Water Fauna of Oligochaeta (Annelida, Clitellata)
Near Hydrothermal Spring of the Frolikha Bay,
Northern Baikal (East Siberia, Russia)
Irina A. Kaygorodova
Limnological Institute, Siberian Branch
of Russian Academy of Sciences
3 Ulan-Batorskaja, Irkutsk, 664033 Russia
The aim of this paper was to describe the oligochaete fauna from underwater hydrothermal vent
affected zone of the Frolikha Bay (Northern Baikal), where the group is dominated in zoobenthic
communities. The paper provides information about a unique community of oligochaetes inhabiting
here. Uncharacteristic for the abyssal zone of Lake Baikal high indices of oligochaete development were
revealed. Possible dependence of species composition and quantitative characteristics of oligochaetes
on the development of bacteria Thioploca in the area were analyzed.
Keywords: Northern Baikal; hydrothermal vent; oligochaetous worms; species composition; species
diversity.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2011 4) 133-141
~~~
??? 599.742.4 (235.32)
?????? ? ???????? ???????
???????????? ??????
?.?. ?????????*,
?.?. ????????, ?.?. ????????
?
???????? ????????????? ??????? ???????????? ?? ???,
?????? 677000, ??????, ?? ??????, 41
?
?????-????????? ??????? ?????,
?????? 677000, ?????????? ??????,
?. ?????-?????, ??. ???????, 7 1
Received 3.06.2011, received in revised form 10.06.2011, accepted 17.06.2011
??????? ????????????? ?????????, ???????????? ? ??????? ? ??????????? ????? ???????
????????, ??????????? ??????? ? ??????????? ?????? ? ???????? ????????? ??????? ????????
??????? ???????????? ??????. ??????????? ???????????? ??????? ?????????? ?????????
? ??????? ????? ????????? ? ?????????, ?????????? ?????????? ????? ??????? ???????? ?
???????????? ? ???????? ????????? ??????? ?????????? ?????, ?????????? ??????? ????????
????????????. ?????? ? ?????? ???????????? ?????????? ????? ???? ?????? ?????????.
????????? ?? ??????????? ???????? ?? ?????? ??????????? ????????? ?????. ? ?????????
???? ?????????? ???????? ???????? ????? ????????? ????????. ?????????? ????????????
????????????????? ????? ??????????????? ????????, ??? ??????????????? ??? ???? ??
?????? ?? ?????????? ?????? ??????????.
???????? ?????: ??????, ??????? ? ?????????? ?????????, ???????????, ?????????????
????????????, ???????, ???????????.
????????? ?????? ? ???????? ???????
???????????? ?????? ??????? ? ??????????
??????? ????????? ???????, ?????????? ??
??????????? ? 1951-1958 ??. ???????????
????????????? ? ??????????? ????? ????????? ??? ??? ??????? ??????????? ?????
????, ????????? ??????????? ? XVIII-XIX
??. (?????????, 1965, 1980; ?????????? ? ??.,
*
1
1971). ????????? ????????? ? ?????? 1960-?
??. ?????????? 0,6-1,0 ???. ?? 1000 ?? ??? ??????? ??????? ???????? 30 %. ???????????
???????? ?? ??????? ?????? ? 1961 ?. ? 19701980 ??. ?????? ??????? ??? ????????? ???
???????? ?????????, ????????? ?????????
???????? 0,8-2,1 ???. ?? 1000 ?? (???????? ?
??., 1985). ???????? ???? ? ???????? ????-
Corresponding author E-mail address: vmsafronov@ibpc.ysn.ru
© Siberian Federal University. All rights reserved
# 133 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????? ? ???????? ??????? ???????????? ??????
?????????, ?????????? ? ??????? ??????
??????, ??????? ????????????. ? ????????? ?????? ??????????????? ????????? ??
????????? ?????????, ????????????, ??????? ? ??????????? ?????? ? ???? ??????? ?
??????? ??????.
????? ?????, ????????? ? ??????
????? ???????????? ?????????? ?
??????-?????????
??????
(64-68°?.?.,
124-132° ?.?.). ?? ??????????????????? ????????????? ?? ????????? ? ???????? ????????? ?????- ? ??????????? ??????????? ??????
???? (???????, 1965).
?????? ????? ???????????????. ??????? ???????? ??????????? ??????? ? ????
?????????? 17,9, ? ?????? ? ????? 39,7 °C.
?????????? ???????? ??????????? ??????? ????????? 35, ??????? ? ????? 63°.
???? ?????? ????? 3 ??????? (????-??????),
???? ? 7 ???????. ??????? ?????? ????????
? ??????? ?? ?????? ??? ? ???????????????
????????? ????????? (30-52 ??). ? ??????????????? ????????? ????? ???????????
??????? ????????????? ????????????? ????? (????????, 1975). ?????? ???????? ?????
????????? ?????????, ?????? ??? ? ?????? ???, ??????? ?? ??????? ?????, ????????
???????? ?????????. ? ??????????? ???????? ????????? ????????????? ???? ? ?????????? ??? ?????????-??????? ????????,
????????????-??????????? ?????????? ?
????????? ?????????????? ? ???????? ???.
??????????? 761 ????? ??????, ?????????? ? 2004-2010 ??. ?? ?????? ??????? ?????????????. ??????? ???????? ?????????? ??
???????? ????? ? ??????? ?????? ???????
???????. ????????????? ????? ????????? ?
5 %-??? ???????? ??????? ??????? ? ???????
24 ?. ??? ????????????? ?????? ????????????
?????? ???????? ??-2 ? ??????????????
?????????????????? ???????? ???-28-02?.
?????????? ?????????? ????? ??????????
?????????????? ?????? ? ????????? ? ???????? (?????????, 1988).
????????????? ???????????? ????? ??????????? ?? ?????????? ?????? ??? ???????????? (???????, 1956). ????????? ????????
???????? ???????????? ??????? ?????????
?????????? ?? ????? ?????, ????????????
?.?. ????????? (1988) ??? ?????? ???????????? ????? ? ??????????? ???????????? ?????????, ???????????? ? ?????? ????? 225260 ???? ????? ?????????????? (???????,
1973). ?????????????? ??????? ???? ?? ??????? ??????????? 193 ???????? ?? ???????????? ????? ???????. ???????? ?????????
????? ????? ?????????? ?? ?? ?????????????
? ?????????? ???????? ????????, ?????????? ? ?????????. ??????????? ????????? ??
?????????? ? ????????????? ????? ?????????????? ???? ? ???? ????? ??? ?????? (?/??)
(????? ? ??., 1968), ?????? ????? ????? ??????????????.
??????? ???????? ? ?? ??????????????
?????? ?????????? ?? ????????, ?????? ??
????????? ??????????? ????? ??? ? ?????
?? ?????? ????????????. ??????????? ?????? ?????????????? ?????????? ????????? ?
???????? ????????? ??? ????????? ?????????????? ????????????. ???????? ???????
??????? ????????? ???????????? ??? ??????
?????????? 0,05 (?????, 1980).
?????????? ? ??????????
? ????? ????????????? ????? ???? ????????? ??????????? ?? ??????????? ??????? ?? 55,2±6,5 ?? 66,1±3,7 %, ? ??????? ???
?????????? 61,6±1,8 % (???. 1). ? ????? 1950-?
??. ???? ?????????? ???????? 29,3 % (?????????, 1961), ? 1980-? ?? ????????? ?? 46,8 %
? ?????????? ????? ? ????????? ?????, ???
? ???????????? ???? ??????? ? ??????????? ????????? ??? ???????????? ????????
# 134 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????? ? ???????? ??????? ???????????? ??????
% 80
70
60
50
40
30
20
10
0
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2005/06
2006/07
2007/08
2008/09
2009/10
n= 58
25
221
188
107
162
????????
?????
????????
???. 1. ????????????? ?????????? ????????? ? ????????????? ????? ?????? ? ????????????? ? 2004-2010
??. n ? ????? ? ????? ?????????? ?????????? ????????????? ?????. ???????????? ????? ? ???????????
??????
(???????, 1973; ???????, 1983; ????????,
??????, 2000; ? ??.). ?????? ? ??? ??? ??????? ???????? ????????????? ???????????
???????????????? ??????, ?????????????
????????-?????????????? ?????????.
????????????? ????? ????????? ?????????? ????? (55,0±2,3 %). ? ????????? ??????????? ????? ?? ???? ??? ????? ????????????? (69,5±2,7 %) ? ????? ????????? ?????
????? (30,5±2,7 %; ?<0,01). ? ????? ???????
????? (n=300) ???? ?????? ?????? ???? ?????????? 29,7±2,6 %, ?? ??????????? ??????? ??? ?????????? ?? 19,5±6,1 ?? 39,7±6,4 %.
? ?????? (n=461) ???? ?????????? ??? ????
(44,0±2,3; ?? 38,6±5,2 ?? 56,1±6,0 %).
??????? ????? ??????? ???????, ???????????? ?? ???????? ?????? ? ??????? ?????????? ???????, ???????? ? ??????? 71 %.
?? ??????? ?????????? ? ?????? ????????????????? ? ??????? ?????? ???????????
??????????. ? ??????????? ???????? ?????-
????? ?????????, ??, ??? ?? ?????, ??????????
???????. ?? ?????? ???? ????? ??? ?????????? 42, ?? ??????? ? 59, ? ? ????????????????
?????? ?? ???? ?? ???? ??? ? 45,4 %. ?????
??????? ???????? ???????? ? ?????????????
???? ??????????? ??????, ??? ? ???????? ?
???????????? ?????? ????????. ?? ??????
?.?. ???????? (1983), ? ???????? ????????
?????????? ????????? ????????? 79 %, ????????? ?????? ? 58, ? ????? ?? ???? ?? ???? ???
??? ??????????? ?? 16-31. ?? ???????? ?????????? ?????????? ????? ????? ??????????? ?
19, ????????? ? ? 12, ? ??????? ????????? ?
?????? ? 6 % (????????, ?????, 1973).
?????????? ???????????? ????? ??????? ???????? ???????? ? ????????????? ????? ??????????? ???????? ??????? ??????
??????? ?????????????. ? ?????????? ????? ?
?????????? ??????? ?????? ???? ??????????? ????? 1+ (46,2±2,9 %) ? 2+ ??? (26,7±2,6 %).
??????? ? ???????? 3 ??? ? ????? ???????? ?
# 135 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????? ? ???????? ??????? ???????????? ??????
% 35
30
25
20
15
10
5
0
1+
2+
3+
4+
5+
6+
7+
8+
9+
n= 135
78
32
18
10
7
5
1
4
?????
10+ ? ???????,
???
??????
2
?????
???. 2. ??????????? ?????????? ????? ????? ???????? ??????? ? ????? ????? ?? 2004-2010 ??.
???????????? ???? 27,1±2,6 % (???. 2). ???????? ?????? ???? ????? ???????? ?????,
????????????????? ??????? ?????????????? ??????????????? ??????? ? ?????? ??????, ??? ? ?????? (???????, ???????, 1976).
? ?????? ???????? ?? ???? ?? ???? ??? ?????
?????????? ???? 10,9 %, ?????? ? 19,0 % ??
??????????? ????????? ???? ?? ????.
?????????? ?????? ??? ????????????
? ??????? ?? ???? ?????????? ????? (n=52)
????????? 3,29±0,12. ? 2006 ? 2007 ??. ??????????? ????????? ????????? ???????????? (3,53±0,27 ? 3,43±0,23), ?? ?????????
?????????????? ?????????? (???. 3). ???????
???????? ????????????? ???????????? ? ????????? ????? ??????, ??? ?? ?????? ?????? ?????????????? ???? ? ?????????????
? 1960-? ??. (2,96±0,05; p<0,01) (?????????? ?
??., 1971). ??? ????, ??? ? ???????? ????????? ?????? ? ???????????? (2,93±04; p<0,01)
(???????, 1971). ?????????? ?????????????
???????? ????????, ?????????????? ????-
?????? ???? ????????? ? ??????????? ??????, ????? ????? ?????????? ???????? ???
??????????????? ????????? ??????, ????????? ?? ?????? ??????.
? ???????? ????? ???????????????????
???? ?????? ?????????? ????? ? ???????? ?? 1
?? 7 ???. ? ???????????? ???????? ??????????
????? ????? ???????????, ? ??? ?????? ?????????????? ???? ? ??????????????? (???????, 1979). ? ????????????? ??????????????
???? ???????? ????? ?? 1 ?? 5 ??? (94,4 % ??
????? ???????? ?????), ????? ??????? ??????????? ????-?????????? ????? (71,9±4,7 %) ?
???????? ??????????????? ?????????????,
??? ??????? ???????? ????????.
????????? ?????, ??????? ???????????? ? ???????????, ????? ?????????
(2,81±0,21), ??? ????? ???? (3,4±0,21; ?<0,05)
? ????-?????? ??? (3,57±0,18; ?<0,01). ????????? ??????????? ??????? ??????? ? ??????????? ??????????????? ?????. ????? ?????????
???? ?????????? ?????????? 41, ????????? ?
# 136 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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% 70
60
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2006/07
2007/08
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25
221
188
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162
?????
???. 3. ????????????? ?????????? ??????- ? ??????????????? ??????? ? 2004-2010 ??.
60, ? ????-?????????? ?????? ? 87 %. ?????? ?? ????? ????? ? ???????? 1+ ????? ????
26,3 %, 2+ ? 29,9, ????-???? ??? ? 40,9, ? ??????????? ? 2,9 ???????? ?????????. ?????
? ??????????? ??????????? 58,4 % ???????? ?????. ???????? ????? ??????????????
???????? (56,2 %) ??????????? ?? ????????
??????????????, ???? ? ????? ???????????? ????- ? ????????????? ?????. ?? ??????
?????? ?????????? ??????? ????? ????????
???????????? ???????????????? ?????; ??????? ?????????? ???????????? ? ??????? ?? ?
??????????? ????????? ?????????????? 4,2-5
? 74-97 % (????????, ?????, 1973; ???????,
1990; ? ??.).
????????? ???????????? ?????? ? ??????? ?? ???? ??????????????? ????? ?????????? 2,72±0,23 (n=29) ? ???? ???? ????????????? ???????????? ?? 17,3 %. ??????? ???????
????? ??????????? ?????? ??? ? ??????????
??????????? ? ???????????? ???? ? ? ??????????? ??????? (16,5 %; ???????, ????????,
1992).
?????? ???????? ???? ??????? ?????????? ?????????? ???????, ????????????
? 81,7 % ????????????? ????????. ???????
??????? ??????? ???????????????? ? 26,4 %
???? (?? 8 ?? 53,8 %), ??????-????? ??????? ?
? 20,7 % (?? 7,3 ?? 25,6 %), ???????-????????? ?
? 18,6 % ???????? (?? 2,4 ?? 29,7 %).
?????? ????????????? ? ?????????? ?
?????? ???? ???????????? ???????. ??????
????? 2007/08 ??. ??? ??????? ? 25,6 % ????????. ????? ???????, ? ????????? ???? ?????????? ????? ??????????? ??????? ????????
? ??????? ??????, ??? ????? ?????????? ?
??????? ??????? ?. ???? (??????, ????????,
1955). ???????????? ??????? ????????????? (????-?????, ???????, ???????? ??????,
?????), ?????????? ?????????????? ??????
?????? (0,5-4,7 % ?????????????).
? ????????? ???? ? ??????? ?????? ????????????? ???????? ???? (20,0 % ?
2004/05 ??., ? ????????? ???? 9,8 %), ?????
?? ????????? ?????????? ????????? ???????
????? ? ?????? ???????.
???????????? ????? ????????????? ?
???? ???????? ????????? (40,9 %) ? ???????? ? ???????? ?????? ????????? ???????? (23,8 %), ????? ???????? (6,7 %) ? ???????? (6,2 %). ? ?????? ??????? 2004/05 ?
2008/09 ??. ??????????? ?????? ??????????? ?
# 137 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????? ? ???????? ??????? ???????????? ??????
????????????? ?????, ?/??
2,5
2
1,5
1
0,5
0
2004/05
2005/06
2006/07
2007/08
2008/09
n= 11
11
70
100
29
2009/10 ?????
31
???. 4. ????????????? ????? ???? ?? ?????? ??????? ????????????? ? 2004-2010 ??.
52 ? 58,5 % ????. ? ????????? ???? ??? ???????????? ?????? (5,1-18,9 %), ? ????? 2006/07
??. ?????? ?? ?????????. ??? ???????? ???????????? ????????? ???????? ??????????? ??????? ????? ????? ???????. ????????,
? ??????-?????? ?????? 2008/09 ??. ???????
?? ?????? ????????????????? ? ???????????
???????? ? ??????????? ?? ???????????, ???
??????, ?? ????????? ??????????. ????????
??????????? ??????? ????????? ? ?????????? ?? ?????????? ?? ????????? ????????
??????????, ??????? ??????????? ????????.
????? ???????? ??????????? ? ??????-??????
??????? 2004/05 ? 2008/09 ??., ?? ?????????? ????????, ??? ??? ?? ???? ???????????
??? ????? ?????????????? ?????? ? ????? ?
????????? ???????????????? ???????????? ??????????? ?????????? (??????, 1957;
???????? ? ??., 1985). ? ???? ? ?????? ????????????? ????????? ????????, ??????? ??????????? ? ????????????? (????????, ????????, 1986), ??????????? ? ???????????????
?????? ????????? ???????? ?? ???????????
?????????? ????????.
? ????? ??????? ????? ? ????????? ??????????? ??????????? ???? ? ??????? ??-
??? ?????????? ? ??????? 35,7±1,7 %. ???
????????? (64,3 %) ?????????? ? ??????????????. ? ?????? ?????? ?????? ???? ???????, ??????? ?????????? ??????? ?????????, ?????????? ?? 18,9±5,1 ?? 54,2±3,6 %
(???. 4). ?? ???????????? ?????????? ??????????? ????? 2007/08 ??., ????? ? ???????
??????????? ??????? ??????? (53,8 % ?? ?????
????????????? ????????).
????? ?????? ????????? ??????
???? ?????? (39,5±2,3 %), ??? ????? ????? (30,0±2,6 %; ?<0,01). ??? ?? ?????, ??????? ????????? ??????????? ?????? ?
????? ????????? ????????? (r=0,88±0,24;
?<0,01,
n=300).
?????????????
????? ???? ?? ????? ?????????? ? ???????
1,44±0,08 ?/??. ?? ????? ??? ??????????? ??
0,75±0,13 ?? 1,83±0,34 ?/?? (???. 4). ? ?????? ???? ?????????? ???? (1,5±0,1 ?/??),
??? ? ????? (1,29±0,1 ?/??). ??????????
?????? ???? ??????? ? ???????????? ????? (1,12±0,16 ?/??), ?????????? ? ? ??????? ?????? (1,58±0,15 ?/??; ?<0,05). ???????? ????? (1,4±0,16 ?/??) ? ??????? ?????
(1,39±0,14 ?/??) ???????? ?? ????? ?????????? ????????????? ?????????.
# 138 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????? ? ???????? ??????? ???????????? ??????
?????????? ??????? ??????????? ??????????? ? ?????? ?????? 2004/05, 2008/09
? 2009/10 ??. ??? ?????????? ???????????
????? ????????? ???????? (??????????????,
52,0, 58,5 ? 18,9 % ????????????? ? ??????).
??????????? ???????? ????? ?????????? ???????? ????? 2005/06 ??., ????? ???????????
?????? ?????? ?? ????????? ????????. ?????????? ??????? ???? ???????????? ?????
???? ? ???? ???????? ???????????? ????????
???????, ?? ???? ?????????, ?? ??????? ? ??????????? ??????? ??????? ????????, ??????? ?? ????????? ?? ??????????? ? ???????? ??????????????? ?? ????????? ????? ???
???????? ? ??????????? ???????????.
??????????
????????? ?????? ? ???????? ???????
???????????? ?????? ??????????????? ??????????? ? ???????????? ??????? ??????????
??????????, ?????????? ??????? ????????
????????, ???????? ????? ? ?????? ????????, ?????????? ????? ?????????. ?????? ????????? ????????? ?????????? ????? 1-5 ???,
????? ??????? ??????????? ??????? ???-
??????? ?????? 1-2 ???. ????????????? ????
?????????? ????????? ??????????????? ?
?????????? ??????????? ???????????????
???? ?? ??????????? ?????????? ??????????
????????? ???????????? ? ?????????????
?????????????? ?????????. ????? ?? ?????????? ???????????? ??????????? ?????? ?
???? ???????? ?????? ?????????? ????????????? ????????????, ?????????? ? ????????? ??????. ? ??????????? ????? ??????????? ?????.
?????? ??????? ?????? ? ?????????????
?????????? ?????????? ???????. ?????????????? ???????? ????????, ????????????? ? ????????? ????, ?????????? ???????????? ????????????????? ????????? ??
??????????, ???????????? ??????????? ??
?????????? ??????????????? ?????????? ?
???????????? ????????. ?????????? ??????? ?????? ???????? ? ???????? ?????. ????????????? ?????????? ?????????????? ??????? ????? ???? ????? ?? ?????? ?? ?????
???????, ??? ? ??????, ?????????? ? ????????????? ?????? ???????? ?????????????
???? ??????.
?????? ??????????
???????? ?.?., ???????? ?.?. (1986) ? ??????????? ??????????????? ?????? ???????
????? ???????? ? ?? ???????? ??? ????????????? ????????????. ??????????, ??????????? ?
?????? ???????. ?????? ???????? XI ??????????? ?????????? «????????????? ???????? ??????», ???. 3. ??????, ?. 7-8.
???????? ?.?., ????? ?.?. (1973) ??????. ???????? ? ?. ???????????. ?: ??????, ??????,
?????. ?.: ?????, ?. 118-131.
????????? ?.?. (1961) ????????? ?? ??????????????? ??????? ?????????? ?????? ? ??????
???-???????? ??????? ???????????? ??????. ?: ??????? ????????? ????????? ??????? ??
?? ????, ????????, ???. 5. ??????, ?. 71-77.
????????? ?.?. (1965) ???????????????? ?????? ? ???? ??????????? ?????????? ???????? ? ?????? ???-???????? ??????? ???????????? ??????: ???????. ???. ? ????. ????. ????.
??????, 22 ?.
????????? ?.?. (1980) ?????????? ???????????????? ?????? ? ??????. ?: ????? ? ???????? ???????? ??????????? ??????? ??????. ??????, ???????? ??????????????? ???????????,
?. 43-78.
# 139 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????? ? ???????? ??????? ???????????? ??????
??????? ?.?., ??????? ?.?. (1976) ????? ??????-?????? ????. ?.: ?????, 256 ?.
??????? ?.?. (1956) ????????? ?? ??????????? ? ???????? ??????? ? ???????. ???????
???????? ?????? ??????. ??. ?????, ???. 16. ?.: ???-?? ??????????? ? ????????????? ?????????? ?? ???????? ?????????, ?. 122-144.
????????? ?.?. (1988) ?????????????? ????????? ????????????? ? ????????????? ?????????????. ?.: ?????, 288 ?.
??????? ?.?. (1965) ?????? ? ????????????? ????????. ?: ??????. ?.: ?????, ?. 29-114.
????? ?.?. (1980) ?????????. ?.: ?????? ?????, 294 ?.
??????? ?.?. (1983) ?????????? ????????? ????????? ?????? (Martes zibellina). ????. ????.
9: 1398-1406.
??????? ?.?. (1990) ?????????? ??????????? ??????????? ??????? ??? ?????? ???????????? ?? ??????. ?: ????????????? ???????????? ???????? ?????? ??????. ?????: ????
??????????? ????????? ? ???????????? ??. ?.?. ???????, ?. 4-9.
??????? ?.?. (1971) ???????????? ???????????? ??????? ???????????? ? ??????????.
????. ????. ????????. 5: 40-44.
??????? ?.?. (1973) ?????? ? ???????? ??????????? ?????????. ?: ??????, ??????, ?????.
?.: ?????, ?. 36-38.
?????? ?.?., ???????? ?.?. (1955) ??????. ?.: ??????????, 403 ?.
??????? ?.?. (1973) ??????? ??????. ?: ??????, ??????, ?????. ?.: ?????, ?. 70-78.
?????? ?.?. (1957) ? ??????? ? ???????????? ????????? ???????? (Pinus pumila (Pall.)
RgI.). ???????????? ????., 42(5): 745-751.
???????? ?.?., ?????? ?.?. (2000) ???????? ??????, Martes zibellina (Carnivora, Mustelidae), ? ??????-????????? ??????. ????. ????. 4: 471-479.
???????? ?.?., ???????? ?.?., ?????????? ?.?. (1985) ????? ?????? ???????? ??????
(Martes zibellina L.) ? ???????? ?????????????. ?: ????? ? ???????? ????????????? ??????.
??????, ?? ?? ?? ????, ?. 24-55.
??????? ?.?. (1979) ????????????? ???????? ????? ??????. ???????????: ?????, 256 ?.
?????????? ?.?., ?????? ?.?., ????????? ?.?. ? ??. (1971) ????????????? ??????. ?.:
?????, 660 ?.
??????? ?.?. (1988) ??????? ? ???????? ?????? ??????????? ???????????? ?????????
?????????????. ?: ?????????????? ??????? ???????? ?????????? ????????: ????????????
????????????. ?????: ???? ??????????? ????????? ? ???????????? ??. ?.?. ???????, ?.127137.
??????? ?.?., ???????? ?.?. (1992) ???????? ??????????? ???????????? ???????. ?: ???????????? ????????????? ???????? ??????. ??????????. ????????? III ?????????????? ??????????????????????? ????????? 11-14 ??????? 1992 ?., ?. 86-87.
????? ?.?., ??????? ?.?., ?????????? ?.?. (1968) ????? ???????????????????? ??????????? ? ???????? ???????? ???????????. ??. ??-?? ???????? ???????? ? ????????, ???. 58.
??????????, 387 ?.
???????? ?.?. (1975) ?????? ?????? ??????-??????? ????. ???????????: ?????, 344 ?.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????? ? ???????? ??????? ???????????? ??????
Sable in the Western Spurs
of the Verkhoyansk Ridge
Valery M. Safronova,
Eugeny S. Zakharova and Afanasy P. Zakharovb
a
Institute for Biological Problems of Cryolithozone SB RAS
41 pr.Lenina, Yakutsk, 677000 Russia
b
Municipal Educational Institution Segen-Kyuel Secondary School,
7 ul. A. Kychkina, Segen-Kyuel settl., Yakutia, 677000 Russia
Population structure, fecundity and participation in breeding of females of different age, feeding trends
and fat condition of sable were studied under the conditions of the cold climate in the spurs of the
Verkhoyansk mountain ridge. A relatively simple age structure and rapid alternation of generations
in the population, increased mortality of mature age females and abundance of junior age cohorts in
the sable stock, larger mean value fecundity were determined. Sables with low fat reserves made up
over two thirds of the population. A change in their fat condition depended on the consumption level of
the animal food items. In some years seeds of the mountain pine were more important as a feed-stuff.
Females of the reproductive age had the least fat reserves that may be regarded as one of the reasons
of their increased death-rate.
Keywords: sable, aliens, age-sex composition, reproduction, potential fecundity, nutrition.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2011 4) 142-157
~~~
??? 581.17: 612.172: 535.372: 53.082.56
????????????? ?????????????? ?????????????
??? ?????? ????????????
??????????? ????????
?.?. ???????*, ?.?. ????????,
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?.?. ????????, ?.?. ???????, ?.?. ?????,
?.?. ????????????, ?.?. ???????, ?.?. ?????????*
?
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ?????????, 79
?
???????????? ??????????????? ??????????? ???????????
??. ????. ?.?.?????-??????????,
?????? 660022, ??????????, ??. ????????? ?????????, 1 1
Received 3.06.2011, received in revised form 10.06.2011, accepted 17.06.2011
???????????? ?????????? ?????????????????? ???????????? ??????????? ??????????
???????-??????????????? ??????? ???????? ??? ?????? ???????????, ???????????????
??????? ? ???????? ???????????????????? ?????????????. ????????, ??? ????? ??????????
??????? ? ?????????????? ???????? ?????? (337 ??) ????????? ?????????????? ??????????????
????????? ? ????????, ????????????? ?????????? ?????? ??????????? ???????????,
?????????? ??????? ?????????????, ?????????? ???(?)?-????????????? ????????? ?
????????? ??????????? ? ?????? ??????????????.
???????? ?????: ?????????? ???????, ???????, ??????, ?????????????.
????????
? ????????? ???? ? ???????? ??????????? ??????????????? ??????? ????????????
????????? ? ?????????? ??????? ???????????? ??????? ??????????? ??????????????
?????????? ??????? ? ??????. ??????????????? ????????? ?????? ???? ??????? ? ??????????? ???????? ??????????? ?? ???????
????????????????, ??????????? ?????????-
*
1
?? ????? ????????????, ???????????? ?????????? ?????? ? ?????? ????????????
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???????? ???????? «?????????? ???????»
(Optical Biopsy) (Bigio, Mourant, 2007). ???
?????????? ???????, ? ??????? ?? ???????
???????, ????? ?? ?????????, ? ???????????
Corresponding author E-mail address: vsalmin@gmail.com
© Siberian Federal University. All rights reserved
# 142 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ???????? ????????????? ?????????????? ????????????? ??? ?????? ?????????????
?? ??? ???? ????? ?????????? ?????????, ????? ??????????????????, ??????????? ??????????? ??? ?????????? ????????? ? ?????
????????? ??????? in situ ? ? ?????? ????????? ???????. ?????? ??????????? ?????????? ???????????? ? ?????????? ???????????
??????????, ???????????? ???????? ???????????????, ?????????????? ? ?.?. ? ????????? ? ????????? ??????? ??????? ??????????
??????? (Newton et al., 2011). ??????? ???????????? ????????? ???????? ???????????
????????? ???????? in situ ?? ????? ??????????????????? ???????????? ??? ??? ??????
???????????????? ?????????????????? ?????? (???????? ? ??., 2002; Khabbaz et al., 2001).
???????? ??????????? ????? ??????????????????? ???????????? ? ?? ????? ???
???????? ??????, ? ???????? ??????? ?????
???????????????? ????????? ????????: 1) ?????????, ???????????? ????????? ?????????
??????????? ? ????????? ? ????? ????????
(1-15 ?); 2) ?????? «?????????», ???????????? ????????? ?????????? ????????? ???
(5-6 ???); 3) ?????? ??????????? ?????????,
????????????????? ??????? ???????????
? ?????????? ????????? ??????????????
(?? 15 ??? ? ??????? ?????? ??????). ?????
????????? ? ???????????? ????????????
??????? ????????????????? ????????????
???????? ????? ?????? ??????????? ???????????????????? ???????????, ??, ????????????? ????????????? ???????? ???????????
???????? ? ????????????? ????????? (????????, ?????????), ???????????? ????????????
?????????????? ???????? ? ??????? ????????????? ? ?????????????? ???????? ????????????? ????????? (Nighswander-Rempel
et al., 2002; Mair, Hammemer-Lercher, 2005).
?????? ? ??? ?????????? ????????? ??????? ???????????? ??????????? ?????????, ?
?????????, ?????????????? ??????? ???????? ???????????? ???????? ? ??????, ???????-
?????? ????????????????? ??? ??????? ?????????? ????????? ??????????, ??? ???????
??????????? ?????? ? ????????????? ????
??????? ? ???????? ??????????? ????????.
?????????? ? ????????????? ?????? ?
??????? ??????????? ?????? ?????????????????? ????????????????? ? ????????????? ??????? ?????????? ???????, ? ????????????? ??????? ?????????: 1) ??????????
????????????? ? ?????????????? ???????????? ????? ? ???????????????? ???????????; 2) ??????????? ????????-???????????
? ??????? ??????????????? ???????; 3) ?????????? ????????????? ???????? ?? ?????
?????????? ??? ?????????? ???????; 4) ??????????? ????????? ????? ????????????
?????????? ?????????? ? ???????????????
??????? ? ??????; 5) ??????????? ???????????? ??????????? ???????????; 6) ??????????
? ?????????????? ?????????; 7) ??????????? ?????? ??????- ? ???????????? ??????? ????????????.
????? ??????? ?????????? ???????, ??????????? ??????????? ?????????? ???????? ?? ????? ?????? ??????, ????? ???????
????? ???????-?????????????? ????????????? (Renault et al., 1985). ? ?????? ?????? ????? ??????????? ????????? ?????????
?????????????????, ????????? ??????????????? ??????????? ??????????? ?? ?????
?????? (Chance et al., 1962, 1965).
????????, ??? ????? ?? ???????? ?????????? ????????? ????????????????? ???????? ?????????? ? ??????????? ? ??????????, ??????????? ? ???????? ? ????????????
(Riess et al., 2002). ??????????? ??? ????????????? ????????? ??????-????????? ????
??????? ???????? ? ????????? ?? ?????????????. ? ?????????? ?????? ???????????
?????????? ????? ???+ ? ??????????????? ????? ???(?)?. ?????????? ???? ?
???????????? ???????????? ???????? ???
# 143 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ???????? ????????????? ?????????????? ????????????? ??? ?????? ?????????????
?????????? ?????????? ? ??????????? ???? ?
????????????? ????????????????, ???????
? ??????? ?? ??????????????????? ??????????? ???+/???? ? ???????????? ?? ???
??????? ??????. ? ???????? ???????? ?????????? ???? ????????? ???????? ????????? ????????? ??????????? ????????? (Riess
et al., 2002; Brachmanski et al., 2004).
???????????????? ?????????? ???????? ? ?????????? ????????? ????????? ????????? ? ???????? ?????????????? ???????
(Grivennikova, Vinogradov, 2006), ?????? ?
????, ? ???(?)? ????? ???????????? ?????????? ????????? ???????, ????????? ? ???????????????? ?????????? ???????? ????
????????? ? ?????????????? in vitro (Takayanagi et al., 1980). ?????????, ??? ????????
?????????????? ??????? ??????????? ????????? ?????????? ???????????, ? ??????????
???????????? ???????????????? ???? ?
??? ????????????? ??????????? ??????????????????????? ??????? ? ????????????????? ??????????? ?????????? (Brachmanski
et al., 2004). ? ???????? ?????????????? ???????? ??????, ?????? ?? ??????, ????????????? ????????????? ?????????? ???? ?
???(?)? ?????????, ??? ???????? ?????????? ????????? ?????????? ??????????? (Xia
et al., 2009).
?????? ?????? ?????????? ????????????????? ?????????? ?????????? ???(?)?
???????????? ???????? ????? ????????? ????????? ? ?????? ? ?????????????? ??????????? ???????. ??? ???????? ???? ?????? ????????????, ? ????? ???????, ?????????????
?????????????? ? ?????????????? ??????????? ???????????????? ?????????? ???
«????» ????????? (Wittenberg, 2003), ? ? ?????? ???????, ??????????? ?????????????????? ????????? (?????????? ??????????????,
?????? ???-????????? ?????????? ????????????? ? ???????? ???????, ????????????-
???? ????????) ? ???(?)?-?????????????
????????? (????????, ???(?)?-????????).
????????, ??? ????????????? ??? ??????????????????? ?????????????? ????????????
?????????? ?? ?????????? ???????????????????? ?????????? ? ????????????? ??????
?????????? ???????????, ??? ?????????????
??????????? ??????????? ??????????. ?????
?????? ?????????? ?????????????? ??????????????? ????????, ??????????? ?????????
????????????? ??????? ?? ???????????????
???? ? ?????????? ? ??????????? ???????
?? ????????????? ???????????? ? ????????.
????? ???????, ?????????? ???????????? ?
???????? ?????????????? ?????? ?????????
??????????? ????????? ?????????? ?????????????????? ????????? ? ????????, ? ?????????? ???? ?????? ????????????????? ????
???????? ? ???????? ???????????????????
???????? ????????, ? ????????, ??????? ????????? ???????????????????? ????????? ?
????????????.
???????? ?? ??????? ?????????? ?????,
??????????? ???????????? ?????????????
????????, ???????? ??????????? ????????????????? ???????? ? ????????? ???????? ????????????? ? ?????????????? ?????????, ??????????? ? ????? ???????? ???
?????? ??????, ??????????? ?????????????
?????????????? ??????????? ??? ????????
????????? ???????? ??? ?????? ? ????????
??????????????? ??????? ????. T. Nishioka
(1984) ????? ?? ?????? ??????????? ?????????????? ????? ???????? ???????? ??? ????????????????? ??????. ???????? ????????????? ???? ????????????? ???????????
Omron Redoximeter. ???? ?????????? ?????
????? ????????? ????????????? ???? ?
?????????????? ?????? ? ?????????????,
???????????? ? ????? ????????????. ???
???? ?????? ? ??????? ?????????????? ?????? ????????????? ???????? ? ??????????
# 144 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ???????? ????????????? ?????????????? ????????????? ??? ?????? ?????????????
??????????? ????????? ? ????????, ? ???????? ?????????????? ? ? ??????? ???????. ?
??????????? ?????????????? ??????????????? ?????? ???????? ????? ??????? ??, ???
?????? ? ????????? ???????? ????? ????
???????? ?????? ? ?????????????? ???????.
????, ??? ??????? ?? ??????????? ?????????
?????? ???, ?????????? ????????????? ????????? ?????????? ? ??????? ????? ???????
??????. ??? ??????? ?? ??????????? ??????? ???????? ???????? ????????????? ??
????? ??????, ????????? ? ?? ????????????? ?? ???? ??????????. ?? ???? ?????????,
???????? ????????? ????????????? ????????????? ?? ????????? ???????????????? ?????
?????????.
???????? ?????-?????????????? ????????????? ???? ? ?????? ???????? ??? ??????????? ?????????? ???????? ?????? 337 ??
??? ???????????????? ???????? ? ????????? ??????????????? ???????????? ? ??????
(Horvath et al., 1994). ? ??? ???????? ??????????? ???????? ? ???????? ? ?????????
????????????? ? ????????? ????? ????????
(???? ????????, ??????????? ? ??????????) ?
??????? ????????. ??????, ??? ? ? ?????????? ??????, ??????????? ???????? ? ???????? ? ????????? ????????????? ?? ????? ?????? ? ????????? ????? ?? ???? ??????????.
??????????? ????????? ? ?????? ?????
460 ?? ??? ??????????? 350 ?? ?????????????? ??? ??????????? ???????????? ????
(Riess et al., 2004). ??? ?????????? ??????????,
????????? ? ???????????? ???????????????
????? ???????????? ???????, ??????????????
????????????? ??????????? ?????????????
????????????? ?? ????? ????? 405 ??. ??????? ???? ?????????? ?? ???????? ????????? ?????????????? I465/I405. ?????? ?????
????? ????? 405 ?? ? ???????? ???????????
??????? ???????????????? ??????? ?? ?????? (Brandes, Bers, 1996) ??? ??????????? ???
????? ?????????? ?????????? ?????????? ??
????? ????????. ? ?????? ???????? ???????
???????? ????????????? ???? ??? ?????? ??? ?????? ????????????, ?????? ???????????? ????????? ?? ???? ???????? ???????????.
???????? ???????????? ???? ??? ????????? ??????????????? ??? ?????? ??????????? ? ??????? ??????????? ????????????? ?? ???? ?????? ???? ? 405 ? 460 ?? ???
??????????? ?????????? ? ?????? ????? 350
?? (An et al., 2005). ??? ????????? ???????
???? ????????????? ?? ????? ????? 5 ??? ?
??????????? ?????????? ?????? ? ???????
????? ??????? ?????? (2 ?). ?????????, ??
???????? ????????????? ??????? ?????????? ??????-?????????? ??? (??????????? 480
??, ????????????? 540 ??), ??????????? ?
?????? ??????, ????? ???????????? ???????.
???, ? ?????? ?????? ?????????? ???????
(? ??????? 5 ???) ???? ?????????????, ?????
???????? ????????? ???? ? ??????? 30 ???,
???????????? ??????? ? 1 ? ? ?????????
?????? ?? ????????? ????????????. ??????
????? ?? ???????? ???????? ????????????
??????????.
??????????? ???????? ?????? ????????
????????? ??????? ???? ? ??? ??? ??????
???????? ??? ????????????? ????????????
????????????????? ????????? ? ??????????
????????? ? ?????? (Aldakkak et al., 2009). ???????? ??????? ? ???????? ?????????????
???? ??? ????????????? ???????????? ?
??? ???????. ??? ???? ??????? ???????????
??????? ? ?????? ????????????? ?? ?????
?????? ??? ????????????? ????????????
? ??? ???. ???????? ????????????? ???
?????? ? ????????? ? ?????? (An et al., 2005),
?????? ?? ????????? ?????? ?? ?????????????? ??????????? ??????? ????? ?????????????? ? ????????? ????????????? ???
? ?????????????? ???????????? ? ??? ???, ?
# 145 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ???????? ????????????? ?????????????? ????????????? ??? ?????? ?????????????
??????? ?????????????? ?????? ?? ????? ??????????.
? ?????? (Brachmanski et al., 2004) ??????????? ???????? ????????????? ????
?? ????? ?????? ??? ????????????? ???????????? ????????? Custodiol ??? ?????????
???????????. ??????? ???????? ??????????
????? «???????», ?????????? ?????????? ??????? ????? ????????????? ????, ???????
?????????? 88 c, ? ?? ????? ??? ??? ????????????? ???????????? ? 43 ?. ?????? ????????? ?? ??, ??? ??? ????????????? ???????????? ????????????? ????????????? ????
????????? ????????? ? ??????? 4-5 ???, ??
?? ???????? ??????? ???????? ?? ????? ?????????? ????????? ??????, ???? ?????????
??? ????? ??????? ??????????????? ? ????????? ?????????????? ?????????? ????? ?? ????????? ??????? ?????? ?????? ?? 30 ???.
??????????????? ?????? ???? ???? ???
?????? ???????? ??????? ???? ? ?????????????, ? ????? ??? ??????????? ??????? ???
??????? ???????? ?????? ? ??????????????
????????????, ??????? ????? ? ???? ????????????? ???????, ? ????? ????????? ?????? ?? ???????? ????????????? ???????? ?
??????? ?????? ?????? ??? ?????????????
???????????? ???????? ??? ? ?????????? ????? ????????????????? ??????.
???? ???????????? ? ??????? ??????????????? ?????? ?????????? ??????? ???
?????? ????????????? ?????????? ??????????? ???????? ? ??????, ??????????????
???????????? ??????????? ??????, ?????????? ???????? ??????????????? ??????? ?????????????? ???????? ? ???????? ???????????
? ?????????? ???? ????????? (????????????),
????????? ?????? ????????????? ???????,
??????????????? ??????? ? ????????????????? ????????? ? ?????????????? ??? ??????
??????? ??? ?????? ??????????? ??????????
?????? ?????????? ??????? ? ?????????????-
?????? ???????? ??? ????? ?????????????????? ?????????????????.
????????? ? ??????
???????????? ????????? ?? 59 ?????
??????????? ??????-?????? ?????? 180-200
?, ???????????? ? ??????????? ????????
???????. ??? ???????????? ????????? ? ??????????? ?????? ????????? ????????? ?
????????????? ?????????.
??????? ?????? ???????????? ????????????? ????????? ????????????? ????????? ????? ????????? ??? ????????????
?????? ????????? 0,4 ?/???, ???????? ??????
????????? ????????? (2,5 ??). ?????? ? ?????? ????????? ???????????? ??????????? ???????????, ????? ?????????????? ???????
???????? ??????????? ?????????? ?????
(?????, ???????? ???????, ??????? ????? ????)
??? ????????? ??????? ?? ???????. ?????????
?????? ???????? ???????? ?????????? ?????? ?? ?????. ??? ???????????? ? ????????? ?????? ? ????? ????????? ??????? ????????????????? ?????????????? ??????? (4 ??)
? ??????????? ? ?????? 1:4. ?????? ?????????????????? ???????? ?????: NaCl 0,9 % ?
200 ??; ???????? 50 ??; KCl 4 % ? 10 ??;
NaHCO3 ? 10 ??, ??????? MgSO4 25 % ? 2,5 ??,
?????????? ???????????? 1 % ? 1 ??, ??????????? 1,5 ??. ????? ???????? ???????? ? ??????? ?????? ????????? ?????????? ? ???????
60-100 ?. ? ????????????? ? ??????????????
??????????? ????????????? ????????? ????????????????? ??????? ? ?????????? ??????, ??????????? ? ???? ??????????????
????- ? ??????????, ??? ?????????????????
???????, ?????????? ??, ??????????? ?
???????? ????????? ??????????? ???????????????????.
????????????????? ????? ????????
?????????????? in situ ? ??????? ?????????????????? ????????? ??????????????????
# 146 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ???????? ????????????? ?????????????? ????????????? ??? ?????? ?????????????
? ?????????????? ????????? ????????? (????? ? ??., 2005; Popov et al., 2006). ??? ??????????? ????????????? ???????????? ????????????????? ?????????????? ???????
????? ? ????????? ??????? ???????? ??????????? (Provorov, Salmin, 1993; Provorov et
al., 2005). ??? ?????? ???????? ????????????? ????????? ?????????? ?????: ????? ????????? ??????? ?? ????? ?????????? ????
???????????? ?? ??????? ???????? ????????
2 ??2, ?????? ?????????? ? ???? ? ????????? ??????????? ? ??????? ????? ??????? ??????. ??????? ????????????? ?????????? ?
????????? 380-550 ??. ??????????? ????????
???????????? 20 ?????. ?? ??????? ????????
??????? ????????? ???? ???????? 40 ???????? ????????????? (?? 2 ??????? ? ??????).
??????? ????? ???????? ??? ????????????? ? ??????????????? ???????????? ????
???????? ? ??????????? ?????? ????????, ?
????? ? ????????????????? ?????? ????????
????? 250 ? 500 ? ????? ?????? ??????. ????????????? ?????????? ????? ???????? ???????????? ? ?????????????? ??????????????
Silent Crusher M (Heidolph, ????????). ????????? ???(?)?-?????????? ??????????
???????? ? ?????????? ???????? ? ?????????????? ???????? ????????????? (????????????????? SOLAR SM2203, ????????) ???
??????????? ????????????? ?? ????? ?????
340 ?? ? ???????? ? ???????? ??????????
?? ?? ????? ?????. ???????????? ???????
? ?????????? ????? ???????? ???????? ???????, ?????????? ?? ?????????????? ???????
? ??????????? ??????????????????? ? ???+
?? ?????????????????????? ????????????
?????????? ??????????????? ????. ? ??????
??????????? ????? ??????? ????????? ?????, 1 ? ?????????? ????? (?? 9,5), 0,56 ?
????????? ??????? (?? 8,6), 0,1 ? ???+ (??
6,75), ????? 5 ??? (????? ?????????????) ???
+37 °? ????????? 2,7 ??????/??? ????????-
???????????, ???????? ?????????? ?????????? ??? 340 ??, ?? ?????????????? ???????
?????????? ???????????? ??????? (?????/?).
???????????? ?????????? ??????????? ???
???????? ??????????? ????????? ??????? ?
?????????? ????? ??????????? ?? ???????????? ?????? ? ??????????????? ????????
(?????????, ????????, 2008). ??? ??????
??????????????? ????????? ? ???????? ????????? ??????????? ?????? ?????? ??????????????? ??????????? ? ????????????
????? (?900).
?????????????? ?????? ??????????
??????????? ??????? ?????? ????????????
?????????? ? ???????? ???????? ? ???????????? ?????????????. ???????? ? ???????????? ????????????? ? ???????? ??????? ????????? ?????????? ? ?????????????? ???????
?????????(P) ?????? MS Excel. ??????????? ?????????? ???????? ????????? ????????? ?? ???? R 2>0,95. ??? ??????????
???????????? ? ???? M±m, ??? ? ? ???????
????????, m ? ??????????? ?????? ????????.
????????? ??????? ???????? ????????????
??????? ? ???????? ??????? ???????????? ?
??????? ?-?????.
??????????
?? ??????????, ??? ? ???????? ??????
???????? ??????????? ??????????? ????????? ? ??????????????, ????????????????? ??????????????? ???????????? ??????.
???, ? ?????? ??????? ????? ?????? ??????
????????????? ????? ??????? ????? ? ?????????? ?????????????, ????? ???????????
??????? ??????????????, ??????????? ??????? ???????????? ? ?????????? ?????? ?????????? ?????????? ?? 1:2 (? ?????) ?? 1:1. ?
250-? ??????? ?????? ???????? ???????????
????????????? ??????? ???????????? ??????, ???????????? ?????????? ??????????.
? 500 ??????? ?????? ???????? ? ????? ??-
# 147 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ???????? ????????????? ?????????????? ????????????? ??? ?????? ?????????????
?
?
?
???. 1. ??????????????? ??????? ???????? ???? ??? ??????. ? ?????? ??????: ????????????? ?????
??????? ????? ? ?????????? ?????????????. ? 250 ?: ?????? ? ??????????? ??????????????? ???????
??????? ?????????????? (???????????). ? 500 ?: ????????????? ??????? «?????????????» ??????????
? ?????????
???????????? ????????????? ??????? ?????????? ?????????? ? ??????? (???. 1).
??? ??????????? ????????????? ??
????? ????? 337 ?? ??????????? ??????
????????????? ???????? ???? ???? ???????? ??? ????????????????? ????? ??????
????? ? ?????????? ?? ????? ????? 470 ??,
??? ????????????? ????????? ????????????? ????????? ????? ???(?)?. ?? ???????????? ?????? ???????? ?????????????
???????? ? ????????? ?? 380 ?? 550 ?? ? ????? 10 ??, ? ?????????? ???????????? 30 ?
? ??????? 20 ??? ??????. ?????? ?????? ?????? ??????? ???????????? ???????? ?????
??????? ????????????? ???????? ? ???????
?????? ??????. ??? ???? ????????? ????????? ? ????????????? ????????. ?????????? ??? ??????? ???????????? ?????????
???????????? ? ??????? ???????? ???????
???????? ??????? ?? ????? ????????? ??????????? ?????????????. ??? ??????????
?? ???? ??????? ?????? ?????? ?????????
????????????? ????????????? ?? ?????? ??
???? ???? ????????????? ????????? ?????-
# 148 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ???????? ????????????? ?????????????? ????????????? ??? ?????? ?????????????
0,9450
0,8306
0,7163
0,6019
0,4875
0,3731
0,2588
0,1444
0,03000
1000
?????(?)
800
600
400
200
380
400
420
440
460
480
O(??)
500
520
540
???. 2. ????? ????????????? ???????? ???????? ????????????? ???????? ???? ?? ????? ?????? ???
????????????? ???????????? (n=10)
?????? ???, ????? ???????????? ????????
????????? 1, ? ??????????? 0. ???????????
? ?????????? ????????? ????? ?????????
???????? ????????????? ?? ??????? ????????? ?? ??? 2.
?? ?????????? ????? (???. 2) ????????
?????? ??????????? ????????????? ????????? ?? ????? ????? ? 20 ?????? ??????
(???. 3). ?? ????????? ????????? ?????????? ? ????????? ?? ?????????? ?????? ???
?????? ???????? ? ??????? ?????????????
???????? ????? ??????? ????????? ???????? ????? ???? ? 390, 440, 490, 540 ??. ???
???? ????????????? ????????????? ?? ?????? ???? 450 ? 540 ????????????? ????????
?????????? ???????? ??????? ????????????? ? ????? ???? ???????????? ? ????????
??????????? ???????. ????????????? ??
????? ????? 490 ??, ??????? ????????????? ???????? ????????? ?? ????? ??????,
???????????????? ???? ??? ????????? ???????????? ????, ? ????????????? ?? ?????
????? 390 ??, ??????? ????????????? ????????, ? ??? ????????? ???????????? ????-
?????? ?????????????. ??????????????, ???
????? ?????????? ?? ????????????? ????????????? ??????????? ??????? ??????, ??????????? ????:
? NADH =
I 490 ,
I 440 + I 540
? ??????, ??????????? ????????????????:
? HbO =
I 390
.
I 440 + I 540
???? ????? ????????? ???????????
????????, ??????????? ??? ????????? ???????????? ????, ??? ? ????????????????? ?
?????? ???????????????? (Viability Index):
?VIA =
I 390 .
I 490
?????? ?????? ???????????? ???? ?????? ???????? ????????????? ?? ????????
?????? ???? ??? ????????? ????????????
??????? ?????????????????? ???????? ??????????, ????????? ?? ????????? ?????????????: ?????????? ?????? ? ??????????
# 149 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ???????? ????????????? ?????????????? ????????????? ??? ?????? ?????????????
0,35
0,3
V(???.??.)
0,25
0,2
0,15
0,1
0,05
0
370
390
410
430
450
470
490
510
530
550
O(??)
???. 3. ?????? ??????????? ?????????? ????????????? ???????? ????????????? ?????????????
???????? ???? ? ??????? 20 ??? ?????? ??? ????????????? ????????????
????????. ??????????? ???? ?????? ?? ??????????? ???????????????? ????????????????? ?????????.
??? ?????? ???????????? ??????????
?? ???????????? ????????????? ?????????? ????? ?????? (20 ???), ??? ????????? ?????? ?????????? ????????. ???????? ????????????? ?????? ???????? ????????????
????????????????? ? ??????????? ??????
???????????? ?? ???. 4. ?????? ????????
????? ???? ??????? ????????????????
?????? ???? ???????????????? ???????.
?????? ????????? ??????????? ? ???????
????????????? ?????????? R 2>0,99 ??????????? ??? ???? ???????????? ??????. ???
?????, ??????? ????????? ????? «?????????» ?????????? (? 2). ???????? ??????????
??????? «?????????» ?????????? ??? ???????? ???????????? ????, ????????????? ? ??????? ???????????????? (VIA) ???
????????? ????? ???????????? ????????? ?
????. 1
???????? ?????? Na2O4S2, ?????? ??????? ???????????????, ???????? ??????????????? ????????????????, ????????? ?
??????????? ??????????. ??????????????,
????? ??????? ???????? ??????????? ????????? ? ????? ???????? ??? ??? ????????????? ? ????????????????? ????????, ???
?????????? ? ?????????? ???????????????
?????????? ??????? (????.1). ? ????????????????? ???????? ????????? ?????? ??
????????? ?????????? ?????????? ???????
??? ????????? ? ?????? ??????????????????
???????? ?? (????.1).
??? ????? ?? ????. 1, ??? ???????????
?????? ?????????? ???????? ??????????
??????? 460-500 ?. ??? ????????? ?? ???????????????? ???????? ????????? ?????? ?????????? ??????????? (???) ? ????? (???. 5).
?????? ?? ??????? ????????? ??? ?????????? ????????? ???????????? ?? ?????? ?????? ??????. ????????, ??? ?? ???????????
????????? ?????????? ?????????? ????????
?????? ???, ??? ??????????????? ?? ????????? ?????????? ????????? ???? ?? ??????????? ??????????????? ??????? ? ??????
?????? ?????? ??????. ?? ???????? ???????????? ??????? ????? ?????? ???????, ???
?????????? ??????? t, ?????? 516 c, ?????? ??
# 150 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
y = A1*exp(-x/t1) + A2*exp(-x/t2) + y0
R^2
CHbO(???.??.)
1,0
y0
A1
t1
A2
t2
0,99902
Value
Error
0,02004
0,00109
0,37759
0,0064
38,00786
1,33188
0,59541
0,00416
485,85172
5,52212
0,5
0,0
0
200
400
600
800
1000
1200
????? (c)
???. 4. ???????? ???? ????????????????? ? ???????? ??? ?????? ??? ??????????? ?????? (??????
???????????? ?-???, n=10). ????? ? ????????????????? ????????, ???????? ????? ? ?????????????
?????? ???? ?????????. ????????? ????????????? ??????? ? ??????? ?????????
??????? 1. ?????????? ??????? «?????????» ?????????? ??? ????????? ??????????? ?????? ???
??????????? ??????? ?????????????????? ????????
??????????? ???????
??????????? ???????
+???????? ??????
??????????? ???????+??
NADH
492±5 (?)
425±10 (?)
853±20 (?)
Hb
486±6 (?)
430±10 (?)
650±5 (?)
VIA
465±5 (?)
406±10 (c)
654±6 (c)
0,6
R^2
0,92549
Value
Error
Slope
-0,00193 3,12808E-4
ln(C/C0)(???.??.)
0,4
0,2
0,0
-0,2
-0,4
-0,6
-0,8
-1,0
-1,2
0
250
500
?????(?)
???. 5. ???????? ????????????? ???????????? ??? ? ???????? ???? (n=10 ??? ??????? ????????
???????) ?? ????? ?????? ?? ???? ????????????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ???????? ????????????? ?????????????? ????????????? ??? ?????? ?????????????
2,0
R^2
Slope
0,95899
Value
Error
0,00199 2,36388E-4
ln(I/I0)(???.??.)
1,5
1,0
0,5
0,0
-0,5
-1,0
0
250
500
?????(?)
???. 6. ?????????? ?????-???????? ? ???????? ???? (n=10 ??? ??????? ???????? ???????) ?? ?????
?????? ???????? ?? ???? ????????????
140
???????(??/?)
120
*
100
80
60
40
20
0
0
250
?????(?)
500
???. 7. ???????????? ??????? ? ????? ???????? ???? (n=10 ??? ??????? ???????? ???????) ?? ?????
?????? ???????? ?? ???? ????????????. * ? ????????????? ??????? ????? ???????? ??? P<0,05
???????? ? ?????????? ??????? ???? ? Hb,
??????????? ? ????. 1.
????????
??????????
???(?)
?-???????? ??????????????? ? ???????????
????? ?? ??????????? ????????? (???. 6).
?????? ???????? ?????????? ??????? t=502 c
????? ?????? ? ?????????? ??? ????, Hb,
???. ???? ?????????? ????-???????? ???
?????? ? ?????? ????????? ???????, ?????????? ? ???????? ????????? ?????????
?????????. ????????????????? ????????? ?????????? ???????? ? ?????????? ?????????
??????????? ????????? ???????, ? ??????
??? (???. 5), ???????????? ?????????????
# 152 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ???????? ????????????? ?????????????? ????????????? ??? ?????? ?????????????
?? ??????????? ?????????? ??????????????
??????? ? ?????? ?????? ?????? ???????????? ??????????????? ???????.
????????? ???????????? ??????? ? ????? ???????? ?? ????? ?????? ????????????
?? ???. 7. ???????? ?????????? ???????? ????????? ?????????? ??????? ?????????: ????????? ????????? ??????????? ????????? ?
?????????????? ???????, ??????????? ??????????? ??????? ????????? ??? ??????????
????, ??????? ? ????????? ???+ ? 500-?
??? ??????.
??????????
???????????? ????? ???????????? ??
???????? ???????????????? ???(?)? ???
?????? ????????? ???????? ? ?????????????
?????????????? ????????????? ?? ??????????? ??????????? ???(?)? ? ????? ??? ??????? ??? ???????? ?????????? ??????????????
??? ???????????? ??????????? (Fomin et al.,
2003; Brachmanski et al., 2004). ? ?????????
????????????? ???? ??????????? ???????
??????? ??????? ????????????? ????????
(?????????, ????????? ???????????) ??
?????????? ????????? ???? ????? (Arai et al.,
1999; Leisey et al., 1994), ?????? ?????? ? ??????? ???????? ????????????? ?? ????????
????????????? ??????????? ? ??????????
??????????? ???????? ??? ?????? ? ?????????????? ???????????? ? ?????????? ??????????? ???????????.
?? ??????????, ??? ??????????
???????-??????????????? ??????? ????????
? ????????????????? ??????, ???????????
???????????? ? ???????? ??????????? ???????? (??????????????????? ?????????????
? ????????????? ? ?????????? ??????), ????????? ????????? ???????????????? ? ??????? ??????????? ??????, ???????????????
? ???????????????? ? ??????????????????
???????????. ?? ??????? ????????, ??? ??-
??????? ?????????????????? ??????????
?????????????????? ???????? ????? ???????? ? ??? ?????? ????????? ?????? ????????? ?????????? ??????? ?? ???????? ???????
????????????? ????????, ????? ??? ??????????? ? ??????? ?????????????????? ????????
????????? ???????? ????????? ?????????????????? ??????, ?????????????? ?? ?????????? ?????????? ???????. ????????????
???? ? ???????????? ?????? ?? ? ?????????
???????????????? ????? ???????? ??????
????????????? ???????????? ?????? ? ???????????? ???????? ??, ??????????? ?? ????
????????? ?38 ????, Akt, ??????? ??????
????? (Fujimoto et al., 2004), ????????? ?????????? ????? (Varadi et al., 2007) ? ??????????????.
??? ?????? ???????? ????????????
????????????? ? ?????? ???? ??????????
? ???????? ???????????????? ???? ? ??????????? ????????? ? ???????????? ? ?????
?? ???????????? ???????? ????????? ???
????????? ????????? ?????????? ? ??????????? ???? ??????????? (Wittenberg, Wittenberg, 2003). ??? ?????????? ???????????
????????? ????????? ???????????? ??????
? ???????????? ??????? ??????????? ???????? ? ??????? ????????????? ? ???????
???????????? ????? (Nighswander-Rempel et
al., 2002).
??????????? ????????????????? ?????? (???????????? ? ????????? ??????) ????????? ??? ??????? ???????? ?????????????
???(?)? ??? ???????????? ??????????
?????????????????? ????????? ? ????????,
? ????? ??? ????????? ???????? ????????? ?
?????? ??-?? ??????????? ????????? ?????????? ???????? ????????? ? ????????????????
???????????? ??? ?????????? ???????????
?????????? ???????? ? ????????? ??????????? ?????????. ???? ?????? ???????????,
??? ????????????? ???(?)? ???????? ???-
# 153 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ???????? ????????????? ?????????????? ????????????? ??? ?????? ?????????????
??????????? ??????????? ?????????? ????????? ? ???????? (Stoner et al., 2004). ?????????????????? ???? ????????? ???? ???(?)
? ?????? ?????????????? ???????? ??????????? ??????????? ? ?????? ??????????,
??????????? ??????? ? ????????? ???(?)
?-?????????? ??????????. ? ?????? ???????????? ?????? ? ???, ??? ??????? ???? ?
???+ ? ???????? ?????????????? ???????????? ????????????? ??????? ?? ??????????????? ???? (Zima et al., 2004), ???????????
????????????? ??????????? ???????????
???????? ???? ???????????? ?????????? ?
??????????? ?????????????? ??????????????
?????????? ?????? ????? ?????????? ??????
??????.
?????????????? ? ??????? ?????? ?????????? ??????? ????????? ???????? ????????????? ???(?)? ???????? ?????????????
?????????????? ??????? ?????????? ??????????????? ??????????? (??????????????
??????????? ?????????, ????????????????
??????????), ??????????????? ???????????
(??????????? ? ????? ??????????) ? ???????? ??????????? ????????-???????????? ?
????????-???????????? ??????? ????????????? ? ?????.
?????????????, ??? ????????? ????????? ???+ ? ???? ? ??????? ???????? ?????
???????????????? ???????? ??? ?????????
???????????????? ???????????? ?????????? 2-?? ???? ?, ??????????????, ???????? ?
????????? ????????????? ? ??????? ??2+ ??
??????????????? ???? (???????????????????? ??????????) (Cherednichenko et al., 2003;
Zima et al., 2004). ??? ????????? ??? ???????
??????????????? ???????????????? ????????? ??????????? ???? ? ???????? ??????????? ?????????????? ?? 250-500-? ????????
??????. ????? ???????, ????????????? ?????? ????????????? ???????, ??????????????? ??????? ? ????????????????? ????????? ? ?????????????? ??? ?????? ??????
??????????????? ? ??????? ???????????????
?????? ?????????? ??????? ??? ???????????
????????? ? ??????????? ????????? ? ????? ???????? ??? ??????. ? ????? ????????
???????????? 500-? ??????? ? ??????? ??????????? ?????? (? ???????? ?????? ????????
????????????????? ?????????) ???????? ?????? ??????????? ??????????? ???????????????????????? ?????????.
??????????
????? ?????????? ??????? ?????????
?????????????? ?????????????? ?????????
? ????????, ????????????? ??????????
???? ??????????? ???????????, ?????????? ????????-???????????? ? ???????????????????? ??????? ?????????????,
?????????? ???(?)?-????????????? ????????? ? ????????? ??????????? ? ??????
??????????????.
??????????? ??????????????? ???????
? ?????????????? ?????? ? ??????????????
???????-??????????????? ??????? ? ???????? ??????????????????? ???????? ????????
?????????? ?????? ???????? ????????????
??????????? ?????? ? ??????? ????????????? ???????? ?????? ????????.
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?????? ??????????
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# 154 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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?.?. ??????, ?.?. ???????? ????????????? ?????????????? ????????????? ??? ?????? ?????????????
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727-741.
Application of Fluorescence Spectroscopy
for Assesment of Myocardial Ischemic Injury
Vladimir V. Salmina*, Alla B. Salminab,
Alexander A. Fursovb, Olga V. Frolovab,
Dmitry I. Laletinb, Maxim A. Fursovb,
Gennady V. Yudinb, Nataliya A. Malinovskayab,
Larissa D. Zykovab and Alexander S. Provorova
a
Siberian Federal University,
79 Svobodny, Krasnoyarsk, 660041 Russia
b
Krasnoyarsk State Medical University
named after Prof. V.F.Voino-Yasenetsky,
1 P. Zheleznyaka str., Krasnoyarsk, 660022 Russia
Data of experimental assessment of a possibility to use laser-fluorescent analysis of myocardium
for detecting alterations caused by ischemia under the conditions of cardiac surgery are presented.
It is confirmed that the method of optical biopsy with Nitrogen laser (337 nm) allows registering
metabolic changes in myocardium associated with alterations in pyridine nucleotides concentrations,
dysfunction of hemoproteids, changes in the activity of NAD(P)H-utilizing enzymes, and development
of contraction and death of cardiomyocytes.
Keywords: optical biopsy, myocardium, ischemia, nitrogen laser.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2011 4) 158-171
~~~
??? 579.6
Microbial Synthesis and Characterization
of Poly (3-Hydroxybutyrate-co4-Hydroxybutyrate) Copolymers
Natalia O. Zhilaa, b*, Tatiana G. Volovaa, b,
Elena D. Nikolaevaa, b and Daria A. Syrvachevab
a
Institute of Biophysics Siberian Branch
of Russian Academy of Science,
50 Akademgorodok, Krasnoyarsk, 660036 Russia
b
Siberian Federal University,
79 Svobodny av., Krasnoyarsk, 660041 Russia 1
Received 3.06.2011, received in revised form 10.06.2011, accepted 17.06.2011
The research is focused on the prospective representative of the family of biodegradable polymers ?
polyhydroxyalkanoates (PHAs), the copolymer poly(3-hydroxybutyrate-co-4-hydroxybutyrate)
(P(3HB-co-4HB). Difficulties with producing this type of PHA are associated with the fact that the
precursor for the 4-hydroxybutyrate (4HB) monomers synthesis, the ?-butyrolactone, is toxic for
the bacteria producers and reduces the total biomass production and the polymer yield when it is
added to the medium. Having used the natural strains of hydrogen-oxidizing bacteria Ralstonia
eutropha B5786 and Cupriavidus eutrophus ?10646, the latter possessing the increased sustainability
to the ?-butyrolactone influence, the conditions for efficient synthesis of P(3HB-co-4HB) have been
determined. A set of highly purified samples of P(3HB-co-4HB) with different content of 4HB (from 8.7
to 24.3 mol %) was produced. It has been determined that the incorporation of 4HB in the copolymer
of a higher degree than 3-hydroxyvalerate and 3-hydroxyhexanoate leads to the decrease of the
copolymer?s crystallinity; samples with the crystallinity of 12 % and 25 % have been obtained. It
has been demonstrated that the average molecular weight of the P(3HB-co-4HB) samples and the
polydispersity do not depend on the ratio of monomers and vary within the wide range from 540
to 1100 kDa and from 1.91 to 2.76 correspondingly. It has been determined that the 4HB content in
the studied range (8.7-24.3 mol %) does not effect the melting temperature of the copolymers (168.9172.5 єC).
Keywords:
hydrogen-oxidizing
bacteria;
?-butyrolactone; physicochemical properties.
*
1
poly(3-hydroxybutyrate-co-4-hydroxybutyrate);
Corresponding author E-mail address: nzhila@mail.ru
© Siberian Federal University. All rights reserved
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Natalia O. Zhila, Tatiana G. Volova? Microbial Synthesis and Characterization?
Introduction
Synthesis of the storage polymers ?
hydroxy derivatives of alkanoic acids
polyhydroxyalkanoates
(PHAs)
?
by
microorganisms has become an extensively
studied subject. PHAs are represented by various
polyesters, consisting of homogenous monomers
with different length of the carbon chain and
by copolymers; there are high-crystallinity
thermoplastic PHAs and thermolabile rubber-like
elastomers (Philip et al., 2007).
Synthesis of PHAs with tailored properties
is a difficult technological task, and production
of a PHA of definite composition should be based
on fundamental knowledge of how a certain PHA
is synthesized and how the chemical structure
of the polymer influences its physicochemical
properties. There are several ways to produce
PHAs consisting of monomers with different
length of the carbon chain: using genetically
modified PHA producing strains, with genes
controlling the key enzymes of PHA synthesis
(?-ketothiolase, acetoacetyl-CoA reductase,
PHA-synthase) taken from microorganisms of
different taxa and introduced into them, and
based on the knowledge of the mechanisms of
PHA synthesis by natural producers, which are
influenced by culture conditions.
A very promising but insufficiently studied
PHA is the poly(3-hydroxybutyrate-co-4hydroxybutyrate) (P(3HB-co-4HB)) copolymer.
PHAs of this type are biodegraded in vivo and
in the environment at high rates and exhibit
better elongation at break and higher tensile
strength than most of the well-known polymers
of this class (Martin, Williams, 2003; Freier,
2006; Chanprateep, 2010). The ability of
such microorganisms as Cupriavidus necator
(previously known as Wautersia eutropha,
Ralstonia eutropha, Alcaligenes eutrophus)
(Nakamura et al., 1992), Alcaligenes latus
(Hiramitsu et al., 1993), Comamonas testosteronii
(Renner et al., 1996), Comamonas acidovorans
(Saito, Doi, 1994), Hydrogenophaga pseudoflava
(Choi et al., 1999), Chromobacterim sp (Kimura
et al., 1999), Rhodococcus rubber (Haywood et
al., 1991), Delftia acidovorans (Hsieh et al., 2009)
to synthesize PHAs of this type when grown on
the media containing 4-hydroxybutyric acid,
?-butyrolactone or 1,4-butanediol as carbon
substrate was reported in a number of studies
in the 1990s. The inhibiting effect of these
substrates, however, reduces both total biomass
production and copolymer yields.
In recent years, this representative of PHAs
has received much attention. Recombinant
Escherichia coli, Aeromonas hydrophila,
Pseudomonas putida, Ralstonia eutropha H16
(Zhang et al., 2009; Li et al., 2010) and wild-type
strains Cupriavidus sp. USMAA1020 (Amirul
et al., 2008), R. eutropha A-04 (Chanprateep
et al., 2008), Cupriavidus sp. USMAA2-4
(Ramachandran et al., 2011), R.eutropha KCTC
2662 (Park et al., 2011) have been described as
novel producers of P(3HB-co-4HB).
The objective of this study was to research
the conditions of the P(3HB-co-4HB) synthesis
by natural strains of hydrogen-oxidizing bacteria
and determined the effect of the composition on
the properties of the copolymer and the resulted
products.
Materials and Methods
Two strains of hydrogen-oxidizing
bacteria ? Ralstonia eutropha ?5786 (Stasishina,
Volova, 1992) and Cupriavidus eutrophus
?10646 (registered in the Russian Collection
of Industrial Microorganisms (VKPM)) were
batch cultured in 1.0-L glass flasks, which were
50 % filled with the culture, under strict aseptic
conditions, on a ?New Brunswick? temperaturecontrolled shaker (USA); the cultures were grown
on Schlegel mineral salt medium. Autotrophic
cultures were grown on the (??2, ? 2, ?2) gas
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Natalia O. Zhila, Tatiana G. Volova? Microbial Synthesis and Characterization?
mixture as carbon and energy source. Air was
evacuated from the flasks with a compressor,
and the gas mixture was supplied from a
gasholder, via airtight hoses. The proportions of
CO2, O2 and H 2 in the gas mixture were 1:2:7
(v/v). Heterotrophic cultures were grown on
fructose or butyric acid. Cells were cultured
in two phases ? in nitrogen-limited medium in
the fi rst and nitrogen-free one in the second, at
pH 7.0 and temperature 30 °C, in accordance
with our previously developed procedure
(Volova et al., 1992). Synthesis of monomers
4-hydroxybutyrate (4HB) was induced by the
addition of ?-butyrolactone (Fluka). The culture
was added with ?-butyrolactone with varied
frequency and in varied amounts. The duration
of the cultivation depended on carbon nutrition
conditions and varied from 72 h to 96 h.
Biomass yield was determined from dry
mass and optical parameters of the culture.
Polymer content and PHA composition was
determined with a gas chromatograph-mass
spectrometer (GC/MS, model GCD Plus, Hewlett
Packard, USA) Percentages of the monomers
in (P(3HB-co-4HB) were quantified based on
mass spectra, and more accurate quantification
was attained by taking 1H-NMR spectra of
copolymer solutions in CDCl3 using an Avance
III 600 NMR spectrometer (Bruker, Germany).
Current concentrations of salts of alkanoic acids
and ?-butyrolactone in the culture were analyzed
using gas chromatography. Fructose concentration
in the medium was measured using the resorcinol
method (Ermakov et al., 1972).
X-ray structure analysis and determination
of crystallinity of PHA samples were performed
using a D8 Advance X-ray spectrometer (Bruker,
Germany) (graphite monochromator on a reflected
beam) in a scan-step mode, with a 0.04° step and
exposure time 2 sec, to measure intensity at
point. The instrument was operating at 40 kV Ч
40 ??. Thermal properties of biopolymers were
examined using differential scanning calorimetry
with a NETZSCH analyzer (Germany). Each
sample of mass ~2 mg was shaped as a disc of
diameter 5 mm, inserted into an aluminum capsule
with a small hole, and heated at 5 K/min. Melting
point, devitrification temperature, and thermal
degradation temperature were determined from
DSC curves at the given heating rate.
Molecular weight was determined by gel
permeation chromatography at a temperature of
40 °C, using a Waters chromatographic system
(a Waters 151 isocratic pump, a Reodyne 7725i
injector, and a Waters 2414 refractometric
detector) and columns Styragel HR4E and HR5.
Chloroform was used as eluent, with the flow rate
0.8 ml/min. The system was calibrated using low
polydispersity polysterene standards supplied by
Sigma (USA).
The number-average molecular weight of
the PHA was determined as
Mn=? (Ni · Mi / N),
where Ni is the number of molecules of mass I; N
is the total number of molecules; Mi is the mass of
molecules of length I.
The weight average molecular weight of the
PHA was determined as
Mw = ? (wi · Mi),
where wi is the portion of the mass (wi = Ni Mi / ?
(Ni · Mi).
Polydispersity, which provides an estimate
of the proportions of fragments with different
polymerization abilities in the polymer, was
calculated from the formula
PD = Mw / Mn.
Experiments were done in three replicates.
The results were analyzed statistically by
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Natalia O. Zhila, Tatiana G. Volova? Microbial Synthesis and Characterization?
conventional methods, using the
software package of Microsoft Excel.
standard
Results
In the autotrophic culture of R. eutropha
B5786 grown in flasks on a shaker under nitrogen
deficiency, using single-carbon substrate (CO2),
the biomass yield and the polymer content after
72-h fermentation amounted to 6.1 g/L and 65.4 %
of dry cell weight (DCW), respectively (Table 1).
The polymer synthesized by R .eutropha B5786
on single-carbon substrate contained 99.2 mol %
of 3HB and 0.8 mol % of 3-hydroxyvalerate
(3HV). Previously we reported that cultivation
of bacteria in flasks for longer periods and the
use of advanced fermentation equipment can
increase polymer yield to 85-90 % of DCW
(Volova, Kalacheva, 2005). The main purpose
of this study, however, was to find out whether
this strain is able to synthesize a PHA containing
4-hydroxybutyrate (4HB) and what growth
conditions are necessary for this. Biomass yield
was considered as a secondary issue, so the
duration of experiments with autotrophic culture
in flasks did not exceed 70 h.
Taking into account the data on the toxicity
of ?-butyrolactone (Nakamura et al., 1992), we
studied the effect of this supplementary substrate
on R. eutropha B5786 growth and PHA synthesis
using ?-butyrolactone concentration from 0.5
g/L to 4.0 g/L. Results of PHA biosynthesis and
incorporation of 4HB into the copolymer, using
mixed carbon source (CO2 + ?-butyrolactone)
are shown in Table 1: ?-butyrolactone inhibited
polymer synthesis and accumulation. Higher
concentrations of ?-butyrolactone in the medium
did not increase 4HB proportion in the copolymer;
moreover, they inhibited transformation of
?-butyrolactone to 4HB. While with 0.5 g/L of
?-butyrolactone in the medium the degree of
its transformation into 4HB reached 0.21, at
concentration of 4 g/L it dropped below 0.02.
The amount of ?-butyrolactone added to the
culture did not affect biomass yield, which was
not higher than 5.0 g/L, or 25 %-30 % lower than
in the control culture (with CO2 as sole substrate).
Investigation of the influence of ?-butyrolactone
on polymer synthesis showed a dose-dependent
reduction in the intracellular concentration of
the polymer; the highest polymer yield (39.9 %
of DCW) was attained in the experiment with
a single addition of 0.5 g/L of ?-butyrolactone.
Polymer synthesized under these conditions
contained three monomers: 3HB as a major
component, 4HB, and minor amounts of 3HV.
Monomer 4HB was detected in all samples, with
the highest percentage amounting to 6.3 mol %
(Table 1).
To reduce the toxic effect of ?-butyrolactone
on the culture, it was added in several portions.
By varying the size of the dose of ?-butyrolactone
added to the culture and the duration of the
subsequent cultivation, we managed to increase
the total yield of bacterial cells and the polymer
yield. When the ?-butyrolactone dose (6 g/L)
was divided into two portions, the polymer yield
reached 55.7 % of DCW and the total culture
productivity was not decreased, but the 4HB
fraction did not grow, amounting to not more
than 6 mol %. When the dose of ?-butyrolactone
of 9.0 g/L was divided into three portions (3 g/L
each) and added every 8 h, the polymer yield was
50.3 % of DCW, the total biomass yield 4.0 g/L,
and the 4HB fraction did not rise above 6 mol %
(Table 1).
In the culture of R. eutropha B5786 grown
heterotrophically, with fructose as the main
substrate, the inhibitory effect of ?-butyrolactone
was less pronounced (Table 1). Doses of
?-butyrolactone added in one portion were 2.0,
6.0 and 10.0 g/L. Inhibition of cell growth was
only recorded with the highest concentration
of ?-butyrolactone: biomass yield was as low
as 4.5 g/L. At the two other concentrations,
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Natalia O. Zhila, Tatiana G. Volova? Microbial Synthesis and Characterization?
Table 1. Content and composition of polyhydroxyalkanoates synthesized by bacteria Ralstonia eutropha B5786
in different conditions (experiments were done in three replicates)
Concentration of
?-butyrolactone, g/l
Hours
Biomass yield,
g/l
Polymer content,
% of DCW
Polymer composition, mol %
3HB
3HV
4HB
Autotrophic conditions (CO2)
0
72
6.1±0.2
65.4±4.2
99.2
0.8
-
0.5
72
4.2±0.3
39.9±1.7
93.3
0.4
6.3
1.5
72
4.3±0.2
24.9±1.9
93. 6
0.4
6.0
2.0
72
4.6±0.2
13.9±0.7
95.7
0.5
3.8
4.0
72
4.4±0.4
12.0±0.9
98.7
0.7
0.6
6.0 (3+3)*
72
5.9
55.7
93.4
0.7
5.9
9.0 (3+3+3)**
72
6.0
50.3
93.5
0.6
5.9
2.0
96
98.6
0.5
0.9
6.0
10.0
Heterotrophic conditions (fructose)
7.5±0.4
57.3±3.4
96
7.0±0.3
50.6±4.1
97.9
0.7
1.4
96
4.5±0.3
56.7±2.9
98.0
0.4
1.6
6.0 (3.0+3.0)
96
7.4
78.1
95.6
0.6
3.8
9.0 (3.0+3.0+3.0)**
96
7.2
65.9
95.3
0.6
4.1
95.1
0.5
4.4
*
Heterotrophic conditions (butyric acid)
6.0 (3.0+3.0)
**
*
96
7.1
81.7
? ?-butyrolactone adding at 24 and 32 hours of cultivation
? ?-butyrolactone adding at 24, 32 and 40 hours of cultivation
**
biomass yield was similar to that in the control,
reaching 7.0-7.5 g/L, after cultivation for 96
h. Assimilation of ?-butyrolactone occurred at
a higher rate than in the autotrophic culture,
and in 34 h after ?-butyrolactone was added,
its concentration in the culture did not exceed
0.1-0.2 g/L. In the experiment with the
inhibitory concentration of ?-butyrolactone,
polymer yield amounted to 56.7 % of DCW,
but the 4HB fraction of the polymer was
small ? 1.6 mol %. With 6.0 g/L and 2.0 g/L of
?-butyrolactone added to the culture, polymer
yield amounted to 50.6 % and 57.3 % of DCW,
but the polymer contained less than 2 mol %
of 4HB. In experiments with ?-butyrolactone
added in several portions, polymer yield was
increased to 65.9-70.1 % of DCW, but the 4HB
fraction grew insignificantly ? to 3.8-4.1 mol %
(Table 1).
The use of butyric acid (butyric acid sodium
salt) instead of fructose increased copolymer
yield and improved its composition. As butyric
acid concentrations above 4.0 g/L inhibited cell
growth, the concentration of butyric acid was
1-2 g/l. Cultivation of R. eutropha B5786 on the
medium containing butyric acid and 6 g/L of
?-butyrolactone (added in two portions) resulted
in polymer yield of 81.7 % of DCW. The fraction
of 4HB, however, remained low ? not higher than
4.4 mol % (Table 1).
Thus, by varying the conditions of carbon
nutrition, including the main carbon substrate
and its concentration, and the mode of feeding the
supplementary substrate (?-butyrolactone) into
the culture, we managed to maximize the yields
of the polymer and total biomass of R. eutropha
B5786 however the content of 4HB in copolymer
was low.
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Natalia O. Zhila, Tatiana G. Volova? Microbial Synthesis and Characterization?
Further experiments were done with the strain
C. eutrophus ?10646 which has the increased
sustainability to ?-butyrolactone is compared to
R. eutropha ?5786. The highest concentration
of ?-butyrolactone for this strain is 15-20 g/L.
In the culture of C. eutrophus ?10646 grown
autotrophically and heterotrophically, a higher
content of 4HB in the copolymer was obtained
(up to 15 mol %), which on the background of the
higher indexes of biomass yield and total yield
of the copolymer made 6-7 g/L and 81-85 %,
correspondingly.
The mode of growing C. eutrophus ?10646
in the medium containing the propionic acid as
the main substrate in the concentration 2 g/L
and ?-butyrolactone provided synthesis of three
component polymer containing 8.1 mol % of
4HB and 35.2 mol % of 3HV besides 3HB. Using
butyric acid with ?-butyrolactone as the carbon
source resulted in higher production of polymer
in the culture of C. eutrophus ?10646 (Fig. 1).
Biomass yield and polymer content varied
depending on the ?-butyrolactone concentration.
The yields were 6.2-8.2 g/L and 72-89.7 % of
DCW, correspondingly; whereas the content of
4HB varied in the range of 8.7 ? 24.3 mol %.
Using the strain of C. eutrophus ?10646 with the
higher tolerance to the ?-butyrolactone, effect of
both the copolymer P(3HB-co-4HB) strain and
the single-carbon substrate allowed not only to
increase the content of 4HB in the copolymer, but
also to reduce the time of bacteria cultivation.
Growing of C. eutrophus ?10646 bacteria
in the medium contained the butyric acid (or
fructose) as the main substrate and addition of
?-butyrolactone in various concentration (from
2.0 to 10.0 g/L) allowed to fulfil the process with
the total yields of biomass and copolymer within
29-32 hours of cultivation, i.e., up to 25-32 g/L
and 88-95 % of DCW, with 15-20 mol % of 4HB
in the copolymer. Composition of the produced
and purified samples of P(3HB-co-4HB) were
analyzed by the 1H-NMR method which proved
the presence of 4HB in the copolymer. The 1HNMR spectra of the copolymer samples with
4HB incorporation at 8.7 mol % and 16 mol % are
presented in Fig. 2.
Thus, the substitution of the strain R.
eutropha ?5786 by the strain C. eutrophus
?10646 and the developed mode of proportions
40
100
polymer
biomass
80
30
60
20
40
10
20
0
Dry cell weight (g/l)
4HB content (mol%)
PHA content (% of DCW)
4HB
5.0
10.0
15.0
?-butyrolactone (g/l)
20.0
0
Fig. 1. Effect of different concentrations of ?-butyrolactone on the P(3HB-co-4HB) production of Cupriavidus
eutrophus ?10646 under heterotrophic (substrate ? butyric acid) conditions
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Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
a
?
Fig. 2. 1H-NMR spectrum of the P(3HB-co-4HB) with content of 4HB 8.7 mol % (a) and 16 mol % (b)
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Natalia O. Zhila, Tatiana G. Volova? Microbial Synthesis and Characterization?
of ?-butyrolactone in the culture provided
significantly higher values of the three key
indexes: biomass yield, copolymer concentration
in cells and content of 4HB.
The properties of the set of synthesized
and highly purified samples of P(3HB-co-4HB)
were studied using modern physical methods.
Using the gel permeation chromatography, the
molecular mass of the set of the synthesized
samples of copolymers P(3HB-co-4HB) was
determined (Table 2). No influence of this
parameter on the molecular mass and the
polydispersity of the copolymer was detected
in the studied range of the monomers ratio in
the copolymer (the incorporation of 4HB varied
from 8.7 to 24.3 mol %). The number average
mass in the set of samples varied in the range
from 230 to 480 kDa, the weight average
molecular mass amounted to 540?1110 kDa; the
polydispersity varied from 1.91 to 2.76 with no
clear correlation with the content of 4HB in the
copolymer.
It has been demonstrated that the
incorporation of 4HB influenced significantly
the ratio of crystalline and amorphous phases in
the copolymer (to the higher extent if compared
to 3-hydroxyvalerate and 3-hydroxyhexanoate),
sufficiently reducing its crystallinity (Table 2). For
the first time we managed to obtain the samples
of PHA with the lowered degree of crystallinity
(from 44 % to 12 %).
The melting temperature of the produced
samples of P(3HB-co-4HB) was lower
(168.9-172.5єC) in comparison to poly(3hydroxybutyrate) (180єC). However, no relation
between the 4HB content in the range from 8.7
mol % to 17 mol % and the melting temperature
has been determined. The temperature of thermal
degradation of various copolymer samples varied
in the range from 264єC to 286єC and did not
depend on the content of 4HB.
Discussion
Analysis of the up-to-date literature
witnesses the activity of the research aimed at the
synthesis of copolymer P(3HB-co-4HB). This
type of PHA characterized by high elasticity and
strength, as well as quicker and regulated speed
of biodegradation, attracts the attention of the
researchers who develop the products intended
for the biomedical field. For the present moment
the company Metabolix (USA) has developed
and produced a line of products made out of this
copolymer in the form of suture material, films
and reticular implants. For successful production
of this type of PHA many research teams isolate
new strains-producers, improve the fermentation
technology using various compounds as growth
Table 2 Physical characterization of P(3HB-co-4HB) produced by Cupriavidus eutrophus ?10646
4HB content,
mol %
0
Mn (Ч 103) Da
PD
Crystallinity, %
Degradation
Melting temperature
temperature (Td), єC
(Tm) , єC
180
280
760 ± 15
1.60 ± 0.03
76
8.7
230 ± 2
2.76 ± 0.03
44
171
286
10.7
480 ± 9
2.32 ± 0.06
43
171.9
268
14.9
320 ± 7
2.65 ± 0.04
44
168.9
264
16.0
370 ± 5
2.59 ± 0.04
43
171.3
279
17.0
370 ± 12
2.31 ± 0.06
25
172.5
271
24.3
285 ± 5
1.91 ± 0.01
12
-
274
- not identified
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Natalia O. Zhila, Tatiana G. Volova? Microbial Synthesis and Characterization?
substrates. Thus, Li with co-authors (2010) has
determined that the recombinant strain E. coli
containing the plasmid with the genes incorporated
into the succinate degradation out of Clostridium
kluyveri, and the directional genes of the poly(3hydroxybutyrate) synthesis out of R. eutropha,
can synthesize with higher yields (up to 70 % of
DCW) of copolymer P(3HB-co-4HB). Among
the natural producers various bacteria have
been described. Recently new highly productive
strains have been isolated from soil in Thailand
and bottom sediments of the lake in Malaysia
which have been identified correspondingly
as R. eutropha A-04 (Chanprateep et al., 2008)
and Cupriavidus sp. USMAA1020 (Amirul et
al., 2008), synthesizing copolymers P(3HB-co4HB) using ?-butyrolactone, 1,4-butanediol or
4-hydroxybutyrate acid. If R. eutropha A-04 is
grown on 4-hydroxybutyric acid the maximum
incorporation of 4HB amounts to 52 mol %
with 22 % of polymer of DCW biomass. The
maximum content of the copolymer (59 % of the
dry biomass) was obtained using the mixture
of 4-hydroxybutyric acid and butyric acid (1:1),
nevertheless, the incorporation of 4HB did not
exceed 24 mol % (Chanprateep et al., 2008).
When the strain Cupriavidus sp. USMAA1020
was grown on ?-butyrolactone, 4- hydroxybutyric
acid and 1,4-butanediol, the best substrate for
growth and synthesis of the polymer (4.96 g/L
and 58.7 % of DCW, correspondingly) for this
strain was 1,4-butanediol with 32 mol % of 4HB
(Amirul et al., 2008). Higher values of 4HB in the
copolymer (over 80 mol %) were obtained using
the mixture of 1,4-butanediol and ?-butyrolactone,
nevertheless, the biomass and the polymer content
reduced therefore to 1.9 g/L and 16 % of the dry
weight (Vigneswari et al., 2010). The copolymer
with the same high content of 4HB was produced
during the cultivation of another strain of bacteria
Wautersia eutropha H16 in the medium which
besides 4-hydroxybutyric acid also contained
amino acids L-alanine and L-threonine, however,
the biomass yield was significantly higher, about
12 g/L, but the content of the polymer did not
exceed 42 % of DCW (Kimura et al., 2008).
Moreover, Nakamura and co-authors (1992)
proved the possibility to produce homopolymer
P(4HB), but the content of homopolymer did not
exceed 2 %. To reduce the cost of the polymer the
research on using cheaper carbon sources for the
copolymer P(3HB-co-4HB) synthesis have been
carried out, including oleic acid and saturated
fatty acids with even amount of carbon atoms
(?12-?18) (Rahayu et al., 2008; Ramachandran
et al., 2011), alkanediols (Vigneswari et al., 2009),
worked out palm oil (Rao et al., 2010), soy bean
oil (Park, Kim, 2011), nevertheless, no significant
yields of the copolymer P(3HB-co-4HB) on these
substrates have been obtained.
Thus, using the recently produced, new
natural strains and recombinant producers the
possibility of synthesizing the copolymer P(3HBco-4HB) with different yields (from 2 % to 70 % of
DCW) of the copolymer with the content of 4HB
up to 100 mol % at various yields of the biomass
yield (from 2 to 12 g/L) has been determined.
In this work the results of studying the
synthesis of the copolymer P(3HB-co-4HB) by
two natural strains of hydrogen-oxidizing bacteria
are presented. The analysis of the capability of
the strain R. eutropha B5786 characterized by
the high yields of poly(3-hydroxybutyrate) (up to
90 % of DCW) at different substrates including
the synthesis-gas (Volova, Voinov, 2004) and
the copolymers poly(3-hydroxybutyrate-co-3hydroxyvalerate) (Volova et al., 2007) and 3- and
4-component PHA formed by the monomers
?4, ?5, ?6, ?7 (Volova et al., 2006) has shown
that content of the monomer in the copolymer
did not exceed 4 mol %, the maximum biomass
production and copolymer content amounted for 6
g/l and 60 % of DCW, correspondingly, due to the
pronounced inhibiting effect of ?-butyrolactone
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Natalia O. Zhila, Tatiana G. Volova? Microbial Synthesis and Characterization?
on this strain. Attempts to change the dosage
mode of ?-butyrolactone (fractional adding)
allowed to increase the total biomass and the
copolymer production, unfortunately, the content
of 4HB did not exceed 6 mol % as in autotrophic,
so in heterotrophic conditions.
As shown in the presented work, for the
strain R. eutropha B5786 with concentration of
?-butyrolactone exceeding 6 g/L in the culture, the
decrease of the yield and copolymer production
is observed due to the inhibiting effect. Another
studied strain C. eutrophus ?10646 selected for
the sustainability to ?-butyrolactone influence
(registered in the collection of the Russian
Collection of Industrial Microorganisms in 2010)
allowed us to obtain higher results. Unlike the
strain R. eutropha B5786, ?-butyrolactone with
concentration up to 15 g/L in the medium did
not inhibited the growth and the synthesis of
the polymer in the strain C. eutrophus ?10646.
The fulfilled mode of growing allowed to obtain
good results in the biomass production (to 7-8
g/L of 72 hours of cultivation) with the yield
of copolymer up to 80-90 % of DCW and the
content of 4HB monomer up to 17 mol %. These
results can be compared with the data of the
other research obtained by Amirul et al. (2008)
during the cultivation of the strain Cupriavidus
sp. USMAA1020 and Chanprateep et al. (2008)
during the cultivation of the strain R. eutropha
A-04.
The samples synthesized in the experimental
amounts with the different ratio of monomers 3HB
and 4HB purified to the homogeneous condition
allowed to study their properties: molecular
mass, degree of crystallinity and temperature
characteristics. At the same time no relation
between the 4HB content and the copolymer?s
molecular mass has been determined. This
confirms the previous results that there is no
correlation between the composition of PHA and
the molecular mass (Volova, Kalacheva, 2005).
The absence of correlation between the molecular
weight of the polymer and the 4HB content
has been also demonstrated for Comamonas
acidovorans (Mitomo et al., 2001). Nevertheless,
Vigneswari et al. (2009) has demonstrated the
inverse dependence between the 4HB content
and the molecular mass for Cupriavidus sp.
USMAA1020.
The crystallinity degree of the copolymers
P(3HB-co-4HB) with the content of the 4HB
ranging from 8.7 to 16 mol % amounted to 4344 % which corresponds to the results of other
authors (Doi et al., 1990; Nakamura et al., 1992;
Tsuge, 2002). Thus, Mitomo et al. (2001) has
demonstrated that the crystallinity degree of
the copolymer with the 4HB content of 19, 38
and 65 mol % amounted for 40, 18 and 27 %,
correspondingly. The authors have assumed
that the crystallinity degree of the copolymer
is minimum when the 4HB content is 40-50
mol % (Mitomo et al., 2001). Nevertheless,
according to the data of the other authors the
minimum crystallinity degree of 17-18 % has
been determined for the polymers with the 4HB
content of 78-82 mol % (Saito, Doi, 1994).
There is uncertainty of the obtained data
in the literature for the melting temperature of
the P(3HB-co-4HB) samples. According to the
data of some authors the melting temperature
of the samples with 4HB content of 7 mol %,
11 mol % and 15 mol % amounted to 173, 169
and 160.8є? correspondingly (Kunioka et al.,
1989; Doi, 1990; Rao et al., 2010), when in the
other papers the melting temperature of the
samples with the similar amount of 4HB is
considerably lower: 114є? and 131.5є? (Xie,
Chen, 2008; Li et al., 2010). In our work the
melting temperature for the set of P(3HB-co4HB) samples amounted to 168-171.9 °C. The
gap between the melting temperature and the
temperature of thermal degradation for the set
of the studied samples made 95.1-115єC. We
# 167 #
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Natalia O. Zhila, Tatiana G. Volova? Microbial Synthesis and Characterization?
have not found any references regarding the
temperature of thermal degradation of P(3HBco-4HB).
Thus, P(3HB-co-4HB) was produced by R.
eutropha B5786 and C. eutrophus ?10646 under
autotrophic and heterotrophic conditions using
?-butyrolactone as source of 4HB. It was shown
that C. eutrophus ?10646 possesses the increased
sustainability to the ?-butyrolactone influence.
The fulfilled mode of growing of C. eutrophus
?10646 allowed to obtain good results in the
biomass production (to 7-8 g/L of 72 hours of
cultivation) with the yield of copolymer up to 8090 % of DCW and the content of 4HB monomer
in it to 17 mol %). A set of highly purified samples
of P(3HB-co-4HB) with different content of
4HB was produced and their physicochemical
properties were studied.
Acknowledgements
This study was financially supported by
the Projects of SB RAS No 93 and 96 and the
Project ?Biotechnologies of novel biomaterials?
(Agreement ? 11.G34.31.0013) in accordance
with Resolution No. 220 of the Government of
the Russian Federation of April 9, 2010, ?On
measures designed to attract leading scientists to
the Russian institutions of higher learning?.
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Volova T.G., Kalacheva G.S., Kozhevnikov I.V., Steinbьchel A. (2007) Biosynthesis of
multicomponent polyhydroxyalkanoates by Wautersia eutropha. Microbiology 76: 704-711
Xie W.P., Chen G.-Q. (2008) Production and characterization of terpolyester poly(3hydroxybutyrate-co-4-hydroxybutyrate-co-3-hydroxyhexanoate) by recombinant Aeromonas
hydrophila 4AK4 harboring genes phaPCJ. Biochem. Eng. J. 38: 384-389
Zhang L., Shi Z.-Y., Wu Q., Chen G.-Q. (2009) Microbial production of 4-hydroxybutyrate, poly-4hydroxybutyrate, and poly(3-hydroxybutyrate-co-4-hydroxybutyrate) by recombinant microorganisms.
Appl. Microbiol. Biotechnol. 84: 909-916
????????? ??????
? ?????????????? ???????????
????(3-????????????????-??-4-????????????????)
?.?. ?????,?*, ?.?. ???????,?,
?.?. ??????????,?, ?.?. ??????????
?
???????? ?????????
?????????? ????????? ?????????? ???????? ????,
?????? 660036, ??????????, ?????????????
?
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ?????????, 79
?????? ????????? ?????????????? ????????????? ????????? ??????????????
??????????????????????
(???)
??????????
????(3-?????????????????-??-4????????????????) (????(3??-??-4??)). ????????? ????????? ????? ???? ??? ??????? ? ???,
??? ?????????????? ??? ??????? ????????? 4-???????????????? (4??) ? ?-????????????
???????? ??? ????????-??????????? ? ??? ??? ?????????? ? ????? ????????? ????? ??????
???????? ? ????????. ? ?????????????? ????????? ??????? ????????????????? ????????
Ralstonia eutropha B5786 ? Cupriavidus eutrophus ?10646 ????????? ???????? ??????????
# 170 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Natalia O. Zhila, Tatiana G. Volova? Microbial Synthesis and Characterization?
????????????? ? ??????????? ?-?????????????, ??????? ??????? ???????????????
??? ???????????? ??????? ????(3??/4??). ???????? ????? ??????????????? ????????
????(3??/4??) ? ????????? ??????????? 4?? (?? 8,7 ?? 24,3 ???. %). ???????????,
??? ????????? 4?? ? ????????? ? ??????? ???????, ?????? 3-??????????????? ?
3-?????????????????, ???????? ? ???????? ??????????????? ??????????; ????????
???????, ??????? ??????? ??????????????? 12 ? 25 %. ????????, ??? ??????? ????????????
????? ???????? ????(3??/4??) ? ???????????????? ?? ??????? ?? ??????????? ?????????
? ????????? ? ??????? ????????, ??????????????, ?? 540 ?? 1110 ??? ? ?? 1,91 ?? 2,76.
???????????, ??? ?????????? 4?? ? ??????????? ????????? (8,7-24,3 ???. %) ?? ?????? ??
??????????? ????????? ??????????? (168,9-172,5 єC).
????????
?????:
?????????????????
????????;
????(3-???????????????-??-4-
???????????????); ?-????????????; ??????-?????????? ????????.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2011 4) 172-182
~~~
??? 579.64:631.46
??????-???????????? ??????????
?????????? ??????? Bacillus subtilis
? ?? ??????? ?? ??????? ???????????
?????? ???????
?.?. ?????????,
?.?. ??????????*, ?.?. ??????????,
?.?. ??????????, ?.?. ?????????
?
?????????? ?????????? ???????? ????
???????? ???????? ? ???????? ????????? ???????? ?????? ???
?????? 450054, ???, ??. ???????, 71
?
???? ??? ?????????? ???????????????? ???????? ???????????
?????? 450001, ???, ??. 50 ??? ???????, 34
?
??? ??? ??????????????? ???????????????
?????????????? ???????? ??. ?????? ????????
?????? 453103, ???????????, ??. ??????, 49 1
Received 3.06.2011, received in revised form 10.06.2011, accepted 17.06.2011
??????????? ??????????? 23 ?????????? ??????? Bacillus subtilis ? ???????????
????????????????? ???????????, ? ????? ???????????? ???????? in vitro. ????????, ???
??????????? ??????????? ????? ?????????? ? ?? ?????????????? ??????? ???????? ?? ????
?????, ?? ??????? ???????????????? ????????. ??? ????????????? ??????? ??????????? ?????,
?????????? ????????????? ???????, ??????, ? ?????, ????? ????? ?????????? ??????
??????. ? ???????????? ? ?????????????? ?????? ????? ??????????? ?????? ???????????
?????????? ???????? ??????? ? ?????????? ??????????? ?????????? ??????? ?????? ?
???????? ? ??????????? ???????. ????????? ?????? ????????? ????? ???????????
??????????, ??? ????????? ?? ?? ??????????? ? ?????????? ???????????? ??????????
???????. ????????, ??? ????????? ????? ??????? ??????????? ???????? ????????,
???????????? ???????, ??????? ????????????? ??????????? ?????? ?????????, ??????????
???????? ? ??????? ? ?????? ? ?????? ?????????? ???????????. ?? ????????? ??????????
?????? ???? ??????? ?????????? ???????? ? ???, ??? «???????????» ???????? ????????
????? ?? ????????, ????????? ????????????????????? ?????????????? ?????? ? ??????????????????? ??????????? ??????????, ??????????? ?????????? ????? ????????.
*
1
Corresponding author E-mail address: krm62@mail.ru
© Siberian Federal University. All rights reserved
# 172 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ??????-???????????? ?????????? ?????????? ??????? Bacillus subtilis?
???????? ?????: ?????????? ?????? Bacillus subtilis, ??????????? ????????, ????????.
????????
?????? ???????? ????? ?? ?????????
??????????? ????????? ? ????? ????????,
??????? ???????? ??? ????????? ?????? ?
????? ?????????????? ??????-???????, ??????????????? ? ???? H 2PO4Ї (Lambers et al.,
2008). ???????? ?? ??????? ?????????? ?????? ??????? ? ?????, ??? ??????????????,
??? ???????, ???????? ???????????? ???????? ??? ?????, ???????? ? ??????????????
????????, ??? ??????? ???????? ??????????????? ??? «????????? ????????» (Bieleski,
1973; Marschner, 1995; Lambers et al., 2006).
???, ???????????? ?????????? ??? ???????? ??????? ? ????????? ???????? ?????????? ????? 1 ?? ? ????? ????????? 10 ??
(Bieleski, 1973; Lambers et al., 2006). ?????????? ??????? ? ??????? ????? ?????, ?
????????, ???????????? ????? ????????,
??? ????????? ? ?????????, ? ? ???????????
??????? ?? ?????? ???????? ???? ?? ?????
(Afif et al., 1993).
? ????? ? ???? ???????? ?????? ?????????????? ?????????? ??????????? ????????,
????????????? ???? ???????? (plant growthpromoting bacteria), ???????? ???????????
????????????????? ????????, ??? ?????????
????? ?? ????????? ?? ??????? ? ???????????????? ???????? ????????????? ????????????? ??????????????? ??? ???????? ???
?????????? ???????????? (Thakuria et al.,
2004; Pйrez-Garcнa et al., 2011).
?????? ?? ???????? ???????? ????????????? ???????? ????????? ?????????????
????? Pseudomonas ? Bacillus (Rodriguez et al.,
1999). ???????? ???? Bacillus ????? ???????????? ? ???????? ??????????? ????????????,
????????? ???????? ?????, ????????? ??????????? ???????????????? ? ??????????
? ???????????? ????????????. ? ??????
???????, ????? ?????????????? ????? ??????? ??????? ?????, ?????????? ?????????? ?
????????????? (????????, Bacillus anthrachis,
B.cereus ? ????????? ??????). ?????? ? ???,
????? ??? ???????????? ???????????? ?????????? ?????? ????????, ????????? ??
?????? ???????? ??????????? ??? ????????
????????? ??????? ? ?????, ?? ? ?????????
???????? ?????????????? ?????? (Selosse et
al., 2004; Pйrez-Garcнa et al., 2011). ??? ???? ??
?????????, ??? ????? ????????????????? ????????, ???????? ?????????? ?? ??????, ????????? ????? ???????? ? ?????????? ??????, ???????? ?????? ???? ? ???????????
???????? ?????????.
? ????? ? ???? ????? ?????? ????? ???????????? ??????????? ????????? ??????? Bacillus subtilis ? ??????????? ????????
in vitro, ? ????? ?? ??????? ?? ???????????
?????? ???????.
???????, ????????? ? ??????
??????? ????????????
??????? 23 ?????? Bacillus subtilis
Cohn, 19 ?? ??????? ???? ???????? ???????????? ??????????? ?????????????
???????????
????????????????
????????? ???????????? ?? ??????????????????????????? (??????????) ??????
???????? ???????, ?.?. ???????? ?? ?????????????? ? ????????. ??????? ????????????? ???? ??????? ??????????? ??
????????????-??????????????? ?????????
? ?????????-????????????? ????????? ???????? ???????????? ?????? (??? ???. ????? ? ??., 1997), ? ????? ??????? ?????????????? ??????? ????? ??????????????
?????????? ??? ? ???????????????????
# 173 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ??????-???????????? ?????????? ?????????? ??????? Bacillus subtilis?
????????????? ????????. ?????? ????????? ?????? (26?, 24D, 11? ? ?1), ?????
??????????, ???????? ?????????????.
????? 26?, ?????????????? ????? ??????
???????????? ???????????-? (??? ???
«????????», ???), ????????????? ? ???????? ???????.
???????? ?????????????? ??? 37 °?
?? ??????? ? ?????? ??????????? ?????? ? ????-????????? ????? (???) ? ????????????? ??????? (???) ???????: ??????
???? (0,5 ?? ???? ?? 1 ? ????), ??????????? ?
1 %, NaCl ? 0,5 %, ? ?????? ??? ?????????
1,5 % ????-????.
????????? ??????-????????????
?????????? ??? ????? ????????
?? ??????? ??????????? ?????
??????????? ??????? B. subtilis ? ??????????? ?????????????? ???????? ????????? ?? ????? ????????? (??????? 10 ?/?,
????????? 1 ?/?, K 2SO4 0,2 ?/?, MgSO4 0,2 ?/?,
?????????? ???????? 0,02 ?/?, ???? 20 ?/?;
?? 6,8). ????????, ??? ??????? ?????????
????????????? ? ?????????? ? ?????? ??????? ????????? Ca2+, Fe3+, ? Al3+ (Delgado,
Torrent, 2000). ??????? ? ???????? ????????????? ???????? ? ????????????? ?? ???????????? ???? ??? Ca 3(PO4)2, ??? FePO4,
??? AlPO4, ??????? ? ???????? ????????????? ????????? ??????? ????????? ? ????? ?
???????????? 5 ?/?. ?????????????? ???????? ???????? ??????? ?? ??? ? ???????????? ? ?????????? ? ??????? ????? ??? 37 є?.
????? ???????? ???????? ????? ? ?????????
? ???????? ?? ?? ????? ?? ?????? ?????????. ????????? ?????????? ??? (????) ?
?????? ??????????? ???????? ????????? ?
??????? 7-10 ???? (????, 1983). ?????????????????? ?????????? ???????? ?????????
????????????? ??? ??????? ??? ???? ??? ?????? ???????? ?????? ?????.
????????? ??????????????????
?????????? ??? ????? ????????
? ?????? ??????????? ?????
??????????? ???????? ?????????? ?????????????? ??????? ????????? ????? ??? ??????????????? ? ?????? ????? NBRIP (National
Botanical Research Institute?s phosphate growth
medium) ???????: ??????? 10 ?/?, MgCl2·6H2O 5
?/?, MgSO4·7H2O 0,25 ?/?, KCl 0,2 ?/?, (NH4)2SO4
0,1 ?/?, ?????????????? ?????? (Ca3(PO4)2, FePO4 ??? AlPO4) 5 ?/?, ?? 7,0 (Mehta, Nautiyal,
2001). ??? ????? ???????? ???????? ????????,
?????????? ?? ???, ??????? ??????????
0,9 %-??? ????????? NaCl, ??????? ????????? ?????? ? ???????????? 109 ??????/?? ??
??????? ????????? ????????. ???? ??????????
????????????? ??????????? ????? NBRIP ?
?????????? ??????, ??????? ????? ????????
?? ?????? ??? 37 є? ? 200 ??/???. C???? ???
????????? ???????????? ? ???????? ????????.
????? 3 ????? ???????? ???????? ???????????????? ??? 27464 g, ???????? ???????????,
? ??????? ?? ?????? ???????? ??????????
?????????? ????????? ???? ???????, ??????????? ? 0,5 ? ???????? ??????? (????
26204-91) (??????, 2001).
????????? ???????
?????????? ?????????
??? ??????????? ??????????? ?????????? ???????? ??????? ?? ???????? ??????
(??? ??????? ???? ??? ????? ?? ????? ?????????) ???????? ?? ???????????? ????? GYEXP (15 ?M (NH4)2SO4, 1 ?M MgSO4,
100 ?M MOPS (????????????????????????? ??????? ?? 7,0), 0,5 ?M FeCl3, 1 ?M NaH 2PO4, 1,5 %-? ????), ? ??????????? 0,2 %-?
???????, 0,1%-???? ?????????? ?????????,
?? 25 ???/?? L-??????????, L-?????????,
L-??????, ? ????? 1 ???/?? ???????, 5 ???/??
?????????????? ? 50 ???/?? XP (5-?????-4?????-3-?????????????, ????????? ????), ?
# 174 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ??????-???????????? ?????????? ?????????? ??????? Bacillus subtilis?
???????????? ??? 37 є? ? ??????? 16 ?. ???
??????? ?????????? ????????? ????????
???????? ??????????? ????? ???????, ????????????????? ? ????????????? ???????????? ?? ???????? 5-?????-4-?????-3??????????????, ???????????? ? ??????
?????? ? ???????? ????????? ??? ?????????. ??? ?????????? ??????????? ?????????? ??????? ?? ??????????? (Payne, Jackson,
1991).
?????? ??????? ?????????? ????????
??????????? ???????? ????????
?? ??????????? ??????
???????????-????????????? ?????????
?????? ?????? ?????? ??????? (Triticum aestivum L.) ????? ????????????? 10 ???????????? ?????????? ???? ???????? B.
subtilis ?? ??????? 20 ? ? ?????? 108 ???/??
?? 1 ? ?????. ????? ????? ????? ????????? ??
?????? ??????? ???????? 80Ч80 ?? ?? ?????????? ??????? (4-5 ??) ??????? ????????
?? 150 ??????????????? ?????. ????? ????????? ? ??????????? ???????????. ???????? ??????????? ????????? ???????. ?????
??????? ? ???????????? ????????. ????? 10
?????? ????? ?????? ? ?????? ??????? ???????? ?? 50 ???????? ?????? ? ???????? ????????,
??????? ????? ????????, ????????? ? ????????? ???? ? ?????????? ?? ?????? ??????? (Trouvelot et al., 1986) ?????????? ?????
??? ???????????? ???????? ???????????????????????? ????????. ?? ???????? ????
?? ??????? ????????? ??????? ?????????????
????????, ??????? ??????????? ?????????,
?????????????????? ???????? ? ???????.
?????????????? ???????
???????????? ????????? ? ???? ????????????? ? ???? ????????????? ????????????.
?????????????? ????????? ??????????? ????????? ? ?????????????? ????????? Micro-
soft Office Excel. ? ???????? ????????? ??????? ???????? ? ?? ??????????? ??????????.
??????????
?????? ??????????? ?????????? ??????? B. subtilis ???????????? ??????? ?? ??????? ????? ????????? ????????, ??? ??? ???
????? ?????????? ?????? ??????, ? ??????????? ? ? ?????? ??????? (????? ??????? ?1,
??, 118??, 962??) (????. 1). ?? ???? ????? ??
??? ???????? ?????????? ?????? ????????.
?????? ??????? ???????? ?????????? ?????? 89??, 922??, 49?? (?? ????????), ??????
?????? ? ?????? 121??, 922??, 171??. ????? ???????, ???????? ??????????? ????? ???????????? ????? 922?? ??????? ????????
?????????? ????? ??????????????????
???????????.
? ????????????? ? ????????????????
??????? ? ?????? ????? NBRIP ???????????
??????? ??????? ??????????? ????????, ???
???????? ?????? ? ????????, ? ???????????
AlPO4 ? ? ????? ?????, ??? FePO4 (????. 1),
??? ??????????? ? ??????? ?????????? (Ahn,
1993). ??? ???? ????? ????????, ??? ??????????? ???????? ??????, ??????????? ?
???? ??????? ?? ??????? ????? ?????????,
? ???????????? ?????? ?????????? ?????? ?
?????? 832??, ? ?? ????? ??? ????????? ???????? ????????? ??????????? ????????????
? ??????????? ????? ??????????. ?????? ??
??????????? Ca3(PO4)2 ????????? ????????
?????? «???????-???????» (11??, 118??,
162??, 24?, 11?), ?????? ??????? ????????? ??
??????????? ?? ??????? ??????????? ?????.
?????? ???????? ???????? ?????? ?????????? ?????? 871??, 962??.
?????? ??????? ??????????? ?????????? ? ?????????? ??????? ?????? ???????,
??? ? ??????? ?? ??????????? ? ???????????
??????????? ???????? ??????????? ?????????? ??????????? ???? ? ??????? 26?,
# 175 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ??????-???????????? ?????????? ?????????? ??????? Bacillus subtilis?
??????? 1. ??????????? ???????? ???????? B. subtilis ? ?????? ?????? (???????????? ????????? ? ????
????????????)
?????
Ca3(PO4)2
???????
??????
?????*
?????**
FePO4
???????
??????
?????
?????
AlPO4
???????
??????
?????
?????
??????????
?????????
26?
139±9,3
2,18
191±9,5
??***
0
1,38
+
24D
132±11,7
2,23
179±16,3
??
0
1,31
+
11?
95±4,8
2,23
166±7,1
??
0
1,38
?
?1
0
2,13
105±3,7
??
0
1,23
?
??
0
2,10
112±8,1
??
0
1,38
?
??2
103±8,6
1,91
105±4,5
??
0
1,23
?
11??
140±9,9
2,41
135±13,2
??
0
1,23
+
11??
108±4,2
1,95
157±12,7
??
0
??
+
49??
156±12,9
2,04
138±6,9
??
0
1,31
?
89??
182±12,4
1,95
135±10,7
??
0
??
?
112??
95±7,7
1,86
162±13,2
??
0
??
+
118??
0
2,23
196±14,0
??
0
??
+
121??
125±6,2
2,04
315±21,4
??
0
??
+
122??
121±8,8
1,91
129±6,3
??
0
??
?
141??
91±5,3
2,00
133±14,6
??
0
1,23
+
161??
81±6,7
2,00
135±8,1
??
0
??
?
162??
135±10,7
2,23
195±15,8
??
0
??
+
171??
113±10,1
1,82
203±19,4
??
0
1,23
+
811??
121±9,2
1,91
160±12,7
??
0
??
+
832??
133±5,4
2,18
181±11,7
1,33
0
??
?
871??
101±8,8
1,95
105±7,1
??
0
1,54
?
922??
156±10,6
2,00
252±18,0
??
0
??
?
962??
0
2,10
161±10,4
??
0
1,46
?
* ?????? ?? ?????? ??? ????, ?? .
**??????? ????????? ?????????? ???????????? ???????? ? ????????? ? ??????????? ??????.
*** ??????? ? ????????? ?????????????.
2
24D, 11??, 11??, 89??, 112??, 121??, 141??,
162??, 171??, 811?? (????. 1).
??? ??????????? ?? ????????? ???????????? ?????? ?????????? ???????? ????? ?????? ????????? ? ????? ????????, ???
?????????????? ?????, ???????????????
??????? ? ?????????? ??????? ?? ??????
?????????? ??????? ??????. ? ????? ? ????
?? ??????????? ??????? ???? ??????????
??????? (11?? ? 26?) ?? ??????????? ???????? ??????? ? ???????????? ???????????-
????????????? ????????. ????? 11?? ???
?????? ???? ??? ????????????? ? ????????
?????? ?????? ???????????? ? ???????????
????????????? ???????????, ? ????? 26? ?
???????? ?????????, ??? ??? ?? ???????? ??????? ?????????? ????????????? ????????????
??????????-?.
????????? ????? ??????? ?????????
??????? ??????? ??? ?????????? ??????????? ?????? (????. 2). ??????? ?????????????
???????? (?.?. ???? ????????, ???????????-
# 176 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ??????-???????????? ?????????? ?????????? ??????? Bacillus subtilis?
??????? 2. ?????????? ??????????? ?????? ??????? ??? ?????????? ????? ??????????? ??????????????????? ???????? B. subtilis (???????????? ????????? ? ???? ????????????), %
?????????? ???????????
????????
B.subtilis 11??
B.subtilis 26?
99,5±0,2
90,7±2,0
82,1±1,8
??????? ????????????? ???????? ? ????????
???????
????????????? ??????????? ????????? ?
???????? ???????
???????? ???????? ? ???????? ???????
???????? ??????? ? ???????? ???????
????????????? ??????????? ????????? ?
???????? ?????????
???????? ???????? ? ?????????????? ?????
????????? ?????????
???????? ??????? ? ?????????????? ?????
????????? ?????????
??? ????????) ??? ???????????? ????????????? ???????, ??? ?? ?????, ??????????? ??
?????????? ??????? ?????? ?? ????????? ?
??????? ????????????. ?????? ????????, ???
? ????????? ????????????? ??? ??????????
????? ????????, ?????????????? ??????????????? ??????? ??? ??????? ???????? ?
??????, ??????? ??????????????? ???????
??????????????? ? ?????? ????? ???? ?????????, ??? ???????????? ???????? ???????? ???????? ?? ????????? ???? ??????????
??????? ?, ??????????, ??????? ????? ?? ????????? ???????? ?????????? ???????????? ???????????. ?????????????, ??? ?????????? ????? ?????????? ????? ?????????,
??? ????????????? ??????????? ?????????,
???????? ???????? ? ???????, ??????????
??????? ???????? ????? ? ????????, ????
???????? ? 10 ??? ???? ? ????????? ? ?????????????????? ??????????.
??????????
?? ?????? ??????????????? ????? ?????????? ??????? ? ????? ????????? ?????????????, ?????? 80-90 % ????????, ???????? ?
???? ?????????, ??????????? ?? ?????????
20,5±1,1
2,6±0,2
1,7±0,1
11,7±1,0
0,1±0,01
0,1±0,04
2,7±0,3
0,1±0,02
0,1±0,01
20,4±1,7
1,9±0,1
2,0±0,1
42,9±3,1
3,1±0,2
1,5±0,1
11,6±1,6
2,3±0,4
3,2±0,5
????????, ??????? ?????? ??? ??????? ??????? ???????? ????????????? ??? ????????
(Jones, 1998a; Gerke et al., 2000). ??? ????????? ??? ??????????? ????????? ??????????????? ???????????? ?? ?????? ???????
???????????????, ????????? ????????????
????????????? ????????? ???????. ????
????????? ??? ??????????? ???????? ???????????? ? ???????? ?????? ????? ?????????
(???????? ? ??., 2008), ?????? ? ?????????????????? ?????????? ?????????? ???????
????????, ????????? ????????? ?? ?????????? ???????????? ????? ? ?????????? ?????
???????????? ? ???, ?? ?????? ????? ??????
????????-???????, ? ????????? ??? ?????????? ?? ???????.
???? ????????, ??? ?? ??????? ??????????? ????? ??? ????????? ?????? B.subtilis
???????? ?????????? ?????? ?????? ? ?????? ??????? (????? ??????? ?1, ??, 118??,
962??) ? ?? ????? ???????????? ??????
???????? (????. 1). ?????????? ???????????
????????? ??????? ? ??????????? ???????? ??????? ????? ???????? ??????????, ????????? ????????, ??? ????????? ??????? ?
???????? ??????? ?????? ??????????? ????
# 177 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ??????-???????????? ?????????? ?????????? ??????? Bacillus subtilis?
????, ???????? ?? ??????????? ??????? (Hinsinger, 2001; Fankem et al., 2006). ????? ??????? ??????, ??? ?????? ??????? ????????????
???????????????? ?????, ??? ?????? ????????????? ???????, ????????? ? ???? ???????? ???????????? ???? ?????? ????????
?? ?????, ? ?? ????? ??? ??? ???????????
???????? ?????? ? ???????? ????? ???????? ????????????????? ? ???????? ???????,
???????????????, ??- ? ????????????? ?????? (Selosse et al., 2004; Pradhan, Sukla, 2006).
? ????? ? ???? ????? ????????, ??? ?????????? ??????, ??????????? ?? ??????????? ???
????, ???????????? ??? ????????? ?????????????????? ???????????????, ?????????
?????? ???????, ?? ????????????? ???????
???? ???? ?? ????????????? ??????, ?????
?????????? ???????????? ?????????? ???????? ? ?????? ????? (Gupta et al., 1994).
? ????????????? ? ????????????????
???????? ? ?????? ?????? Ca3(PO4)2 ????????? ? ??????????? ????? ??????? ????????,
??? ?????? ?????? ? ????????, ? AlPO4 ? ?
????? ?????, ??? FePO4. ??? ???? ???????????? ??????-??? ??? ????????? ???? ?
? ??????????? ????? (????. 1). ????????, ???
??????????? ?????? ????? ???????? ?????????????? ?????????? ? ??????????? ???????? ? ???????? ???????? (????????, ??????), ??? ????? ????? ?? ?????? ? ?????????
?????????????? ?????????? ????????, ?? ?
? ?? ??????. ??? ?? ?????, ?? ????????? ?????????? ?????? ????? ?????????, ??? ??????????? ???????? ???? ????????????? ?
???????????? ????????????????? ???????
???????, ??????, ? ????????? ????? ? ???????? ? ?????? ????. ?????????? ????????
????????? ????????????, ??? ? ??????? ???
???????? ????? ? ????? (?????????) ??????
? ??????? ?????????? ??????? ????? ?????????? ????????? ??? ???????? ?????????
?????????? ??????, ????????? ????????????
??? ?? ????, ????????? ? ???????? ??? ???????, ?? ?? ? ?????????. ????? ????????, ???
??? ??????? ????????? ???? ? ????? (?????????) ? ???????? ?????? ???????????????
?????????????? ?????? ?????? ???????. ?????????? ?????? ????????? ????? ?????????
?????????????, ??? ????????????? ??????
???? ???????????? ?? ?????? ????????? ??
?????????? ????????, ?? ? ??????? ????????
???????????????? ????????? ??????????
????????? ??????? ??????????????????
?????????? ????????? ??????.
????????, ??? ???????????? ???????,
??????????????? ?????????? ???????????????? (Hinsiger, 2001; Fankem et al., 2006),
????? ?????????????? ???????? ? ????????? ??????? ??? ???????????, ??? ? ???????????? ????????, ????? ???? ??????????
????????? ?????????????? ????????????
????????? ?????? ??????????? (Lambers et
al., 2006; Plassard, Dell, 2010). ???????????
?????????? ???????? ???? ? ??????? 26?,
24D, 11??, 11??, 89??, 112??, 121??, 141??,
162??, 171??, 811?? (????. 1). ??????????
??????????? ?????????? ? ?????? ???????
????? ????????? ???, ??? ??? ???? ???????????????? ????????? ?????? ???????? ?????????? ?????????? ???????????? ?? ????? ????????? ?????????????? ???????? (Moura et
al., 2001), ???????, ??-????????, ??????????
? ???. ? ????? ? ???? ????????? ?????? ??????????? ??????????????, ? ??????? ????????? ????? Sinorhizobium meliloti RD64 ? ?????????????? ??????????????? ???????, ??
????? ? ???????????? ??????????? ???????
?? ?????? ??? ???????? ?????????? ??? ??????????? ????????????? ?????, ?? ? ???????
??????? ??????????? ?????? ?????????, ??????? ???????? ? ???????? ??????? (Bianco,
Defez, 2010). ??????? ?? ?????????, ??? ?????????? ?????? ???????? ??? ???????????
?????? ???????????? ??????, ??????????
# 178 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ??????-???????????? ?????????? ?????????? ??????? Bacillus subtilis?
???????????? ?????????? ???????????????
???????, ? ????? ??????? ? ??????? ????????????, ??? ? ????????? ????????, ????? ????????????? ????????? ? ???????????? ??????????? ??????????? ??????????? ????????,
????????, ?? ???? ????????? ????????????
??????.
??????, ?? ??? ??????, ??????????????
???????? ??????? ?? ??????????? ???????? ? ??? ??????????? ?????????? in vitro
?? ????? ??????, ????????? ? ????????? ?????? ?????????? ????????? ??????-???????
?????????????? ?????? ????????????? ????
? ???????? ?????????? ????????? ??????????????? (Hinsinger, 2001).
??????????? ???????? ? ??????? ??????????? ?????????? ? ?????????? ???????
B. subtilis ????????? ???????????? ???????????? ???????? ???? ??????????????? ?
??????????? ???????? ???????? (Gyaneshwar
et al., 2002; Pйrez-Garcнa et al., 2011). ?????? ?
???????????? ???????? ???????? ????? ???????????? ???? ???????. ??-??????, ??? ?????????? ??????? ???????? ??????????? ?
???????? ?????????, ? ? ??????? ????????
?????????? ????? ???????????? ???????????? ???????, ??? ?????????? ?????? ??????
??????????? ????????? ?????????? ? ??????????? ???????? ???????? (Lambers et al.,
2006). ??-??????, ????????? ??????????
?????????? ????????????? ???? ??????
???????????? ?????? ?????? ??????????? ??????????? ?????? ?????????? ??????? ????
???????? ? ??????????? ???????? ???????? ?? ?????. ?, ???????, ?? 90 % ????????
???????? ????????? ???????? ? ??????????????????? ???????, ? ?????????? ????
????????? ??????? ????????-?????? ???????
?????????? (Selosse et al., 2004).
????????, ??? ??? ??????????? ??????
?????????? ??????? ???????? ????? ????????? ???????? (Rouached et al., 2010). ???
??????? ??????????? ???, ??? ? ???????? ?????????? ? ???????? ??? ????????? ??????
?????????? ??????? ?????? ????????? ??????????, ?????????? ??????????? ?????????? ??? ????????? ??????????? ?????????
?????? ?????????? ??????, ??????????, ???
? ?????????? ??????? ??????????? ????????
?????? (Hammond, White, 2008).
??? ??? ??????????, ??????? ? ???????????? ????????, ????? ??????????????
??????????? ?????????? ?? ?????????
???? ? ????? ? ??????????? ???????? ???.
?????? ?? ?????, ????? ????????????, ??? ?
????????, ??????? ?? ????????? ?????? ???????????? ?????????? ? ?????????? ???? ??
????????? ????? ??????? ? ?????????, ??????????? ? ??????????? ???????? ????????
????? ?? ???????? ??????????? ? ??????????? ???????, ??????? ????, ??? ?????????
??????????? ?????????? ????????? ????????????? ??? ??????????? ? ??????? ?? ????
?????? ? ?????? ????????. ????? ????????
????? ???? ????? ?? ???????????? «?????????????» ???????? ??????????? ????????
????????? ? ???????? ??????????? ?????????? ????????????? ??????????? ??????
?????. ???????? ???????? ?? ?? ????????? ?
??????????, ??? ?? ??? ? ?????????????????
??????, ?????????????? ??? ????????????? ??.
????????????? ????? ???????? ????????????? ?????????? ???????? ???????????? ?? ???????? ??????? ?????????? ?????
??????????? ???????? ?????? ?? ??????????????? ? ???????? ???????? ? ?????? ????????. ????????? ????? ??????? ?????????
??????? ??????? ??? ?????????? ??????????? ??????, ?? ??????????? ??????? ????????????? ???????? ? ???????? ???????, ??????? ????????? ????????????? (????. 2). ???
????????? ????????????, ??? ?????????????
???????? ??????????? ?? ?????? ?? ??????-
# 179 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ??????-???????????? ?????????? ?????????? ??????? Bacillus subtilis?
????? ???????? ? ???????????? ????????
?? ????????? ?????? ??????????? ??????????????? ????? ????????? ? ??????. ??????
?????? ?????? (?? ????????? ? ?????????)
?????????? ????????????? ??????????? ?????????, ???????? ???????? ? ??????? ? ???????? ??????? ? ????? ????? ???? ??????????
????????? ??????? ???????? ??????? ???????????? ?????????? ? ????? ?? ?????????
?????????? ???????? (???????, ?????????,
2006).
??????????? ?????? ??????????? ??????? ?????????? ???????? ?? ?????????????
?????? ???????, ??-????????, ?????????
??????????? ??????????? ?????????? ????????? ????????????? ?????????? ??????,
???????? ?????????????????, ????????? ???????? ? ??????????? ??????????? ???????
????????? ???? ?????????????? ?????? in
vitro (????????? ? ??., 2009). ?????? ? ???
?????????? ?????? ? ??????????? ????????? ? ??????????? ????????????? ???????? ????? ??????? ?????? ?????????????? ???????????? ???? ????????. ? ?????
? ???? ???? ??????????? ????? ?????????
???????????? ????????? ????? ??????????????? ?????????? ???????? ? ??????????? ???????.
?????? ??????????
?????? ?.?. (2001) ????????? ?? ?????????. 2-? ???. ?.: ???-?? ???, 689 ?.
??????? ?.?., ????????? ?.?. (2006) ???? ??????? ? ????????? ?????? ???????? ????
????????? ? ???????? ??????? ?? ?????????? ?????????? ??????? Bacillus subtilis. ??????
??????? ?????????????? ?????????? «?????????? ??????? ?????? ????????: ???? ? ????????? ? ??????????». ??????. ?. 93?94.
???????????? ???????? ?????? (1997) ???. ???. ????? ??., ???? ?., ???? ?., ?????? ??.,
??????? ?., ?.: ???, ?.1-2.
???????? ?.?., ??????? ?.?., ??????? ?.?. (2008) ????????? ??????????????? ????????????? ????? ???????? ??? ???????????? ??????????? ????????. ???????? ?????????? ???????????????? ????????????. ????? «????????. ????????». 1: 68-71.
???? ?. (1983) ?????? ????????? ?????????????. ?.: ?????, 162 ?.
????????? ?.?., ?????? ?.?., ???????? ?.?., ???????? ?.?., ??????????? ?.?. (2009) ????????????? ????? ?????????? ??????? Bacillus subtilis ? ????????? ???????????? ??????? ? ????????. ??????? ????????????? ???????????????? ????????????. 2: 133-137.
Afif E., Matar A., Torrent J. (1993) Availability of phosphate applied to calcareous soils of West
Asia and North Africa. Soil Sci. Soc. Am. J. 57: 756-760.
Bianco C., Defez R. (2010) Improvement of phosphate solubilization and medicago plant yield by
an indole-3-acetic acid-overproducing strain of Sinorhizobium meliloti. Applied and Environmental
Microbiology. 76: 4626-4632.
Bieleski R.L. (1973) Phosphate pools, phosphate transport, and phosphate availability. Ann. Rev.
Plant Physiol. 24: 225?252.
Fankem H., Nwaga D., Deubel A., Dieng L., Merbach W., Etoa F.X. (2006) Occurrence and functioning of phosphate solubilizing microorganisms from oil palm tree (Elaeis guineensis) rhizosphere
in Cameroon. African J. Biotechnol. 56: 2450-2460.
Gyaneshwar P. Naresh Kumar G., Parekh L.J., Poole P.S. (2002) Role of soil microorganisms in
improving P nutrition of plants. Plant Soil. 238: 83-93.
# 180 #
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?.?. ????????, ?.?. ?????????? ??????-???????????? ?????????? ?????????? ??????? Bacillus subtilis?
Hammond J.P., White P.J. (2008) Sucrose transport in the phloem: integrating root responses to
phosphorus starvation. J. Exp. Bot. 59: 93?109.
Hinsinger P. (2001) Bioavailability of soil inorganic P in the rhizosphere as affected by rootinduced chemical changes: a review. Plant Soil. 237: 173-195.
Lambers H., Shane M.W., Cramer M.D., Pearse S.J., Veneklaas E.J. (2006) Root structure and
functioning for efficient acquisition of phosphorus: matching morphological and physiological traits.
Ann. Bot. 98: 693-713.
Lambers H., Chapin F.S., Pons T.L. (2008) Plant Physiological Ecology. Second Edition. Springer,
604 p.
Marschner H. (1995) Mineral nutrition of higher plants. London, Academic Press, 889p.
Mehta S., Nautiyal C.S. (2001) An efficient method for quantitative screening of phosphate-solubilizing bacteria. Curr. Microbiol. 43: 51-56.
Moura R.S., Martin J.F., Martin A., Liras P. (2001). Substrate analysis and molecular cloning of
the extracellular alkaline phosphatase of Streptomyces griseus. Microbiol. 147: 1525-1533.
Payne M.S., Jackson E.N. (1991) Use of alkaline phosphatase fusion to study protein secretion in
Bacillus subtilis. J. Bacteriol. 173: 2278-2282.
Pйrez-Garcнa A., Romero D., de Vicente A. (2011) Plant protection and growth stimulation by microorganisms: biotechnological application of Bacilli in agriculture. Curr. Opin. Biotechnol. 22: 1-7.
Plassard C., Dell B. (2010) Phosphorus nutrition of mycorrhizal trees. Tree Physiol. 30: 11291139.
Pradhan N., Sukla L.B. (2006) Solubilization of inorganic phosphates by fungi isolated from
agriculture soil. African J. Biotechnol. 5: 850-854
Rodriguez J.B., Self J.R., Soltanpour P.N. (1999) Optimal conditions for phosphorus analysis by
the ascorbic acid-molybdenum blue method. Soil Sci. Soc. Am. J. 58: 866-870.
Rouached H., Arpat A.B., Poirier Y. (2010) Regulation of phosphate starvation responses in plants:
signaling players and cross-talks. Mol Plant. 3: 288-299.
Selosse M.-A., Baudoin E., Vandenkoornhuyse P. (2004) Symbiotic microorganisms, a key for
ecological success and protection of plants. C. R. Biologies. 327: 639?648.
Thakuria D., Talukdar1 N.C., Goswami1 C., Hazarika1 S., Boro R.C., Khan M.R. (2004)
Characterization and screening of bacteria from rhizosphere of rice grown in acidic soils of Assam.
Curr. Sci. 86: 978-685.
Trouvelot A., Kough ?.,Gianinazzi-Pearson V. (1986) Mesure du taux de mycorhization VA d?un
systиme radiculaire. Recherche de mйthodes d?estimation ayant une signification fonctionnelle. In: Gianinazzi-Pearson V., Gianinazzi S. (eds.) Physiological and Genetical Aspects of Mycorrhizae. INRA
Press, Paris, p. 217-221.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ??????-???????????? ?????????? ?????????? ??????? Bacillus subtilis?
Phosphate-Mobilizing Activity
of the Endophytic Bacillus subtilis Strains
and their Effect on wheat Roots Micorrhization Ratio
Anna A. Egorshinaa,
Ramil M. Khairullin , Mikhail A. Lukyantsevb,
Zilya M. Kuramshinac and Yuliya V. Smirnovac
a
Institute of Biochemistry and Genetics
71 Prospekt Oktyabrya, Ufa, 450054 Russia
b
Bashkir State Agrarian University
34 50-letiya Octyabrya St., Ufa, 450001 Russia
Sterlitamak State Pedagogical Academy named Zainab Biishevoy
c
49 Lenin st., Sterlitamak, 453103 Russia
a
The ability of 23 endophytic Bacillus subtilis strains to mobilize sparingly soluble mineral and organic
phosphates in vitro has been researched. Detection of such activity and its ratio has been shown to
be depended on the methods used. When solid medium with insoluble phosphates has been used most
of strains solubilized iron phosphate better than the others. In experiments with liquid media the best
mobilization of calcium phosphate has been observed but bacteria didn?t solubilize iron and aluminum
phosphates except a few strains. Some strains displayed phosphatase activity so one can assume
their ability to degrade organic phosphorus compounds. For the first time we advance a hypothesis
that phosphate solubilization could be one of the factors reducing the efficiency of endomycorrhiza
fungi competition with phosphate-mobilizing endophytic bacteria colonizing internal plant tissues.
Wheat grain treatment with endophytic phosphate-mobilizing strains has been shown to reduce some
parameters of root mycorrhization confirming authors? hypothesis.
Keywords: endophytic Bacillus subtilis strains, phosphate mobilization, mycorrhiza.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2011 4) 183-196
~~~
??? 630.*811.4
????????? ?????? ???????????? ???????????
????? ???????? ????????? ??????
? ???????? ?????? ? ??????? ???????
?.?. ???????????*, ?.?. ?????????????,
?.?. ??????????, ?.?. ?????????
?
????????? ??????????? ???????????,
??. ????????? 79, ??????????, 660041 ??????
?
???????? ???? ??. ?.?.???????? ?? ???,
?????????????, ??????????, 660036 ?????? 1
Received 3.06.2011, received in revised form 10.06.2011, accepted 17.06.2011
???????? ? ??????? ??????? ????????? ???????????? ? ?????????????? ???????. ??????
?????? ??????? ? ??????? ?? ?????? ?????? ?? ????????????? ?????????? ???? ?
?????? ?? ????????? ? ????????. ??? ??? ??????? ??????????? ??????????? ??????????
? ??????????? ????? ???????? ?????? ? ???????? ??????. ????? ????? ???? ??????????
?????? ??? ?????? ???????????? ??????????? ????? ????? ???????????, ??? ???????
??????????? ?????????? ????? ?????????. ????????????? ?????????? ???? ???????????
?? ???????? ???????? ????? ?????????? ??????? ??????, ???????????? ??????? ? ?? ???????
??????????? ??????? ????????? ??????. ???????????? ??????????? ??????????? ????????????
? ??????? ?? ???????? ??????? ? ??????? ???? ????? ??????? ??????. ? ????????? ????????
??????? ???? ?????????? ???? ????????? ?????????? ??????? ? ??????????? ????????????
??????-?????. ????? ????????? ??????????? ???????? ?? ????????? ????????? ???????
?????? ?? ??????? ?????? ????????? ?????? ?? ??? ????????, ??????????????? ?????? ?
??????? ?????????. ?? ?????????? ?????????? ??????? ????? ?????? ? ??????? ?????????? ?
??????? ????????????? ???????????? ???????????: ? ?????? ????????? ????????? ??????????? ??????, ? ? ??????? ????????? ?????? ??? 0,25.
???????? ?????: ????????? ????, ??????? ???????, ??????????? ????????? ?????????
?????????, ??????????? ??????? ????????? ?????, ?????? ???????????.
????????
?????? ? ????????? ???????????? ?????????? ????????? ????????? ??????? ??????-
*
1
?????? ??? ????????? ???????? ??????????
???????? ?????????? ?????????? ? ????????
????????????? ? ???????????????? ???????
Corresponding author E-mail address: isvider@mail.ru
© Siberian Federal University. All rights reserved
# 183 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ????????????? ????????? ?????? ???????????? ??????????? ????? ???????? ????????? ?
? ???????, ??????? ??????????? ? ????????????? ??????????? (????????, ???????
? ??., 1981; Liang et al., 1997; Kirdyanov et al.,
2003; Panyushkina et al., 2003). ???????????
?????? ????????????? ??????????? ???, ???
????????? ????????? ??????? ????????? ???
???????? ????????????????, ?????????????
? ?????????????? ????????. ????????????
??????????? ????????? ???????? ????????
??????? ? ??????? ??????, ??????? ???????????, ????????????? ???????, ????? ??? ??????????, ????? ????????? (????????, ??????????, 1979; ????????? ? ??., 2001; Carlquist,
1988). ????? ????, ?????? ???????????????
?????????????? ????????? ??????? ????????
??? ??????? ??????? ? ??????????????? ???????????????? ?? ?????????? ????????, ?
?????? ? ???????????? ????? ??????????
???????? ????????, ?????? ? ???????? ?????????? ?????? (?????????, 1969; Panshin et
al., 1970; Siau, 1984; Zobel et al., 1989). ??????
???????????? ??????????? ???????? ??????
? ?????? ? ??????? ??????? ????????????
?? ?????? ???????? ??????? ???????, ?? ?
??????????? ???????????????? ? ?????????????? ?????????????, ??????????????
?????????????? ????????? ??????? ? ????????????? ????? ? ?????????????? ??????????? ???????????.
??????? ????? ??? ?? 90 % ???????
?? ???????, ??????? ???????????? ?????
???????, ?????? ?????????, ????????? ???????????????, ????? ?????? ? ????????
????????, ?.?. ???? ??????????????? ??? ?????????? ?? ??? ???????? ??? ?????????????
????????? (????, 1969; Zimmermann, 1983). ?
??????? ???????? ????????? Pinaceae ? ?????????? ? ????????? ????? ? ??????? ?????
?????????? ???????? ???? ????????, ????????? ? ???????????? ??????? ????? ? ????? ?
???????????????? ???????? ????????? ? ??????? ??????. ??????????? ??????????? ??-
??????? ???????? ????? ???????? ??????????? ??????????????? ?????????? ????????
??????? ? ?????????? ???????????, ???????
???????????? ????????????? ???????? ???????, ? ?? ????????????? ?? ???????? ??? ????????????????? ????????????. ? ??????????
??????? ??? ???????? ???????????? ??????????? ??????? ???????, ??????????? ???????
?????? ? ??????? ????????? ??????, ????????? ????????????? ? ?????????? ??????? ??????? ????????? ??????????? ??????.
????? ???????? ??????????? ????????????
???????? ??????? ? ???????? ??????? ? ??????? ????????? ????????????? ????? ??????
? ??????? ?????????. ????????????? ?????
?????????? ????? ?????? ? ??????? ????????? ?????? ????????????? ?????? ???????? ???????? ????? ? ????????? Pinaceae ??
????????? ?? ??????? ?????? ?? ????????
Araucaricaceae ? Podocarpaceae (Jagels et al.,
2006). ? ?????? ?????? ????? ???????????
???????? ???????? ??????????? ??????? ?
??????? ???????? ? ???????? ???????? ?
?????? ?????????. ????? ? ????? ?????? ??????????? ???????? ???????? ???????????
??????? ? ????? ???????? ???????? ? ???????? ???????? ? ??????? ????????? (??????????, 1979; ??????? ? ??., 1985; Panshin et al.,
1970; Vysotskaya, Vaganov, 1992). ???? ??????
?????? ????????? ???? ???????? ??????????? ??????? ???????, ??????????? ???????
?????? ? ??????? ????????? ?????? ???????????: ? ??????? ?? 8 ?? 70 ?????? ? ??????????? ??????? ????????, ?? ????? ???????
?? 10 ?????? ? ??????? ????????? ??????,
?? 1 (??? ???? ????? ???????) ?? 50 ??????
??? ?????? (??????????, 1979; ??????? ? ??.,
1985; Vysotskaya, Vaganov, 1992).
? ??????? ?? ??????? ???????????????,
???????? ???????????? ????????? ??? ??????????????, ??? ? ???????????? ???????.
?????????????? ??????????? ???????? ????-
# 184 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ????????????? ????????? ?????? ???????????? ??????????? ????? ???????? ????????? ?
??? ?? ???????????????? ?????? ? ????????
???. ???????? ????????? ??????-????????
???????? ???????? ?????? ???????? ?? ?????? ????? ??????????????? ???????? ???????
??? ??????????? ???????. ?????? ??? ????????????? ???????? ??????? ?? ??????????? ??-??
????????????? ???????? ??? (Schulte et al.,
1988; Tyree, et al., 1991; Zwieniecki et al., 2001;
Lancashire et al., 2002; Aumann et al., 2005;
Sperry et al., 2005; Pittermann et al., 2006?).
????????? ?????????? ?????????? ??????? ???? ?????????? ??? ?????????????
????????, ?? ???????? ? ????????? ?????????? ?????????, ??? ????? ? ????? ?????????
? ??????????? ???????? ? ??????????? (????????) ??????????? ??????. ????????????
? ?????????, ????????? ?????? ????????,
??? ????, ??? ?????? ?????????????? ???????????????? ????????? ? ??? ??????
?????????? ??????? ?????? (Zimmermann
1983; Sperry et al., 1990). ???????????? ? ?????????, ????????? ? ???????????-???????????
???? ? ???????????, ??? ????, ??? ??????
??????? ?????? ??????? (Davis et al., 1999;
Pittermann et al., 2003; Pittermann et al., 2006?).
??????????? ????????????? ???????? ????????????? ??? ??????????? ????????? ??????
??????? ? ????????? ??????? ?????????
?????? ? ???????? ?????? (Hacke et al., 2001;
Pittermann et al., 2006?).
?????? ?????????? ??? ?????????? ???????? ??????? ????????? ??????, ??? ?
????????? ??????? ?????? ? ??????? ?????? ????? ????????????? ???????????: ?????????? ??????? ???????? ?? ????????????
? ?????????????? ????????, ? ?????????????? ????? ? ?????, ?.?. ?????????? ?? ?????????????? ????????? (Zimmermann 1983;
Zobel et al., 1989; Hacke et al., 2001). ???? ?
??????????? ??? ????? ???????? ??????????
???????, ??????????? ?? ???????????? ?????????????? ????? ??????, ?????? ? ??????,
???????????? ? ???? ????????????? ????????????????? ??????????????? ????? ??????? ??????????, ???????????? ? ???? ?????
??????, ??? ??????????? ??????????????? ?
????????????? ??????????? ?? ???????????? ??????? (Gartner, 1995; Lacointe, 2000).
?????????, ????? ??????? ????????? ??????: ?????????? ????????? ??????? ??????
? ??????? ?????? ???????? ??????????????
???????? ????????, ??????????? ?????????????? ?????????, ?? ??? ???? ?????????? ???????????? ?????????, ? ????? ?????????????
? ????????? ? ????????.
??? ?????? ??????????? ????? ????????? ??????? ????? ???? ????????????? ?????????? ???????? ??????????????? ??????
? ????????? ??????? ????, ????????????
???????????? ???????, ? ??? ???????. ???????? ??? ???????? ?????????????? ???????
? ?????????? ????????, ??? ???????????? ?
?????????????? ??????? ??????????? ??????? ???????? (Ewers, 1985; Carlquist, 1988).
????? ?????????? ? ??????? ? ???? ???????????? ???????????? ??????? ? ?????????????? ?????????? ?????????? ?????????? ?
???????????? ???????. ???????????? ??????? ?? ?????? ??????????????? ?????? ?????????? ????? ???????, ??? ??????? ??????????? ?????????? ???? ??????? ????????, ?.?.
???????????? ?? ?????????? ??????, ? ??????
????? ?????????? ????????? ????????? ????????? ??????. ????????? ???? ?????????
?????? ????????????? ???????? ??????,
?? ????????? ????????? ?????? ????????? ?????? ??????????? ???????? (??????)
????????, ? ??????, ???????? ??????????,
?? ???????? ???????????? ????????, ? ???
????? ??????????? ????????????? ????????
(Barnet, 1981; Larson, 1994). ??? ????????, ???
????????????? ???????????? ????? ???????
??????????. ?????? ??????? ??????????????? ??????? ???????? ??? ? ??????????,
# 185 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ????????????? ????????? ?????? ???????????? ??????????? ????? ???????? ????????? ?
??? ? ? ???????? ???????? ??????????, ???
??????? ???????? ??????? ?????????? ??????????? ????? ????? ???????????????
??????????????? ???????????? ? ???????
????????????? ? ???????????? ???????????
??????????.
????? ????? ?????? ???? ?????? ???????????? ??????????? ????? ??????????
???????? ?????? ? ???????? ?????? ????????, ??? ??????? ??????????? ?????????? ????? ?????????????? ? ????????????
????????? (1); ? ?????????? ??????????????? ???????? ??? ?????????? ???????? ?????
??????? ?? ???? ?????? ? ??????? ?????????
?? ?????? ??????, ??????????? ?????????????? ? ????????????? ??????????????
?????????? ?????? ??????? ??????? (2).
????????? ? ??????
?????????? ??????? ??????? (???) ? ?????? (???), ??????? ????????? ?????? (???)
???? ???????? ? ???????? ??????? ???? ????? ???????, ??????? ??? ????? ???????
?????? (????. 1). ??????? ?????????? ?? ????????????? ???????, ??????? ? ???????????
??? ??????? ????. ?????? ?????????????
???????? ????? ??????????? ?? 0,1 ?? 3,0 ??,
????? ????, ??? ????? ???????? ?? ??????
????? ??????, ?????????? ? ??????? ?????????. ??????? ????????? ???????????? ?????:
?????, ?????? ??????? ?? ??????? ????????
?? ?????? 1,3 ? ??? ??????? ? ????? ???????????????, ??????? ?????????? ?????????
?? ??????? ?? ??? ?? ??????. ????? ??????
? ???? ? ??????? ?????????? ?????. ???????,
?????? ?????????, ??????????? ? ????? ???????, ???? ? ????????? (1:1:1) ? ??????? 3?6 ?????. ?????????? ????? ???????? 10?15 ???,
?????????? ? ??????? ??????? ?????????,
?????????? ?????????? ??? ???????????
?????, ????????? ? ????????? ? ????????
??? ????????-???????? ??? ????????? ????-
??. ? ???????? ??????? ??? ????????? ???????? ??????? ????????? ?? ???????????
????????? ?????? ??????????? (Spiecker et
al., 2000).
????????? ??????????? ??????? ??????
? ??????? ????????? ?????? ????????? ?
??????? ??????-?????????? ? ???? ??????
?????????? ??? ?? ?????????????????? ???????????? ?????????????????? ????????? ???
?????????? ?? 225 ?? 1500? (??????? ? ??.,
1983), ??? ?? ??????? ??????? ???????????.
??? ????????? ? ???????? ?????? ????????
?? ???? ?? ?????? ?????????? ????? ???????
?? ??????? ????????????? ????????, ?? ??????????????? ? ?? ????????????? ?? ????
?????????? ?? ?????????? ??????? ??????
? ???????. ?????????? ?????? ???????? ???????? ??? ?????????? ????? ??????????
??????????? ?? ???????? ?? ??????????????
???????. ??????? ????????? ?????? ?????????????? ??? ??????? ??????? ??? ???? ??????????????? ?????????? ?????? ????????.
?????????? ?????? ?????? ???????? ?????????? ?????????? ???????? ??? ?? ???.
????? ? ?????? ???, ??? ? ??? ???????????? ??? ??????????? ???????? ??????, ? ?
????????? ???????? D, h ? l ? ??? ??????????? ??????????? ??????? ???????, ???????
?????????? ????????? ?????? ? ???????????
??????? ?????? ??????????????.
??????????
???????? ??????
? ?????? ???????? ???????????? ?????
?????, ???????????? ?????? ? ??????????
??????????? ? ???? ?????? ???????? ??????
??????? (???. 1). ?? ?????????? ?????????????? ??????????? ?????????? ??????
???????? D = 2h+l. ???????, ??? 2h = aD,
l = (1?a)D ? 2h
# 186 #
a
� l , ??? 0 < a < 1 ? ??(1 a )
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ????????????? ????????? ?????? ???????????? ??????????? ????? ???????? ????????? ?
?????? ?????????, ??????? ???????????? ????? ????????? ????????? ??????? ?????????? ????????? ?????? ? ??????????? ???????
????????. ? ????? ???? ???????? ?????????
????? ????? ???????? ?? ?????????? ???????? ????? ?? ???? ??????????. ????? ????? ???? ????? ??????????? ???????? ????????? a, ??????? ????????? ????? ????? ???
?????????????? ????????: ?????????? ?????? ???????? D, ?????????? ?????? ??????
l ? ?????????? ??????? ????????? ??????
h. ????????? ??? ?????????????? ??????????? ??????????? ?????????? ??????
???????? ???????????? ?? ????, ??? ????????
????????? ????????? ????????? ??????, ??
?????????? ?????? D ?????????? ????????
???????????? ??????? ?????? l ? ???????
????????? ?????? h, ? ????????? ????????
????? ?????????? ????? ?????????? ?? ????
? ?????????? ????????? a.
???????????? ????????? ??????, ?? ????????, ??? ??????????? ??????????? ?????
???????? ????????? ?????? ? ???????? ?????? ???????????? ??????????? ??????????? ????? ????? ??????????? ??????????????? ????????????: ???????????????
? ???????????? ??????????. ?????????????
??????????? ?????????? ???? ??????????
???????????? ??????????? ????????????
???????????, ? ???????????? ?????? ?? ????????????? ?????? ???? ???????? ? ?????????? ?????? ??????.
?????? ???????????? ???????? ????????? ??????? ????????? Pinaceae, ? ??? ????? ??????????? ??????????????, ??????? ?
?? ???????? ?????, ??????? ? ???? ???????
?????? ??????????? ? ????? ????????? ?????? ? ????? ?????? ???????? (Diaz-Vaz et al.,
1975; Zobel et al., 1989). ? ?????? ?? ??????????? ? ?????? ?????????????? ??????????? ??????? ???????, ??????? ???? ????????? ??????
? ?????? ????????, ? ??????, ? ????????????
D
t
l
h
h
???. 1. ?????????? ??????? ????????? ????????:
D ? ?????????? ?????? ????????; l ? ??????????
??????? (??????) ??????; h ? ??????? ?????????
??????; t ? ????????????? ?????? ????????
????????? ????????, ?????????? ????? ???????????????? ????????? ??????? ??????????? ??????? ????????? ?????? ? ?????
??????? ??????????? ??????? ????????.
????????????? ?????????? ???????????? ??????? Vcond (a) ??????????? ?? ???????? ???????? ?????? ???????? ? ?????????
????????. ?? ????????? ??????-????????
???????? ???????? ??????????? ?????? ???????? ?? ??????????????? ????????? ??????????????? ????????? ??????? ??? ???????,
???, ??? ?? ?? ?????, ?????? ??????? ???????
??????????? ???????. ? ???????????? ?????????????? ?????? ???????? ???????? ??????? ??????????? ??????? ?????? ??????????
??????????? ?????? ??????????? ???????
??????, ??? ??? ????????????? ?????? ???????? ??????????. ????? ???????, ????????????? ?????????????? Vcond ???????????????
?????? ??????? ??????????? ??????? ??????
V cond ~ l2 ~ (1-a)2D2 .
(1)
????????????? ?????????? ???????????? ??????? Vmech ??????????? ?? ???????
??????????? ??????? ????????? ??????. ????????? ????????????? ?????? ???????? ??????????, ?? ??????? ??????????? ???????
# 187 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ????????????? ????????? ?????? ???????????? ??????????? ????? ???????? ????????? ?
n
????????? ?????? ??????????????? ??????
??????? ??????? ?????? h:
V mech~ 2h~ aD.
(2)
????? ???????, ?? ????????????? ??????? ??????????? ????????????? ??????????
???? ??????????????? ???????, ???????????? ? ????????????, ?? ????????? ?????????
a. ?????????????? ???? ???? ??????? ?????
??????????????? ? ?????? ????????????????
???????, ??????? ????????????? ??? ?????????? ??? ??????? ???? ????????????? ?????
? ????? ??????? ???????? ? ?????????????
????????????? (????, 1993; ????????????,
2004; Nowak, 2006).
? ?????? ????? ??????? ????? ????????
??????????????? ??? ???????, ??????? ???????????? ????????? ? ?? ???????????? ???????????? ??????? ????? ????????????????
????????? Vi(x) ????? ???????, ????? ??????????????? ???? ????????????. ??? ????????? ??????????????? ??????? Vi(x) ?????????? ???????????? ????????? ??????????
??? ?????????????, ? ?????? (?????) ?????????? V(x) ?????????????? ?????????????
??????????? ???????:
V ( x) = ? Vi ( x) ,
i =1
??? x ? ????????, ??????????????? ????????? ??? ?????????, ???????? ?????? ??????,
?????? ???? ??? ?????? ??????, ??????? ???????? ???????????, n ? ????? ???????????
???????. ????????????? ????? ???? ??????? ?? ?????. ??????????, ???? ??? ??????????? ? ?????? ?????????? ? ??????????????? ??????? ??? x??. ? ?????? ?????????
????? ???????????? ????????? ?????????????, ?.?. ????? ???? ?? ??????? ??????????,
? ?????? ??????????? ??? ?????????? ????????? x, ????? ??????? ????????????? ?? ??????? ?? x ?????????, ? ????? ???? ????????
(???. 2). ????? ?????????????? ????? ????????? ?????????? ????????????, ??????????? ?? ?????? ?????????? ??????? ???????????.
??? ????? ?? ????????? (1) ? (2), ???
?????????? ????????? a ????????????? ?????????????? ???????????, ? ?????????????
?????????? ???????????? ??????? ?????????????. ??? a = aopt ????? ??????? ?????????? ????????? ?????????????, ?.?. ????????????, ????????. ????? ?????????? ????????
aopt, ????? ?????????? ???? ???????? ???-
???. 2. ???????????? ????????????? ??????????? ??????? ?????????? ?? ????????? ????????? ? ??????
???????????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ????????????? ????????? ?????? ???????????? ??????????? ????? ???????? ????????? ?
??? ??????????? ????? ??????? ??????????
l = (1?a)D ? 0,67D
dV
da
h
0 ?? ????????? ????????? a. ????????,
? ??????? ????? ??????? ???????????????
?????????? ????????? ?????????, ? ?????
??????????? ?????????, ?.?. ? ???????????
???????? 2h/D=aopt.
?????? ???????????? ???????????
??????? ?????????? ?? ????????????? ??????????? ???? ????????: ?????????? ????????
????????? a ?? ????????? ? ?????????????, ?
Vmech ? Vcond ?? ???????? ????????????????.
?? ??? ??????? ???????????, ???? ????????????? ???? ???????? ??? ????????? a. ?????
?????????? ??? ??????????? ? ??????, ?? ????????????? ?????? ????? ???????? ? ?????
??? ????? ??????? ?? ???. 2, ??? ???????????
??????????? ??????? ?????????? ?? ????????? ?????? ? ????????. ? ????? ????????
????????????? ????? ??????? ??????????
????? ???????? ????????? ???????:
V (a ) = ?Vmech (a ) ? ?Vcond (a ) =
= ? aD ? (1 ? a ) 2 ? D 2 ,
??? ?, ? ? ????????????. ??? ????????? ???????? ????????? ? ?????????? ??????? ??????????:
R = ln V (a ) = ln ? + ln Vmech (a ) +
+ ln ? + ln Vcond (a ) .
????? ?????? ??????????? R ?? a, ???????????? ?????????? ????????? ? ????:
dR 1
2
= ?
= 0.
da a 1 ? a
(3)
????? ??????? ????????? (3) ?? ???????? aopt ? 0,33, ? ??????, ????????? ??????????? ???????? ???????? ????????? ???????:
h = 0,5aD ? 0,165D,
a
l | 0,25l .
2(1 a )
(4-6)
????????? ? ????????? (3) ????????
?????? ???? ?????????? ? ???????? a, ?? ?
???????????? ?????? ??????????? ????????
aopt ???????? ?????????? ?????????, ?? ????????? ?? ?????????? ???????? ???????
?????? h ? ??????????? ??????? ???????? D
??? ??????? ?????? l. ? ??????????? ????????, ? ??????? ????? ??????? ??????????
????????? ????????????? ????????, ????????? ??????? ?????????? ????????? ??????
?????????? ???? ????? ?? ??????????? ??????? ???????? ? ???????? ?? ??????????? ??????? ??????. ????? ???????, ???????????
???????? ??????????????? ?? ????????????
?????????? ????????? ??????????? ???????, ??????? ????????? ?????? ??? ??????, ? ???????????? ????? ????. ???????
???????, ??????????? ????? ???? ???????? ? ?????????? ???????? ??? 50, ??? ? 15
???, ?????, ????? ????? ? ????? ????????
????????? 0,67 ?? ??????????? ???????, ???
??????? ?????? ?????????? ???????? ??????? ??????.
????????? ? ????????? ???????
???????? ??????????? ??? ?? ??? ?
??? ???????????? ?? ???. 3. ?? ???? ??????? ???????? ????? ?????, ??????????????? ?????????? ??????????? ????????
(h ? 0,165D ??? h ? 0,25l). ?? ????????? ??????????? ??? ?? ??? ??? ??? ?? ??? ?????
????????? ??????????? ???????? ?? ???????? ?????? ?????????? ??? ?????? ?? ???????? ???????, ?, ????????, ??????????????
?????? ?? ??????????. ?? ???? ?????????
???????? ??????? ???????? ??????????? ??
??? ???????? ?? ????????? ? ????? ????????? ??????????? ????????: ???? ??? ????
# 189 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ????????????? ????????? ?????? ???????????? ??????????? ????? ???????? ????????? ?
???. 3. ???????? ?????? Pinus sylvestris ?? ?????????? ???????. ??????????? ????? ??? ?? ??? (?????)
? ??? (??????), ?????? ?? ???? ?????????? ?????, ????? ?? ?????????? ????????????? ?????????
??????????? ????????
????? ?????????, ? ??????? ??????? ?????
?????? ????????????? ????? ????? ?????????? (???. 3).
?????????? ???????????? ????????????
??? ?? ???, ??? ? ???????? ???? ???????
? ?????????? ?????? ???????? ????? ??????????, ??? ??????, ??????? ?????????????
?? ????????? ??? ?? ???, ??? ???? ?????
?????????, ????????? ? ??????? ??????? ?????????, ? ??????, ????????????? ????, ? ?
?????? ????????? (???. 4). ?? ???? ?????????
???????? ??????? ??????, ??????? ?????????? ?? ?????????? ?? ????? ????? ?????????, ???? ??????????? ??????, ??? ??????
???? ??? ???? ???? ?????.
??????????? ???????? ???????????,
???? ????????????? ?? ????????? ????????
??????, ? ?????? ??? 10 ???????? ?????
(???. 5). ??????? ???????????? ?????? ??
???? ????????? ?? ????????? ? ??????? ???
????????? ???????? ????? ??????????? ?
???, ??? ?????? ? ?????????? ?????? ? ??????? ??????? ???-???? ????? ???????? ??????? ? ??????? ?????? ??????, ??????? ??????????? ????? ???? ? ????? ????? ? ?????
????????? ??????????? ????????.
? ????. 1 ????????? ??????? ????????
??? ????????? ???/??? ????????? ???
?????? ? ???????? ??? ?????? ? ??????? ????????? ? ???????? ???? ?????. ????????? ?
???????????? ?? ?????? ??????????? ????????? h/l = 0,25 (????????? 6) ??????????, ???
? ??????? ????????? ??????? ???????? ???/
??? ???????? ? ??????? ???? ????, ??? ???????????? ???????????, ? ? ?????? ?????????
???????? ? ??????? ???? ????. ??????? ??
?????? ???????? ????????? ???/??? ? ????
????????? ????? ????? ? ????????? ????????, ?? ??? ??? ??????? ?????? ????????????
????????, ?.?. ? ???????? ????????? ??????
?????????? ??????? ???? ?? ??????? ??????. ?????? ? ?????? ???????? ? ????. 1, ???
?????? ???????? ????? ???????? ???????????, ???????????? ??????????? ???????? ???????? ? 95 %-? ????????????? ????????. ???
???????? ???????? ??????????.
???????? ?? ???? ???????? ????? ??????? ???????? ??????????? ???/??? ? ????
????? ???????? ?????. ??? ??????????? ??????? ???????? ???/??? ?? ??????????? ????????????? ??????? ????? ???????? (p > 0,05).
??? ??????????????? ? ?????? ????, ??? ????-
# 190 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
???. 4. ???????????? ?????????? ???, ??? (??????? ??? ???? ?????????? ?????) ? ????????????? ??
????????? ???????????? ???????? ??????? ????????? ?????? h ????? ??????? ??????
???. 5. ????? ????? ??? ? ???. ?????? ??? Pinus sylvestris, 10 ???????? ?????, ????? 1179 ??????.
??? ?????? ????? h=0,25D ? h=0,125D ????????????? ?????? ?????????????? ??????????? ???????
????????? ????? (????????? ? ??????). ????? h=0,165D ????????????? ????????? ???????????
????????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ????????????? ????????? ?????? ???????????? ??????????? ????? ???????? ????????? ?
??????? 1. ??????? ???????? ?? ??????? ? 95 %-? ????????????? ???????? ??? ????????? ???/
??? ? ???????? ??????? ???? ????? ??????? ? ??????? ? ????????? ??????. ??????????? ????????
?
h
l
0,25
???-?? ??????????
???????? ?????/
????????
??????
? ?????
??????
?????????
???????
?????????
Pinus sylvestris L., ????????? ???.,
50? ?.?. 115? ?.?.
17/3
0,18±0,04
0,10±0,06
0,36±0,11
Pinus sylvestris L., ??????????? ?-?,
58? ?.?. 94? ?.?.
13/3
0,19±0,04
0,08±0,02
0,43±0,11
Pinus sylvestris L., ????????
???????????,
56o ?.?. 92o ?.?.
11/3
0,19±0,04
0,11±0,05
0,46±0,13
Larix sibirica Ldb., ?????????,
66o ?.?. 87o ?.?.
62/5
0,21±0,04*
0,06±0,02
0,46±0,10
Picea obovata Ldb., ?????????,
66o ?.?. 87o ?.?.
11/5
0,14±0,04
0,08±0,04
0,33±0,04
??? ? ?????? ?????????????,
????????? ??????????
????? ???/??? ???????? ?????? ? ?????????? ??????????????? ????????? ?????????
??????? ? ????????????? ????? ???????.
??????????
???????????? ???????? ?????? ?? ????????? ???/??? ??? ???????? ????? ????
????? ???????, ?????????????? ? ???????
? ????????? ?????? ? ?????? ????????????
??????????????? ??????, ??????????, ???
?? ???????? ?????? ??????? ?? ????????????? ???????, ??? ???? ???????????? ???????? ??????????????? ???? ????? ? ?? ????????????????? ? ????????????? ????????.
???????? ?? ???, ??? ???????????? ??????
?? ????????????? ???????? ???????? ? ???????????
???????????? ?????? ?????????, ???
??????????? ???????? ??????? ??????? ?????????????? ?? ?? ?????? ????????? ??????,
? ?? ???????? ?????? ? ?????? ?????? ????????? ???? ????????, ? ??? ?????? ??????????
? ?????? ????????? ??????? ? ??????? ??????????????, ? ? ??????? ????????? ????? ??-
???????? ??????????? ???????????? ???????. ??? ??? ???????????? ??????? ???????
??? ?????? ???????? ?? ????????? ????????
???????? ??????? ? ???, ??? ?????? ????????
???????????? ??????????. ??-??????, ???
?????????? ???????? ?????????????? ??????? ??????????????? ??????????. ??-??????,
? ?????? ?? ???????? ?????? ????????????
???????, ???????? ?? ???????????? ????????? ?????? ? ??????????????, ????? ???
???????????? ????. ??????????? ????? ??,
??? ? ???????? ???????? ?? ?????????????
?????????? ???????? ?????? ??-?? ????????? ????????????? ??????, ??? ?????? ???????????????? ????????? ??????????? ??????? ????????? ?????? ? ?????.
???????? ?????? ?????????????? ??, ???
?????? ????? ????????? h ? 0,165D ?? ????????? ??????? ??? ?? ??? ??? ????? ????????? h ? 0,25l ?? ????????? ??????? ??? ??
??? ????????? ????????? ?????? ????????
????? ?? ??? ?????????????? ????????, ??????????????? ???????????? ?????????????
????? ?????? ? ??????? ?????????.
# 192 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ????????????? ????????? ?????? ???????????? ??????????? ????? ???????? ????????? ?
??????????? ?? ??????? ?????????????,
? ????????? ?? ????????? ??????-????????,
????????? ????????????? ???????? ?????
?????????? ?????? ? ??????? ?????????
??????? ??????????? ? ?????????? ? ???????????????? ? ????????? ??????????????
?????? (Zimmermann, 1983; Gibson et al., 1985;
Tyree et al., 1991). ?????? ?????????, ????????? ?? ?????? ? ???????? ???????? ? ??????? ????????, ????? ????????????? ??? ?????????? ?????????????? ??????? (Domec
et al., 2002).
?????? ? ???????? ????????? ?????????? ???????? ????? ??????? ?? ???
????, ?????? ? ??????? ?????????, ??????????????? ??? ????????????, ???????????? ?????? ???????? ??????????????
????????? ?????????, ?? ??????? ??? ????? ??????????? ?????? (Denne, 1989; Park
et al., 2006). ?????????? ? ????????????
????????? ????????? ??? ??????????? ??????? ????? ?????? ? ??????? ??????????
? ???????? ??????? ???????. ????????,
????? ??????? ???????????? ?? ??????????
??????? ????????????? ???????? ?????????
?????????, ??????????? ????????????? ????????????? (Schweingruber, 1988). ????????? ?????? ??????????? ??????????????
? ????????????? ????????, ????? ??????????? ?? ???????? ???????????? ??????????? ??????? ? ??????? ????????? ??????
????? ??????? ?????? (Mork, 1928; Studhalter
et al., 1963; Creber, 1984). ???????? ??????????? ????????????? ??????????? ??????
? ??????? ????????? ???? ?????????? E.
?????? (1928). ?????????? ??? ????????-
????? ????????????? ?????? (Denne, 1989):
? ??????? ??????? ????????? ???>0,25·???
??? ???>0,125·???. ?????? ?????, ??????????????? ???? ???? ?????????????? ??????????? ?????, ???????? ?? ???. 5. ?????,
??????????????? ???????????? ??????????? ????? ??? ? ???, ??????????? ?????
????. ??? ???????????? ??????????? ?????? ? ??????? ?????????, ??????? ??????????? ?????, ????? ???????????? ???????? ?
?? ????? ??? ????? ????????????? ??? ??????????????? ??????????????? ????. ?????????? ?????????? ???? ?????????????
?????? ??? ??????????? ??????? ????? ?????? ? ??????? ?????????? ?? ?????????????? ????????????????? ??????? ? ?????????? ?????????????? ???????????? ????????
??? ??????????? ??????? ????? ?????? ?
??????? ??????????.
?? ?????????? ???????????? ???????????? ??????????? ???????? ??? ????????? ??? ? ??? ??? ???????? ??? ??????????
???????? ????? ??????? ????????? ? ?????????? ????????????? ??? ?? ?????? ?
??????? ?????????: ???? ?????? ?????????
????????????? ?????, ????? ????????? ???/
??? ?????????? ?????? ??? 0,165 ???, ???
?? ?? ?????, ????????? ???/??? ?????????? ?????? ??? 0,25. ???????????? ?????????
???????? ???????? ?? ?????????? ?? ???????????????. ??? ????? ??????????? ? ?????
???????? ????????????? ??? ???????????
? ????????? ? ????????? ?????? ????????, ??
???????? ??????? ?????? ???????, ??? ??? ?????????? ??? ???????????? ??????????????
????????? (Zobel et al., 1989).
?????? ??????????
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??????? ?.?., ?????????? ?.?., ?????????? ?.?., ?????? ?.?. (1983) ??????????????????
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Model Estimation of Optimal Ratio
Between Cell Wall Thickness and Lumen Size
for Tracheids of Conifers in Temperate Zone
Irina V. Sviderskayaa, Vladislav G. Soukhovolskyb,
Ekaterina Y. Radostevaa and Aleksandr V. Kirdyanovb
1
Siberian Federal University,
79 Svobodny, Krasnoyarsk, 660041 Russia
2
V.N. Sukachev Institute of Forest
of the Siberian Branch of the Russian Academy of Sciences,
50 Akademgorodok, Krasnoyarsk, 660036 Russia
The tracheids in conifer xylem both provide mechanical strength and transport water to the crown.
Diameter and wall thickness of conduits influence xylem conductivity and protection against cavitation
and implosion. There is a trade-off between these functions. We aimed to develop an optimality model
and calculate the optimal value of the ratio between lumen diameter and secondary wall thickness
when the compromise between the two functions is reached. The efficiency of conductance function
was evaluated by the volumetric speed of water flow. The efficiency of mechanical function was
evaluated by the area of wall cross-section. The calculated optimal ratio was compared to actual data
for three species growing in Middle Siberia. In the examined rings there were found few tracheids
with the optimum ratio but the straight line of the equation for optimal tracheid separated cells in the
diagrams of wall thickness vs tracheid or lumen diameter into two clusters corresponding to earlyand latewood. We proposed to define the latewood as tracheids in the outer layer of the ring with wall
thickness-to-lumen diameter ratio more and the earlywood as tracheids in the inner layer of the ring
with values of ratio less than the optimal value (0,25).
Keywords: conifer xylem conductance, optimization of wood cellular stru?ture, Mork?s definition of
latewood, trade-off model.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2011 4) 197-208
~~~
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?????????? ???????????? ????????????,
?????? 660041 ??????????, ?????????, 79
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?????????? ????????? ?????????? ???????? ????,
?????? 660036, ??????????, ?????????????, 50/50 1
Received 3.06.2011, received in revised form 10.06.2011, accepted 17.06.2011
???????????? ????????? ????????? ? ?????????????? ????????????? ???????????? ???????,
??? ???????, ?? ?????????????? ?????????? ?? ??????? ?? ????????????. ??? ????????
????? ???? ?????? ??????????? ?????????? ????????????? ? ?????????????????
???????. ?????? ???????????? ?????? ????????? ?? ??????? ??? ??????????? ????????
??????? ? ??????? ?????????, ????????? ?? ?? ??????? ???????, ??? ???????????? ??????????
?????? ??????????? ? ???????????? ?? ?????????. ??? ???????????? ????????????? ?
??????? ???????? ?????????? ?????????? ? ??????? ?????????. ?????? ?????? ?????????
??????? ??????????? ???????????? ????????? ?????????? ?? ????? ????? ? ?????? ??????
? ????????? ????????????. ???????? ????? ?????? ?? ???????????? ???????? ???????????
?????????? ??????????? ?????????? ? ??????????????? ?????????. ???????????????
?????? ????????? ?????????? ? ????? ???????????? ????????: ???????????? ?
????????????. ??? ?????? ?????????? ??????? ????????, ??? ? ?????? ????????? ??????
???????????? ?????????? ????? ????????????? ? ?????? ????? ???????? ? ??? ?????.
???????? ?????: ?????????????? ?????????????, ????????? ??????????, ????????????.
????????
? ????????? ????? ? ????? ? ????????? ??????? ???????? ?? ???????? ????????? ???????????? ?????? ???????????? ??????????? ?????????????? ??????????? ??
*
1
??????????. ???????? ????????? ??????
?????? ???????????? ????????? ?????????.
????????? ??????????? ?????? ???????????? ?????????? ?????????? ????? ????????
?????? ? ?????????????, ?????????? ? ???-
Corresponding author E-mail address: saltykoff.mixail@yandex.ru
© Siberian Federal University. All rights reserved
# 197 #
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?.?. ????????, ?.?. ??????? ??????????? ???????????? ??????? ????????? ????????? ?? ????? ?????
??????? ???? ??????????, ?? ????????, ???
????????????????? ?????? ?????????? ???
?????????? ?????????? ????. ????????????
?????? ?????? ???????????? ????????? ? ?????????? ? ???????????? ?? ?????????????
???????.
?????? ????????? ?????? ? ??????
???????????? ???????? ????????? ???????????? ? ????? ???????????. ????? ?? ????????? ??????????? ???????? ???????????????
??? ???????????? ???????????? ??????????
?? ????? ???????? ? ??? ????? ? ?????? ???????? ????????? ? ?? ???????????? ?????????????? ???????1. ?????????????, ? ?????
???????, ? ???????????? ??????? ?????????
??? ?????????? ?????????? ????????????????? ????? ??????????? ??????? ???????????? ? ???????????? ?????????? ?????? (May,
1971). ????? ????, ?????????? ??????? ????????????? ?????????? (?????, 1933), ????????, ??? ????? ??????????? ????????????
?? ????? ??????????? ?????? ????? ?? ?????
????????? ????? ????????? ?? ???????????
??????????. ? ?????????? ??????? ????????????? ?????????? ??? ????????. ???????? ??????????? ??????????????? ??????????
????? ?? ????? ??????????? ?????????, ????
?? ????? ?? ????????? ?????????? ??????????????????? ?????????????? ????
???????? (????) ? ??????? (???????????,
1978; 1979). ????? ??????????? ??????????
? ???? ??????? ????????, ?????????? ???
????????????? ????, ? ????? ???????? ???????? (????, 1982).
? ?? ?? ????? ????????????????? ?????? ???????????????, ??? ? ?????????
??????????? ??????? ????????????? ?????????? ????? ?? ???????????. ????????
????????? ?????? ????????? ????????
????????????? ?????????? ? ???????? ?????????, ????????????? ? ???????????????
?????????? ????? ?????????, ??????????
????? ??????????? ?????????? (Hutchinson,
1961). ?????? ??????? ??????????? ?? ?????? ? ??????, ?? ? ? ???????? ???????????
(Jonathan, 2009). ??????????? ?????????, ???
???????????? ???????? ????????? ??????
? ??????????? ????? ???????? ? ??? ?????
(????????, ???????, 1978; Winemiller, 1990;
Polis, 1991). ?????? ???????? ?????????????
??????????, ??? ?????????? ???????????? ?
?????? ????? ????? ?????????? ????? ??????????? ? ???????, ?????? ??????? ? ?????
?????? ?????? ???????? ??????????? (Ives,
Carpenter, 2007).
??? ?????????????? ?????? ?????? ? ?????????? ???????, ????? ????????? ??????????? ????????????? ??????????. ???????????? ?????? ????????? ????????? ??
???????? ? ??????????? ?????????, ?? ????
??????????? ????????? ???????????? ??????????? ? ?????? ???????? ?????????. ???
???? ????????? ??????? ????? ??? ???????
???????? ? ??????? ????????????. ???????????? ?????????????????? ?? ????????
? ?????? ???????? ??????????????????
??????????????. ????? ?????? ??????????
????????? ????????? ??????? ??????????,
??, ??? ??? ??????????, ?? ??????? ??? ?????
??????? ?????????.
???????? ????????? ??????????? ???????????? ??????? ???????? ???????? ????????? «???????????? ???????» («predator
switching»), ??? ???????? ? ?????????? ??????? ???????????? ?????? (Pelletier, 2000).
?? ????? ??????????? ??????? ????????????, ??? ??? ??????????? ?? ???????????
????????? ? ????????? ???????????? ?????
????????? ??????????, ??????????? ???????? ???????????? ??????????? ???????
?????????. ? ????????? ???????? ? ??? ?????? ????????? ??????? ???? ????? ??????
?????? ????????? ??? ?? ?????????? ??
????? ??????.
# 198 #
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?.?. ????????, ?.?. ??????? ??????????? ???????????? ??????? ????????? ????????? ?? ????? ?????
?? ???????????? ??????, ??????????
?? ???????? ???????????? ???????????????? ?????????. ?????? ?????? ???? ??????
?????????????? ? ???????? ? ?????????????
???????? (???????? ? ??., 1982; ???????? ?
??., 1990).
??? ????? ???????????? ?????? ?????????? ? ?????? ????????? ?????? ???????
????????? ??????? ?? ??? ?????????? ??????????????? ???????? (????????? ? ??.,
1978). ? ?????????, ????????????? ?????????? ??????????????? ????????? ??????????? ??????? ?? ?????????????? ?????????
????? ?????????? ?????????? ???????????????? ???????? ????????? ???????? «????»
?????? ? ???????????? ?? ??????????, ???
???? ??????????? ?????? ????? ????? ??????, ??? ?? ?????? ?????????? ???????
? ????? ??? ?????????? ?????? ????? ????????????.
??? ????? ??????? ??????? ????????
????? ????? «?????». ? ?????? ?????? ??????
??? ????????? ???????? ???????????? ??????????? ????? ???????????? ?????????????
???????? ? ?????????? ? ???????? ????????
????? ?????? ?? ???????????. ?????? ???????? ?????? ??-?? ??? ???????? ??????? ??
???????????? ????????? (????????, 1987) ?
???????? (????? ?????????? ??????) ???????
? ??????????? «????????????? ????????»,
??????????? ??????????? ?????????????
????????? ?????????? ???????????? ?????????? ? ?????? ????? ???????? ? ??? ????? (????????, ??????? 1978; Ives, Carpenter,
2007). ?????? ???????? ?????? ??? ????, ????? ??????? ????????? ?????????? ??????-
? ???? ???????????? ?????????. ? ??????
?????? ?????? ???? ????????? ?????????
?????????? ??????????????? ????????? ???
?????? ?????????, ????????????? ? ???
?? ???. ?????? ???? ????????? ?????????????? ?????????? ??????, ?? ???????? ?????
????????? ?????????? ?????? ???????????
?????????????? ??????? ????????????????
????????? ? ??????? ?????????? ???????????
???????. ? ????? ?????????? ?????????????? ????????????? ??????????????? ??????: ???????????? ?????????, ??????????????
????????? ???????????, ?????????. ????
???????? ?????????? ?? ???????????????
?????????????????, ?? ????? ????????????????? ??? ???????????.
???????, ????????? ???????????? ?????? ?? ????? ???????????? ?? ????????
?????????????? ???????. ?????? ????????-
?? ????????????? ?????????? ?? ?????? ???????????.
??????? ? ???????? ????????????? ???
???????????? ????????? ???? ???????????
?????? ? ??????? ????????????, ??????????? ????????? ????????????? ???????, ?????????????? ????? ????????? ?????????
? ????????? ?? ???? ??????????? ???????:
??????????? ? ???????????.
??????, ??? ???????????? ???????????? ?????????? ???????? ????????????????? ? ????????? ?????? ?? ???????????
?? ????? ? ????????? ???????????. ?????????? ???????????? ????? ?????????
?????????, ?? ???????? ??????? ????????, ?
???????????? ????? ???????? ? ??????????
???? ?????? ?? ???????? ????????? ???????????. ??????? ?????????? ???????? ??????????? ????????? ??????? ?? ?????????
????? ?????? ?????? ????? ???????????? ??????????? ???????????? ?????????
?? ????? ????? ? ?????? ?????? ??????????
???????????. ? ?????? ?????? ?????? ???
???????????? ??????????? (? ?????? ??????????? ??????????? ??????????) ??????????
??????????? ?????????? ?????? ?????????
???????????? ??????????.
????????? ? ??????
# 199 #
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?.?. ????????, ?.?. ??????? ??????????? ???????????? ??????? ????????? ????????? ?? ????? ?????
???? ???????? ? ??????????? ??????????
???????, ????????? ??????? ???????? ??????????? ??????? ?????????. ????? ??????
????????????? ????????, ?????? ?????????????? ?????, ???????, ? ????????,
????? ??????????????? ? ????????? ????????????, ???? ?????????? ? ????????????
?????? ??????????? ?????? ?????????, ?
??????? ?? ?????????? ?????? ???? ??????????? ????????. ? ??????? ????? ??????
??????????????????? ???????? ??????????? ?? ??????, ?? ??? ???? ???????????
?????????? ????????? ?????????? ??????????? ?? ???????????? ??????????, ??????????? ? ?????? ??????????? ???????. ???
???? ????????? ????? ?????????? ?? ?????
???????????? ? ??????. ????? ??????????-
? ????????? ????? ??????????. ????? ???????, ? ?????? ??????????? ????????? ? ???
??????? ????????, ??????????? ??? ???????
???????????, ?.?. ??? ?????????? ????????
????? ?? ?????????? ????????? ??? ???????
???????????. ????? ??????????? ?????????? ??????????? ??? ?????????????? ????????????? (????????, ???????, 1978). ???????????? ?????? ??????????? ???????????
????????? ???????????? ????? ?? ????????? ? ????????? ?? ????????????. ??? ???????? ???????? ??????????? ???????????
???????????? ?????????????? ???????????
??????? ???? ????, ? ??????? ?????????
????????? ????????? ?? ????? ???????????? ??????????, ??? ???????? ? ?????????
??????? ?????????:
­ dx i
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i
??? xi ? ??????????; yj ? ??????????; Ai ? ???????????? ????? ???????????; B0 ? ?????
?????????? ????????????? ???? ????????;
? ji ? ???????????? ???????? ???????????
??????????? ??????????; Kj ? ????????? ?????????????; dj ? ????????? ????????? ???????????.
? ?????? ?????? ?????????? ??????????? ????? ???????? ?????????????? ???????.
???????? ???????? ?? ????????? ?????, ???,
????????, ????????????? ???????? ?????
??? ?????????????? ?????????????? ???????????? ???????????? ???????????? ?????????? ????????? ?????????? ?????????, ?????, ???? ? ????????, ??? ????????? ???????
?????????? ??????????????? ???? ??????????????? ???????????.
???????????????? ????????????? ??????????????????? ????????? ? ??????????-
??? ?? ????????? ?????????????, ?????????
?????????? ??????, ????????????? ??????
????, ?? ????????? ????????????? ?????
???? ?????????? ???????????????? (Clark,
2010).
??????????? ???????? ????????? ? ???
??? ?????? ??????????? ????? ?????????
????? ???????????????? ?????????? ???????????????? ????? ?????? ????????????
?????????. ????? ???????, ??? ??????????
???????????? ?????? ?? ?????????? ??????????? ? ??? ??????????? ?? ???? ????????
??????? ????? ?????????? ? ????????????
? ??????????????? ??????????? ???????.
??????, ?????????? ?? ?????????????????
?????????????? ? ??????????? ??????????,
?????????? ?? ???????? ??????? ??????????,
??????????? ??????? (1), ????? ?????????
???:
# 200 #
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?.?. ????????, ?.?. ??????? ??????????? ???????????? ??????? ????????? ????????? ?? ????? ?????
­ dxi
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k
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,
P ji xi
y ji d j y ji ¦J jk xk y ji ¦J ji xi ykj
K ji xi
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i zk
??? yji ? ?????????? (?????? ?????? ????????
????? ????????????? ????, ?????? ? ???????????? ??????); Kji ? ????????? ?????????????; ?ji ? ????????? ???????? ????? ???????????? ????????; dj ? ????????? ?????????
???????????. ????????? ??????????? ????????? ?????? (1).
?????????? ? ??????????
????????? ?????????? ? ??????? (1) ???????? ????? ??????????, ?? ???????? ???????? ????????????? ?????????? ????????
???????????? ????????????? ?????? ??????
??????????, ????? ?????? ? ??????? ????????? ?????????? ????? ??????????, ??? ? ???????? ?????????? ?????????????? ????????????? (???. 1).
(2)
? ?????? ???????, ?????????? ?????????????? ????????????? ????????, ??? ? ?????? ?????? (2) ? ???????? ?????? ?????????? ????????? ???????????? ??????? ???
???? ??????????. ????? ????, ? ?????? ?????
?????-??????????? ??????????? ??????????
??????? ????????????? ??????? ? ?????????? ??????? ???????????? ???????. ??????????? ???????? ???????? ????????????
??????? ? ????????? ?? ????? ????? ??????????? ? ?????? ?????? ?????????? ????????
?? ???. 2.
????? ???????, ? ?????? ?????? (2)
?????????? ????????? ???????????? ??????????????? ?? ????? ??????????? ??????
???????? ????? ?????, ??? ????????? ?????
??????????? ???? (? ?????? ?????? ????
???. 1. ??????????????? ??????? ???????? ?????? ??????? (1) (??? ??????????, ???? ?????????) ?
??????????? ???????? ????????? ????? ??????????? A2 ? A1. ????????? ???????: B0 = 2; ?11 = ?12 = 1;
K1 = K2 = 1; d1 = 0,1. ?????????? ?? ???????? ?????????? ?????????? ?????????? ??????? ???????
????????? ??? ???????? ??????????: ??????? ? ???????????? ??????????????? ???? ?????; ????? ?
???????????? ????????? ? ?????????? ????? ?????; ????????? ? ????????? ? ?????????? ????? ?????;
?????? ? ????????????? ??????????????? ???? ?????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????? ??????????? ???????????? ??????? ????????? ????????? ?? ????? ?????
???. 2. ?????????? ??????? ???????????? ????????? ??????? (2) ? ??????????? ????? ?????-???????????.
??????????: ????? ?????????? ????????????? ???????? (B0) ? ???????? ???????? ????? ???????
?????????? (A1). ????????? ???????: A2 = 1; A3 = 2; A4 = 3 ?11 = ?12 = ?13 = ?14 = 1; K11 = 1; K12 = 0,1; K13 = 0,2;
K14 = 0,3; d1 = 0,1; ?11 = ?12 = ?13 = ?14 = 20. ??????? ??????????: ??????? ? ???????????? ???????????????
???? ?????; ?????? ? ?????????
??????????? ???????? ??????????? ? ????
?????????). ? ?????? ???????, ???????????? ?????????, ???? ??????????? ??????
?????????? ????????????? ??? ?????????
????.
????????? ? ?????????? (2) ?????????,
??????????? ? ????? ?? ??????????? ???????, ????? ??????????????? ??? ?????????
????-??????????, ????????? ?????? ? ???,
????? ?? ???? ?????????? ????????????
??????????????? ????? ? ?????? ????????
??????????? ?? ???? ???????? ??????? ?????????? ????? ?????????????????? ???????????. ????????? ???????????? ??????? (2)
??????? ????????? ? ????? ???? ???????
?? ????????????? ??? ????? ????????? ???????? ?ji .
? ????????????? ????? ?????? ?????????? ???????? ???????? ????? ????????????
???????? ? ???? ???????? ????????????
# 202 #
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?.?. ????????, ?.?. ??????? ??????????? ???????????? ??????? ????????? ????????? ?? ????? ?????
???. 3. ??????????? ???????????? ???????? ?????? ??????? (2) ?? ???????? ????????? ???????? ?????
??????????????? ????????. ??????????: ????????? ???????? ????? ???????????? ???????? (?ji) ?
????? ?????????? ????????????? ???????? (B0) ????????? ???????: A1 = 1,5; A2 = 1; ?11 = ?12 = 1; K11 = 1;
K12 = 0,1; d1 = 0,1; ?11 = ?12 = 20. ????????? ?11 ? ?12 ????????????? ??? ???? ????????. ????? ?????????
???????? ? ????????????? ???????? ??? ?ji, ??????????? ? ????. ??????? ??????????: ??????? ?
???????????? ??????????????? ???? ?????; ?????? ? ????????????? ??????????????? ???? ?????,
????? ? ???????????? ??????????????? ? ?????????? ????? ?????
????????, ??????? ? ??????? ?????????????? ?? ????????? ???????????????????????? ?????. ????? ????????????, ???
????? ???????? ?????????? ????? ?????????
? ?????????? ??????? ????????????? ??????? ?, ?????????????, ????????? ??????????? ????????????? ??????????. ?? ???????
3 ?????, ??? ?? ?????????? ?????????? ?????????? ???????? ?????????? ????????? ?ji
??? ??????? ????????? ?0 ????? ?????????
? ?????????? ??????? ????????????? ???????.
??? ???? ????????, ? ?????? ?????? (2)
??????????? ????????? ???????? ?????, ???????????? ? ???????????? ??????????????? ?????????? ?????-??????????? ?? ?????
??????????? ?????????. ??? ?????? ???????? ?????????? ??????????????? ??????????? ?????????? ??????????? ???????????
??????????????? ?????????? ????? ? ????????? ????????????, ??????????? ?? ???????
??????? ????????. ?? ??????? 4 ??????? ??????????????? ??????? ???????, ?????????
?? ???? ??????????? ? ???? ??????????? ?
??????????? ???????????? ????????? ?????
???????????.
?? ??????? 4 ?????, ??? ??????????????? ???? ??????????? ? ????? ?????? ???????? ?????? ??? ?????????? ????? ?????????? ?? ????????????? ??? (????????,
???????, 1978). ????? ??? ?????????? ?????
????? ??????????? ????? ??????????? ????????? ?????????? ??????? ????????????? ??????????????? ??????????? (???. 5).
?? ??????? 5 ???????? ???????????????
???????? ?????? (2) ??? ?????? ???? ??????????? ? ?? ?????? ?? ??????? ???????????. ?????? ????????? ????? ?????????
??????? ???????? ??? ??????? ??????????,
?????? ????????? ? ??? ??????? ??????????. ??????????????? ??????????? ?????????? ????????? ?????? ? ?????? ???? ? ?????
# 203 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????? ??????????? ???????????? ??????? ????????? ????????? ?? ????? ?????
???. 4. ??????????????? ??????? ???????? ?????? ??????? (2) ? ??????????? ???????????? ?????????
????? (?22 ? ?11). ????????? ???????: A1 = 1,5; A2 = 1; A3 = 2; ?12 = ?13 = 1; ?21 = ?23 = 1; K11 = K12 = K13 = 1;
K21 = K22 = K33 = 1; d1 = d2 = 0,1; ?11 = ?12 = ?13 = ?14 = 20; ?21 = ?22 = ?23 = ?24 = 20. ??????? ??????????: ??????? ?
???????????? ??????????????? ???? ?????; ????? ? ???????????? ??????????????? ? ?????????? ?????
?????
???????????. ??? ??????? ? ??????? ?????????? ? ???????? ????????? ????? ??????????? ???????, ??? ? ?????? ???????????,
???????????? ???????????? ??????? ?????????, ??? ???????? ????? ????????? ????????
«????????». ? ??????????? ?????? ?????????? ????????????? ???????? ??????????????? ??????????? ? ????? ??????? ????? ?????????? ?????????.
????? ?????????????? ?????? ?????????
???? ????? ???????? ???????? ????? ??????????????? ????????, ??? ?????????????
??????????? ???????? ????? ??????????????? ?? ??????????????????, ??? ????? ??
??????? ????????. ????? ????????????? ????????? ????????? ??????, ?? ??? ?????? ??????????? ????????????? ??????? ????????
?????, ?????? ?????, ?? ?????????. ?? ?????
??????????? ???????, ??? ??????? ????????
???????? ??????????????? ??? ???????, ??????? ???????????? ???????????? ????????
??? ????? ? ??????????? ???????? ? ?????
?????????????? ????? ?????? ???? ????-
???. ?????? ?????????? ???????????? ?????
??????, ????? ???? ???????? ? ????? ??????
??????? ????????, ????? ???? ??????????
??? ????? ???????? ????????? ????? ????
?????? (???. 6).
?? ???????? 5 ? 6 ???????? ???????????
??????????? ? ?????????? ?????????? ?
??????? ???????? ????????? ?????. ??? ?????? ???????????? ???????????? ??????????? ????????? ???????? ???????????? ???????????? ??? ??????? ???????????? ???
?????????? ???????? ???????? ????? ??????????????? ?? ??????????. ? ?????? ??????????? ???????????? ???????????? ????????? ?
?????????? ??????? ????????? ?????????
???????????? ????? ????????? ??? ?????????? ? ??????? ?????????? ? ???????? ????????? ????? ???????, ??? ? ???????????????
??????????.
??? ????? ??????? ?????????? ????????
????? ???????????? ???????? ??????? ????????????? ??????????????? ???????????
?????????????, ?????? ????????? ?????? ??
# 204 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????? ??????????? ???????????? ??????? ????????? ????????? ?? ????? ?????
???. 5. ??????????????? ???????? ??????? (2) ? ?????? ???? ???????????. ??????????: ?????
?????????? ????????????? ???????? (B0) ? ???????? ???????? ????? ??????? ?????????? (A1). ?????????
???????: A2 = 1; A3 = 2; A4 = 3; ?11 = 4; ?12 = ?13 = ?14 = 1; K11 = K12 = K13 = K14 = 1; ?22 = 4; ?21 = ?23 = ?24 = 1;
K21 = K22 = K23 = K24 = 1; d1 = d2 = 0,1. ??????? ??????????: ??????? ? ???????????? ???????????????
???? ?????; ????? ? ???????????? ????????? ? ?????????? ????? ?????; ????????? ? ????????? ?
?????????? ????? ?????; ?????? ? ????????????? ??????????????? ???? ?????
??? ??? ??????????? ??????????????? ?????????? ???????? ?????????.
? ?????? ???????, ???? ?????????? ?
????????? ?????? ? ???? ?? ???????? ???????? ??????????? ???? ????????? ???????,
??? ???????? ???????? ????? ???????????
?????? ?? ????????. ?????, ??? ????????
??????????, ? ?????? ?????????? ???????
???????? ???????? ????????? ????? ??????????? ? ?????? ?????????? ?????????-
???? ???????? ???????????? ??????? ???
?????????? ?????????? ??????????? ?????????????.
????? ???????, ????? ??????????, ???
? ?????? ????????????? ?????? ??? ?????????? ??????????? ???????????? ??????????
? ???????? ?????? ?????????? ? ?????? ?
??? ????? ?????-???????????. ?????? ???????? ????????, ????? ??????????? ??????
????-?????????? ? ?????? ?????????? ???-
# 205 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????? ??????????? ???????????? ??????? ????????? ????????? ?? ????? ?????
???. 6. ??????????????? ???????? ??????? (2) ? ?????? ???? ???????????. ??????????: ?????
?????????? ????????????? ???????? (B0) ? ???????? ???????? ????? ??????? ?????????? (A1).
????????? ???????: A2 = 1; A3 = 2; A4 = 3; ?11 = 4; ?12 = ?13 = ?14 = 1; K11 = K12 = K13 = K14 = 1; ? 22 = 4;
? 21 = ? 23 = ? 24 = 1; K 21 = K 22 = K 23 = K 24 = 1; d1 = d 2 = 0,1; ?11 = 20; ?12 = ?13 = ?14 = 2. ??????? ??????????:
??????? ? ???????????? ??????????????? ???? ?????; ????? ? ???????????? ????????? ? ??????????
????? ?????; ????????? ? ????????? ? ?????????? ????? ?????; ?????? ? ?????????????
??????????????? ???? ?????.
??????? ????? ???????? ? ????????? ?????
?????-???????????.
?????????: 1). ? ?????? ?????????????
?????? ?????????? ??????????? ?????????? «???????? ?????????» ? ???????????????
?? ????? ????????? ?????????? (? ?????????????? ???????????? ?? ???????) ????????????????. ???????? ????? ?????????,
???? ??????? ????????? ? ?????? ??????????
??????????? ?????????????? ?????? ?????????????? ?? ????????? ????. ??? ???-
??????????? ????????????? ???????????????? ??????????? ? ?????? ????????? ????
???????? ???????? ?ji, ???????????????? ???????? ??????? ?????????? ??????????????
??????????? (???. 3). 2). ??? ???? ????????
????, ?????? ???????? ???????????? ????
????????? ? ?????? ????? ????? ??? ?? ???????? ????????????????? ??????. ???? ???????? ?????????? ? ?????? ?????? (2) ???????? ????????????? ?????????? ?? ??????
??????????? (???. 4) ? ??? ?? ???????????
# 206 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????? ??????????? ???????????? ??????? ????????? ????????? ?? ????? ?????
??????? ???????????? ?????????? ? ???????????? ????? ??????????? (???. 5, 6).
??????????
? ?????? ?????? ?????????? ??????????? ? ???????? ?????? ??????????? ?????????? ???????????? ?????????? ? ?????? ?????
??????????? ?????, ??? ????????? ???????????? ??? ?????? ???????? ???????????. ?
?????? ?????? ??????????? ??????????? ??????????? ????? ????????? ? ?????????????
??????????????? ????? ?????-???????????,
????????????
??????????
???????????
???????????? ????. ? ?????? ???????, ?
??????? ?????? ??????????? ????? ???????
????????????? ?????????? ?? ?????? ???????????, ????????? ? ??????????? ?????????? ???????????.
????? ???????, ????????????? ??????
????? ???? ????????? ??? ???????? ??????? ????????? ????????????? ??????, ?????????????? ????? ????????? ?????????.
????????? ??????????? ????? ????? ????????????? ??? ???????? ????? ????? ?????????? ??????????????????? ??????????????
???? ????????.
?????? ?????????? ?????????????? ???????? ?? ??? ? 50.
1
? ???????? ???? ???????????? ????? ? ????? ????? ??????????? ?????? ??????? ????????? ??????? ?? ????
?????????? ??????? ? ???????????? ?????????? ??????.
?????? ??????????
????? ?.?. (1933) ?????????????? ?????? ? ????????? ?????? ?? ?????????????. ????.
????. 12: 170-177
??????????? ?.?., ???????? ?.?., ??????? ?.?. (1978) ?????? ?????????????? ????
????????? ????????? ?? ???? ????????????? ? ??????????? ????????. ???????? 2:
91-94.
??????????? ?.?., ???????? ?.?., ??????? ?.?., ??????? ?.?. (1979) ???????? ??????????? ??????????????? ?????????? ? ?????????? ??????? Saccharomyces cerevisiae ? ????????? ????????. ???????? ?? ?? ????, ????? ????????????? ????. 5(1): 62-68
????????? ?.?., ???????? ?.?., ??????? ?.?. (1978) ?????????????? ????????? ??????.
?.: ?????, 308 ?.
???????? ?.?., ???????? ?.?., ????? ?.?. (1982) ?????????????? ????????????? ?????????? ?????????? ?????????. ?.: ?????, 272 ?.
???? ?. (1986) ????????. ?.: ???, 376 ?.
???????? ?.?., ???????? ?.?., ?????????? ?.?. (1990) ????????????? ??????? ?????????????. ???????????: ?????. 173 ?.
???????? ?.?., ??????? ?.?. (1978) ???????????? ????????????? ?????????. ?.: ?????,
352 ?.
???????? ?.?. (1987) ?????????? ?????, ????????????? ????????? ? ?????????? ? ????????. ?.: ?????, 368 ?.
Andersen T., Elser J.J., Hessen D.O. (2004) Stoichiometry and population dynamics. Ecology
Letters 7: 884-900
Bartsev S.I. (2004) Stoichiometric constraints and complete closure of long-term life support
systems. Advances in Space Research 34: 1509-1516/
# 207 #
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?.?. ????????, ?.?. ??????? ??????????? ???????????? ??????? ????????? ????????? ?? ????? ?????
Clark J.S. (2010) Individuals and the variation needed for high species diversity in forest trees.
Science 327: 1129-1132
DeMott W.R. (2003) Implications of element deficits for zooplankton growth. Hydrobiologia 491:
177?184.
Hutchinson G.E. (1961) The paradox of the plankton. The American Naturalist 95(882): 137-145.
Ives A.R., Carpenter S.R. (2007) Stability and diversity of ecosystems. Science 317: 58-62.
Levine J.M., HilleRisLambers J. (2009) The importance of niches for the maintenance of species
diversity. Nature 461: 254?257
May R.M. (1971) Stability in multi-species community models. Mathematical Biosciences 12:
59-79.
Pelletier J.D. (2000) Are large complex ecosystems more unstable? A theoretical reassessment
with predator switching. Mathematical Biosciences 163: 91-96.
Polis G.A. (1991) Complex trophic interactions in deserts: an empirical critique of food web theory. The American Naturalist. 138: 123?155.
Winemiller K.O. (1990) Spatial and temporal variation in tropical fish trophic networks. Ecol.
Monogr. 60: 331?367.
Dependence of Closed Ecosystem Models?
Stability on the Number of Species
Mikhail Yu. Saltykova,
Sergey I. Bartsevb and Yuly P. Lankinb
a
Institute of Fundamental Biology and Biotechnology
of Siberian Federal University,
79 Svobodny, Krasnoyarsk, 660041 Russia
b
Institute of Biophysics of Siberian Branch
of Russian Academy of Science,
50 Akademgorodok, Krasnoyarsk, 660036 Russia
Investigations of natural ecosystems by only empirical methods usually are not possible due to their
iniquity. This problem may be solved by combination of empirical and theoretical investigations.
However traditional ecological models can not describe large complex ecosystems correctly. These
models showed that stability of ecosystems should decrease with increasing their complexity. It
contradicts with fact of existence in nature great amount of stability large complex ecosystems. This
paper is devoted to estimation of dependence of closed ecosystem stability on numbers of species in
terms of adaptive metabolism model. In contrast with classical models, in considered one species
consume nutrients in non-fi xed proportion. Considered model ecosystem has two trophic levels:
producers and consumers. With numerical analysis show that in term of it ecosystems stability may
increase with increasing number of included species.
Keywords: mathematical modeling, closed ecosystems, stability.
# 208 #
???????? ?????? ?????????????? ??, ???
?????? ????? ????????? h ? 0,165D ?? ????????? ??????? ??? ?? ??? ??? ????? ????????? h ? 0,25l ?? ????????? ??????? ??? ??
??? ????????? ????????? ?????? ????????
????? ?? ??? ?????????????? ????????, ??????????????? ???????????? ?????????????
????? ?????? ? ??????? ?????????.
# 192 #
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?.?. ??????????, ?.?. ????????????? ????????? ?????? ???????????? ??????????? ????? ???????? ????????? ?
??????????? ?? ??????? ?????????????,
? ????????? ?? ????????? ??????-????????,
????????? ????????????? ???????? ?????
?????????? ?????? ? ??????? ?????????
??????? ??????????? ? ?????????? ? ???????????????? ? ????????? ??????????????
?????? (Zimmermann, 1983; Gibson et al., 1985;
Tyree et al., 1991). ?????? ?????????, ????????? ?? ?????? ? ???????? ???????? ? ??????? ????????, ????? ????????????? ??? ?????????? ?????????????? ??????? (Domec
et al., 2002).
?????? ? ???????? ????????? ?????????? ???????? ????? ??????? ?? ???
????, ?????? ? ??????? ?????????, ??????????????? ??? ????????????, ???????????? ?????? ???????? ??????????????
????????? ?????????, ?? ??????? ??? ????? ??????????? ?????? (Denne, 1989; Park
et al., 2006). ?????????? ? ????????????
????????? ????????? ??? ??????????? ??????? ????? ?????? ? ??????? ??????????
? ???????? ??????? ???????. ????????,
????? ??????? ???????????? ?? ??????????
??????? ????????????? ???????? ?????????
?????????, ??????????? ????????????? ????????????? (Schweingruber, 1988). ????????? ?????? ??????????? ??????????????
? ????????????? ????????, ????? ??????????? ?? ???????? ???????????? ??????????? ??????? ? ??????? ????????? ??????
????? ??????? ?????? (Mork, 1928; Studhalter
et al., 1963; Creber, 1984). ???????? ??????????? ????????????? ??????????? ??????
? ??????? ????????? ???? ?????????? E.
?????? (1928). ?????????? ??? ????????-
????? ????????????? ?????? (Denne, 1989):
? ??????? ??????? ????????? ???>0,25·???
??? ???>0,125·???. ?????? ?????, ??????????????? ???? ???? ?????????????? ??????????? ?????, ???????? ?? ???. 5. ?????,
??????????????? ???????????? ??????????? ????? ??? ? ???, ??????????? ?????
????. ??? ???????????? ??????????? ?????? ? ??????? ?????????, ??????? ??????????? ?????, ????? ???????????? ???????? ?
?? ????? ??? ????? ????????????? ??? ??????????????? ??????????????? ????. ?????????? ?????????? ???? ?????????????
?????? ??? ??????????? ??????? ????? ?????? ? ??????? ?????????? ?? ?????????????? ????????????????? ??????? ? ?????????? ?????????????? ???????????? ????????
??? ??????????? ??????? ????? ?????? ?
??????? ??????????.
?? ?????????? ???????????? ???????????? ??????????? ???????? ??? ????????? ??? ? ??? ??? ???????? ??? ??????????
???????? ????? ??????? ????????? ? ?????????? ????????????? ??? ?? ?????? ?
??????? ?????????: ???? ?????? ?????????
????????????? ?????, ????? ????????? ???/
??? ?????????? ?????? ??? 0,165 ???, ???
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Model Estimation of Optimal Ratio
Between Cell Wall Thickness and Lumen Size
for Tracheids of Conifers in Temperate Zone
Irina V. Sviderskayaa, Vladislav G. Soukhovolskyb,
Ekaterina Y. Radostevaa and Aleksandr V. Kirdyanovb
1
Siberian Federal University,
79 Svobodny, Krasnoyarsk, 660041 Russia
2
V.N. Sukachev Institute of Forest
of the Siberian Branch of the Russian Academy of Sciences,
50 Akademgorodok, Krasnoyarsk, 660036 Russia
The tracheids in conifer xylem both provide mechanical strength and transport water to the crown.
Diameter and wall thickness of conduits influence xylem conductivity and protection against cavitation
and implosion. There is a trade-off between these functions. We aimed to develop an optimality model
and calculate the optimal value of the ratio between lumen diameter and secondary wall thickness
when the compromise between the two functions is reached. The efficiency of conductance function
was evaluated by the volumetric speed of water flow. The efficiency of mechanical function was
evaluated by the area of wall cross-section. The calculated optimal ratio was compared to actual data
for three species growing in Middle Siberia. In the examined rings there were found few tracheids
with the optimum ratio but the straight line of the equation for optimal tracheid separated cells in the
diagrams of wall thickness vs tracheid or lumen diameter into two clusters corresponding to earlyand latewood. We proposed to define the latewood as tracheids in the outer layer of the ring with wall
thickness-to-lumen diameter ratio more and the earlywood as tracheids in the inner layer of the ring
with values of ratio less than the optimal value (0,25).
Keywords: conifer xylem conductance, optimization of wood cellular stru?ture, Mork?s definition of
latewood, trade-off model.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2011 4) 197-208
~~~
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Received 3.06.2011, received in revised form 10.06.2011, accepted 17.06.2011
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??????? ? ??????? ?????????, ????????? ?? ?? ??????? ???????, ??? ???????????? ??????????
?????? ??????????? ? ???????????? ?? ?????????. ??? ???????????? ????????????? ?
??????? ???????? ?????????? ?????????? ? ??????? ?????????. ?????? ?????? ?????????
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? ????????? ????????????. ???????? ????? ?????? ?? ???????????? ???????? ???????????
?????????? ??????????? ?????????? ? ??????????????? ?????????. ???????????????
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????????????. ??? ?????? ?????????? ??????? ????????, ??? ? ?????? ????????? ??????
???????????? ?????????? ????? ????????????? ? ?????? ????? ???????? ? ??? ?????.
???????? ?????: ?????????????? ?????????????, ????????? ??????????, ????????????.
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? ????????? ????? ? ????? ? ????????? ??????? ???????? ?? ???????? ????????? ???????????? ?????? ???????????? ??????????? ?????????????? ??????????? ??
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Corresponding author E-mail address: saltykoff.mixail@yandex.ru
© Siberian Federal University. All rights reserved
# 197 #
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??????? ???? ??????????, ?? ????????, ???
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?? ????? ???????? ? ??? ????? ? ?????? ???????? ????????? ? ?? ???????????? ?????????????? ???????1. ?????????????, ? ?????
???????, ? ???????????? ??????? ?????????
??? ?????????? ?????????? ????????????????? ????? ??????????? ??????? ???????????? ? ???????????? ?????????? ?????? (May,
1971). ????? ????, ?????????? ??????? ????????????? ?????????? (?????, 1933), ????????, ??? ????? ??????????? ????????????
?? ????? ??????????? ?????? ????? ?? ?????
????????? ????? ????????? ?? ???????????
??????????. ? ?????????? ??????? ????????????? ?????????? ??? ????????. ???????? ??????????? ??????????????? ??????????
????? ?? ????? ??????????? ?????????, ????
?? ????? ?? ????????? ?????????? ??????????????????? ?????????????? ????
???????? (????) ? ??????? (???????????,
1978; 1979). ????? ??????????? ??????????
? ???? ??????? ????????, ?????????? ???
????????????? ????, ? ????? ???????? ???????? (????, 1982).
? ?? ?? ????? ????????????????? ?????? ???????????????, ??? ? ?????????
??????????? ??????? ????????????? ?????????? ????? ?? ???????????. ????????
????????? ?????? ????????? ????????
????????????? ?????????? ? ???????? ?????????, ????????????? ? ???????????????
?????????? ????? ?????????, ??????????
????? ??????????? ?????????? (Hutchinson,
1961). ?????? ??????? ??????????? ?? ?????? ? ??????, ?? ? ? ???????? ???????????
(Jonathan, 2009). ??????????? ?????????, ???
???????????? ???????? ????????? ??????
? ??????????? ????? ???????? ? ??? ?????
(????????, ???????, 1978; Winemiller, 1990;
Polis, 1991). ?????? ???????? ?????????????
??????????, ??? ?????????? ???????????? ?
?????? ????? ????? ?????????? ????? ??????????? ? ???????, ?????? ??????? ? ?????
?????? ?????? ???????? ??????????? (Ives,
Carpenter, 2007).
??? ?????????????? ?????? ?????? ? ?????????? ???????, ????? ????????? ??????????? ????????????? ??????????. ???????????? ?????? ????????? ????????? ??
???????? ? ??????????? ?????????, ?? ????
??????????? ????????? ???????????? ??????????? ? ?????? ???????? ?????????. ???
???? ????????? ??????? ????? ??? ???????
???????? ? ??????? ????????????. ???????????? ?????????????????? ?? ????????
? ?????? ???????? ??????????????????
??????????????. ????? ?????? ??????????
????????? ????????? ??????? ??????????,
??, ??? ??? ??????????, ?? ??????? ??? ?????
??????? ?????????.
???????? ????????? ??????????? ???????????? ??????? ???????? ???????? ????????? «???????????? ???????» («predator
switching»), ??? ???????? ? ?????????? ??????? ???????????? ?????? (Pelletier, 2000).
?? ????? ??????????? ??????? ????????????, ??? ??? ??????????? ?? ???????????
????????? ? ????????? ???????????? ?????
????????? ??????????, ??????????? ???????? ???????????? ??????????? ???????
?????????. ? ????????? ???????? ? ??? ?????? ????????? ??????? ???? ????? ??????
?????? ????????? ??? ?? ?????????? ??
????? ??????.
# 198 #
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?? ???????????? ??????, ??????????
?? ???????? ???????????? ???????????????? ?????????. ?????? ?????? ???? ??????
?????????????? ? ???????? ? ?????????????
???????? (???????? ? ??., 1982; ???????? ?
??., 1990).
??? ????? ???????????? ?????? ?????????? ? ?????? ????????? ?????? ???????
????????? ??????? ?? ??? ?????????? ??????????????? ???????? (????????? ? ??.,
1978). ? ?????????, ????????????? ?????????? ??????????????? ????????? ??????????? ??????? ?? ?????????????? ?????????
????? ?????????? ?????????? ???????????????? ???????? ????????? ???????? «????»
?????? ? ???????????? ?? ??????????, ???
???? ??????????? ?????? ????? ????? ??????, ??? ?? ?????? ?????????? ???????
? ????? ??? ?????????? ?????? ????? ????????????.
??? ????? ??????? ??????? ????????
????? ????? «?????». ? ?????? ?????? ??????
??? ????????? ???????? ???????????? ??????????? ????? ???????????? ?????????????
???????? ? ?????????? ? ???????? ????????
????? ?????? ?? ???????????. ?????? ???????? ?????? ??-?? ??? ???????? ??????? ??
???????????? ????????? (????????, 1987) ?
???????? (????? ?????????? ??????) ???????
? ??????????? «????????????? ????????»,
??????????? ??????????? ?????????????
????????? ?????????? ???????????? ?????????? ? ?????? ????? ???????? ? ??? ????? (????????, ??????? 1978; Ives, Carpenter,
2007). ?????? ???????? ?????? ??? ????, ????? ??????? ????????? ?????????? ??????-
? ???? ???????????? ?????????. ? ??????
?????? ?????? ???? ????????? ?????????
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?????? ?????????, ????????????? ? ???
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????????? ? ??????? ?????????? ???????????
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????????? ???????????, ?????????. ????
???????? ?????????? ?? ???????????????
?????????????????, ?? ????? ????????????????? ??? ???????????.
???????, ????????? ???????????? ?????? ?? ????? ???????????? ?? ????????
?????????????? ???????. ?????? ????????-
?? ????????????? ?????????? ?? ?????? ???????????.
??????? ? ???????? ????????????? ???
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?????? ? ??????? ????????????, ??????????? ????????? ????????????? ???????, ?????????????? ????? ????????? ?????????
? ????????? ?? ???? ??????????? ???????:
??????????? ? ???????????.
??????, ??? ???????????? ???????????? ?????????? ???????? ????????????????? ? ????????? ?????? ?? ???????????
?? ????? ? ????????? ???????????. ?????????? ???????????? ????? ?????????
?????????, ?? ???????? ??????? ????????, ?
???????????? ????? ???????? ? ??????????
???? ?????? ?? ???????? ????????? ???????????. ??????? ?????????? ???????? ??????????? ????????? ??????? ?? ?????????
????? ?????? ?????? ????? ???????????? ??????????? ???????????? ?????????
?? ????? ????? ? ?????? ?????? ??????????
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???????????? ??????????? (? ?????? ??????????? ??????????? ??????????) ??????????
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# 199 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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???? ???????? ? ??????????? ??????????
???????, ????????? ??????? ???????? ??????????? ??????? ??
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