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120.Журнал Сибирского федерального университета. Сер. Биология №1 2013

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Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Журнал Сибирского федерального университета
2013
Journal of Siberian Federal University
6 (1)
Биология
Biology
???????????? ?????:
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???????? ??? ?.?. ?????????
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???????? ??? ?.?. ???????
??.-????. ???, ?-? ???.-???. ????
?.?. ???????
??.-????. ???, ?-? ????. ????
?.?. ??????
??.-????. ???, ?-? ???.-???. ????
?.?. ????????
??.-????. ???, ?-? ???.-???. ????
?.?. ????
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
Executive Editor for Biology:
Elena S. Kravchuk
CONTENTS / СОДЕРЖАНИЕ
Ю.А. Дарикова, Е.А. Ваганов,
Г.В. Кузнецова, А.М. Грачев
p=?,=???/L !%?2 C!,","%* *??!%"/. ?%??? (Pinaceae)
" 3??%", . *!=??% !?*%L ???%?2?C,
?3?
Б.Э. Богданов, Т.В. Свердлова, И.Б. Книжин
h?????,"%?2? , 2=*?%?%?,???*,L ?2=23? ?,K,!?*,. C%C3? ?,L
C??2!%?%?%?% C%?*=????,*= Cottus poecilopus complex
(Scorpaeniformes: Cottidae)
? 18 ?
М.Т. Макенов, О.А. Михайлова
r*3?/ ????L ?%K=*=?,: %K?= .=!=*2?!,?2,*=
? 32 ?
И.Н. Третьякова, Е.В. Ворошилова,
Д.Н. Шуваев, Н.С. Лукина
nK!=?%"=?,? *=??3?= , ,??3*?, ?%?=2,???*,. ?=!%?/??L
" *3??23!? in vitro 3 Pinus sibirica Du Tour
? 44 ?
Alexi M. Grachev, Eugene A. Vaganov,
Steven W. Leavitt, Irina P. Panyushkina,
Eugene P. Chebykin, Vladimir V. Shishov,
Natalia A. Zhuchenko, Anastasia A. Knorre,
Malcolm K. Hughes and Mukhtar M. Naurzbaev
Methodology for Development of a 600-Year Tree-Ring MultiElement Record for Larch from the Taymir Peninsula, Russia
? 61 ?
???????? ?.?. ?????? ????????? ?.?. ???????
???????????? ??????? ?.?. ?????????
????????? ? ?????? 22.03.2013 ?. ?????? 84x108/16. ???. ???. ?. 9,0.
??.-???. ?. 9,5. ?????? ???. ?????? ????????. ????? 1000 ???. ????? 1467.
?????????? ? ?? ???. 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
О.В. Колмакова
q%"!?????/? ??2%?/ %C!??????,
",?%?C??,-,??/.
K,%??%.,?,???*,. -3?*?,L K=*2?!,%C?=?*2%?=
? 73 ?
Т.А. Зотина, Е.А. Трофимова,
А.Я. Болсуновский, О.В. Анищенко
}--?*2,"?%?2? 2!%-,???*%?% C?!??%?= !=?,%=*2,"?/.
, ?2=K,???/. ,?%2%C%" ??2=??%" * !/K=?-K??2%-=?=?
!. e?,??L
? 96 ?
М.И. Гладышев, О.П. Дубовская
p?????, ?= ?%?%?!=-,? `.`. o!%2=?%"= &f,??? "
?,?!%?-?!?. n??!*, C% %K??L ?,?!%K,%?%?,,[. $ j,?":
`*=???C?!,%?,*=, 2011. $ 704 ?.
? 108 ?
????????????? ? ??????????? ???
?? ? ??77-28-725 ?? 29.06.2007 ?.
????? ???????? ? «???????? ??????? ????????????? ??????? ???????? ? ???????, ? ??????? ??????
???? ???????????? ???????? ??????? ?????????? ??????????? ??
????????? ?????? ??????? ??????? ?
????????? ????» (???????? 2010 ?.)
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2013 6) 3-17
~~~
??? 630. 228.7:582.475.4(571.51)
?????????? ???? ???????? ???????? ????? (Pinaceae)
? ???????? ???????????? ?????????
?.?. ?????????*, ?.?. ????????,
?.?. ??????????, ?.?. ???????
?
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ?????????, 79
?
???????? ???? ??. ?. ?. ???????? ?? ???,
?????? 660036, ??????????, ?????????????, 50, ???. 28
Received 29.03.2012, received in revised form 15.03.2013, accepted 22.03.2013
???????? ?????? ??????????? ????? ? ?????????????? ??????? ??????????????????
???????? ????? ?????????? (Pinus sibirica Du Tour) ? ????? ???????????? (Pinus cembra L.)
???? ??????????, ???????? ?? ?????? ????? ???????????? (Pinus sylvestris L.). ??????????
??????? ???????? ????? ?????????? ?? ????????? ? ?????? ??????????? ?? 23 % ????,
??? ???????? ?????????? ?? ??????? ????????????. ?????????? ??????? ?????? ?????
???????????? ??????? ?? ???? ??????: ??????? P. sylvestris (?????? ??? ????? ??????????)
???? ???????? ??????????? ???????? ?? 20 % ? ????????? ?????? ? ?? 15 % ? ? ?????????
??????????, ? ?? ????? ??? ??????? ?? ???????? ????? ???????????? (?????? ??? ?????
????????????) ?? 10 % ??????, ??? ? ???????????? ????????? (?????????? ????????). ???
????? ????????????? ??? ????????? ??????? ??????. ?????? ?????????????? ???????
?????? ? ?????? ??????????, ??? ?? ??????? ?? ??????????? ??????? ????????? ???????
??????????? ???????? (????????????? ????? ? ???????????? ??????). ?????? ??? ????????
?? ??????? ??????????, ?????????????? ????? ????????????? ??????? ???????????
????????, ???????? ???? ?????????: ? ??????? ??????????? ????????????? ??????, ?
??????? ???????? ??????? ???????????? ??????????? ? ???????????? ???????? ???????
??????????? ???? ? ???????????? ? ? ???????????? ?????????? ???????? ???? ?????.
???????? ?????: ??????, ??????, ???? ?????????, ???? ???????????, ????? ????????????,
?????? ????????? ??????, ????????????? ??????.
????????
???????????? ?????? ????????? ???????? ????? ?????????????? ???????? ????????-
*
??? ????????? ?????????? ???????? ?????????? ? ??? ????????? ????????? ? ???????????
?????????. ?????? ? ????????? ?????????
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: jualeksa@rambler.ru
#3#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????????? ???? ???????? ???????? ????? (Pinaceae) ? ?????????
?? ?????? ???????? ????? ?????????? ??????? ????????? ????????????? ???????????
? ?????????, ?????????? ????????? ?????? ???? ?? ?????? ?????????????????? ??
??????, ? ????? ???????? ?????????????????
?????. ???????? ????? ????????? ????????? ?????? ? ??? ?????? ???????? ?? ???????
???????? ????????? ????? ? ?? ????????????? ? ???????? ???????? ???????????????
????????, ??????????? ?????, ??????? ????????????? ????? ???????????? ???????? ?
???????????? ????????????? ???????? (?????????, 2003, 2007; ?????, 2004; ???, 2011).
???????? ??????, ???????? ????????????? ? ??????? ????????? ????????, ???????? ???????? ???????????? ????????????
???????? ? ?????????? ??????? ????? ????????? ????????. ??? ?? ????? ???????? ?????????? ?????? ?????????????? ??????????? ????????????????? ? ??????????????
???????? ????????? ?????????? ? ????????
???????????? ????????? ?????. ???????
?????? ??????? ???????????? ?????????????????? ???????? ???????? ??? ?????? ???????????? ??????????, ??????? ? ??????
????????? ???????, ????????????? ??? ????????? ??????.
??????????? ???? ???????? ????????
????????? ??????? ????? ?????? ??? ?????,
??? ? ??????? ??????????? ???????? ????????. ????????? ???????? ???????????? ?????
????????? ???? ??????????? ????????? ?????????? (?????? ? ??????), ???????????, ????????? ?????? ? ??????? ??????????? ????????,
????????????? ??????????? ????????.
???????? ??????????? ?.?. ??????????
? 2010 ???? ?? ???????????? ???????? ???????? ?????, ?????????????? ?? ???????????
????????? ? ???????????? ???? ????????????? ????, ???????? ??????????? ???????? ????????, ????????? ????????? ?????
??????????. ??? ???? ????? ??????? ????-
???????? ? ???????? ????????? (??????????????) ? 55 %, ??????? ? ? ??????????? ????????? ? 42 %. ????? ???????? ???????? ?????
???????????? ??????????? ?????????, ? ??????? ???????????? ?????????? 22 %. ? ?????
? ???? ?????? ??????? ? ????? ??????????
???????????? ????? ?????? ????????, ??????? ???????? ??????? ???????, ???????? ??
????????????????? ????? ????????.
??????????? ????? ? ???????? ????????
????? ??????????, ??????????????? ? ???????????? ???? ????????????? ????, ?????????
????? (?????????, 2001, 2003, 2007; ?????????
? ?????, 2004, ????? ? ??., 2004). ? ??????
?????? ? ?????? ??????????? ????? ????????
????? ?????????? ?????? ? ????? ???????????????? ??????? ????? ? ??????, ?????????? ? ?????? ???? (2002-2010). ????? ??????? ?????? ??????????? ????? ???????? ?????
????????????, ??????????????? ?? ??? ??
?????????. ????????? ???? ????? ????? ????
??????????? ??????????? ????????????? ????? ?????? ? ??????????? ?? ???? ??????.
???????? ????????????????? ??????
? ??????? ?????? ?? ????????? ????? ??????, ?????????? ???????? ????? ?????????? (Pinus sibirica), ??? ? ???????? ?????? ?????????????? ????? ????????????
(Pinus sylvestris), ?????? ????? ??????????
?? ????????? ? ?????????????? ??????????. ??? ???? ?????????? ???????? ?? ?????
?? 25.5 %, ? ???????? ?????? ? 56.7 % ????
???? ? ?????, ????????? ?? ????? (???????? ? ???????????, 2006). ??????? ???????
P. sylvestris ?? ???? ??????? P. sibirica ????
??????????? ??? ? ?????? ????????? (1958),
??? ????????? ?????????????? ???????? ???????? ????? ?????????? ? 6-10 ??? ????????? ??????? ?????????????? ????????, ?????????????? ? ??? ?? ????????.
?????? P. cembra, ??????????? ? ???????? P. sylvestris ? P. griffithii, ????? ???????
#4#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????????? ???? ???????? ???????? ????? (Pinaceae) ? ?????????
?????????????? ??? ???????? ???????????
?????????.
?????????? ?? ????? ?????????, ??? ???????
?? ?????? ??????, ????? ?????, ????? ? ????
????. ? ?? ?? ????? ?? ??????? ?????? ???? ?
????? P. nigra ? P. strobus ? ??????? P. cembra
?????????? ?????? ???????? ?????????????
?????????? (Holzer, 1970). ?????? ????? ?????????? (P. elliottii) ?? ??????? ????????????????? ????? ????? ???????, ??? ?? ??????? ????? ?????????? ? ????????????????
????????? ????? (Schmidtling, 1986).
?????????? ??????????? ??????????,
??? ??????? ???? ??????? ?????????? (????????, ??????? P. patula ?? ??????? P. radiata),
??????????? ??????????? ???????????????
????? ???????????? ???????? (Dyson, 1975).
??????? ?????? ?? ?????? ????? ?????????
? ????????? ?????????? ?? ?????? ?? ????????? ??? ????????? ??????????? ???????
????????. ????????, ?????? ????????????
??????? ??? ?????????? ???????? ?? ??????
???????? ???????????? ???????? ???????? ?? ????? ???????????? (Pinus sylvestris)
??????? ????? ???????? ????????? (Pinus
sibirica) (?????????? ????? ?????? ?????????) ? ????? ???????? ??????????? (Pinus
cembra) (????? ?????????? ?????? ???????????). ???????? ???????? ????? ???? ??????? ? 1963?1965 ??. ? ???????????? ?????????
?? ?????????? ?????????????????? ???????? ????????? ???????????? ??? (56°22? ?.?.,
92°57? ?.?). ? ?????? ?????? ?????? ????????????????????, ??????? ????? ??????? ?????????? 410 ??, ????????????? ???????????
1.3є?, ????? ?????????? ???? 5 є? ?????????? 1968, ??????? ????????????????? ?????????????? ??????? 149 ????, ?????? ????????
?????? ??????????? ? ????? ????????, ???????? ?? ?????? ????? ?????????????? (Pinus
densiflora) (Schmidtling, 1983).
??? ???????? ??????? P. sylvestris ?
P. sibirica ?????????? ? ??????? ???????, ?
???????? ???????????, ??????? ?????????
????????? ?? 20-? ??? ????? ??????????
???????? (????????, 1958; 1968).
??????????? ??????????? ?????? ?
??????? ?????? ?? ?????? ??? ??????? ????? ????????. ??????? ???????????? ???????
???????? ?????? ??????? ???????? ??????
Pinus sibirica ? Pinus cembra ?? ??????????
???? ?????? P. sylvestris. ????? ??????? ?????? ??? ????? ?????? ?? ??????: ????? ??
????????????? ??????????????? ???????????? ????????? ????????, ??????????????
????? ?????????? ??????? ?????? ? ??????, ????????????? ?? ???????????????? ?
????????? ???????????? ?????????? ?????
??????? ???????? ??????????? ??????????
???????? ?????, ??????? ????? ????????
?????????? ?? ????????? ?????? ???, ? ????? ? ?????? ????????.
??? ?????????????? ??????? ???? ???????????? ???????? ??????? ????? ?????????? ???? ?????????? ? ???????? ?????????????? (???????????? ????) P. sibirica (??) ?
??????????? ?? ????????? ??????? P. sibirica
(??), ? ????? ????? ???????????? ???????????
????????? (?? ?????-??????????? ???????
???????) P. cembra. ?????? ?? ???? ?????
???????????? ????? ?????????.
? ???????? ??????????? ???????? ????
???????????? ?????????? ??????? ?????
???????????? ?? ????????????? ?????????? ?
????????????????? ???????? ????? ????????
?????????, ?????????????? ????? ? ??????????? ??????????, ?.?. ? ??????? ????????????? ? ????????????? ???????? ? ?????????
?????????. ??????????? ?????? ????????
?? ?????? ????????. ?? ?? ???????? ? ???????? ???????? ??????? ????? ????????????,
??? ??? ?? ?????????? ??? ????????? ? ????-
????????? ? ??????
#5#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????????? ???? ???????? ???????? ????? (Pinaceae) ? ?????????
?? ????????????? ???????? ????????. ?????
???????????? ? ????????? ????????? ????
????????? ?????????? ???????. ??????? ??????? ???????? 43-45 ??? ? ???????? ???????? ?
???????????? ????????? ????? ????????????
? 35 ??? ? ??????????? ???????? ????? ??????????. ????????, ??? ??????????? ?????????
????? ?????????? ????? ???????, ?? ???????????? ??? ????????? ? ???? ??????? ???????? ? ??????? ??????????? ???????? ??????
?????????? ????????? ??? ????? ? ???? ???????????? ??????? ???????? ?? ???? ????????.
?????? ? ??????? ????????-????????? ?????????? ?? ????????????? ?????????? ??? ????
?????????? ??????: ? 1976 ?? 2010.
????? ????????? ?????? ? ?????? ?????????? ?????????? ??????? ?? ???? ???????? ??
?????????? 15 ?? ????? ? ???? ?? ???? ?????????. ?????????? ?????? ????????? ? ????????? ?????? ???????? ????? (???) ??????????? ???????? ???????? ??????? ? ?????????
(2000). ??? ???????? ? ????????? ?? 0.01 ??
? ??????? ????????? «Lintab 3.0».
? ?????? ?????????????? ???????, ???
???? ????? ????????????? ??????? ?? ?????????? ??????? ???????? ???????????????
??????? (???????, ???????????? ?????????,
??????????? ????????-????????? ??????? ?
??.), ?????????? ?????????? ???????? ?????? ???????? ? ??????? ??????-???????
(???????? ??????? ???????). ?????? ???????? ????? ??????????????? ??? ??????
?????????????? ?????????? ????? ???????
??????????? ???????? ??? ?? ??????????????????. ???????? ????????? ?????????????? ????????? ?????? ???????? ? Cook
et al. (1990). ?? ?????????????? ????????????? ???????????? ???????????? ???????????? ? ????????????????, ??????? ??????
???, ??????????? ??????????, ?????????
??????? ? ????, ???? ?????????, ????????????? ?????? ???????????, ? ??????????????
??????? ??????? (??????, 1986; ??????? ? ??.,
1996). ????????? ??????? ? ???? (SNR) ????????????? ????????? ????????????, ?????
??? ???? ?????????? (nR), ? ??????????????
????????????, ????????? ??? ?????? ?????????? (R), ? ??????????? ????????
SNR = n · R/(1 ? R),
??? n ? ????? ?????????????? ??????????
(Cook et al., 1990). ???????? ???????????? ?????????? ????????????? ????????? ?????????? ???????????? ????????. ???????????????? ? ????????????? ???????? ????????
????????? ?????? ???????? ????? ??? ??
????, ?????????, ? ????? ??????? ???? ????????? ????? ????????? ??????? ???????
????????. ?????????????? ?????????? ??????? ??????? ??????????? ????. ????????
??????????? ??????, ?????? ???????????????
???????? ??? ? ?????????????? ????????????? ??????????? ??? ?????? ?????????
ARSTAN (Holmes, 1983).
??? ?????? ?????????????? ???????
?????????????? ?????????????? ?????? ??
??????????? ? ??????? ????????? ???????????? (???. ????????????? ?????????????
????), ????????????? ? 20 ?? ?? ????? ?????
???????? ?? ?????? 164 ? ?.?.?. ?????? ??????????? ? ??????? 2001 ???? ?? ????????
2010 ??????????? (?????????? ???????????
?????? ????????????) ??????????? ??? ?????? ???????? ????????? ? ??????????????
?????? ???????????? ??????????, ????????????? ? 40 ?? ?? ????????????? ????. ??????
????????????? ?????????? ??? ? ???????????? ? ???????? (??? ???? ??????????????
??????????????? ???????? ?????????? ??????????, ? ??????? ??????? ??????????????
? ?????????? ?????), ? ????? ?????????????
?????? ????????? ? ????????? Statistica 8.0
(?????????, 2001).
#6#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????????? ???? ???????? ???????? ????? (Pinaceae) ? ?????????
? ?????? ?????? ??? ???????? ?????
?????? ??? ??????? ?????????????? ???????
???????? ????????. ???????? ?? ????, ???
?????? ?????? ????????? ?????? ?????? ???
?????? (Ind???) ???????? ? ???? ????????? ????????????:
Ind??? = Ind??? (??????) +
+ Ind??? (????????) + ?,
??? Ind??? (??????) ???????????? ????? ???????????? ??????? ???????? (???????????)
???????? ?? ????????????? ???????, Ind???
(????????) ??????????? ???????? ?????? ??
?????? ??? ????????, ? ? ????????? (?????????????) ??????????. ????????? ??????, ????
?? ? ???????????? Ind??? (????????) ????????????? ????????????? ????? ????????
???, ???? ????????? ??????????????? ?????????????? ????????? (????? ? ??., 2011).
?? ?????? ????? ??????????????? ????????
??? ???? ????????????? ? z-scores (???????? ?????????????) ??? ??????????? ??????? ???????? ? ????????? ????????? ?????.
????? ?? ??????????????? ?????????? ?????? ??? ?????? ?????????? ???????????????
??????????????? ?????????? ???????????
???????? (??? ?????? ? ?????????? ?? ?????,
??? ?????? ? ?? ?????). ? ? ?????????? ?????????? «??????????» ?????????? ????????? ?? ??????? ?????????? ? ??????????????
?????????, ??? ?? ??? ? ???????? ?????????
??????????.
???????? ????????????? ?????? ???
????????? ???????? ? ???????? ??? ?????
????????.
??????????
??? ???? ???????? ???????? ?????????? ?????????? ??????????? ???????? ?????? ??? ??????? ??? ??????????? ??? ?????
(???. 1). ? ????????? ???? ?????? ???????-
???? ??????. ?????? ???? ?? ?????????
????? ?????? ? ???????????, ??? ?????
???????? ? ?????? ????, ??? ??? ????????
??????????????? ?? ?????, ? ???????????
?????????. ??????? ? ??? ????? ???????
? ??????? ????????? ? ??????? ????? -0.05
? +1.1 ?? ? P. sibirica (??) ? P. cembra, ? ?
P. sibirica (??) ????? -0.05 ? +0.6 (???. 1). ?
??????? ??? ?????? ????????? ??? ?????? ?? 0.07 ?? ? P. sibirica (??), ?? 0.25
?? ? P. sibirica (??) ? ?? 0.27 ? P. cembra
(????. 1).
?????? ???????? ???????? ??? ??????
? ?????? ??????? ????????? ?? ?????. ???
?? ????? ??? ?????????????? ?? ???. 1 ?????? ?? ?????????? ???????? ????? ?????
?????, ?? ??????????? P. cembra ? P. sibirica
(??) (r=0.55, p<0.05). ???, ??????????? ?????????? ????? ??????? P. sibirica (??) ?
P. sibirica (??) ????? 0.28, ? ??????????????? ???????? ??? P. sibirica (??) ? P. cembra
?????????? 0.30.
? P. sibirica (??) ? P. cembra ??????
????????? ?????? ? ?????????? ???????? ?? ?????????? ????? ??????? ?????,
??? ? ?????????? ????????? ??????????
13 ? 11.6 % ??????????????. ? P. sibirica
(??) ??????????? ??????????? ???????
?? ???????? (???. 2). ?? ????????? ? ?????? ??????????? P. sibirica P. cembra ????? ??????????? ??????? ??? ??????, ???
? ??????. ??? ???? ? ??????? ??????????
??? ?????? P. cembra ?? 0.5 ??, ? ??????
?? 0.6 ?? ???? ??????????????? ??????????? ???????? P. sibirica (????. 1), ? ?????????? ??????? ??? ??????, ??? ? ??????
?????? ?? 23 %.
????????, ??? ??????????? ?????????
????? ?????????? ?????? ???????????? ????????? ????? ???????????? ?? 8-13 ???, ????????????? ???? ???????? ?? ??? ?? ????????? 15 ???, ?.?. ????? ??????? ???????? ?? ????
#7#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
??? ?????? - ??? ??????, ??
P. sibirica (??)
P. sibirica (??)
P.cembra
???. 1. ??????? (??) ? ?????? ????????? ?????? (???) ????? ??????? ? ??????? ? ?????????? ????????
P. sibirica ? P. cembra
??????? 1. ?????????????? ?????????????? ??????????? ???????? P. sibirica (?????????? ??????,
??????????) ? P. cembra
??????????????
1-?? ???????
?????????
??????? ? ????
???? ?????????,
??????????? 1-?
???????????, %
??????? ????????
???*, ??
P. sibirica (??)
???????????
????????????????
P. sibirica (??)
???????????
????????????, %
????????
???????????
??????????
?????????????? ??????????????
??????
0.14
77
0.13
0.22
5.4
49
2.30±0.44
??????
0.16
81
0.17
0.23
9.4
56
2.23±0.39
??????
0.14
86
0.17
0.36
11
64
2.37±0.55
??????
0.16
82
0.19
0.29
5.2
49
2.12±0.55
?????????
????????
??????
0.16
82
0.18
0.22
8.4
58
1.83±0.46
??????
0.19
82
0.20
0.19
8.2
55
1.56±0.37
????????
P. sibirica
0.16
77
0.14
0.20
8.8
41
1.20±0.62
????????
P. sylvestris
0.11
79
0.13
0.21
6.1
48
1.85±0.47
P. cembra
*??????? ???????? ??? ??????????? ???????? P. sibirica ?????????? ?? ????????? 15 ??? ?????, ????? ??? ????
????????? ? ?? ???? ????????????? ?????? (35 ???).
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????????? ???? ???????? ???????? ????? (Pinaceae) ? ?????????
?????????? ???????, ??
P. sibirica (??)
P. sibirica (??)
P. cembra
???. 2. ?????????? ??????? ?????? (???????? ?????) ? ?????? (?????????? ?????) ? ??????????
???????? P. sibirica (??) ? (??) ? P. cembra
???? ???????????. ?????????, ??? ? ??????? ??? ?????????? ???????? ????? ??????
??? ?????????? ???????? ????? ?? 0.65 ??
(????. 1). ?????? ??? ????????? ???????????
???????? ? ????????? ?????????? ??????????????? ?????????: ??????? ?????? ????????
????? (?? ??????????? P. sibirica (??)) ????????? ??????? ?????? ????? ???????????? ??
??? ?? ?????????? ??????????? ??? ?????
(???. 1, 2). ????????, ??? ??? ???????????????
? ???????? ??????? ??????????? ????????
???? ?? ?????.
?????????? ??????? ????? ????????????, ??????? ???????? ??????? ??? ????? ??????????, ????????? ??????? ????????????
????????? ?? 20 % ? P. sibirica (??) ? ?? 15 % ?
P. sibirica (??), ? ?????? 35 ??? ????? ?????????? ?????????? ???????? ???????????
(???. 3). ????? ????????????, ?????? ??????? ??? ????? ????????????, ?? ??? ??????????? ??????????? ??????? ? ??????? ????????
?????? 17 ??? ????? ??????????, ????? ????
??????????? ??????????????? ???????????
????? (???. 3).
?????????? ?????????????? ??????? ??????????, ??? ?????? ?????????????? ?
?????? ??? ?????? ????????????? ??????????? ???????? ? ???????? (???) ??????????? ???????? ? P. sibirica (??) (????????
?????? F=16.8, p<0.001) ? P. cembra (F=8.7,
p=0.003) ??? ????????????? ????????? 0.95. ?
P. sibirica (??) ??????????? ???????? ? ?????????? ???????? ????? ??????? ? ???????
(F=1.2, p=0.27). ???????? ?????? ????????? ??????, ????????????? ??????? ???????????????, ????? ????????????? ??????????
(F=182.4, p<0.001). ???????? ? ??? ?????
????????? ????????? ? ????????? ???????????? ????????? ????????????? ???????
(F=24.4 ? 57.6 ??? p<0.001 ??? P. sylvestris ?
P. sibirica ??????????????).
???????????????? ? ??????? ?????? ????? ???????????? ? ???? ????? ?????????
???????????????? ???????????? ?????????
#9#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????????? ???? ???????? ???????? ????? (Pinaceae) ? ?????????
???. 3. ????????? ?????????? ???????????? ??? ? P. sylvestris: 1 ? ?????? ??? P. sibirica, 2 ? ?????? ???
P. cembra, 3 ? ???????????? ?????????
????? ???????????? (????. 1), ??? ?? ??? ?
???????????????? ?????? ????? ?????????
???????????????? ???????????? ?????????
?????????, ?? ??????????? P. sibirica (??). ?
???? ????? ???????????????? ???????? ?????? ????, ??? ??????. ?????????? ?????????
?????????? ???????????????? ??? ??????,
??? ? ?????? ??? ????? ?????? ?????????
(???????) ?? ????????? ? ???????? (??????),
????? ??? ??????? ???????? ?????? ???????-
?? ?????????? ??? ????? (???????) ? ?????
(???????) ??? ?<0.05 ? ???? ????? (0.63-0.64)
????, ??? ????? ???????????? ?????????
????? ? ????? (0.55). ????? ?????? ??????????
????? ???? ??????????? ?????????????? ?????? ? ???????, ????????????? ?? ??????????
??????????????? ? ?????????? ?????. ?????????? ??????????????? ?????? ???????
????? ? ??????????? ???????? ??????????
???????? ?????? ? P. sibirica (??) (0.18), ? ??
?? ?????? ????? ???????? ?????????. ?????????????? 1-?? ??????? ????? ????????? ?
???? ??????????, ????? P. sibirica (??), ? ???????? ?????????? ??????? ????, ??? ? ?????? ?????????? (????. 1). ????????? ???????
? ???? (5-11), ?.?. ?????????? ???????????????
?????????????? ??????????, ? ???????????
?????????? (0.11?0.19) ??????????????? ? ???,
??? ?????????? ?????????? ???????? ?????
?????? ? ???????? ????????????? ???????.
???? ?????????, ??????????? ?????? ???????????, ????????? ?? 41 ?? 64 % ??? ?????????? ??????????.
??????????????? ????????-?????????
?????????? ????????? ?? ???. 4. ???? ? ?????
??? ????????, ??? ? ??????????? ????????
?????????? ??????? ????????????. ???????????? ?????????? ????? ??????????????-
????? ??? ? P. sibirica (??) ? P. cembra ???????????? ????? 0.55 ? 0.57 ??????????????. ????? ????? ?? ????? ??????????? ? ??????????????? ?????????? ??????? ?????
???????????? ? ?????????? ???????????
???????????? ????????? ????????? (r = 0.37;
0.51; 0.55 ? ??????????? ?? ??????????????
???????, ??????????? ? ????????, ????????? ? ????). ??????????? ???????? ???????
???????????? ????? ??????????????? ???????????? ? ?????? ?????? (77?86 %). ?????????????, ?????????????? ?????? ????????
?????? ???????????.
??????????????
??????
????????
??? ???????? ?????????? ? ????????????
? ???????? ??????, ??? ?? ?????????? ???????? ?????? ???????? ??????? ?????????
????????? ??????????? ?????? (???. 5). ???
# 10 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????????? ???? ???????? ???????? ????? (Pinaceae) ? ?????????
??????? ?????? ????????? ??????
P. sibirica (??)
P. sibiric? (??)
P. cembra
????????
???. 4. ??????????????? ????????-????????? ?????????? (???????? ????? ? ???? (??????), ?????????? ?
????? (??????)) ? P. sibirica ?????????? ?????????? (??) ? ?????? (??), P. cembra, ? ????? ???????????
???????? P. sibirica (???????? ?????) ? P. sylvestris (??????????)
????? ?????????? ? ?? ??????, ?????? ?? ???
????. ? ?????? ??????????????? ?????????? ?????????? ??????????? ??????????
??????????? ???????: ?????? ????? ?? ?????
???????? ?????????????? ?????????????
?? ? ????????????, ?? ? ????????, ?.?. ind???
(????????) ?? ???????? ????????????? ??????????. ? ?? ?? ????? ind??? (????????)
?????? ???????? ????? ???????????? ????????? ??????????. ??? ????????????? ??????? ??????????? ???????? ?? ???????????
??????? ???????? ??????? ??????????? ????
? ????????????? ?? ??????????? ??? (? ???
P. sibirica (??) ? ??????????) ???????? ????
?????. ? ???????? ?????????? ?????? ?????
?????????????? ?????? ????? ??????????
????????? ??????? ??????? ???????????,
?? ??? ????????? ??????????, ?????????????? ????? ????????????? ??????? ??????????? ????????, ???????? ?????? ????? ?
?????? ????????. ????? ???????, ????????
????? ????????? ??????????? ???????? ?????? ? ???????? ????????????? ??? ????? ?????????? ?????????? (???. 5).
??????????
????????????? ?????? ???????? ???????? ???????? ????? ?????????? ????????
????????????? «????????» ???????????
????? ?????? ? ?????? ? P. sibirica ??????-
# 11 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
P. sylvetris 2?
1
P. sibirica (??????????)
P. sibirica 2?
P. sibirica (??????)
???????????? ??????????
P. cembra
3?
P. sibirica (??????????)
3?
P. sibirica (??????)
P. cembra
??????
???. 5. ???????????? ?????????? ? ???????????? (?????) ? ???????? (???????) ????????
??? ??????????? ???????? ????? ? ????? (1); ??????????????? ???????? ?????????? (2A, 2?) ?
??????????????? ?????????? ?????????? (3?, 3?): ? ? ????? (??????), ? ? ???? (??????). * ? ????????????,
???????? ??? p<0,05. ????_?, ????_? ? ????????? ?? ?????? ??????????? ????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????????? ???? ???????? ???????? ????? (Pinaceae) ? ?????????
????? ?????????, ??? ??????????? ?????????????????? ? ?????????? ???????? ??
???? ???????????? ?????????? ??????
????????? ?????? ??????????? ???????? ? ?????????? ????? ????? ???????????
??????? ?? ????????. ???????????? ????
????? ??????? P. cembra ? P. sibirica ?????????????? ????????? ? ??????? ?????
???????????? ??????????? ???, ??? ? ???????????? ???????? ???? ????????? ? ????
??????????? ?????? ????????? ?????. ????
?? ????????????? ???????? (? ???????????
?? ???? ??????) ?????? ? ?????? ??????????? ????????????, ?? ?? ???????? ? ?????? ?????, ???????? ??? ????????????????
?????, ?????????? ???????? ??????????
?????, ? ?? ????? ??? ?????? ????? ??? ??????? ??????, ? ?????????? ???? ???????????
??????????????? ??????????? ????????.
?????? ???? ????? ??????????? ????????
???????? ????? ?? ?????? ?? ??????????????? (????????? ? ??., 2001).
?????? ??????????? ????????, ????
????????? ????????? ? ????? ????? ????????????, ???????? ?? ?? ???, ??? ????????,
P. sylvestris ????? ?????????????? ??? ??
????????? ? P. sibirica, ? ???? ??????????
?? ??????????? ???????? ?? ?? ????, ?????
??????? ???????? ?? ???? ??????????, ???
????????????. ????? ???? ?????? ??????????, ??? ?????????? ??????? ????????
???????? ????? ?????????? ???? ????????
??????????? (??????????) ????????. ?????
?????????? ?????? (????????, 1958; 1968;
????? ? ??., 2004) ????? ????????, ??? ?????? ????? ?????????? ?????? ???????, ???
???? ?????????, ???????????? ?? ?????.
??????? ???? ??????? ????? ?????????? ??????????? ???, ??? ???????? ??????? ??????
?????? ???????????? ???? ? ????? ??????.
??? ???????, ???????? ????? ????????????
(??????? ????, ???????????? ? ?????????-
???? ????????, ??????????????? ? ????????? ????????, ????????????????? ? ??????
???????? ?????????????), ?????? ?? ?????????? ??????? ??? ????? ?????????? ? ???
????? ???????????? ??? ??????????????
?????.
?????????? ??????? ???????? ?????
???????????? ?? ????????? ? ??????????
????? ?????????? ????? ???? ??????????
??????????????? ?????????????: ? ???????? ????????????? ???? P. cembra, ? ???????
?? P. sibirica, ????? ???????? ? ???? ?????????. ??????? ? ????? ?????? ????????? ?????
??????? ?????? ? ????????? ???? ?????????? ?? 8 ?? 20 ???? ? ??????????? ?? ???????? ??????? ????? (?????????, 2007). ?????
??????????? ? ????? ????????? ??????????? ?????????? ??????? ???????????????
????????????? ?????? ????? ?????: ???
?????????? ? ???? ????????? ??? P. sibirica
??????????? ????? ????????????? ?????????? ???? 5 є? ?????????? 1823?2015, ? ???
P. cembra ??????????? ?????? ? 2200?2350
(?????????????, 1982). ????? ????????????, ??? ??? ???? ??????????? ??????? ????????? ????? ?????? ????? ????????????
????? ?????????????, ?????????, ????????
????????? ?????????? ?????????, ?????? ??
???????? ???????????? ???????????? ?????
? ??? ??????, ????? «??????????» ?????????
?????????? ? ???? ?????????. ???????? ??
?? ??? ???? ??????????? ?? ??????????????
? ????? ? ???????? ?????-????????????????
??????? (??????? ??????? ???? ????? ????
????????? ?? ??????, ??? ??????? ?????????? ????? ?????? ?????? ? ????????????
?? ?????? 1500?2000 ? ??? ??????? ????),
????????????? ?????? ??????? P. cembra ?
P. sibirica ????? ??????.
??????????? ???? P. sylvestris, ?????????? ??????? ??? P. cembra, ????? ????
?????????? ??????? ???????? ??????. ??-
# 13 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????????? ???? ???????? ???????? ????? (Pinaceae) ? ?????????
???????? ? ?????? ??????????? ????????????????? ??????? ?????? ??? ????????? ??
??????. ??????????? ? ?????? ????????????
???????, ???????? ???????????? ?????????? ??? ???????????? ???????? ?????, ????????? ? ??????????? ?? ?????????????? ?????????? ??????.
???????? ????? ?????????? ?????????????? ? ??????????? ????????????? ??????????? ?? ????? ??????????? ?????????, ???
??????????????? ? ????????????? ???????
?????? P. sylvestris. ???????? ? ??????????
??????????? ???????? ???????? ??????
?????????? ????? ????????? ? ??????? ?????????? ????????. ??????????? ?????? ???????? ????? ???? ??????? ? ???????? ??????????? ???????? ???? ?? ?????, ??????????
???????? ???????? ?????????????? ?????
?????? ? ???????????? ?? ??????, ?????????? ? ??????? ?????????? ??????.
? ?????? ????????????? ???????????? ? ?????????????? ??????????? ???????? ?????? ? ?????? ?????????? ????????
?????????. ??????????????? ??????????
?????? ??????? ???????, ??????????? ??????? ? ??????????? (??????????) ????????
?????. ? «??????????» ?????????? ??????
????????????? ???????????? ????????? ?
????? ?? ??????? ????????????? ????????.
??????????????? ?????????? ?????? ??
????????????? ??????? ????? ?????? ??????????? ? ???????????? (???????????)
?????????, ? ?????????? ?????????? ??????????, ??? ? ???????? ??????????? ???????? ???? ???????? ????????????, ????????????? ???????? ??????. ??? ????????????
??????????????? ???????? ?????????????
???????? ??????????? ????? ??????????? ???? ? ????????????? ???????? ????????? ??????????? ? ??? ???????? ????
?????, ??? ?????????? ??? ?????????????
??????? ????????? ???????? ????? ??????-
??? ??????? ? ???? ???????? ??????? (????????, 2003).
??????????
?????? ????? ? ?????????? ???????????? ???????? ???????? ????? ??????????
(P. sibirica) ? ???????????? (P. cembra), ?????????????? ? ???????? ???????????? ?????????, ???????, ??? ?????????? ?????????
???? ??? ??????, ??? ? ?????? ?? ?????????
? ???????????? (???????????) ?????????
??????????????? ?????. ??? ???? ?????????? ??????? ??????? ????? ???????????? (P. sylvestris), ??????????? ? ????????
P. sibirica, ?????????? ?? 20 % ? ????????
??????????? ????????? ? ?? 15 % ? ??????????????. ?????? P. cembra, ???? ????? ??????? ?????????? ??????? (?? ????????? ?
???????? P. sibirica), ????????? ? ???????
??????? ????? ?? 10 %.
?????? P. sibirica ? P. cembra ?????????
?????? P. sylvestris ?? ???????? ???????????
?????, ??? ???????? ???????? ??????????????? ????? ???????????? ????????. ????????
????????????? ?????????? ???? ? ???????
????? ???????????? ??????????? ? ???????
P. sibirica ????????? ??????. ????? ???????
?????????? ??????? ????? ???????????? ??
????????? ? ?????? ????????? ??????????
????????????? ????????????? ??????? ????
? ?? ?????? ? ???????? ????????????? ????????.
????????????? ??????? ??????? ????? ???????????? ?????????? ???????
?????????? ????????, ??? ??????? ??????? ???????? ???????????? ??????????? ?
???????????? ??????. ? ??????? ?? ??????? ?????? ????? ????? ??????? ???????
?? ????????????? ???????. ??? ? ????????
??????????? ??????????? ?????????????
???????? ????????? ?????????, ? ????? ???????? ??????.
# 14 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????????? ???? ???????? ???????? ????? (Pinaceae) ? ?????????
?????????????
?????? ????????? ??? ????????? ????????????? ????????????? ??????? ????????? ???????????? ??????????? ? ?????
?? ? ???????????? ????????? ?? ??????????? «???????? ???????? ?????????? ?????? ?????» (?????????? ???????????????
???????????? ? ?????? ????????????? ????? ?????????? ???????????? ????????????,
?????? ? 1.7.09 «?????? ????????????????
?????????? ??????????? ???????? ????????? ????????? ? ????????? ????????
? ??????????? ????????? ????????», ???????????? ???????? ?.?. ???????), ? ?????
??????? ? ??-65610.2010.4 ? ???? ? 1104-00033-?. ?????? ???????? ????????????? ?.?. ????? ?? ??????? ? ????????? ?
?????????? ???????????.
?????? ??????????
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
????????? ?.?. (2001) STATISTICA: ????????? ??????? ?????? ?? ??????????: ??? ??????????????. ???.: ?????, 656 ?.
??????? ?.?., ?????? ?.?., ?????? ?.?. (1996) ??????????????????? ???????????? ?
?????-????????? ??????????. ???????????: ?????, 246 ?.
??? ?.?. (2011) ???????-?????????????? ?????????????? ????? ??????????: ???? ???????????? ex situ: ???????. ???. ? ????. ????. ????. ?????.
????????? ?.?. (2001) ??????????? ????? ? ???????? ???????? ????? ?? ???????????? ???????? ??????? ??????: ???????. ???. ? ????. ????. ????. ??????????.
????????? ?.?., ????? ?.?. (2004) ? ??????? ? ?????????? ??????????????? ????????
????? ?????????? ?? ????? ????????????. ??????? ?????. ???. ??-??. 10: 45-46.
????????? ?.?. (2003) ??????????? ? ???????? ????? ?????? ????? ?????????? ???????
????????????? ?? ????????? ? ???????????? ?????????. ??????????? 6: 42-48.
????????? ?.?. (2007) ???? ???????? ???????? ????????? ???????? ????? ? ????????????
?????????. ??????? ?????????? ???? 2-3: 217-224.
???????????? ????????????? ???????? ?????????????? ????? ? ???? (1982) ??? ???. ???????????????? ???????? ???? ?? ??????? ?????????. ?.: ?????? ??????????????, 368 ?.
????? ?.?., ???? ?., ??????? ?.?., ??????? ?.?. (2011) ????????? ?????????????? ???????
?? ?????????? ?? ????????-????????? ?????????? ? ?????? ?????????? ?????-?????????
???????. ??????????? 1: 28-37.
???????? ?.?., ??????????? ?.?. (2006) ????????????? ????? 18-??????? ????????? ????? ?????????? (???? 91/55) ? ??????????? ?? ???? ??????. ??????? ?????????? ???? 2:
220-222.
????? ?.?., ???????? ?.?., ??????? ?.?., ????????? ?.?. (2004) ??????? ???????? ?????
?????????? ?? ????????? ????????????? ????????. ?????? ????????? 5: 36-38.
???????? ?.?. (1958) ???????? ????? ?????????? ?? ?????. ???????? ???? ?? ???? 3-8.
???????? ?.?. (1968) ??????????? ???????? ??????? ?????. ?????? ????????? 2: 58-62.
???????? ?.?. (2003) ?????????? ????????? ??????? ? ?????????? ??????? ???????????
?? ?????? ??????? ?????? ? ??????-??????? ?????? ? ??????? ????????: ???????. ???. ?
????. ????. ????. ??????????.
????? ?.?. (2004) ????????????? ??????????????? ???????? ?????. ???????: ?????,
165 ?.
# 15 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????????? ???? ???????? ???????? ????? (Pinaceae) ? ?????????
16. ?????? ?.?. (1986) ???????????????? ??????? ??????? ???? ?? ?????. ?.: ?????, 137 ?.
17. ?????? ?.?., ??????? ?.?., ???????? ?.?. ? ??. (2000) ?????? ????????????????. ????? I.
?????? ????????????????. ???? ? ????????? ????????-????????? ??????????: ???????????. ???????. ??????????: ??????, 80 ?.
18. Cook E.R., Briffa K.R., Shiyatov S.G., Mazepa V.S. (1990) Tree- ring standartization and growthtrend estimation. Methods of Dendrochronology. Application in the Environmental Sciences.
Dordrecht, Netherlands: Kluwer, ?. 104-123.
19. Dyson W.G. (1975) A note on dwarfing of Pinus patula grafts. Silvae Genet. 24: 60-61.
20. Holmes R.L. (1983) Computer ? assisted quality control in tree-ring dating and measurement.
Tree-Ring Bulletin 44: 69-75.
21. Holzer K. (1970) Experiments in heteroplastic grafting of Pinus cembra L. Silvae Genet. 19:
164-170.
22. Schmidtling R.C., Ronald C. (1983) Rootstock influences flowering, growth, and survival of
loblolly pine grafts. For. Sci. 29(1): 117-124.
23. Schmidtling R.C. (1986) The effect of genetically related rootstock on survival, growth and
flowering in grafted slash pines. Proc. IUFRO conference, a Joint Meeting of Working Parties on
Breeding Theory, Progeny Testing and Seed Orchards. Williamsburg, Virginia, ?. 422-428.
Radial Growth
of Siberian Pine Heterografts (Pinaceae)
in Krasnoyarsk Forest-Steppe
Yulia A. Darikovaa, Eugene A. Vaganova,
Galina V. Kuznetsovab and Alexi M. Gracheva
a
Siberian Federal University,
79 Svobodny, Krasnoyarsk, 660041 Russia
b
V.N. Sukachev Institute
of Forest of the Siberian Branch
of the Russian Academy of Sciences,
50/28 Akademgorodok, Krasnoyarsk, 660036 Russia
Analysis of the radial growth and of the climatic response of heterografts of Siberian pine (Pinus
sibirica Du Tour) and Swiss pine (Pinus cembra L.) from three provenances was conducted. In all of
the cases the rootstock was Scots pine (Pinus sylvestris L.). Radial growth of the grafts of Siberian
pine was 23 % higher than for Swiss pine due to species peculiarities. The radial increment of the
rootstock of Scots pine depends on the type of scion: the diameter growth of P. sylvestris (rootstock
for Siberian pine) is greater than for the control trees by 20 % for the Surgut provenance and by
15 % for the Emelianovo provenance, whereas the diameter growth of Scots pine (rootstock for Swiss
pine) is 10 % less than for the control stand (ungrafted trees). This can be considered to be a result
of the scion influence. Analysis of climatic response of scion and rootstock shows that their reaction
# 16 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ?????????? ???? ???????? ???????? ????? (Pinaceae) ? ?????????
to air temperature is identical to the reaction of the control trees (positive correlation with April
temperature). However, when the component representing the climatic reaction of the control trees is
subtracted out, other results are obtained: the climatic response is absent for the rootstocks, whereas
the indexes of the growth of scion are positively correlated with the temperature of autumn period and
negatively correlated with the temperature of April-May.
Keywords: scion, rootstock, Pinus sibirica, Pinus cembra, Pinus sylvestris, tree ring width, climatic
signal.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2013 6) 18-31
~~~
??? 597.556.31
???????????? ? ??????????????? ??????
????????? ????????? ???????????? ????????????
Cottus poecilopus complex
(Scorpaeniformes: Cottidae)
?.?. ?????????*, ?.?. ??????????, ?.?. ???????
?
??????????????? ???????? ?? ???,
?????? 664033, ???????, ??. ????-?????????, 3
?
????????? ??????????????? ???????????,
?????? 664003, ???????, ??. ?. ??????, 1
Received 27.01.2012, received in revised form 19.12.2012, accepted 22.03.2013
????? ?????? ???????? ?????? ????????????? ? ?????????????? ???????????? ???????????????
????????? ???????????? ???????????? Cottus poecilopus complex ? ???????? ????????? ????????
??????? ??? ????, ??? ? ???? ??? ????????? ???????????????? ??????? ???????????????
??????????? ????. ?????????? ?????????? ???????? ?????????? ?????????????? ????????
???????? ?? ????????????? ? ???????????? ????????? ? ??? ???????? ? ??????? ?????????,
??? ????????? ?? ?? ????????????????. ??????????? ??????????? ?? ???????? ?????
???????? ????????????? ? ????????????? ????????????? ????????? ????? ??????????? ?
????????-????????? ????????, ??? ????????? ???????? ? ?????????? ?????? ??? ??????? ??
??? ? ??????. ?????? ???????????? ?????? ?? ???????????? ????????????? ? ????????????
????????????? ????????? ??????????????? ? ??????? ?????? ????????????? ???????????????
????????? ? ?? ?????????????? ? ?????? ??????????????? ????. ?????? ????? ? ???, ???
????????????? ??????????? ?? ????????? ????? ?????? ????????????? ????? ????????:
C. p. altaicus Kaschenko, 1899, C. p. szanaga Dybowsky, 1869, C. p. kolymensis Sideleva et Goto, 2012
? C. p. volki (Taranetz, 1933), ??????????????? ?????? ??????? ? ????????? ????? ???????.
???????? ?????: Cottus poecilopus complex, ??????????????? ??????, ?????????, ??????.
????????
???????????? ???????????? ???????????? Cottus poecilopus Heckel, 1837, ?????????? ??? ?? ?????? ??????????? ????????????
*
??????????, ???????? ??, ??? ?? ????? ?????
???????? ? ??? ???? ??????????? ?????.
? ?????? ?????? ?????? ??????????? ?????
???????? ?????????: ?????? ???????? ????-
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: bakht@lin.irk.ru
# 18 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ???????????? ? ??????????????? ?????? ????????? ????????? ?????????????
???? ?????????? ???, ??????????? ? ????????? ?????, ?????, ????? ? ???????; ?????? ?
??????? ???????? ? ??????????? ????????
???????????? (????, 1949). ?? ????? ???????? ??? ??????? ???????????? ????????????
? ????????? ??? ???????? ?. ????, ??????? ?.
????? (???????? ? ??., 2006). ?? ?????? ????????? ????? ?????? ?? ???????? ????????
????????? ??? ???, ????? ? ????. ? ????????
?????? ???????? ??? ????? ??? ????????: ? ?.
?????? (????????????, 2000; ???????? ? ??.,
2006; ????????, ????, 2009), ?. ???????????
???????? (????????????, 2000;) ? ????????
????? ???????? (???????, ???????, 1976; ???????, 1988). ????? ?? ?????? ??? ???????????
??????????? ? ????????? ??? ??????????? ????? ?? ??????? (???????, 1940; ??????????,
1965; ?????? ? ??., 1999) ?? ??????? (????????, 1982), ??????? ????? ??? ??????? ?. ????
(???????, 1928; ????, 1949; ????????, 1972). ?
???????? ????????? ???? ??????? ? ?. ????
?? ???????? (????, 1949). ? ??????????? ????? ?????????????? ???????? ??????????? ??????????? ?? ?. ??????? (????????, 1982) ??
?????? ?? ??? ?????? ??????-????? (???????,
1933; ????, 1949) ?? ???. ??????? ? ?????? ?
?????????? ???????? ?. ???? ?? ???? ?????????? ???????? (????, 1949). ???? ????? ?
????? ???????? ????? ?. ??????? (??????,
??????, 2003).
??????? ????????????? ? ????????????? ?????? ???????????? ????????????
???????? ???????? ???????????? ? ???????????? ??????????? ??????????????? ????????????? ???????????, ??? ????? ????????? ? ??????????? ?????????????? ?
??? ??????????? (????????, 1982; ??????,
??????, 2003; ????????, ??????, 2004; ??????, ????????????, 2007; ????????, ????,
2009, 2012; Hol?ik, Pivni?ka, 1969; Freyhof et
al., 2005; Kottelat, 2006). ? ????????? ?????
???????? C. poecilopus ????????????? ??????
????????????? ? ?????????? ?????? ?????????. ? ????????? ????? ?????? ????????????
??? C. szanaga Dybowsky, 1869, ????????? ?
???????? ?. ???? ? ????? ????????? ????????, ? C. kuznetzovi Berg, 1903 ? ? ???????? ?.
????. ?????? ?? ???????? ?????? C. altaicus
Kaschenko, 1899 ?? ???????? ????? ? C. volki
(Taranetz, 1933) ?? ??? ?????? ????????, ????? ????????? ? ????? ????????. ?????????
??? ?????????? ???? ??????? ??? ????? ??? ?
????????? ??????????? C. kolymensis (????????, ????, 2012). ??? ???? ??????????????
????, ? ????? ?????????????? ?? ???????????? (Cottus sp.) ?????????????????? ???????????? ???????????? ? ???? ??????????????? ? ????????????? ???????? ?????????? ?
?????? ????? ??? ???????? poecilopus.
?????????? ???????, ?????????? ????
????????? ????????? ??????????? ?? ??????
?????, ??? ???????? ????????????? ???????????? ?? ???????????????? ????????? ??????
??????? ?????????? ????????????. ?????
????????? ?????? ???????? ?????? ????????????? ? ?????????????? ???????????? ??????????????? ????????? ???????????????
??????????? ???????????? ????????????,
??????????? ??????? ?????????? ???? ????????? ???????? ??????? ??? ????, ??? ? ????
??? ????????? ?? ???????????????? ???????.
????????? ? ??????
? ?????? ?????? ??????? ????????, ????????? ? ??????? ??????? ??????? 19872006 ??. ??? ??????????? ???????? ??????????? ?.?. ?????????? (????????? ? ??., 1981) ?
????????? ?????? ???????? ???? 10 ? 12 ??.
??????????? ??? ??????????? 4 %-??? ????????? ?????????????.
????? ???????????????? ?????? ??????? ????? ?????? 309 ???. ? ????????
???? ???? ??????? ???? ???????: ? ???????
???????, ? ?????? ??????????? ???????-
# 19 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ???????????? ? ??????????????? ?????? ????????? ????????? ?????????????
??? (54°00?10??.?. 105o42?50??.?.) ? 55 ???.; ?
??????? ??????? ?. ??????? (56°17?16??.?.
107°34?31??.?.) ? 61 ???.; ??. ???????? ? ??????
?. ???? (56°36?50??.?. 117°27?24??.?.) ? 30 ???;
??. ???????, ? ?????? ?. ??????? (57°08?54??.?.
119°28?13??.?.) ? 50 ???; ??. ????????, ? ?????? ?. ???? (56°59?57??.?. 119°33?21??.?.) ? 22
???. ? ???????? ???? ???? ? ???? ???????:
?. ???????, ??????? ?. ?????? (51°20?43??.?.
113°27?08??.?.) ? 33 ???.; ? ???????? ?. ??? ?
??? ???????: ????? ?. ??? (51°47?12??.?.
87°14?41??.?.) ? 30 ???. ? ?. ???????, ??????? ?.
??? (50°37?41??.?. 87°54?17??.?.) ? 28 ???.
????????????? ???????????? ?????????
?? ???? ????????????? ? 28 ????????????
?????????: ????? ????? ? ?????? (D1) ? ?????? (D2) ???????, ??????? (P) ? ????????
(A) ????????? ? ???????? ??????? (sp.br.);
????? ?????? (?), ????? (L), ?????? (H) ? ?????? ???????? (B), ????? (pl) ? ?????? (h)
?????????? ??????, ??????????????? (aD),
??????????????? (pD), ????????????????
(aV), ????????????? (aA), ?????????????????? (PV) ? ??????????????? (VA) ???????????,
?????? ????????? 1-?? (lD1) ? 2-?? (lD2) ??????? ? ????????? (lA) ?????????, ?????? ?????????? ????? ? 1-? (hD1) ? 2-? (hD2) ???????, ???????? (hA), ??????? (lP) ? ???????
(lV) ?????????, ????? ???? (ao), ???????????
???????? ????? (o), ????????????? ?????????? (op), ?????? ? ??????? (hcz) ? ? ?????????
???????? ????? (hco), ?????????????? ?????????? (io), ?????? ??????? (lmx) ? ??????
(lmd) ????????.
????????????? ?????? ???????????????? ?????? ???????? ?? ??????????? ????????? (??????????, 1980) ? ??????????????
?????? ???????? Microsoft Excel 9.0. ????????? ?????? ??????? ??????? ?????????
(PCA) ???????? ? ?????????????? ??????
???????? SPSS 12.0 (SPSS Inc.). ??? ???????????? ???????????????? ???????? ?????-
???? ??????????? CD (????, 1971). ??????
??????????? ???????????? ????? ?????????
???????? ??????? ? ????? ??????? ???????????? ?????????? ???????? ? ???? ???????????? ???????? ? ?????????? ????????
?????? ?????????????? ???????? ???????????? ???????? ? ???? ??????????? ????????????? (??????????). ?????????????? ????????
????????? ??? ????????? ???????? ???????
???????? CD > 1.28, ??????????????? ???????? ?????? ?????????????? ???????? ?????
90 % (????, 1971).
??????????
???????????????? ???????? ???????????? ???????????? ????????????? ????????
???????????? ? ????. 1. ????????? ??????
(PCA) ???????, ??? ?????? ? ?????? ???????
?????????? ????????? 55,5 % ????? ????????? ????????????? ? 34,6 % ????????? ???????????? ?????????. ?? ???. 1 ????????????
????????????? ????????? ?? ?????????????
???????? ? ???????????? ?????? ???? ??????? ????????? ?? ???????????? ?????????.
?????????? ????????????? ????????
?? ?????? ??????? ?????????? ???? ?????
?????? ? ??????????????? ??????????, ? ????????????? ? ?????????????? ??????????
? ????? ?????????? ??????. ?? ?????? ??????? ??????????: ????????????? ? ??????
??????? ???????? ???????? ? ??????????????
??????????, ????????????? ? ?????? ?????????? ?????? ? ?????? ? ???????. ?? ????????????? ????????? ?? ??????????? ??????
??????? ???????? ???????? ???????? ???????????????? ???????? ?????????????? ????????? ?????? (????. 2). ? ?????? ??????? ??
?. ??? ? ??. ??????? ?????????? ?????? ?? ?????? ?????????? ??????, ? ? ??? ??? ??????? ?
??????? ? ?? ????? ??????. ?? ??????????
????????? ????? ?????????? ?????????????? ???????? ??????? ????????? ??????? ??
# 20 #
aD
B
h
H
L
c
sp.br.
A
P
D2
D1
SL, ??
TL, ??
1
9.0 ± 0.08
8-10; 0.65
17.7 ± 0.10
16-19; 0.80
14.1 ± 0.07
13-15; 0.55
14.5 ± 0.10
12-16; 0.80
5.7 ± 0.09
4-7; 0.71
29.8 ± 0.17
26.3-33.6; 1.30
71.7 ± 0.26
65.3-77.1; 2.01
20.1 ± 0.15
17.0-22.5; 1.15
6.8 ± 0.06
5.9-8.3; 0.44
18.1 ± 0.18
14.7-20.8; 1.37
33.5 ± 0.16
30.6-36.9; 1.24
9.0 ± 0.05
8-10; 0.40
18.5 ± 0.09
17-20; 0.68
14.0 ± 0.05
13-15; 0.40
14.7 ± 0.08
13-16; 0.58
5.5 ± 0.08
4-7; 0.63
29.7 ± 0.17
27.3-32.5; 1.23
73.3 ± 0.18
70.5-77.2; 1.30
19.9 ± 0.13
17.4-22.0; 0.98
6.1 ± 0.07
5.1-7.7; 0.55
17.6 ± 0.14
15.1-20.7; 1.06
32.6 ± 0.17
29.6-35.5; 1.27
81.8
74.9-94.8
67.7
62.4-78.5
??. ???????
(n=50)
5
9.5 ± 0.11
9-10; 0.50
18.8 ± 0.13
18-20; 0.60
14.0 ± 0.08
13-15; 0.37
14.6 ± 0.10
14-15; 0.49
5.3 ± 0.09
5-6; 0.45
27.9 ± 0.22
25.8-30.1; 1.20
75.7 ± 0.42
70.8-79.7; 2.33
20.5 ± 0.44
15.5-25.2; 2.44
5.7 ± 0.08
4.8-6.8; 0.45
17.2 ± 0.51
13.2-23.0; 2.84
32.5 ± 0.35
27.1-36.6; 1.96
28.6 ± 0.13
26.7-30.7; 0.94
74.7 ± 0.23
70.9-77.5; 1.62
16.7 ± 0.18
14.2-19.7; 1.26
5.7 ± 0.04
4.9-6.2; 0.31
14.9 ± 0.12
12.9-16.7; 0.86
33.5 ± 0.17
30.9-37.1; 1.22
27.7 ± 0.18
26.1-28.9; 0.83
74.4 ± 0.35
69.3-76.7; 1.65
17.1 ± 0.23
15.0-18.6; 1.09
5.6 ± 0.10
4.9-6.8; 0.46
16.2 ± 0.24
14.5-18.4; 1.12
31.3 ± 0.17
29.6-32.9; 0.82
???????????? ????????, % ?? ??????????? ????? (SL)
8.8 ± 0.09
7-10; 0.65
17.8 ± 0.11
17-19; 0.76
14.0 ± 0.04
13-15; 0.32
13.1 ± 0.08
12-14; 0.54
5.9 ± 0.11
4-7; 0.75
90.3
81.7-118.0
75.5
67.9-98.8
??. ????????
(n=22)
6
????????????? ????????
65.9
54.8-84.5
54.4
45.8-70.6
83.2
62.9-116.2
69.8
52.9-99.0
83.2
68.0-99.3
68.7
55.5-83.0
9.1 ± 0.12
8-11; 0.64
18.0 ± 0.14
16-20; 0.78
14.1 ± 0.16
12-15; 0.88
13.6 ± 0.16
12-15; 0.90
5.7 ± 0.12
4-6; 0.64
??. ????????
(n=30)
4
??????? ?. ????
?. ???????
(n=61)
3
?. ????
(n=55)
2
??????? 1. ???????????????? ???????? ???????????? ???????????? C. poecilopus
26.8 ± 0.20
23.5-28.0; 1.09
77.4 ± 0.50
72.4-87.5; 2.75
18.9 ± 0.22
16.7-20.8; 1.22
7.7 ± 0.11
6.8-9.2; 0.58
16.3 ± 0.20
14.6-18.8; 1.09
30.5 ± 0.25
27.1-32.3; 1.39
8.7 ± 0.10
8-10; 0.57
18.1 ± 0.13
16-19; 0.70
13.7 ± 0.09
13-15; 0.51
14.0 ± 0.14
13-15; 0.75
4.6 ± 0.13
3-6; 0.71
92.5
65.5-125.0
77.2
53.2-106.0
26.0 ± 0.16
24.5-27.3; 0.87
77.2 ± 0.30
73.6-80.9; 1.58
18.5 ± 0.21
16.2-20.9; 1.10
7.2 ± 0.09
6.4-8.0; 0.47
15.2 ± 0.19
13.2-18.1; 1.00
30.1 ± 0.15
29.0-31.9; 0.77
8.9 ± 0.06
8-9; 0.31
18.6 ± 0.09
18-19; 0.48
13.7 ± 0.09
13-14; 0.45
14.1 ± 0.11
13-15; 0.56
4.8 ± 0.12
4-6; 0.62
75.2
60.0-97.0
62.7
49.8-82.5
?. ???????
(n=28)
8
?????? ?. ???
?. ???
(n=30)
7
30.9 ± 0.25
28.1-35.0; 1.42
72.0 ± 0.32
67.3-74.3; 1.82
20.0 ± 0.19
17.6-22.0; 1.08
6.3 ± 0.07
5.7-7.4; 0.42
18.3 ± 0.21
15.2-20.9; 1.20
33.9 ± 0.29
30.8-38.5; 1.65
8.6 ± 0.10
7-9; 0.55
17.8 ± 0.08
17-19; 0.43
13.9 ± 0.05
13-14; 0.29
13.7 ± 0.09
12-14; 0.52
5.4 ± 0.10
4-6; 0.55
68.5
55.4-98.4
57.6
46.3-83.2
??????? ?. ????
?. ???????
(n=33)
9
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
ao
lV
lP
hA
lA
hD2
hD1
lD2
lD1
VA
PV
pl
aA
aV
pD
1
3
9.3 ± 0.14
6.9-11.9; 1.07
30.5 ± 0.34
24.0-35.9; 2.64
55.9 ± 0.27
51.4-60.9; 2.10
15.1 ± 0.21
11.7-18.1; 1.61
6.3 ± 0.13
4.0-10.8; 0.99
27.1 ± 0.29
21.0-31.7; 2.24
21.2 ± 0.27
15.8-26.0; 2.10
37.6 ± 0.25
33.2-42.6; 1.97
8.1 ± 0.12
5.4-10.1; 0.94
11.9 ± 0.15
8.5-15.0; 1.19
30.0 ± 0.23
25.0-33.2; 1.79
13.3 ± 0.16
9.9-15.8; 1.27
32.2 ± 0.30
25.2-36.5; 2.37
24.0 ± 0.30
20.5-34.2; 2.37
31.5 ± 0.30
24.4-38.6; 2.36
2
11.0 ± 0.13
9.1-13.7; 0.96
29.1 ± 0.24
24.5-32.8; 1.79
55.6 ± 0.16
53.2-58.3; 1.17
15.2 ± 0.15
12.1-18.6; 1.15
6.0 ± 0.11
4.4-8.2; 0.79
27.2 ± 0.24
23.1-31.5; 1.81
20.3 ± 0.19
17.2-24.6; 1.38
37.1 ± 0.23
32.9-42.1; 1.72
9.3 ± 0.11
7.6-11.6; 0.81
12.6 ± 0.11
10.7±15.0; 0.82
29.9 ± 0.18
27.3-33.6; 1.34
13.3 ± 0.17
10.5-15.8; 1.25
29.5 ± 0.19
25.5-32.8; 1.41
25.6 ± 0.27
22.2-29.0; 1.98
29.0 ± 0.30
24.9-33.8; 2.24
??????????? ????. 1
11.8 ± 0.16
9.5-14.3; 1.15
27.3 ± 0.32
24.1-32.9; 2.27
56.2 ± 0.21
52.5-59.0; 1.45
16.7 ± 0.12
15.3-18.9; 0.86
5.0 ± 0.09
3.4-6.6; 0.65
29.2 ± 0.27
24.7-35.0; 1.94
19.7 ± 0.30
13.3-25.4; 2.14
36.7 ± 0.27
32.0-42.9; 1.94
9.0 ± 0.12
6.8-10.9; 0.87
13.9 ± 0.15
9.3-17.0; 1.08
27.0 ± 0.22
22.4-31.5; 1.59
13.0 ± 0.18
9.8-16.0; 1.27
29.1 ± 0.20
26.1-32.3; 1.44
25.0 ± 0.25
21.6-28.3; 1.77
5
12.2 ± 0.19
10.3-13.6; 0.89
27.0 ± 0.44
24.0-30.8; 2.07
54.0 ± 0.27
51.4-56.6; 1.26
17.1 ± 0.16
16.3-19.3; 0.74
4.7 ± 0.10
3.8-5.8; 0.47
27.3 ± 0.46
23.2-31.7; 2.16
19.3 ± 0.31
16.4-22.1; 1.45
37.3 ± 0.37
33.3-40.0; 1.75
8.7 ± 0.17
7.2-10.5; 0.78
13.2 ± 0.20
11.6-14.8; 0.92
29.2 ± 0.25
26.5-31.5; 1.19
12.1 ± 0.22
10.0-14.1; 1.02
27.1 ± 0.19
24.3-28.7; 0.91
22.6 ± 0.37
19.2-25.9; 1.75
6
29.2 ± 0.53
23.3-35.7; 2.95
27.3 ± 0.20
23.3-29.9; 1.40
28.5 ± 0.35
25.3-31.2; 1.65
???????????? ????????, % ?? ????? ?????? (?)
12.5 ± 0.25
10.3-15.3; 1.42
25.6 ± 0.35
21.2-29.1; 1.94
53.0 ± 0.33
47.1-55.9; 1.86
17.3 ± 0.27
14.1-20.1; 1.48
5.4 ± 0.17
3.5-7.0; 0.92
28.5 ± 0.44
22.9-33.7; 2.44
19.9 ± 0.43
15.3-24.1; 2.39
38.0 ± 0.31
34.9-42.8; 1.73
8.5 ± 0.17
6.5-10.9; 0.96
12.7 ± 0.25
10.0-16.5; 1.37
29.8 ± 0.32
24.6-32.4; 1.79
11.1 ± 0.20
7.4-13.3; 1.10
27.9 ± 0.25
25.3-31.4; 1.40
23.0 ± 0.43
18.6-27.6; 2.40
4
28.1 ± 0.35
24.2-31.8; 1.91
11.9 ± 0.23
9.8-16.2; 1.26
26.0 ± 0.33
22.2-31.1; 1.81
55.0 ± 0.34
50.4-59.3; 1.84
16.8 ± 0.19
14.6-19.0; 1.04
5.8 ± 0.13
4.1-7.0; 0.69
29.3 ± 0.39
25.4-34.0; 2.11
21.6 ± 0.31
16.3-24.3; 1.71
37.1 ± 0.40
29.3-43.4; 2.21
8.4 ± 0.19
6.1-10.4; 1.02
12.3 ± 0.19
10.2-14.3; 1.05
28.0 ± 0.32
24.4-32.0; 1.74
12.9 ± 0.24
10.2-16.2; 1.29
29.3 ± 0.34
24.4-32.3; 1.85
25.5 ± 0.49
19.0-30.0; 2.67
7
28.6 ± 0.21
26.7-31.0; 1.08
12.5 ± 0.17
11.0-15.2; 0.90
25.5 ± 0.46
21.7-30.0; 2.42
52.6 ± 0.33
48.7-56.0; 1.73
17.8 ± 0.18
15.2-19.7; 0.97
5.1 ± 0.14
3.8-6.8; 0.74
27.2 ± 0.33
23.9-30.2; 1.74
22.5 ± 0.25
20.1-25.1; 1.31
37.2 ± 0.24
33.8-39.1; 1.26
8.9 ± 0.13
7.5-10.9; 0.70
12.2 ± 0.16
10.5-14.2; 0.83
28.3 ± 0.28
25.2-31.9; 1.48
11.3 ± 0.20
9.5-13.8; 1.03
28.7 ± 0.26
26.1-31.8; 1.39
24.1 ± 0.40
20.7-29.0; 2.14
8
27.8 ± 0.36
24.4-32.7; 2.06
9.4 ± 0.23
7.1-13.8; 1.32
29.9 ± 0.46
24.9-35.2; 2.66
55.6 ± 0.34
51.0-58.4; 1.92
16.3 ± 0.22
13.3-18.6; 1.26
6.4 ± 0.13
5.2-8.0; 0.72
25.8 ± 0.37
21.2-29.2; 2.12
22.4 ± 0.32
17.5-25.1; 1.81
39.7 ± 0.28
36.2-42.3; 1.62
8.3 ± 0.18
5.9-10.5; 1.02
13.0 ± 0.19
10.4-16.2; 1.11
30.0 ± 0.25
27.0-33.2; 1.45
12.4 ± 0.23
9.7-15.1; 1.30
32.2 ± 0.53
23.2-39.1; 3.05
23.1 ± 0.53
18.6-32.6; 3.05
9
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
3
23.4 ± 0.29
19.3-29.9; 2.28
49.4 ± 0.36
41.1-55.9; 2.79
63.2 ± 0.53
52.4-74.5; 4.16
50.2 ± 0.48
41.7-60.0; 3.75
11.3 ± 0.19
8.4-15.1; 1.50
41.9 ± 0.44
34.3-50.5; 3.44
49.2 ± 0.44
42.4-58.2; 3.40
2
23.8 ± 0.24
20.0-28.9; 1.76
49.2 ± 0.31
42.9-54.1; 2.33
59.0 ± 0.46
49.5-65.0; 3.38
47.9 ± 0.51
40.4-56.0; 3.81
10.7 ± 0.23
7.2-15.2; 1.74
40.5 ± 0.36
35.4-46.7; 2.64
48.6 ± 0.25
43.8-52.2; 1.85
4
28.6 ± 0.57
20.3-33.6; 3.19
47.7 ± 0.45
42.0-51.6; 2.48
62.6 ± 0.71
53.4-70.0; 3.98
45.8 ± 0.70
39.5-54.4; 3.92
8.7 ± 0.33
4.7-12.3; 1.83
40.3 ± 0.59
33.3-47.8; 3.27
48.0 ± 0.45
42.3-53.3; 2.53
5
26.6 ± 0.20
23.7-29.7; 1.41
47.5 ± 0.30
42.7-52.2; 2.10
54.6 ± 0.40
47.9-61.5; 2.83
44.7 ± 0.40
38.8-51.1; 2.82
8.4 ± 0.15
6.4-10.2; 1.04
42.2 ± 0.33
36.9-47.3; 2.33
48.3 ± 0.30
43.4-53.0; 2.12
6
24.0 ± 0.37
19.6-27.0; 1.75
47.7 ± 0.51
42.8-53.3; 2.40
55.1 ± 0.66
50.5-60.7; 3.09
44.0 ± 0.58
39.4-51.1; 2.72
7.7 ± 0.20
6.4-10.3; 0.93
43.0 ± 0.52
38.9-48.1; 2.46
50.5 ± 0.48
46.8-54.1; 2.24
7
22.8 ± 0.29
19.8-26.7; 1.60
49.1 ± 0.29
45.9-53.5; 1.58
63.3 ± 0.61
55.0-69.1; 3.35
47.5 ± 0.63
41.6-54.6; 3.45
7.0 ± 0.21
4.7-10.2; 1.18
39.1 ± 0.36
35.7-43.1; 2.00
45.0 ± 0.34
38.7-48.0; 1.86
8
25.0 ± 0.30
22.2-28.1; 1.56
47.0 ± 0.30
42.9-50.6; 1.60
62.6 ± 0.57
55.3-67.4; 3.03
45.2 ± 0.53
39.6-51.0; 2.78
6.7 ± 0.25
4.7-11.3; 1.33
37.5 ± 0.40
34.0-42.1; 2.09
45.0 ± 0.32
41.4-50.6; 1.70
9
24.1 ± 0.28
20.5-27.7; 1.63
49.2 ± 0.44
44.5-54.3; 2.55
57.5 ± 0.57
49.3-64.0; 3.30
42.0 ± 0.69
33.9-50.0; 3.95
10.3 ± 0.33
6.7-16.1; 1.91
39.6 ± 0.52
31.8-45.7; 2.98
47.9 ± 0.58
39.4-57.0; 3.31
??????????. ??????????? ????????? ? ??. ?????? «???????? ? ??????». TL ? ????? ?????, SL ? ??????????? ?????. ??? ?????? ? ??????? ???????? ? ??? ??????, ??? ?????? ?
??????? ???????????? ???????? ? ?????????????????? ??????????.
lmd
lmx
io
hco
hcz
op
o
1
??????????? ????. 1
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ???????????? ? ??????????????? ?????? ????????? ????????? ?????????????
3
1
2
3
4
5
2
6
7
8
PC 1
1
0
-1
-2
-3
-2
-1
0
1
2
3
PC 2
???. 1. ????????????? ??????? ???????????? ???????????? C. poecilopus ?? 28 ???????????? ?????????
? ???????????? ?????? ???? ??????? ?????????: 1 ? ?. ????; 2 ? ?. ???????; 3 ? ??. ????????; 4 ? ??.
???????; 5 ? ??. ????????; 6 ? ?. ???; 7 ? ?. ???????; 8 ? ?. ???????
??????? 2. ???????????????? ???????? ???????????? ???????????? C. poecilopus, ?? ???????
???????????????? ???????? ????? ???????? CD>1.28
??????
???????
??. ????????
?. ????
?. ??????? ??. ???????? ??. ??????? ??. ????????
H, B
H, h, B
??. ????????
c, H, h, io
H, h, pD,
lP, io
io
?. ???????
?. ???????
c, h
c*, L,
B, aD, pD
h
??. ???????
?. ???
?. ???
c, io
c, aD, io
c, L, aD,
pD, io
A
A
h
h*, o, hcz
h, hcz, lmd
h
c, h, aD, hcz
h, lmd
H, B
c, H, PV
?. ???????
* ? ????????, ? ????????????? ??????? ???? ??????.
??????. ?? ????????? ????????? ?? ???????????? ????????? ??????????? ??? ??????. ?????? ?????????? ??????? ?? ?. ????,
??????? ? ???????; ?????? ? ?? ?. ??? ? ???????; ?????? ? ?? ??. ???????? ? ???????.
??????????? ????????? ???????? ??????? ??
??. ????????, ??????? ??????????? ???????? ? ????????? ?? ???? ???? ????? ?, ?????
???????, ?????????????? ????????????????
????? (???. 1, ????. 2).
# 24 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ???????????? ? ??????????????? ?????? ????????? ????????? ?????????????
??????????
?????????? ?????????? ???????? ?????????? ?????????????? ???????? ???????? ??
????????????? ? ???????????? ?????????
????? ????????? ?? ?. ??????? ???????? ?.
????, ?. ????, ?. ??????? ? ??. ????????, ???
????????? ?? ???????????????? ?????????.
????????, ? ???? ?????? ????? ????????? ????????? ????? ???????? ? ??? ???????????
????? ?? ?????? ?? ???, ?? ???????? ??????? ? ??????????? ????????????? ?. ????
??????????? ????? (????????, 1972; ???????,
???????, 1976).
??????????? ??????????? ???????
???????? (???. 2?) ?????????? ?? ?????????
????????-????????? ??? (???. 2?, 2?) ?? ?????????????? ???????? ??????, ?????? ?????,
??????? ?????? ?????? ? ??????? ???????
?????????? ?????? (? ????????? ??????? ? ???
????????????), ? ????? ?????????? ???????
??????????. ??? ??????? ????????????? ????????? ?????? ?????????? ?????? ? ?????
?????? ??? ?????? ??????????????? ??????? (????????, 1982; ????????, ????, 2009;
McAllister, 1964; Freyhof et al., 2005 ? ??.).
?? ???. 3 ???????????? ?????????, ?????????????? ?????? ??????????? ? ??????????? ?
????????? ????????? «?????? poecilopus».
?? ??? ?????, ??? ??????? ????????????
??????? ???????? ? ???????? ?? ??? ???? ?
??????? ??????????? ??????? ???? ??????
?? ????????? ?????? ? ???????? ???????,
??????? ???????????? ????? ? ??????????
????????????. ??? ???? ??????? ?????? ??
?
?
?
???. 2. ??????????? ????????????: ?) C. poecilopus altaicus; ?. ???; ?) C. poecilopus szanaga, ?. ????;
?) C. poecilopus szanaga, ?. ?????
# 25 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ???????????? ? ??????????????? ?????? ????????? ????????? ?????????????
40,0
35,0
30,0
25,0
20,0
15,0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
???. 3. ????????? ???????????? ????????? ?????? ?????????? ?????? ? ????? ?????? ( % ?? ?????
??????) ? 14 ???????? ????????????? «?????? poecilopus» (?? ??????????? ???????? ????????).
??????????? ???????: C. p. szanaga: ??. ??????? ? 1; ??. ???????? ? 2; ??. ???????? ? 3; ?. ??????? ? 4;
?. ???? ? 5; C. p. kolymensis: ?. ?????? (????????, 1982) ? 6; ?. ???? (????????, 1982) ? 7; ?. ??????????
(????????, 1982) ? 8; ?. ??????? ? 9; C. volki: ???? ???????? (????????, 1982) ? 10; C. p. altaicus: ?.
??????? ? 11; ????? ?. ??? ? 12; ??????? ?. ?????? (???????, 1899) ? 13; Cottus p. poecilopus: ?. ???,
??????? ????? (????????, 1982) ?14. ???????????? ?????? ?????????? ??????? ???????????? ????????,
?????? ? ??? ??????? ????????
??????? ???????? ? ????????????? ????????? ? ???????? ??????????? ????????????? ?? ?????????? ???????????. ???????????
??????????? ?? ???????? ????? ?????????
???? ??????, ?, ????? ???????, ????????????
?????????? ???????? ????? ?????????? ????????. ???????? ?? ????????, ??? ????????? ????????? ???????? ????????????? ?
????????????? ????????????? ?????????
????? ??????????? ? ????????-?????????
?????????????, ????? ??????? ????? ? ?????????? ?????? ?? ??????? ?? ??? ? ??????.
??? ?????????????? ???????????? ????????,
??????????? ?.?. ??????? ? ???????????
? ???????? ?????????? ???????????? ????????????: «???? ????? ???????? ??????????
????? ??????????? C. poecilopus ? ????????????????? C. szanaga. ????? ????????, ???
???? ????????? ??? ? ? ????? ?????? ?????
????? ??????? ?????? ???????? C. poecilopus»
(???????, 1989. ?. 152).
??????????? ???????? ??????????? ?
???????????? ???????????? (?????????????)
?????????, ? ????????? ???????????? ???????? ??????????????? ??????? ? ??????? ???????????? ?????? ??????, ????? ???????????? ? ??????????????? ???????.
????????????? ?????? ? ??????????????? ?????????? ??????? ????? (c.l.l.) ?????
????????? ?? ????. ???, ??????? ????????
??? ??????????? ????????? (? ??? ????? ???
????????????? ???????) ??????? ?????????
??????? ????????????: 24-26 (????????, ????,
2009); 26-34 (????????, 1982); 14-37 (??????,
??????, 2003); 10-38 (Witkowski, 1984). ? ????????? ?????? ????????? ???????? ????????
????????? ??????????? ??????? ? ?????????
?? ?????? ???????????, ?? ? ????????? ??????
# 26 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ???????????? ? ??????????????? ?????? ????????? ????????? ?????????????
?????? ? 12-25 ? ????? ???????? ????? ???????? ??????? (????????, 1982), 15-19 ? ?. ???
(??????? ?. ????), 20-30 ? ???????? ?. ?????
(????????, ????, 2009), 20-28 ? ???????? ?. ?????? (???????, 1989), 22-23 ? ?. ???? (Hol?ik,
Pivni?ka, 1969), 24-27 ? ???????? ???????? ?
???????? ??????? ?. ???? (????????, ????,
2009) ? 19-28 ? ???????? ??????? ??????? ?.
????, ????? ???????? ????? ???????? ? ?.
??????? (??????, ??????, 2003). ? ???????????? ????? ??????? ????? ????? ????
?????? ??? ???????? ? ?????? ??? ?? 22 ?? 47
(????????, 1982; ??????, ??????, 2003). ?????????????, ?? ????? ???????? ?????? ?????? ?? ????? ?????????? ??????? ?? ??????
???? ???????????? ????????????.
?? ?? ???????? ? ?????????? ??? ?
????????????-??????????????
??????
(c.pm.). ????????????? ??? ????? ???? ????
??? ???, ? ??????????? ?? ????, ???????????
?????? ????? ? ?????? ?????? ? ?????????
????? ??? ???. ??? ???????????? ????? ?????????? ????????? ????????????? ????, ???
?????? ??????????? ????????????? ? ???????, ???? ???????????? ??????????? ?????
? ? ????????? ????? (????????, 1982; ????????, ????, 2009; Freyhof et al., 2005). ?????
???? ?????????? ?????? (pm5) ????? ?????
???? ??????? ??? ?????????. ? ?????? ?????????? «?????? poecilopus» ???????? ??????????? ????? ? ??? ??? ???? ?? ???????????. ?? ? ???? ???????????????????
???????????? ???? ??????? ?????? ???????
???????????????. ?.?. ???????? ? ?. ????
(2009), ????? ?????????????? ????? ????????
? C. kuznetsovi, ? ?????? ??????? ?????????,
??? pm5 ??????, ?????? ??????, ???? ???????? (????????, ????, 2009). ?.?. ????????
(1982) ?????????, ??? ? ?????? ?????? ??????
? ??? ????? ???? ?????? ???? ????? ???? ????????? (??????? ?. ?????), ????? ?????????
(??????? ?. ??????), ?????? ??????? (????
?????????????? ????????), ?????? ???????
(?. ????). ??? ????????? C. szanaga ?????? ????????????? ????????? ?????? ??????? pm5
(????????, ????, 2009) ??? ?????? ?????????
pm5 (??????, ??????, 2003). ? ????????????
????? ???? ??????? ?? ????? ????????, ?????? ?????????? ?????????? ???????? ??????
???? ? ????? ? ?????? ?????? ?????? (??????, ??????, 2003).
???? ?? ?????? ?????? ??????????
??? ???????????? ????? ? ?????????????
??????????? ? ?????????? ?????????? ????????????. ? ?????? ?????????? «??????
poecilopus» ?????? ??????? ?? ??????? (????????, 1982; ??????, ??????, 2003).
????? ???????? ???????????? ?????????????? ???????? ????????? ??????????????? ? ??????? ?????? ????????????? ??????????????? ?????????, ??? ????????? ??
?? ?????????????? ? ?????? ??????????????? ????. ? ?????? ?????????????? ??????? ? ?????????? ???????????-????????????
????????????. ???????????????? ?????????? (??????, ????????????, 2007; Kontula,
Vдinцlд, 2003; Yokoyama et al., 2008) ???????? ???????????????? «?????? poecilopus» ?
?????? ???? Cottus ? ?????? Cephalocottus,
??? ??????? ???????????? ?????? ???????
(????????, ??????, 2004). ? ?? ?????? ??????
?????????, ?????????? ?? ????? 8 ???????????? ?????, ????????????? ????????????
? ?????: 1) ?????? ????????; 2) ????????
????? ????????? ???? ? ???????? ???????
??????; 3) ????????? ????????, ?????? ????????, ??????? ? ???????? ??????? ?????;
4) ???????? ??????? ?????; 5) ???????? ?
???????? ??????? ????; 6) ?????; 7) ??????;
8) ????????????.
?????? ????? ????????????? ???????????? ?????. ????? ?? ???????????????
?????? ?? ?????????? ?????? ?????? ?????????, ??? ????????????? ? ?? ????????-
# 27 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ???????????? ? ??????????????? ?????? ????????? ????????? ?????????????
???????? ????? ???????? ????????? ????????? ? «?????? poecilopus». ?? ???????????
??????????????? ???????????? (????????,
1982; ??????, ??????, 2003; ????????, ????,
2009) ????????, ??? ??? ???????? ????? ????????????? ????????? (???? ?? ?????? ?????? ? ???????, ?? ?????????? ? ???????
?????????? ????????????-??????????????
????? ??????????????? ??????? ?, ??? ???????, ?????? ??????? ?????), ??????????
?? ?? ????????? ???????????. ?????????
??????????? ??????????? ? ??????????????
???????? ????????????? ????????? ?????
????????????? ????? ? ??????? ????????????????? ??????? «?????? poecilopus».
??????????
??????????? ???????????? ? ??????
???????????? ?????? ????????, ??? ???????????? «?????? poecilopus» ????????????
????? ??????????????? ????????????? ???????????, ? ??????? ??? ??????? ???????
????????? ????????? ?????????????? ?????????. ? ???? ??????? ???????? ?????????
?????????????? ????????? ??????, ??? ????????? ????????????? ????????? ?????????????? ??????? ? ????? ???????? ? ??? ??????, ????? ??????? ???? ?????? ?. ???? (1971),
?. ?. ????????????? ????????? ?????? ????,
?????????? ????? ???????? ?????? ? ?????????????? ???????? ?? ?????? ?????????
???? ?? ????.
?????? ?? ??????????????, ???????????
??????????? ???????? ?????? ???????????????? ?????????????? ?????, ????????-
????? ????????????? ?????? ????????? ???????? ????????????? ????????????? ? ?????
????????. ? ????????? ????? ?????? ?? ??????????? ?????????: ????????? ???????????
????????????? ? C. p. altaicus Kaschenko,
1899; ????????-????????? ??????????? ????????????? ? C. p. szanaga Dybowsky, 1869;
????????? ??????????? ????????????? ?
C. p. kolymensis Sideleva et Goto, 2012 ? ??????????? ????????????? ????? ? C. p. volki
(Taranetz, 1933). ????? ?????????? ???????
?????????? ???????? ???????? ??? ???, ????,
????????? ??????? ?. ????? ? ???????? ?
?????. ??????????? ??????????? ?. ??????
?????, ????????, ??????????? ? ???????
???????. ????? ????????-?????????? ??????? ???????? ????????? ???????? ??? ???? ?
????. ????????, ? ???? ?? ????????? ????????? ??? ??????????? ????? ?? ?????? ??
???. ????????? ?????? ???????? ???? ?????????? ????, ? ??????????? ???????????
????? ? ???? ?????? ????????.
?????????????
?????? ???????? ???????????????
?.?. ????????? (???????? ???????? ???????? ??. ?.?. ???????? ???), ?.?. ????????,
?.?. ?????????, ?.?. ?????? (????????? ??????????????? ???????????) ?? ??????? ? ????? ?????????. ?????? ????????? ??? ????????? ?????????? ????????? ???? (???????
98-04-49428-?, 99-04-63057-?, 02-04-63165-?),
??? «??????????» (?????? ?0788) ? ???????????? ????? ?????????? ?????????? (FCT
POCT1/33364/BSE/2000).
?????? ??????????
1.
2.
???? ?.?. (1949) ???? ??????? ??? ???? ? ???????????? ?????. ?. ? ?.: ???????????? ??
????. ?. 3. 452 ?.
???????? ?.?., ?????? ?.?. (2004) ??????? ???????????? ? ??? ??????? ? ???????????
??? ?????? ? ??????????????? ? ???????????????? ?????????????. ?.: ????????????
??????? ??????? ???, 389 ?.
# 28 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ???????????? ? ??????????????? ?????? ????????? ????????? ?????????????
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
??????? ?.?. (1928) ???? ???? ????. ??. ????. ?????. ?? ????. 9: 1?181.
???????????? ?.?. (2000) ???? ??????: ??????????. ???????????: ?????, 188 ?.
????????? ?.?., ???????? ?.?., ???????? ?.?., ?????????? ?.?. (1981) ???? ?????????
?????. ???????????: ?????, 150 ?.
??????? ?.?. (1899) ?????????? ????????? ????????????? ?????????? 1898 ?. ???????????. ?????: ???????????? ???????? ??-??, 158 ?.
???????? ?.?. (1972) ???? ??????. ?.: ?????, 360 ?.
??????? ?.?., ??????? ?.?. (1976) ?????-???????????? (Cottoidei) ?? ????? ????? ?????
????????. ????. ??????????. 16(3): 553?555.
??????? ?.?. (1940) ????? ?????? ? ??? ????????????????? ????????. ??. ??-?? ?????.
??????????, ?????????????? ? ???????. ???-??. ???. «???????. ???.-??». ???. 11. ????????????????? ???????? ?????????? ????, ?. 7?71.
???? ?. (1971) ???????? ????????????? ???????????. ?.: ???, 454 ?.
?????????? ?.?. (1965) ????? ?????????? ???? ??????. ????. ??????????. 5 (2):
261?278.
?????? ?.?., ?????????? ?.?., ???????? ?.?., ???????? ?.?., ???????? ?.?., ???????
?.?., ???????? ?.?., ?????????? ?.?., ?????? ?.?. (1999) ???????????? ??? ???????:
???????????, ???????? ????????? ????? ??? ?????? ??????????????? ? ??????? ???????,
??????????? ????????? ? ???????? ?????????????? ???????????. ?.: ?????, 207 ?.
?????????? ?.?. (1980) ????????? ?????????. ?.: ???-?? ???, 151 ?.
??????? ?.?. (1988) ??????????. ?: ??????? ?????????? ????????????. ?????.: ???-??
?????. ??-??, ?. 199?236.
15. ???????? ?.?. (1982) ??????????????? ??????? ? ???????? ??????????? ???????????????
???. ???????????: ?????, 147 ?.
16. ???????? ?.?., ?????? ?.?., ???????? ?.?., ??????? ?.?. (2006) ??????? ???????? ????????? ?????????????? ????????? ???. ????? ????????? ???? (Osteichtyes). ????? ???????????????? (Scorpaeniformes). ???????? Cottoidei. ????? II. ?: ???????????? ????? ?????.
?. 57 (65). ???.: ????. ??-?, ?. 225?349.
17. ???????? ?.?., ???? ?. (2009) ??????? ?????? ? ???????????? ???? ????? ?????? Cottus
poecilopus (Cottidae) ???????. ????. ??????????. 49 (5): 617?631.
18. ???????? ?.?., ???? ?. (2012) ????? ??? ???????????? Cottus kolymensis sp. nova
(Scorpaeniformes: Cottidae) ?? ??? ?????????? ????. ????. ??????????. 52 (3): 288?294.
19. ??????? ?.?. (1933) ? ????????? ????? ???????????? ????? ?? ???????????????? ????.
????. ?? ????. 2: 83?85.
20. ???????? ?.?. (1982) ??????????? ???? Cottus (Cottidae) ?? ???????? ???? ???? (??????????? ???????). ????. ??????????. ?. 22. ???. 1: 15?26.
21. ?????? ?.?., ?????? ?.?. (2003) ????? ?????? ?? ???????????? ?????????? ??? ???????? ??????? ??????. ?: ?????? ?????? ?.?. ??????????. ???. 2. ???????????: ?????????,
?. 320?333.
22. ?????? ?.?., ???????????? ?.?. (2007) ???????????????? ????? ???????????? ?????
Cottus Volki Taranetz, 1933 (Scorpaeniformes, Cottidae) ???????? ??????????? ??????? ???????????? ??????? ???????????????? ???. ????. ??????????. 47(1): 26?30.
# 29 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ???????????? ? ??????????????? ?????? ????????? ????????? ?????????????
23. Freyhof J., Kottelat M., Nolte A. (2005) Taxonomic diversity of European Cottus with description
of eight new species (Teleostei: Cottidae). Ichthyological Exploration of Freshwaters. 16 (2):
107?172.
24. Hol?ik J., Pivni?ka K. (1969) Notes on a collection of fish from Mongolia with description of
Microphysogobio tungtingensis anudarini ssp.n. and discovery of some new or little known fishes.
Annot. Zool. Botan. Bratislava. ? 56: 25.
25. Kottelat M. (2006) Fishes of Mongolia. A check-list of the fishes known to occur in Mongolia with
comments on systematics and nomenclature. The World Bank. Washington, DC. Fish Mongolia.
i-xi + 1-103.
26. McAllister D.E. (1964) Distinguishing characters for the sculpins Cottus bairdii and C. cognatus
in Eastern Canada. J. Fish. Res. Board Canada. 21(5): 1339?1342.
27. Kontula T., Vдinцlд R. (2003) Comparative phylogeography of freshwater cottid fishes: continentscale versus European affinities in the Cottus poecilopus and Cottus gobio complex. Acad.
dissertation. Helsinki: 1?15.
28. Witkowski A. (1984) Morphological variability in lake and river populations of Cottus poecilopus
Hekkel, 1836 (Pisces: Cottidae). Acta ichtiologica et Piscatoria. 14: 43?57.
29. Yokoyama R., Sideleva V.G., Shedko S.V., Goto A. (2008) Broad-scale phylogeography of the
Palearctic freshwater fish Cottus poecilopus complex (Pisces: Cottidae). Molecular Phylogenetics
and Evolution. 48: 1244?1251.
Variability and Taxonomic Status
of Siberian Population of Alpine Sculpin
Cottus Poecilopus Complex
(Scorpaeniformes: Cottidae)
Bakhtiar E. Bogdanova,
Tatiana V. Sverdlovab and Igor B. Knizhinb
a
Limnological Institute SB RAS,
3 Ulan-Batorskaya str., Irkutsk, 664033 Russia
b
Irkutsk State University,
1 Karl Marx str., Irkutsk, 664003 Russia
The main goal of investigation is the analysis of ecological and geographical variability of morphological
characters of alpine sculpin Cottus poecilopus complex in the Siberian rivers ? Lena, Ob, Amur for
identification of their taxonomic status. The data obtained have shown absence of taxonomically
significant differences in phenotypes of fishes in the Amur and Lena populations. The sculpin from Altai
reservoirs occupies geographically and phenotypically intermediate position between the European
and East Siberian population groups. It allows making conclusion about subspecies level of its
differences from those and others. The analysis of the literary data on molecular phylogeography and
characters variability is testify to high intergradations level of allopatric populations and belonging
# 30 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ???????????? ? ??????????????? ?????? ????????? ????????? ?????????????
to the same one species. It is possible to conclude that population groupings from the Asian part of a
general inhabiting area correspond to subspecies rank: C. p. altaicus Kaschenko, 1899, C. p. szanaga
Dybowsky, 1869, C. p. kolymensis Sideleva et Goto, 2012 and C. volki (Taranetz, 1933) and their
taxonomic status is overestimated.
Keywords: Cottus poecilopus complex, taxonomic status, population, subspecies.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2013 6) 32-43
~~~
??? 591.557.2.2
????? ????? ????????:
????? ??????????????
?.?. ????????*, ?.?. ??????????
?????? ??????-????????????????? ????????
????????-???????? ???????? ????????????????,
?????? 644080, ????, ??. ????, 7
?
????? ??????? ? ????????????? ?? ?????? ???????,
?????? 644116, ????, ??. ???????? 27-?, 42?
?
Received 27.12.2011, received in revised form 06.03.2013, accepted 22.03.2013
? ?????? ???????? ?????? ????????? ??????? ?. ????? ?? ??????????? ??????? ? ????????
?? ?????? ????? ? ?????? 2006-2010 ??. ?? ?????? ??????????, ??????????????? ????
«????? ??????? ? ????????????? ?? ?????? ???????». ????? ?? ????????? ?????? ? ?. ?????
???????????????? 11 849 ?????? ????? ????????. ??????? ?????????? ????????? 2901,3 ?????
? ???, ??? ????????????? 256,9 ?????? ?? 100 000 ???????. ?? ?????? ????????????, ? 99 %
??????? ???? ???????? ????????????? ??????, ?????? ? 66 % ??????? ???? ???????? ?????????
??????????? ????????, ????? ???????? ????? ????????? 0,36 %. ???????? ??????????
???????? ???????????: ???????????? ?????????? ????????? ?? ??????????? ??????? ?
???????? ?? ?????? ????? ?????????? ?? ?????? ?????? ? ? ??? ?? ?????? ????????????,
??????????? ? ? ?????? ??????, ? ?????? ?? ???????. ???? ???????? ?? ?????? ????? ????,
??? ????????, ??? ???? ???????????? ????? ??????? ?????? ? ??????? ?????? ? ??? ? 19-44 %.
???????? ????? ???? ????? ?????? ?? ???? ??? ?? ???? ? 87 % ?? ???? ??????. ?????? ??????
????, ??? ??????, ? 217,6 ? 304,0 ?????? ?? 100 000 ??????? ??????????????.
???????? ?????: ??????, ?????, ????????, ???????? ????????.
????????
?????? ???????? ?????? ?? ????? ?????????????? ???????? ???????? ? ??????
(????????, 2007). ??????????? ????????????? ????? ? ??? ??????????? ???????? ? 5258 ??? ?????? (American Veterinary?, 1997),
? ????????? ? ????? 3,9 ??? ?????? (McHarg
*
et al., 1995), ??????????? ????? ?? ???? ????,
?? ??????? ???????? ?????????, ?????????? ???????? 400 ??? (Coppinger, Coppinger,
2002). ????? ?????????? ?? ??????, ????????
? ??????????, ?????????, ? ???? ??????, ??
?????????, ???????????, ???????????? ?
????????? ????? ??????????, ??????????
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: makenovm@mail.ru
# 32 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????????. ????? ????? ????????: ????? ??????????????
????????. ????? ?????? ?????????????? ????
????? ????????? ?????? ?? ????? ????????????? ????????? ??????????? ????????,
?????? ??????? ? ?????????? ???????, ??????? ??????? ???? ?????????? ? ???????? ?
????????? ????? ?? ????????. ?? ?????????
???????, 1 ?????? ?? 20 ???? ?? ??? ?? ????
????? ?????? ???????? (Griego, 1995), ? ???
???????????? ??????????????? ????? ?????? ????? ??????? ???? ???????? ????????
(Presutti, 2001). ????????? ????? ???????????? ????, ???????? ? ????????????? ??????,
? ????? ???????? ? ? ????????? ???????, ??????? ????? ????? ???????? ?????? ????????
(Sacks et al., 2000; Ozanne-Smith et al., 2001).
????? ????, ???????? ????? ????? ???? ???????? ????????? ??????????????? ???????, ?
??????????? ??? ????????? ?? ????? ? ?????????? (Schalamon et al., 2006).
??????? ????????? ?????? ?? ???????? ????????????, ??? ?? ?????? ??????? ??
??????????? ?????, ??????? ?? ??????????,
???????????? ????????? ????? ? ????????
? ?.?. ???????? ?? ???????? ????????????
?????? ????????????, ??????????? ??????????? ????????, ???? ????????? ????? ??
????? ? ?????????? ??????? ???????? ???????????? ?????????.
? ???? ????? ???? ?????? ?????? ????
?????????????? ????????? ???????: ???????????????? ????? ???????? ????? ??
??????? ?????????????? ?????????, ????????? ? ??????? ?????????? ??????.
????????? ? ??????
???????????? ????????? ? ?. ????? ?
2010-2011 ??. ? ?????? ???? ???????????? ????????? ?? ?????? ??????? ?. ????? ? ????
2006 ?? ???? 2010 ?., ??????????????? ??????????? ????????? ??????????? ??????????????? «????? ??????? ? ????????????? ??
?????? ???????». ?????????? ???? ??????
???????? ? ???? ??? ?????? ????????? ??????? ? ?????????? ??????????????? ?. ?????
? ????????? ??????? ??????????? ??????
????? ?????? ????????. ? ???? ???????????
???, ??????? ? ????? ?????????? ?????????????, ???? ?????, ?????????????? ??????????
?????????, ??????????? ?????.
??? ???????? ??????? ?????? ?? ???????
????????? ????????????? ??????, ??? ? ???????? ??????? ???????????? ????? ????. ???
??????? ?????????? ???? ????????? ????????????????? ??????:
xt = tr t · st · ?t,
??? tr t ? ?????; st ? ???????? ??????????; ?t ?
????????? ??????????.
??? ????????? ?????? ???? ??????????? ???????? ???????????? ?????????? ????
?? ????????????????? ?????? ? ?????????????? ??????????? ???????? (?????? ?????????? ? 12 ???????) ? ??????????? ????????????? ????????.
??? ??????? ?????????????? ?????????
???????????? ???? ???????????? ???????? ???????????????? ?????? ???????????
?????? ??????????????? ?????????? ??
?????? ??????? ? ??????????? ???????? ?
???????? ????????? ?. ????? ?? ??????????
??????? ?? ??????????? ??????. ??? ????????? ??????? ? ??????? ????????? ?????? ????????????? ??? ????????? ???????.
?????????? ???? ???, ???????????? ??
?????? ?????, ????? ? ??????? ?-????????
?????? (??????? ??????????????) ????????? ????????????? ???????? ????? ??????????? ?????? ???? ???????. ???????????
??????? ??????????? ????????? ?? ?????????? ???????.
??? ????????? ? ??????? ?????? ????
???????????? ?????? ???????? Microsoft
Excel ? SPSS Statistics.
# 33 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????????. ????? ????? ????????: ????? ??????????????
??????????
?????? ?????
???? ???????????? ??????? ??????????
?????? ????????? ?? ????? ?? ????????? ?
???????? ? ?????????? ????????? ???????? ??????: ???????? ????????; ???????? ???????? ?????????; ???????? ???????? ???????????. ?????? «????????» ????????? ???
????? ??????????? ??????, ?????? «????????
?????????» ? ???? ???????????? ???? ??????,
?? ????????? ??? ??? ??? ?? ????????, ??????
«???????? ???????????» ? ???? ??????????
??????, ??? ??? ??????? ????????????? ??????,
?? ?? ?? ????, ??? ??? ????? ? ??? ?? ????????.
? ????? ? ??? ??? ????????? ????????? ??
2006 ? 2010 ??. ????????, ??? ??????? ????????????? ???????? ???? ???????????? ?????? ?
2007 ?? 2009 ??. ???????????? (????. 1). ?????
?? ????????? ?????? ???????????????? 11 849
?????? ????? ????????. ??????? ??????????
?????? ????????? 2901,3 ?????? ? ???, ??? ????????????? 256,9 ????? ?? 100 000 ???????.
??? ??????? ?????????? ? ????????,
???????? ????, ???? ????????, ??? ? ??????????? ???????, 99,6 %, ????? ????????
??????, ??????? ??????? («???????? ?????????» ? «???????? ???????????»); «????????», ?? ?????? ????????????, ?????? ?
0,3 % ??????? (????. 1).
?????????? ??????
??? ???????? ???????? ???????? ???????????? ????????? ????? ?? ????????? ?
???????? ?? ???????????? ??????? ??????????? ?????????? ?????????? ?????? ??
??????? ????. ??? ???????? ???? ????????
?????????? ?????? ???? ????????? ?? ?????? ??????: ????, ?????, ????, ????? (????. 2).
????????????? ?????? ?????????? ???????
??????????? ?????????? ?????? ?? ??????
(F = 35,22; p < 0,001).
??? ???? ????? ?????? ???????? ???????????, ?????? ?? ?????? ???? ????????????
? ???? ?????????? ???? (???. 1).
??????? 1. ?????? ????????? ?????, ?? ?????? ???????????? (n ? ?????????? ??????????; % ? ??
?????? ????? ?????? ?? ???)
?????????? ??????
???
?????
?????????
??????????
????????
n
%
?????????
????????????
????????
n
%
n
?????????
????????
???
??????
%
2006
(????-???.)
1918
1352
70,5
530
27,6
31
1,6
5
2007
3233
2174
67,2
1054
32,6
4
0,1
1
2008
2762
1977
71,6
781
28,3
3
0,1
1
2009
2709
1905
70,3
798
29,5
3
0,1
3
2010
(???.-????)
1227
766
62,4
459
37,4
1
0,1
1
11849
8174
69,0
3622
30,6
41
0,3
11
?
m ± SD*
??. ???-??*
?????? ??
100 ???.???.
2901,3±288,5
2018,7±139,3
877,7±152,9
3,3±0,6
256,9
178,7
77,7
0,3
*? ??????? ???????? ?????????? ?? ?????? 2007-2009 ??.
# 34 #
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??????? 2. ????????????? ?????????? ?????? ? ??????????? ?? ??????? ???? (N ? ?????????? ?????????
??????? ? ??????)
????? ????
N
???????
???????????
??????????
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????
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143,3
34,9
109
223
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12
272,9
83,8
161
408
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278
383
255
408
?????
12
196,4
43,6
114
?????
49
241,2
92,5
109
???. 1. ?????????? ????????? ??????? ?. ????? ?? ??????????? ??????? ? ???????? ?? ?????? ?????,
???? 2006 ?. ? ???? 2010 ?.
?? ??????? ????? ???????? ?????? ???????? ????????????? ? ??????????????
????????????? ?????. ? ??????? ???? ???????????? ?????????? ????????? ?? ??????????? ??????? ? ???????? ?? ?????? ?????
?????????? ?? ?????? ?????? ? ? ??? ?? ?????? ????????????, ??????????? ? ? ??????
??????, ? ?????? ?? ???????.
?????? ? ???????? ??????????? ???????
???????????? ? ?????. ?????? ???????????
????? ?????????? ???????? ?????????? ????
??????? ??????? ?????????????? ??????, ??????? ??? ??????????????? ???????? ??????? (???. 2):
y = 284,01 ? 1,76x; R 2 = 0,40;
??????????? ???????????? ? ????????????
????????? ????????? ????????????? ???????
(p < 0,001).
?????? ???????? ????????????? ??????
??????? ??????? ???????? ? ???????? ?????????? ?????? ? ?????? ? ?????? 2007 ?. ???????, ????????, ??????????? ???, ??? ?
???????????? ?????? ?????????? ????, ?????? ?????? (tr t) ? ???????? ????????????
(st), ???????????? ????? ??????????? ?????????? (ct) ? ?????????? ??????? ?????????????? ??????? ? ?????. ?????? ??? ??????
# 35 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????????. ????? ????? ????????: ????? ??????????????
???. 2. ????????? ??????? ?. ????? ?? ??????????? ??????? ? ???????? ?? ?????? ?????, ???? 2006 ?. ?
???? 2010 ?., ????? ???????? ???????????? ?????????? ???? (? ?????? ?????? y = 284,01 ? 1,76x)
??????????? ?????????? ?????????? ?????????????? ??????????, ? ????? ???????????? ???? ?????? ???? ?? ?????? ????, ???
?? ????????? ???????????????? ???????????
?????????.
?????????????? ?????????
????????????
???? ???????????, ??? ?????? ??????
?????? ????, ??? ??????: ? ??????? ?? ???
???? 304,0 ? 217,6 ????? ?? 100 000 ??????? ?????????????? (? = 9,0; p<0,01). ??? ???????????? ?? ?????????? ??????? (???. 3) ????????, ??? ?????? ?????? ????????? ??????????
????, ??? ???????, ? ???????? ?? 5 ?? 15 ???
(? ? 2,5; p<0,01) ? ?????? ? ???????? ?? 20 ??
40 (? ? 2,7; p<0,01) ? ?? 70 ?? 75 ??? (? = 1,7;
p<0,05) ????, ??? ?????? ???? ?? ????????.
? ????????? ?????????? ??????? ????????
???????? ????? ???????, ?????????? ???????? ? ????????, ?? ???????????. ? ?????
?? ????????? ???? ????? ???????? ?????? ????? ??????????? ???? ? ??????? ???? ? ???????? ?? 5 ?? 19 ??? (???. 3).
??????????? ??????
??? ???????????? ??????????? ??????
???? ???????????, ??? ? ??????????? ??????? ?????? ?????? ?? ?????????? ? 87 %
(???. 4). ???? ????? ????? ??????????? ? ??????? ????????, ??? ? ??????, ?????? ???????? ??????? ?? ????? ???????????? ?????
? ?????? ? ??? ?????????? ? ??????? ? 8,3 %
???????.
??? ???????? ??????????? ????? ? ??????????? ?? ???? ????????????? ???? ????????, ??? ? ???????, ?? ?????? 2007-2009 ??.,
?????? ?????? ?? ???? ????, ??? ??????,
? ?????? ?? ???? ?????? ????, ??? ??????
(p<0,05).
? ????????? ???????????? ??????????? ?????? ????? ???????????? ?????????
????????? (???. 4). ??? ????? ?????? 10 ???
?????????? ???????????? ??????? ???????
?????? ? ?????? ? ?? 44 ?? 19 % ? ? ??????
???? ? 8-10 %, ?? ???? ????? ?????? ???????????? ????? ? 16-40 %. ? ????? ?????? 65
??? ?????????? ???? ?????? ?? ???? ? 50 % ?
?????.
# 36 #
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?.?. ???????, ?.?. ?????????. ????? ????? ????????: ????? ??????????????
???. 3. ?????????????? ????????????? ???????????? ?? ?????? ?????, ??????? ???????? ?? 2007-2009 ??.
???. 4. ??????????? ?????? ? ??????????? ?? ???????? ????????????
??????????
?????????? ? ???? ???????????? ???????? ?????????? ?????? ???????? ? ?. ?????
???????? ???????????? ???????? ? ? ???????
2901,3 ????? ? ??? (256,9 ??????? ?? 100 000
???????). ????? ????????????? ??????????
????????????? ????????????, ???????????
???????? ?????? ????? ? ?? ??????, ??? ????? ?? ???????, ??????? ?? ???????? ?????????? ?????? ?????? ? ???????????? ?????????? ??????? (????. 3).
??? ?????, ???????? ??????? ??????????
?????? ???? ???????????????? ? ?. ?????????
? 1993 ?. ? 214 ??????? ?? 100 000 ???????
# 37 #
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?.?. ???????, ?.?. ?????????. ????? ????? ????????: ????? ??????????????
??????? 3. ?????? ? ?????????? ?????? ????? ???????? ? ?????? ???????
???????????
?????????, ???.
?????????? ??????
????? ?? 100 000 ???.
??????
????????????
????? (??????)
115 ???.
160
1986-1987 ??.
?????? (???????)
560 ???.
150
1984-1988 ??.
????????? (???)
313 ???.
214
589*
1993 ?.
Chang et al., 1997
??????? (??????)
380 ???.
58
2000-2002 ??.
Ostanello et al.,
2005
??????? (??????)
1 213 ???.
149
2003-2006 ??.
Vucinic, 2008
????? (??????)
??????
Szpakowski et al.,
1989
Rufino Gonzales,
1990
* ? ????????????????? ?????, ??????? ???? ?????????? ????? ????? ?????? ?????, ?? ???????????? ?? ???????????
???????.
(Chang et al., 1997). ? ??????? ?????, ??????
? ??????? ????????? ? ??????? ????? ??????????? ??????????? ???? ? ???????? 150160 ??????? ?? 100 000 ????????? (Szpakowski
et al., 1989; Rufino Gonzales, 1990; Vucinic,
2008).
?????????? ?????? ????? ??????? ??
?????? ??????, ????? ???????: ???????????
? ????????? ????????? ??????, ???????????
????????? ?????, ??????????? ?????????
?????????, ??????? ?????????? ????? ? ??.
?? ?????? ????? ???????????? ??????????
???????????????? ????? ??????? ????????
????? ??????, ?????????? ??? ?. ?????, ??
?????????????? ?????????, ????????? ?????????? ??????? ???????????? ????????, ???????????????? ??????????? ????????? ?????
? ??????? ? ?.?.
???? ?? ???????? ???????? ???????????,
?????????? ??? ??????? ??????, ??????????? ? ???, ??? ?у????? ????? ?????? ????
????????, ?? ?????? ????????????, ?????????????? ????????. ?????? ? 69 % ???????
???????????? ?? ?????? ? ???? ?????????
?????? ???????? ????? «???????? ?????????», ? ??? ??????, ??? ??? ???? ???? ????
??????????? ???? ??????, ???? ???????? ?
???????????, ?????????, ????????? ?????????? ? ???????? ??????. ? ????????? 30,6 %
??????? ???????????? ?????????? ????? ???
«???????? ???????????», ?????? ???????
?????? ????????, ???? ???? ???????? ????????, ?????????? ??????? ???????????? ??
????. ????? ????????? ????? ?????????? ???
????????????? ???????? ??????? ?? ???????
?? ????? ???????? ??? ??? ??? ??????????
????????? ??????, ????? ?????? ????? ????????, ??? ????????????? ?????????, ???????? ?????????? ?????????????.
??????????? ?????????? ????????? ?? ??,
??? ?????? ????????? ????? ?? ???????? ???????????? ???????? ? ???, ????? ???????
?????????? ??????????? ????????? ?? ?????
????????, ??? ??? ???????????, ????????????, ??????????, ?????????. ? ???????????
??????? ?????? ? ?????????? ?????????????.
???, ? ?????? ?????????? ???????, ????????? ????? ????? ? 1972-1973 ??. ? ?????????, ???? ???????????, ??? ? 5,9 % ???????
???????? ?????? ????? ??????????? ????????? ??? ????? ?? ????? (Beck et al., 1975). ?
64,8 % ?? ???? ????????????? ???????? ?????????? ???????????? ???????????????
# 38 #
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?.?. ???????, ?.?. ?????????. ????? ????? ????????: ????? ??????????????
????? ?????? ????????????????? ? ???????,
?? ???????? ??? ?????????????? ???????. ?
25,4 % ??????? ???????????? ????? ??????
????????????????? ? ???????. ???? ????? ???
???? ???? ? ???????, ????? ??????, ?????????? ????????? ????????, ??????????????
??????????, ? ????? ???????????? ????????? ?? ????? ??? ???? ??????, ?????????? ??
?? ????? ???, ???????? ????? ?? ????, ?? ??????? ? ??. (Beck et al.,1975).
?? ?????? ???? ? ??????????
(Weiss et al., 1998), ? ??? 58 % ???? ??????
????? ???? ????????, ???????? ? ???? ????. ?
???????????? ?????? ????? ?? 17 ??? ? ?. ????
(???????) ???? ????????, ??? 73 % ???? ?????? ???????? ??????, ??????? ????????????
????: ? 24 % ??????? ?????? ????????????
????? ????? ?????????????, ? 14 ? ????????????, ? 20 ? ?????, ? 15 ? ??????? ? ???? ?
15 ? ?????? ??????????? ?????? (Schalamon et
al., 2006).
??????? ????????, ??? ?????????? ?????????? ???????????? ????? ???????????
?????????? ?? ?????????? ???? ????? ??????????????? ? ????? ? ????????????? ??????????? ??????????, ????????? ? ???????????? ?????????? ?????, ????????? ? ??? ?
?.?. ? ??????????? ???? ???????????? ????
???????????, ??? ??????????? ?????? ?????? ?????, ?? ?????? ????????????, ???? ?
0,36 % ??????? ? ? ??????? 3,3 ????? ? ???. ???
?????????, ?? ?????? ?????????? ??????????????, ???? ??????, ?????????? ???????????? ????????, ??????????? ???? ? ??????????
10-14 %, ??? ?????????? ????? 200-350 ??????? ? ??? (Parrish et al., 1959; Beck et al., 1975).
? ???????? (2003-2006 ??.) ?????????? ??????,
?????????? ????????? ???????? (stray dogs),
?????????? ????????? ?????????? ??????
????????????? ????? ? 1507,5 ? 832,8 ????? ?
??? ??????????????, ??? ?????????? 95,6 ? 52,8
?????? ?? 100 000 ??????? (Vucinic, 2008).
????? ?????????????? ?????, ?????????? ???? ??? ?. ?????, ????? ???????????
????? ?????????: ????????????? ?????????
??????????? ????? ?? ????????? ? ????????
? ????????? ???????????? ????????? ??????????? ????? ? ????????? ????. ???????????
?????????? ??????????, ??? ???????????
?????? ?????? ????????, ????????? ????????
????????????????? ???????? ? ???????????
?????????. ??????????? ????????? ??? ????????? ? ???? ???, ??????? ? ????????????
??????, ????? ? ???????? ? ??????? ????????, ?????? ???????? ?????????????? ???????? ???????? (?????? ????? ?????, ??????)
??????????, ????? ????????????? ?????????
? ????.
?????? ????????????? ???????????? ??
?????????? ?????? ????? ?????? ? ??????????? ??????????? ?????. ????????? ?????
??????????? ????????? ?????-????? ? ?. ????? ????????? ? 2005 ?. (???????, ??????,
2006), ?? ??? ?????? ?????????? ?????????
??????????? ????? ? ????? ???????????? ?????????? ?????????? ????? 14 ???. ??????.
?????? ?? 2006-2010 ??., ?? ?????? ???????????? ?????????? ????????? ?????????????
?. ?????, ???????????? ???????? ???? ????????? 54 487 ???????? ?????, ?????? ???????? ??????????? ???????? ?????? ?????????? ??????????? ????? (???. 5). ??? ??????,
? ????? ?????????? ??????????? ?????? ??
??????? ????????? ????????? ?? ????????
??????????? ????????? ?????-????? ? ?? ????????????????? ?? ?????????? ??????. ???,
????????, ? ???????? ????? ?? ?????? ?????
??????????????? ?????????? ??????, ?????????? ???????????? ????????.
?????????? ?????????? ??????, ???????????? ??? ??????? ?????????? ????, ?????
????????? ????????? ??????. ? ????????
????? ???? ??? ???? ????? ?????? ?????????? ???? ???????, ??????? ????????? ??-
# 39 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????????. ????? ????? ????????: ????? ??????????????
???. 5. ?????????? ?????, ??????????? ????????????? ???????? ???????????? ?????????? ?????????
?. ????? ? 2006 ?? 2010 ??. (? ?????? ?????? y = 17077,7 ? 2060,1x)
??? ????????? ?????????? ????? ????????
????????????????? ?????. ??? ?????????
??????????? ???????? ????? ? ?????????,
???????????? ??????. ????? ????, ? ???
????? ???? ??????? ?????? ??????? ????????????? ???????? ????, ??? ??? ???? ?????????? ??????? ?????? ??????, ???????
??????? ????????? ? ??????? ??????. ????????? ?????, ????????, ?????????? ? ?????
?? ????????, ??? ? ?????, ?? ??? ?? ???????? ? ??????, ? ???? ?? ?????????? ?? ??????????? ???????. ?????, ????????, ?
????? ? ??????? ????????, ????????? ?????????? ????? ????? ????? ???????? ?????
?? ?????????, ? ??? ????? ? ????????? ?????
?? ?????? ?????????, ???? ????? ??????
?????? ? ??? ??? ??????????? ???????????
????????? ?????.
???????????? ?????? ? ???????? ????????? ?????? ????????. ???, ? ??????? (??????),
????? (???????), ???????? (??????), ? ????? ?
?????? ?????? ?????????? ???? ?????? ????? ? ?????? ??????: ? ??? ?? ??????, ? ?????
? ??????? (Ostanello et al., 2005; Schalomon et
al., 2006; Vucinic et al., 2008). ???????????? ?
?. ???-???? (???) ???????? ??????????? ??
??????? (Patrick, O`Rourke, 1998). ???? ? ?????????? (Weiss et al., 1998), ?????? ?????????
? ??????? ????? ? ???, ?????? ?????????????? ????????? ? ?????????? ?????? ? ?????
? ?????? ??????, ??????, ???????? ?? ?? ???
??????????? ?????? ???? ????????????????
? ????? ??????, ????????????????? ????????????? ?????? ?? ?????? ???? ????????
????? ? ?? ????? ??????????? ???????.
??? ???????? ???????? ??????????? ???????????? ? ??????????? ????? ???? ???????????, ??? ?????? ?????? ?????? ????
(Szpakowski et al., 1989; Sacks et al., 1996;
Patrick, O`Rourke, 1998; Ostanello et al., 2005;
MacBean et al., 2007). ? ????????? ???????????? ???? ???????? ????? ?? ??????????????,
?????????? ?????????? ??????, ? ???????
??????????? ??????????? ???????.
???????? ????, ??? ? ?????????? ?????? ?????? 15 ??? ????????? ?????? ??????
????, ??? ???????, ????????? ?????, ???????? ??????? ? ?? ?????????. ???????? ? ????
???????? ???? ??????? ?????? ? ????, ??????? ????? ?????????? ????? ? ?????????????
?? ????????: ??????? ?? ??????????, ???????,
???, ???? ? ???, ???? ? ???????? ???????-
# 40 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????????. ????? ????? ????????: ????? ??????????????
?????, ???????, ??????????? ???????????
???????????. ????? ????, ???????? ?????
?????????? ???????? ????? ??? ??????????
????? ???????? ?????? ???? (???????? ??????, ?????? ?? ????? ? ?.?.). ?????? ? ???????? ?? 20 ?? 40 ??? ?????? ?????? ????, ???
??????, ????????, ??????, ??? ??????? ????? ??????? ? ?????? ?? ????????? ? ?????
????????, ???? ?? ??????????.
?????? ?????? ?? ?????????? ???????
???????, ??? ?????? ???? ????? ?????? ?????
? ??????? ????? ? ???????? ?? 5 ?? 19 ???.
??????????? ?????????? ???? ???????? ?
??????????? ???????????? ?????????? ???????, ??????????? ? ?????? ??????? ? ???????.
? ????????? ??????? ???????? ???????????
???? ? ?????????? ????????????, ??, ???????? ?? ???, ????? ??????????????? ????? ?????????????? ? ????? ? ?????????? ??????
?????? ????, ??? ???????? (Sacks et al., 1996;
Ostanello et al., 2005; MacBean et al., 2007). ??
?????? ???? ? ?????????? (Weiss et al., 1998),
? ??? ???? ????? ?????? ????? ?? 9 ???, ????? ??????? ??????? ???? ???????? ? ????????? ?? 5 ?? 9 ??? ? 607 ?????? ?? 100 000 ???????. ??????? ?????? ???? ???????? ??????
? ?????????? (Sacks et al., 1996) ? ??????????
?????? ? ??? ????????? ??? ???????? (?????? 18 ???) 200 ?? 100 000 ?????????, ? ??? ????? (?? 14 ???) ? 640 ?? 100 000 ???????.
???????? ??????? ?????? ???????????? ?????? ????? ??? ??????? ????????, ?? ?????? ?????????? ? ??????????
(Rosenberg, 1995), ????? ??????????? ? ?????????: ????????? ???? ????? ?????? ??
????? ????????? ??? ?????, ? ?????????? ?
?????????? ???????? ?????? ?????? ?????????? ????? ??????????? ????? ???????, ???
???????? ???????????? ??? ??? ?????? ???
????????.
?????????? ??????????? ?????????????
??????????? ?????? ????? ????? ?????????
???????. ??????? ??? ?????? ???????? ???
????????? ?????, ??? ??? ????????????? ??
?????? ????? ?????? ? ?????? ??? ???????
??????. ? ??????? ??? ????? ?????????, ???
???????, ? ??? ???????, ????? ????????????
????? ???? ? ?????? (???????? ?????????, ?????????, ????????, ???????? ?????????????
??? ??????????? ?????????) ??? ?? ?????????? ?? ????????? ??????, ?????????? ?????. ?? ?????? ?.?. ????????? ? ?????????
(2010), ? ??????? ?????? ? ??? ???? ???????? ????? ??????????????? ?? ?????????????
?????, ??? ??? ?? ??????? ?? ??? ????????? ?
?? ???????? ????????? ?????.
?? ???????????? ??????, ???????????
?????? ????? ? ???????????? ??????? ?????????. ?????? ????? ???????? ??? ?????
??????????????, ?????????? ?? ?????? ?????????????: ?????? ? ??????????? ?????
??????? ???? ???? (Szpakowski et al., 1989;
Patrick et al., 1998; Schalomon et al., 2006),
????? ???? ????? ?????? ? ??????, ???? ???
??? (Szpakowski et al., 1989; Gershman et
al., 1994; Weiss et al., 1998; Ostanello et al., 2005;
Schalomon et al., 2006). ? ????? ?? ???? ?????????? ??????? ??????? ????? ?????? ?????????? ?? ???? ? 26-48 % ? ?? ???? ? 18-36 %,
? ? ?????? ???? ?????? ????? ? 4-10 % ????
??????? (Szpakowski et al., 1989; Gershman
et al., 1994; Patrick et al., 1998; Ostanello et
al., 2005; Schalomon et al., 2006). ???????????
???????? ?????? ??????????? ? ???????????
???? ???????.
??????????
? ?. ????? ???????????????? ???????????? ??????? ?????????? ?????? ???????? ???????? ? ? ??????? 2901,3 ?????? ? ???, ??? ????????????? 256,9 ????? ?? 100 000 ?????????.
??????????? ?????, ???????? ????, ???????????? ?????: 69 % ???? ???????? ????????? ?????????? ???????? ? 30,6 % ? ?????-
# 41 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????????. ????? ????? ????????: ????? ??????????????
???? ????????????, ???????? ?????? ??????
????? ? ? 0,36 % ??????. ? ???????-??????
?????? ?? ??????????? ??????? ? ????????
?? ?????? ????? ???? ?????????? ?????????? ????, ??? ????? ? ??????. ???? ????????
?? ?????? ????? ????, ??? ????????, ??? ????
???????????? ????? ??????? ?????? ? ???????
?????? ? ???. ???????? ????? ???? ????? ?????? ?? ???? ??? ?? ????. ?????? ?????? ?????? ????, ??? ??????.
?????? ??????????
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
???????? ?.?., ??????? ?.?., ??????? ?.?., ???????? ?.?. (2010) ????????? ????? ? ??????
?? ?????? ???????? ?? ? ?????? ??I ????. ?????????? ???????????? ??????. ?????? ???????? ? ????? ????????. ?3: 2-6.
??????? ?.?., ?????? ?.?. (2006) ???????????? ????????? ?????-????? ?. ?????. ?: ???????? ???????????? ???????? ??????. ????????? ?????????. ???.???. ?.?. ???????,
???? ??. ?.?. ????????? ???, ??????, ?.123-138.
???????? ?. (2007) ?????? ? ?? ?????????. ?: ?????????? ??????????. ? 10, ?.35.
American Veterinary Medical Association. (1997) US pet ownership and demographics
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Public Health Reports. 90 (3): 262-267.
Chang Y.F., MacMahon J.E., Hennon D.L., LaPorte R.E., Coben J.H. (1997) Dog bite incidence in
the city of Pittsburg: a capture-recapture approach. American Journal of Public Health. 87 (10):
1703-1705.
Coppinger R., Coppinger L. (2002) Dogs: A new understanding of canine origin, behavior, and
evolution. Chicago: University of Chicago Press, 352 p.
Gershman K.A., Sacks J.J., Wright J.C. (1994) Which dog bite? A case-control study of risk
factors. Pediatrics. 93(6): 913-917.
Griego R., Rosen T., Orengo I., Wolf J. (1995) Dog, cat, and human bites: A review. J. Am. Acad.
Dermatol. 33: 1019-29.
MacBean C.E., Taylor D. McD., Ashby K. (2007) Animal and human bite injuries in Victoria,
1998-2004. Medical Journal of Australia. 186 (1): 38-40.
McHarg M., Baldock C., Headey B., et al. (1995) National people and pets survey. Melbourne:
Urban Animal Management Coalition.
Ostanello F., Gherardi A., LaPlaca L., Passini A., Prosperi S. (2005) Incidence of injuries caused
by dogs and cats treated in emergency departments in a major Italian city. Emergency Medicine
Journal. 22: 260-262.
Ozanne-Smith J., Ashby K., Stathakis V.Z. (2001) Injury Prevention. 7: 321-326.
Parrish H.M., Clack F.B., Brobst D., Mock J.F. (1959) Epidemiology of dog bites. Public Health
Reports. 74: 891-903.
Patrick G.R., O`Rourke K.M. (1998) Dog and cat bites: epidemiologic analysis suggest different
prevention strategies. Public Health Reports. 113: 252-257.
Presutti R.G. (2001) Prevention and treatment of dog bites. Am. Fam. Physician. 63: 1567-72.
Rosenberg N., Dos Santos L., Abramo T., Schunk J. (1995) Dog bites. Pediatric Emergency Care.
11: 313-316.
# 42 #
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?.?. ???????, ?.?. ?????????. ????? ????? ????????: ????? ??????????????
18. Rufino Gonzales J.F. (1990) Prevenir las mordeduras caninas. Rev Enferm. 13: 15-21.
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fatal human attacks in the United States between 1979 and 1998. JAVMA. 217(6): 836-840.
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Analysis of dog bites in children who are younger than 17 years. Pediatrics. 117(3): 374-379.
22. Szpakowski N.M., Bonnett B.N., Martin S.W. (1989) An epidemiological investigation into the
reported incidents of dog biting in the city of Guelph. Canadian Veterinary Journal. 30: 937-942.
23. Vu?ini? M., Dordevic M., Radenkovi? B., Jankovi? L., Mirilovi? M. (2008) Bites to humans
caused by stray and owned dogs in Belgrade. Acta Veterinaria (Beograd). 58 (5-6): 563-571.
24. Weiss H.B., Friedman D.I., Coben J.H. (1998) Incidence of dog bites injuries treated in Emergency
Departments. The Journal of the American Medical Association. 279 (1): 51-53.
Biting Humans by Dogs
(General Description)
a
Marat T. Makenova and Olga A. Mihaylovab
Omsk Research Institute for Natural and Focal Infections,
7 Mira av., Omsk, 644080 Russia
b
Omsk Regional Center for Hygiene and Epidemiology
of Federal Service on Customers? Rights Protection
and Human Well ? Being Surveillance,
42a 27 Severnaya st., Omsk, 644116 Russia
The analysis of appeals for medical help concerned with dog bites made by dwellers of the Omsk City
in 2006-2010 has been performed, using the materials of the ?Omsk Regional Center for Hygiene and
Epidemiology?. In total, 11849 cases of human biting by dogs were registered in this period of time.
The mean number of cases is 2901,3 bites per year that is equal to 256,9 cases per 100 000 individuals.
In opinion of the dogs? victims, 99 % of bites were made by owned dogs, in 66 % humans were bitten by
animals familiar to them. The percentage of bites by stray dogs is only 0,36 %. Seasonal relationship
has been found: the maximum amount of appeals for medical help is registered in summer months
(May-August). The minimum amount is observed in the winter period, from November to February.
Children suffer from dog bites more frequently than adults. The percentages of bites in the head and
neck regions are relatively high (19-44 %). Adults are most often bitten in arms or legs (87 % of all
registered bites). Females are rarely bitten than males (217,6 and 304,0 bites per 100 000 individuals,
respectively).
Keywords: dogs, bites, aggression, stray animals.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2013 6) 44-60
~~~
??? 581*581.1: 582.475.2
??????????? ??????? ? ????????
???????????? ????????? ? ???????? in vitro
? Pinus sibirica Du Tour
?.?. ???????????*, ?.?. ???????????,
?.?. ???????, ?.?. ???????
?
???????? ???? ??. ?.?. ???????? ?? ???,
?????? 660036, ??????????, ????????????? 50/28
?
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ?????????, 79
Received 05.12.2011, received in revised form 28.01.2013, accepted 17.03.2013
? ???????? in vitro ?? ???????? ???????????? ????????? ?????, ????????? ? ???????? Pinus
sibirica (???? ?????????), ?????????????? ? ???????????? ????????? ????????? ????? ? ??
???????? ?????????, ??? ??????????? ?????????????? ??????. ???????? ???????????? ???
? ?????????? ????????? ?????, ?????????? ? ?????????? ??????????????? ???????? ??????
??????? ??????-??????????? ? ?????????? ?????? ???????? ??????? ?????. ??????????????
???????????? ????????, ??? ?? ?????? ???????? ??????? ? ???? ?????????? ???? ???????
????????? ?????? ??????????. ?? ?????? ???????????? ? ???????????? ???????????? ?????
????? ????????? ????? ?????? ????????? ??????, ????????? ??????? ????????-???????????.
??? ???? ?????????? ?????? ???????????? ???????? ? ??????????? ????????????? ????????,
?? ??????? ??????????? ??????? ???????????? ????????? ? ????????????? ??????.
???????? ?????: ???? ?????????, ?????? ??????-???????????, ????????????
????????, ????????? ????????, ??????, ???????????? ?????, ???????????? ????????.
????????
Pinus sibirica Du Tour (???? ?????????) ?
???????????? ??????????? ??? ? ????? ????? ?????? ? ???????????? ????????????
??????????? ? ?????????? ?????????????
????????. ?????? ? ??? ???????????? ????????????? ????? ?????????? ?????????? ??-??
????????????? ??? ??????? ???????????, ??-
*
????? ?????????? ?? ?????? ??????????????
????????? Pinaceae:
? ??????? ???????????????????? ?????, ?? 16 ????????? ? ????? ??????????????, ????????? ?????????? ?????????????
(4-7 ???.) (??????????, 1990);
? ?????????????? ? ????????? ?????????? ????? ?????????? ?????????? ??????-
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: culture@ksn.krasn.ru
# 44 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
??? (0,2 %) ? «??????-???????????», ? ???????
??????????? ??????????? ????????????? ????? (?? ???????? ?? ?????????????? ????????
2 ???. ?????? 14-15 ???. (?????????, 1974;
??????, ?????????, 1979). ?????? ??????????? ????? ???? ???????????? ????? ?????????? ?????????? ???????????? ??????????????? ? ?????????? ???????? (??????????,
1990, ?????????? ? ??., 2004).
??? ???????? ??????????????? ????????
????? ?????????? ? ?????????? ??? ?????????? ???? ???????? ????????????? ???????????? ???????? ?????????? ?????????????
?? ??????????????? ???????? ? ????????? ????????????-?????? ????????, ???????
????? ?????????? ex situ ? ????????????
?????????????? ???????? ????????????? ?
???????? in vitro ????? ???????????? ???????????.
???????????? ???????????, ????????
26 ??? ????? ? Picea glauca (Hakman et al.,
1985), ???????? ???????????? ???????? ???????????????? ???????????, ??????????
?? ??????????????? ???????????? ??????
(???????, 1964). ? ????????? ????? ???? ??????? ?? ?????? ???????????? ??? ?????? ???
???????????? ????????? ??????????? (???????????????, ????????????, ???????????????) ???????????? ??????, ??????????????? ?
???????????-???????????? ?????????, ?? ?
????????? ???????? ??????????? ??????????
???????????? ???????? ?????????? ????
? ????????? ?? ???????? ?????????????. ?
?????????? ??????? ???????????? ??????
? ???????????? ???????? ???? ???????????? ? 16 ????? ???? Pinus ? ? ????? ???????? ?? ??? ???????? ????????-???????????
(Klimaszewska, Cyr, 2002). ? ???????? ????????? ???????????? ?????? ??? ????????
????????????? ???????????? ????????????
?????????????? ? ???????? ???????? ??
?????? ??????? ????????. ???? ????????,
??? ?????????????? ?????????? ??? ????????
?? ? ???????? ????????? ???????? ??????? ??
????????????? ?????? ???????? ????????
(???????? ???????????????) ? ??????????????? ?????? ???????? (???????? ?????????)
(Lelu et al., 1994; Klimaszewska et al., 2001; ?
??.). ??????? ????????? ????????????? ???????????? ? ??????????? ????????? ?????
???? ?inus ?????? ???????????. ???, ? P. taeda
??? ?????????? ?? 2 (Becwar et al., 1990) ??
25 % (MacKay et al., 2006), ? P. strobes ? ?? 52,9
(Finer et al., 1989) ?? 76 % (Klimaszewska et al.,
2001), ? P. pinea ? ?? 0,5 ?? 7,2 % (Carnerous
et al., 2009), ? P. pinaster ? ???????? 15 %
(Bercetche, Pagues, 1995). ???????, ??? ????????? ????????????? ??????? ? ???????????
???????????? ????????? ? P. strobus (Carin et
al., 1998; Klimaszewska et al., 2001), P. monticola
(Persy et al., 2000), P. taeda (MacKay et al.,
2006), P. sylvestris (Nikanen, 2004; Lelu-Walter
et al., 2008), P. pinea (Klimaszewska et al., 2009)
?????????? ??? ???????????? ?????????.
??? ???? ???? ????????, ??? ? P. sylvestris ???????????? ???????????? ?????? ?? ???????
????????? ????????????? ???????. ? ????????, ?????????? ??? ??????????? ?????????, ????????? ??????? ? ???????? in vitro
???????? ???????????? ????????, ???????
????????? ????????????? ??????? ??????????? ????, ??? ? ??????????? ??????????
(Lelu-Walter et al., 2008).
?????? ?????????? ?? ???????? ????????????? ???????????? ?? ???????????????
? ????????????? ???????? ???????????? ????????? ? ????? ?????????? ???? ????????
???? ? 2009 ?. ??? ???? ???? ????????, ???
??????? ????????? ??????? ??? ? 58 % ?????????? (??????????, ?????????, 2009).
????? ?????????? ???????????? ???????? ???????? ????????????? ??????? ?? ????????? ????????? ????? ????? ??????????
? ????????? ???????????? ??????????????
# 45 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
????? «?????????» ? «???????????» ????????, ????????????? ? ????????????? ?? ??????????????? ????????.
????????? ? ??????
??????? ????????????
???????? ???????????? ??????? ????? ????? ??????????, ?????????????? ??
??????????? ????????? ???????-?????????
???????? ??????? ????????? (???). ????????
??????????? ????????? ????????????? ?? ?????????? ??? ????????? ??????????? ??????
???. ??????????? (53°16`?.?. ? 92°23` ?.?., ?????? ??? ??????? ???? 300-320 ?). ?????????
???????? ??????? 10 ??. ?????? ???? ??????????? ?????????? ???????? ???????? ????? ?????????? ?? ????????????? ?????????
????????? ?????, ?????????? ?.?. ??????????? ? ????????????, ? ??????? ?? 12-15 ???????? (?????).
? 2007-2011 ??. ?? ???? ???????????????
14-20-?????? ?????? (277/22, 275/20, 280/25,
281/26, 002, 145/4, 153/13) ????????? ?????
?? ??????????????? ????????. ??? ???????? ?????? ???????? ?????? ? ???? ??????????????????? ? ?????????? ?????? ????????
??????? ????? (??????? 106 ? 107) ? ?????
???????? ????????, ????????????????? ????????? ??????? ???????? ???????????????
??????? ?????, ???????? ?????????????????? ????????. ??? ??????? ????????????
? ???????????? ????????? ????????? ?????.
??????? ???????? 100-110 ???. ?? ???? ??????? ????????, «?????????» ? «???????????»
(?????), ????????? ???? ?????, ??????????
? ?????????? ?????????? ????????, ? ?????
??? ?????????????? ???????? ?????? ? ??
????????????. ????????? ?????? ???? ???????? ?? 49 ????????? ???????????????
????????. ? ????. 1 ????????? ???????? ???-
??????? 1. ???????? ??????????????? ???????? ? ??????????? ????? ????? ?????????? ?? ????????
????????? ???????-????????? ???????? ??????? ????????? (????????? ? ????? ???????????? ?????,
??????????? ? % ????????????? ????? ?? ????? ????????????)
????
???????????????
275/20
277/22
145/4
153/13
280/25
281/26
002
2006/2007 ??.
106
16/80
492
14/78
277
2/40
24/60
2008/2009 ??.
106
66/94
21/68
13/50
40/56
15/37
??2
55/88
1/10
3/100
27/66
9/43
357
29/10
17/100
????????????
31/56
6/100
4/100
8/27
5/25
13/50
9/56
2010/2011 ??.
106
14/47
8/29
12/40
5/41
13/45
17/62
15/45
107
9/56
11/79
5/22
13/76
12/75
5/55
12/67
??1
10/83
10/83
7/47
8/61
11/48
10/55
17/80
??2
10/67
5/62
24/80
21/95
14/82
7/43
14/82
????????????
13/43
10/58
8/40
10/53
15/68
5/28
12/70
# 46 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
???????????? ???????? ? ??????????? ????????? ?????; ? ?????? ????????? ???????? ?? 2 ?? 66 ?????, ?????? ??????? ????
???????????? ? ????????????? ?? ????????
?????.
???????????? ????????
? ???????? ?????????? ??? ????????
??????? ???????????? ???????????? ???????? ?? ??????????????? (I ?????? ????)
? ??????????? (III ?????? ???? ? ??????
???????) ???????. ? ???????? ??????? 20-50
????????? ?? ??????? ???????? ??????????????? ????????. ?????????????? ????????? ?? ????? ? ????????????? ???????????? ?????? ??? ????????? ???? ? 90 %-???
??????? (1:3) ? ??????????? ???????????
??????????? ? ?????????? ???????????????? ????. ? ?????????? ???????? ???????? ????????? ?? ??????????????? ? ????-
???? ?? ??????????? ????? Ѕ LV (Becwar
et al., 1990) ? ??????????? ???????????
(0,1 ?·?-1), L-????????? (1,0 ?·?-1), ????????????: 2,4-????????????????????? ???????
(2,4-?) (0,5-3 ??·?-1) ? 6-?????????????????
(6-???) (0,5-1,0 ??·?-1), ???????? (30 ?·?-1), ?
????? ????? (7 ?·?-1). ????? ????, ? 2011 ?. ????? ????? ?????? ??????????? ? ??????????? ?????? ??? ??????????? 0?2 є?, ? ???????
1-2 ???????, ? ? ????? ??????? ????????????
??????? ? ???????????? 300 ??·?-1. ???????????? ????????????? ????? ??? ?? ??? ??
????? ? ?????????? ????????????? 6-???
(0,25-0,5??·?-1) ? ???????? (20 ?·?-1). ??????????????? ????????? ? ??????? ??? 25°?±1°?.
??? ????????? ????????????? ????? ???????
???????????? ?????? ?????????? ????? 10,
20, 30 45 ? 60 ?????. ? ????. 2 ???? ????? ?????????? (?????????), ? ??????? ??? ??????????? ???????.
??????? 2. ????? ?????????? (???????? ????? ?????? ????? ??????????), ?? ??????? ??? ???????
??????
????
???????????????
275/20
277/22
145/4
153/13
280/25
281/26
002
2006/2007 ??.
106
12
492
14
277
8
17
2008/2009 ??.
106
25
8
14
15
16
??2
21
6
9
10
11
357
11
10
????????????
12
4
12
9
13
7
8
19
21
24
31
2010/2011 ??.
106
22
15
107
14
17
11
16
20
22
15
??1
16
12
17
8
16
13
26
??2
13
9
11
15
19
14
17
????????????
7
10
14
11
6
8
7
# 47 #
29
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
?????????????? ??????
??? ??????????????? ??????? ???????????? ????????? ???????? ?????????. ??????? ????????? ?????????? ? ???????????
????? ???????????? ??????. ???????? ???????? ???????????? ?? ?????????? «????????-6» (????, ??????). ?????? ?????? ?
????????????? ???????? ?????? ??? ??????
??????-??????????.
?????????????? ??????
??? ?????? ????????????? ???????? ???????????? ????????????? ?????????????
1??
1??
?????? ? t-???????? ????????? ? ????????
?????? (??? p ?0,05) (????????, 1973).
??????????
???????????? ????????? ?????,
????????? ? ????????
?? ????????????? ?????????
??????????????? ?????? ?????????
?????, ????????? ? ???????? ????? ?????????? ?? ????????????? ?????????, ?????????
? ???????? in vitro, ??? ??????? ??? ?? 4-8-?
????? ? ????????? ? ???????????? ??????? ?
??????? ???????????? ??????? (???. 1?). ??
?
1??
?
1??
?
?
?
E
1??
1??
???. 1. ??????????? ??????? ?? ?????????? ????????? ????? ????? ?????????? ?? ????? 1/2 LV: ? ?
???????? ??????? ? ??????? ???????????? ??????? (7-10-? ????? ???????????????); ? ? ???????????
??????? ?? ???? ????? ?????????? (30-? ????? ???????????????); ?, ? ? ??????????????? ??????? ??
50-60-? ?????; ?, ? ? ????????? ???????? ? ??????? (60-? ????? ???????????????)
# 48 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
30-? ????? ??????????????? ??????? ?????????????? ?? ???? ????? ?????????? (???. 1?).
??????? ??????????? ? ????????????? ??????????? ?????????? (75-80 %), ????????? ?
???????? ?? ??????????????? ? ???????????
??????? ???????? ????????????? ????????,
????? ??? ????? ?????????? 2 ?? (I-III ??????
????).
?????????? ?? ????????? ????? ????????????? ??????? ?? ????? 1/2LV ????????, ???
??????? ????????? ?????????????????? ? ????????? ?????, ????????? ? 15 ???????? ????? ??????????, ?????????????? ? ???????????? ?????????, ?????????? ?? ??????????
??????????? ????????? ?????. ?? 28-? ?????
??????????????? (????? ?????? ?????????) ?
??????????? ?????????? ????? ??????? ?????????? ?? 0,29 ?? 0,79 ? (???. 2). ??? ?????????
???????? ?? ???????????????? ????? ????
?? ???????????. ????? 60 ???? ??????????????? ???????? ???????????? ??? ??????? ?
?????????? ?????? 358, ????? ??????? ???????? ????????? 0,98 ?. ???????? ?? ??????????
????????????? ? ?????????? ?????? 358 ??
????????? ? ??????? ?????????-????????
???????? ????????????? ????????? (F=16,87;
p < 0,05).
??? ??????????? ?????????? ???????
??????? ? ?????????? ?? ????????????? ????????? ?????????? ? ? ??? ?????????? ????????????? ??????????? (???. 1?, ?). ? ??????????
???? ???????? ? 277, 257 ? 358 ? ???????????? ??????????? ? ??????? 7-10 ???????, ????? ???? ??????? ?????? ? ????????. ?????
???????, ? ?????????? ????? ?????????? ??
????????????? ????????? ????????????? ?????????????? ???????.
???????????? ? ????????? ?????,
????????? ? ??????
? 2010 ?. ?? ???????? ????????? ??????
????? ??????????? ????????????. ???? ?
??????? ?????? ?????????? ??????????? ?????, ???????? ????? ??????? ???? ???????
? ???????? in vitro ?? ?????? ????????? ?
????????? ????????? (?????? ? ?????? ??????
1
0,9
0,8
?????,??.
0,7
7???.
0,6
14???.
0,5
28???.
0,4
56???.
0,3
0,2
0,1
69
70
35
8/
70
4
35
7/
47
6
47
4
43
5
43
4
43
3
42
9
42
8
42
7
39
0
36
0
28
3
27
7
0
???. 2. ???????? ????? ???????, ??????????? ?? ?????????? ????????? ????? ????? ?????????? ??
????????????? ?????????, 2009 ?.
# 49 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
3,00
2,50
?????, ??.
2,00
20???.
35???.
1,50
45???.
1,00
0,50
?
0,00
277/22
002
145/4
153/13
3,5
3
?????, ??.
2,5
30???.
2
45???.
1,5
60???.
1
0,5
?
0
277/22???. 275/20???. 280/25??? 281/26???.
002???.
145/4???.
153/13???.
???. 3. ???????? ????? ???????, ??????????? ?? ?????????? ????????? ????? ?????? ????? ??????????
(????????? ????????), ? ? 2010 ?., ? ? 2011 ?.
????). ? ?????????? ????????? ????? ?????????????? ?????? ??????????? ??????? ??
?????????? ?? ????????????? ????????, ?????????? ?? ???????? ?? ????????????? ?????????. ???????? ???????? ???? ??????? ???
? ????? 277/22. ?? 1 ????? ????????? ?????
??????? ????? ????? ????????? 1,96 ?, ? ?????
??? ?????? ?? ???????????????? ????? ???
????? ??????????? ? 2 ???? (???. 3?). ? ????
?????? ?????? ?????????? ????? ??????? ???
??????????? ?????? ? ?? 60-? ????? ??????????????? ??????? ?????? ? ???????????????.
? ????? 277/22 ???? ??????? ??? ??????????
?????????? ??????????? ? ??????? 12 ???????, ????? ?????? ???????. ??????? ??????-
?? ????? ?? ??????????????? ??????????? ?
???????? in vitro ???????? ????????????? ???????? (F=5,5; p < 0,05).
? 2011 ?. ? ???????? in vitro ???? ???????
???????? ????? ?? ?????? ????????? ????????? (?????? ?????? ????), ?????????? ??
????????????????? ??????????? ?????? ????? ??????????. ?????????? ?? ????????????
???????? ?? ?????????? ????????? ????????,
??? ???????????? ?? ??? ?????. ????? ??????? ? ????? ?????? ????????? (30-? ?????)
? ?????????? ?? ?????? ?????? ?????????? ??
0,01 ?? 0,66 ? (???. 3?). ???? ? ????? 153/13
????? 1 ????? ??????????????? ????? ??????? ????????? 0,7 ?. ?? ?????? ????????????
# 50 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
????? ?????? ??? ?? ???? ????????????????? ?????? (277/22, 002 ? 153/13). ? ?????
??????? ??????? ?? ???????????????? ?????
??????? ?? ????????????????? ??????????
?????????????????.
???????????? ? ?????????
????????? ?????
? 2007 ?. ???? ??????? ? ???????? in
vitro ????????? ???????? ?? ?????? ????????? ????????? (?????? ?????? ????), ?????????? ? ?????????? ??????????????? ???????? ??????. ???? ???????? ????????? ?????,
?????????? ? ?????????? ???????? ??????
277/22, 280/25 ? 281/26, ??? ?????? ??????
(???. 4). ???????? ???????? ??????? ????????? ????? ??? ??????? ? ?????? ?????? ?
????????? ??????????????? ???????? ? ?????????????? ?????? ??????-?????????? 106.
????? ??????? ??????? ?? ????????????????
????? ??????? ??????? ???????? ???????????
?? ??????? ? ??????? ???????????????. ???????????? ???????????? ?????? (???. 1?, ?).
????? ???????, ??????????? ?? ?????? 106, ?
2 ???? ????????? ????? ??????? «?????????»
???????? ?? ????????????? ????????? (???. 4).
????????????? ??????? ?????? ???????????????? (106) ?? ??????????????? ??????????? ? ???????? in vitro ?? ????????? ? ??????? ??????????? ???????? ?????????????
????????. ?????? ??????? ????????? ???
????? 277/22 ????? ?????????? ????????
277/22x106 ? 277/22x492.
? 2009-2011 ??. ? ???????? in vitro ????
??????? ????????? ????????? ?????, ?????????? ? ?????????? ??????????????? ???????? ?????? ??????? ????????-???????????
106 ? 107, ???? ???????? ???????? ? ????????????? ??????.
? 2009 ?. ???????????? ? ?????????? 18
????????? ????????? ?????, ???????? ??????? ???? ??????? ? ???????? ?? ?????? ???????? ????????? (??????), ??? ????? ?? ????????? ? ??????? ? 2007 ?. ? ????? ???????
????????? (30-? ?????) ????? ??????? ? ????????? ?????? ?????????? ?? 0,07 ?? 0,38 ?.
3
2,5
?????,??.
2
10???
20???
1,5
30???
45???
1
60???
0,5
0
277/22?106
280/25?106
277/22?277
281/26?492
277/22?492
???. 4. ???????? ????? ??????? ?? ?????????? ????????? ????? ?????? ????? ??????????, ?????????? ?
?????????? ??????????????? ???????? ?? ???????? ??????????? ?????????, 2007 ?.
# 51 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
1,2
?????,??
1
0,8
30???.
45???.
0,6
60???.
0,4
0,2
15
3/
13
15
*1
3/
06
13
*1
53
/1
3
00
2*
10
14
6
5/
4*
14
10
5/
6
4*
14
5/
4
28
0
28 /25
*3
0/
57
25
*2
77
/2
28
2
1/
26
*1
06
20
*2
77
/2
20
2
*2
75
/2
0
27
5/
27
5/
27
7/
22
*2
77
0
???. 5. ???????? ????? ???????, ??????????? ?? ?????????? ????????? ????? ?????? ????? ?????????? ?
?????????? ??????????????? ???????? ?? ???????? ??????????? ?????????, 2009 ?.
???????????? ??????????? ???? ? ??????????, ?????????? ?? ???? ?????? 280/25 ? 002,
????????? ??????? ????????? ?????? 2??.
?????? ???????????? ?????????? ???? ?
?????????? ????????????? ?????? (???. 5).
???????? ??????????????? ??????????? ?
???????? in vitro ??? ?????? ????????? ???????? ???? ????????????? ??????? (F=311,7;
p < 0,05). ?????? ???????????? ? ?????????? ?
2009 ?., ????????, ????? ????????? ???????
?????? ????? ????? ? ????????? ????????? ?
???????? ?? ?????? ?????? ?????????.
? 2011 ?. ? ???????? in vitro ???? ???????
????????? ????????? ????? ?? ?????? ????????? ????????? (?????? ?????? ????) ?? 18
????????? ??????????????? ????????, ????????? ??????? ???? ??????-???????????
106 ? 107 ? ?????????? ?????? ???????? ?
??????? ????????? ?????? 2??, ? ????? ????????????? ??????. ?????????? ???????????? ????????, ??? ?? ?????? ????????? ?????
??????? ? ?????????? ????????? ????? ? ?????????? ?? 0,08 (??????? ???????? 153/13?107)
?? 0,5-0,6 ? (??????? 145/4?1??, 153/13?1??,
153/13?106, 153/13?153/13). ?????? ??? ??????????? ???????? ?? ????? ? ??????????
??????????? 6-??? ???? ???????? ???????
??????????. ???, ? ???????? ??????????????? ???????? 277/22?107 ?? 45-? ????? ?????
??????? ????????? 1,74 ?, ? ?? 60-? ????? ???
??????????? ?? 2,75 ?; ? ???????? 275/20?106
?????????????? 0,81 ? 2,9 ? ? ?.?. ????????
???????? ???????????? ??? ??????? ? ??????????, ?????????? ? ?????????? ????????
?????? ??????? ????????-???????????. ?
2011 ?. ???????? ? ?????????? ????????????? ??? ???????? ?????? ??????? ??????????????????? ?? ???????? ????????????? ?????????. ??? ?????????? ???????????????
???????????? ??? ???????? ?????????? ? ????? 153/13, ???????? ? ???????? ??????????????? ???????? 153/13?107 (??????-??????????)
(???. 6). ??????? ??????? ???????? ?????????
?????? ?????????, ??????????? ??? ???????????? ?????, ? ? ??? ??????????? ????????
??????????? ???????????? ????? (??).
# 52 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
3,5
3
?????,??
2,5
2
30???.
1,5
45???.
60???.
1
0,5
0
???. 6. ???????? ????? ??????? ????? ??????????, ??????????? ?? ?????????? ????????? ????? ??????
????? ?????????? ? ?????????? ??????????????? ???????? ?? ???????? ??????????? ?????????, 2011 ?.
????????? ?? ????????????? ??????,
????????? ? ????????, ?????????????????
?????? ???????????????????. ? ????? ?????? ????????? ????? ??????? ? ?????????
????????????? ?????? ????????? ????? 0,1 ?,
?? ??????????? ????? 153/13, ? ???????? ????? ??????? ?????????????? ???????? ? ?????
????? ??????????? ?? 0,5 ?. ??????? ????????,
??? ?????? ? ????? ????? ? ??????? ??????????? ???????????? ??. ??????? ?????????
???? ????????????? ?????? ????????????????? ????? ??????? ?? ??????? ?????? ????????????.
????? ???????, ?????????????? ???????? ?????? ????? ?????????? ?? ????????
??????????? ????????? ???????????? ????????? ?? ????? ????????? ???????. ????????
??????? ?????????? ????? ??????? ???? ???????? ? ????????? ??????????????? ???????? ?????? 002 ? 153/13, ????????? ???????
??????-???????????, ?? ?????? ???????? ??
???????? ????????????? ?????????.
??????????????????? ????????????
??????????? ???????
? ???????????? ?????????
?????????????? ???????????? ????????,
??? ?? 20-? ???? ??????????????? ??? ???????????? ?????? ????????????? ?? 200-300
??? (???. 7?). ?????????? ?????? ???????
?????????? ??? ????????? ?? ????? ?????????. ??? ????????? ???????? ?? ???????????????? ????? ? ????????? ?????? 277/22 ?
153/13 (2007, 2011 ?.) ? 002 (2011 ?.), ?????????
??????? ???????? ??????-???????????, ?????????? ?????? ???????????? ?????????????
??????? ? ??????????? ????????? ?????? ?
????????????? ???????? ? ? ??????? ?????? ? ????????????? ??????. ??? ???? ??????
??????? ????????: ?? ??????????? ??????
?????????????? ??????? ???????????? ?????????, ? ?? ????????????? ?????? ????????????? ??????? ????????? ?????????
? ?????????????? ????????????? (???. 7?,
?). ?????? ??????? ?????????? ??????????-
# 53 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
?
?
0,1 ??
?
?
0,1 ??
0,1 ??
?
0,1 ??
???. 7. ???????????? ???????????? ?????????: ? ? ????????? ???????????? ?????? ??????????;
? ? ????????????? ?????? ? ?????????????; ? ? ??????? ????????????? ???????? ? ??????????????
???????? ?? ?????????? ?????; ? ? ?????? ???????????? ????????? ?? ?????? ???????; ? ?
??????????????? ???????????? ????????
???? ?????????????? ????????. ????????????? ??, ??????? ???????? ?? ??????
???????????? ????????? ? ????????????
?????? ? ???????????. ????????????? ??????? ???????????? ??????? (???. 7?). ??
50-60-? ????? ??????????????? (1-? ?????
????????????) ? ????????? ???????? ???-
?????? ??????????????? ???????????? ???????? (???. 7?).
????? ???????, ???????????????????
???????????? ????????, ??? ???????? ??????????? ????????????-???????????? ?????
? ???????????? ? ??? ???????????? ????????? ??????????? ???? ? ???????? ??????-
# 54 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
????????? ???????? ?????? ??????? ???????????????.
??????????
???????????? ?????? ? ????????????
???????? ???? ???????????? ??????????? ??????? ? 16 ????? ?????: P. sylvestris,
P. pinaster (Lelu et al., 1999, 2008; Nikanen et
al., 2004; Park et al., 2006), P. strobus (Garin
et al., 1998; Park et al., 2006) ? P. banksiana
(Park et al., 1993, 1998, 2006), P. caribae (Laine,
David, 1990), P. radiata (Smith, 1996), P. nigra
(Salajova et al., 1999), P. roxburghii (Arya et al.,
2000), P. monticola (Persy et al., 2000), P. pinea
(Carnerous et al., 2009), P. taeda (?ercwar et al.,
1990; MacKay et al., 2006) ? ??. ??????? ????????? ????????????? ??????? ???????? ??
??????-??????, ?????? ???????? ????????? ?
????????, ??????? ??????????? ?????, ???????? ? ????? ????????, ?????????? ??????? ?
?????? ???????? (Park et al., 2006). ??? ????
?? ?????? ? ????????? ? ???????????? ??????? ??????????? ????????????? ??????? ?
?????? ????? Pinus ?????????? ?? 2 ?? 76 %
(?ercwar et al., 1990; Klimaszewska et al., 2001;
MacKay et al., 2006; ?arneros et al., 2009).
? ????? ?????????? ????????? ???????
?? ????? Ѕ LV ??? ? 75-80 % ????????? ??
????????, ?????????????? ? ????????????
?????????, ?? ???????? ????????? ? ???????????? ??????????????? ????????.
??????? ????????????? ??????? ?????????
? ???????? in vitro ????????????? ???????
???????. ???????? ??????????? ???????????
??????? ???? ???????? ? ?????????, ?????????? ? ?????? ????????? ???????????????
???????? ? ???? ??????: 277/22, 002 ? 153/13.
?????? ?????????? ??????? ?????, ??? ???????, ??????? ??????? ????????? ? ???????
6-10 ??????? ??????????????? ? ????? ???????????? ???????. ???????????? ????????
? ?????? ????? ?????????? ???? ??????????
? ????????? ?????, ????????? ? ???????? ??
???? ?????? (002 ? 153/13), ????????? ??????? ????????-???????????. ?????? ???????
?????????????? ? ??????? ????? ????????
???????? ? ?????? ??????????, ??????? ?????, ? ? ??? ??????????? ???????????? ??????????????? ??.
????????, ??? ??????????? ?? ? ???????????? ????????? ? ????? ??????? ???? ???
??????? ???????????? ????????? (Pullman,
2007; Carneros et al., 2007). ???????? ?????
?????? ????????? ????????-??????? ????????? ?????????? ???????????? ???????????? ?????? ? ???????????? ????????
(Klimaszewska et al., 2009; Lelu-Walter et al.,
2008; Tretyakova, Barsukova, 2010). ???????????? ???????? ????????? ????????????? ???????????? ??? ??????? ? Picea glauca (Park et
al., 1993), Pinus sylvestris (Niskanen et al., 2004;
Lelu-Walter et al., 2008), Pinus taeda (MakKay,
2006). ????? ?? ??????????????? ????????
P. sylvestris, ??? ? ???????? ???????????? ?????? ???? ???????????? ??????????? ? ????????? ??????? ? ?????? ???????? ?? ?????????? ?? ???????????? ???????????, ????????
??????? ???????????? ??????? ?? ????????
????????????? ????????????. ????????????
????? ? ???????????? ??????????? ???? ???????? ?????? ??? ????????, ???????????? ?
?????????????? ???????? ??? ???????????.
??? ???????????? ?????? ???????? ??????????? ???? ?????? ????????? ?????????????
????????????, ? ??????????? ??????? ?? ??????????? (Nikanen et al., 2004), ??? ??????
??????? ??????? ?????????? ? ????????????
???????????? ???????, ???? ??????? ?? ????????? ?? ????????? ? ???????????? ?????????.
? ????? ?????????? ???????????? ???
? ????????????? ??????????? ??????????.
?????? ? ????????? ?????, ?????????? ??
????????????? ??????, ???? ??????? ???
# 55 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
????? ?????? ??? ??????????? ??????? ?????? ?? ???????????. ???????????? ??????
? ??????????? ???????????? ????????? ????
?????????? ?????? ? ???? ??????, ????????? ??????? ????????-???????????. ??????
????? ???????? ??????? ??????????????
???????????. ????????, ??? ? ????? ?????????? ??????????? ???????????? ?????????
??????????? ????????? ??????.
??????????? ?? ? ???????? ? ??? ???????????? ????????? ? ????? ?????????? ? ???????? ??? ?? ?????, ????????? ??? ?????? ????? ??????? (von Arnold, Hakman, 1988; Lelu
et al., 1994; Klimaszewska et al., 2001; Stasolla,
Yeung, 2003; Lelu-Walter, Pagues, 2009). ???????????? ???????? ????????? ???? ????????? ? ????????????. ?????? ????? ?????? ????????????? ???????????? ????? ??????????,
??? ? ? ?????? ????? ???????, ?????????? ?
?????????? ?????????????? ??????? ???????????? ?????? ??????????, ??????? ???????
????????? ??????? (2,4-?) ? ?????????? (6???) (von Arnold, ??kman, 1988; Stasolla et
al., 2003; ???????????, ??????????, 2008). ????? ????? ?? ????????????? ??????? ?????
? ?????? ????????????? ???????????? ????
????? ???????? (????????, 1999). ????????,
??? ????????????? ??????? ???????? ??? ?
?????? ??????? ??????, ??????? ???????? ?
??????????? ???? ???????? ??????: ????????? ????????????, ??????? ???? ?????? ????????, ? ??????? ????????? ??????, ??????
?????? ?????? ?????? ? ??????????. ??? ????
??????????, ?????????? ? ??????, ? ?????????? ?? ??????? ?????????????? ? ??????????
???????? ???????. ?? ?????????, ??? ????????????? ??????? ? ?????????? ????????
????????? ??????????, ?????????????
??????? ?????? ?? ???? ????????????: ????????????? ??? ?????????????.
? ????? ?????????? ????????????? ???????? ???????????? ?????????????? ????-
???? ? ??????????? ??????? ????????????
?????????, ?????????? ????????????, ?????????? ?? ??????-??????. ?? ??????????? ?????? ???????? ??????????? ???????????? ????????? ???? ???????? ??????.
????? ??????? ???????????? ?????????
????? ?????????? ???????????? ? ??????????????? ?????????. ???????????? ???????
??????? ?????.
??????? ????????? ?????????????
???? ??????? ? ?????, ? ??? ????? ? ????? ??????????. ? ?????????? ????????????? ???????? ????????, ?????????? ?? ????? ??????????????? ??????????, ??????????? ??
?????? ?????????? ????????????? ???????, ?????? ?? ??????? ???? ?????? ???????
?????????-????????? (??????????, 1990).
????? ????, ? ????? ?????????? ???????????
??????? ? ??????? ???????? ?????????????.
? ????? ???????? ??????????? ????????? ??????? ?????????? (? ????? 2-3) ? ?? ??????????????. ?? ?????? ??????????????? ?????????? ?????? ???????? ? ?????????? ???????
?????????? ?????? ??????????? ????????.
????? ???????, ? ??????????? ?????? ?????
????????? ? ????? ?????????? ????? ??????????? 12-16 ????????? (??????????, 1990).
????? ????????????, ??? ? ??????? ?????
???????????? ??????? ???????, ??????? ?
???? ??????????????? ?????? ??????????? ?
???????? in vitro ? ??????????????? ????????????? ????? ????? ????????? ??????? ?
?????????? ????????????? ?????? (??????) ?
????????????? ?????? ??? ????????? ????????. ??????? ????????, ??? ??????? ? ???????
???????? ????????????? ????? ???????
?????? ? ?????????????? ??????? ?????,
?.?. ????? ??????? ????? ??????? ?????????????? ?????????. ????? ????????????,
??? ????????? ????? ???????? ????? ???????
??????????? ? ???????? ????????? ??? ????????? ???????????? ????????? ? ????????.
# 56 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
??????????
? ?????????? ??????????????? ?????
?????????? ? ???????? in vitro ??????? ????
???????? ???????????? ?????? ? ???????????? ???????? ?? ???????? ??????? ????????? ????????? ?????, ????????? ???????
??????-???????????. ????? ????????? ???????????? ????????? ??? ?????? ? ?????????
?????. ?????????????, ??? ????????? ????????????? ??????? ? ????????? ????????????
????????? ?????????? ?????????? ??????
????? ?????? ?????????, ?????????? ?????????? ???????????? ? ???????????. ?? ?????????, ??? ?????? ?????????? ????????
??????? ? ??????? ?????????????? ?????-
??????, ??? ??? ???? ???????? ???? ???
????? ??????????? (?????????? ? ??., 2011).
??? ???? ???????? ????????????? ???????????? ??????? ?????????? ????? ?? ???????????? ? ?????????????? ?????? ????????????????? ? ?????????? ?????? ????????
??????? ?????. ?????????? ????? ?? ???????? ????????? ????????? ? ???????? in
vitro, ????????? ?? ??? ????????????? ???????, ???????????? ????????? ? ???????????????????? ????? ?????????????? ????????? ?????????????????? ?????? ?????
? ????? ?????????, ?? ??????? ????? ???????? ?????? ??? ???????? ????????????
????????? ??????? ????.
?????? ??????????
1.
2.
3.
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??????????? ?.?., ?????????? ?.?. (2008) ??????????? ???????????? ???????????? ????????? ? ??????????? ?????????: ???????????????? ???????. ????????? 39(2): 1-10.
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????? ???????? ?????????? ???????? ?????? «?????????? ????????- ???????????????
?????? ???????? ? ????????????? ?????????????». ????????? ????????. ?. 2. ??????
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(1998) Somatic embryogenesis in jack pine (Pinus banksiana Lamb). In: Somatic embryogenesis
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35. Stasolla C., Yeung E. (2003) Recent advances in conifer somatic embryogenesis: improving
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36. Smith D.R. (1996) Growth medium, US Pat., p. 5, 565, 355
37. Tretyakova I.N., Barsukova A. (2010) Somatic embryogenesis of Siberian larch. Abstract of
conference «Advances in somatic embryogenesis of trees and its application for the future forests
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Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????, ?.?. ??????????? ??????????? ??????? ? ???????? ???????????? ????????? ? ?????????
Formation of Callus and Induction
of Somatic Embryos in vitro
Culture in Pinus sibirica Du Tour
Iraida N. Tretyakovaa, Elena V. Voroshilovab,
Denis N. Shuvaevb and Anastasya S. Lukinaa
a
V.N. Sukachev Institute of Forest SB RAS,
50/28 Akademgorodok, Krasnoyarsk, 660036 Russia
b
Siberian Federal University,
79 Svobodny, Krasnoyarsk, 660041 Russia
In culture in vitro nonembriogenic callus was inducted from immature zygotic embryos of seeds of
Siberian pine trees (Pinus sibirica), growing in natural stands of West Sayan and in a clonal plantation.
The active callus development goes in explants hybrid seeds, getting as results of control pollination
of clones by pollen of Siberian pine trees with annual cycle development of female cones. Cytological
study shows that the strong elongation in hypocotyl cells goes at the stage of callus induction. Only
explants of embryo seeds of some clones, which were pollination of pollen trees-accelerate reached
the stage of proliferation and formed embrygenic mass. Globules of somatic embryos and embryonal
tubes are developed from embryonal initiales.
Keywords: Siberian pine, pollen of annual cedar, zygotic embryos, hybrid embryos, callus,
embryogenic mass, somatic embryos.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2013 6) 61-72
~~~
??? 550.47+630*561.24
Methodology for Development
of a 600-Year Tree-Ring Multi-Element Record
for Larch from the Taymir Peninsula, Russia
Alexi M. Gracheva*, Eugene A. Vaganova,
Steven W. Leavittb, Irina P. Panyushkinab,
Eugene P. Chebykinc, Vladimir V. Shishova,
Natalia A. Zhuchenkoc, Anastasia A. Knorred,
Malcolm K. Hughesb and Mukhtar M. Naurzbaevd
a
Siberian Federal University,
79 Svobodny, Krasnoyarsk, 660041 Russia
b
Laboratory of Tree-Ring Research, University of Arizona,
Tucson, AZ 85721, USA
c
Limnological Institute SB RAS,
3 Ulan-Batorskaya, Irkutsk 664033 Russia
d
V. N. Sukachev Institute of Forest SB RAS,
50/28 Akademgorodok, Krasnoyarsk, 660036 Russia
Received 17.05.2012, received in revised form 15.03.2013, accepted 22.03.2013
We developed a long (600-year) dataset for the concentrations of 26 elements in tree rings of larch
from the Taymir Peninsula, the northernmost region in the world (ca. 72°N) where trees grow.
Tree rings corresponding to the time period from 1300 to 1900 A.D. were studied. Eleven wood
strips, each from a different larch tree, were cut into ca. 100 mg samples usually consisting of ten
consecutive tree rings (but occasionally fi ve). Between 19 and 40 consecutive samples resulted
from each tree, yielding a total of 277 samples. The replication of each time interval ranged
from three (for periods 1300-1400 A.D. and 1600-1700 A.D.) to six (for 1450-1600 A.D.). Wood
samples were digested with concentrated HNO3 for measurement of Li, B, Na, Mg, Al, Si, P, Cl,
K, Ca, V, Cr, Mn, Fe, Co, Ni, Cu, Zn, As, Rb, Sr, Y, Zr, Nb, Mo, Ag, Cd, Sn, Sb, I, Ba, La, Ce, Nd,
W, Au, Pb, Bi, Th, and U using solution Inductively Coupled Plasma Mass Spectrometry (ICPMS). Fourteen elements (V, Co, As, Y, Nb, Mo, Sb, La, Ce, Nd, W, Au, Th, and U) with extremely
low concentrations were eliminated from consideration as unreliable. Here we report our
sample preparation and measurement procedure, as well as the observed concentrations in tree
rings, emphasizing considerations for developing representative and reliable denrodochemical
datasets.
*
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: amgrachev@juno.com
# 61 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Alexi M. Grachev, Eugene A. Vaganov? Methodology for Development of a 600-Year Tree-Ring Multi-Element Record?
Keywords: Dendrochemistry, Larix gmelinii, elements, tree rings, Taymir, ICP-MS.
Introduction
This work investigates the multi-elemental
composition in tree rings of Dahurian larch
(Larix gmelinii) from the Siberian Arctic near
Khatanga at ca. 72°N, 102°E (Fig. 1). It has long
been recognized that the tree-ring archive can
capture changes in the chemical environment
related to atmospheric chemistry, biogeochemical
cycling and climate influences (see e.g., Baes,
McLaughlin, 1984; Hantemirov, 1991; 1992;
1996; Padilla, Anderson, 2002 and references
therein).
Seasonal and long-term tree growth of larch
at the site of this study is strongly limited by
temperature (Naurzbaev et al., 2003). Elemental
composition of tree rings has not been studied
at this location since the 1980?s (Adamenko
et al., 1985; Chetverikov, 1986), and of course
the analytical capabilities have improved
tremendously since then. Siberian Arctic tree
rings have recently been used to extensively
investigate other tree-ring variables: tree-ring
width (Vaganov et al., 1996, 1999; Hughes et
al., 1999), maximum latewood density (Briffa
Taymir
Peninsula
75o
75o
Khatanga
70o
70o
Norilsk
60o
60o
70o
80o
90o
100o
110o
120o
130o
140o
150o
Fig. 1. Approximate location of sampling site on the Taymir Peninsula marked with a star immediately south of
the town of Khatanga. The city of Norilsk is to the southwest of the sampling site
# 62 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Alexi M. Grachev, Eugene A. Vaganov? Methodology for Development of a 600-Year Tree-Ring Multi-Element Record?
et al., 1998, 2001, 2004; Esper et al., 2010),
tracheid dimensions (Silkin, Kirdyanov, 2003;
Panyushkina et al., 2003) and stable isotopic
composition (Kirdyanov et al., 2008; Sidorova
et al., 2010). It is desirable to precisely measure
as many parameters as possible on each tree ring
and to understand controls on each parameter
to make more accurate reconstructions of past
environmental conditions (Hughes, 2002).
Solution inductively coupled plasma mass
spectrometry (ICP-MS) used in this study allows
a wide range of elements to be measured at the
same time. However the dissolution of samples
is a very laborious procedure involving many
steps, including the need to weigh samples with
high precision multiple times at different stages.
The method was applied to tree rings for the fi rst
time in the work of Hall et al. (1990). The fi rst
report of application of another high-sensitivity
method, solution inductively coupled plasma
atomic emission spectrometry (ICP-AES) to
tree-ring analyses in Russia was made in 1990
by R M. Hantemirov (Hantemirov, 1992). Since
then a number of studies using the solution
ICP-MS method have appeared in literature
(e.g., Yu et al., 2007; Rothpfeffer, Karltun,
2007; Kirchner et al., 2008). Solution ICP-MS
should be distinguished from laser ablation ICPMS (e.g., Monticelli et al., 2009), which is less
laborious but less precise.
We measured samples over the last 600
years, making the element series reported in this
work the longest ever developed. The dataset
presented here was obtained to investigate multielemental composition in tree rings over a long
period in a very remote site in the Siberian Arctic
to document inter-tree variability in long element
time series. This paper reports the methodology
and presents concentrations of elements in tree
rings, whereas a subsequent paper (Panyushkina
et al., in prep.) will analyze and interpret the
observed trends of the elements.
Materials and Methods
Field sampling
In 2006, cross-sections (discs) of 11 larch
(Larix gmelinii) trees were collected from
the Taymir peninsula near Khatanga. All tree
samples were from dead trees collected at a site at
70°53?N 102°55? E (ca. 350 m a.s.l.) on the Kotuy
River (Fig. 1). This location has a strong seasonal
climate, with the mean annual temperature of
-13°C, January temperature of -42°C (minimum)
and -20.4°C (maximum), and July temperature
of 18.3°C (max) and 7.9°C (min) based on data
from the nearest meteorological station Khatanga
(71°98?N 102°47? E, 30 m a.s.l.) for the period
1947-2006. This area receives an average of
254 mm of precipitation annually, ca. 105 mm
of which falls in the growing-season months of
June, July and August.
Laboratory subsampling
Since 2006 the cross-sections were stored
in a wood depository (dry conditions, room
temperature) until the chemical processing. The
work proceeded in several stages, the first of which
involved surfacing each of the 11 cross-sections
with progressively finer sandpaper to increase
visibility of rings. Subsequently one wood strip
was cut out of each cross-section (Fig. 2). Ringwidth measurements and dating were performed
on each of eleven wood strips from the crosssections on the Lintab workstation (Rinntech®,
Germany) with TSAP-Win crossdating software
(Rinntech®, Germany). The chronology used
in this work is the same as in Naurzbaev and
Vaganov (2000).
The eleven wood strips were subsequently
transported to the Institute of Geochemistry
SB RAS (Irkutsk) where they were processed
in the Grade-1000 clean room according to the
following procedure. First, each surface on the
wood strips cut from the cross-sections was
cleaned with a stainless-steel scalpel. It should be
# 63 #
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Alexi M. Grachev, Eugene A. Vaganov? Methodology for Development of a 600-Year Tree-Ring Multi-Element Record?
Fig. 2. Photographs of the 11 wood strips that were analyzed in this work. The labels indicate the time range
covered by each wood strip
noted that the blades on the scalpel were replaced
after cleaning a wood strip and subsequently
cutting it into pieces (see below). Contamination
of wood samples by razor blades was investigated
in the work of Sheppard and Witten (2005), who
found that laser-trimmed cores had less metal
contamination than razor-trimmed cores. Because
the razor blades were from the same batch from
a single manufacturer and the physical contact of
blades with the wood samples was similar for all
samples, any contamination was deemed to be
equal for all of the samples, and thus not affecting
the trends observed.
Ten-year or five-year pieces were cut out
from each wood strip with a scalpel under the
microscope. Five-year pieces were cut only in
cases where tree rings were wide (i.e., cumulative
ring width of the five years exceeding ca. 3 mm).
If 5-year tree rings were narrow, it was very
difficult to make an accurate cut and dry weight
of an obtained piece was markedly lower than the
desired 100 mg for the analysis. Hence, 10-year
# 64 #
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Alexi M. Grachev, Eugene A. Vaganov? Methodology for Development of a 600-Year Tree-Ring Multi-Element Record?
samples were taken in most cases. Each sample
was placed in a pre-weighed sterile polypropylene
tube. Measuring element concentrations in 5 or 10ring samples has two advantages over measuring
concentrations in single-year samples: i) reducing
the influence of any potential contamination due
to a greater sample mass, ii) removing biases that
may affect a single ring due to elemental mobility
by effectively averaging the concentrations over
5 or 10 rings.
A total of 277 samples were obtained with
the following number of samples cut from each
wood strip: 1) 23 samples with average ages 1703
to 1896 A.D., 2) 19 samples with average ages
1706-1896 A.D., 3) 20 samples with average ages
1706-1896 A.D., 4) 20 samples with average ages
1706-1896 A.D., 5) 20 samples with average ages
1706-1896 A.D., 6) 40 samples with average ages
1306-1696 A.D., 7) 30 samples with average ages
1406-1696 A.D., 8) 25 samples with average ages
1456-1696 A.D., 9) 30 samples with average ages
1306-1596 A.D., 10) 30 samples with average ages
1306-1596 A.D., 11) 20 samples with average ages
1406-1596 A.D.
Chemical preparation and analysis
The next step required digestion of the
samples following the methodology described
in Sheppard et al. (2008). Assuming that the
carbon content in the samples is ca. 50% and that
oxidation proceeds according to the reaction 4
HNO3 + C = 4 NO2 + CO2, we calculated the triple
molar excess of nitric acid. For 100 mg of wood,
the required amount of 70% HNO3 is ca. 4.5 g
(or ca. 3.2 ml). All the chemical treatment of the
samples henceforth was done at the Limnological
Institute SB RAS (Irkutsk). The required volume
of 70% nitric acid was added to each wood sample,
and they were left at room temperature for 2.5
days. After this treatment the solutions became
intensely yellow and the wood had the appearance
of an almost transparent sponge, hardly visible in
the solution. The closed tubes with the samples
were then placed in an oven for 3 hours at 70°C.
At the end of each hour, the tubes were taken out
from the oven and placed into the ultrasonic bath
for 15 minutes. It should be pointed out that the
dissolution appeared to be nearly complete after
the first 1 hour in the oven. The tubes were then
cooled to room temperature and the lids on them
were slightly opened to release excess pressure
from the gases that accumulated during the
oxidation reaction. The tubes were then weighed
in order to determine the loss of mass (as gas
emitted). The loss of mass of the reactive solution
was in most cases no more than 4%. Subsequently,
1 ml aliquots of the solutions were placed in 2-ml
polypropylene tubes and were centrifuged for 10
min at 10000 g. An aliquot of 120 ?l was placed
into other pre-weighed 2-ml sample tube, into
which 1.8 ml of deionized water was then added,
and the total mass of solution was determined. A
solution of internal standard, indium, [In]=193
ppb, of 100 ?l volume was added to the solutions.
The total coefficient of dilution after all steps
amounted to ca. 500.
The concentrations of Li, B, Na, Mg, Al, Si,
P, Cl, K, Ca, V, Cr, Mn, Fe, Co, Ni, Cu, Zn, As,
Rb, Sr, Y, Zr, Nb, Mo, Ag, Cd, Sn, Sb, I, Ba, La,
Ce, Nd, W, Au, Pb, Bi, Th, and U were measured
on an Agilent 7500 quadrupole inductively
coupled plasma mass spectrometer (ICP-MS)
with the inert sample inlet system consisting of
polypropylene spray chamber, polypropylene
microflow concentric spray nebulizer (180 ?l/min)
and a platinum injector. A special configuration
of the ICP-MS needed to measure Hg was not
available, and therefore this element was not
measured.
The element Se was also measured, but the
observed concentrations did not significantly
exceed the background values (the concentration
in blank samples), and therefore the measured
values are not reported. In order to calibrate the
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Alexi M. Grachev, Eugene A. Vaganov? Methodology for Development of a 600-Year Tree-Ring Multi-Element Record?
mass spectrometer, we used a multi-element
standard solution ?2A Standard? (concentrations
in the standard were the following: [Ag], [Al],
[As], [Ba], [Be], [Ca], [Cd], [Co], [Cr], [Cs], [Cu],
[Fe], [Ga], [K], [Li], [Mg], [Mn], [Na], [Ni], [Pb],
[Rb], [Se], [Sr], [Tl], [U], [V], and [Zn]= 10.08 ppb,
[In]= 10.07 ppb, [Th]= 10.11 ppb). Additionally a
standard of Lake Baikal water was used (for Na,
Mg, Si, S, Cl, K, Ca), referred hereafter as the
control sample (Suturin et al., 2003). All other
elements were calibrated using the algorithm of
semi-quantitative analysis based on empirically
selected sensitivity factors. The measurements
were run in series as follows: control sample, 12
samples, control sample. Thorough rinsing of the
whole system (3 min) was performed before and
after each control sample. Rinsing was not done
between the samples because of the similarity of
composition of the samples from adjacent wood
pieces. Short rinsing of the autosampler needle
(20 seconds) was done after each measurement.
Metals having concentrations in excess of 0.1
ppb in solutions (or 0.05 ppm referenced to the
initial starting mass of dry wood sample) were
measured with the uncertainty of no more than
30% (Na, Mg, Al, K, Ca, Mn, Fe, Ni, Cu, Zn,
Rb, Sr, Pb, Bi), which is typical for this method.
The manufacturer of the ICP-MS instrument
specifies that the uncertainty for the other
elements (B, Si, P, S, Cl, Ti, V, Co, As, Br, Zr,
Nb, Mo Rh, Pd, Ag, Cd, Sn, I, Hf, W, Au) may
in some instances exceed 30%. All element
concentrations are reported in units of ppb (?g
per kg of dry wood).
Results and Discussion
A total of 11 wood strips from different
trees were analyzed to obtain the 600-year
elemental concentration record. The mean
values and the range of concentrations for each
element during the time interval 1300-1900 A.D.
are given in Table 1. The concentration trends
are shown in Fig. 3. Number of replicate trees
corresponding to each year varied from three to
six, i.e. each datapoint in Fig. 3 is an average of
at least three values obtained independently for
different trees. The large variability of values
among different trees could be an indication
that the microenvironmental effects on growth
of specific trees (or differences in tree behavior
more generally) may influence the level of
element concentrations in tree rings to a greater
extent than the broad changes in environmental
conditions in the region. However, additional
analyses are necessary to check contributing
of possible flaws in our sample protocol to the
discrepancies in concentration series among
trees. In order to establish the nature and
magnitude of variability of trace elements
circumferentially and perhaps longitudinally in
tree rings, a study such as that done for isotopes
by Leavitt and Long (1984), is needed for
microelements in tree rings.
Since the present paper is methodological,
the discussion of the observed concentration
trends, as well as of the sapwood- hardwood
effect, age effect, etc. is outside the realm of this
paper and will be addressed in a subsequent paper
(Panyushkina et al., in prep.). The latter paper
will also report statistical techniques helpful in
interpreting the results presented here.
Conclusions
The longest published record of elements in
tree rings (600 years) was obtained for the time
period of 1300 to 1900 A.D. The set of the elements
that was measured (40) is substantially greater
than any previously reported. Our sample set is
represented by eleven trees, which allowed good
replication of each time interval between 1300 and
1900 A.D. (by 3-6 trees). We found that time series
for most elements generally do not correlate well
from tree to tree because of different individual
physiological responses to environmental factors
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Alexi M. Grachev, Eugene A. Vaganov? Methodology for Development of a 600-Year Tree-Ring Multi-Element Record?
Table 1. Means and ranges (11 trees) of concentrations (ppm or mg per kg of dry wood) of elements in tree rings
of larch from the Taymir Peninsula collected in 2006. Time interval is 1300-1900 A.D
Element
Mean
Minimum
Maximum
Ca
770
440
1200
K
300
220
350
Cl
270
190
360
Mg
250
150
370
Si
140
110
150
Na
42
24
75
P
32
26
39
Mn
14
6.2
20
Fe
13
8.2
62
Ba
12
8.0
15
Sr
4.2
3.4
5.2
B
3.2
2.3
5.1
Zn
3.2
2.5
4.7
Cu
2.4
0.65
8.7
Al
1.7
1.0
3.2
Bi
1.0
0.25
1.8
Ni
0.59
0.21
1.9
Rb
0.35
0.29
0.43
I
0.21
0.10
0.38
Pb
0.21
0.093
3.6
Cr
0.19
0.12
1.7
Li
0.11
0.063
0.42
Ag
0.031
0.014
0.070
Cd
0.015
0.008
0.036
Zr
0.015
0.006
0.041
Sn
0.014
0.006
0.048
and chemical uptake or perhaps because of
shortcomings in the experimental protocol. The
studied tree-ring series show no trend related to
tree age. Our subsequent paper (Panyushkina et
al., in prep.) will contain a detailed discussion of
this dendrochemical dataset based on statistical
treatment.
Acknowledgements
Because the results presented in this paper
would not have been possible without the
initial idea of this work largely contributed by
our friend and colleague Eugene L. Goldberg
(deceased), we dedicate this paper to his
memory.
We thank the U.S. Civilian Research and
Development Foundation and the Russian
Foundation for Basic Research for the support
of this project (CRDF grant RUG1-2950KR-09, RFBR grant 09-04-92515-??_? to
S. W. Leavitt and E. A. Vaganov). This work was
also supported from the Fundamental Research
Program ?23 of the Presidium of the Russian
Academy of Sciences (project ?23.6, principal
# 67 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Fig. 3
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Alexi M. Grachev, Eugene A. Vaganov? Methodology for Development of a 600-Year Tree-Ring Multi-Element Record?
Fig. 3. The average concentrations of elements in tree rings of larch from the Taymir Peninsula collected in 2006
(bars indicate standard deviations of mean values)
investigators E. A. Vaganov and T. V. Khodzher),
and a 1-year grant from the program of the
Russian Ministry of Education and Science
?Development of the Scientific Potential of the
Institutions of Higher Learning? (awarded to the
Siberian Federal University in 2011, principal
investigator A. M. Grachev). We are grateful to
Prof. R. M. Hantemirov (Institute of Ecology
of Plants and Animals UB RAS) and Prof.
N. N. Sushchik (Institute of Biophysics SB RAS)
for reviewing this paper and providing valuable
critical comments.
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Alexi M. Grachev, Eugene A. Vaganov? Methodology for Development of a 600-Year Tree-Ring Multi-Element Record?
??????????? ????????? 600-??????
??????????????? ????????
?? ?????? ???????? ????? ???????????
? ??????????? ??????, ??????
?.?. ???????, ?.?. ????????,
?.?. ???????, ?.?. ??????????,
?.?. ????????, ?.?. ??????, ?.?. ????????,
?.?. ???????, ?.?. ?????, ?.?. ??????????
?
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ?????????, 79
?
??????????? ???????? ???????? ????? ????????,
??????????? ???????,
?????, ??????? 85721, ???
?
??????????????? ???????? ?? ???,
?????? 664033, ???????, ??. ????-?????????, 3
?
???????? ???? ??. ?.?. ???????? ?? ???,
?????? 660036, ??????????, ?????????????, 50/28
???? ??? ??????? ?????????? ?????? ?????? (600 ???) ???????????? 26 ????????? ? ????????
??????? ??????????? ? ??????????? ??????, ?????? ????????? ??????? ? ???? (????? 72° ?.?.),
??? ???????? ???? ????????. ????????? ???????? ??????, ??????????????? ??????????
??????? ? 1300 ?? 1900 ??? ?.?. ??????????? ????????? ???????, ?? ?????? ??? ??????
???????????, ?????????? ?? ??????? ?????? ????? 100 ??, ???????, ??? ???????, ???????? ??
?????? ???????? ????? (?? ? ????????? ??????? ?? ????). ?? ??????? ?????? ???? ???????? ??
19 ?? 40 ???????????????? ????????, ??? ???? ? ????? ????????? 277 ????????. ???????????
??? ??????? ?????????? ????????? ??????????? ?? ???? (??? ???????? 1300-1400 ?.?.?. ?
1600-1700 ?.?.?.) ?? ????? (??? ??????? 1450-1600 ?.?.?.). ????????? ??????? ?????????? ?
????????????????? HNO3 ??? ???????????? ????????? Li, B, Na, Mg, Al, Si, P, Cl, K, Ca, V,
Cr, Mn, Fe, Co, Ni, Cu, Zn, As, Rb, Sr, Y, Zr, Nb, Mo, Ag, Cd, Sn, Sb, I, Ba, La, Ce, Nd, W, Au, Pb,
Bi, Th ? U ??? ?????? ????-????????????? ? ?????????? ????????? ??????? (ICP-MS) ???
?????????. ???????????? ????????? (V, Co, As, Y, Nb, Mo, Sb, La, Ce, Nd, W, Au, Th ? U) ? ?????
??????? ?????????????? ???? ????????? ?? ???????????? ??? ?????????????. ? ??????
??????, ???????? ????? ??????? ???????? ????????? ???????? ????????? ????????????????
? ??????????? ???????????????? ??????, ?????????? ?????????????? ???? ?????????
??????????????? ? ?????????, ? ????? ?????????? ???????????? ? ???????? ???????.
???????? ?????: ???????????, Larix gmelinii, ????????, ???????? ?????? ????????, ??????,
ICP-MS.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2013 6) 73-95
~~~
??? 574.583+57.083.1+577.2
??????????? ?????? ???????????
??????????????? ???????????????? ???????
?????????????????
?.?. ?????????*
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ?????????, 79
???????? ????????? ?? ???
?????? 660036, ??????????, ?????????????, 50/50
Received 03.09.2012, received in revised form 28.01.2013, accepted 22.03.2013
? ?????? ??????????? ??????????? ?????? ??????????? ???????????????? ???????
????????? ????? (????????) ????????????????? ? ?????? ???????????. ????????????
??????, ?????????? ??????? ???????????? ? ????????? ?????, ?? ????????? ??????????
??????????
???????????????
????????????????
???????
?????????
????????
?????????????????. ????????????? ???????? ???????? ??????????????? ?????????
??????????????????? ????????? ? ????????? ??????????? ??????? ? ?????????? ? ??????
????????, ??? ??????????????, ????????? ??????????? ????????????????????, ? ?????
??????????????? ??????????????? ????-?????????????. ????????? ??????????? ?????
??? ?????????? ??????? ????????? ?????????? ???????????????? ??????? ?????????
????? (????????) ????????????????? ? ??????????, ??? ?????????? ??? ????????? ?????
???????????????? ?????? ?????????.
???????? ?????: ????????????????, ???????????????? ???????.
????????
????????? ??????? ???????? ????????
???????? ????????? ? ??????? ?????????
?????????, ?.?. ??????????? ???????? ??????? ? ????????????? ????, ??????? ?????????
?????? ??? ? ??????????. ?? ????????? ????????? ?. ?. ???????????, ?????? ????????????? ??? ? ??????? ????????? ??????????,
?????? ??? ???????? «????????????? ????-
*
??», ?.?. ?????????? ???????????? ???? ???????? ? ??????? ? ??????????? ?? ??? ??????
????????????? ????????, ??????? ??????????? ???????? (??????????, 1978). ? ??????
??????????? ??????????? ???????? ???????
? ?????????????? ???? ????????? ?????
??????????? ? ??????????? ? ??????????,
?????????????? ???????? ? ??? ? ???????
?????????????? ? ??????? ?????????? ??-
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: neverlandia@gmail.com
# 73 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
???????????. ?????? ? ??? ???????? ??????
?????????? ????? ?????? ?????????, ??? ?????????, ???? ?? ????????? ??????? ?????????? ??????????? ?????????? ???????.
? ???????????? ??????????? ????? ??????????, ??? ????????, ???????????? ??????????????????. ????????? ???????? ??????????? ??????? ?????????????? ???????? ?
??????? ??????????? ??????? ???? ??????
??????????? ? ??????????????? ?? ???????
??????????? ??????. ?????? ??? ??????????? ????????? ???????????????? ???? ????
? ??????????? ???????????, ???????? ??????? ???????? ???????? ? ????????????? (????????, 2003). ????? ??????????????? ????
????? ???????????? ???????????, ?????????
?? ??????? ??????????? ?????? ?????????
?? ????? 1-3 % ??????????????? ??????????? ????????, ???????????? ?????? ??????
(???????? ? ??., 1977; Cole, 1982; Amann et
al., 1995). ??? ???? ??????? ????????? ???????, ??? ?????? ?????? ????? ???????????? ???????? ??????????? ?????? ????????
???????? ??????????? ? ???????????????,
??????? ????? ????????? ???????? ????? ???
??? ???????????? ???????? (????????, 2003).
?????? ???? ????????????????? ?????????? ?????????????? ?? ???????????? ??????
(Cases, de Lorenzo, 2002). ????? ???????, ??
????????? ??????? ??????? ?????? ???????????? ??????????? ????? ?????? ????????
????????? ??????????? ???????????.
????????, ??? ???? ?????????? ?????????? ??????? ?????????????? ????????? ?
???????? ???? ??? ?? ???????, ?? ? ?? ???????
??????? ??????????????? ??????? ???????
???? ? ??????????. ? ????? ? ??????????????
????????????? ????????? ????? ????????????? ?????????? ????? ?????? ????????,
???????????????? ???????????? ?????
???????????????? ???????? (Ouverney,
Fuhrmann, 1999). ???????? ????????? ???????
?????????????? ????? ?????? ??????????????? ? ????????? ???????????, ???????????
? ?????????? ???????????? ??????? (?????????, ??????, ???????????, ????????????, ?????????? ??????, ?????????????, ?????????
??????? ? ??.), ? ??????????? ????? ? ??????
????????? (???????? ? ??., 1977; Weiss, Simon,
1999; Grover et al., 2000; Rosenstock, Simon,
2001; Pйrez et al., 2003; Zubkov et al., 2008).
?????? ???? ????????? ????? ? ??????????
???????????? ??????? ????? ?????????? ??????????? ? ????? ? ??????????? ??????? ????????????? ?????????????????. ? ?? ?? ?????
?????? ?????? ????????? ?? ?????????????
??????????? ???????????????? ?????????????? ?????????????????, ????????????
? ??? ??? ???? ???????????????? ????????? (Grover et al., 2000; Pйrez et al., 2003). ????? ??????????? ???????????? ??????????
???????????-???????????? ??????? ??????????? ??????? ?????????????? ????????,
??????????? ? ????????? ??? ???????????.
???????????-???????????? ??????, ????? ??? ?????? ?????????????????? ???? 16S
????????????? ???, ?????????????? ??????????? ??????????????????? (MLST) ? ??????
???????????? ?? ????? ?????????????? ????????? ???????? (MLVA), ?????? ???????????? ??? ????????????? ?????????????????
????????????????? (Hцfle et al., 2008). ? ?????
?????? ????????????? ??????? ????? ??????????????????? ?????, ????????????? ?????? ?????????, ? ??????? ?????? ?? ?????
???????? ? ???????????????? ?????????????? ?????? ?? ???????? (????????, 2003). ???????????? ?????????? ?????????????????
? ??? ????? ??? ???????????????? ???????
? ????????? ??????? ? ??????? ??????. ?????? ???? ???????? ???????? ?????????????????, ??????? ?????????? ???????????
???????????? ??? ????????? ? ??????????
???????? ? ???????????? ????????.
# 74 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
???????? ?? ??????? ?????????????? ??
?????????? ????????: 1) ???? ?????? ???????? ? ???????????????? ?????? (Nystrцm,
2001); 2) ???????????????? ??????????? ???????????? ??????, ??? ??????? ???????
??????????????? ??? ?? ????????? (Amann
et al., 1995; Suzuki et al., 1997). ????? ???????,
???????????????? ???? ????????????????? ?????????? ???? ?? ??????? «???? ??? ??
???????????????» (Yokokawa, Nagata, 2010).
?? ????????? ??????, ????? ????? ????????
?????????? ???????? ? ?????? ????????, ????????? ? ??????? ??? ?????????? ?????? ?
?????? ?????????? ? ??????? ???????????
(Connon, Giovannoni, 2002). ? ????? ??????,
?? ?????? ?. ?. ???????????, ??????????
«????? ???????» ?? ?????? ???? ?????????
???????? ???????? ??????? ? ????????????
????????????? ????????? ?????, ? ?? ?????
??? ??????? ???????? ????? «??????? ???»,
?????????????? ?????????????? ??? ??????? ????? ? ????????????? ????????????
????? (??????????, 1978). ???? ???? ???????
????????, ??? ????????????? ? ?????????????????? ??? ????????????????? ? ???????????? ???????? ?????????? ??????????
????????, ??? ?? ????????, ??? ? ?????????
?????????? ???????, ?? ???????? ?? ??? ???????, ?????? ??? ????????? ?? ?? ???????.
?? ???? ? ?? ?? ????????? ??????? ?????,
???????? ?? ?????????? ???????????? ???????????? ????????, ??? ????? ???????????
??-???????, ? ????????c?? ?? ????, ???? ??
?????? ???? ?????? ????????????? ????, ?
????, ????? ??? ???? (Lawrence et al., 2012).
??????????? ??? ????? ?????? ????????????? ???????????? ???????? ????? ????????????? ???? ??????????, ???????????? ?? ?
???????????? ????????. ???, ?????? ????????, ??? ? ???????????? ???????? ??????????
?????????? ?????? ???????? ????? ???????,
??? ??????????? ???, ? ??? ?????????? ??-
?????????????? ??????? ?????????. ?????
???????, ???? ? ???????????? ???????? ??
???????????? ????? ????????? ???????, ???
?????????????? ?????, ??????????? ?? ???????, ??????????? ? ?????? ??????????????,
?? ??????????????? ?????????????? ???? ??
????????? ?????????? ??????????? ?????????? ? ????????????????? ?????? ??????????????? ? ????????? ??????????? (Maron et
al., 2007).
? ????????? ????? ??????????????????
???????????? ?????????? ? ??????????????
???? ????? ???????. ??-??????, ??????????? ???????????? ?????? ???????????????.
??????????? ????????? ???????? ? ??????
???????? ???????? ??????????? ????????
?? ????????????? ?? ?????? ?????????????????????? ????????? (????????????
???????? ??????, 1997). ??-??????, ??? ????
???????????? ???????????-???????????? ??????, ????????? ??????? ? ??????????????
?????????? ????????????? ????? ?????? ?
??????????????????? ????????????? ????? ?
???????????? ????????? ????????????? ????????? ?? ?????? ????????? ????????????
??????????????? ?????? (Hцfle et al., 2008).
??????????? ????????: ????????????? ??????????????? ????????????? ???????? ???????????? ?? ?? ???????????????? ??????? ?
??????????????? ?? ??????????? ??????, ?
??, ??? ?????????????? ? ????????????????
?? ?????????, ?????????? ??? ???????? ??????????????. ? ????????? ?? ????? ?????
???????????????? ???????? ???????? ???????????? ????????, ?? ?????? ?????????? ??
???????. ?????? ? ??????? ????????? ? ????????? ????????????????? ?????????? ???
??????????????? ?? ??????? ?? ?????????? ??????? ?????????? ????? ? ?????????
???????????????? ?????????, ????????????
? ?????????? (Zehr, 2010). ????? ???????, ?????????? ?????????? ??????, ???????????
# 75 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
?????????? ??????????????? ?????????????
(????????????????) ??????? ? ?????? ??????????? ???????????????? ????????, ???????????????? ?????????? ???????????? ????????. ?????? ???????? ??????? ?????????
?????? ??????.
?????? ?????????????? ?????,
???????????? ? ??????
?????? ?????? ??????? ??????? ??????
?????????????? ????? ? ????????? ?????????? ????? ???????????????????? ??????????,
?.?. ?????????? ?? ????????? ?????????????
??????????? ???????????? ????????????????? ????????? ?? ??? ???? ????????????????
???????.
?????? «?????»
??? ??????? ? ?????? ??????????, ?????????????????, ???????????? ? ?????????? ?
???????????? ? ???????????? ?????????? ??????????? «?????» ?? ????????? ????? ?????
????????. ?????? ??????? ??????? «?????»
?? ???? ????????? ??????? ??????????? ???????????????? ??????? ?????????????????
??????? ? ???, ??? ??? ???? ?????????????
???????????? ??? ???? ?????????????? ?????????, ??????? ???????????? ????? ??????????? ? ?????????? (?aron et al., 2007).
????????????
?? ??????? ??????????? XXI ?. ???????? ??????? ???????????? ????????????
?????, ???????????? ??????????? ???????
???????????????? ??????? ??????????????????? ?????????, ? ???????????? ??????.
???????????? ??????? ???????????? ???????? ???? ?????? ?????? ?????????? (Logue et
al., 2008).
????? ????????????? ??????? ? ????????? ????? ???? ???????????????: ??????
??? ?????????? ?? ????????? ????? ????
? ??????????? ?????????? ???????? ? ??????????? ??????????????? ??? ????????????
?????????? ? ???????????? ????????????
?????????????????? (Nardini, 2010). ??????
?????????????? ?????? ????????? ?????????
????????? ????????? ?? ??????? ? ??????????? ??????? ??????? ???????? ??????????
?????? (Schuster, 2008). ????????? ?????????
?????????????? ?????????? ???????????? ????? ????????? ???????????? ????????????
???????????????????, ????????????? ?????? ?????? ??????????, ?? ??????????????
????? ?????????????? ?? ?????? ? ???????
????? ?????????????????? ?????. ?????? ??
???? ?? «?????»-??????? ?? ???????? ??????????, ????? ????????? ??? ?????????? ?? ???
???? ????????. «?????»-?????? ??????????
???? ????????????? ???????? ? ??????? ?????? ??????????.
???????????? ?????? ??????? ??????????? ??? ????????? ?????????????
??????? ????????????????? ????????????????? ? ??????????. ????????, ? ?????? ????????????????? ? ??????????? ?????? ????
??????? ????, ????????????? ?? ??????????? ????????? ???????? ? ???????????????
???? (Hцfle et al., 2008). ???????????? ????
??????????, ??? ??????????????? ???????????????? ??????? ????????????? ????
??????????? ?????????????? ??????????
?????????????????. ?????? ? ?????????
??????? ??????????? ?????? ??????????
???????????? ???????? ??????????????
????????????? ???????????????????? ?????????? (Mou et al., 2008). ????????, ??????????? ????? ??????????????? ? ???????
??????????? ????????? ?????? ????????????????? ? ???, ??? ?????? ???????? ???????????? ?????????????? ??? ?????????
???????, ?.?. ??? ?????? ????? ??????? ??
?????, ??? ? ???????. ?????? ?????? ???????, ?????????? ??? ???????????????, ???-
# 76 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
?? ????????? ??????? ?? ???? ? ??? ??????????, ? ??????????? ?? ??? ?? ???????????
???????? ????? ?? ????? (Fuhrman, Steele,
2008). ??????? ???? ? ????????? ? ???????
?????????????? ?????????? ?? ????????, ???
?? ??????????????? ? ?????????? ???????
???????? (Maron et al., 2007). ????? ???????,
??????????? ?? ?????? ????? ?????? ???????????? ????????????, ?????? ?????? ??????, ??? ????????? ?????????? ????????????? ????????? ? ?????? ?????????? ???????,
??????????????? ???????????? ?????.
??? ????????????? ??????? ???????? ????? ?????????????????? ???????????.
?????? ?????? ????? ??????????? ?????
??????????? ???? ??? ???????? ??????? ?????? ? ??????????, ? ??????????? ????????,
??????? ????? ?????? ???????? ?????????
???? ? ??????? ???????????? ???????,
???????????? ?? ????????? (Grossart, 2010).
????? ???????, ????????? ????????????? ??????? ? ????? ?????????, ??????? ??
????? ??????????? ????? ????????. ??????
????????, ???? ???????????????, ????????
??????? ????? ???????, ?????????????? ?
??????????? ?????, ?????????? ????????
??????? ? ????? ?????????. ?? ???? ???????????? ?????? ?????? ???? ?????????????????, ??????????? ? ??? ????? ? ??? ????????????? ???????, ?? ????????? ????????
? ?????? ????? ???????? ? ????? ????????
«??????? ?????» ?????????????? ?????????
?????????? ? ?????????????? ????? ???????????????? (Grossart, 2010). ?????????? ?
?????? ???????????? ????????? ? «????????? ?????» ????????????? ??????? (Hцfle et
al., 2008), ???????? ????????? ?????? ??????????????????? ????? ?? ????????? ??????????????? ??????????, ?????????? ? ?????????? ??? ?????? ????? ???????????????
????????? ? ?????????????? ???????????????????? (Kunin et al., 2008).
???????????? ???????????? ????? ???????? ?????????? ??? ?????? ????? ?????.
?????? ?? ?????????? ? ???????? ???????????????????? ??????????, ?????????? ??
????????? ???????????? ??????, ?????????? ???????? ????????? ? ???????????????
???????? ????????? ?????????. ?????????? ? ?????? ??????????? ????????????? ???????: 1) ?? ?????? ??????? ??????? ???????????? ?????????? ??????????????????? ???
?????? ????? ???? ??????????, ???????? ???,
??? ???? ? ?????????? ????????; 2) ?????????? ????????????? ???????????????? ????,
??????? ???????? ????? ? ????????????
????????? (Heidelberg et al., 2010).
???????????????????
???????? ?????????? ?????????? ????
????????????? ??????? ??????????? ? ???,
??? ??????? ???? ?? ??????????????? ? ???
??????????. ????????? ????? «?????» ? ??????????????????? ? ??????? ?????????? ??
???????????? ???, ??? ????????? ?? ??????
?????? ??????????, ? ??? ??????????? ? ????????? ???. ??????? ?????? ???????????? ? ??????? ???????? ??????????????? ??
?????????? ?????, ?????????? ?????????????? ????? (Logue et al., 2008). ????? ???????, ??????????????? ???????????? ?????
?????? ?????????? ? ?????????? ????????,
???????? ?????????????? ??????????.
??? ?? ????? ??? ???????????? ??????????????????? ???????????? ?? ???????????? ??? ???????? ????????????????
??????? ?????????????????. ? ????? ???????????? ????????? ??????? ??????????
????, ????????? ????????? ???????????????? ???? ? ???????? ????????? ??????
?? ????? ??????????, ????????? ????????
???????????? ??????? ????? ? ?????? ??????????, ?????? ?????????? ????? ???????
???? ? ???????? ???????????????? ???-
# 77 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
?? (Poretsky et al., 2005; Maron et al., 2007).
??????? ?????????? ??? ??????? ?????????
????????? ??? ????? ???????. ????? ????????? ??? ? ?????????? ???????????? ????
???????? ???????? ???????????? ????
? ????, ??????? ? ??????????? (Poretsky
et al., 2005). ?????????? ????????? ???????????????????? ??????? ? ???????????
???-??????????, ?? ?? ????????????????
? ??????? ???????????? ?????????? ??????
?????? ?? ???? (Nardini, 2010).
??? ? ?????? ????????????, ??????????
???????????????? ??????? ????????? ????????? ???????? ??????????? ??? ????????????? ????? ??????? ? ???????? ?????????????????, ?? ?? ??????????????? ? ??????????
???? ??????? ? ????????? ???????????.
??????????????
?????????????? ? ???????? ???? ??????
?????????? ?????????? ???????????????.
??? ??? ?????, ?????? ????????, ????????? ?
?????????????? ????????, ??????????????
?????? ???????????? ????? ?????????? ???????? ??? ?????? ??????? ???????????????????? ?????????? (Maron et al., 2007). ????
??????????? ???? ????????????? ????????
?? ????????, ??? ???? ??? ??????????????? ?
?????, ?? ??????? ?????? ????? ??? ?????
? ??????? ??????????? ?????????????????
? ?????????? ?????????? ????????????????
???????. ????? ????????? ?? ?????? ???????? ? ???????????? ?????????? ?????? ?
???? ?????????? ?? ????????????? ???????
????????????????? ????????? ????????? ?
??????? ????-????????????? (Maron et al.,
2007).
????????? ?????? ???????, ??? ?????????????? ???????????? ????????? ???????? ????? ????? ????????????, ???? ??
????????? ??????????? ? ??????? ???????
?????????? (Wilmes, Bond, 2006). ??????, ???
? ??? ?????? «?????»-??????, ?????????????? ?? ????? ???? ?????? ?? ??????: ????? ???
???????? ????????? ?????????? ????????
?????? ???????????? ????? ???? ???????
??????????? ? ?????? ??????? ????????
???????????????? ???????, ????????????
?????????? ? ?????? ?????????, ????????? ? ??????????.
????????, ??? ?? ????????? ? ?????????????? ??????????? ?????????????? ?????????? ? ?? ??????????? ?????????? ????
??? ????? ?????? (Schneider, Riedel, 2010).
??????? ???????????? ? ???????? ????????????? ?? ??????? ????????? ???????????,
????? ??? ?????????? ????????? ??? ??? ?????????? ???????? ?????????? (Wilmes, Bond,
2006). ?? ??????? ??????????????????? ????????? ?????? ???????? ??? ????????????
??????. ???????? ????????? ??????????????? ??????? ? ??? ??????? ?????? ???????
?????????? ?????? ? ????????????? ???????
? ???????? ???????????? ?????????????? ????????????? ?????????. ????????, ? ???????
????????? ???????????? ?????? ?????????
?????? ?????? ? ???????????? ? ??????????
????-????????????? ? ????????? ????????
?????? ?????? ?????????????? ???????
?????????? ????????? ? ??????????? ?????????????? ????????????. ?????? ???? ?
????? ?????? ?????????????? ?? ?????? ??????????????? ??????? ??????????? ??????????????? ???????????????? ???????
?????????????????.
????????????
???????? ?????????? ? ??????? ?????? ???????????, ????????????? ? ?????????, ? ???????? ?????????????? ???? ?
????, ?????????????? ? ?????? (Tang, 2011).
???????????? ????????? ?????? ?????????????? ????????, ?????? ??? ???????????? ????????????? ??????? ?????? ??????
# 78 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
???? ? ??????????? ????????? ?????????????? ???????, ???????????? ? ??????. ?????? ??????????? ?????????? ? ???????
??????? ??????-?????????? ????????? ? ????????????????? (Tang, 2011). ?????? ????????
?????? ??????????? ?????????? ?? ????? ????
??????????? ?????????? ? ??????????????
???? ?????????????????. ???????????? ????? ???? ????? ?? ?????????? ??????????????? ???????????????? ??????? ????????????????? ???? ??? ?????????????? ?????.
???????????? ???????
????????? ??????
??? ??? ??????????, ?????? ?????????????? ???-?????????? (????????????)
?? ???????? ??? ?????? ??????? ?????????
?????? ???????? ?????????? ?????????? ??????, ??? ?? ????????? ??????, ?????? ????
???????? ??????????? ???????????? ? ?????????? ?????????????? ???. ???????????? ??? ????????? ?????? ???????? ?????
???????? ? ??????? ??? ?????????? ???????????? ??????? ????????? ?????? (Nardini,
2010). ????????????? ?????? ????????? ????????????, ????????????????? ??? ??????????? ????????????? ?????????????? ??????. ????? ???????? ??? ?????? ??????
??????? ????????????? ? ?????????? ??
???????????? ??????????????????. ?????
?????????, ??? ?? ?????? ??????? ????????
??? ???????? ??? ?? ????? ?????????????
??????, ? ????? ? ???? ??? ????? ??????????? ???????????? ???????????? ???????????????????.
???????????? ??????? ????????? ?????? ????????? ?????????? ??????????????
???? ?????????? ????? ????????????????
???????? ? ????? ?????????????? ??? ?????????? ? ??????? ??????????? ???????????????? ??????? ?????????????????, ???? ? ??????? ?????? ?????.
?????? ? ?????????
??????????? ???????
????? ???????? ?????? ??????????
???????????????? ??????? ?????????????? ????????????????? ? ???????? ???????????? ???????? ? ?????????? ??? ???? ?
?????????, ????? ???? ??? ??????????. ?????? ?????? ????????, ??? ? ??????? ?????
??????? ? ????????? ???????????? ???????
????? ???????? ?????? ?????????? ? ??????????????? ???????????????? ???????
??????????????? ? ??? ??????, ???? ??????
?????? ???????????? ? ????????? ? ???????
????????????????? ?????????????? (??????????) (Schдfer et al., 2001; Carlson et al.,
2002; Giovannoni, Stingl, 2005; ???????, ????????, 2006; Zubkov et al., 2008). ???? ??????
????????????????? ?????????????? (???)
??????? ? ???, ??? ???????????????? ? ???
????????????? ??? ????? ?????????? ?????????? ????????? ? ???? ????????????
????????????? ???????????????? ???????
??????????? ??????????? ??????????? ??????????? ???????. ????????????? ??????????
? ???????????????? ?????? ?????????? ??????? ??????, ??? ???????????????? ?????????????? ????????, ? ????? ??????????? ????????? ???????????, ???????? ??????????,
? ???????? ????? ?????? ?? ?????????? ?
?????????? ??? ??? ???? ????????????????
??????? ?????????? ????? ?????????????????. ?????? ? ???????? in situ ????? ???????, ??? ???????? ????????? ???? «?????????» ??????? (Hцfle et al., 2008).
???????? ??????????? ???? ?????????
???????? ???????? ? ?????????? ?????????
??? ????, ??? ? ?????????? ?????????????
??????????? ??????? ??????????????? ????
????? ????? ??????????????? ?????????? ?
???????????? ????? ?????????? (White, 1995).
???????? ????????? ? ????????? ?????????? ??????? ?????? ?? ??????????? ???????,
# 79 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
??????? ?????????? ?????????? ????? ????????????? ????????????????. ???????????
?????????? ?????? ???????? ???????????
??????? ?????????? ?????? ??? ??????????
????? ???????? ? ???????????? ???????, ? ??????, ?????? ?????? ?????, ??? ???????? ???????? ???? ????????? ? ? ??????? ????????.
??????????????, ??? ????????? ?????????? ?????????? ?????????? ? ???????????????? ??? ??, ??? ?????????? ?? ? ???????????? ??????? ??? ??????????? ?????????????
(Caron, Countway, 2009). ?????????????????
?????? ???????? ???? ???????????? ??????
?????? ?????????? ??????????, ? ??? ? ????????? ????? ??????????? ??? ???????????.
??? ????????? ? ???? ??????? ??????
???????????? ????????? ? ????????????????
???????????????????? ?????????? ? ??? ?
????????? ??????????? ???????. ???? ?????????? ????????? ???? ??????? ? ??????????, ????? ??? (??? ???? ??????, ? ??????????? ?? ????????????????? ??????????
??????) ????????????????? ????????? ??????????? ????????.
??????? ??????????? ?????????
??????????? ???????? ???????? ??????????? ?????????????????? ?????????
?????????? ???????? ?? ??????? ??????????? ????????? (SIP ? stable isotope probing)
(Radajewski et al., 2000; Hцfle et al., 2008).
?????? ? ???????? ????? ?????????? ??????? 13C ??? 15N ? ??????????, ?? ???????????? ????? ????????????? ? ??????? ?????.
????? ???????? 2H ?? ?????????, ?????????
??????? ????? ????????? ? ?????????????
?????? ?? ?????? ?? ??????????? ???????,
?? ? ?? ???? ????????????? ????? (KreuzerMartin, 2007).
????? ?????????? ? ?????, ??????????
??????? ????????, ????????, ???????????
???, ????? ?????????? ?????????? ????????
??????? ? ????? ???????. ?????? ???????????? ????????? ????? ? ???, ??? ??? ?????????? ?????? ??????, ????? ??? ??????
??????? ???????? ???????, ???? ????????????? ??????????, ????????? (Adamczyk
et al., 2003; Kreuzer-Martin, 2007). ???????
??? ??? ??? ?????????? ?? ????????? ?
??????? ????????????????? ? ?????????
?????????, ? ????? ????????????? ???????
?????? ????????, ??????????? ??????????
?????? (Kreuzer-Martin, 2007). ?????? ???????????? ????????????? ????????? ?????????????????? ??? ??????? ???????-?????????????????? (Gray, Head, 2001).
?????? ???????
?????? ??????? ?? ?????????? ????????? ????? ????? ????????????, ??? ??????
??????????? ?????? (Kreuzer-Martin, 2007).
??? ?????? ???????, ??? ???????, ??? ? ???,
?????????? ?????? ? ??????????? ???????
??????????????. ???? ??????? ??????????
???? ????????? ???????? ? ?????????? 13? ?
?????????. ???????????????? ???????? ???????, ????? ????????????????? ??????? ??
????????? ???????? ???????? ?????? ????????? ?????. ? ????? ???????? ????????? ??????????? ????????????????? ????????????,
?????? ?????????? ???? ????? ?? ??????????? ????, ? ????? ??????????? ??????????
?????????????? ?????????? ????????????
?????. ??????????? ????????????????? ???????????? ? ??????????? ?????????? ?????????????? ?????????? ???????????? ?????
????? ?????????????? ????????? ??????????
?????, ???????????? ??? ?? ????????????? ?
??????? ??????????? ??????. ? ????? ??????
???????????????? ?????? ???????.
???????? ??????, ?????? ?????????? ???????????????? ?????????????? ????????,
??????????? ??????? ????????. ?????????? ?????? ??????? ????? ???????????????
# 80 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
??????? ??????, ? ??????? ?????? ??????
????????? ?????????? ????? ?????????? ?
??????????? ?? ??????? ?????????? ?????.
?????? ???? ?????????? ?????????? ?????????????? ?????????, ? ???????? ???????
???????????????? ????????, ??? ???????,
??????????. ??????????? ????????? ?????
? ?????? ??????? ?????????? ????????????????? ???????????? ? ???? ??? ?????????,
????? ????????????? ??????? ????????????
????? ?????????? ??? ??? ????????? ??????????? ??????????????? ?????? ???????.
?????????? (Kreuzer-Martin, 2007). ?????
?????????? ????????? ??? ????????? ?????
???????? ? ??????? ?? ??????? ??????????
?????. ??? ?????? ?????????? ???????????,
??? ?????? ???????????, ??? ???????????
??????????? ???????? ????????????????
?????????? ?? ?????????? ??????????, ????????? ??? ??????????? ??????? ??????
?????????????????. ??? ??????????, ??????????? ?????????? ??????? ???????, ?????
???????? ??????? ? ????? ???????? ?????? ?
??? ????????, ??????????? ??????? ??????????? ????????.
?????? ???
??????????? ????? ? ??? ???????????????? ? ??????????? ???????, ?????
????????????????? ???????????? ?? ???????? ????????????. ???????????? ????????????? ??????? ??? ??????????? ????????? ??????? ??, ??? ??????????? ????????
????????? ??? ????????? ? ???? ???? ?????
??????? ????????????? ????????? ????? ?????????? (Gray, Head, 2001). ????????????
??????? ?????????? ??????? ??? ? ?????????????? ????????? ??????? ????????? ??????????? ??? ????? ??????????? ?????????
??????? ???????????????? ????????, ??????????? ????????????? ????????????????
???????. ?? ????????? ?????????? ??????
????? ?????????????? ????? ??? ????? ???????, ??? ?????????????? ???????????? in situ
(Kreuzer-Martin, 2007). ????????? ????????????? ????????, ??? ?????? ??????? ? ????????? ??????? ??? ???????????? ???????????
?? ????????? ????????? ? ??????? ???????
??? ??????????? ? ???????? ?????????
(Kreuzer-Martin, 2007). ?????? ???? ??????????? ???????, ??????????? ???????? ??????? ?????????? ??????? (Chauhan, Ogram,
2006). ??????????? ??????? ??????? ??? ???????? ????????????? ????????? ???????????
?????????? ????? ??? ????, ????? ?????????
?????? ???
?????? ???, ???????? ???????????
?????????, ????????? ? ???? ???????????
????????????? ?????????????????, ???????????? ??????? ????????, ? ????????????
?????????????????? ??????? ?????????.
??? ????????? ???????? ? ???? ?????? ????????? ????? ???????, ??? ??? (Adamczyk
et al., 2003). ???????? ????????? ????? ? ???
?????? ????????? ? ??????????? ?? ????????? ????????: ???? ?????????, ????????????
??? ? ??????????? ????????? ???????????.
????????????? ??????? ?????????? ????????? ??? ?????? ????????????????? ????????????. ? ??????????? ??????? ???????
??? ????????? ?? ??????????????.
????? ?????????? ???? ???????, ?????????? ?? ???????? ??????????? ??????? ?
???????? ???????, ??????? ? ???, ??? ???????? ????????? ?? ?????? ???????, ?? ?
????????? ????????? ?????????, ??????????
???? ??????????? ????????????? ??????????
?????, ???????????? ? ??? (Radajewski et
al., 2000). ????? ?????????? ??????????? ???
???????? ?????????? ????????? ?????, ?????????? ????????? ??????? ???????? ? ?????????? ?????????? ?? ????????? ? ???????????? ????????????? ????????? ? ????????????
# 81 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
?????????? ????? ????????? ??? ??????????
?????? ????????? ?????. ?????????? ???????
???????????? ???????????? ???????? ?????
?????????????? ???? ???????????? ??????????, ??????? ? ????????? ???????? ?? ?????
??????????? ??????? ??????????????? (Gray,
Head, 2001). ?????????? ??????? ?????????
????? ????? ???????? ? ??????????? ???????
???????????, ??????? ?????????????? ?????????? ?????????????. ??? ?? ????? ????? ?????????? ????? ????? ?? ??????? ????
?????????? ??????????, ??????????? ???????
????? ????? ????????? ?????????? ???????
????? ?????????? ??????? ??????????. ?????
?????? ???? ???????? ??? ? ???????? ??????????????? ??????? ?????????? ????? 16S
???? ?? ????????? ? ??????? ???????. ?????
?????????? ???????????? ?????????????????? ?????????? ????? 16S ???? ????????,
??????????? ?????. ????????? ??????????
?????????????????? ????? ? ?????? ??????,
???????? ???????????????? ?????? ? ?????????????? ?????????, ??????? ??????????????? ????????????? ??????? ?? ??????????
?????????. ????????? ??????????? ? ????????
???????? ?????????????? ???????? ?????????? ???????????? ????????? ????? ???????
???????????????? ???????????? ???? ??????????? (Gray, Head, 2001).
???????? ?? ????????? ???????????,
????????????? ?????????? ???????? ? ???????? ????? ??? ??????????? ??????????????? ???????????????? ??????? ????????????????? ? ????????? ???????????.
?????????? ?????????????????
? ?????????????????? ???????????
??? ??????????? ????????????????
??????? ?????????????????
? ???????? ????? ??????? ????????????
???????????? ????????? ?????? ????????? ???????? ???????????????? ? ?????-
????????????? ??????????, ??????????
???????????? ?????????????? ??????????
????????????? ??????. ?????? ????????
???????????????? ??????????? ? ????????????????? ??????????????? ? ???????? ?????
???????????? ? ??????????? ???????????
?????????. ????????????? ???????? ?????? ?????????? ????????????????, ??????
??? ????? ????????????? ??????? ??????, ???
?????? ????????????????? ??? ???????? ?
???? ???. ???, ?????????? ????????????????, ??? ?????? ???????? ?????????????
????? ???? ???????? ?? ???, ??????? ?? ???????? ?????? ?????? ???????? (Gray, Head,
2001).
?????????? ?????????????????? ???
????????????? ???????? ?????? ??????????
?????????? ??????????????? ???????????????????? ???????. ??? ????? ???????????? ???????????????????????? ?????????????? ???????? ??? ????????????? ????????
????????? ????????. ????? ???????? ?????????????????? ??? ?????? ???? ??????????? ???????????????? ???????? ? ?????????????? ????????????? ???????, ????????
??????????.
?????????????????? ??????????, ??? ?
????????????????, ????? ???? ???????????
??? ??????????? ? ??????????? ????????????? ???????? ?????? ?????????? ??????????, ??????????? ?? ?????????????????
??????? ??????? (Gray, Head, 2001). ????????????????????? ????? ???????????? ???
???????? ??????? ????????????????? ?? ?????????? ?????????? ?????????? ??????????. ??? ??????????? ?????????? ????????
????????????? ???? ????????????????? ?
???????????? ????????????? ??????????
????????? ?????? ??????. ???????????????
????????? ????????? ???????? ?????????
???????? ??????, ??? ????????? ?????????
?? ?????????????????? ? ????????? ??????-
# 82 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
??? ??? ???????????? ??????? ??????????,
???????????? ??????? ??? ??????? ??? ??????????? ?????????.
?????????? ?????? ? ??????????????
?????????????????? ??????????? ? ????????????????? ??????? ? ???, ??? ?? ????????? ??????????? ?????????? ????????,
?????? ??????????? ?? ?????????? ?????????? ????????? ?????????. ??????????
?????????????????? ??????????? ?????????? ???, ??? ??? ?????????????? ??????? ??
?????????????? ??????? ?????????????? ?
????????? ????????-???????????? ?????.
????????? ???????? ????? ????? ?? ??????????????? ?????????????????? ??????????
??? ?????? ??? ? ????????????? ???????????
??? ?????????? ?????? ??????????????????. ???????????? ???????????? ?????? ?
??????????? ????????????????? ????????
??, ??? ?? ??? ????????????? ????????? ????????? ???????? ??????????, ???? ?????
??? ????????????? ???????. ? ????? ??????
??????? ???????????? ??????????? ???????? ?????? ?? ??????????? ???????????????? ? ?? ??????????????? ??? ?????????????
????????. ????, ?????????? ????????????????? ? ?????????????????? ??????????? ???
??????????? ???????????????? ???????
????????????????? ???????? ????? ????????????? ?? ????????? ? ??????? ?????????? ????????.
??????, ??????????
?? ??????????????
???????????? in situ
?????????????? ???????????? in situ
(FISH) ????????? ????????????????, ?????????? ??????????????? ? ??????????
????????? ????????? ?????? ? ????????
(Schramm, 2003). ?????? ????????????? ?
????????????? ????????? ??????????????????
???????,
????????????????
????????????? ???????? ???? 16S ????,
????? ???? ??????????? ??? ???????????.
?????, ??????????? ??? ???????????? ????????, ????????? ? ????? ?????? (Pernthaler,
Amann, 2005).
?????????? FISH ??? ???????? ????????????????? ????? ??????? ?????????
???????????? ?????????, ????? ??? ??????
??????, ??????? ??????? ????????????? ?
??????????? ??? ???????? ?????? (Schramm,
2003). ????? ????, ???????? ????????? ??????????? ????? ??? ???????????????? ???????????????. ?????? ???????????? FISH
???????? ???????????? ????? ?????? ?? ????????????? ???. ???? ???? ?????? ?????????????, ? ?? ???????????????? ??????????
??????, ??? ? ?????????????? ????? ???
????? ???????????? ?????????? ????????
?????????????? ??????, ????? ??? ?????????
?????????????? ??????? (Gray, Head, 2001).
??? FISH ????? ????????? ????????? ?????,
??????????? ????????????? ??????? ??????
? ??????????? ?????????? ?????? ????????????? ??????????? (Yokokawa, Nagata,
2010). ????? ????, ????????????? ???????????? ????????? ? ????????? ????????, ???
????? ???????? ? ??????????????? ??????
? ?????? ????????? ??????????. ? ???????
FISH ???????? ?? ?????????????? ?? ????, ?
?????????? ?? ?????????????? ? ????? ??????? ????????.
??? ?????????? ???????? ? ???????????????? ?????? ?????? ?? ??????????????
?????? ?????????? ????????? ?????? ??????
??????????? ?????????????? ????????????
???????????? ?????????? CARD-FISH. ?????
CARD-FISH ??????? ?? ?????????? ???????
??????? ????????. ???????? ? ??? ????????? ??????????, ???????????? ??? ???????????? ??????????????? ??????? (Pernthaler
et al., 2002). ???????? ???????????? ???????? ????????????? ? ????? ????????????.
# 83 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
?????????? ?????????????? ??????? ? ?????
???????????? ???????? ? ???????? ?????????? ???????????????? FISH ?? ????????? ?
???????? ???????. ??? ???? ?????????????
?????? ? RING-FISH ? ???????? ?? ?????????? ???????????? ???????? ???????????????? ??????, ??????? ???????????? ? ??????? ???? ?????? ??????? ??????????????????
(Logue et al., 2008).
FISH ?????? ???????????? ??? ????????????? ????? ???????? ????????? ????????? ????????? ? ????????? ? ????????, ???????????? ?????????? ???????????
???????????? ??????????.
?????????? ????????????????????
? ??????????????
????????????? in situ
???????????????????? ????????? ?????????? ??????????? ?????????? ????????
???????, ???????? ?????????????? ????????? 3H, 14C, 33P ??? 35S (Gray, Head, 2001;
Hцfle et al., 2008). ?????????? ???????????????????? ?????????????? ??? ????????
??????????? ????????? ?????????? ??????????????? ?????????? (Andreasen, Nielsen,
1997). ?????????? ?????????? ? ?????????????? ????????????? in situ ?????????
????? ???????????????? ????????????????
????????, ??????????? ??????? ????????
(Ouverney, Fuhrmann, 1999). ????????????
????????? ????????????? ?????????????
????? ????? ???????: ?????????? ???????????????????? ? FISH (MAR-FISH ? MicroFISH) ? ?????????? ????????????? ???????? ??????????????? ? FISH (STAR-FISH)
(Hцfle et al., 2008).
???????? ?????? ???????????? ???????????????????? ? ??? ??????????? ?????????? ????????? ????? ??????? ????????
????????????? ??????????? (Hцfle et al.,
2008). ? ??????????? ?????? ???????????-
????????? ? ?????????? ? FISH ?????????
??? ????????????, ??? ???????????? ?????????? ?????????? ??????????? ????????????? (Adamczyk et al., 2003). ???????
?????????, ?????????????????? ?????????? ?????????????, ????? ???????????????
??????? ??????????? ????????? ???????
????, ??????? ?? ?????????? ?????????????? ??????????? ????????. ????????, ???????????? ??????????, ????? ????????
???????? ?????????????? ?????????, ???
????? ???????? ?????????????? ? ??????
???????. ??????? ????? ????????? ???????, ?? ?? ????????? ???????? ???????????
?????????? ??????????, ?? ?????????????
???????? ??????????? ????????.
? ????????? ??????? ?????????????
?????? ???????????????????? ????????? ?
????????????? ???????. ????????, ?????????? ????????????????????? ???????????
????????? ????????? ? ?????????????? ???????????? in situ (STAR-FISH) ?????????????? ??? ????????? ???????????????? ??????? ??????? ??????????? ????? (Ouverney,
Fuhrman, 2000). ????? ?????????? ?????
3
?-??????? L-??????????? ? ????? ????
????? 60 % ????? ???????? ?????????????
?????, ? ?? ???? ????????? ??? ???? ???????? ? ????????????. ?????? ???????????
???????????? ????????, ??? ????? ???????? ?
???????? ??????????????? (Fuhrman, Steele,
2008). ????????? ???????? ????????? ??????????????? ?????? ?? ?????????? ???????????? ??????? ?????????, ? ?? ? ???????????
????? ??? ???????? ?????? ? ????? ? ?????? ???????? ??? ???????? ???????? ? ???????? ?????????? ???????? (Andreasen, Nielsen,
1997). ??? ?? ????? ????????????????????????? ???????????? ??????????? ?????????
???? ?????? ? ???????????????? ????????
????????? ????????? (Cottrell, Kirchman,
2000).
# 84 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
?????????? ?????????????
??????????????? ????????? ?????,
????????? ?????
? ?????????????? ???????????? in situ
????????? ??????? ?????????? ?????????? ???????? ???????????????? ????????
????? ???? ? ??????? ?????????????????
??????? ? ??????? ??????????? ??????????
FISH, ?????????? ???????? ? ?????????????????? ??????????????? ????????? ????? (?
???????????? ?????????? ?????????? ???????????? Raman-FISH, ?? ??????? ?????? ??
????????????????? ??????? ??????????????? ?????????). ? ????????????? ??????????????? ????????? ????? (?????-?????????????)
???????????? ???????????? ?????, ?????
???????? ???????????? ??????? ?????????? ?????? ? ???????. ?????????? ???? ?????? ???????????? ???????????, ? ??????????
?????? ??????????????? ????????? ? ??????????? ?????? (Neufeld et al., 2007).
????????????? ??????????????? ????????? ????? ????????? ?????????? ????????? ?????????? ?????? ? ????????????? ????????? ????????? ????????? ?????? (Huang et
al., 2007). ????????? ??????????? ??????? 13?
? ????????? ?????? ???????? ????????????
????????? ???????????? ???????????? ??????? ?? ????????? ? ??????? ????????, ???????????? ???????? ? ??????? 12?. ?????????
??????? ????????? ??????? ???????????????
??????????? ????? ??????? ?????, ?????
????????? ??????? ???????? ????????????
????? ????????????? ? ?????????? ??????????? ?????????????? ????????? ?????. ???
??????????? ????????? ????? ???? ???????
??????? ????????? ??-?? ??????????? ?????
??????? ???? ? ??????? ???????????????
????????? ?????. ???????????? ????? ????????, ??? ???????????? ?????? ?? ??????? ??
???? ??????????? ???????? ? ??????????? ?
??????????? ???????? ????? ? ??????. ?????
???????, ?????????????????? ??????????????? ????????? ????? ???????????? ????? ?????????????? ????? (Neufeld et al., 2007).
????? ? ?????????? (Huang et al., 2007)
?????????????????? ??????????? ????????????? Raman-FISH ??? ???????????? ???????????????? ??????? ??????????????? ??
?????? ????? ?????? ? ??????????? ??????,
????????? 13?-??????? ????????. ?? ????????? ? MAR-FISH Raman-FISH ????? ??????
??? ??????? ?????????? ? ?? ????????? ?
?????????? ????????????? ???????? ? ???????? ?????. Raman-FISH ????????? ????????????? ????????????? ????????? ?????????? ???????? ?? ?????? ????? ??????.
???????? ???????????? ??????????
?????-?????????????, ????????? ????? ?
?????????????? ???????????? in situ ??????????? ? ?????????: 1) ????? ??????? ???????????????? ?????????? ?? ????????? ?
MAR-FISH (???????? 1 ???); 2) ???????????
???????????? ?????????? ??????? ? ????????
?????; 3) ????????? ?????????? ? ???, ? ????? ????????? ?????? ?????????? ????????.
? ??????????? ??? ?????????????????
?????? Raman-FISH ?????????? ????????,
???????? ?? ??????? ???????? ?????????,
???????? ??????? ??????????? ?????????
(????????, 15N), ? ??????????, ????? ?? ????????????? ?????? ??? ?????? ???? ??????????? ????????? ? ???????? ?????????.
???? ??????? ???????? ????????? ?? ???????? ??? ???????? ????? ???????????????
? ??????????, ????? ????? ???????? ??? ???????????????? ???????? ? ????????? ?????
(Huang et al., 2007).
? ?????? ? Raman-FISH ????????? ???????????, ????? ??? ???? ????????, ?????????? ?? FISH: ??????????????? ?????? ?????? ???? ???????????? ???????????????
??? ???????????????? ???????? (Neufeld,
Murrell, 2007).
# 85 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
???????????????
???????????????
????-?????????????
?????????? ????-??????????????
? ??????????????
????????????? in situ
????? ?????????????? ??????? ??????????? ????????? ????????? ????? ?
?????? ???????? ??????????????? ??????????????? ????-????????????? (MIMS)
(Hцf le et al., 2008). ??? ????? ?????????
????????? ?????? ????-?????????????
(SIMS), ???????????? ? FISH, ?? ???????
?????? ??????? ? ??????????? ??????-
????-????????????? ????????? ? ??????? ????????????????? ???????? ????????????? ????????????? ???????? ??
????????? (Neufeld et al., 2007). C ???????
????-?????????????? ????? ??????????
????? ????????, ??????????? ?????????????? ????????. ?????? ? ???????? ??? ??????????????? ???????? ????? ?????????????? ???????? ???????????? ???????????
?-?????? (Laniece et al., 1998). ???????????
?????? ???????? ???????????? ?????? ???????????????? ?????????? ????? 10 ???, ???
?? ????????? ????????? ????????? ??????.
?????? ????-????????????? ???????????? ????????? ? ?????????????????????????
??????? ??? ??? ????????, ???????????
??????? ????????????? ???????? ? ???????
???????????.
????? ? ?????????? (Gieseke et al., 2005)
????????? ????????? ???????????????????? ? ????-?????????????? ? FISH ???
??????, ????????????? ?????????????
??????????? (Lechene et al., 2006). ?????
????????? ? ???????????, ???????????
?????????? ???????, ?????????? ????????
? ???????, ??????????? ?????, ????????????? ? ????? ??????? ????????????????
? ???????? ???????????, ? ??????? ????????????????? ? ??????????????????.
?????????? MIMS ????????? ????????????? ?????? ?????????? ? ?????????????
???????? ????????? ?????? ? ??????? ??????????? 50 ??, ????????????? ?????????????????? (Neufeld et al., 2007). MIMS
? 1000 ??? ??????????? ?? ???????????????? ???????????????????? ? ????? ???????? ????? ?????????? ???????? ±1 %.
??? ? Raman-FISH, ?????????? MIMS ?
FISH ????????? ????????? ???????????????? ? ????????? ?????? ?????????? ??????? ? ???? ????????????. ?????? ? ?????????? (Behrens et al., 2008) ??????????
??????????? ??????????? CARD-FISH
? MIMS ? ???????? ?? ???????????? ???
???????????? ???????????????? ???????
???????????????? ????????.
???????????????
???????????????
????-????????????? ???? ?? ?????????
??????? ??? ???????? ????????????????
??????? ?????????????????, ???? ?? ????????? ???????????????? ??? ????????, ??????????? ????????, ?? ????.
???????? ??????????? ?????????? ??????????, ??????????? ?????????. ???????????? ???????????????????? ?????????
?????????? ??????????? ????????? ?????????? ????????, ? ????-?????????????
???????????? ??? ??????????? ??????????
????????????? ???????? ????????. ?????? ?
?????????? (Treude et al., 2007) ??????? ??????????? ?????????????? ?????????????? ?????? ?? ??????? ???? ?????? ? ??????????? ????, ???????? 14?. ?????????? ??????
???????, ??????? ????-?????????????
??? ???????? ?????????? ?????????????????? ???????? ????????, ????????? ?????? ????-????????????? ??? ???????????? ????????????? ???????? ???????? ?
CARD-FISH, ????????? ??????? ????????? ????????? ????????????. ??? ?????-
# 86 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
????????? ????? ? ????????????????????
????-????????????? ??? ???????? ??? ???????????? ?????????? ???????????????
??????????????? (Collins et al., 2007).
??-??
???????
????????????????? ?????????? ? ????????? ????? ????????????????? ???????? ?????? ??? ?????????? ? ?????? ?????????????????
????????? ???????????, ????? ??? ?????????? ?????????, ????????????? ????? ?
?????????? ?????? ???????? ??????????
(Neufeld et al., 2007).
????????? ????
????????????? ?????????? ??? ????????
???????????????? ??????? ???????? ? ????
?? ????? ?????????????????????? ???????, ???????????????? ?? ????????? ????????????? ??????????? (Ward, 2005). ????
???????????? ????? ?????????? ????????,
?? ??????? ???????? ?????????????? ?????
(??????????????? ??? ???-????????) ? ????
????????? ????????? ????? (Logue, 2008).
??????????? ??????? ?????????? ???????? ??? ???????????????? «?????????» ??
???????? 100 ??? ? ???????? 20 ??? (Stahl,
2004). ??? ????????????? ?????????? ????????, ?????????????? ? ?????, ?? ???????? ??????? ?????? ?????? ?????? ????, ??????????????? ????????? ??????????. ?????
??????????????? ??? ??? ??? ?? ?????
?????????????????, ??????? ??????????????
? ?????, ??????????? ???????? ? ?????????
??????, ????????????? ? ?????????? ??????????????????? ?? ???? ? ??????????? ? ??????????????? ??????, ????????? ???????
????????? ?? ??????? ??????? ????? ? ????? ?? ?? ????????? ?? ????. ????? ??????
???????? 16S ???? ??????????? ?????????
???????????? ??????????? ????????? ?????????? ? ??????????? ??????????? ????????? ??? ??????? (Adamczyk et al., 2003).
????? ????????? ????? ????? ??? ???????????. ??? ? ? ?????? ? ???????? ??????? ??????????? ????????? ? ?????????? ???????????????????? ? FISH, ??????
?????? ???????? ????????? ????????????
????????? ?? ???????????????, ???????
????? ?? ????????? ?????? ??? ?? ??????????????? ????????? ????????????. ? ????????? ????????????? ????????? ???????? ??????? ????????????????? ??????????
??????????, ????????? ??? ????????? ????????? ??????????????????, ????????????
????? ??? ?? 12 % (Wilmes, Bond, 2006). ???
????????? ????????, ??? ?????? ??????,
???????? ????????? ???????????????????
????? 16S ????, ?????????? ???????????? ??????????????????, ?????????????
?? ???? ????????. ?????? ? ???????????
????????? ????? ?? ???? ??????????? ??????? ? ??????????? ???????? ??? ????????????? ???????? ?????????? ??? ??????????? ???????. ???????????? ? ?????????
???????? ????????? ??????? ? ?????? ???????????? ???????? ????? ????????????
???????????? ??? ? ??????????. ????????? ?????? ???????????, ??? ? ???????
????????? ?????????? ???????? ????????
??????? ????????? ????????? (Sharkey et
al., 2004).
? ?????? ???????, ? ?????? ?????????
????? ???? ????????? ?????????? ???????????. ? ??????? ?? ??????? ??????? ??????????? ?????? ??????????? ????????? ????????? ???? ????????? ???????? ????????
????????? ????????? ? ??????????? ?????????????, ????????? ???? ??????????? ????????? ?????????????????? ??????????.
?? ????????? ? MAR-FISH ????????? ????
???? ??????????? ????????? ????? ??????
???????????, ??? ???????? ???????, ??? ???
?????? ???????, ??????? ????????????????
??????? ????????? ?????????.
# 87 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
????? ?????????????
(«?????????? ???????»)
?????????????? ????????? ? ???????
?????? ??????? ????????? ? ??????? ????????????? ????????? ? ?????? ???????????
??? ???????? ????????????? ?????????????,
? ??? ????? ?????? (Yan, Yu 2010). ????? ??
????????? ?????????? ?? ???????????? ??????????? ??????? ?????????? ???????? ?????????????????. ???????????? ???????????? ????? ??????? ??????????, ?? ???????
????????? ?????? ??? ????????????? ?????
????????? ????????. ???? ????? ??????????
???????????? ???????? ????????? ?????????
???? ???? ? ?????? ???????????? ????????? ? ??????????, ??????????????, ??? ???
???????? ??????????? ????????? (???????,
????????, 2006).
? ????????? ????? ???????? ????? ???????????? ??? ???????????? ?????? ????????? ???????? ??????????????? ????????
??????????????/????????????? ??????????? ????-???????????? (DGGE/TGGE), ?????????????? ??????????????? ??????????? (SSCP), ????????? ????????????
??????????? ??? (RAPD), ???????????
???? ?????????????? ?????????? (T-RFLP),
?????????????? ?????? ????????????????? ????????????? ??? (ARDRA), ??????
????????????? ????????? ???????????
(RISA) ? ?????????????? ?????? ????????????? ????????? ??????????? (ARISA) (Yan,
Yu 2010). ??? ??? ?????? ????????????? ??????? ? ??????????????? ?????? ???????
????????? ? ???????????? ????????? ??????????? (Logue, 2008). ??????????????????????, ????????? ??????? ????????????? ???????????? ????????? ?????????, ????
??????????? ??????????? ??????????????
????????? ? ?????????? ?? ???????. ?????????? ???????? ??? ???????????? ???????????????? ??????? ????????? ????? ??????-
?? DGGE, TGGE ? SSCP (Nocker et al., 2007),
????????? ????? ???????? ??????? ????, ?????????? ??? ????????? ????? ????????,
??? ??????????? ??????????? ?? ???????????? ??????????????????. ?????????????? ?
?????????????????? ?????, ??? ??? ?????????? ? ??? ??????? ??????? ?????????
?????????? ????????????? ?????????, ????????? ??? (????????????? ????????????
????? ?????? ????????, ??????????? ???????? ???????????????????). ?????? ?? ??????? ??????????, ??????????????? ?????
?????? ????? ????????, ???????? ?? ????????????? ? ??????????????? ??????????
? ????????????? ????? ?????? ????? ?????????????? ????????? (Logue, 2008, Zajec et
al., 2012). ?????????, ??? ?????????? ?? ???
???????????? ???????? ??????? ???? ? ??????????, ? ???-???????????? ????????????? ????????????? ???????? ?????? ?????????? (Brettar et al., 2012).
?????????????? ?????????????????
?????????? ???? 16S ????????????? ??? ?
??????? DGGE ? ???????? ?????? ??? ??????????? ????????? ??????? ???????????? ????? ?????? ????????????? ?????????
? ????????????????? ???????????????? (????????, Schдfer et al., 2001; Ovreеs et al., 2003;
???????, ????????, 2006; ?????????, ???????, 2011; ??????? ? ??., 2012). ????? ????,
?????????????? ????????? ??????? ???????
????????????? ????????? ?? ?????? ?? ??????? ???????????? ???????, ?? ? ?? ????????? ?????? ??????????, ?????????? ? ??????????? ???????? ????? (Brettar et al., 2006;
Brown et al., 2012).
???????????? ???? ??????????? ??????? ?? ?????????? ????????? ???????????????????? ?????????? ?? ????? ????????
??????????? ???????????? ????????. ??????
????????, ??????????? ???????? ???????????? ???????????, ????????? ???????????????
# 88 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
? ????? ????, ?????????? ??? ???????????
??????????. ?? ?????????? ??????? ?????
?? ???????, ??????????????? ????? ? ???????????? ??????? ????????, ?????? ?? ????????? ?????????????? ?????????? ? ????
????????????. ?????????, ?? ????????????
??????????? ????????, ???????? ???????????, ????????? ??????????????? ?? ?????
?? ??????? ?? ??????????. ????????? ????????????? ?????????? DGGE ????????? ?
??????? ????????, ???????????? ????????
????????????? ???????? ????? ??????????,
????????, ???????? ???????????????? ? ???????? ????? (Hamasaki et al., 2007). ??????????? ???????????? ??????????????????
???, ?????????? ?? ????? ??????, ?????????
??????????????? ????????????????? ????????, ???????????? ??? ????????? ?????????????????? ?????? ???? 16S ????. ?????
? ??????? ??? ? ???????? ??????? ?????
?????????? ??????????? ????????? ?????
? ?????????? ?? ?????????? in situ ? ? ????????????????? ????????. ???????? ?????
???????????? ?????? ?????????????? ??? ??????? ????????????????? ?????? ??? MicroFISH. ???????? Micro-FISH ????????? ? ????????????????, ????? ???? ?? ???????????,
??? ????????? ?????????????????, ??????????? ???? ???? ? ??????????, ??????????
??????????? ????????.
??????????
C ????????? ???????????-????????????
??????? ???????????? ????????????????
??????? ????????????????? ????? ?? ?????
???????. ? ???????? ?????????????????, ????????? ????????????????? ??????????????
? ????????? ???????? ??????????????????
???????, ???????? ??? ?????? ????? ?????
(Logue et al., 2008).
?????? ??????????? ??????????????? ???????????????? ??????? ????????-
????????? ????? ??????? ?? ?????????
?????????: 1) ???????????????? ??????????
??????; 2) ??????????? ??????? ??????? ???
????? ????? ? ???????? in situ; 3) ??????????? ??????????? ???????? ?? ???????; 4)
??????????? ??????? ?? ????????? ????????????? ???????????. ????????? ???????????????? ??????????? ???????? ??????
??????????? ??????, ???????? ???????????
?????????, ? ?????? ?????????????. ???????,
???????? ??????? ? in situ, ????????? ???????
??????, ??????????????? ? FISH (????????????????????, Raman-FISH, ??????????????? ??????????? ????-?????????????), ??
??????????? ? ?????????? ??????? ????????? ????????? ??? ???????? ??? ?????????
??????????????????. ????????? ???????
??? ???????? ???????? ?? ??????? ????????
????????????. ??????? ?? ???? ???????????
??????? ????????? ????????? ??????. ???????? ?????????? ??? ???????? ???????????????? ??????? ????????????????? ????????
????????? ?????????? ???????, ???????????
? ??????????? ?? ???????? ? ???? ????????????. ? ?????? ??? ???????? ????, ?????
??? ????????????????? ?????????? ??????
????????, ????? ?????????? ????????? ?
????????????????? ???????????????? ? ???????? ???????????? ????????, ?????????
????????????? ????????, ???????????? ?
??????????????? ??????????????? ????????????????? ? FISH. ???????????? ? ??????????? ???????????????? ????????
???????? ????????? ???????? ????????? ?
??????????. ?????????? MIMS ??????????,
??? ??????????? ???? ???? ????????? ????????????? ?????????? ??????????? ????????. FISH ??????????? ?????????? ??????????? ?????????, ???? ??????? ?? ???????
?????????? ???????? ? ???? ????????????.
???? ????????????????? ?????????? FISH
????? ????????????, ????? ?????????? ???
# 89 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
?????????? ????????? ? ????????? ?????????
??? ????????? ??????? ??????? ? ???????????????? ????, ??????????? ???????????
????????.
?????? ???????????? ???????????????? ???? ????????????????? ????????????? ?? ????????? ??????????? ????????? ?
??????? ?????????? ????????????????????
??????????, ???? ? ????? ?????? ?????? ?????????????? ????? ?????????????? ?? ????????? ????????????? ??????????, ??????? ????? ????????????????, ?????????? ?
?????????? ?? ???????????????? ???????
? ?????? ??????????? (Pernthaler, Amann,
2005). ?? ?????? ?????????? «??????????????????? ??????????» ???????? ??????
????????????? ????????, ??? ???????? ????????????? ???????? ?? ????? ??????????,
??????? ? ???????????? ????????. ??????????????????? ?????????? ????? ????????????? ??? ????????? ???????? ?????????
?????????, ?????????????? ??????? ?????
?????????? ? ???????? ???????.
????????? ?????????? ??????? ?? ???????????? (??????????? ?????????? ????????? ???? ????????? ? ??????????????
???????? ??? ??????????? ??????? ????????), ???????????? (??????????? ?????????? ???????????? ? ???????? ??? ???????
? ???? ????????? ????? ??????????? ???????????) ? ?????????????? ????????????
(?????????? ??????? ??????????? ???????
???????????) (Maron et al., 2007). ????????????????? ??????????????, ?????? ????????? ??? ???????? ? ????????, ???? ???
???? ??????? ? ????????? ????????. ??????????????? ????????????? ?????????????
????????????????? ??????? ? ????????
??????????????? ? ?????????????? ????????????? ????????? ?????????, ???????
?? ?????????? ??????? ???? ??????????
????????????????. ??????????? ??????
??????????? ?????????????? ???? ????????? ???????? ????????????????? ???????
?????? ????????? ??????? ? ?????????
????????, ??????????? ? ????? ????? ???????????? ? ?????????????? ????????????? ? ????????????? ??????? ???????????????? ? ??????????????? ???????????? ?
???????????? ?????????.
???????????? ? ???????????????? ? ????????? ??????????? ?????????? ????????? ?
??????????? FISH ???????? ?????????? ???
?????????? ???????????? ??????????????
????????? ????????? ?????????????????:
????????? ?????????????, ????????????
???????? ???????? ????? ?? ?????? ????????? ? ??????????? ???????????. ???????????? ?????????????? ?????????? ???
?????????? ?????????????? ???????, ????????? ?????????????? ???????? ????????? ???
? ????????? ?????????? ?????? ?????????
(???????????, ????????, 1995). ? ?????????
????? ?? ?????? ??????? ???????? ????????? ??? ????????????? ????????????????
??????????? ??? ?????????????? ?????????? (????????, 1992; ????????, 1999; Cottrell,
Kirchman, 2000; Degermendzhi, 2010), ???
?? ????????? ????????? ?????????? ?????????????? ???? ???????????? ?????. ?
???????-?????????????? ??????? ????????
????????? ??? ?????????????? (????????????????) ???? ????? ? ?????????? ????????????? ? ???? ????????? ???????? ???????????? ???????? ?????????????, ???????
????????????? ???????? ???????????? ???????? ????????????? ???????, ??????????
? ???????? ??????????. ?????????????????
??????????? ???????????? ?????????????
????????????????? ?????? ????????????
?????? ?? ???????????? ???????.
????? ???????, ?????? ??????????????
?????? ? ??????????????? ???????????????? ???????? ????????????????? ? ?????-
# 90 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ??????????? ?????? ??????????? ??????????????? ???????????????? ????????
??? ????? ????? ??????????? ???????????
???????? ???? ????????? ???????????????
???????????????? ????????? ?????? ???-
??????, ???, ? ???? ???????, ???????? ???????? ???????? ???????? ? ?????????? ????????? ????????? ???.
?????? ?????????? ???????? ?-15 ?????????? ???????????? ????????????, ???????????? ? ?????? ???????????????? ??????? ???????????? ??????????? ? ????? ??????????
?????????.
?????? ??????????
1.
2.
3.
4.
5.
6.
7.
8.
9.
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Contemporary Methods for Identification
of Bacterioplankton Species-Specific
Biogeochemical Functions
Olesya V. Kolmakova
Siberian Federal University,
79 Svobodny, Krasnoyarsk, 660041 Russia
Institute of Biophysics of Siberian Branch
of Russian Academy of Sciences,
50 Akademgorodok, Krasnoyarsk, 660036 Russia
Contemporary identification methods of biogeochemical functions of individual bacterioplankton
species (taxa) in aquatic ecosystems are considered in this review. Currently popular metagenomic
methods are not suitable for determination of species-specific biogeochemical functions of various
bacterioplankton taxa. Cultivation of naturally occurring bacterioplankton communities amended
with nutrients is quite promising in combination with such methods as fingerprinting, some
modifications of microautoradiography and multi-isotope imaging mass-spectrometry. Appropriate
method or a combination of methods allows identification of biogeochemical functions of individual
bacterioplankton species (taxa) in the community, which is essential for understanding of principles of
aquatic ecosystems functioning.
Keywords: bacterioplankton, biogeochemical function.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2013 6) 96-107
~~~
??? [577.34:597.08:621.31](28)(477)
????????????? ???????????? ????????
????????????? ? ?????????? ???????? ????????
? ?????-?????????? ?. ??????
?.?. ??????*, ?.?. ?????????,
?.?. ????????????, ?.?. ????????
???????? ????????? ?? ???,
?????? 660036, ??????????, ?????????????, 50/50
Received 22.05.2012, received in revised form 22.06.2012, accepted 29.03.2013
? ?????? ??????????? ?????????? ????????????? ? ?????????? ???????? ???????? ?
??????????? ??????????? ???? ?. ??????, ?????????? ??????????? ???? ???-??????????:
????? (Leuciscus leuciscus baicalensis) ? ??????? (Thymallus arcticus). ?? ?????? ???????
????????????? ?????????? (??) ??????????? ??????????? ???????? ????????????? ?
?????????? ???????? ???????? ? ??????????? ?????: ?????? ??? (Fontinalis antipyretica) ?
????????? (Gammaridea), ?????????-????, ?????????-??????. ? ?????????? ????????, ???
??????????? ??????? ???????? (Fe, Zn, Cu, Mn, Cr, Co, Pb, Cd) ? ??????????? ?????????????
(65Zn ? 137Cs) ? ????? ??? ?? ??? ???????? ??? ???????????? (??<1). ??????????? ???????
? ?????? ??????????? ????? ???????? ?????? ??? ????? (?? = 2.6-3.4) ? ??? ??????????
??????? 40K (?? = 2.6-2.8). ? ?????? ??????? ???????? ??????????? ?????????? K, Zn ? Cd
?? ???????? ???????? (?? = 3.2, 3.0 ? 2.1 ??????????????). ? ???????? ???????? ????????
?????????? ???? ? 65Zn ?? ??????? ??? (?? = 2.9 ? 2.2-8.7 ??????????????). ???????
?????? ????????????? ?????????? ? ????????????? ???????? ? ?????? ??????????? ????
????????????. ????????? ?? ?????????? ? ????????????? ???????? ?????? ? ???? ??
???????? ????????, ??? ? ????????????? ??????????? ????? ????????????? ????????
????????? ?????? ??? ???? ????????????????? K ? 40K. ????????????? ????????
????????????? ?????????-???????? 40K ? 137Cs ? ????????????? ??????????? ?????
??????????? ???????????.
???????? ?????: ????, ??????, ?????????, ?????? ???.
???? ?????? ?????????? ????????????
?????????????? ? ?????????? ???????????
*
?????? ?????-??????????? ????????? (???)
????????, ??????? ?????????? ?? ?????? ??-
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: t_zotina@ibp.ru
# 96 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
???? ????, ? 60 ?? ???? ?. ???????????. ?????
????????? ??????????? ????????? ?? ???
? ?????? 1990-? ??. ?????????? ???????????
????????????? ? ???? ? ????? ?. ?????? ??????????? ????????? (?????, ?????????, 1996,
????????? ? ??., 2008). ?????? ??????? ? ?????????? ??????????? ???????????????? ? ???????? ?????? ??????????, ??????? ???, ? ??
????? ?????? ???????? ? ????????? ????????
?????????? (????????????, ?????????, 2004;
?????? ? ??., 2012). ?? ????? ?????? ??? ?
?????? ?????????? ???? ?????????? ???????
?????????? ???????????? ??????????? ????????????? (Bolsunovsky, Bondareva, 2007). ?
??????? ? ??? ???? ? ??????? ????? ??????
????????? ?????? ?????????????? 60Co, 137Cs,
152
Eu, 241Am, ??????? ???????? (Bolsunovsky,
Bondareva, 2007; ?????????, ????????????,
2008). ??? ???? ???????????? ???? 60Co, 152Eu
? 241Am ????????? ? ???????????? ???????????? ????? (?????????, ????????????, 2008).
????? ???????, ?????? ????????? ?????????
?????????? ????????????? ??? ????? ?. ?????? ? ??????? ??????? ???????.
?????????? ????????? ?????? ??????????? ???? ? ????????? ? ??????? ?????????
?????????. ???? ????? ??????????? ??????????? ???????????? ?? ???? ? ???? (?????,
2004). ???????? ????? ??????? ????? ???
?. ??????, ??????? ??????????? ????????
??? ????????? ? ??????? ? ?????, ????????
?????????. ? ?????? ??????? ??????? ??????
????? ??????? ???????? ?????????? ?. ??????,
?? ?????? ??????? ?????????? ????????? ?
??????? ?????????? (???? ? ??., 2011). ?????
?????? ?????? ???????? ?????? ????????????? ???????????? ???????? ???????????
????????????? ? ????????? ???-??????????
(??????? ? ?????) ? ?. ??????. ????????, ???
?????????? ????????? ???????? ?????? ??-??
?????????? ???????????? ?? ???????????
???????? (???????, 1981). ???????-??????? ?
????????-??????? ?????? ???????????? ???
???????? ????????? ????????????? ? ??????
??????????? (IAEA, 2009; Varga et al., 2009,
Yankovich, 2009). ? ???? ????? ??? ????? ??????? ???????????? ???? ????????? ????????????? ???????? ????????????? ? ??????????
???????? ???????? ? ??????????? ?????, ??????? ? ?????-?????????? ??????.
????????? ? ??????
????? ????? ? ???? ?. ?????? ???????? ?? ???????, ????????????? ?? ?????????? ?? 20 ?? ?? ??? ? ??????????????? 2009
? 2010 ??. ???? ???????? ? ??????????? ??
?????????? ????? 5-10 ? ?? ??????. ??????
??? (Fontinalis antipyretica Hedw), ???????????? ?? ?????? ?? ??? ????, ???????? ?
??????? 1.5?2 ? ??? ?????? ???????????????? ??????? ???????. ??????????????
??????????, ????????, ???????? ? ???????? ??????? ???. ? ?????? ???????? ?????????? ??? Philolimnogammarus viridis Dyb.
???-??????????, ????? (Leuciscus leuciscus
baicalensis Dyb.) ? ??????? (Thymallus arcticus
Pallas), ?????? ?????? ? ??????? ? ???????????? ????. ?????????? ????? ????? ???????
?????????? 124?295 ??, ????? ? 165?222 ??.
????? ????? ??????? ?????????? 102?348 ?,
????? ? 51?99 ?. ??????? ???????? ? ??????,
???????????? ?? ??????? ??????? ?? ?????,
????????? 2+ ? 3+ ?. ????? ???? ???????????????? 61 ???. ??????? ? 55 ???. ?????. ???????
????????????? ?????????? ? ??? ?????????
?? ???????????? ????? (???????????? ? ??.,
2002), ??????????? ??????????? ???????????? ?? ??????? ???????? (????????????,
2002). ?????? ??????? ???????? ???????????, ??? ??????? ????? (???? ? ??., 2011).
??? ????????? ?????????? ????????????? ????? ??????? ??? ? ?????????? ??????????? ????????? ????????????? ????? ? ?????? ??????? ?? ???????, ????? ? ?????????
# 97 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
????? ??? 80 °? ?? ?????????? ?????. ?????
???? ??? ?????????? 200-300 ?, ???????? ? 1370 ?. ???? ??? ????????? ?? ?????? ? ?????,
??? ??????? ???? ????? (?????? ? ??., 2012).
??? ??????????? ??????? ???????????? ???????? ????? (????? ???? ??????) ? ??????
???, ?????? ????? ????????? ????? ???? ?????? ?? 6-11 ???. ????? ????? ??? ???????
? ????? ???????? ???????? (30 %) ? ???????
??????? (????.) ? ????? ??????????????? ??
100 ??.
?????????? ????????????? ? ?????? ????? ???????? ?? ?????-???????????? ?? ??????????? ??????????? ?????????? (Canberra,
???), ??????? ????????????? ? ???????
????????????
???????????
Genie-2000
(Canberra, C??). ???????? ?????????? ????????????? ?? ???? ?????? ????. ???????? ?????????? ????????????? ? ?????? ?????????? ? ??/?? ????? ????? ? ?????? ?????????
?????????? ????? (????. 1).
??? ????????? ???????? ??????????
???????? ????? ???????? ??? ? ??????????
(????????) ????????? ???????????????? ?????, ??? ??????????? ?? ????????? ???????
??????????? ??????. ????? ???? ??????????? ????? ???????? (Millipore, RAWP, ????.
47 ??) ? ???????? ??? 1.2 ???. ??? ????????????? ???? ??? ???????? ????? ? ??????;
?????? ????? ????????? ????? ????? ?????.
??? ????? ????? ?????? ??? 105 °?. ???-
?? ?????????? ? ?????????? ?????? ? ?????
??????? ?????? 1-2 ?. ?????? ????????????
?????????. ????? ???????? ????????????????? ????? ??? ?????????, ??? ??????? ?????
(???????? ? ??., 2009). ???????????? K ?????????? ?? ????????? ????????? FLAPHO-4
(Carl Zeiss, Jena) ? ????????-??????????
??????? (???? 30504 ? 97, ?? 52.24.391-95).
?????????? Fe, Mn, Zn, Cu, Cr, Cd ? Pb ? ??
??????-?????????????
????????????????
«??? ????? 2?» (???? 30692 ? 2000, ???
8288-86). ??????? ??????????? ??? K ?????????? 0.1; ??? Fe, Ni, Pb ? Co ? 0.01; ??? Cr ?
0.006; ??? Cu ? Mn ? 0.003; ??? Zn ? 0.001; ???
Cd ? 0.0003 (??/?). ??????? ????????? ? ????
????????????. ? ???????? ???????? ???????????? ????????? ???????????? ??????????????? ??????????? ??????? (????????? ?????
????????????, ??????). ??????? ?????????
? ??????????????? ????????????? ??????????? ????????? ????????? ?? ??? (????????
???????????? ? ????.RU.???.086.324). ?????????? ????????? ? ??/?? ??? ?/?? ?????
?????.
???????????? ????????? ?????????
??? ???? ???? ? ???????? ????? ?????? ??
????????? ? ?? ????????? ????? ????? (????? ??????????? ??? 105 °?) ? ???????? ? %.
???????????? ?????????? ????????????? ?
??????? ???????? ? ??????????? ????? (??)
???????????? ??? ????????? ???????? ??-
??????? 1. ?????????? ???? ? ???????? ???????????? (?????????, % ?? ????? ?????), ??????? ????. ±
?????. ??????.
???????????
?????????
????? ????
?????? ??? (F.antipyretica)
80.1±1.5
5
????????? (Philolimnogammarus viridis Dyb.)
78.0±1.2
4
?????? (Th. arcticus) ?????
72.5±3.8
20
?????? (Th. arcticus) ??????
70.6±7.4
19
???? (L. l. baicalensis) ?????
74.9±2.3
5
# 98 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
???????? ??? ???????????? ? ???????? ?????????? ? ???????? ?????????? ??? ???????????? ? ???????? ??? ???????? ?????????
(???????). ?? ????? ????? ? ????????? ???
???????????? ??? ????????? ????????????
??? ???????? ?????????? ? ????? ????????
??? ? ???????????? ??? ???????? ??????????
?? ??????? ????? ????. ???????????? ?????????? ???????? ???????????? ??? ???????
???????, ?????????? ?? ???? ?????? ???????????? (?????????????? 2009 ? 2010 ??.).
??????????
?????????? ?????????????
? ???????? ????????????
?? ???? ?????? ????? ?????????????
????????? ?????? 40K, ???????? ??????-
?
??/??
30
100
Ce (?1/2 = 284.89 ???.), 152Eu (?1/2 = 13.5 ???)
? ??????? ???????? ??????? 137Cs (?1/2 = 30.1
???). ????? ??????? ???????? ????????????? ??????????????? ? ???????? ??????? ???
(???. 1?). ? ?????? ???? ??????? ? ????? ??
??????????? ????????????? ?????????? ???????????????? 65Zn ? 137Cs (???. 1?, ?), ? ?????? ?????? ? ?????? 40K (?? 4.3 ??/??).
???????? ?????????? 60Co ? 65Zn ? ???????? ??????? ??? (???. 1?) ?????? 2010 ?.
144
?
100
30
0
Be-7
50
0
20
K-40
10
K-40
10
0
0
Co-60
Zn-65
Cs-137
?
Ce-144
Eu-152
Cr-51
Co-60
Zn-65
?
200
160
??/??
Cs-137
Eu-152
160
200
??/??
100
80
3.0
0
0
2.0
100
80
3.0
00
K-40
1
1.0
K-40
1
2.0
1.0
0.0
0.0
Zn-65
2.5
2.0
1.5
1.0
0.5
0.0
(???. 1). ?? ??????????? ????????????? ?
?????? ????? ?????????? ??????? ? ?????????? ??????????? 51Cr (?1/2 = 27.8 ???.),
54
Mn (?1/2 = 312.3 ???.), 58Co (?1/2 = 70.82 ???.),
60
Co (?1/2 = 5.27 ???), 65Zn (?1/2 = 243.9 ???.),
??/??
50
20
?????? ???????????? ???????? ??????????;
? ?????? ??? ??????????????? ????? 7Be
???.09
Cs-137
???.09
Zn-65
Zn-65
???.09
???.10
Cs-137
???.10
Cs-137
???.10
???. 1. ???????? ?????????? ????????????? (± ?????. ??????. ???????? ????????? ?????) ? ?????? ?????
???????? (??/??) ??????? ??? (?), ???????? (?), ???? ???????? (?) ? ???? ?????? (?), ?????????? ? ?.
?????? ? ??????????????? 2009 ? 2010 ??.
# 99 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
???? ????, ??? ?????? 2009 ?. ???????? ?????????? ??????????? ????????????? ? ???????? ???????? ? ?????? ??? ?????? 2009 ?
2010 ??. ?????????? ???????? ?? ????? ??????
(???. 1?-?). ?????????? 65Zn ? ?????? ?????
?????? 2010 ?. ???? ???? ???????? ??????????? (???. 1?).
???????????? ????????
? ???? ? ???????? ????????????
???????????? ?????????????? ? ?????? ???? (n = 5) ??????????? ? ??????? ??????? (0.6-0.8 ??/? ??? K, 2.6-2.9 ??/? ??? Na,
15-29 ??/? ??? Ca, 2.7-3.7 ??/? ??? Mg), ???
?????????????? (0.01-0.04 ??/? ??? Fe, 0.0030.010 ??/? ??? Zn, 0.0008-0.0015 ??/? ??? Cu,
0.002-0.005 ??/? ??? Mn, 0.00001-0.0007 ??/?
??? Cr ? Pb) (????. 2). ???????????? ????????
? ???? ? ? ??????????? ???? ???? ? ??????
??? ???? ???? ???????? ?????????????? ??????????? (????. 2). ???????????? ?????? ?
?????? ?? ???? ??????, ????? ??????? ???, ??????? ?? ????????? ???????? ?????????????? ???????????. ?????????? ???????? ? ?????? ???? ? ?????, ????????? ? 2009 ? 2010
??., ??????????? ?? ??????????. ??????? ???
??????? ?? ?????????????? ??????? ????????
?? ???? ??????, ??????????? ? ????. 2.
???????????? ??????????
????????????? ? ????????
?????????????
?????? ?? ????????????? ??? ????, ????????? ? 2009 ?., ????????? ? ?????????????? ???????? ??????????? 137Cs ? 120; 60Co ?
4.6; 65Zn ? 3.3 ???/? (????????? ? ??., 2008),
??? ????, ????????? ? 2010 ?. ? 82; 0.8 ? 0.3
???/? ??? ?? ?????????????, ??????????????
(????????????, ????????????, 2012). ???????? ?? ????????????? ? ???????? ???????
??? ?? ???? ?????????? ? ???????? 5000?
12000 ??? 60Co, 900?3200 ? ??? 65Zn, 60?140 ?
???
Cs (???. 2?). ???????? ?? ????????
???? ?? ???? ?????????? ?? 1420 (??? Pb) ??
233 000 (??? Mn) (???. 3?).
? ??????????? ???? ?????? ???????????? ???????? ??????? ?? ??????????
??? 65Zn (2.1-8.7), ?? ????????? ????????????? ?? ????????? ??????? (???. 2?). ?? ???????? ???????? ??????? ???????? ?? ? ????
??????????? ???? ???????? ??? ???? (2.9),
?? ?????? ????????? ???? ?????? ???????
(???. 3?).
????? ??????? ???????? ?? ????????????? ? ?????? ????? ? ??????? ?? ????????? ? ???????? ???????? ???????? ??? 40K
137
(???. 2?, ?). ???????? ?? 40? ? ?????? ??????? ?? ???????? ?? ????????? 0.1. ????????
?? 65Zn ? ?????? ??? ?? ????????? ? ???????? ???????? ?????????? ? ??????? 0.13
? 0.21 ??? ??????? ? ?????, ??????????????,
? ?? 137Cs ? 0.21. ????? ??????? ????????
?? ???????? ? ?????? ? ?????? ??? ?? ????????? ? ???????? ???????? ???????? ???
K (???. 3?). ?? ????? ? ?????? ??? ?? ???????? ???????? ???? ?????? ? ???????. ??
????? ? ?????? ? ?????? ??????? ?? ????????
? 2-3 ???? ????????? ??????? (???. 3?).
??????????
? ?????????? ???????? ? ???????? ??????????? ?? ???? ??? ?? ???????? ?????????
????? ????????????????? ???????? ??, ??????????? ???????. ??? ???????????? ???????? ?? ????????? ???????, ??? ???? ????????????? ??????????. ??????????? ?????
??????????? ???????????? ? ??????? ??????? ??? ?? ????, ??? ? ????, ? ?????? ??????
?????? ?????????. ???????????? ???????? ?
???? ?. ??????, ?????????? ????, ????????
? ???????? ????????????? ???????? ????, ????????? ?? ?????? ???????? ?? ?. ???????????
(???????? ? ??., 2010). ???????????? ??????????? ???????? ? ???????? ????????????
# 100 #
4
5
?????? (??????)
????
0.00065± 0.00011
4.10±0.79
3.31±0.87
4.30±1.46
1.26±0.27
2.58±0.57
K, ?/??
0.023±0.011
64.21±60.94
5.30±1.00
6.80±0.50
60.32±35.24
982±231
Fe, ??/??
0.001±0.000
2.71±0.99
0.39±0.08
0.49±0.13
11.02±2.08
3.79±0.75
Cu, ??/??
* ?.?. ? ???????? ?????? ???????? ?????????????? ???????????.
4
5
?????? (?????)
5
???? (?????)
4
?????????
?????
????
?????? ???
???????????
0.005±0.003
26.29±5.01
7.42±1.48
10.38±2.56
13.02±1.34
15.23±4.18
Zn, ??/??
0.003±0.001
1.97±0.65
0.42±0.36
0.65±0.29
22.82±9.21
593±239
Mn, ??/??
?.?.
?.?.
?.?.
?.?.*
0.13±0.03
1.24±0.18
Co, ??/??
0.052±0.061
0.031±0,023
?.?.
0.054±0.053
0.40±0.06
Pb, ??/??
0.0002± 0.0003 0.0003± 0.0003
0.25±0.13
0.26±0.03
0.22±0.03
0.45±0.11
7.40±2.85
Cr, ??/??
?.?.
0.015±0.018
0.001±0.003
0.005±0.005
0.008±0.006
0.04±0.01
Cd, ??/??
??????? 2. ?????????? ???????? ? ?????? ????? ???????? ???????????? ? ???? (??????? ????. ± ?????. ??????.) ?. ??????, ?????????? ? ???????-??????? 2009
? 2010 ??.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
?. ??????, ????????????? ????, ??????????
????????????? (????. 2). ????????? ?????? ??????? ? ?????????????. ? ?????? 2010 ?. ????????? ????????? ??????????? ????????????
?? ???, ????? ???? ?????????? ???????????
??????????? ????????????? (????? ??? 60Co ?
65
Zn) ? ???? ?. ?????? ??????????? ?????????
(????????????, ????????????, 2012), ?????????????, ???????? ????? ????????????? ? ???????? ???????????? ?? ????.
?
??
?
??
100000
10
10000
8
1000
6
100
4
10
2
0
1
Co-60
Zn-65
?
??
K-40
Cs-137
Zn-65
Co-60
4
3
3
2
2
1
1
0
Eu-152
?
??
4
Cs-137
0
K-40
ZN-65
CS-137
K-40
ZN-65
CS-137
34012
???.09
???.09
K-40
???.09
???.10 ZN-65 ???.10
???.10
??.????.±?????.??????.
CS-137
???. 2. ???????????? ?????????? ????????????? ? ???????? ??????? ??? ?? ???? (?) ? ? ???????????
????? ?????? ????????????? (?), ???????????????? (?????) (?), ?????????????? (?????) (?)
?
??
?
??
3.5
1000000
1
3.0
100000
2
3
4
2.5
2.0
10000
1.5
1.0
1000
0.5
100
K
Fe
Zn
Cu
Mn
Cr
0.0
Pb
K
Fe
Zn
Cu
Mn
Cr
Co
Pb
Cd
???. 3. ???????????? ?????????? ???????? ? ???????? ??????? ??? ?? ???? (?) ? ? ??????????? ?????
(?): 1 ? ?????? ?????????????, 2 ? ???????????????? (?????), 3 ? ?????????????? (?????), 4 ?
???????????????? (??????)
# 102 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
?? ????????????? ???? ?????????? ?????? ??? ??????????? ??????????? ???????????? ? ??????? ???????????? ?????????????????? ??????? ?? ????. ????? ?????????
??????? ????? ?????? ????????? ???????????
?????? ?? ??? ? ?????? ???????? ???????
??? (???????????????? 2010 ?.) ?? ?????????? ?????????????? ????????, ????? ??? 24Na
(?1/2 = 14.96 ?.), 51Cr ? ??., ? ????? ???????????
????????? ?????????? ?????? ???????? ? ?????????? ???????????. ???, ?????????? 60Co
? 65Zn ? ???????? ??? ????????? ? ?????????
??????? 2010 ?. ? 3-4 ???? ?? ????????? ? ??????????? ???????? 2009 ?. (???. 1?). ?????????? ????????????? ? ???????? ? ????????
??? ?? ???? ????? ??????????, ? ??? ??????????????? ??????? ???????? ?? (???. 2?,
3?). ????????? ??????????? ?????????? ??????????? ??????? ??????? ? ????????????
?? ????, ?????? ??? ?????? ????????????
??? ????????? ???????????? ??????????? ?????????.
?????????? 65Zn ? ???????? ????????
?????? 2010 ?. ?? ??????????? ?? ?????????
? ?????? ??????????? ???? (???. 1?), ?????????? 60Co ????????? ? ??????? ???????, ???
??? ??????? ???, ? ??????? ? ??????? ????. ????? ???????, ????? ?????????????? ????? ??
?????????? ???????? ?????????? ?????????
?? ????????? ? ??????? ?? ??????????, ???,
????????, ??????????????? ? ????????????
???????? ???? ??????????? ???? ????????
? ????????? ???????? ??? ??????. ????????, ? ?????????? ??????? ??????? ????????
?????????? 65Zn ? ?????????? ?? ???? ??????? ???????. ??????? ???????? ????????
? ?. ?????? ??????, ? ????????, ?????????,
? ????? ???????? ??????????. ? ???????
?????????????? ???????? ???? ????????,
??? ?????????, ????????? ? ??????, ?????
??????????? ? ???? ????? ???????? ???????
??? (Kalachova et al., 2011). ?????????? 65Zn
??????????? ?? ???????? ??????? ??? ?????
?????????? (???. 2?). ??? ???? ???????? ??
65
Zn ? ???????? ???????? ???????? ????? ????????? ???????? ?? ???? ???????? ??????????
????? ??????? ? ?????? ???. ?? ??????????
???????? ???????? ? ??????????? ???? ?????? ????????????? ???????? ???????????
?????????? ???? (???. 3?). ????? ??????????
? ??????????? ???????????? ??????????
???? ??????????? ?? ?????????? ?. ??????
(???????? ? ??., 2009). ??????? ?????? ??????????? ? ????????? ????????????? ? ???????? ? ???? ???????? ?? ???????? ???????
??? ????????????.
????????? ???????? ???????? ??????????? ??????? ??????? ? ?????-???????
?????? (???? ? ??., 2011). ??? ??????????? ?
?????????? ???????? ????????????? ????
???????? (???? ? ??., 2011). ????????????
?????? ??????? ??????? ?????? ????? ???????? ??? ??????? ????? ? ???????? ???????
?????? ? ???????? ? ??????? ??????. ???????? ???????????? ????????????? ?????????? ? ???????? ?????? ???? ????????? ????
(1.8-3.7), ? ???????????? ??????????? (1.81.9) ? ??????? ????, ??? ? ????????. ?????????? ??????????? ????????????? ? ??????
???? ???? ????? ??? ???? ????????. ? ?????? ???????? ?????????? 65Zn ?? ?????????
??????? ??????????? ? ??????? ? ????????,
?? ???? ???????????????? ? ??????? 2010 ?.
(???. 1?), ??? ?? ????? ???? ????????? ?????? ??????? ?? ????????? ? ???????????
????? ????????, ?.?. ?????? ???? ? ????????
???????? ? ??????????????? 2010 ?. ??? ???????. ????? ?? ???????? ???????????? ???????? ? ????????, ???????????? ?????????????
?????????? (3-4) ? ??????????? (1.4-1.6) ???????? ? ????????????? ????????.
???? ?????? ????????, ??? ? ??????
???-?????????? ???????? ??????????? ?????????? ????? ? 40K ?? ????????. ?????? ??-
# 103 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
???? (???????? ? ??., 2009) ????? ????????
??????????? ?????????? ????? ? ?????? ??????? ?? ???????? ? ?????? ??????????????
?????????? ??????. ???? ?? ????????? ??,
?????????? ??????????? ????????????? ?
?????? ???-?????????? ?? ???????? ???????? ????????????. ?????????? ???????????
??????????? ????????????? ? ????? ??????? ?????? ???????? ????? ???? ?????? ??????? ??????? ???? ????. ???, ????????, ???
??????? ????? ????????? ? ???? ??? ?? ????
????? ????? ? ??????????????? ????? (?????, 2002).
???????????? ???? ?????????? ????????, ??????????? ? ????? ??????????? ???
????????????????? ???, ? ?????? ???????
????????? ???????????? ? ?????? ? 8.7
??? ??? ??????, 6.9 ??? ??? ????, ? 4.7 ??? ???
????????, ? 3.5 ??? ??? ?????. ????? ???????
???????????? ?????????????? ???????? ?
?????? ?? ????????? ? ??????? ??????????
??? ?????? ????? ???????????? ??? ??????? ???????? (???????, ?????????, 1998; ????????? ? ??., 1998; ?????, 2002; ???? ? ??.,
2009). ????? ???????, ??????? ????? ?????????? ????????, ???????? ?? ????? ? ?????? ???, ????????? ?? ?????????. ? ??????
??????? ???????? ??????????? ??????????
?????, ????? ? ?????? ?? ???????? ????????
(???. 3?). ??????? ?????? ???????? ? ??????
??? ??????????????. ???????????? ? ?????? ??? ??? ?????????? ?? ???????, ???? ?
????? ???? ? ?????? 2010 ?. ??? ???? ???????????????? ?? ?????????? ??????? ????????
?????? (?????? ? ??., 2012).
????????????? ?? ????????????? ????????? ????????????? ? ?????????? ???????? ? ?????????? ???????? ?? ?? ??????????
????????, ???? ???????? ????? ???????, ???
???????????????? ???????? (???????, 1981).
? ????????????? ???? ??????????? ?????
????????????? ?????????? ????? ????????
? ?????????????? ?????????? 40K, ??????? ?
????????? ????? ???????? ?????????? 0.012 %.
????? ????????? ????? ????????, ?????????
??????????? ???????? ????? ???????? ? ??????? ?????????. ?????????? ????? ? ???????? ? ??????? ???? ?????? ????????????? ?
????????? ??? ????? ??????????, ??? ?????????? ????????????? ???????? 60Co ? 65Zn.
? ??????????? ????? ????????????????
(?????) ? ?????????????? (?????), ????????, ?????????? ????? ????? ??????????, ???
??????? 65Zn (???. 2?, ?; ???. 3?). ? ???????
?? ?????????????? ?????????? ?????? ????? ????? ?????????? ????????????? ?? ???
???????? ? ?????? ??????? ? ????? ?????????? ? ? ?????? (???. 3?). ? ????? ?????
??????????????, ??? ??????????? ???????
????????????? ?????????? ????????, ? ????? ?? ????????? ????????????? ???????? ?
????? ???-?????????? ?. ?????? ?? ???????? ??????????????.
? ????????????? K ? Cs ??????????????? ??? ????????-??????? (IAEA, 2009; Varga
et al., 2009). ???? ?????? ????????, ??? ???????????? ???? ?????????? (?? 70 %) 40K ?
137
Cs ? ????? ???????? ????????? ? ??????
(?????? ? ??., 2012). ??????? ???????? ?????????? ??? ???????????? ??? ??????? ???????? (?????? ? ??., 2008; Yankovich, 2009).
????? ???????, ????? ?????????????? ? ????????? ??? ??? ??, ??? ??? ?????? ? ?????.
?????? ????????????? ?????????? 137Cs ?
?????? ???-?????????? (???????? ? ??????)
?? ?????????? ? ?. ?????? ???? ???????????
??????, ??? 40K ? K. ????????, ??? ????????????? ?????????? ?????????-???????? ???????????? ??????????????? ???????????????????? ? ????? (Yankovich et al., 2009). ??
?????? ??????? ??? ?? ?????? ????????????
????????? ???????????, ??? ?????????? 137Cs
????????????? ?????? ???????????? ??????????? ? ????????????? K+ ? ??????????
# 104 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
????? (Rowan, Rasmussen, 1994). ????????,
??? ???????????? ????????????? ??? ????????????? ????? ? ?. ?????? ???????????
????????? ???????????? ?????.
?????? ?????? ???????? ???????????
????????????? ?????????? 137Cs ????????????? ?????? ?? ???? ?? ???????. ? ?????????, ????????, ??? ????? ??????? ????????
?????????? 137Cs ?????????????? ? ??????
(??????) ?????? ????? ??? (?????, 2004;
???????? ? ??., 2009; ??????? ? ??., 2009).
?? ????? ???????? ??????????? ???????????? ???????????? ???????? 137Cs ? ?????
???? ?? ??? ???????? ? ?????? ? ?. ??????
(????????? ? ??., 2012). ?????????????, ? ????? ???-?????????? 137?s ??????????? ?????
????????? ????? ??? ?????? ???????????
???????.
??????????
?????? ???????? ??????????? ????????????? ? ???????????? ???????? ? ??????????? ??????????? ????? ?. ??????, ??????? ? ?????-?????????? L. l. baicalensis,
Th. arcticus, ???????, ??? ??????????? ??????? ???????? (Fe, Zn, Cu, Mn, Cr, Co, Pb, Cd)
? ??????????? ?????????????, ?????????????? ? ?????? ??? (65Zn ? 137Cs), ?? ?????? ??
???????? ??????????? ??????? ??? ? ???????? ? ???????????? (??<1). ???????????
?????????? ? ?????? ??????????? ?????
???????? ?????? ??? ????? ? ??? ??????????
?????????????? ??????? 40K. ? ?????? ??????? ???????? ??????????? ?????????? ?????,
????? ? ?????? ?? ???????? ????????. ? ???????? ???????? ???????? ?????????? ????
? ???????????? ????? (65Zn) ?? ??????? ???,
??????? ?????? ????????????? ?????????? ?
????????????? ???????? ? ?????? ??????????? ???? ????????????.
????????? ?? ?????????? ? ????????????? ???????? ????????, ??? ? ?????????????
??????????? ????? ????????????? ?????????? ????????? ?????? ??? ???? ?????????
???????? K ? 40K. ????????????? ??????????
???????? ?????????-???????? 40K ? 137Cs ? ????????????? ??????????? ????? ?? ??????????? ??????????? ?????????.
?????? ??????????
1.
2.
3.
4.
5.
???????????? ?.?., ???????????? ?.?. (2012) ???????????? ? ???? ?. ??????. ?: ?.?.
???????? ? ??. (???.) ????????????? XXI ????: ????????? ????????????? ?????????????????? ???????????, ??????????, 14-16 ??? 2012 ?., 190-194 c.
???????? ?.?., ???????? ?.?., ??????? ?.?., ????? ?.?., ?????????? ?.?. (2009) ????????????? ? ???????? ???????? ? ??????????? ????? ?????????? ???? ?????? ? ??????
?. ???????????. ?????? ??????? 36 (5): 623?632.
???????? ?.?., ???????? ?.?., ??????? ?.?., ???????? ?.?., ?????????? ?.?. (2010) ?????? ?????????????? ??????????? ?. ?????? ?? ?????????? ???????? ? ???????? ??????????? ?????????? ?? ????????, ????????????? ???? ? ???? ?. ???????????. Journal of
Siberian Federal University, Biology 3(1): 82-98.
???? ?.?., ???????? ?.?., ?????? ?.?., ???????? ?.?. (2009) ??????? ??????????? ????????????? ??????? ???????? ? ????????? ??? ????????? ???????. ?????????. ????. 45 (3):
55?64.
???????????? ?.?., ????????? ?.?. (2004) ????????????? ??????????? ?????? ??????????
???? ?????? ? ???? ??????? ????????????????? ?????????. ??????. ????????. ????????????? 44 (3): 361?366.
# 105 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
6.
7.
8.
????????? ?.?., ???????????? ?.?. (2008) ???????? ???? ?????????? ??????????? ????????????? 60Co, 137Cs, 152Eu ? 241Am ? ?????? ?????????? ?. ??????. ?????????? 50 (5):
475?480.
????????? ?.?., ???????????? ?.?., ??????????? ?.?., ??????????? ?.?. (2008) ????????????? ????? ???????? ???????????????? ????????????? ????????? ? ?????? ???? ????
??????. ????. ?? 423 (4): 479? 482.
???????????? ?.?., ???????? ?.?., ?????? ?.?., ???? ?.?. (2002) ????????? ?? ??????????. ??????????: ??????, 127 ?.
9. ???????????? ?.?. (2002) ??????? ??????? ?????????. ??????????: ??????, 320 ?.
10. ??????? ?.?. (1981) ??????? ?????????????? ???????????????? ????????. ?.: ?????,
247 ?.
11. ?????? ?.?., ?????? ?.?., ??????? ?.?., ?????? ?.?. (2008) ???????? ?????????? ? ????????????? ???????? ?????????????? ????????????? ? ??? ???? ?????????? ????????????? ???. ?????????. ????. 44 (3): 95?113.
12. ??????? ?.?., ????? ?.?., ?????? ?.?. (2009) ??????????? ?????????? 137Cs ? ????????? ????? ??? ?????????? ????????????? ?? ??????????? ?????. ?????????. ????. 45 (5):
110?116.
13. ?????? ?.?., ????????? ?.?., ???????????? ?.?. (2012) ???????????? ? ??????? ????????? ?? ???????????-???????????? ??????? ???????? ??????? ?. ??????. ??????. ????????.
????????????? 52 (3): 305-311.
14. ???? ?.?., ???????? ?.?., ???????? ?.?., ?????? ?.?., ????????? ?.?., ?????? ?.?.,
15.
16.
17.
18.
19.
20.
21.
22.
?????? ?.?. (2011) ??????? ??????? Tymallus sp. ? ??????? ??????? ?. ??????. Journal of
Siberian Federal University, Biology 4 (3): 281?292.
??????? ?.?., ????????? ?.?. (1998) ??????????? ????????? ????????????? ??? ???????
??????????? ??????? ????????. ?????? ??????? 25 (5): 603?610.
????? ?.?., ????????? ?.?. (1996) ?????? ???????????? ?????????? ?? ?. ?????? ?????
?????? ? ???????????? ??????????? ????????? ????????????? ???. ??????? ??????? 81
(3): 226?232.
????????? ?.?., ??????? ?.?., ????????-?????? ?.?., ?????? ?.?. (1998) ??????? ??????? (Zn, Fe, Cu, Mn, Pb) ? ?????? ? ??????? ???????????? ?????. ???. ????. ????. 5:
477?483.
???????? ?.?., ?????????? ?.?., ?????? ?.?., ????? ?.?. (2009) ???????? ??????????
137
Cs ? 40K ? ?????? ??????????? ????? ??? ??? ????? ? ????? ? 1995-2007 ??. ??????.
????????. ????????????? 49 (6): 721?728.
????? ?.?. (2002) ?????? ?????????????? ????????? ???????? ???????? ??????????????.
???????????: ???????. ???. ??-?, 270 ?.
????? ?.?. (2004) ????????????? ??? ???????? ? ???? ??????? ?????? ?? ?????????????
???. ?.: ???????????? ??????? ??????? ???, 215 ?.
????????? ?.?., ?????? ?.?., ???????????? ?.?. (2012) ?????? ???????? ???????????
????????????? ? ??????????? ????? ???? ??????. ???. ????. ????. 4: 497-504.
Bolsunovsky A., Bondareva L. (2007) Actinides and other radionuclides in sediments and
submerged plants of the Yenisei River. J. Alloy. Compd. 444-445: 495?499.
# 106 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
23. IAEA (International Atomic Energy Authority) (2009) Quantification of radionuclide transfer in
terrestrial and freshwater environments for radioecological assessments. TECDOC-1616. Vienna:
IAEA, 622 p.
24. Kalachova G.S., Gladyshev M.I., Sushchik N.N., Makhutova O.N. (2011) Water moss as a food
item of the zoobenthos in the Yenisei River. Cent. Eur. J. Biol. 6 (2): 236?245.
25. Rowan D.J., Rasmussen J.B. (1994) Bioaccumulation of radiocesium by fish ? the influence of
physicochemical factors and trophic structure. Can. J. Fish Aquat. Sci. 51 (11): 2388-2410.
26. Varga B., Leclerc E., Zadyvai P. (2009) The role of analogues in radioecology. J. Environ.
Radioactiv. 100: 802?805.
27. Yankovich T.L. (2009) Mass balance approach to estimating radionuclide loads and concentrations
in edible fish tissues using stable analogues. J. Environ. Radioactiv. 100: 795?801.
Efficacy of Trophic Transfer of Radioactive
and Stable Isotopes of Metals
to Zoobenthos-Feeding Fish of the Yenisei River
Tatiana A. Zotina, Elena A. Trofimova,
Alexandr Ya. Bolsunovsky and Olesia V. Anishenko
Institute of Biophysics SB RAS,
50/50 Akademgorodok, Krasnoyarsk, 660036 Russia
Accumulation of radioactive and stable isotopes of metals in trophic chain of the Yenisei River leading
to zoobenthivorous fish species, dace (Leuciscus leuciscus baicalensis) and grayling (Thymallus
arcticus), have been investigated. Concentration factors (CF) of radionuclides and metals in trophic
pairs, aquatic moss (Fontinalis antipyretica) ? gammarids (Gammaridea), gammarids ? dace and
gammarids ? grayling, were estimated. Dietary transfer of metals (Fe, Zn, Cu, Mn, Cr, Co, Pb and
Cd) and artificial radionuclides (65Zn and 137Cs) from gammarids to muscles of fish species was not
effective (CF<1). Effective transfer of potassium (CF = 2.6-3.4) and its natural isotope 40K (CF = 2.62.8) is possible in these trophic pairs. Effective accumulation of K, Zn and Cd in liver of grayling is
possible from biomass of gammarids (CF = 3.2, 3.0 and 2.1, respectively). Accumulation of copper
and 65Zn is possible in biomass of gammarids from aquatic moss (CF = 2.9 and 2.2-8.7, respectively).
Dietary transfer of other stable and radioactive isotopes of metals in this trophic pair is not effective.
Efficacy of dietary transfer was similar for analogue isotopes K and 40K only. Efficacy of diet transfer of
isotopes of analogue elements, 40K and 137Cs, differed considerably among the studied trophic pairs.
Keywords: aquatic moss, dace, gammarids, grayling, Fontinalis antipyretica, Gammaridea, Leuciscus
leuciscus baicalensis, Thymallus arcticus.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2013 6) 108-111
~~~
???????? ?? ??????????
?.?. ????????? «????? ? ??????????.
?????? ?? ????? ?????????????».
????: ???????????????, 2011. ? 704 ?.
?.?. ????????, ?.?. ?????????*
???????? ????????? ?? ???,
?????? 660036, ??????????, ?????????????, 50/50
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ?????????, 79
Received 08.03.2013, received in revised form 15.03.2013, accepted 22.03.2013
????? ?.?. ????????? «????? ? ??????????. ?????? ?? ????? ?????????????» ??????? ?????????? ???????? ?????????????
????????? ? ?????????????, ????????, ?????????, ?????????? ? ?????????????? ????????????? ??????????????. ??-??????, ??????
??? ????????????? ???????? ?????????? ??
?????????? ???????????? ??????? ? ?????????? ?? ????? ????????????? ? ????????????? ? ????? ????? ????? ?????????. ????????, ?????? ??? ????? ???????????? ?????
??????????????? ????, ?????????? ???????????, ?????? ?? ?????? ???????????? (????????, ????????????? ????????? ?????? ??
?.?. ???????? ? ?.?. ?????????), ?? ? ??????????? (????????, ??????????????? ? ??????? ?????????? ? ??????????). ? ????? ?????
??????????? ???????? ????????????? ??????????, ????????, ?????? ? ?????????????
????????????? ????? ????????, ??? ?????????????? ? ????????????. ? ?????????????
????? ??????????????? ?????????????? ??
?????? ?????? ?? ?????????? 2000-2010 ??. ?
*
?????? ?????????? ?????????? (????? 800
??????????!), ? ??????? ?????? ?? ??? ????
?????????? ????? 31 % (?? ??? ?????????? ?
????? 10 %). ??? ??????, ??? ??? ? ????? ????? ??????? ????? ?????? ?????????? ?????? ?? ???????????? ??????, ??????? ? ?????
XIX ????, ??? ?? ? ??? ????? ?????. ?????
???????? ????????, ????????? ? ?????????,
??????????? ??????????, ??? ??? ?????????? ???????? ? ?????????? ????????? (??????? 1.1, ??????????? ???????? ???????
?????? ???????? ? ???????? (?. 20)). ? ?????
????? ???????? ??????? ????????? ? ??????? ??????????????? ???????.
? ????? 1 ????? ???????? ???????? ????
???????????? ??????? ????? ????????????? ?
????? ?? ? ??????? ???????????? (?????????
? ??????? ??????? ????????? ????). ? ?????
2 ??? ????? ?????????? ?? ??????? ?????????? ? ????? ? ???, ? ?????????, ??????????? ????????????? ? ???????? ??????????,
?????????????? ???????? ??????, ???????
????????? ????? ???????????? ? ??????
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: dubovskaya@ibp.krasn.ru
# 108 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????. ???????? ?? ??????????. ?.?. ????????? «????? ? ???????????
????? ? ???????. ??? ????? ????????? ???????? ?? ??????? ??????? ???????, ???? ??
?? ??? ??????? ????????? ??? ???? ????????????? ?? ???????? ??????????, ? ????????
?????????, ? ???????? ???????????? ????? ?
??????????. ????? 3 ???? ?????? ? ?????????
??????????? ?????????? ? ???????????? ????? ??????? ??????? ? ???????? ?????? ?
?????? ???????? ? ? ???????? ?????????????? ? ???????????. ? ???????, ???????????
???????????? ??????????, ?????? ??????? ?
?????????????? ?????????????? ??????????? ? ????? ??????? ????????? ?? ????????
??????????????. ????? ???????? ?????, ???
? ??????????, ??? ?????????? ?????????????. ????? 4 ? ?????????? ??? ????? ????? ?
????? ????????????, ???????? ??????????
?? ??????????, ???????????, ??????????
???????? ?????? ? ??????????????? ???.
???????? ?????????? ????????, ?? ???????
????????, ??????? ????? ? ??????????. ? ????? 5 ??????????? ???????? ??????? ? ????????? ????????????, ? ?????????, ????????????? ???????? ????????, ????????????,
????????? ?????, ??????? ???? ??????????????? ????????? ??????????-?????????? ?
?????? ?????? ????????. ????? ?????????
????????? ??????? ???????????? (?? ???????? ??????? ????, ????? ???? ? ??.), ?? ???????? ? ??????? ???????????. ???????????
??????????? ?????????? ?????????? ?????
????????? ??? ??????? ???????????, ?????????, ????? ?????????????? ?????????? ?
????????????. ????? 6, ????????? «????????????? ??????????? ????????????», ????????? ???????? ???????????? ????? ?????
?????????? ??? ???????, ??????, ?????????,
????????, ?????? ? ?????????? ??? ???????? ? ????????, ???????, ??????????, ?????????; ? ????? ?? ????????. ????? ????????
????????? ? ???? ???????? ?????? ????????????? ? ??????????, ?????????, ?????????-
???, ???????????, ????????? ???????????.
????? 7 ? ?????????? ???????????? ? ???????????? ????????? ????????? (??????????)
? ????????????? ????? ?????????, ?? ?????????, ??????, ????????? ????????? ? ???????????? ?????????, ????????? ?????????
?????????. ?????? ??????????? ????????????? ? ??????????, ? ????????? ? ??????????. ????? 8 ? ??????????, ?????????????,
????????? ? ?????? ???????? ? ?????????
??????????, ????????? ??????????, ?????????? ??????, ????????? ?????? ?????????.
???????? ????? ???? ????? ????????? ?????????? ??????? ????????????? ? ?????? (???????? ?????????) ? ?????????? ? ??????????
????????? ??????????????? ?????????? ??
???? ??????: ????????????? ????? ?????? ?
?????, ?????????-?????????? ?????, ??????? ?????????-????????????, ????????????
(????????? ????), ??????????????-????????,
??????????????? ???????? ? ??????????.
???????? ?????? ????? ?????? ???????????????, ?? ??????? ????? ????????? ??????
?? ??????????? ?????????? ? ???????????.
? ????? ????? ????????? ?????????? ?????
??????? ? ????????? ? ?????????? («???????????? ??????????»), ????????? ?????????
????????? ?????????? ? ????. ?????????????? ????? 9 ???????????? ????? ????? ???????
????????????? ?? ????? ?????, ???????????.
????? ???????? ???????? ????????????? ?
?????. ? ????? ????? ????? ??????? ? ????????? ??????????, ?????????? ????????? ????? ? ???????? ????????????? ??????.
?? ???? ?????? ????? ? ??????? ???
??????? ??????? ????????? ???????????
???????? ??????, ?????????? ??????? ?? ??????????? ????, ??? ????????????? ? ??????????. ??? ? ????????? ??? ?????, ???
??? ?? ???????? ??????? ??????? ???????????? ?????????, ? ?????????? ? ???????? ?
?????????? ???????????? ??? ? ????????.
# 109 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????. ???????? ?? ??????????. ?.?. ????????? «????? ? ???????????
?. 145 ?????? ?????? ?????? ? ?????? «????????????» ?????? ???? «???????????»;
???. 4.17 (?. 253), ?? ????????, ?????????
? ?????? ?????????, ??? ????????? ???
x ?????? ???? ????????? ??-???????;
?. 261, ????? 2, 15-? ?????? ?????? ??????
???? «? ???????????? ??????» ?????? «?
?????????? ??????»;
?? ?. 301 ????? «???????? ????????? ???
??????????» ???????? Mo, ?? ??????????? Mg;
?. 384, 10-? ?????? ????? ? ?????? ????
??????? ?/?3, ? ?? ??????? ??/?3, ??? ???
????????????? ????????? ??? ??????????? 3 ??? ???/?3 ????? ?? ????? ????????? ???????? ???????? ??/?3;
?. 512, 16-? ?????? ?????? ????????? ???????? «? ?????????????????? ?????????»
?????? «? ?????????????? ?????????»;
?. 583, 21-? ?????? ?????? ? ?. 609, 11-?
?????? ????? ?????? «?????????????»
????? ?? ???????? «???????????», ???
??? ????? ?????? ????, ??? ????? ???
?????? «???????????» ?????????????
?????? (?. 617, 12-? ?????? ?????) ?????????.
?????????? ????? ??? ????????? ???-
?????? ?????????? ????????? ???? ???????? ??????? ????????? ??? ?? ??????????? ?
???????. ???? ???????????? ????????????
?????????, ???????? ? ????????????? ???????????, ?? ????? ???????? «??????????»
????????????. ?????????? ???? ??????????? ?????????. ???????? ??? ???????? ????????????, ????????? ? ????????? ?????????,
????????? ???? ? ?????????. ????????? ??
??????? ???????? «?????????????» ? «????????» ?????????????? ??????????????? ?
???? ??????????. ?????? ???????? ???????? ?????? «?????????????» ??? ???????
«?????????????», ?.?. ????? ? ???????????????? ???????? ? ??????????, ????????
???????? ????????? ? ??????????. ?????
«?????????» ?????????? ?? ?????? ????????? ???????????? ??????? ???????? ? ??????? ? ?????? ??????????? ? ???????????
?????????? ???? ???????????? ? ????????,
?????????? ? ????? ????????? ????? «?????????????» (?????????????????? ??????, Hydrobiologia, International Review of
Hydrobiology), «??????????» (Limnologica,
Limnology and Oceanography), «????????»
(????????, ????????? ????????????? ??????, Aquatic Ecology, Ecological Modelling,
Microbial Ecology, Oecologia), ?????? «????????» (???????? ?????????? ???, ????????
????, ?????? ????? ????????, Freshwater
Biology) ? ???? ? ?????? «???????» ????????????? (?????? ???????, Marine and
Freshwater Research, Water Research). ??
? ????? ?? ???????? ??? ?? ??? ????? ??????: «??, ??????????, ??? ???????». ???
??????????????, ??? ???? ?? ????????????
?? ????????? ????? ???????, ? ???????? ?
???????????? ???????????? ????????????????? ????????.
??????? ???????? ?????????? ?????? ?
????????, ????? ??????? ??????? ? ??????????????:
????????? ?? ???? ???????? ??????? ??? ???????????? ??????????? ?? ?????????????,
?????? ? ???? ????? ?????? ? ??? ???????
????????? ????????. ????????, ??????????? ??????????? ???????? «??????? ??????» ?
«??????? ?????????», ??????? ????? ???????????? ? ??????. ??? ???????? ??, ????? ? ????? 3 (????????? ??????????? ??????????) ?
???????? ? ???????? ???? ?? ?????? ???????
????????, ?? ? ?????????????? ? ????????
???????????? ? ?????? ???????, ??? ??? ?????????? ?? ???????? ???? ?? ?????? ??????????? ? ?????? ? ?????? ?? ???????, ? ?????
??????????????? ?????? ????????? ??????
????????.
# 110 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????. ???????? ?? ??????????. ?.?. ????????? «????? ? ???????????
? ?????????? ????????? ???????, ???????, ?? ??????? ??????, ???????? ??????:
?????????? ????????? ?????????? ? ???????? ??????????? ? ??????????? ?????? ???
??????? ????????????? ???????? ? ????????????????? ???????????. ?? ???????????
??????????? ?????????? ?????????? ?????????? ??????????? ????? ? ?? ?? ????? ???????????? ??? ???????????????? ???????????! ? ????? ??????????? ??????????????
?????????? ?????? ? ???????? ???????????????? ????????? ?? ???????????????? ???????: ?????????? ???????.
??????? ???????????, ??? ??????????
?????????? ?? ???????????? ????????? ?????????? ?????. ??????? ?????????, ??? ???
????? ????????? ? ???? ????????????? ????? ??????? ??????, ?????? ? ????, ???????
????????? ??????? ???????? ?????????? ?
?????????????? ??? ???????????? ?????????????????? ????????, ??? ? ???????? ??????? ???????? ? ???????????? ????????, ????????, ????????? (??? ??? ?????????? ???
??????) ? ?????? ????? ???????? ? ????????
???? ????? ? ? ?? ??????? ? ???????????? ?
????????? ??????.
?????? (IAEA, 2009; Varga et al., 2009,
Yankovich, 2009). ? ???? ????? ??? ????? ??????? ???????????? ???? ????????? ????????????? ???????? ????????????? ? ??????????
???????? ???????? ? ??????????? ?????, ??????? ? ?????-?????????? ??????.
????????? ? ??????
????? ????? ? ???? ?. ?????? ???????? ?? ???????, ????????????? ?? ?????????? ?? 20 ?? ?? ??? ? ??????????????? 2009
? 2010 ??. ???? ???????? ? ??????????? ??
?????????? ????? 5-10 ? ?? ??????. ??????
??? (Fontinalis antipyretica Hedw), ???????????? ?? ?????? ?? ??? ????, ???????? ?
??????? 1.5?2 ? ??? ?????? ???????????????? ??????? ???????. ??????????????
??????????, ????????, ???????? ? ???????? ??????? ???. ? ?????? ???????? ?????????? ??? Philolimnogammarus viridis Dyb.
???-??????????, ????? (Leuciscus leuciscus
baicalensis Dyb.) ? ??????? (Thymallus arcticus
Pallas), ?????? ?????? ? ??????? ? ???????????? ????. ?????????? ????? ????? ???????
?????????? 124?295 ??, ????? ? 165?222 ??.
????? ????? ??????? ?????????? 102?348 ?,
????? ? 51?99 ?. ??????? ???????? ? ??????,
???????????? ?? ??????? ??????? ?? ?????,
????????? 2+ ? 3+ ?. ????? ???? ???????????????? 61 ???. ??????? ? 55 ???. ?????. ???????
????????????? ?????????? ? ??? ?????????
?? ???????????? ????? (???????????? ? ??.,
2002), ??????????? ??????????? ???????????? ?? ??????? ???????? (????????????,
2002). ?????? ??????? ???????? ???????????, ??? ??????? ????? (???? ? ??., 2011).
??? ????????? ?????????? ????????????? ????? ??????? ??? ? ?????????? ??????????? ????????? ????????????? ????? ? ?????? ??????? ?? ???????, ????? ? ?????????
# 97 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
????? ??? 80 °? ?? ?????????? ?????. ?????
???? ??? ?????????? 200-300 ?, ???????? ? 1370 ?. ???? ??? ????????? ?? ?????? ? ?????,
??? ??????? ???? ????? (?????? ? ??., 2012).
??? ??????????? ??????? ???????????? ???????? ????? (????? ???? ??????) ? ??????
???, ?????? ????? ????????? ????? ???? ?????? ?? 6-11 ???. ????? ????? ??? ???????
? ????? ???????? ???????? (30 %) ? ???????
??????? (????.) ? ????? ??????????????? ??
100 ??.
?????????? ????????????? ? ?????? ????? ???????? ?? ?????-???????????? ?? ??????????? ??????????? ?????????? (Canberra,
???), ??????? ????????????? ? ???????
????????????
???????????
Genie-2000
(Canberra, C??). ???????? ?????????? ????????????? ?? ???? ?????? ????. ???????? ?????????? ????????????? ? ?????? ?????????? ? ??/?? ????? ????? ? ?????? ?????????
?????????? ????? (????. 1).
??? ????????? ???????? ??????????
???????? ????? ???????? ??? ? ??????????
(????????) ????????? ???????????????? ?????, ??? ??????????? ?? ????????? ???????
??????????? ??????. ????? ???? ??????????? ????? ???????? (Millipore, RAWP, ????.
47 ??) ? ???????? ??? 1.2 ???. ??? ????????????? ???? ??? ???????? ????? ? ??????;
?????? ????? ????????? ????? ????? ?????.
??? ????? ????? ?????? ??? 105 °?. ???-
?? ?????????? ? ?????????? ?????? ? ?????
??????? ?????? 1-2 ?. ?????? ????????????
?????????. ????? ???????? ????????????????? ????? ??? ?????????, ??? ??????? ?????
(???????? ? ??., 2009). ???????????? K ?????????? ?? ????????? ????????? FLAPHO-4
(Carl Zeiss, Jena) ? ????????-??????????
??????? (???? 30504 ? 97, ?? 52.24.391-95).
?????????? Fe, Mn, Zn, Cu, Cr, Cd ? Pb ? ??
??????-?????????????
????????????????
«??? ????? 2?» (???? 30692 ? 2000, ???
8288-86). ??????? ??????????? ??? K ?????????? 0.1; ??? Fe, Ni, Pb ? Co ? 0.01; ??? Cr ?
0.006; ??? Cu ? Mn ? 0.003; ??? Zn ? 0.001; ???
Cd ? 0.0003 (??/?). ??????? ????????? ? ????
????????????. ? ???????? ???????? ???????????? ????????? ???????????? ??????????????? ??????????? ??????? (????????? ?????
????????????, ??????). ??????? ?????????
? ??????????????? ????????????? ??????????? ????????? ????????? ?? ??? (????????
???????????? ? ????.RU.???.086.324). ?????????? ????????? ? ??/?? ??? ?/?? ?????
?????.
???????????? ????????? ?????????
??? ???? ???? ? ???????? ????? ?????? ??
????????? ? ?? ????????? ????? ????? (????? ??????????? ??? 105 °?) ? ???????? ? %.
???????????? ?????????? ????????????? ?
??????? ???????? ? ??????????? ????? (??)
???????????? ??? ????????? ???????? ??-
??????? 1. ?????????? ???? ? ???????? ???????????? (?????????, % ?? ????? ?????), ??????? ????. ±
?????. ??????.
???????????
?????????
????? ????
?????? ??? (F.antipyretica)
80.1±1.5
5
????????? (Philolimnogammarus viridis Dyb.)
78.0±1.2
4
?????? (Th. arcticus) ?????
72.5±3.8
20
?????? (Th. arcticus) ??????
70.6±7.4
19
???? (L. l. baicalensis) ?????
74.9±2.3
5
# 98 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
???????? ??? ???????????? ? ???????? ?????????? ? ???????? ?????????? ??? ???????????? ? ???????? ??? ???????? ?????????
(???????). ?? ????? ????? ? ????????? ???
???????????? ??? ????????? ????????????
??? ???????? ?????????? ? ????? ????????
??? ? ???????????? ??? ???????? ??????????
?? ??????? ????? ????. ???????????? ?????????? ???????? ???????????? ??? ???????
???????, ?????????? ?? ???? ?????? ???????????? (?????????????? 2009 ? 2010 ??.).
??????????
?????????? ?????????????
? ???????? ????????????
?? ???? ?????? ????? ?????????????
????????? ?????? 40K, ???????? ??????-
?
??/??
30
100
Ce (?1/2 = 284.89 ???.), 152Eu (?1/2 = 13.5 ???)
? ??????? ???????? ??????? 137Cs (?1/2 = 30.1
???). ????? ??????? ???????? ????????????? ??????????????? ? ???????? ??????? ???
(???. 1?). ? ?????? ???? ??????? ? ????? ??
??????????? ????????????? ?????????? ???????????????? 65Zn ? 137Cs (???. 1?, ?), ? ?????? ?????? ? ?????? 40K (?? 4.3 ??/??).
???????? ?????????? 60Co ? 65Zn ? ???????? ??????? ??? (???. 1?) ?????? 2010 ?.
144
?
100
30
0
Be-7
50
0
20
K-40
10
K-40
10
0
0
Co-60
Zn-65
Cs-137
?
Ce-144
Eu-152
Cr-51
Co-60
Zn-65
?
200
160
??/??
Cs-137
Eu-152
160
200
??/??
100
80
3.0
0
0
2.0
100
80
3.0
00
K-40
1
1.0
K-40
1
2.0
1.0
0.0
0.0
Zn-65
2.5
2.0
1.5
1.0
0.5
0.0
(???. 1). ?? ??????????? ????????????? ?
?????? ????? ?????????? ??????? ? ?????????? ??????????? 51Cr (?1/2 = 27.8 ???.),
54
Mn (?1/2 = 312.3 ???.), 58Co (?1/2 = 70.82 ???.),
60
Co (?1/2 = 5.27 ???), 65Zn (?1/2 = 243.9 ???.),
??/??
50
20
?????? ???????????? ???????? ??????????;
? ?????? ??? ??????????????? ????? 7Be
???.09
Cs-137
???.09
Zn-65
Zn-65
???.09
???.10
Cs-137
???.10
Cs-137
???.10
???. 1. ???????? ?????????? ????????????? (± ?????. ??????. ???????? ????????? ?????) ? ?????? ?????
???????? (??/??) ??????? ??? (?), ???????? (?), ???? ???????? (?) ? ???? ?????? (?), ?????????? ? ?.
?????? ? ??????????????? 2009 ? 2010 ??.
# 99 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
???? ????, ??? ?????? 2009 ?. ???????? ?????????? ??????????? ????????????? ? ???????? ???????? ? ?????? ??? ?????? 2009 ?
2010 ??. ?????????? ???????? ?? ????? ??????
(???. 1?-?). ?????????? 65Zn ? ?????? ?????
?????? 2010 ?. ???? ???? ???????? ??????????? (???. 1?).
???????????? ????????
? ???? ? ???????? ????????????
???????????? ?????????????? ? ?????? ???? (n = 5) ??????????? ? ??????? ??????? (0.6-0.8 ??/? ??? K, 2.6-2.9 ??/? ??? Na,
15-29 ??/? ??? Ca, 2.7-3.7 ??/? ??? Mg), ???
?????????????? (0.01-0.04 ??/? ??? Fe, 0.0030.010 ??/? ??? Zn, 0.0008-0.0015 ??/? ??? Cu,
0.002-0.005 ??/? ??? Mn, 0.00001-0.0007 ??/?
??? Cr ? Pb) (????. 2). ???????????? ????????
? ???? ? ? ??????????? ???? ???? ? ??????
??? ???? ???? ???????? ?????????????? ??????????? (????. 2). ???????????? ?????? ?
?????? ?? ???? ??????, ????? ??????? ???, ??????? ?? ????????? ???????? ?????????????? ???????????. ?????????? ???????? ? ?????? ???? ? ?????, ????????? ? 2009 ? 2010
??., ??????????? ?? ??????????. ??????? ???
??????? ?? ?????????????? ??????? ????????
?? ???? ??????, ??????????? ? ????. 2.
???????????? ??????????
????????????? ? ????????
?????????????
?????? ?? ????????????? ??? ????, ????????? ? 2009 ?., ????????? ? ?????????????? ???????? ??????????? 137Cs ? 120; 60Co ?
4.6; 65Zn ? 3.3 ???/? (????????? ? ??., 2008),
??? ????, ????????? ? 2010 ?. ? 82; 0.8 ? 0.3
???/? ??? ?? ?????????????, ??????????????
(????????????, ????????????, 2012). ???????? ?? ????????????? ? ???????? ???????
??? ?? ???? ?????????? ? ???????? 5000?
12000 ??? 60Co, 900?3200 ? ??? 65Zn, 60?140 ?
???
Cs (???. 2?). ???????? ?? ????????
???? ?? ???? ?????????? ?? 1420 (??? Pb) ??
233 000 (??? Mn) (???. 3?).
? ??????????? ???? ?????? ???????????? ???????? ??????? ?? ??????????
??? 65Zn (2.1-8.7), ?? ????????? ????????????? ?? ????????? ??????? (???. 2?). ?? ???????? ???????? ??????? ???????? ?? ? ????
??????????? ???? ???????? ??? ???? (2.9),
?? ?????? ????????? ???? ?????? ???????
(???. 3?).
????? ??????? ???????? ?? ????????????? ? ?????? ????? ? ??????? ?? ????????? ? ???????? ???????? ???????? ??? 40K
137
(???. 2?, ?). ???????? ?? 40? ? ?????? ??????? ?? ???????? ?? ????????? 0.1. ????????
?? 65Zn ? ?????? ??? ?? ????????? ? ???????? ???????? ?????????? ? ??????? 0.13
? 0.21 ??? ??????? ? ?????, ??????????????,
? ?? 137Cs ? 0.21. ????? ??????? ????????
?? ???????? ? ?????? ? ?????? ??? ?? ????????? ? ???????? ???????? ???????? ???
K (???. 3?). ?? ????? ? ?????? ??? ?? ???????? ???????? ???? ?????? ? ???????. ??
????? ? ?????? ? ?????? ??????? ?? ????????
? 2-3 ???? ????????? ??????? (???. 3?).
??????????
? ?????????? ???????? ? ???????? ??????????? ?? ???? ??? ?? ???????? ?????????
????? ????????????????? ???????? ??, ??????????? ???????. ??? ???????????? ???????? ?? ????????? ???????, ??? ???? ????????????? ??????????. ??????????? ?????
??????????? ???????????? ? ??????? ??????? ??? ?? ????, ??? ? ????, ? ?????? ??????
?????? ?????????. ???????????? ???????? ?
???? ?. ??????, ?????????? ????, ????????
? ???????? ????????????? ???????? ????, ????????? ?? ?????? ???????? ?? ?. ???????????
(???????? ? ??., 2010). ???????????? ??????????? ???????? ? ???????? ????????????
# 100 #
4
5
?????? (??????)
????
0.00065± 0.00011
4.10±0.79
3.31±0.87
4.30±1.46
1.26±0.27
2.58±0.57
K, ?/??
0.023±0.011
64.21±60.94
5.30±1.00
6.80±0.50
60.32±35.24
982±231
Fe, ??/??
0.001±0.000
2.71±0.99
0.39±0.08
0.49±0.13
11.02±2.08
3.79±0.75
Cu, ??/??
* ?.?. ? ???????? ?????? ???????? ?????????????? ???????????.
4
5
?????? (?????)
5
???? (?????)
4
?????????
?????
????
?????? ???
???????????
0.005±0.003
26.29±5.01
7.42±1.48
10.38±2.56
13.02±1.34
15.23±4.18
Zn, ??/??
0.003±0.001
1.97±0.65
0.42±0.36
0.65±0.29
22.82±9.21
593±239
Mn, ??/??
?.?.
?.?.
?.?.
?.?.*
0.13±0.03
1.24±0.18
Co, ??/??
0.052±0.061
0.031±0,023
?.?.
0.054±0.053
0.40±0.06
Pb, ??/??
0.0002± 0.0003 0.0003± 0.0003
0.25±0.13
0.26±0.03
0.22±0.03
0.45±0.11
7.40±2.85
Cr, ??/??
?.?.
0.015±0.018
0.001±0.003
0.005±0.005
0.008±0.006
0.04±0.01
Cd, ??/??
??????? 2. ?????????? ???????? ? ?????? ????? ???????? ???????????? ? ???? (??????? ????. ± ?????. ??????.) ?. ??????, ?????????? ? ???????-??????? 2009
? 2010 ??.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
?. ??????, ????????????? ????, ??????????
????????????? (????. 2). ????????? ?????? ??????? ? ?????????????. ? ?????? 2010 ?. ????????? ????????? ??????????? ????????????
?? ???, ????? ???? ?????????? ???????????
??????????? ????????????? (????? ??? 60Co ?
65
Zn) ? ???? ?. ?????? ??????????? ?????????
(????????????, ????????????, 2012), ?????????????, ???????? ????? ????????????? ? ???????? ???????????? ?? ????.
?
??
?
??
100000
10
10000
8
1000
6
100
4
10
2
0
1
Co-60
Zn-65
?
??
K-40
Cs-137
Zn-65
Co-60
4
3
3
2
2
1
1
0
Eu-152
?
??
4
Cs-137
0
K-40
ZN-65
CS-137
K-40
ZN-65
CS-137
34012
???.09
???.09
K-40
???.09
???.10 ZN-65 ???.10
???.10
??.????.±?????.??????.
CS-137
???. 2. ???????????? ?????????? ????????????? ? ???????? ??????? ??? ?? ???? (?) ? ? ???????????
????? ?????? ????????????? (?), ???????????????? (?????) (?), ?????????????? (?????) (?)
?
??
?
??
3.5
1000000
1
3.0
100000
2
3
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2.5
2.0
10000
1.5
1.0
1000
0.5
100
K
Fe
Zn
Cu
Mn
Cr
0.0
Pb
K
Fe
Zn
Cu
Mn
Cr
Co
Pb
Cd
???. 3. ???????????? ?????????? ???????? ? ???????? ??????? ??? ?? ???? (?) ? ? ??????????? ?????
(?): 1 ? ?????? ?????????????, 2 ? ???????????????? (?????), 3 ? ?????????????? (?????), 4 ?
???????????????? (??????)
# 102 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
?? ????????????? ???? ?????????? ?????? ??? ??????????? ??????????? ???????????? ? ??????? ???????????? ?????????????????? ??????? ?? ????. ????? ?????????
??????? ????? ?????? ????????? ???????????
?????? ?? ??? ? ?????? ???????? ???????
??? (???????????????? 2010 ?.) ?? ?????????? ?????????????? ????????, ????? ??? 24Na
(?1/2 = 14.96 ?.), 51Cr ? ??., ? ????? ???????????
????????? ?????????? ?????? ???????? ? ?????????? ???????????. ???, ?????????? 60Co
? 65Zn ? ???????? ??? ????????? ? ?????????
??????? 2010 ?. ? 3-4 ???? ?? ????????? ? ??????????? ???????? 2009 ?. (???. 1?). ?????????? ????????????? ? ???????? ? ????????
??? ?? ???? ????? ??????????, ? ??? ??????????????? ??????? ???????? ?? (???. 2?,
3?). ????????? ??????????? ?????????? ??????????? ??????? ??????? ? ????????????
?? ????, ?????? ??? ?????? ????????????
??? ????????? ???????????? ??????????? ?????????.
?????????? 65Zn ? ???????? ????????
?????? 2010 ?. ?? ??????????? ?? ?????????
? ?????? ??????????? ???? (???. 1?), ?????????? 60Co ????????? ? ??????? ???????, ???
??? ??????? ???, ? ??????? ? ??????? ????. ????? ???????, ????? ?????????????? ????? ??
?????????? ???????? ?????????? ?????????
?? ????????? ? ??????? ?? ??????????, ???,
????????, ??????????????? ? ????????????
???????? ???? ??????????? ???? ????????
? ????????? ???????? ??? ??????. ????????, ? ?????????? ??????? ??????? ????????
?????????? 65Zn ? ?????????? ?? ???? ??????? ???????. ??????? ???????? ????????
? ?. ?????? ??????, ? ????????, ?????????,
? ????? ???????? ??????????. ? ???????
?????????????? ???????? ???? ????????,
??? ?????????, ????????? ? ??????, ?????
??????????? ? ???? ????? ???????? ???????
??? (Kalachova et al., 2011). ?????????? 65Zn
??????????? ?? ???????? ??????? ??? ?????
?????????? (???. 2?). ??? ???? ???????? ??
65
Zn ? ???????? ???????? ???????? ????? ????????? ???????? ?? ???? ???????? ??????????
????? ??????? ? ?????? ???. ?? ??????????
???????? ???????? ? ??????????? ???? ?????? ????????????? ???????? ???????????
?????????? ???? (???. 3?). ????? ??????????
? ??????????? ???????????? ??????????
???? ??????????? ?? ?????????? ?. ??????
(???????? ? ??., 2009). ??????? ?????? ??????????? ? ????????? ????????????? ? ???????? ? ???? ???????? ?? ???????? ???????
??? ????????????.
????????? ???????? ???????? ??????????? ??????? ??????? ? ?????-???????
?????? (???? ? ??., 2011). ??? ??????????? ?
?????????? ???????? ????????????? ????
???????? (???? ? ??., 2011). ????????????
?????? ??????? ??????? ?????? ????? ???????? ??? ??????? ????? ? ???????? ???????
?????? ? ???????? ? ??????? ??????. ???????? ???????????? ????????????? ?????????? ? ???????? ?????? ???? ????????? ????
(1.8-3.7), ? ???????????? ??????????? (1.81.9) ? ??????? ????, ??? ? ????????. ?????????? ??????????? ????????????? ? ??????
???? ???? ????? ??? ???? ????????. ? ?????? ???????? ?????????? 65Zn ?? ?????????
??????? ??????????? ? ??????? ? ????????,
?? ???? ???????????????? ? ??????? 2010 ?.
(???. 1?), ??? ?? ????? ???? ????????? ?????? ??????? ?? ????????? ? ???????????
????? ????????, ?.?. ?????? ???? ? ????????
???????? ? ??????????????? 2010 ?. ??? ???????. ????? ?? ???????? ???????????? ???????? ? ????????, ???????????? ?????????????
?????????? (3-4) ? ??????????? (1.4-1.6) ???????? ? ????????????? ????????.
???? ?????? ????????, ??? ? ??????
???-?????????? ???????? ??????????? ?????????? ????? ? 40K ?? ????????. ?????? ??-
# 103 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
???? (???????? ? ??., 2009) ????? ????????
??????????? ?????????? ????? ? ?????? ??????? ?? ???????? ? ?????? ??????????????
?????????? ??????. ???? ?? ????????? ??,
?????????? ??????????? ????????????? ?
?????? ???-?????????? ?? ???????? ???????? ????????????. ?????????? ???????????
??????????? ????????????? ? ????? ??????? ?????? ???????? ????? ???? ?????? ??????? ??????? ???? ????. ???, ????????, ???
??????? ????? ????????? ? ???? ??? ?? ????
????? ????? ? ??????????????? ????? (?????, 2002).
???????????? ???? ?????????? ????????, ??????????? ? ????? ??????????? ???
????????????????? ???, ? ?????? ???????
????????? ???????????? ? ?????? ? 8.7
??? ??? ??????, 6.9 ??? ??? ????, ? 4.7 ??? ???
????????, ? 3.5 ??? ??? ?????. ????? ???????
???????????? ?????????????? ???????? ?
?????? ?? ????????? ? ??????? ??????????
??? ?????? ????? ???????????? ??? ??????? ???????? (???????, ?????????, 1998; ????????? ? ??., 1998; ?????, 2002; ???? ? ??.,
2009). ????? ???????, ??????? ????? ?????????? ????????, ???????? ?? ????? ? ?????? ???, ????????? ?? ?????????. ? ??????
??????? ???????? ??????????? ??????????
?????, ????? ? ?????? ?? ???????? ????????
(???. 3?). ??????? ?????? ???????? ? ??????
??? ??????????????. ???????????? ? ?????? ??? ??? ?????????? ?? ???????, ???? ?
????? ???? ? ?????? 2010 ?. ??? ???? ???????????????? ?? ?????????? ??????? ????????
?????? (?????? ? ??., 2012).
????????????? ?? ????????????? ????????? ????????????? ? ?????????? ???????? ? ?????????? ???????? ?? ?? ??????????
????????, ???? ???????? ????? ???????, ???
???????????????? ???????? (???????, 1981).
? ????????????? ???? ??????????? ?????
????????????? ?????????? ????? ????????
? ?????????????? ?????????? 40K, ??????? ?
????????? ????? ???????? ?????????? 0.012 %.
????? ????????? ????? ????????, ?????????
??????????? ???????? ????? ???????? ? ??????? ?????????. ?????????? ????? ? ???????? ? ??????? ???? ?????? ????????????? ?
????????? ??? ????? ??????????, ??? ?????????? ????????????? ???????? 60Co ? 65Zn.
? ??????????? ????? ????????????????
(?????) ? ?????????????? (?????), ????????, ?????????? ????? ????? ??????????, ???
??????? 65Zn (???. 2?, ?; ???. 3?). ? ???????
?? ?????????????? ?????????? ?????? ????? ????? ?????????? ????????????? ?? ???
???????? ? ?????? ??????? ? ????? ?????????? ? ? ?????? (???. 3?). ? ????? ?????
??????????????, ??? ??????????? ???????
????????????? ?????????? ????????, ? ????? ?? ????????? ????????????? ???????? ?
????? ???-?????????? ?. ?????? ?? ???????? ??????????????.
? ????????????? K ? Cs ??????????????? ??? ????????-??????? (IAEA, 2009; Varga
et al., 2009). ???? ?????? ????????, ??? ???????????? ???? ?????????? (?? 70 %) 40K ?
137
Cs ? ????? ???????? ????????? ? ??????
(?????? ? ??., 2012). ??????? ???????? ?????????? ??? ???????????? ??? ??????? ???????? (?????? ? ??., 2008; Yankovich, 2009).
????? ???????, ????? ?????????????? ? ????????? ??? ??? ??, ??? ??? ?????? ? ?????.
?????? ????????????? ?????????? 137Cs ?
?????? ???-?????????? (???????? ? ??????)
?? ?????????? ? ?. ?????? ???? ???????????
??????, ??? 40K ? K. ????????, ??? ????????????? ?????????? ?????????-???????? ???????????? ??????????????? ???????????????????? ? ????? (Yankovich et al., 2009). ??
?????? ??????? ??? ?? ?????? ????????????
????????? ???????????, ??? ?????????? 137Cs
????????????? ?????? ???????????? ??????????? ? ????????????? K+ ? ??????????
# 104 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
????? (Rowan, Rasmussen, 1994). ????????,
??? ???????????? ????????????? ??? ????????????? ????? ? ?. ?????? ???????????
????????? ???????????? ?????.
?????? ?????? ???????? ???????????
????????????? ?????????? 137Cs ????????????? ?????? ?? ???? ?? ???????. ? ?????????, ????????, ??? ????? ??????? ????????
?????????? 137Cs ?????????????? ? ??????
(??????) ?????? ????? ??? (?????, 2004;
???????? ? ??., 2009; ??????? ? ??., 2009).
?? ????? ???????? ??????????? ???????????? ???????????? ???????? 137Cs ? ?????
???? ?? ??? ???????? ? ?????? ? ?. ??????
(????????? ? ??., 2012). ?????????????, ? ????? ???-?????????? 137?s ??????????? ?????
????????? ????? ??? ?????? ???????????
???????.
??????????
?????? ???????? ??????????? ????????????? ? ???????????? ???????? ? ??????????? ??????????? ????? ?. ??????, ??????? ? ?????-?????????? L. l. baicalensis,
Th. arcticus, ???????, ??? ??????????? ??????? ???????? (Fe, Zn, Cu, Mn, Cr, Co, Pb, Cd)
? ??????????? ?????????????, ?????????????? ? ?????? ??? (65Zn ? 137Cs), ?? ?????? ??
???????? ??????????? ??????? ??? ? ???????? ? ???????????? (??<1). ???????????
?????????? ? ?????? ??????????? ?????
???????? ?????? ??? ????? ? ??? ??????????
?????????????? ??????? 40K. ? ?????? ??????? ???????? ??????????? ?????????? ?????,
????? ? ?????? ?? ???????? ????????. ? ???????? ???????? ???????? ?????????? ????
? ???????????? ????? (65Zn) ?? ??????? ???,
??????? ?????? ????????????? ?????????? ?
????????????? ???????? ? ?????? ??????????? ???? ????????????.
????????? ?? ?????????? ? ????????????? ???????? ????????, ??? ? ?????????????
??????????? ????? ????????????? ?????????? ????????? ?????? ??? ???? ?????????
???????? K ? 40K. ????????????? ??????????
???????? ?????????-???????? 40K ? 137Cs ? ????????????? ??????????? ????? ?? ??????????? ??????????? ?????????.
?????? ??????????
1.
2.
3.
4.
5.
???????????? ?.?., ???????????? ?.?. (2012) ???????????? ? ???? ?. ??????. ?: ?.?.
???????? ? ??. (???.) ????????????? XXI ????: ????????? ????????????? ?????????????????? ???????????, ??????????, 14-16 ??? 2012 ?., 190-194 c.
???????? ?.?., ???????? ?.?., ??????? ?.?., ????? ?.?., ?????????? ?.?. (2009) ????????????? ? ???????? ???????? ? ??????????? ????? ?????????? ???? ?????? ? ??????
?. ???????????. ?????? ??????? 36 (5): 623?632.
???????? ?.?., ???????? ?.?., ??????? ?.?., ???????? ?.?., ?????????? ?.?. (2010) ?????? ?????????????? ??????????? ?. ?????? ?? ?????????? ???????? ? ???????? ??????????? ?????????? ?? ????????, ????????????? ???? ? ???? ?. ???????????. Journal of
Siberian Federal University, Biology 3(1): 82-98.
???? ?.?., ???????? ?.?., ?????? ?.?., ???????? ?.?. (2009) ??????? ??????????? ????????????? ??????? ???????? ? ????????? ??? ????????? ???????. ?????????. ????. 45 (3):
55?64.
???????????? ?.?., ????????? ?.?. (2004) ????????????? ??????????? ?????? ??????????
???? ?????? ? ???? ??????? ????????????????? ?????????. ??????. ????????. ????????????? 44 (3): 361?366.
# 105 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
6.
7.
8.
????????? ?.?., ???????????? ?.?. (2008) ???????? ???? ?????????? ??????????? ????????????? 60Co, 137Cs, 152Eu ? 241Am ? ?????? ?????????? ?. ??????. ?????????? 50 (5):
475?480.
????????? ?.?., ???????????? ?.?., ??????????? ?.?., ??????????? ?.?. (2008) ????????????? ????? ???????? ???????????????? ????????????? ????????? ? ?????? ???? ????
??????. ????. ?? 423 (4): 479? 482.
???????????? ?.?., ???????? ?.?., ?????? ?.?., ???? ?.?. (2002) ????????? ?? ??????????. ??????????: ??????, 127 ?.
9. ???????????? ?.?. (2002) ??????? ??????? ?????????. ??????????: ??????, 320 ?.
10. ??????? ?.?. (1981) ??????? ?????????????? ???????????????? ????????. ?.: ?????,
247 ?.
11. ?????? ?.?., ?????? ?.?., ??????? ?.?., ?????? ?.?. (2008) ???????? ?????????? ? ????????????? ???????? ?????????????? ????????????? ? ??? ???? ?????????? ????????????? ???. ?????????. ????. 44 (3): 95?113.
12. ??????? ?.?., ????? ?.?., ?????? ?.?. (2009) ??????????? ?????????? 137Cs ? ????????? ????? ??? ?????????? ????????????? ?? ??????????? ?????. ?????????. ????. 45 (5):
110?116.
13. ?????? ?.?., ????????? ?.?., ???????????? ?.?. (2012) ???????????? ? ??????? ????????? ?? ???????????-???????????? ??????? ???????? ??????? ?. ??????. ??????. ????????.
????????????? 52 (3): 305-311.
14. ???? ?.?., ???????? ?.?., ???????? ?.?., ?????? ?.?., ????????? ?.?., ?????? ?.?.,
15.
16.
17.
18.
19.
20.
21.
22.
?????? ?.?. (2011) ??????? ??????? Tymallus sp. ? ??????? ??????? ?. ??????. Journal of
Siberian Federal University, Biology 4 (3): 281?292.
??????? ?.?., ????????? ?.?. (1998) ??????????? ????????? ????????????? ??? ???????
??????????? ??????? ????????. ?????? ??????? 25 (5): 603?610.
????? ?.?., ????????? ?.?. (1996) ?????? ???????????? ?????????? ?? ?. ?????? ?????
?????? ? ???????????? ??????????? ????????? ????????????? ???. ??????? ??????? 81
(3): 226?232.
????????? ?.?., ??????? ?.?., ????????-?????? ?.?., ?????? ?.?. (1998) ??????? ??????? (Zn, Fe, Cu, Mn, Pb) ? ?????? ? ??????? ???????????? ?????. ???. ????. ????. 5:
477?483.
???????? ?.?., ?????????? ?.?., ?????? ?.?., ????? ?.?. (2009) ???????? ??????????
137
Cs ? 40K ? ?????? ??????????? ????? ??? ??? ????? ? ????? ? 1995-2007 ??. ??????.
????????. ????????????? 49 (6): 721?728.
????? ?.?. (2002) ?????? ?????????????? ????????? ???????? ???????? ??????????????.
???????????: ???????. ???. ??-?, 270 ?.
????? ?.?. (2004) ????????????? ??? ???????? ? ???? ??????? ?????? ?? ?????????????
???. ?.: ???????????? ??????? ??????? ???, 215 ?.
????????? ?.?., ?????? ?.?., ???????????? ?.?. (2012) ?????? ???????? ???????????
????????????? ? ??????????? ????? ???? ??????. ???. ????. ????. 4: 497-504.
Bolsunovsky A., Bondareva L. (2007) Actinides and other radionuclides in sediments and
submerged plants of the Yenisei River. J. Alloy. Compd. 444-445: 495?499.
# 106 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ?????????? ????????????? ???????????? ???????? ????????????? ? ?????????? ?????????
23. IAEA (International Atomic Energy Authority) (2009) Quantification of radionuclide transfer in
terrestrial and freshwater environments for radioecological assessments. TECDOC-1616. Vienna:
IAEA, 622 p.
24. Kalachova G.S., Gladyshev M.I., Sushchik N.N., Makhutova O.N. (2011) Water moss as a food
item of the zoobenthos in the Yenisei River. Cent. Eur. J. Biol. 6 (2): 236?245.
25. Rowan D.J., Rasmussen J.B. (1994) Bioaccumulation of radiocesium by fish ? the influence of
physicochemical factors and trophic structure. Can. J. Fish Aquat. Sci. 51 (11): 2388-2410.
26. Varga B., Leclerc E., Zadyvai P. (2009) The role of analogues in radioecology. J. Environ.
Radioactiv. 100: 802?805.
27. Yankovich T.L. (2009) Mass balance approach to estimating radionuclide loads and concentrations
in edible fish tissues using stable analogues. J. Environ. Radioactiv. 100: 795?801.
Efficacy of Trophic Transfer of Radioactive
and Stable Isotopes of Metals
to Zoobenthos-Feeding Fish of the Yenisei River
Tatiana A. Zotina, Elena A. Trofimova,
Alexandr Ya. Bolsunovsky and Olesia V. Anishenko
Institute of Biophysics SB RAS,
50/50 Akademgorodok, Krasnoyarsk, 660036 Russia
Accumulation of radioactive and stable isotopes of metals in trophic chain of the Yenisei River leading
to zoobenthivorous fish species, dace (Leuciscus leuciscus baicalensis) and grayling (Thymallus
arcticus), have been investigated. Concentration factors (CF) of radionuclides and metals in trophic
pairs, aquatic moss (Fontinalis antipyretica) ? gammarids (Gammaridea), gammarids ? dace and
gammarids ? grayling, were estimated. Dietary transfer of metals (Fe, Zn, Cu, Mn, Cr, Co, Pb and
Cd) and artificial radionuclides (65Zn and 137Cs) from gammarids to muscles of fish species was not
effective (CF<1). Effective transfer of potassium (CF = 2.6-3.4) and its natural isotope 40K (CF = 2.62.8) is possible in these trophic pairs. Effective accumulation of K, Zn and Cd in liver of grayling is
possible from biomass of gammarids (CF = 3.2, 3.0 and 2.1, respectively). Accumulation of copper
and 65Zn is possible in biomass of gammarids from aquatic moss (CF = 2.9 and 2.2-8.7, respectively).
Dietary transfer of other stable and radioactive isotopes of metals in this trophic pair is not effective.
Efficacy of dietary transfer was similar for analogue isotopes K and 40K only. Efficacy of diet transfer of
isotopes of analogue elements, 40K and 137Cs, differed considerably among the studied trophic pairs.
Keywords: aquatic moss, dace, gammarids, grayling, Fontinalis antipyretica, Gammaridea, Leuciscus
leuciscus baicalensis, Thymallus arcticus.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2013 6) 108-111
~~~
???????? ?? ??????????
?.?. ????????? «????? ? ??????????.
?????? ?? ????? ?????????????».
????: ???????????????, 2011. ? 704 ?.
?.?. ????????, ?.?. ?????????*
???????? ????????? ?? ???,
?????? 660036, ??????????, ?????????????, 50/50
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ?????????, 79
Received 08.03.2013, received in revised form 15.03.2013, accepted 22.03.2013
????? ?.?. ????????? «????? ? ??????????. ?????? ?? ????? ?????????????» ??????? ?????????? ???????? ?????????????
????????? ? ?????????????, ????????, ?????????, ?????????? ? ?????????????? ????????????? ??????????????. ??-??????, ??????
??? ????????????? ???????? ?????????? ??
?????????? ???????????? ??????? ? ?????????? ?? ????? ????????????? ? ????????????? ? ????? ????? ????? ?????????. ????????, ?????? ??? ????? ???????????? ?????
??????????????? ????, ?????????? ???????????, ?????? ?? ?????? ???????????? (????????, ????????????? ????????? ?????? ??
?.?. ???????? ? ?.?. ?????????), ?? ? ??????????? (????????, ??????????????? ? ??????? ?????????? ? ??????????). ? ????? ?????
??????????? ???????? ????????????? ??????????, ????????, ?????? ? ?????????????
????????????? ????? ????????, ??? ?????????????? ? ????????????. ? ?????????????
????? ??????????????? ?????????????? ??
?????? ?????? ?? ?????????? 2000-2010 ??. ?
*
?????? ?????????? ?????????? (????? 800
??????????!), ? ??????? ?????? ?? ??? ????
?????????? ????? 31 % (?? ??? ?????????? ?
????? 10 %). ??? ??????, ??? ??? ? ????? ????? ??????? ????? ?????? ?????????? ?????? ?? ???????????? ??????, ??????? ? ?????
XIX ????, ??? ?? ? ??? ????? ?????. ?????
???????? ????????, ????????? ? ?????????,
??????????? ??????????, ??? ??? ?????????? ???????? ? ?????????? ????????? (??????? 1.1, ??????????? ???????? ???????
?????? ???????? ? ???????? (?. 20)). ? ?????
????? ???????? ??????? ????????? ? ??????? ??????????????? ???????.
? ????? 1 ????? ???????? ???????? ????
???????????? ??????? ????? ????????????? ?
????? ?? ? ??????? ???????????? (?????????
? ??????? ??????? ????????? ????). ? ?????
2 ??? ????? ?????????? ?? ??????? ?????????? ? ????? ? ???, ? ?????????, ??????????? ????????????? ? ???????? ??????????,
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© Siberian Federal University. All rights reserved
Corresponding author E-mail address: dubovskaya@ibp.krasn.ru
# 108 #
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