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58.Журнал Сибирского федерального университета. Сер. Биология №1 2011

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Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Журнал Сибирского федерального университета
2011
Journal of Siberian Federal University
4 (1)
Биология
Biology
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??.-?. ???, ?-? ???.-???.????
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CONTENTS / СОДЕРЖАНИЕ
С.А. Николаева,
С.Н. Велисевич, Д.А. Савчук
n?2%????? Pinus sibirica ?= ??%-"%?2%*? g=C=??%-q,K,!?*%L
!="?,?/
?3?
С.Н. Велисевич, С.Н. Горошкевич
b%?!=?2?= ,?????,"%?2? ?%!-%?????= ????!=2,"?/. C%K??%"
*??!= ?,K,!?*%?%
? 23 ?
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:
Nadezhda N.Sushchik
М.П. Тищенко
p=?2,2????/L C%*!%" %K%?,? ="2%?%K,???/. ?%!%? "
23??!%"%L , 2=???%L ?%?=. g=C=??%L q,K,!,
? 36 ?
Julia A. Darikova,
Yulia V. Savva, Eugene A. Vaganov,
Alexi M. Grachev and Galina V. Kuznetsova
Grafts of Woody Plants and the Problem of Incompatibility
Between Scion and Rootstock (a review)
? 54 ?
А.Е. Безруких,
Е.Н. Есимбекова, В.А. Кратасюк
???C?!=23!?=
,?=*2,"=?,
K,-?!???2?%L
?,?2??/
?"?2 ?,.?
K=*2?!,L
NADH:FMN-%*?,?%!??3*2=?=???,-?!=?= " ???=2,??
? 64 ?
???????? ?.?. ?????? ????????? ?.?. ??????????
???????????? ??????? ?.?. ?????????
????????? ? ?????? 18.03.2011 ?. ?????? 84x108/16. ???. ???. ?. 8,4.
??.-???. ?. 7,9. ?????? ???. ?????? ????????. ????? 1000 ???. ????? 4235.
?????????? ? ?? ???. 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
????????????? ? ??????????? ???
?? ? ??77-28-725 ?? 29.06.2007 ?.
????? ???????? ? «???????? ??????? ????????????? ??????? ???????? ? ???????, ? ??????? ??????
???? ???????????? ???????? ??????? ?????????? ??????????? ??
????????? ?????? ??????? ??????? ?
????????? ????» (???????? 2010 ?.)
Е.И. Пономар?в,
Д.М. Исмаилова, Д.И. Назимова
qC32?,*%"/L ?%?,2%!,?? ?%!?/. ????/. .*%?,?2?? q= ?
? 75 ?
Denis Yu. Rogozin,
Marina V. Pulyayevskaya, Ivan V. Zuev,
Olesya N. Makhutova and Andrey G. Degermendzhi
Growth, Diet and Fatty Acid Composition of Gibel Carp
Carassius Gibelio in Lake Shira, a Brackish Water Body in
Southern Siberia
? 86 ?
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2011 4) 3-22
~~~
??? 581.14:581.4
????????? Pinus sibirica
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?.?. ?????????, ?.?. ??????
???????? ??????????? ?????????????
? ????????????? ?????? ?? ???
?????? 634055, ?????, ??. ?????????????, 10/3 1
Received 4.03.2011, received in revised form 11.03.2011, accepted 18.03.2011
??????? ????????? ?????? ???????? ?????????? ????? ?????????? ??????? ?????????????? ????????? ?? ??????? ???????? ????????? ??? ???????-????????? ????? ????????????????-??????????? ???? ??????????? ????????????. ???????????????? ????????? ???????????????? ?? ???????????? ? ?????????????? ????????? ???????????? (? ??? ?????
???????? ? ??????, ?? ??????? ? ??????? ??????????? ??????? ??????) ? ????????????
(??????? ????????? ? ??????????? ?????- ? ??????????????) ????. ?????????? ?????????
???????? ? ???? ???? ????: 1) ?????????????? ?????, ??????????? ? ?????? ????????????
????????????? ????????? ? ????????????? ???????????????????? ????????? ? 2) ????????
????? ? ??????????? ????????.
???????? ?????: ???? ?????????, Pinus sibirica, ?????????, ????????????????-?????????? ???
??????????? ????????????, ??????? ??????? ? ????? ?????.
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? ??????? ???? ???????-?????????
??????? ???? ????????? (Pinus sibirica Du
Tour) ????????? ? ???? ? ?????? ????????
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??? ????? ?????? ???? ???????????. ????????
??????? ??????????????? ????? ??????????, ?????????????? ????? ?????????????
????????????????-?????????? ???? (??????????, 1990). ? ??????????? ??????? ????????? ???? ????? ???????? ? ??????? ??????
????? ?????????, ??????? ????????? ?????
*
1
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Corresponding author E-mail address: sanikol1@rambler.ru
© Siberian Federal University. All rights reserved
#3#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
(???????, 1969; ??????, 1975; ????????, 1988
? ??.). ???????? ? ??????????? ???????? ?????
?????????? ??????? ??? ?????????????? (?????????, 1972), ???????????????? (???????,
1964, 1975; ????????? ? ??., 1967; ?????????,
2002 ? ??.), ????????????? (?????????, 1985?;
???????? ? ??., 1989; ??????????, 1992; ??????????, ?????????, 1996, 2000; ?????????,
??????????, 1997; ?????????, ??????, 2002?;
??????? ? ??., 2009 ? ??.) ? ????????????????? (?????????, 2007) ?????? ??? ????????.
? ??????? ?????????? ?????????? ?????????? ????? ???????? ?????????? ?????????
?????, ??????, ???????? ?????? ? ????????
?????????????? ???????? ????? ? ?????????
???????????????? ???????? (????????????,
1956; ?????????, 1964; ???????, 1963; ?????????, 1963; ?????????, 1965; ?????????,
1974; ????????, 1981; ????????, 2002 ? ??.).
?????? ??????????????? ? ???????????? ?????????????? ??????????? ???? ????????????
??????? ????????? ??? ?? ?????????????.
?? ?????? ?????? ???????????? ?????? ????? ?????????? ?????, ??????????? ??
???????? ??????? ???????????? ????????? ??????, ????? ? ????????????, ??????????? ?.?. ?????????? (1976) ?? ???????
????????????? ??????????? ???????????
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??????????? ???????? ????????. ?????? ????????????? ??????? ??? ???????? ?????????? ? ???????? «??????? ???????» ?? ????????? ????????, ? ?????????????? ?????????
(?.?. ???????????? ???????? ? ??????????
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?? ?????? ?????? ????????????????? ?????????), ? ????? ??????????? ???????????? ?
???????????? ????????? ??? ?????????????
??????, ?? ?????? ??????????? ? «?????????????» ? ????????? ?????, ??????? ?????-
????????? ????????? ? ?????????? ???? ????? ? ??????? ? ???????????? ?????. ??????
????????? ?????? ? ?? ?????? ????????????
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?????, ???????
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??????? ??????? ?. ??? (? ???????? ???????
???????): 1) ????-????????? ?????????? ?
2) ???-??????? ?????????? ? ??????????? ?
???? ??????????. ?????? ???????????? ?????
?????????????????? ??????? ? ??????????? ???????? ?? 200 ?. ?????? ????? ???????????????. ?????? ????????? ? ??????????
??????? ???? ???????-?????????? ????? ?
??????????????? ?????????? ???????????
? ??????????? ???????????????????? (??????????, 1988 ? ??.).
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??????????) ?????????? ? ????? ????? ??????? ??????? ?????. ? ?????? ???????????
??????? ???? (70 %), ??????? ??????? ??????? ? ??????????. ?????????? ???? (????????? ? ????????) ? ??????????? ??????? ???????? ?????????????? ????????????. ?????
??????????? ?????????? ???????????? ?
????????? ???? (????????..., 1989). ?? ?????????? ????? ?????????? ??????????? ????
??? ????????? ?? ???????? ????????????? ??
????? ?? 500-??????, ? ??????? ???????????
????????? ??????? ????? ??????? ?????????.
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#4#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
??????????? ?????????? ????, ??????? ??????? ?????????. ????? ????????????? ?????????? ??????? ????????? ???????????.
???????? ????? ????? ??????????? ????????
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? ?????????????. ?????? ????? ? ?????????
?????? ???????? ????????????? ??????????,
?????????????? ???????? ???????? ????? ????? ????????????? ????? ?????. ???
???????????? ????????????? ? ????????????? ??????. ? ????? ? ??????? ????????????? ???????????? (?????, ??????, ???? ????? ? ?.?.) ?? ??????????? ?????????? ???????
?????? ??????? ? ????????? ????? ?????? 200
??? ??????????? ?? ??????????? (???, 1974;
???????????, ????????????, 1978).
?????? ???????????? ? ???? ????????? ?? ??????????? ? ???????? ?????????
????????????????-??????????? ???? ??????????? ???????????? ?????????????? ????????????? ?? ????-????????? ??????????
(1) ? ?? ????????????, ????????????? ? ?????????? ????????? ????????? ?????????????
(?????????, ?????? ????????, ?????????????
??????????) ? ?????????? (??????????) ?????
???? ?? ???-??????? ?????????? (2).
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? 9, 3, 1). ??????? ????? ? ??????????? ???????
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??????? ????????? (?? ? 29) ??????????,
???????????????? ?? ???????? 13-??????
????????, ? ????? ??? ??????? 60-85-??????
?????????? (?? ? 14, 26) ??????????????
?????????????. ???????????? ? ???????
???????????? ??????? ?????????? ? ???????????????? ??????????? 80-95- (?? ? 15, 16)
? 120-150-??????? (?? ? 11, 17, 9) ????????,
? ????? ? ???????????-???????? ??????????? 155-180-??????? (?? ? 6, 7, 22) ????????. ???????????????? ???????????? ???????
????????????? ? 180-240-?????? ????????????????????? ??????????? (?? ? 13, 12,
21, 23, 3) ? ?????? ???????????? ??????? ? ?
325-360-?????? ???????-??????????????? ??????????? (?? ? 1, 2, 5). ?? ???-???????
?????????? ????? ? ???????? ????? ????????? ? 180-?????? ??????????. ??????? ?????
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??????? ? ???????????? (50-70-?????? ????????? ? ????????? ?????), ?????????????
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??????????). ? ?????? ???????????? 22 ??????? ??????? ?? ????????? ????-??????????
? 13 ? ???-???????? ??????????.
?? ?????????? ? ????????? ???????
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? ??? ???????? ?????? ? ??????? ?????? ?
????????? (??????? ???????) ? ?? ??????
1,3 ? (???????? ???????), ????????????? ?
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?? ??????? (??????? ?? 5 ??? ? ??????? ???????? ? ?? 10 ??? ? ???????? ????????). ? ???
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?? ?????? ?? ????? ?? ???? ??????? (???????? ? ??., 1990), ????? ????? (?????) ? ????????? ?????? ??? ??????????? ????????. ??
?????????? ??????? ???????? (?? ? 26,
#5#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
27, 28, 9, 3, 1) ???????? ????????? ???????, ?
??????? ????????? ????? ? ??????? ?????,
??????? ??????? ??????, ???????? ?????????
? ????????? ?????? ?????????, ???????? ?????????? ? ????????? ??????? ? ???????
???????????? ??????? ? ????? ??????, ? ??????? ?????? ????????????? ???????????
???????? ???????. ???????? ????? ????????
??????????? ?? ?????????? ????????? ????????? ?????? ? ??????, ?? ??????? ? ???????
??????.
???????????????? ????????? ????? ????????? ?? ???????????? ? ?????????????
?????????? ???????? ???????????? (??????,
1975; ???????? ?, 1989; Smirnova et al., 1999
? ??.). ????????? ? ????????????? ???????????? ?????????? ????????? ?????? ????????
?????? ????????? ???? ????????, ????? ????????? ???? ????????? ? ???????????? ???????? ?? ??????? ?????? ??????? (??????,
1970; ?????????, 1972; ???????, 1989). ???????? ?????????? ???????????? ??????? ? ?????
?? ????????? ??? ????? ? ?????? ????????????? ???????? ?????? 400-??????? ????????
?? ???????, ?????? ???????? ?????????? ??
??????????????? ???????? ??? ?????????
????????????????? ???????. ? ???????? ????????? ???????????????? ?????????????????
????????? ???????? ????? ???? ???? ?????
??????????????? ????????, ????????????
????????? ???? ????? ? ??????????? ???? ?????????? (?????????, ??????, 2002?, 2009).
? ?????? ????????????? ??????? ?? ???????????? ????????? ???????? ?? ??? ?????? ??????????? ? ??????????, ??????????
? ?????? (???????? ?, 1989; Smirnova et al.,
1999). ? ?????? ?????? ??????????????? ????????? ???????? ????? ??????? ??????????????
?????????, ??????? ?? ?????? ???????????? ?????????? ? ?????????? ?????? ???????????. ????? ??????? ?????????????? ????????? ??????? ?????? ??????????? ? ????
??????????? ????????? ????? ?????????
???? ??????????????. ??? ?? ???????? ?
???????? ???????????????? ?????????.
?????????? ? ??????????
?????? ????????? ???????? ????? ?? ?????? ???????: ?????????, ???????????????,
???????????? ? ????????????????, ??????? ??????? ?? ???????????????? ?????????
(??????, 1975; ???????? ?, 1989; Smirnova et
al.,1999), ??? ?????? (????????, 1976). ????????? ? ???? ??????? ?????????????? ?? ????????? (???????? ?, 1989), ??? ???? (????????, 1976). ? ?????????? ????? ??????????
?? ???-??????? ???????-????????? ???????
???? ???????? 4 ??????? (?????????????,
???????????????, ???????????? ? ????????????????) ? 11 ???????????????? ?????????.
????????????? ?????? ???????? ???
?????????: ????????????? ? ?????????.
????????????? ????????? (em) ? ???????????? ? ???????? ????????, ???????? ? ??????? ?????????????? ?????????? ?? ???????
??????????? ????? ????????. ???? ??????? ?
????? ?????? ?.?. ?????????? (1972) ?? ????? ??????? ????????? ????? ?????? ????? ?
???????? ?????? (?? ??????? ???????). ??
?????????????? ? ?????? ????????????? ????? ???????? ?????. ?????? ?????? ???????????? ?? ????? ?????, ????? ? ?? ????????? ????? ? ????? ? ???????? ?????????. ?
???????????? ????? ??????? ???????? ??????
????: ??????????????, ????????????????,
????????????? ? ?????????????????. ???????????????? ??????? ? ?????? ???????
????? ????? 5-7 ??, 8-14 ?????????, ?????????? ? ???????? ?????? (???. 1, sm). ???????????? ???????? ???????? ? ??????????? ??
???????? ??????? ?????????? ?? 7 ?? 12 ??????: ???????? ? ????? ?????? ?????? ????
?? ????? ????-?????? ???????? (?????????,
1972; ???, 1974). ?? ????? ??????, ? ??????-
#6#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
???. 1. ??????? ??? ?????? ????? ?????????? ??????? ????????????????? ????????? ? ?????????????????????????? ???? ??????????? ???????????? ? ??????? ??????? ?????. ?? ??? ??????? ? ????????????????
?????????: sm ? ?????????? ??????, p ? ??????????????? (????: p-1 ? ?????????????, p-2 ? ?????????,
p-3 ? ?????), j ? ?????????? (???? j-1 ? ?????????? ??????, j-2 ? ??????????? ??????), im ? ??????????,
v ? ????????????, g1 ? ??????? ????????????, g2 ? ?????? ????????????, g3 ? ?????? ????????????; ?? ???
??????? ? ?????? ???????? (?, ?, ? ? ??; ? ? ?)
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
????? ???????????? ????? ???????? ? ??????? ?????????? 4-6 ??, ????????-????? ?????
?????? ?????? (??????) ????? 0,191-0,211 ? ?
????? ????? ? 0,155-0,400 ? ? ??????? ?????.
??? ????????????? ?????? ?????? ???????
(?????????, 1972; ????????, 1984 ? ??.), ?????????? ??? ???? ?? ???????.
?????????? ? ??????????? ????? ?????
???????????? ???????? ???????? ????????
? ??? ?????????????????. ?????????? ?????
?????????? ? ????????? ????? ???????? ?
????? ??? ?????? ????? 2/3 (?? ??????, 1970).
???? ????? ????? ? ?????? ???????? ????????? ? ????? ???? ?????????? ? ?????? ???????????? ?? 8 ?? 90 % ?? ???? ?????????????
?????. ????? ????????? (? ???????????????
???? ?? 31 %) ??? ??????????? ? ????? ?????
?????????????????? ????? (??????, 1970;
?????????, 1972) .
????????? ????????? (sm) (??? ????????? ?????? ?? ???????????? ????????????
??????????) ? ????. ??? ?????????? ? ?????
???? ? ??????? ??????????? ????? ????????
? ????? ? ????????????? ?????? ?????????? ???? ???????????? ?????. ?? ?????? ?.?.
??????? (1970), ????? ??? ?????????????
????? ?????????? ????? ? ??????????? ???
????? ????????, ??? ????? ? ?????????????
?????? ? ???????. ??????? ? ??????? ??????????? ?????? ????????? ? ?????????? ?????
??? ?????? ???????? ???????.
????????????????? ?????????????? ??????? ?????????? ???? ????? ????.
??????????????? ?????? ????????
?????? ???????????????? ?????????: ???????????????, ??????????, ?????????? ?
????????????. ??????????????? ?????????
(p) (?????????, ??? ????? ? ??????? ??????
???????) ?????????? ? ??????????? ?????? ?
????????????? ????????????? ??????. ???
???????? 4 ????: 1) ?????????, 2) ????????????? ? ??????????? ??????, 3) ????????? ?
????????????? ???? ? ??????? ? 4) ????? ?
??????? ? ???????????? ??????? ???????
(??????????, 1981; ???????, 1989). ????????
????? ? ?????? ??? ????? ??????? ?.?. ??????? (1964, 1975), ???????? ?? ???? ?????????
????. ??? ????????????? ?????????? ??????
???????????? ??????? ????? ???????????
???????? ??????? (???. 1, p-1). ? ???? ????????? ?????? ????????? ???????, ???????
??????????? ? ????? (???. 1, p-2), ????? ?????????? ?????? ?????? ? ??????????. ?????
????? ?????????? ??????????? ? ???????????? ?????????? ??? ??????????? ????????????? ?????????. ? ???? ?????? ??????????
????????????? ??????????, ???????????? ????????? ?? ???????? ?????? ? ???????????
??????????, ???????????? ???? ???????? ????? ? ?????????? ???? ??????? ??????. ???????????? ????? ????? ??????, ??????????,
???????? ? ???????????? ????? (???. 1, p-3).
??????????? ???? ?????????? ? ?????????
? ??????? ????. ????????????????? ????????? ? ????? 3-4 ?????? (? ????? ??? ?? ?????
????).
?????????? ????????? (j) ?????????? ?
????? ????????????? ? ???????????? ????? ??????????? ?????? ? ?????????? ????
?? ???. ????????? ???? ???????? ????????
?????????????? ??????????????? ???? ???????? ???????, ??? ??????? ????? ???????? ?????????? ? ?????? ?????? ??????????
???????????? ??????? ? ??????????? (Mirov,
1967), ?? ??? ????????? ???????????? ?? ???
????, ??? ?????????: ?????????? ? ??????????? ??????.
?????????? ??????. ?????????? ?????
??????????? ??????????????? ?? ????? ??????
? ?????? ????????????? ????? ? ???????????? ????? ??? ? ?????????? ?????. ?? ?????
???????????, ??????????? ?????? ???????
????? ???????? ????? (?? 4-6 ???? ? ????????????? ????? ?????? ?? 1-3 ? ??????????).
#8#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
???????????????? ?????????? ?????? ? ????? ??????? ?????????????? ?????? ????? ?????????? ?????? 5-15 ??, 9-13 ?????????, ??????????????? ?????????? ????? ? ??????????
????? ? ???????????? ???????? ??????, ?
????? ???????? ???????, ????????? ?? ???????? ? ??????? ?????? (???. 1, j-1). ?? ??????????? ?????? ??????? (????????? ? ??.,
1967; ???????, 1975), ? ?????? ????? ?????????? ??? ?????? ????? ??????? ??? ? ??????????? ???????. ? ??????? ??????? ?????
?????? ???????? ??? ?????????? ??? ???????
???????? ?????????? 1-5 ??, ????????-?????
????? ???????? ????????????? ? 1,5-2 ????
(0,28-0,67 ?), ???????? ?????? ?????????????
(????. 1). ???? ??????????? ?????? ???????????? ? ?????? ???? ???????? ???? ?? ?????
?????????? ????.
??????????? ??????. ?????? ?????????? ???? ???????????? ????? ?????????????,
???????? ?????? ?????? ?????. ?? ??? ????????? ??????????? ?????????? ?????? ? ?????????. ? ??????? ?????????? ??????????
??????? ??????????? ?????? ? ????? ????????? ???? (???????????). ???? ???? ??????????????? ????????, ??????? ??????? ??????
(??? 0-?? ??????? ?????????) (???. 1, j-2). ?
??????????? ? ???????? ??????? ????????
?????????? ? ?????? ????? ???????? ?????
???????? ? ??????????? ? ???. ??? ???? ?
??????????? ?????? ?????? 6 ??? ?????????? ??????????? ????? ? ?????? ????????????
?????? ? ?????? ????? ????????. ? ???????? ??????? ??????????? ? 7-8 ????? ??????????? ???????????? ???????? ???????, ?
??????? ??????????? ????? ??????? ?????
??????????? ??????? (?????????, 2002).
??????????? ????? ?????????????? ?????????? ? ??????? ?????????? ???? ? ?????????? ? ?????????? ????? ??????? ?????????
?????? (???????, 1975; ?????????, 2002) ?,
??-????????, ????????? ????? ??????????-
??? ?????? ????????? ????? ? ???????? ?????
(??????????, ???????, 2002). ??? ??????????
???????????? ????? ?? ????? ????????? ?????
? ??????? ????? ?????????????? ??????????? ??????????? (??? 1-?? ???????). ?? ?????????? ???? ???? ?????? ????? ?????????
??????, ??????? ? ?????????????? ????????
???????? (????. 1). ????? ??????? ? ???? ???
(0,45-2,55 ?) ? 2-3,5 ???? ????????? ????? ?????????? ? ? 3-6,5 ???? ????? ?????.
??????????? ????? ?????????? ? ??????? ???????, ????? ???????-??????????????.
????????????????? ????? ????????? ??????
5-15 ???. ? ??????????????? ???????? ??? ???????????? ?? 25-40 ???, ? ????? ????? ?????
???????? ?? ????????????? ???????. ??????
?????? ??????????? ??????????? ??????????
? ???????? 3-4 ????, ?? ? ???? ?????? ?????????? ???????????? ????? ?????????????????? ?? 5-10 ???. ???????????????? ??? ??????? ? ??????????????? ????????? ????????
???????, ??? ??????????? ???????? ???????
???????????? ????? ? ?????????? ?????????????.
?????????? ????????? (im) ??????????
? ????????? ???????? ? ?????????????????
?????????? ????????? ????? (??????, 1975;
??????????, 1981; ?????????, 1989; ?????????, 2002). ????? ????? ????????????? ???
???????????????????, ?????? (???. 1, im).
? ?????? ???????? ?????? ?? ????????????
?????? ???????? ??????????? ???????????
??????????? (1-3 ????? ? ????? ? ???????).
?????????? ??????????? ????????? ????????? ?? ???? ?????????? ??????? ???? ?? 2-3.
???????? ??????? ?????? ???????: ????? ????????? ?????? ??????????? ? ??????? ???????? ? ? 2-3 ???? ????????? ?????? ?????.
?????? ? ??????? ???????? ?????????. ????????? ??????? ? ?????? ????????? 13-17 ?? ?
?????, ?????? ??? ??????????? ??????? ????
(????. 1) ??-?? ?????? ???????????? ???????
#9#
0,01-0,14
0,4-0,9
1-12
10-15 (12)
9-14 (12)
lim
27
28
11-15
59-115
9
lim
2,6-19
8-14
12-19
12-18
3,4-6,5
2,6-4,3
0,2-2,4
1
4,5-28
8-17
12-28
9-19
10,5-19
8-12,5
5-93
?
7-103
4,5-8
0,4-5
1,2-4
1,3-5
0,4-2
0,6-1,2
< 0,6
0,01-0,6
?
???????2, ??
0
0
0,02-1,7
0,3-1,4
0,4-1,7
0,02-0,7
0,06-0,17
?
2,4-13
4-8
7-13
7-12
0,7-4,3
?
3,5-4,3
?
1,3-2,8
?
0,7-1,4
3,0-5,6
5-8
?
2,4-4,1
???????????? (v)
0,2-2,1
0,4-2,2
0,5-2,1
0,2-0,9
0,2-0,7
?????????? (im)
0
0
0
0
???????, ?
?????????? ( j)
?????????????, ?
?????
3,5-33
3,5-33
5-21
13-33
8-25
24-30
13-19
?
3-14
?
3-14
?
4-13
0,1-6
1-6
0,1-1
? ??????, ??
0,6-5,0
1,0-3,2
0,6-5,0
1,4-5,0
1,2-3,2
1,7-2,8
0,7-1,2
?
0,1-0,9
?
0,1-0,9
?
0,1-0,5
0,05-0,35
0,09-0,33
0,05-0,35
?? ???????, ??
???????? ??????4
0,8-18
1-8
1-18
1,5-12
2-8
1,5-3
0,8-1,8
?
0,01-0,9
?
0,1-0,9
?
0,011-0,084
0,0004-0,026
0,003-0,026
0,0004-0,007
?? ???????, ??2
??????????. ? ?? ? ????? ??????? ???????, ? ??????? ? ??????? ???????? (? ???????) ????????, ??????? ? ?????????? ??????, 0 ? ?????????? ????????, lim ?
??????? ???????? ???????? ? ???????? ?????? ????????????????? ????????? ?? ?????? ?????????; 2 ? ??????? ?????? j-im ?????? ??? ? ?????????, v-g ?????? ?
?? ?????? 1,3 ?; 3 ? ? ????????? ??? ??????? ? ????????? ??????, ? ??????????? ? ?? ?????? 1,3 ?, 4 ? ???????? ? ??????, ?? ??????? (??? ?????? ??????) ? ??
??????? ?????? (??? ??????? ??????????? ??????? ??????) ???? ??? ??????? ?????????????? ?? 5 ??? ? j-im ?????? ? ?? 10 ??? ? v-g ????????.
88-106 (98)
100-115 (109)
17
82-94 (86)
59-81 (72)
15
77-103 (92)
69-78 (73)
26
11
38-108 (63)
14
16
7-10,5
9-47
lim
0,5-2,4
14
0,5-2,3
15-42 (31)
27-47 (37)
26
0,2-1,1
0,01-0,14
0,01-0,05
1
2-12 (7)1
28
??????, ?
28
???????, ???
?
??
??????? 1. ?????????????? ???????? ????? ?????????? ???????????????? ??????? ?????????? ? ?????????? ??????????? ? ??????????? ????-??????????
??????????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
? ?????. ? ?????????? ?????? ?? ?????????
? ??????????? ? ??????? ????????????????? ????? ????????? ????? ????????? ????????????? ???????? ????? ?????? ?? ??????? ?
??????? ?????? (?????????, 2002; ?????????, ??????, 2009).
? ???? ????????? ???????? ??? ???? ?
?????????? 1 ? ?????????? 2, ????????????? ???????? ???????????????? ????? (???????????????? ?????? ? ??????????????
??????? ???????), ????????? ????????? ?
?????? (3-10 ? 10-25 ??) ? ??????????? ?????? ?????????? (???????-?????????????? ?
????????????? ??????????????). ????????????????? ????????? ? 25-50, ??? ?????????
????????? ? ?? 80 ???. ?????????? ?????????
?????? ??????????? ? ???????? 40-60 ???.
???????????? (???????? ????????????)
????????? (v). ??? ??????? ??????????? ?
?????? ???????? ??????? (???. 1, v). ?????
???????????? ?????????? ????????? ?????????, ??????????????? ?? ??????? ????? ?????????????, ? ?????????? ?????????? ? ?????????? ????????????? ??????.
? ????? ????????? ? ????????????? ??????
?????? ????????? ?????? ?? ????????? ????? ??????????? ????????? «????????»
??????? ? ????????????? ????????? ? ?????????. ???? ?? ?????????? ?????? ?????????? ???????? ??????????? ????????? ?????
???????????? ?????????? «?????????» ??
??????? ????????? ??????? ?????????, ??
?? ???? ????? ?????? ? ?????? ???????????? ????????? ????? ????? ?????????, ???
?????????? ???????? ?????????? ??????????, ??? ???? ??????????? ??? ?????? ???????????? ????????? ??????? ?????????. ????? ???????????? ????? ???????????????
???????? ?????? ??????? ?????? ? ??????
?????? ????? ? ?????????? ????????????,
????????????? ???????? ??????????? ????????? ?????.
????????? ??????? ????????? ??????????? ?? 4-?? ???????. ?? 10-30 ??? ?? ???????? ???????????? ???????? ? ???????
???????????? ? ??? ?????????? ???????????
? ???????? ???? ???????????? ??????? ??
?????? ?? ????? ? ????????? ? ???????. ??????? ? ?????? ????????? (?????? 3-7 ?, ??????? 3-8 ?? ? ?????) ???????? ? ???????????
????? ?????????, ? ?? ?????? ???????? (8-19
? ? 8-28 ?? ??????????????) ? ?? ?????? ????
???????????-??????????? ????. ?????????
??????? ? ?????? ?????????? ????????????
?? ???? ????????? (?? 25-50 ??), ? ????????????????? ???? (?? 20-33 ??) (????. 1). ????????????? ???????????? ???????? ?????
????????? ???????? ??, ??? ??????????????
????????, ??????????????? ????????????
??????????? ???????, ? ????? ???????? (??????? ? ?????? ? ?? ???????) ????? ??????
????????? ???????????? ??????? ? ?????
????????????? ??? ?????? ???????? ????????????? ????????? (?????????, 1998; ?????????, ??????, 2002?,?, 2009).
? ???? ????????? ???????? ??? ????
(???????????? 1 ? ???????????? 2), ??????? ??????????? ??????????? ?????? ??????????. ??????? ?????? ???? ? ?? ?????????????? ????? ????????? ? ?????? ?????
?????????? ?????, ??????? ?????? ???? ??
??????????? ????? ????????? ? ????????.
????????????????? ????????? 30-80 ???, ???
?????? ??????????? ? ???????? 70-160 ???.
??????? ????????, ??? ????? ?????, ?????????? ?? ????? ??????????? ???????????? ?
??????? ? ??????, ?? ?????????? ????? ???????????????? ??????? ?????? ? ???????????
??? ?? ???????, ????????? ????????? ????????? ? ?? ??????? ?? ??? ???, ???? ?? ??????
? ???????? ?????. ????????????????? ???????????????? ??????? ? ??????? ??????? ?????
????????? ?? 70-120 ??? ??? ??????? ??????
?? 100-160 ??? ??? ??????? ?????.
# 11 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
???????????? ?????? ???????? ??? ???????????????? ?????????: ??????? ????????????, ?????? ? ?????? ????????????.
??????? ???????????? ????????? (g1).
???? ????? ????????? ?????????? ?????
????? c ???????????? ?????? ???????? ? ?????? ????????? ? ???????? ? ????? (???. 1, g1).
??????? ????? ????? ???????????? ?????????? ????????? ?????????, ?????? ? ??????
?? ?????, ? ????????? ???????? ??? ???????????? ????????? ??????, ???????? ???????????? ? ????????????? ? ???????? «?????»
????????? ??????. ??????????? (???????? ?
??., 1989; ??????????, ?????????, 2000), ??? ?
??????? ????? ????? ??????????? ? ???? ??????? ? ?????? ????????? ????? ???????????.
? ????????? ??????????? ???????? ??????
????? ? ?????? ? ?? ???????, ????????? ???????????? ????????, ???????? ??????????
????????? (????. 2).
? ??????????? ???????? ? ?????? ????????? ?????? ????????? ??????? ?????
(?????????????, 1-4 ??. ?? 10 ???) ????????????? ?? ?????? 1-?? ??????? ? ??????? 5-7
???????? (??????????, ?????????, 2000), ?
?????????? ???????????? ?????????? ?????
?????????? ? ???????? (????. 2). ?????? ??????? ????? (?????????????) ?????????? ?
??????? ? ?????? ?????? ????????? ????? ?
?????????????? ??????????? ? ? ??????????
????????????. ? ???????? ????????? ?? ????? ?????????? ?? 7-8 ?????. ??????? ????????,
??? ??????? ???????????? ???????, ?? ????????? ?? ???????, ???????? ?? ?????? ?????
?????? ???????????? ? ????????? ?????,
?? ? ??????? ???????? ?? ???????????? (????? 50 % ? ????????????? ? 80 % ? ???????
???????????). ?????????? ?????? ???????
?????????? ?? ?????? «????????» (?????????,
1972; ???????? ? ??., 1989; ?????????, 1998;
?????????, ??????, 2002?), ??? ??????? ? ??????????? ??????????? ?????? ? ???????-
?????? ?????????????? ??-?? ????????????
???? ????????? ??????? ???????????? ???????? ? ?????????? ?????? ??????????? ????? ????? ?????? ???????? (?????????, 1998).
??????????? ???? ?????????? ? ?????? ????
????????? ? ???????????-?????????? ???? ?
???????? ???? ? ???????????? ????. ????????????????? ????????? ? 50-90 ???, ??????????? ??? ? 150-210 ???.
?????? (????????????????) ???????????? ????????? (g2). ????? ????? ??????????
?? ?????????????? ??? ???????? ? ????????
???????? ? ?????? ????????? ?? ??????????????????????????? ?? ?????? ??? ????????
(???. 1, g2). ????????? ???????? ?????? ???????????? ????????? ?????? ? ????? ???????
????? ????? ? ???? ?????? ??????????? ??????????? ??? ????? ???????????????? ???????. ???? ????????? ??????? ? ????? ????????? ?????????? ????? 80 %. ??????? ????
????????? «??????» ???????????, ?????????? ??????? ????? ?????? ?????? ??????? ???????? ?????. ? ?? ??????? ??????????
??????? ???????????? ?????? (??????????,
?????????, 2000). ??????? ????????? ??????????? ????????? ??? ??????? ?????? ?????? ?????? (22-31 ?), ????????????? (16-23 ?) ?
???????? (6-7 ?) ?????. ???????? ? ?????????? ???????? ?????? ?????????? ?????????,
? ???????? ?? ??????? ?????? ???????????
?? ??????? ?????? (????. 2).
????????????, ??? ???????, ??????????
? ????????. ? ?????? ???????? ??????????? ????? ????????? ????? ????????? ???????????? ???????: 70-280 ???? ?? ??????
(????. 2). ? ???????????? ?????? ?????????
????? 1-3-?? ???????? (??????????, ?????????, 2000). ? ???? ????????? ??? ????????
????????? ?????????????? ???????? ???????????? ????????? (?? ????? 1/3) ?? ??????
? ?????? ???????? ? ??????????. ????????? ?????????????? ????????? ????? ???-
# 12 #
21-27
320-380 (349)
330-400 (363)
360-460 (393)
320-460
1
2
5
lim
?????????? ? ??. ????. 1.
24-29
150-270
lim
20-29
20-29
19-31
22-28
20-23
233-257 (242)
210-270 (244)
3
24-31
19-22
24-26
20-26
22-24
15-25
18-25
18-21
12
225-242 (234)
150-170 (160)
6
23
150-158 (156)
7
172-190 (181)
71-200
22
lim
184-213 (200)
170-200 (182)
6
21
19-24
143-163 (153)
140-164 (157)
7
13
15-21
147-162 (153)
9
17-24
15-18
95-116 (106)
106-130 (114)
15-19
9
71-96 (84)
11
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?
17
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??.
33-62
33-59
45-62
34-62
24-59
32-59
34-39
30-59
24-42
33-44
30-52
32-42
15-45
21-36
18-33
24-45
19-31
15-21
25-33
16-28
???????,
??
12-17
13-16
11-19
11-23
14,5-18
3,3-5,7
1,9-7,2
1,9-6,2
4,0-7,2
2,5-6,6
3,3-6,3
3,1-6,7
3,6-6,3
12-23
14-21
15,5-21
12-23
1,9-9,0
1,9-6,1
3,5-6,1
2,0-9,0
?????? ???????????? (g3)
11-23
11,5-19
15-18
?
1,7-5,9
3,0-5,4
1,7-4,6
2,3-5,8
3,5-4,0
?
2,9-4,7
2,0-5,9
?????? ???????????? (g2)
6,5-18
6,5-18
6,5-12
9-18
11-14
12
???????,
?
??????? ???????????? (g1)
?????????????,
?
?
3,0-16,0
?
4,5-14,0
3,0-16,0
1,8-14,0
2,0-9,5
6,0-11,0
1,8-14,0
2,0-5,0
4,5-9,0
5,0-8,5
4,0-5,5
0,3-7,9
0,7-5,0
0,3-3,5
2,0-6,0
1,2-3,5
0,3-1,5
0,3-7,9
3,5-7,5
?????????????
???????????? ?????,
?
??????,
????? /
??????
11-274
18-92 (56)
31-233 (98)
11-274 (76)
9-276
27-108 (62)
68-190 (135)
30-118 (56)
9-74 (31)
32-243 (95)
60-276 (116)
63-156 (96)
0,1-59
0,1-25 (8)
0,1-59 (19)
7-44 (26)
0,6-17 (9)
0,2-4 (1)
0,3-27 (9)
1,2-23 (10)
? ??????,
??
1-5
1-5
2-4,5
1,5-4
2,5-11
?
3-5
2,5-6
3-5
4-7
5-11
8,5-10
4-28
4-13
8-22
12
6-16
10-28
5-16
9-26
???????? ??????
0,4-1,6
0,6-1,0
0,5-1,5
0,4-1,6
0,8-5,6
0,8-2,4
1,2-1,6
0,5-3,2
1,0-3,3
0,9-2,0
1,2-5,6
2,3-3,4
0,8-8,4
0,8-4,4
1,6-4,0
1,4-4,6
1,4-3,4
1,4-4,0
1,7-4,0
2,7-8,4
??
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??
?????
3-14
3-7
4-13
3-14
4-30
4-20
6-8
4-17,5
4-12
4,5-11
8,3-30
10,5-20
3-24,5
3-22
6-15,5
5-23
4-14
4-9
7-14
6-24,5
??
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??2
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??????????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
??????? ??????? (????? 33 ???. ??/??????)
??????????????? ?? ???? ??????? ?????????
????? ?????. ??????????? ???? ?????????? ?
???????? ?????????. ????????????????? ????????? 90-110 ???, ??????????? ??? ? ???????? 270-320 ???.
?????? ???????????? ????????? (g3).
????? ???????? ????????? ?? ????? ??-?? ??????????? ? ???????? ?? ????????? ?????????. ? ??? ???????, ????? ????? ??-????????
???????????,
???
?????
?????????????????????????? ????? (???. 1, g3). ????????? ??????? ????????? ???????????
?????? ? ???? ????? ?????????????? ???? ?
????? ??????? ????? ?????, ? ???????? ??
????? ?????????? ????????? ??????? ?????????. ???????? ????????? ?????????????
???????? ????????? ?????? ???????? ??????? ??????????, ? ????? ???????? ??????
???????????? ? ????? ???????. ? ???? ??????
?????????????? ???????? ????????? ??????
?????? ?? ?????? ?????? ????????? ?????
(??????????, ?????????, 1996; 2000; ?????????, ??????????, 1997). ? ?????????? ??????
???????? ?????????? ????, ? ??????? ?????? ?
??????? ????? ?????????? ???????? ?????????????. ???????? ????????? ? ?????? ? ??
??????? ?????? ????????? ?? ??????, ???????????? ??? ?????????? ? ???? ??????????
??????, ?? ??????? ?????? ? ?????????????
?? ????????? ? ???????? ? ??????? ????????????? ??????? (????. 2).
???????? ?? ???????? ?????????? ????????? ????? ?????, ????????? ??????? ????? ???????? ???????? ??????? ? ??????????. ???? ?????? ??????, ??? ? ? ??????
????????, ?? ????????? 1/3 ?? ????? ???????????? ??????????????. ? ?????????? ??????
(90-270 ????? ?? ??????) ? ???????? ? ??????
???????? ?????? ??????????? ?? ?????? ?????? ???????? (????. 2). ?????????? ?????????????? ????????? ?? ??????, ??????????-
?? ??? ??????? ???????????? ???????? (?????
8 ???. ??/??????). ?????????? ?????????????
???????? ? ???? ????????? ?????? ????????
?????? ????????? ????? ????? ? ??????? ??????????? ?? ???? ??????????? ?????? ????????? ?????, ???? ???????? ????? ???????
????? ??????????? ?? ????????? ???????,
? ? ????????? ?????????????? ???? ????????? ? ????????? ?????? ????? ???????? ?????????. ??????????? ???? ?????????? ? ???
???????? ????????? ???????. ??? ?????????
?????? 80-150 ??? ? ??????????? ? ????????
????? 400-460 ???.
? ??????? ??????? ????? ????????????
?????? ? ?????????? ????? ???? ? ????? ?????????? ?? ????? 300 ???. ?? ??? ??????????
???? ??????? ?????????????? ????????? ???????? ?????????? ? ???????????? ??????
?????????, ???????? ???????? ????? ?????????, ? ? ????? ????????? ? ??? ???????? ???????.
??????????????? ????????????? ????????? ? ???????? ???????? ????????? ?????? ????????? ????????????? ??????? ??
????- ? ???????????? ??????????? ?????,
?.?. ?? ??????? ?? ????????? ????????????
?????????? ?????? ?????????. ???????????
?????? ????? ????????? ???? ?????????
???? ?????? ??? ????? ??????????? ?????
??? ?? ????????? ??????? ????????????? ?????????. ???????? ???????? ?????????????? ?????????? ????????? ????????? ? ???????? 260-350 ??? ??-?? ????????????? ?????????
??????? ??????? ? ?????????????? ???????????? ???????? ?????? ???????? (???????,
2007 ? ??.). ?? ????-????????? ??????????
????????? ?? ???????? ????????? ????? ?????????? ? ???????????? ???????? ? ????????
60-180 ???, ????????? ?? 30 % ??????. ? ???????????? ???????? ??????? ?????? ? ???????
?????????? ?????? ????????????? ? 60-70 %
? ???????? 200-240 ??? ?? 70-100 % ? ????????
# 14 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
270-460 ???. ??????? ??? ??????????? ???????? ????? ????????? ? ???????? 450-500
??? ? ???????? ????? (???????, 2007) ? 480520 ??? ?? ????-????????? ? 500-550 ??? ??
?????-??????? ??????????? (???, 1974; ???,
????????, 2007) ??????????? ?????? ????????? ??????? ?????.
????????? ????????????????? ???????
? ???????? ????? ????????????. ???????? ??
??, ??? ? ???????????? ???????? ????? ????? ???? ???? ??????? ???????-???????????
700-??????? ???????? (?????????, 2007), ??????? ?? ?????? ??????????????? ??????????????? ????? ????????? ????????? ?????????? ????????? (s), ???????? ???????????? ?
????????????? ???????????????? ?????????
(??????, 1975; ???????? ?, 1989; Smirnova et
al., 1999) ??? ?? ???, ????????? ????????????
??????? ? ????? ???? ???????????, ??? ?? ???????????? (?????????, 1972; ????????, 1981;
?????????, 1985?; ? ??.), ??? ? ?? ????? ?????? (?????????, 2007; ??????? ? ??., 2009),
?? ????????? ??? ?????. ??????? ??????? ????? ????????????????? ??????? ?????????? ?
?????????? ???????????, ???? ?? ????????
?????? ?? ?????????? ???????????? ???????,
? ?????? ???????????? ?? ????????. ? ?? ??
????? ????? ????????????????? ??????? ?????????? ?? ???????????? ?????? ????????????? ????????? ????????? ??? ???????????????? (??????, 1975; ???????? ?, 1989). ? ????
?????? ?? ????? ?????????, ??? (1) ????????
????????? ?????, ?????? ?? ?????? ??????
??????? ? ?? ????????? ????? (??????????, ?????????, 2000), (2) ?????? ????????
????????? ?? ??????? ??????????? ???????
?????? ? ????????? ??????????? (????????? ?
??., 2008), (3) ??????? ??????? ?? ????????
?????? ??? ?????????? ?????????????? ????
?????????? ??????? ? ?????? 1/4-1/3 ??????, (4) ?????? ????????? ?? ????????? ????,
??????????? ????? «?????» ? ???????, ?
????? (5) ?????? ????????? ?????????? ???????????? ??????? (??????? ???????? ???????????, ??????? ? ?????????? ???????? ??
????????? ?????? ??? ???????????), ????????? ?? ????????????? ???????? ????? ??????
400-460 ??? ??????? ????? ??????? ? ??????
????????????????? ???????.
???????? ???????????? ??????? ???????? ?????, ???????? ??? ????????? ? ?????? ???????????? (???, 1974; ???, ????????,
2007), ????????????????? ?????????????????
??????? ???????????????? ????? ??????????
80-120 ???, ? ???? ????????? ???? ?????????
? ??????????? ???????????? ??????? ?????
????????? ? ???????? 480-550 ??? (????. 3). ??
?????????? ????? ??????? ???? ??????? ?????????????? ????????? ? ?????? ??????????? ??????????? ?????????? ?????????, ?
?????????? ? ????????? ??????? ?????, ????? ? ??? ??????????? ?????????.
? ??????? ????? ??????? ????? ???????
? ???????? ??????????? (???????-????????)
????????????? ????????? ????? ??????
??????? ??????????? ????? ??????? ?????????? ???????????? ? ???????????? ????. ?
???????? ???????????????? ??????? ????????? ???????? ??????????? ???????? ????????????, ????????? ???? ? ??? ????? ??????
??? ??????? ?????????? ?????, ? ???????
????? ??? ?????? ?????????? ????????? ????????? ?? ??????? ?????? ? ???????? ?????.
??? ????? ????????? ? ?????????? ????????????????? ???????????????? ??????? ??
200 ??? ? ?????, ??, ??-????????, ?? ????????
???????????? ???????? ???????? ?????????? ????????? ???????? ??? ???????? ? ??????????? ?????? ??????????. ?? ???????????? ????? ?????????? ? ???????? ??????????
??????????? ?? ????????? ? ??????????
??????????? ??????? ????????? ? ?????????? ???????????? ???????, ? ???????? ?????? ??????????? ? ????? ????????? ??????
# 15 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
??????? 3. ????????????????? ?????? ?????????? ????? ?????????? ?????????? ? ??????????
??????????? ?? ???-??????? ???????-????????? ??????? (? ???????? ??????? ???????)
??????? ?????, ??? ????
?????? ? ????????????????
?????????
?????????????
1) ?????????????
2) ?????????
???????????????
3) ???????????????
4) ??????????
5) ??????????
6) ????????????
????????????
7) ??????? ????????????
8) ?????? ????????????
9) ?????? ????????????
????????????????
???? ?????????
???????
???????
(????????????)
0,83
0,83
0,15-0,25
0,15-0,25
0,75
0,83
70-1601
70-160
0,25-0,33
0,25-0,33
5-20
5-20
25-50
38-60
30-80
70-160
290-300
400-450
50-90
150-210
90-110
270-320
80-150
400-450
80-120
480-550
480-550
480-550
?????
?????????????
?????? ????????
0,83
0,83
0,15-0,25
0,25
0,75
0,83
50-70
50-70
0,25-0,33
0,25-0,33
4-15
4-15
0,83
0,83
0,15-0,25
0,25
0,75
0,83
20-35
20-35
0,25-0,33
0,25-0,33
0,17-3
0,33-3
20-40
40-70
15-20
19-37
?
?
50-80
100-140
(>80)
(>190)
>13-21
(>40)
?
?
?
?
?
?
?
??????????. ? ????????? ? ?????????????????, ? ??????????? ? ??????? ????????? ????????? ???
???????, ???; 1 ? ?????? ??? ???????? ??????? ?????????????? ?????????. ???????????? (ss) ? ?????????
(s) ????????? ? ??????? ?? ???????.
?? ??????? ??????????. ???, ? ?????????????
???? ?????????? ?????? 240 ??? ???????????
???????? ????????? ? ???????????? ???????
(v/g2, v/g3), ? ??????? ? ?????? ?????? ?????????? ????????????? ?????????????, ?? ?????? ? ????????? ??????????? ????????????
??????.
? ????? ????? ? ???????????? ???????????, ??????? ??????? ?? ?????????
200 ???, ????????????????? ???????????????? ????????? ??????????? ?????? ?? ??????? ????????????? ?????????. ?????????????
?????????? ?????? ????????????? ? ?????????? ?????????????? ???????? ???????? ?????, ???????? ????? ? ????????? ????? (???,
2004). ????????? ????? ?????????? ???????
????? ? ???? ??????????? ???????? ??? ?????????? ???????, ? ?? ?????????????????
# 16 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
????????????? ????????????? ??? ???????
(????????????) ?????????. ????? ????, ???
??????? ????????? ???????? ???????? ????????????? ????? ?????????? ? ????????????
????, ??????? ??? ????? ??????? ??? ??? ????? ? ????????, ????????? ??????? ??????????? ??????????? ?????? ?????????? ?? ????????? ? ???????? ????????????. ? ??????
????????? ??????????? ??????? ???????? ??
?????? ???????????, ?? ? ????????? ?????
??????????, ??????? ???????? ??????? ?????
??????????? ? ????????????? ??????? ?
???? ???????? ??????????????. ? ???????????? ?????? ????? ??????? ???????? ? ???????? 40-70 ??? ? ????????????? ??????????? ? ?
19-37 ??? ? ?????? ????????? (????. 3). ? ??????????? ????????????? ??????????? ??????
???????????? ??????? ????? ????? ???????
140-190 ???. ? ?????????? ???????? ???????????? ? ?????? ??????? ???? ????????? ???????
????? ???????? ????? ????????, ?? ?????????
??????? ????????????? ?????????.
?????????? ?????????????? ?????????
?????????????? ??????????? ???????? ?????,
? ?????? ???????, ???????? ????? (??????? ?
??????, ?? ???????, ??????? ?????? ???????
? ?.?.) ? ???????, ?? ?????? ????? ??????????
?? ?????????? ????????? ????????? (????. 1,
2). ??? ????????? ?????????? ???????? ?? ???????????? ? ?????? ??????????? ? ??????????? ?????????????? ???????????? ??????
???????? ?????????? (????????????, 1964;
?????????, 1986) ? ?????????????? ??????
??? ???????, ??????? ???????????? ???????? ???? ??????????? ? ???????? ? ?????????
???????????. ???????????? ????? ????????
?????????? ???????? (????????????, 1964;
?????????, 1986) ???? ???????? ????????????? ?? ?????????? ????????? ??????, ?? ??
????????? ? ????? ??? ????????????? ????
??????????? ??????????. ????????? ????? ?
?????-?? ???? ?????????? ???? ?? ?????? ??-
?????????????? ??????? ????????? ????????????????????? ????? ?????????? ????????
(?????????, 2002; ?????????, ??????, 2002?,
?, 2009; ???, ?????????, 2008).
?????? ?????????????? ???????????
???????????????? ????????? ????????, ?????????? ??????????? ??????? ?? ?????? ?????????, ? ???? ?? ???????????, ??????????
?? ??????????????? ?????? ?? ??????????
???????? ?? ??? ?? ?????????, ???????? ?????????? ???????? ????????? ???????? ?????
??? ???????? ?? ?????? ?????????????????
????????? ? ??????? ?? ????? ? ??? ????????.
?? ?????? ????? ??????? ???? ????????????
????? ????????? ???????? ????? ? ??????????? ? ?????????? ???????? ????? (???. 2 ?).
? ??????? ????? ??????????? ????????? ?
??????????? ???????? ?????? ??????? ?????
?????????? ?????????? ? ?????????? ??????????? ????????? ? ???????????? ? ?????? ???????? ????????????? ?????????, ??
??????? ?????? ? ? ??????? ? ?????? ???????????? ??????????. ? ?????????? ????????????? ???????? ?????? ? ????????? ?????
(????????????? ? ???????) ?????????? ??????? ? ?????? ? ?????? ???????????? ??????????, ?????? ??????? ?????? ????? ????????????? ??????????? ?? ????? ?????? ??????.
?????? ???????????? ?????????? ?? ??????
? ?????? ???????? ??????? ???????????? ?????????. ??????? ????? ?? ????????????? ??????????? ?? ????????? ?? ??????????????
?????????? ????? ??????? ????????? ???????? ?? ??????? ? ??????? ?????? ? ??????? ???????????? ?????????.
?????????? ????????? ??????????????
????? ????? ?????????? ??? ???????????????? ????? ?????? ?????????? ???????? ? ????? (???. 2?), ??????????? ? ?????? ???????????? ????????????? ????????? ? ?????????
(?? ??????? ????? ?????????????) ??? ????????? (?? ????????? ?????), ? ????? ???-
# 17 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
???. 2. ????? ?????????? ????????? ?????????????? ????? (?), ???????? ????? ? ??????????? (?)
???????? ????? ?????????? ?? ???-??????? ???????-????????? ???????. ? ? ???????? ?????: ?????????
(1, 3), ? ????????? (2, 4), ????? (1,2), ???????? (3,4). ? ? ????????: 5 ? ? ??????, 6 ? ?? ???????, 7 ? ??
??????? ??????; 8 ? ??????. ?? ??? ??????? ? ???????????????? ????????? (?????? ? ?????), ?? ???
??????? ? ?????????? ????? ? ???????? ????????, % ?? ???????????? ???????. ???????? ????? ?
???????????, ?????????? ? ?????????????? ?????????, ????????????? ???????? ????????? ????????
?????????? ?????? ?????? ?? ???? ???????? ? ??????? ???????????? ??????. ????????????????
?????????: ss ? ????????????, s ? ?????????, ????????? ????????? ??. ?? ???. 1
?????????? ???????????????????? ?????????. ??????? ???????? ????? ????? ???????
??? ????? ?? ????????????? ?????????? ??
?????????? ????? ???????????????? ??????
?? ??????????????????? ???????? ?? ??????
???????? ?????.
? ??????? ???????? ???????? ??????????? ??????????? ????? ???????? ????????
?????????? ????????? ?????????, ?????????
? ??????? ? ??????? ???? ? ??????? ????????????. ??????????, ???????????? ? ??????? ???????????? ????????? ???????????????
????????????? ???????????? ????????? ?
????????? ?????? ?????? ?????. ? ??? ?????
???????? ???????? ????? ???????? ? ?????????? ? ????????? ?????. ?????????????
???????? ????????? ????????? ?????? ? ??????? ???? ???????? ?????? ?????????????.
?????? ????? ? ???????????? ? ???????
???????????? ???????? ???????? ????????????? ????????? ?????, ?????????? ???????
????????????, ?????????????? ? ??????????
????????? ????????? ??????. ?? ????????
???????????? ?????????? ???????????? ??????????? ?????? ??????? ????? ??????? ???????? ?????????? ?????????? ????????????? ?????????? ????? ? ???? ????????????
????????? ????????????????-?????????? ????????. ??? ???????? ?? ???????? ????????????? ????????? ? ??????? ???????????? ????????? ? ???????????? ????? ???????????.
????????? ????? ????????? ????????????
????????, ??, ???????????? ?????????????
????????????? ??????, ???????????????? ?????????????? ????????. ?????????? ????????
???????, ????????? ? ?????????? ???????-
# 18 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
?? ?????? ????????????? ?????. ??????????
??????????? ?????????? ?????, ??????? ?????????????? ?????????? ????????? ??????
?? ??????? ????? ????? ? ????????? ??????
?? ??????.
? ?????? ???????????? ???????? ??????????????????? ????????? ?????????????? ????????? ????????????? ?????????,
?????????? ? ???????? ?????????? ??????
????????? ????????? ?????? ???????? ????? ?????, ? ????? ? ???? ???????. ????????
????????? ????????? ????????? ????? ???????????? ?????????? ????????? ??????????????????? ????? ????????? ??????, ??????? ???????? ????? ?? ???? ??????????????
???????. ????? ?? ?????????? ?????????
???????????????????? ?????? ????? ?????????? ???????? ?????????? ?????? ?????????? ????? «??????» ????????? ??????. ?????? ????? ????? ??????????? ?? ?????? ?
?????????? ???????????, ?? ? ? ??????????.
?????????? ????????? ??????? ?????????
?????? ????? «??????????» ???? ?? ??????
? ??????????? ?? ????????? ?????????. ????????????? ????? ????? ??????? ????????
?????? ?? «??????» ????????? ?????? (?????????, 2007).
??????????
?????? ????????? ???? ????? ?????????? ?? ???-??????? ???????-?????????
??????? ???????? 4 ??????? ? 11 ???????????????? ?????????. ? ??????????? (???????????? ??? ????) ? ??????????????? (???????????? ??? ????) ???????? ?????????????
??? ????????? ???????? ?????? ??? ??????
??????? ?????????? ?????????? ??????????
???????? ?????? ???????? ??????? ????????????? ????????? ?? ???????????? ?????? ???
??????? ?????. ? ?????????? ?? ????? ??????? ????????????? ????????? ???????? ????????? ???????. ??????? ?????????????????
??????? ??????? ????? ?????????? ? ???????????? ???????? (????????? ??? ????).
?????????? ????????? ???????? ? ???? ????
????: 1) ?????????????? ?????, ??????????? ? ?????? ???????????? ?????????????
????????? ? ????????????? ???????????????????? ?????????, ? 2) ???????? ????? ?
???????????.
????? ???????? ????????? ?? ????-????????? ?????????? ??????? ??????? ???????? ?????? ???????? ?????
????????? ???? ? ????????? ?? ??? ??? ???????????? ?.?.-?.?. ?.?. ????????.
?????? ????????? ? ?????? ????????? ??????????????? ???????????? ?? ??? (??????? VI. 44.2.6 ? VII.63.1.4).
?????? ??????????
???????????? ?.?. (1964) ???????? ????????????? ???????. ?: ??????? ???????????.
?.-?., ?????, 3: 300-447.
?????????? ?.?. (1981) ?????????? ???? ? ??????????? ???????? ????????? ????????. ???.
????. 66(1): 86-98.
??? ?.?. (1974) ?????????? ?????? ???????. ???????????: ?????, 212 ?.
??? ?.?., ???????? ?.?. (2007) ?????????? ????????? ? ?????????????? ???????? ???????????? ? ???????????-???????????? ??????????? ??????? ????? ???????? ??????. ???????
?????. ???. ??-??. ????????. 1: 119-137.
??? ?.?., ????????? ?.?. (2008) ?????? ?????????? ????????? ????????? ????????, ?????????????? ??-??? ?????? ?????????? ?????. ?????? ???-??. 5: 21-23.
# 19 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
????????? ?.?. (1998) ?????????? ???????? ???????????? ????????? ????? ????? ?????????? (Pinus sibirica Du Tour): ???????. ????. ? ????. ????. ????. 06.03.01 ? ?????? ????????,
???????? ? ?????????????. ??????????. 23 ?.
????????? ?.?. (2007) ? ????????????????? ????? Pinus sibirica (Pinaceae). ???. ????.
92(6): 877-884.
????????? ?.?, ?????????? ?.?. (1997) ??????????? ???????? ????????? ????? Pinus
sibirica Du Tour. ???. ???. ????. 54(3): 261-265.
???????? ?.?., ????????? ?. ?., ?????????? ?.?. (1989) ???? ? ??? ????? ??????????. ???????????: ?????, 167 ?.
???????? ?.?., ?????????? ?.?., ?????? ?.?. (1990) ??????????????? ???????? ????????
??????? ??????????? ????? ??????????. ?: ???????? ???????????????? ? ??????????????????. ??????????, ?. 33?34.
????????? ??????? ??????? (1988) ?????? ?.?. (???.). ?????: ???-?? ?????. ??-??,
246 ?.
??????????? ?.?., ???????????? ?.?. (1978) ????????? ????? ???????? ??????. ???????????: ?????, 190 ?.
?????????? ?.?., ????????? ?.?. (1996) ????????? ? ???????? ????????? ????????? ????? ????? ??????????. ?????????. 27(1): 53-61.
?????????? ?.?., ????????? ?.?. (2000) ????????? ????? ????? ?????????? (Pinus sibirica
Du Tour) ?? ???????????? ????? ??????????. Krylovia: ???. ???. ????. 2(1): 110-122.
?????????? ?. ?., ??????? ?. ?. (2002) ?????????? ????????? ????? ???????? ? ????? ?????????? ?? ??????? ??????? ??????????????? ? ????? ????????? ?????. ????????. 4:
243?249.
???????? ? ????? ?????????? ????????? ?????? ????????. ??????? ? ?????????? (1989).
????????? ?.?., ??????????? ?.?., ?????????? ?.?. ? ??. ?. 1: 102 ?.
???????? ????????? ? ??????????????? ????????????? (1986). ??????????? ?.?., ???.
?.: ????, 74 ?.
????????? ?.?. (1974) ??????????? ????????? ????? ??????????. ?: ???????????? ????????? ???????? ??????. ??????????: ???? ?? ????, ?. 77-103.
????????? ?.?. (1985?) ???????????????? ???????? ? ???????????? ????? ??????????. ?:
???????? ???? ??????. ???????????: ?????, ?. 8-40.
????????? ?.?. (1985?) ??????????????????? ???????????. ?: ???????? ???? ??????.
???????????: ?????, ?. 132-150.
??????? ?.?. (1969) ???????? ?????? ????????. ?., ?????, 312 ?.
???????? ?.?. (1976) ???????????? ? ???????? ?????????? ???????????-????????????
?? ???????? ?????. ?: ????????????????? ? ?????????? ???????? ????? ?? ????? ? ????????.
??????????: ??? ?? ????, ?. 104-113.
???????? ?.?. (1981) ???????-????????????? ?????????????? ????? ?????????? ??? ???????? ???????????????? ???????? ?????. ?: ??????????????? ???????? ????? ?? ????? ? ? ???????? ??????. ??????????: ??? ?? ????, ?. 3-12.
???????????? ?.?. (1956) ??????????? ????? ? ???????? ??????? ? ????? ? ???????????
??????. ???. ????. 41(8): 1194-1206.
# 20 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
???????? ?.?. (1984) ????????????? ??????????????????? ????? ? ????? ????? ?????????? ? ??????- ? ??????????? ??????? ???????? ??????. ?: ???????? ????????? ???????????
??????? ??????. ??????????: ???? ?? ????, ?. 39-51.
????????? ?.?. (1964) ???????? ???????? ??????? ????? ??????????, ?? ???????? ??? ????????? ??????? ?????. ?: ??. ?? ????. ???-?? ??????, ???????????, ????. ??-? ?? ????, 8:
145-155.
????????? ?.?. (1972) ????????????? ?????? ??????????? ????? ??????????. ???????????:
?????, 274 ?.
????????? ?.?. (2002) ????????? ????? ?????????? Pinus sibirica ? ???????? ??????? ?????. ???. ????. 87(3): 62-71.
????????? ?.?., ??? ?.?., ?????? ?.?. (2008) ?????? ?????? ????????????????-??????????
???????? ???????????-???????? ????? ?? ????????????????????? ?????? (?? ??????? ?????????????? ??????????). ??????? ?????. ???. ??-??. 309: 180-184.
????????? ?.?., ?????? ?.?. (2002?) ????????????? ? ????????????? ??????? ????? ?????
?????????? ? ???????? ???????? ??????? ???. ?: ???????-???????????????? ???????????? ?
???????? ???. ?????: ??????? ??? ?? ???, ?. 328-350.
????????? ?.?., ?????? ?.?. (2002?) ?????????????? ???????? ????????? ????? ??????????. ?: Enviromis 2002. ?????: ??? ?? ???, ?. 239-243.
????????? ?.?., ?????? ?.?. (2009) ??????????? ?????? ? ???????? ????????????? ?????
? ???????? ???????? ? ?????? ?????????. ??????? ?????. ???. ??-??. ????????. 2(6): 111-125.
??????? ?.?. (1989) ?????????? ????????. ?.: ????. ??., 464 ?.
??????? ?.?. (2007) ???????? ?????????? ????????? ?????: ???????? ? ???????????????
???? ?????. ???????????: ?? ???, 181 ?.
??????? ?.?. (1963) ???????? ? ????????????? ????? ??????????. ?: ???????????? ?????
?????????? ? ????????? ??????. ?.: ?? ????, ?. 5-21.
???????? ?????. ??????????? ????????????? ? ??????????????? ???????? (1989) ???????
?.?., ???????? ?.?., ????????? ?.?. ? ??. ?????: ??? ?? ?? ????, 158 ?.
???????? ?.?. (2002) ????????? ? ???????? ??????????? ??????. ???????????: ?? ???,
253 ?.
????????? ?.?. (1965) ?????????? ??????? ????????? ???????? ???????? ????? ?????????? ??? ???????????? ??????????? ?? ????????. ?: ???? ????????????????? ??????????????
????? ?????? ? ???????? ???????. ?.: ?????, ?. 131-143.
?????????? ?.?. (1992) ????????? ????????????? ??????? ???????????? ???? ????? ??????????. ?: ???????? ?????. ???????? ???????? ?????. ?????: ???? ?? ???, ?.5, ?. 131-137.
?????????? ?.?. (1990) ???????-?????????????? ?????????????? ? ???????? ????????
????? ????? ? ???????-????????? ??????? (???????-??????????????? ?????? ???????????
?????????). ??????????: ??? ?? ????, 286 ?.
????????? ?.?., ??????????? ?.?., ????????? ?.?. (1967) ?????????? ???????? ?????. ?.:
?????, 123 ?.
??????? ?.?., ????????? ?.?., ?????????? ?.?., ?????? ?.?., ??????? ?.?. (2009) ??????????? ???????????????? ????????? ????????????? ??????? Pinus sibirica (Pinaceae) ? ?????
???????????? ?????. ????. ???????. 45(1): 3-12.
# 21 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ????????? Pinus sibirica ?? ???-??????? ???????-????????? ???????
?????? ?.?. (1975) ?????????? ?????? ????????????????? ??? ??????? ??????? ? ?????????????? ???????? ?????????. ????. ?????. 2: 7-34.
?????? ?.?. (1970) ? ????????????? ???????? ?????????? ????? ????? ? ???????????.
?: ??. ???????. ????????. ??-??, ?. 21, ?. 339-343.
???????? ?.?. (1988) ????????? ???????? ?????? ????????. ?.: ?????, 560?.
??????? ?.?. (1964) ???????? ????? ?????????? ? ?????? ????? ??????. ?.: ?????? ??????????????, 100 ?.
??????? ?.?. (1975) ?? ???????????? ? ??????????????? ???????? ??????????? ??????????? ?????? ? ???????? ????? ?????????? ? ?????? ????? ?????????? ?????. ?: ??. ???. ???????. «??????», ?.10, ?. 61-67.
????????? ?.?. (1963) ???????????? ????? ?????????? ? ??????????? ????? ??????????
????. ?: ???????????? ????? ?????????? ? ????????? ??????. ?.: ?? ????, ?. 120-126.
Mirov N.T. (1967) The genus Pinus. NY.: Ronald, 602 p.
Smirnova O.V., Chistyakova A.A., Zaugolnova L.B., Evstigneev O.I., Popadiouk R.V., Romanovsky
A.M. (1999) Ontogeny of a tree. ???. ????. 84(12): 8-20.
Ontogeny of Siberian Stone Pine
(Pinus Sibirica du Tour)
in Southeastern West Siberian Plain
Svetlana A. Nikolaeva,
Svetlana N. Velisevich, and Dmitry A. Savchuk
Institute for Monitoring of Climatic and Ecological Systems SB RAS
10/3 Academichesky prospekt, Tomsk, 634055 Russia
The complete ontogeny of Siberian stone pine (Pinus sibirica Du Tour) trees of the first after-fire
generation is described for the regeneration and aged series of green mossy Siberian stone pine
forests which are typical in West Siberian taiga. Classic scheme of ontogenetic periods is used. The
ontogenetic states are described using quantitative and qualitative vegetative (including height,
radial, and basal area increments) and generative (initiation and preservation of microstrobili and
macrostrobili) parameters. The age changes are generalized into two schemes: i) morphological
structure of the crown taking into account the patterns of origin of their elements and prevailing form
trends and ii) tree growth and cone development rate.
Keywords: Siberian stone pine, Pinus sibirica, ontogeny, regeneration and aged series, green mossy
Siberian stone pine forests, middle subzone of taiga, southern subzone of taiga.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2011 4) 23-35
~~~
??? 631.524.82: 630*181.65: 630*181.522:582.475.2
?????????? ???????????? ???????????
???????????? ??????? ????? ??????????
?.?. ?????????*, ?.?. ??????????
???????? ??????????? ?????????????
? ????????????? ?????? ?? ???
?????? 634055, ?????, ??. ?????????????, 10/3 1
Received 4.03.2011, received in revised form 11.03.2011, accepted 18.03.2011
?? ??????? ??????? ? ??????? ??????? ? ??????? (140 ???), ????????????????
(235 ???) ? ?????? (346 ???) ???????????? ???????? ????? ?????????? (????-?????????
??????????, ???-????????? ????? ???????? ??????) ??????????? ??????? ???????? ??????
?? ??????????? ???????????? ? ???????????? ?????????. ???????????, ??? ????????
?????????, ??????????????? ???????????? ???????? ??????? ???????, ??????????? ?
??????? ??????????????? ??????? ?????????? ???????. ???????? ????????????? ????????
??????? ? ????????????? ????????? ? ???? ???????????? ?????????? ?? ????????, ????????
???????????? ???????? ? ???????????????? ????????. ??????????? ????????? ??????????
? ? ????????? ??????? ???????. ?? ???? ??????????? ??????????? ?????? ? ????????
????? ????? ????????????? ? ????????????? ?????????? ???????? ???????: ??????????
???????????? ??????????? ? ??????? ???????? ???????????? ??????????? ?? ???????????
????????? ?????, ???????? ?????? ????????????? ???????? ? ??????? ???????? ????????????
?????? ?? ???????????? ?????, ? ?????????? ???? ????????? ? ???????? ???????? ????????
???? ????????? ??? ?????. ????????? ?????????????, ??? ??????? ??????????? ??????????
???????? ???????????: (1) ?????????? ??????? ????? ???????????? ? ????????????
????????????? ? ???????? ????? ???????? ??????? ? (2) ????????????????? ???????????
????????????? ?????????????? ??????????.
???????? ?????: ?????, ???????????, ????, ?????????, ???? ?????????, Pinus sibirica Du Tour.
???? ????????? ???????? ???????? «???????» ?????????, ?? ?????? ???????? ?????????????? ?????????????? ????? ?? ??????????? ????????? ???????? ? ????????????
? ????????????. ? ??????? ?? ??????????????? ????????, ? ??????? ??? ?????????? ?
*
1
???????????? ???????????? ????, ? ??????????? ??????????????? ????????, ? ??????? ????????? ??? ??????? ????????? ????,
?????????????? ???? ???????? ???????????
?????????? ????????????? ??????? ????????????? ? ????????????? ???????????.
Corresponding author E-mail address: velisevich@imces.ru
© Siberian Federal University. All rights reserved
# 23 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ??????????? ?????????? ???????????? ??????????? ???????????? ??????? ????? ??????????
????????? ??????, ??? ? ????????? ?????? ????????, ??????????????? ????????????? ??????: ?????????? ???? ?????????
????????? ??????? ?????????????, ?????????
????????? ?, ???????, ????????? (??????,
1963; Kozlowski, 1971). ???? ??????? ???????????????? ??????? ? ?????? ? ????????????????? ? ????? ?????????? ????????????, ???
? ? ??????? ??????????????? ??????? ???????????? ???????? ??????????? ??? ?? ??????????????, ??? ? ????????, ?.?. ??????????
?? ????????. ???????? ?????, ??? ???? ????
????????? ? ????? ?????????? ?? ?????????
?? ???????: ?????????? ? ??????? ??????????? ?????????? ?? ???? ?????? ????????
????? (????????, 1983; ?????????, ???????,
2006).
? ????? ???? ????? ?????????? ????????? ????? ????? ? ????????????? ????????
????? ?? ?????????? ???????? ????????-
?????? ?????????? ????? ?????????? ?? ??????? ?????????????, ?????? ?? ?????? ???????? ? ???????. ?????? ? ??????????????
????????????? ?????? ?????? ? ???????? ??
???????????? ??????? ????? ???????? ? ???????????? ?????????. ?? ?????????????? ??????? ? ?????????? ?????? ????? ??????????
?? ???????????????. ????????? ?????? ???????? ?????????? ???? ?????? ? ?????? ?????
??????? ???????? ??????? ???????? ?????? ??
??????????? ???????????? ? ????????????
????????? ?? ?????? ??????. ? ??????? ??
?????????????? ????????????, ??????????? ?? ??????? ???????????? ??????? ???????? ?? ???????????????? ?????????? ? ????????????? ???????????, ??? ??????? ?????
??????????? ?? ??????? ???????? ????? ??-
???????? ??????????????? ????? ???????? ???????? ????????? ???????: ?????????????
??????????? ???????? ???????? ?? ???? ?????????? ??????????? ????????????; ?????????? ???? ??????? ????????? ??????????
????? ?????????? ???????? ????? (??? ??????
???????????? ??????????? ???????????? ?
???????????? ?????????); ?????????? ?????????? ????? ???????? ? ??????????, ? ?????
? ??????????? ????????? ??????????????
????????. ?? ???? ?? ?????????? ????? ?????????? ???????????? ? ???????????? ???????? ????? ??????? ????? ?????.
??????????? ??????????? ???????????
? ?? ?????? ????????? ????????? ????? ?
?????? ????????? (??????????, ?????????,
1996, 2000), ?????? (??????????, 1990) ? ??????? (??????????, 1991; 2006). ? ?????????,
???????? ?????????? ??????? ????????? ?????? ? ???????? ????? ?????????? ??????????
???????? (?????????, ??????????, 2004) ????????, ??? ??????? ??????? ????? ??????? ?
????????? ? ??????
?????? ??????????? ???? ?? ??????? ???????????.
????? ???????????? ?????????? ? ???????????? ????? ???????-????????? ??????? ? ?????????? ???? ? ?????? (????????????? ????? ??????? ???????, ???????
«??????????? ???», ??????????? ???. ??????, 58є13'?.?., 84є32' ?.?., ?????? ??? ???????
???? 104-116 ?). ?? ??????????? ??????????? ?????????? ??????????? ????????? ??????????? ????????????? ? ?????????????
?????? ?? ??? (????? ??????? ?????????
???? ?? ???) ?? ???????? ?????, ????????? ?????????, ????????? ????????????? ?
????????? ???????????? ??????? ??? ??????
?????? ??????? ??? ??????? ???????. ???
??? ???????????? ????? ?????????????
????????????????-?????????? ??? ????????
???????????? ?? ?????? ????????????????
? ?????? ?????? ??????????? ?????????????????????????.
?? ?????? ???????????? ??????? ??????? ? 1 ???? 140-?????? ????????? ? ??-
# 24 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ??????????? ?????????? ???????????? ??????????? ???????????? ??????? ????? ??????????
????????? ??????????? ????? ???????
????????? ????? (1-? ????: 9??1?+?, 2-? ????:
8?2?+?). ???????? ????? ????? ???????????,
? ??? ????????? «????». ?????? ?????????
?????, ??????? ? ???? ???????? ????????????, ?????? ???????? ? ??????, ?? ?????????
?? ?????? ?????. ??????? ????????? ????????
?????, ??????????????? ?? ???????? ????????? ??????? ?????? ?? ??????, ? ???????
140 ??? (lim 117-160), ?????? ?????? 16 ?; ??????? ?????? 17 ??.
??????? ??????? ? 2 ? ? 3 ? ??????????? ?????? ???????? ????? ?????? ??
?????????? ???????? ????? 100 ? 200 ??? ??????????????. ?? ??????? ??????? ? 2 ????????????? ????????? ?????????? ????????
?????? ?????????, ? ????? ??????? ?? ????????? ? ?????? ??? ? ????? ??? «??????» ?
???????, ??????????????? ????????, ?????
(1-? ????: 7?2?1?+?+??). ??????? ???????
????????? ???????? 235 ??? (lim 229-261), ?????? ?????? 22,8 ?, ??????? ?????? 34,1 ??.
?? ??????? ??????? ? 3 ?????? ????????? ??? ? ????? ??? ???????, ? ?????? ????????????? ?? ?????? ????? (1-? ????: 10?+?,
2-? ????: 5?4?1?). ?????? ? ???????? ? ????
????????? ?? ????????? ?????? ???????.
??????? ??????? ????????? ???????? ?????
346 ??? (lim 325-360); ?????? 24,5 ?; ???????
?????? 43,6 ??.
????????
?????
?????????????
????????????????-?????????? ???????? ??????????? (??????????, 1990), ??????? ??????? ?1 ????????? ? ??????? ???????????? ??????????? ?????-???????-????????
?????????? ????? ?????????????? ???????
?????????? ????? ?????????. ??????? ??????? ?2 ????????????? ??????? ??????????
????????? ??????? ? ??????????? ????????.
??????? ??????? ?3 ????????? ? ?????????????? ???? ???????? ??????????, ????????
??????????? ? ?????? ????????? ????????.
???????? ?????????????, ???????? ? ???????????????? ?????????? ????????? ????????
(??????, 1975), ????????? ??????? ???? ??????? ???????? ????????????? ??????? (g1),
???????????????? (g2) ? ?????? (g3) ???????????? ????????.
?? ????? ???????????? ? ??? ?????. ??????? ???? ?????????? ??????? ???????????? ?????????? 100 ???????? ?????? ??????? ???????. ????? ?? ?? ????? ????????
?? 10 ??????? ?? ????? ? ???????? (????? 30
????????). ? ??????? ?????? ??????? ?? 15
??????? ? ??????? ?????? ?? ??????????????? ???????????? ?????? ?????. ????????
????????? ?????? ?????? ???? ????????????
????????? ?????????????? ???????? ??? ??????? ?? ?????????? ????????? 15 ???. ???
??????????? ??????? ???????? ??????? ??
?????????? ??????? ???????????? ????????????????? (??????? ?? ????????? 12 ???)
???????? ??????? ????????. ????? ???????
??????????? ?????? ????????? ???????? ?
?????? ????????? ?????? ? ????????? ?? 150
??????? ?? ?????? ??????? ???????. ??? ?
?????????? ???????????? ????????? ????????????? ?? ?????? ?? ??????? ??????? ??
???? ??????? (Vorobjev et al., 1994; ????????,
??????????, 1989).
????????, ??? ?????????? ?????? ?????? ???????? ????????? ?????? ?????????????? ???????????? ???????, ????? ?????????? ??????????? ???????????? ??????
?????????????? ?? ???????? ? ????? ??????????. ??????? ??????, ??????? ????????????? (?????), ??????????? ?? ???????
?????? ? ??????? ????? ?????. ??? ???????,
???????? ????? ??????? ?? ???? ???????????? ? ????????? ? ???????. ???????? ?????
??????????? ?? ???????? ?????, ??? ??????????????? ??????????? ????? ??????????,
? ????? ?????????? ??????????, ? ???????
??????? ? ????????????? ??????????? ?????-
# 25 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ??????????? ?????????? ???????????? ??????????? ???????????? ??????? ????? ??????????
???????? ???????????, ??????????? (??????
??????????? ?????????????, ????????? ?????
? ????????? ??????, ????????????? ?? ???) ?
?????. ????????? ????? ? ???????? ?????
????????? ?????? ?????????????? ???????????? ????? ??????? ?????? ? ?? ??????????
? ??????? ?????????? ?????????????? ??????
(??????????, 1994). ?????? ????? ????????
???????? ? 10 ??? ?????? ?????????, ??? ???????? ?????, ? ????? ???? ??? ???? ???????? ??????? ? ??????????? ? ???????????.
????? ???????????? ?? ???????? ?????? ??
??? ???? ????????, ??????? ?????????? ??????? ????????????? ????????? ???????????
?? ???? ?????????? ????? ???????????? ??????? ??????. ????????? ?????????? ?????
????????????? ? ?????????? ??????????? ??
???? ?????????? ???? ??????? ????? ???????? ? ?????? ????????. ??? ?????????? ? ???
?????? ????? ?????, ??????? ?????? ??????
(Lanner, 1978; Greenwood, 1980).
??? ??????? ??????? ??????? ????????????? ???? ?????????? ??????? ??????????
? ?????? ??????? ????????? ?????? ???????
?? ?????????? ????? ??????? ????????? ??????, ????????? ?????? ???? ?????? ?????? ?
?????????? ? ??????????? ????? ? ???????
??????? ? ?????????? ??????????? (????????
? ??., 1989).
??????????
??????? ??????. ?????? ???????????? ?
????? ??????? ??????? ??????????, ??? ??????? ??????? ?????????? ?? ????????????????
? ?????? ???????????? ??????????? ?????????? ? ???????? ???????? ?????????? ???????? ??????? ?? ??? ??????? (???. 1). ???
??????????? ?? ?? ???????? ?????, ????????? ????????, ??? ?? ????????? ????? ??????
? ?????? ???? ?????? ????????? ??? ?????????? ???????????? ? ????? ?????????, ??? ?
????? ?????????? (Lanner, 1978). ? ??????-
?????????? ???????? ????? ?????? ??????????? ????????? ?????????? ?? ?????? ?????
?????????, ?? ? ????? ??????????. ? ?????? ???????? ??????? ??????????? ??????????????? ?? ???? ????? ?????????, ?????
?????????? ??? ???? ???????? ???????????
?? ??? ?? ??????, ??? ? ? ????????????????
????????.
?????????? ???? ??????? ???????, ? ??????? ?? ?????????, ????? ???? ???????? ?????????? ?????????: ?????? ??????? ??????
??????? ??????? 235-??????? ?????? ?? 14 %
??????, ??? ? 140-???????, ? ?? 17 % ??????,
??? ? 346-??????? (???. 1). ?? ???????? ?????????? ?????? ??????????? ???????????
?????????, ???? ????? ???????? ? ???????
????????? ??????????? ??????????? ???????? ?? ???? ???? ?????????.
? ????????? ?????????? ? ??????????? ????????? ????????? ? ??????? ???????
(???. 2). ???? ?? ???????? ?????? ????? ???????? ??????????? ??????????? ? ?????????, ??
?? ?????? ?????? ? ???????????????? ???????? ????? ????????? ????????????? ?? 44 %
?? ????????? ? ???????? ? ? ???????? ????????? ?? 21 %. ?????????? ????? ??????????
??????????? ? ? ????? ????????? ??????? ??????. ???, ? ??????? ???????? ??? ?????????
????? ?????????? ????????? ???? ???????????? (1,55 ± 0,22) ? ???????????? (1,51 ± 0,14)
????? ??????. ? 235-????? ????? ????????
??????????? ??????? ?????? ?????????????,
?? ???? ???????????? ??????????? ?? 34 %
(1,86 ± 0,12) ?? ???? ?????????? ??????????
???????????? (3,71 ± 0,30). ? ?????? ????????
????? ????? ????????? ??????????? ??? ??????????? ?????????? ????? ????????????
(2,90 ± 0,30) ? ???????????? (1,52 ± 0,10).
??? ?????????????? ?????????? ???????????? ?????????????? ????????? ??????
?? ??????????? ??????????????? ???????????? ???? ???????? ????? ????????? ? ???-
# 26 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ??????????? ?????????? ???????????? ??????????? ???????????? ??????? ????? ??????????
????? ??????, ? ?
??? ?? ?? ??? ?? ?
?????? ?, ? ?.
12
8
??? ??
? ? ??? ????? , ? ?
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???. 1. ????????? ??????????? ????? ? ???????????? ??????? ??????? ? 140-, 235- ? 346-?????? ????????
????? ?????????? ? ????-????????? ??????????. ?? ??? ??????? ? ??????? ????????, ???
50
40
30
20
10
0
140
? ???????
? ?????
235
346
???. 2. ??????????? ????????? ????????? ? ??????? ??????? ? 140-, 235- ? 346-?????? ???????? ?????
?????????? ? ????-????????? ??????????. ?? ??? ??????? ? ??????? ????????, ???; ?? ??? ??????? ?
????? ?????????, ??.
?????????, ???????????? ? ?????. ?????????? ?????????? ?????????? ?? ? ??????
?????? ?? ?????????, ????????? ???? ???????
????? ?????????? ? ????????? (??????????,
?????????, 1992). ?? ????? ??????, ?????????? ???????????? ? ???????????? ????????? ?? ?????????? ?????????????? ???????
(????. 1). ?? ????????? ?????, ????????,
???????????? ?????????? ???????????????
??????? ? ?????? ???????, ????????????? ?
????????????????.
?????? ?????????????? ????????????
????? ???????????? ? ???????????? ???????????? ??????? ??????? ??????????, ???
? ???????????????? ???????? ????? ???????????? ????? ???????????? ??????? ? ???????? ?????? ?????? (r = 0,52, n = 150, ? < 0,05).
? ??????? ???????? ?????????? ???? ??????-
# 27 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ??????????? ?????????? ???????????? ??????????? ???????????? ??????? ????? ??????????
??????? 1. ?????????? ???????????? ?????????????? ????????? ?????? ? 140-, 235- ? 346-??????
???????? ????? ?????????? ? ????-????????? ??????????
???????
???????, ???
235
27,5 ± 1,05?
346
15,7 ± 0,73?
????? ????????????, ??.
140
41,5* ± 1,65**?
????? ????????????, ??.
1,58 ± 0,16?
0,40 ± 0,07?
0,20 ± 0,04?
????? ?????, ??.
1,35 ± 0,12
2,15 ± 0,12
1,75 ± 0,08?
?
?
* ??????? ????????, ** ??????. ??????? ??????? ?????????? ??????????? ????????, ? < 0,05.
?????? ????????? ????? ?? ?????? (r = ? 0,58,
n = 150, ? < 0,01), ? ? ?????? ???????? ?? ?? ????????? ????????????? ??????? ?? ???? ???????. ????? ????, ?? ???? ?????????? ???????
??????? ?????????? ????? ?? ?????? ?????????? ???????????? ?????? ?? ?? ?????????
? ????????? ???? (???. 3). ?????????? ??? ??
???????????? ???????????? ???????, ?????????????? ???????????? ?????? ? ????????
????? ????? ?????? ? ????? ??????? (???????? ? ??., 1989). ?? ????????, ??? ????? ???????? ????????????????????? ????????????
???????? ????? ??????? ????????, ???????
?? ????????? ?? ?????????? ???????? ???????, ???????????? ??????????? ?? ??????????? ????????? ?????. ????????? ????????
???????????? ? ??????? ???????? ????? ???????????? ?????? ?? ???????????? ?????, ?
?????????? ????????? ? ???? ???????? ???????? ???? ????????? ??? ?????.
??????? ?????????? ?????? ? ???????,
??????????????? ?????? ?????????? ????????? ?????? ?? ????????????? ????? ???
???????? (?????????, 1974; O'Reilly, Owens,
1988). ?? ????? ??????, ??????? ???????
?????????? ?? ?????????? ????????????? ???????????? (??? ?? ????????? ?? ????? ???????? ????????, ???????????? ? ?????) ??????? ?????????? ?? ???? ???????? ????????,
??? ?????? ???????, ???? ?????? ????????????
?????????? ???????? ???????? ??????????.
????? ???????? ?????????? ?????? ? ??????
???????????? ?????, ????????, ??????????? ? ?????????. ??? ??????????????? ? ???????? ??????????? ???????????? ???????
?? ???????????? ??????, ??????????????
???????????? ??????????. ?????? ???????
?????? ??????? ??????? ?? ???? ??????????
??????? ???? ????? ???????????? ??????? ?
??????????? ???????????? ?????.
??????? ??????. ??????? ?????? ??????????? ?? ?????? ??????? ? ?????? ??????
????? ? ??????????? ???????? ??????? ??????? ?? ???????. ????? ???????? ??????
?????? ???????????????, ?.?. ??? ?? ????????
????????????? ? ??????? ?????? ?????????????? ??????. ????? ????? ?????????? ?????????? ?? ??? ?????? ??????????????? ????????????? ????? ?????????????? (???????
?????), ????? ???????????? ? ????????????.
?????? ?????????????? ??????? ??????? ? ???????? ???? ????? ???????, ??? ?
????????? ????? ?????? ???????????, ??????
?? ??? ???????????, ??? ??? ?? ?????????
? ??????? ???????. ?? ????? ???????? ???
???????? ????? ???????? (1,31 ± 0,03) ?
????????????????? (1,28 ± 0,07) ?????????,
? ???? ? ?????? (1,05 ± 0,07) ?????????? ??????????? ?????????? ????? ???????. ?????
????????? ?????? ???????????? ???????????
??????????? ????? ??????????????, ???????
???????????? ???????? ????????? ? ???????????????? ???????? (???. 4). ????? ???????????? ?????????? ?????????????? ?????
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??????????. ??????? ??????? ???????????? ?????????? (n = 150 ? ?????? ?????????? ??????), * ? ????????????, ??????????? ??? ?< 0,05
0,42
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???. 4. ????????? ??????? ??????? ? 140-, 235- ? 346-?????? ???????? ????? ?????????? ? ????-?????????
??????????. ?? ??? ??????? ? ??????? ????????, ???
??????????????: ? ??????? ? ? ?????? ???????? ?? ??????, ??? ? ????????????????.
?????????? ?????? ??????????????? ?
???, ??? ? ????????? ?????????? ?? ???????
?????? ??????? ???????, ??????? ?? ????????? ?? ????? ???????? ???????????. ???, ?
??????? ???????? ??????????? ????????????
????? ??????????? ? ???????????? ???????,
??? ?????????? ? ?????????? ???? ?????????????? ? ?????????? ???? ????????????.
??????????????? ??????? ??????????? ? ??????? ? ?????? ????????, ??? ??????????????? ? ????????????? ???????????? ?????????????? ?? ???????????.
??????????
????????????? ? ????????? ?????? ?????????????? ? ?????????? ??????????????
??????? ??????? ????????? ? ????????????
? ????? ????? ????????? ????? ? ????????
?????????????? ? ??????????. ???, ????????
???????? ??????????? ???????? ??????????
????????????? ???????? ?? ???????? ????????
?????? ????????, ???? ?????? ?? «?????????» ?? ???????????????? ?????? ????? ? ??
??????? ????????????? ???? ???????????????
????????????? ?????? ?? ?????????? ?????. ??????? ??????? ????????? ???? ?????????, ? ??????? ????? ?????? ?????????????
???????????????? ? ????? ? ????? ? ??????
???????.
????????, ?????????? ????????? ?????
?? ??????, ? ????? ???????? ??? ????????????
?????????, ??????? ??????????????? ???????
? ?????????????? ????????????? (?????????? ???? ? ????????? ???????????? ?? ?????? ??????), ????? ????????????? ????? ?
????????? ??????. ???????? ????????? ?????
?????? ??????????? ???????????? ??????
????? (????????, 1983 ? ??.), ?? ????? ??????????? ???? ????????? ??????? ? ????????? ???????: ??????? ??????? ????? ?????????? ? ?????? ????? ?????????? ?????,
???????????????? ? ?? ???????, ? ?????? ? ??
# 30 #
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?.?. ?????????, ?.?. ??????????? ?????????? ???????????? ??????????? ???????????? ??????? ????? ??????????
?????????? ????? ?????????? ?????? ?????????? ???????????. ? ???? ?????? ?????????
?????????? ????????????? ???????????? ??????? ??????? ? ??????????????? ??????????
? ??? ????????????? ?? ????????? ????? ????????? ? ????? ??????????????? ?????????????? ?????????. ???, ??????, ?? ?????????
?????? ?????????? ???????? ? ?????????????? ???????, ?, ?????? ?????, ????????????
???????????? ?????? ???????? ????????????.
?????????? ?????? ????? ???????, ??
?????? ??????, ??????? ?????? ? ??????????
???????????? ????????? ???????? ???????.
? ?????? ???????? (Greenwood, 1980), ??????????? ?? ??????? ????? ????????, ????
????????? ?????????????, ??? ???????? ?????????? ????? ? ??????? ???????? ????????
??????? ??????????????? ?????? ???????????? ???????????? ???????, ? ??????? ? ???????????? ???? ?????????? ??????????????
? ?????????? ????? ??????? ????????? ???????? ?????. ????? ???????? ????? ??????????????? ???????? ??????. ?????????? ??????? ????????????? ???????????? ? ?????????
?????? ?????????? ? ???, ??? ? ??? ??????????
?????? ?????? ???????, ???????? ???? ?????? 2-3 (Boyer, 1970). ? ????? ?????????? ??
??????? ?????? ? ????? ???????? ??????????
? ??????? 1-?? ???? ??? ???????????? ?????? ?
???????? ? ??????. ??? ?? ?????, ??????? ???????? ???????? ????? ???? ???????????? ?
??? ????? ????.
??????????? ?????, ? ??????? ??????? ?
??????? ???????? ???????????? ????, ??????????. ????????? ?????? ????????? ??? ?
???????????? ?????? ??????????? (Giertych,
1967; Owens, Molder, 1979). ? ????? ?????
????????????? ??? ???????? ??????, ? ??
?????????????? ?????????? ????? ????????? ????? ??????????. ??????? ??????????
????? ?????????? ?????? ? ????????????-
????? ??????? ???????????? ????. ??????
? ???????????, ? ??????? ????? ????????
???????????? ????????? ? ?????????? ?????
????????????????? ???????? ?? ?????????
??????? ? ????????? ???????, ?????? ???????? ?????? ?? ????????????? ????????? ? ???????????? ????? ???? ?????????? ? ???????
?????????. ???????????? ?????? ? ?????????? (Owens, Pharis, 1971; Owens, Molder, 1974)
????????, ??? ? ??? ?????????? ? ???????????? ??????????? ??????? ? ????????? ??????? ????? ??????????? ?????? ? ??????? 2-3
?????? ?????? ???? ? ?? ?????????? ??????? ???????????? ?????? ????????, ? ??????
????? ????????? 3-4 ??? ???????. ??? ??????? ????????? ????????????, ??? ? ????? ??????? ???????????? ???? ????????? ??-?????.
?????? ????? ?????????, ??? ? ? ???? ??????
????????? ??????, ? ??????? ???????? ???????? ???????? ????????? ????????????
???????, ?????? ? ????? ???????? ???? ??????, ??????????? ? ??? ????? ?? ????????????
?????? ????????.
? ????? ?????????? ???????????? ?
????????? ????? ? ????? ??????? ?????
???????-????????? ??????? ??????????
?? ?????? ???????? ???? (?????????, 1973).
?????????? ?????????????? ????????????
(??????????, ?????????, 1992) ????????,
??? ? ?????? ???????? ? ??? ????? ????? ??????????? ??? ????????? ?????????????
???? ???????. ?? ? ???????????? ????????
?? ?????? ???????? ???? ???????????? ??????????? ?????????? ???????? ??? ??????
?? ???? ???????????? ???????? ???????. ?
????? ? ???? ???? ????????? ?????????????:
??????? ??????? ?????????? ????????????? ????? ????????????? ?????????? ?????
??????? ????? ?? ?????? ?????????? ? ???
???????? ??????????. ?????????? ? ????????
????????? ????? ?????? ????????, ????????, ?????????? ? ?????????? ??????????????
# 31 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ??????????? ?????????? ???????????? ??????????? ???????????? ??????? ????? ??????????
????????? ?????? ???????????? ? ????? ??
???????? ??? ??????? ???????. ?????????
???????????? ??????? ???????????? ??????
?????, ????? ?????????? ??????????? ??????? ??? ??????? ?????????????. ??????????
????? ? ??????? ???????? ????? ?????????
???, ??? ????????? ???????????? ??????? ????????????? ??????? ??????, ????? ??????,
????????????? ??? ????????? ?????, ???
????????.
???????? ???? ????????, ????????
??????? ???????? ?? ????? ????????????
? ???????????? ????????? ?????? ? ??????? ???????? ????? ??????? ?????????
??????????????? ?????? ?? ?????????. ?
?????????? ????? ? ??????? ??????? ?????? ?????????? «????????????» ?????????
??????????? ? «????????????» ?? «????????????», ??? ? ?????????????? ????? ?????????????? ?????????? ????? ?? ??????. ?
???????????????? ???????? ???? ????????????? ?????? ?????, ????? ?????????? ????????????? ??????? ??????? ??? ???????
?????????????. ???????? ????????? ?????
?? ??????? ??????????????? ? ?????????????????? ?????????? ?????? ???????????? ?
????? ??????? ? ????????? ????????.
????? ????????????, ??? ?????????
? ?????????? ????????????????? ???????
????????????? ???????????? ??????????? ??
?????????? ????? ?????????? ????? ??????.
??????? ? ?????? ???????? ?????? ????????
????? ?????? ??????? ? ????????? ???????????? ???????, ??? ??????????? ?? ?????????? ????? ??? ??????? ? ?????????? ????????? ???????????? ???????. ????????
?????, ??? ????? ?????????? ?????? ???????????? ???????? ?? ?????? ????????????,
?? ? ???????????? ?????????, ?.?. ? ??????
???????? ???????? ????? ?????? ?????????
???????? ????? ? ?????????? ???????, ?
??????? ???????? ?????????? ????????? ??-
???, ??? ?????????? ?? ?????????? ?? ???????????.
?? ???????? ? ???????? ???????? ????? ????????????, ??? ? ??????? ????????
????? ????????? ???????? ??????? ???????
???????? ?????????? ??? ????????? ??????????????. ??? ????????, ??? ??????? ?????
???????? ?????????? ???????????? ?????????????, ? ??????? ? ???????? ????????????.
??? ????????????? ?????? ????????? ???????? ? ??????? ?????????? ? ??????? ???
??????? ???????? ? ?????? ???????? ? ??????? ????? ?????????? ???????????????? ???
???????????.
??? ????? ???????? ?????? ??????????? ???????? ?????? ???????????? ?? ??????
???????????? ???????, ?? ? ???? ???????, ???????????? ????? ??????, ???????? ?????????? ????? ????? ?????? ? ???????? ????????
(???????, 1969). ????????, ??? ???????? ????
? ????????? ???????? ?????? ??????????????
?????????? ????? ??????? ? ????????, ?.?.
???????? ?????????? ????? ? ???????????
???????? ????? ???? ?????????? ????????
???????? ???????. ???????? ???? ????????? ???????? ?? ????? ???????????? ??????
???????????? ?????????? ????? ?????? ??-??
??????????? ??????????? ? ??? ???????????, ?????????? ?????????? ?????? ???????? ? ????????? ??????? ??????, ???????????, ??????? ?????? ? ?.?., ??? ???????
????? ????????????????? ???????? (Ryan,
Waring, 1992; ??????, 1994). ????? ????, ????????????? ???????????????? ???????? ??????? ???????? ? ?????????? ???????????
?????????? ? ????????? ?????? ? ????????
?????????? ?????????? ???????? ???????,
??? ????? ???????????? ???????? ????? ???????? (???????, 1973; ???????, 1982; ????????, 1988 ? ??.). ? ??????????????? ????????
???????? ????????? ????? ??????, ? ???????
?????? ?????????????? ??????. ? ????????,
# 32 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ??????????? ?????????? ???????????? ??????????? ???????????? ??????? ????? ??????????
???????? ????? ???, ?? ???? ?????????? ?????? ?????? ????? ????? ???????? ? ???????
????? ??????????, ?? ???? ??????? ????????????? ?? ??????? ? ?????????? ??????????,
???????? ? ???????? (???????, 1969).
??????? ?????????????, ???????????
?????????? ???????? ??????????? ???????????? ??????? ???????????????? ?????
????????? ?????????. ? ??????? ????????
?????????? ??????????????????? ? ???????????????????? ???????????? ? ???????? ?????
??????????
?????? ????????? ???????????? ????????
????????? ??? ?????????????. ? ???????????????? ???????? ??????????? ????????? ????
???? ???????? ???????????? ????????????
????????? ????? ? ????????????. ? ?????? ???????? ????????? ????????????? ??????????
? ????? ??????? ? ????????????????? ?????????? ???????????? ???????? ?????? ???????????? ?????????????? ???????? ???????? ?
???????? ?? ?????? ?????????????, ?? ? ????-
???????????? ??????? ???????? ???????? ????? ?????????? ?? ?????????? ??
???????????? ??????? ????????, ??? ????????, ??????????????? ???????????? ????????, ??????????? ? ??????? ??????????????? ??????? ?????????? ???????. ????????
????????????? ???????? ? ?????????????
????????? ? ???? ???????????? ?????????
???????????? ???????? ? ???????????????? ????????, ?????? ???????????? ??
???????????? ??????? ??????????? ? ???????? ?????? ? ???????? ?????? ?????
???????????? ? ???????????? ????????????. ??????????????, ??? ??????? ??????????? ?????????? ???????? ???????????
??????????? ???????? ???? ????????: (1)
?????????? ??????? ????? ????????????
? ???????????? ????????????? ? ????????
????? ???????? ??????? ? (2) ???????????????? ?????????? ?????-???????? ???????-
????????? ???????? ???????.
??????? ??????????.
?????? ??????????
????????? ?.?., ?????????? ?.?. (2004) ??????????? ???????? ? ???????????? ????????? ???????? ????? ?????????? ?????????? ????????. ??????????? 6: 66-69.
????????? ?.?., ??????? ?.?. (2006) ???? ? ?????????? ? ???????????? ???????? ???????????? ????????? ? ???????????????? ??????? ????? ??????????. ?????? ????????? 3: 39-40.
???????? ?.?. (1983) ????????????? ?????? ???????????? ????????????? ???????? ?????.
???????????: ?????, 253 ?.
???????? ?.?., ????????? ?.?., ?????????? ?.?. (1989) ???? ? ??? ????? ??????????. ???????????: ?????, 167 ?.
???????? ?.?., ?????????? ?.?. (1989) ???????? ???????????????? ???????? ????????
???????? «????????» Pinus sibirica (Pinaceae). ???????????? ?????? 74 (4): 554-557.
?????????? ?.?. (1994) ? ??????????????? ????????? ? ???????? ??????? Pinus sibirica
(Pinaceae). ???????????? ?????? 79 (5): 63-71.
?????????? ?.?. (2006) ? ????????? ???????? ??????? ????? ??????????. ??????? ?????????? ???? 2: 43-54.
?????????? ?.?. (1991) ???? ? ????????????? ??????? ????? ??????????. ??????????? 1:
70-75.
?????????? ?.?. (1990) ????? ????? ? ???? ? ???????? ?????? ????? ??????????. ?????????
21 (2): 207-214.
# 33 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ??????????? ?????????? ???????????? ??????????? ???????????? ??????? ????? ??????????
?????????? ?.?., ????????? ?.?. (1992) ? ??????????? ????????????? ? ?????????????
???????????? ? ????? ??????????. ?????????. 23 (3): 268-279.
?????????? ?.?., ????????? ?.?. (1996) ????????? ? ???????? ????????? ????????? ????? ????? ??????????. ????????? 27 (1): 53-61.
?????????? ?.?., ????????? ?.?. (2000) ????????? ????? ????? ?????????? (Pinus sibirica
Du Tour) ?? ???????????? ????? ??????????. ????????? ?????. ????. 2 (1): 110-122.
??????? ?.?. (1969) ???????? ?????? ????????. ?.: ?????, 312 ?.
?????? ?.?. (1994) ??????????????? ?????? ??????????????? ???????? ????????. ?.: ?????, 269 ?.
??????? ?.?. (1973) ??????????, ?? ????????? ? ???????. ?.: ?????, 263 ?.
????????? ?.?. (1973) ??????? ???????? ????????? ??????????? ???????. ?: ????? ????.
??-?? ?? ?? ????. ???????????: ?????, ?. 40-52.
??????? ?.?. (1984) ???????????? ????????? ?????????? ???????? ?: ???? ??????? ? ????????? ????? ? ???????? ????????. (?.?. ???????, ???.). ?.: ?????, ?. 87-100.
?????? ?. (1963) ?????????? ????????. ?.: ???-?? ??????????? ??????????, 603 ?.
????????? ?.?. (1974) ??????????????????? ?????????????? ???????? ????? ???, ?????,
??????????? ? ????? ? ??????? ??????? ????? ???? ????. ?: ????????????? ???????????? ??
?????????????? ??????????? ????? ????. ?????????, ?. 47?53.
?????????? ?.?. (1990) ???????-?????????????? ?????????????? ? ???????? ????????
????? ????? ? ???????-????????? ??????? (???????-??????????????? ?????? ???????????
?????????). ??????????: ???-?? ??? ?? ????, 288 ?.
?????? ?.?. (1975) ?????????? ?????? ????????????????? ??? ??????? ??????? ? ?????????????? ???????? ?????????. ????. ?????. 2: 7?34.
???????? ?.?. (1988) ????????? ???????? ?????? ????????. ?., ?????, 560 ?.
Boyer W.D. (1970) Shoot growth patterns of young loblolly pine. Forest Sci. 16 (4): 472-482
Giertych M.M. (1967) Analogy of the differences between male and female strobiles in Pinus to
the differences between long- and short-day plants. Can. J. Bot. 45 (3): 1907-1910
Greenwood M.S. (1980) Reproductive development in loblolly pine: I. The early development of
male and female strobili in relation. Amer. J. Bot. 67 (10): 1414-1422
Kozlowski T.T. (1971) Growth and development of trees (V.1). New-York, Academic press, 443 p.
Lanner R.M.(1978) Development of the terminal bud and shoot of slash pine saplings. Forest Sci.
24 (2): 167-179
O?Reilly C., Owens J.N. (1988) Reproductive growth and development in seven provenances of
lodgepole pine. Canad. J. Forest Res. 18 (1): 43-53
Owens J.N., Molder M. (1974) Cone initiation and development before dormancy in yellow cedar
(Chamaecyparis nootcatensis). Can.J. Bot. 9: 2075-2084
Owens J.N., Molder M. (1979) Sexual reproduction of Larix occidentalis. Can.J. Bot. 57 (23):
2673-2690
Owens J.N., Pharis R.P. (1971) Initiation and development of western red cedar cones in response
to gibberellin induction and under natural conditions. Can. J. Bot. 49: 1165-1175
Ryan M.G., Waring R. (1992) Maintenance respiration and stand development in a subalpine
lodgepole pine forest. Ecology. 73(6): 2100-2108
# 34 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ??????????? ?????????? ???????????? ??????????? ???????????? ??????? ????? ??????????
Vorobjev V.N., Goroshkevich S.N., Savchuk D.A. (1994) New trend in dendrochronology: 2.
Method of retrospective study of seminiference dynamics in Pinaceae. Proceedings of International
workshop on Subalpine Stone Pines and their environment: the status of our knowledge; 1992
September 5-11; St.Moritz, Switzerland. Gen. Tech. Rep. INT-GTR-309. Ogden, UT: U.S. Department
of Agriculture, Forest Service, Intermountain Research Station, ?. 201-204
Age-Related Variability
of Generative Shoots Morphogenesis
of Siberian Stone Pine
Svetlana N. Velisevich and Sergey N. Goroshkevich
Institute for Monitoring of Climatic
and Ecological Systems SB RAS
10/3 Academichesky pr., Tomsk, 634055 Russia
On the example of male and female shoots in young (140 years), middle-aged (235 years) and old
(346 years) generative trees of Siberian stone pine (Ket-Chulym rivers, south-eastern part of Western
Siberia) the effect of tree age on the ratio of vegetative and generative processes is studied. The
traits characterizing vegetative development of generative shoots are reduced linear during the
reproductive period of ontogenesis. The traits of the generative development of shoots and indirectly
related vegetative traits reached maximum values in middle-aged trees. At the peak of reproduction
increases the direct and indirect relationship between vegetative and generative traits of shoots:
increased vegetative organogenesis in young trees has effected the cone initiation, reduced vegetative
development in middle age has a positive effect on the cone formation and continuation this trend
due to tree aging even strengthens this relationship. It is suggested that observed differences due to:
(1) changing the balance between vegetative and generative organogenesis in the seasonal cycle of
development of shoots and (2) ontogenetic changes in the root-leaf functional correlation.
Keywords: shoot, organogenesis, growth, ontogenesis, Siberian stone pine, Pinus sibirica Du Tour.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2011 4) 36-53
~~~
??? 581.524.34(571.5)
???????????? ?????? ?????? ????????????? ?????
? ????????? ? ??????? ????? ???????? ??????
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??????????? ????????? ???????????? ??? ?? ???,
?????? 630090, ???????????, ??. ???????????????, 101 1
Received 4.03.2011, received in revised form 11.03.2011, accepted 18.03.2011
??????????? ?????????????? ????????????? ?????????? ?????? ??????????, ?????????????
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? ???????????? ??????? ???? ? ?? ????????? ????. ??????? ?????????????? ??????
????????? ??????, ?????????? 257 ????? ?????? ?????????? ????????, 8 ????? ???? ? 4
??????????. ??????????? 43 ???? ? ??????? ??????????? ???????????, ???????????
?????????? ??????? ? ????????? ??????. ?? ???????? ??????? ?????????? ??? ??????????
? ??????? ????????????? ??????????, ?? ????? ??????? ? ??????? ????, ???????????????
?????. ?? ????? ??????? ?????? ??????????? ??????????? ???? ?? ??????????? ????, ?????
??? ???????????? ????????, ? ??? ????? ??????????? ???????????, ????????? ????????
?????????? ?????? ?? ????? ??????? ????????????? ??????????. ???????????? ??????
?????? ??????????? ????????????, ???????????? ?? ?????? ??????? ????????? ?????????.
?? ???????? ??????????????? ???????? ???? ???????? ?????? ?????????, ??????????? ???
?????? ??? (??????) ??????????????. ? ?????? ? ?????????? ???????? ?????? ???????????
??????????? ???????-???????? ?????? ? ???????????? ??????-???????????? ??????.
? ?????????? ??????? ????? ?????????? ? ???????? ??????? ???????? ? ??????????
???????? ???????? ??????????? ? ????????????. ?????, ??????? ? ??????? ???????? ?????,
???????? ???? ? ?????????? ?????? ?????? ??????? ????? ? ???????. ???????? ??????? ???????
?? ???????? ? ??????? ???????? ????? ????????? ????????????, ??? ??????? ?????????, ?
???? ???????????? ???????, ?????? ?????? ??????????? ??????? ????? ?? ?????.
???????? ?????: ???????, ??????????? ??????????, ??????????????, ???????-?????????
???????.
????????
????????????? ????????????? ?????????????? ? ?????????? ?? ??????????????
????? ????????? ??????????? ????????? ?
*
1
????????? ??????????? ????????? ????? ???????? (????????????, 1979; ????????, 1988;
???????? ? ??., 1999; ???????, 2005; ??????????, 2006; ?????????, ?????????, 2006
Corresponding author E-mail address: tishenko-1957@mail.ru
© Siberian Federal University. All rights reserved
# 36 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????? ?????? ????????????? ????? ? ????????? ? ??????? ????? ???????? ??????
? ??.). ? ?????????, ??????????? ?????????
????????????? ????????? ? ????????????
?????????????? ?? ????? ????????? ??????????. ?????? ????????????? ????? ? ???? ??
???????? ???????????????? ????? ????? ??????????. ???????? ?? ??? ?????????????
???????? ???? ???????? ? ????????? ??????
????? ?????????? ? ??????????????? ???????????? ??????????? ? ???? ??????????????
??????????????? ? ???????????? (??????????????, 1963; ?????, 1971; ?????????, 1985;
??????, 1994; ????????, 2002). ?????? ??????????? ????? ??????????? ????? ???????????? ?????, ?? ??????? ????????????
???????????? ??????????? ? ?????????. ?
???????? ?????? ???????? ???????????? ??
???????????.
?? ?????????? ???????? ?????????????? ????????????? ?????????? ?????? ??????????, ????????????? ?? 55 ?? 67є ?.?. ? ???????????? ??? ???? ??????????????: ???????
(??????? ??????? ????????, ????? ?????,
??????? ?????, ???????? ????? ? ??????????)
? ?????????. ?? ???? ????? ?????????? ?????????? ???????????? ????? ???????? ?????? 1-2 ? ???????, ?????????? ?????????
???????, ?? ??????? ???????? ???????????
????????.
?????????? ?????? ???????? ?????? ??
???? ?????? ?????????????? ???????? ?????.
?????????? ???????? ? ???????? ???????????? ???????????? ??????????? ? ?????????
??????? ?? ???????????? ???????. ??????? ????????? ??? ??????? ???????? ????????????
??????????, ????????????? ??????????????
?????? ??? ??????? ????????? ? ????????
??????, ????????????? ??? ??????. ? ???????? ?????? ???????????? ???????-?????????
??????? ???????? ??????? ?????????? ????? ???????????? ????????????? ???????
????????????????. ???????? ?????, ?? ????????? ? ??????? ???????????? ???????????
(???????????, ????????? ????????, ???????? ?? ????????? ????? ? ?????), ????? ???????????? ?? ??????-?????????? ?????????
??? ????????????? ??????????. ?? ??????,
?????? ??????????, ???? ????? ????????????? ??????????, ???????? ? ??????? ????? ??????.
??????? ???????? ????? ??????? ?? ??????? ????????? ????? ??????. ??-??????,
???????????? ?????? ? ????? ?????????????? ???????? ? ?????? ?????? ????????
?????? ????????? ??????? ????????????
??????? ? ???????????? ? ???????????? ??????????? (??????????, 1997; ?????????????, 2003). ??? ?????? ?????????? ?????? ?????: Festuca rubra, F. ovina, Bromopsis
inermis, Poa pratensis ? ??. ??-??????, ???????? ??? ????????????? ?????????? ????? ????????? ?????????? ?????????? ??????? ?????. ?-???????, ?? ???????????? ????? ??????
?????????? ????????? ????? ????? ?? ????
???????.
???? ??????: ???????? ????????????
?????????? ?????? ????????? ? ?????????
????? ? ???????? ?????????????? ???????????????? ??????? ? ?????????? ????????????? ?????, ??????????? ? ??????????,
? ?????????????? ?????????? ??? ?????????.
????????? ? ??????
??????? ???????????? ??????????? ?
2010 ?. ?? ???????? ??????????, ????????????? ?? ?. ????????? ????????? ???. ??
???. ????????? ?????-????????? ??????????? ?????? (???. 1). ? ??? ?? ????? ??????
?????????? ??????? ???? (??????? ???????
????????, ?????, ??????? ? ???????? ?????)
? ??????? ? ??????? ????? ???????????????
?????. ?????????? ????? ??????? ??????????,
????????????? ? ??????? ?? ????? ? ????????
????????????? ??????? ? ??????????? ???.
# 37 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????? ?????? ????????????? ????? ? ????????? ? ??????? ????? ???????? ??????
???. 1. ????? ????????????
??????? ? ???. ??????? ?????-????????? ??????????? ??????. ????? ????? ????????????? ?????????? ????? 1500 ??. ? ????? ??????
??????? ?????? ????? ??????? ?? 1 ? 1,5 ?,
??????? ?????????? ???????, ???????? ?????? 3 ? 5 ? ??????? ? ??????, ??? ???????,
??????????? ?????. ??????????? ?????????????? ?? ????????, ??????? ?????? ??????
?? ??????.
? ?????? ?????? ???????? 107 ??????????????? ????????, ??????????? ?? ????-
??????? ?????????. ???????? ?????? ??
????????? ????????????? ????? ?? ????
???????????? ??????? ?? ?????? ???????
??????. ?????? ???????? ??????????? ??
4 ?? 7 ?, ? ??????????? ?? ?????? ??????,
?????? ????? ???????? ???, ????? ???????
???????? ???? ????? 100 ??. ?. ??? ???????? ??????????? ??????? ?? ?????? ?????
??????????, ????? ???? ????? ???????????, ?
????? ???????? ?? ?????????? ?????????,
?? ??????????? ??? ???????, ????? ??? ??-
# 38 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????? ?????? ????????????? ????? ? ????????? ? ??????? ????? ???????? ??????
???? ???????????? ?????? ?????????? ???
?? ??????????? ??? ?????? ????????? ?????
(????????, ? ????????? ?-?? ????????? ???.
??????? ??????? ?????). ??????? ? ??????
??????????????? ???????????????? ?? ?????? ?????????? ?????????? ??? ????? ????????? ??????????? ????????.
??????????????? ???????? ???? ?????????? ? ??????? ????????? ?.?. ???????
IBIS (??????, 2007) ? ????????????? ?? ???????? ??????????????? ????????.
?????????? ??????? ??????? ????? ???
????????? ???? ? ??????? ?????? ???????
????. ???????? ?? ????? ????? ? ????????
?????????? ? ???? ??????? ? ????? ? ?? ????????? ???????????. ???? ? ??????? ????????????? ? ????????, ????????????? ???
???????-????????? ??????? (??????, 1979;
?????????????, 1985). ????????? ????????
?????? ?????????? ???????? ???? ?? ??????
?.?. ?????????? (1995), ???? ? ?? ?????? ?.?.
???????? ? ??. (2006).
?????????? ? ??????????
?? ???????? ????????????? ?????????
???? ???????? 257 ????? ?????? ??????????
????????, ??????????? ? 162 ????? ? 37 ??????????, ? ????? 8 ????? ???? ? 4 ???? ??????????. ??????? ????????? (Asteraceae, Poaceae,
Rosaceae, Fabaceae, Cyperaceae, Salicaceae,
Apiaceae, Caryophyllaceae, Polygonaceae ?
Ericaceae) ???????? 69,6 % ???????? ??????.
14 ???????? ???????????? 1? 2 ??????, 121
??? ???????? ?? ?????? ????.
??????? ?????? ????????? ?????? ?? ????? ? ???????? ?????????????? ??????????????? ??? ??????????? ?????, ???????? ?????????????? ?.?. ?????? (1982). ????????? 5
??????????? ???? ?????????? ???????????
?? ????????? ? ??? ?? ????? (????. 1). ?? ????
??????????? ?????? ??????????? ???? ? ????????? ??????? ? ??????????????. ??? ??
?????, ? ?????? ???? ? ??????? ???????????
????????, ???????, ????????? ?? ????????, ?
???? ????? ???????-????????????? ???????????? ????????? ? ???? ?????????????? ??????? ??????????? ? ?????? ???????? ????
? ???????????? ?????????. ??? ??????????
?? ???????? ??????????? ??????????, ????????????? ?.?. ??????? (1982).
? ?????????? ???????????? ?????????
??????? ?????? ??????????? ?????????? ????? ?? ?????? ??????????? ????????????? ?
?????? (???????, 1973; ???????, 1974; ???????, 1980; ????????, 1988). ?? ????????????
?????? ?.?. ????????? (1988) ? ????????????
??????????? ?????????? ????? ??? ???????????? ????????????? ? ???????? ???????????? ????????. ????, ?????????? ?????????? ??????????? ??????????? (1,0 ? ????),
????????? ? ????. 2 ? 6. ?? ?????? ?????, ???
????? ???????? ?????, ??? ? ??????? ???????????? ??????????? ????, ????????????? ?
?????? ?? ??.
????? ?? ???????? ????????????? ????????? ???????? 43 ???? ? ??????????? ??????????? 1,0 ? ????. ? ?? ????? 19 ?????
??????? ? ?????????? ???????? (????. 7) ? 24
???? (???????? ? ????. 2 ? 6 ?????? ???????) ? ?????? ? ????? ???? (???????). ????????
??????? ??????????????? ????? Equisetum
arvense, Chamaenerion angustifolium, Bromopsis
inermis ? Poa pratensis, ??????? ??????? ???????? ??????? ????????? ?? ????? ???????:
?? ?????? ?? ????????. ??? ?????? ???????????? ?? ?????? ?????????? ? ??????? ??????? ????????. ??????? ????? ????????????
????????????, ??? ???? ????? ??????????
?????????? ??????????? ?????????? ? ???????? ???????, ??? ?? ???????? ???????????
??????????? ??????????? ??????????.
?? ????. 7 ?????, ??? ?????????? ???????? ????, ????? ??? ?????????? ??? ? ??????, ????? ????????? ?? 2 ?????? ? ????????
# 39 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
??????? 1. ?????????? ????? ?? ???????? ?????????, ???????-????????? ???????, 2010 ?.
???????????????
??????
??????
??????????
???
?????
??????
??????????
????????
?????
???????
?????
????? ?????
? ????????
257
27
64
122
118
179
8
2
6
5
3
?
4
?
3
2
?
?
?????????
??????? 2. ?????????? ???????? ???? ?? ???????? ????????? ? ????????? ???? ???????-?????????
???????, 2010 ?.
????
1. Equisetum arvense L.
?????????????
???????
???????????
????????, %
???????????
???????????
??????????
1,00
27,4
27,4
2. Deschampsia cespitosa (L.) Beauv.
1,00
5,4
5,4
3. Poa pratensis L.
1,00
4,0
4,0
4. Bromopsis inermis (Leyss.) Holub
1,00
3,0
3,0
5. Chamaenerion angustifolium (L.) Scop.
1,00
2,6
2,6
6. Calamagrostis lapponica (Wahlenb.) C. Hartm.
1,00
1,7
1,7
7. Carex aquatilis Wahlenb.
0,57
2,5
1,4
8. Calamagrostis langsdorffii (Link) Trin.
0,57
2,0
1,1
??????????. ?????? ??????? ? ???? ? ??????????? ???????? (????. 2 - 6) ???????? ????, ???????????
??????? ??????????? ?????????? ?????? ? ????? ???? (???????) ??????????????.
??????? 3. ?????????? ???????? ???? ?? ???????? ????????? ? ????????????? ??????? ???????????????? ???????, 2010 ?.
????
1. Equisetum arvense L.
?????????????
???????
???????????
????????, %
???????????
???????????
??????????
20,0
1,00
20,0
2. Ceratodon purpureus (Hedw.) Brid.
0,65
11,2
7,3
3. Chamaenerion angustifolium (L.) Scop.
0,85
6,5
5,5
4. Equisetum sylvaticum L.
0,45
9,9
4,4
5. Poa pratensis L.
0,60
6,6
4,0
6. Calamagrostis langsdorffii (Link) Trin.
0,75
4,1
3,1
7. Festuca ovina L.
0,60
4,8
2,9
8. Calamagrostis lapponica (Wahlenb.) C. Hartm.
0,65
3,5
2,3
9. Deschampsia cespitosa (L.) Beauv.
0,85
2,2
1,9
10. Polytrichum juniperinum Hedw.
0,70
1,9
1,3
11. Bromopsis inermis (Leyss.) Holub
0,55
1,9
1,0
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
??????? 4. ?????????? ???????? ???? ?? ???????? ? ??????? ???????? ????? ???????-?????????
???????, 2010 ?.
????
1. Festuca rubra L.
?????????????
???????
???????????
????????, %
???????????
???????????
??????????
0,85
18,4
15,6
2. Ceratodon purpureus (Hedw.) Brid.
0,71
15,6
11,1
3. Equisetum sylvaticum L.
0,47
12,2
5,7
4. Polytrichum juniperinum Hedw.
0,60
9,5
5,7
5. Equisetum arvense L.
0,60
8,7
5,2
6. Chamaenerion angustifolium (L.) Scop.
0,96
4,0
3,8
7. Bromopsis inermis (Leyss.) Holub
0,93
3,8
3,5
8. Calamagrostis lapponica (Wahlenb.) C. Hartm.
0,78
3,9
3,0
9. Calamagrostis langsdorffii (Link) Trin.
0,58
4,6
2,7
10. Betula pubescens Ehrh.
0,78
2,9
2,3
11. Festuca ovina L.
0,29
7,2
2,1
12. Poa pratensis L.
0,20
6,6
1,3
13. Equisetum palustre L.
0,24
4,5
1,1
??????? 5. ?????????? ???????? ???? ?? ???????? ? ??????? ??????? ????? ???????-?????????
???????, 2010 ?.
????
1. Calamagrostis epigeios (L.) Roth
?????????????
???????
???????????
????????, %
???????????
???????????
??????????
0,92
11,8
10,8
2. Equisetum arvense L.
0,62
9,9
6,1
3. Chamaenerion angustifolium (L.) Scop.
0,92
4,6
4,2
4. Equisetum sylvaticum L.
0,62
6,6
4,1
5. Hieracium umbellatum L.
0,85
3,9
3,3
6. Trifolium pratense L.
0,77
4,1
3,2
7. Calamagrostis langsdorffii (Link) Trin.
0,54
5,0
2,7
8. Agrostis gigantea Roth
0,38
7,0
2,7
9. Poa angustifolia L.
0,38
6,4
2,4
10. Elytrigia repens (L.) Nevski
0,62
3,6
2,2
11. Poa palustris L.
0,46
4,7
2,2
12. Bromopsis inermis (Leyss.) Holub
0,54
3,8
2,0
13. Phleum pratense L.
0,54
3,6
1,9
14. Petasites frigidus (L.) Fries
0,31
5,2
1,6
15. Poa pratensis L.
0,38
4,0
1,5
16. Crepis tectorum L.
0,62
2,4
1,5
17. Amoria hybrida (L.) C. Presl
0,38
3,4
1,3
18. Rubus idaeus L.
0,46
2,5
1,2
19. Festuca pratensis Huds.
0,54
2,1
1,1
20. Artemisia vulgaris L.
0,92
1,2
1,1
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
??????? 6. ?????????? ???????? ???? ?? ???????? ? ???????? ????? ????? ? ???????? ???????????????? ???????, 2010 ?.
?????????????
???????
???????????
????????, %
???????????
???????????
??????????
0,83
5,5
4,6
2. Agrostis gigantea Roth
0,85
5,4
4,6
3. Tussilago farfara L.
0,92
4,1
3,8
????
1. Agrostis tenuis Sibth.
4. Elytrigia repens (L.) Nevski
0,92
4,0
3,7
5. Salix dasyclados Wimm.
0,25
13,3
3,3
6.Bromopsis inermis (Leyss.) Holub
0,67
4,0
2,7
7. Poa palustris L.
0,75
3,6
2,7
8. Poa angustifolia L.
0,17
15,0
2,6
9. Calamagrostis epigeios (L.) Roth
0,67
3,8
2,5
10. Cirsium setosum (Willd.) Bess.
2,2
1,00
2,2
11. Trifolium pratense L.
0,75
3,0
2,2
12. Achillea millefolium L.
0,83
2,7
2,2
13. Equisetum sylvaticum L.
0,58
3,7
2,1
14. Deschampsia cespitosa (L.) Beauv.
0,42
4,2
1,8
15. Secale cereale L.
0,08
20,0
1,6
16. Festuca pratensis Huds.
0,83
1,9
1,6
17. Trifolium medium L.
0,92
1,7
1,6
18. Equisetum arvense L.
0,42
3,6
1,5
19.Urtica dioica L.
0,42
3,6
1,5
20. Trifolium arvense L.
0,50
2,8
1,4
21. Pimpinella saxifraga L.
0,75
1,8
1,4
22. Plantago major L.
0,92
1,4
1,3
23. Chamaenerion angustifolium (L.) Scop.
0,67
1,9
1,3
24. Amoria repens (L.) C. Presl
0,92
1,3
1,2
25. Potentilla anserina L.
0,58
2,0
1,2
26.Sonchus arvensis L.
0,83
1,4
1,2
27. Phleum pratense L.
0,83
1,3
1,1
28. Poa pratensis L.
0,50
2,0
1.0
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????? ?????? ????????????? ????? ? ????????? ? ??????? ????? ???????? ??????
??????? 7. ????????? ??????????? ?????????? ?????, ????? ??? ?????? ??? ? ?????? ??????????????
?????? ????????? ???????-????????? ???????, 2010 ?.
??????????? ??????????? ??????????
????
??????
??????????
????????
?????
???????
?????
????? ?????
? ????????
1. Equisetum arvense
27,4
20,0
5,2
6,1
1,5
2. Deschampsia cespitosa
5,4
1,9
?
?
1,8
3. Poa pratensis
4,0
4,0
1,3
1,5
1,0
4. Chamaenerion angustifolium
2,6
5,5
3,8
4,2
1,3
5. Calamagrostis langsdorffii
1,1
3,1
2,7
2,7
?
?
2,9
2,1
?
?
7. Bromopsis inermis
3,0
1,0
3,5
2,0
2,7
8. Calamagrostis lapponica
6. Festuca ovina
1,7
2,3
3,0
?
?
9. Ceratodon purpureus
?
7,3
11,1
?
?
10. Polytrichum juniperinum
?
1,3
5,7
?
?
11. Equisetum sylvaticum
?
4,4
5,7
4,1
2,1
12. Calamagrostis epigeios
?
?
?
10,8
2,5
13. Trifolium pratense
?
?
?
3,2
2,2
14. Phleum pratense
?
?
?
1,9
1,1
15. Agrostis gigantea
?
?
?
2,7
4,1
16. Elytrigia repens
?
?
?
2,2
3,7
17. Poa palustris
?
?
?
2,2
2,7
18. Poa angustifolia
?
?
?
2,4
2,6
19. Festuca pratensis
?
?
?
1,1
1,6
??????????. ?????? ??????? ???????? ?????????? ???????????? ??????????? ?????????? ???
??????? ????.
?? ???????? ??????? ?????????? (? ??????,
?????????? ? ???????? ?????) ? ???????? ??
????? ??????? (? ??????? ?????, ????? ?????
? ????????). ??????????? ?????, ???????? ?
?????? ??????, ????? ??????? ???????????????. ??? ???????? ??????? ? 4 ? 5 ????????, ?? ?????????? ???????? ???????????
?????????? ????? ? ???????? ???????. ????
4 ???? (Festuca ovina, Calamagrostis lapponica,
??? Ceratodon purpureus ? Polytrichum
juniperinum) ?? ?????? ???????? ???? ??
???????? ????? ???????? ?????. ????????
?? ???? ????????, ??? ????? ?????????? ???????? ????? ???? ?????? ?????????? ??????????? ??????????? ????-??????????,
??????? ?? ?????????? ??????????????
? ??????? ??????? (?????, 1971; ????????,
2002). ?????? ?????? ???????????? ????????
??????, ??????????????? ???????, ???????
?? ???????? ?? ???????? ? ???????? ???????
???? ????? ??? ????? ?????? ??????????.
?????????, ??? ????????? ??????? ????
????? ?? ???????? ? ??????? ????? ???? ????? ??????? ??????????, ??? ? ????? ????? ? ????????, ???? ??????? ??????????????
??? ??????? ? ?????? ??????? ???????????. ?
???? ??????, ??? ?????????? ????????????
??????????? ?????????????, ??????????? ????????? ?????????? ????? (Crepis tectorum,
Trifolium arvense, Secale cereale).
# 43 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????? ?????? ????????????? ????? ? ????????? ? ??????? ????? ???????? ??????
??????? ?????, ??????????? ???????
??????????? ?????????? ?????? ? ????? ???????????? ???? (???????), ??????????????? ?
????????? ??????????????? ???????? ???????????? ?????????????? ?? ????? ?????????
??????????. ???, ? ???????? ????? ?? ???????
????????? ?????? ??????? Betula pubescens,
? ??????? ????? ??????? Rubus idaeus ?
Hieracium umbellatum, ? ? ????? ????? ? ???????? ?????????? ????????? ? ??????? ?????:
Agrostis tenuis, Achillea millefolium, Trifolium
medium ? ??. ? ??????? ????? ???????? ????????? ??????? ?????????? ????????????
???????? (Crepis tectorum, Artemisia vulgaris),
???? ??????? ? ???????? ??????? ????????.
?? ????? ??????? ?????????? (????? ?????
? ????????) ????? ???????????? ????? ??????????? ???????????. ???????? ???????
????? Cirsium setosum, Sonchus arvensis, Urtica
dioica, Trifolium arvense. ? ?????? ??????????? ????? ?????? ???????? ?????? ???????????? ???????? ????, ??? ??????? ? ???????
??????????????? ?????????? ? ?????????????? ???????, ??????????????? ??????????? ? ??????? ?????????. ? ?????? ???????
?????????? ????????? Carex aquatilis, ? ???????? ????? ? Equisetum palustre, ? ??????? ????? ? Petasites frigidus, ? ????? ????? ? ???????? ? Tussilago farfara ? Salix dasyclados.
??????????? ?????? ???????????????
? ???, ??? ?????? ??????????? ???? ??????
????????? ?? ?????? ???????????? ?????????
???????????????. ????????? ?????? ? ???????? ??????? ???????? ?????? ?????????? ?
???????? ?? ???? ????????? ????? ? ??????? ????????????? ?????????? ? ????? ???????????? ????????????, ? ? ???????????
? ????? ??????? ?? ???????? ???????????
??????? ?????, ??????????????? ?????????????????. ? ?????????? ? ???????? ????????
????? ??????????? ????????? ?????? ???????? ??? Ceratodon purpureus ? Polytrichum
juniperinum. ?? ????? ??????? ?????? ???????????? ??????????? ???? ?? ??????????? ?????, ??????????? ??????????? ?????
??????? ????? ?????? ??????????. ???????????? ???????????? ??????????? ???????
??? ??????????? ?? ????? ????????? ?????
???????? ?????? ?????? (??????????????,
1963; ?????, 1971; ?????????, 1985; ????????, 2002).
?????????????? ?????? ?????????????
????????? ???????????? ????????????, ??????? ?????????????? ? ???? ?????????
?????????. ??? ??????????????? ????????????? ???????????? ???????? ??????? ?????? ??? (??????) ?????????????? ? ????????
??????? ????????????. ??? ?????? ? ?????????? ?????????? ???????? ? ?????????
?????????? ?? ???????????? ?.?. ????????? (1964), ??? ??????????? ??????????? ????????????? ????? ?????. ????? ???????????
???????? (???) ???? ????????? ??????????
?? 20 ?? 60 %. ? ??????? ???? ????? ?????????? ?? ???????? ?????? ???????????
????????? ?????????? ???????? ???? ? ???
80-100 %, ???? ????????????? ????? ?????,
??-????????, ??????????.
??? ???????? ?????? ??????, ?????????
????????????? ???????? ??????????????
(??????????, 1975; ?????????, 1985; ????????, 1988; ???????? ? ??., 1999), ?????
???????? ????? ??????????????, ????????
?? ???????? ? ??????????? ????????? ????????? ? ??????????? ?????????? ????????? ????????. ????? ????????, ???????? ??
?????????? ????????????? ??????????????
????????????? ???????, ???????? ?????????? ? ?????-?????????? ???????? ?????????,
??? ?????????????, ??????? ???????????? ?????????? ???????? ? ???????????? ??????????? , ??????? ???????? ?????????? ????????????. ????????????? ?????????? ?????????
? ???????????????? ???????? ?? ?????????-
# 44 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????? ?????? ????????????? ????? ? ????????? ? ??????? ????? ???????? ??????
??? ??????????????, ????? ???????? ??????, ????? ????????????? ??? ??????????, ???
?????? ??????????? ?????????, ??????? ????????? ????????? ????????? ????? ????????
??????. ?? ??????? ????, ????????? ??????
???????????? ????????? ? ????????? ???????????, ????? ?? ??????????????? ????????? ??????? ????????? ?????? ???????????
???????? (?? 15 %), ????? ???????? ?? ????.
? ????? ? ??????? ????????? ?????????? ?
??????? ? ?????? ?????? ?????? ??????????
?????? ?????????? ????????????. ???????,
??????????????? ??????????? ? ?????????
???????, ???????? ?????, ???????????? ???????? ????? ? ???????? ????? ?????. ?????,
??? ???????, ??????????? ???? ?????????
?????????? ????????? ? ??????????? ?????, ????????????? ?? ??????????? ?????????. ??? ?????? ?? ????????? 10-15 %. ???-
??????. ?? ??????????????? ???????????
? ???????????? ??????? ???????? ? ?????
? ???? ?????????????? ??????? ?????????????? ??????????? ???? Tripleurospermum
perforatum ? Polygonum aviculare. ????? ??????????? ? ???????????? ???? ??????????
????? Deschampsia cespitosa, Poa pratensis,
Bromopsis inermis ? Calamagrostis lapponica.
? ??????, ????? ??????????? ????? ?????? ????????????? ?????? Calamagrostis
neglecta, ??????????? ???????????? ????????????? ????????: Carex aquatilis, Eriophorum
scheuchzeri ? ??. ? ??????? ????????? ??????
??????????? ????????? ?????????? ????????????? ?? (Salix phylicifolia, ???? S. glauca)
? ?????????? ??????? Betula nana. ???????
???????????? ??????????????? Chamaenerion
angustifolium, ???? ?????? ??? ???? ????? ????????. ? ?????, ?????? ??????????? ? ????-
??? ??????? ??? ???????????? ???????????
???????????? ?? ??????? ??????? ??????,
???????? ? ??????, ???????? ???????????
?? ?????, ??? ??????????? ????? ?????? ???????? ????????. ??????? ?? ???????????
?????????, ???????? ?? ???????? ???????
??????????, ????? ???? ????????? ????????
?? ????????, ??????????? ???????? ?????????????? ?? ????????? ????????.
?? ???????? ???????? ???????? ???????
???????? ?????? ?????????, ?????????????
?? ???????? ????????? ? ?????? ????? (????????) ?????????????? (????. 8).
?? ???????? ????????? ? ????????? ????
(????????? ?-? ?????-????????? ??????????? ??????), ??? ???????????? ??????????????
???????????? ?????????? ? ???????????????????-????????????? ????????, ???????? ?????? ???????-???????? ??????????.
??????? ????????? ?? ? ??????? 15 ????? ??
100 ??. ?, ??? 35 ? 60 %. ???????? ?????????? ??????????? ????????? Equisetum
arvense, ???????? ???????? ? ??????? ?????
??????? ????? ??????????????? ? ?? ??????
???????? ????. ??????????? ???? ??????
??????? ?? ????????? ? ?????????? ???????????? ???????????????. ????????? ????????? ???????????, ???????????? ???? ????????? ?? 1? 2 ????? (Polytrichum juniperinum
? Ceratodon purpureus), ???????? ??????? ??
????????? 5 %.
?????????? ??????????? ???????????? ??????????? ? ????????????? ???????
(???????????, 1954; ?????????, 1985; ????????, 1988) ? ???, ??? ?? ????????????? ?????????? ???????????? ?????? ? ?????????
???????? ???????? ?????? ??????????? ??????????????? ???????? ??????????????. ??
?????? ???? ???????, ? ???? ????????? ????????? ????? ??????????????? ???????????
???????????? ? ?????????? (Calamagrostis
neglecta) ?????? ??????????? ??????????
?????????? (Poa pratensis) ??????????.
????????? ????????????? ???? ?????????
??????? ? ????????????? Equisetum arvense,
???????? ?? ???????? ??????? ??????.
# 45 #
????????
?????
??????????
??????
????
(???????)
???. ?????????? ?
???. ???????
???. ??????????
?????????? ?-??
? ???. ???????
?????????? ?-??
????????? ?-?
?????-?????????
??, ???. ???.
???????
????????? ?-?
?????-?????????
??
????????? ?-?
????????? ?-?
?????-?????????
??
??????????????
?????????
17
16
14
24
13
15
??. ???-??
????? ?? 100
??. ?
40 ?100
30 ? 90
30 ? 90
20 ? 60
20 ? 60
35 ? 60
???
?????????
?????, %
10 ? 90
5 ? 60
10 ? 45
5 ? 50
4?8
1?5
???
????????
?????, %
??????????? ????
Deschampsia cespitosa, Calamagrostis
lapponica, Poa pratensis,
Tripleurospermum perforatum (Mйrat) M.
Lainz
Equisetum arvense, E. sylvaticum,
Bromopsis inermis, Carex aquatilis,
Calamagrostis langsdorffii, C. lapponica Galium mollugo L., Polygonum aviculare
L., Amoria repens, Eriophorum
scheuchzeri Hoppe
Ceratodon purpureus, Equisetum
Calamagrostis langsdorffii, C. neglecta,
arvense, E. sylvaticum, Calamagrostis
Ceratodon purpureus, Betula nana L.,
langsdorffii, C. neglecta (Ehrh.) Gaertn., Festuca ovina, Poa pratensis, Luzula
Festuca ovina, Poa pratensis, Tanacetum multiflora (Ehrh.) Lej., Rubus arcticus
bipinatum (L.) Sch. Bip., Chamaenerion L., Antennaria dioica (L.) Gaertn.,
angustifolium, Hierochloe arctica C.
Vaccinium uliginosum L., Pyrola minor
Presl
L., Hieracium umbellatum, Euphrasia
parviflora Schag., Carex globularis L.
Betula pubescens, Ceratodon purpureus,
Equisetum arvense, E. sylvaticum,
?
Calamagrostis langsdorffii, Phleum
pratense, Chamaenerion angustifolium,
Bromopsis inermis
Festuca ovina, Deschampsia cespitosa,
Festuca rubra, Ceratodon purpureus,
Polytrichum juniperinum, Calamagrostis Arctostaphylos uva-ursi (L.) Spreng.
langsdorffii, C. lapponica, Equisetum
arvense, E. sylvaticum, Descampsia
cespitosa, Chamaenerion angustifolium,
Bromopsis inermis, Hieracium
umbellatum
Festuca rubra, Ceratodon purpureus,
Calamagrostis epigeios, Pinus sibirica
Polytrichum juniperinum, Calamagrostis Du Tour
langsdorffii, Festuca ovina, Poa
pratensis
Equisetum arvense, Deschampsia
cespitosa, Carex aquatilis
?????????
??????? 8. ?????????????? ???????????? ????????? ?? ???????? ????????? ???????-????????? ???????, 2010 ?.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????
????? ?
????????
???????
?????
????????
?????
????????,
?????????,
????????,
????????????
?-?? ?????????
???.
???. ?. ?????????
??. ???????????
??????????? ?-??
?????-??????????
?? ? ?. ??????
?. ???????????
? ??. ????????
????????? ?-??
????????? ???.
???. ?. ????????
?????????? ?-??
??????????? ????. 8
56
47
46
19
24
30 ? 90
80 ? 90
60 ? 100
30 ? 60
20 ? 60
?
?
?
0?5
5 ? 45
Salix dasyclados, Equisetum sylvaticum,
Elytrigia repens, Bromopsis inermis
Tussilago farfara, Equisetum sylvaticum,
E. arvense, Trifolium arvense, T.
pratense, Deschampsia cespitosa,
Pimpinella saxifraga, Poa palustris, P.
angustifolia, Calamagrostis epigeios,
Ranunculus acris L., Pinus sylvestris,
Melilotus officinalis (L.) Pall., Secale
cereale
Equisetum arvense, Crepis tectorum,
Chenopodium album L. Chamaenerion
angustifolium, Calamagrostis epigeios,
Trifolium pratense
Equisetum sylvaticum, E. arvense,
Bromopsis inermis, Phleum pratense,
Calamagrostis langsdorffii,C. epigeios,
Hieracium umbellatum, Rubus idaeus,
Poa palustris, P. angustifolia, P.
pratensis, Agrostis gigantea, Amoria
hybrida, Trifolium pratense, Petasites
frigidus, Festuca rubra
Ceratodon purpureus, Equisetum
sylvaticum, E. palustre, Rumex
acetosella L., Festuca rubra,
Calamagrostis lapponica, C. epigeios,
Trifolium pratense, Carex globularis,
Chamaenerion angustifolium, Bromopsis
inermis, Puccinellia borealis Swall.
Lepidotheca suaveolens (Pursh) Nutt.,
Picris hieracioides L., Leucanthemum
vulgare Lam., Erigeron acris, Prunella
vulgaris L., Trifolium arvense, Veronica
chamaedrys L., Deschampsia cespitosa,
Potentilla anserina, Linaria vulgaris L.,
Geuim aleppicum Jacq.
?
Elytrigia repens, Equisetum sylvaticum,
Salix caprea L., S. cinerea L., Bromopsis
inermis, Phleum pratense, Festuca
pratensis, Calamagrostis langsdorffii,
Cirsium setosum, Pinus sylvestris,
Taraxacum officinale Wigg., Rubus
idaeus, Poa palustris, P. angustifolia, P.
pratensis, Agrostis gigantea, A. tenuis,
Stellaria graminea L., Vicia cracca L.,
Lathyrus pratensis L. ? ??.
Melampyrum pratense L., Equisetum
sylvaticum, E. palustre, Trifolium
pratense, Amoria repens, Populus tremula
L., Pinus sylvestris L., Salix myrsinifolia
Salisb., Picea obovata Ledeb., Festuca
pratensis, Dactylorhiza maculata (L.)
Soу , Orthylia secunda (L.) House, Carex
globularis, C. aquatilis
Crepis tectorum, Chenopodium album,
Senecio vulgaris L., Erigeron acris L.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????? ?????? ????????????? ????? ? ????????? ? ??????? ????? ???????? ??????
? ????????????? ??????? ????????????
?????????????? ???????????? ? ????????
?????????? ???????? ? ??????????????
????????????. ?? ???????? ????? ????? ??????????? ?????????? ? ??????????????
?????? ? ??????. ?????? ?????? ? Equisetum
arvense, ?????????????? ? ????????? ????,
?? ?????? ??????????? ? ????? ???????????? E. sylvaticum. ?? ?????? ??????? ???????
? ???????????? ?????????? Calamagrostis
langsdorffii, C. lapponica ? Deschampsia
cespitosa. ??????? ????????????? ?????????
Tripleurospermum perforatum, Salix phylicifolia,
Polytrichum juniperinum.
?????????? ?????? ????????? (????????? ?-?) ? ?????????? (????????? ?-?) ???????? ?????????? (???. 1) ?????? ? ??????
??????? ????? ???????????? ????????.
?? ????????? ??????? ??????????? ???????? ??????????, ??????? ? ??????????
? ????????? ????. ???????? ??????????? ?
??????? ???????????? (? ??????? 13 ?????
?? 100 ??. ?.), ??????????? ???????? (20 ?
60 %), ??????? ? ??????????? ????????????
?????, ?????? ??????????. ??????, ?????? ?
??????????? ????? ????????? ?????????, ?
Poa pratensis ?????? ???????? ? ?????? ??????????. ???????? ????????? ????????? ???????? ? ???????? Calamagrostis lapponica
? Bromopsis inermis. ????? ??????????? ????? ????? ??????? Carex aquatilis, Galium
mollugo, Polygonum aviculare, Amoria repens,
Eriophorum scheuchzeri.
??????????, ????????????? ?? ???????? ??????? ??????, ?????????? ?????
??????? ??????? ?????????? (? ???????
24 ???? ?? 100 ??. ?), ????????? ??????? ???????? ??????? (??????????? ???????? ??
50 %) , ??????????????? ???????????????
Ceratodon purpureus, ??????? ????????????
(??? ????? ??????) ???? ??????????? (Betula
nana, Salix glauca, S. cinerea), ????????????
(Vaccinium uliginosum) ? ???????? ???????????. ??? ????????? ????? ?????????? 20 ?
60 %. ????????? ??????????? ?????????
???????? Equisetum arvense, E. sylvaticum,
Calamagrostis langsdorffii, C. neglecta,
Festuca ovina, Poa pratensis, Chamaenerion
angustifolium, ?????? Tanacetum bipinatum.
??????? ??????????? ????? Luzula multiflora,
Rubus arcticus, Antennaria dioica, Pyrola minor,
Hieracium umbellatum, Euphrasia officinalis,
Carex globularis, Petasites frigidus, Pedicularis
labradorica ? ??. ??????? ?????????????
?????????? ?? ?????? ???????? ?????? ????????? ?.?. ???????? ? ??. (1999). ?????
????????, ??? ?? ???????? ??????? ?????????? ????? ??????? ?????????? ??????? ?????
? ?????????? ???????????? ??????????????,
??? ????????? ????. ?????? ??? ????????
??????? ?????????? ?? ??? ??????? ?????????? ? ?????????? (??????????????? ?????????????? ? ??????????????? ??????, ??????????? ????????????? ??????? ? ???????
?????????? ????????).
?????????????? ?????? ??????? ???????? ????? ???????? ????????????. ????? ???????? 4 ?????? ?????????, ?????????????
????? ????? ?? ??????????????? ???????.
?????? ?????? ?????????? ??????????,
???????????????? ? ???????? ????? ???????,
?? ??????????? ? ???????????? ???. ???????.
???????????? ?????????????? ????????????
?????????????
???????-??????????????
??????, ?? ??????? ??? ??????????? ??????
????. ??? ????????? ???? ?????? ?????????? ????????? ??????? ????????? ? ? ???????
14 ????? ?? 100 ??. ?. ??? ?????????? ?? 30
?? 90 %. ????????????? ?????? ?????????
???????? ??????? ?????????? ??????, ????????????? ????????? ?????? Betula pubescens
? ???????????? 1 ? 2 ? ? ??????. ??? ???????, ??????? ????????? ?????? ? ???????
????? ?????? ??????. ???????????? ?????
# 48 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????? ?????? ????????????? ????? ? ????????? ? ??????? ????? ???????? ??????
????????? ??????? ???????????, ???? ???????????? ?????: ??? ? ?????. ?????? ???
Salix cinerea, S. phylicifolia. ? ???????? ?????
?????????? Equisetum arvense, E. sylvaticum,
Calamagrostis langsdorffii, Bromopsis inermis,
Chamaenerion angustifolium, ?????? Phleum
pratense. ? ??????? ???????????? ??????????? Festuca rubra, Calamagrostis lapponica.
? ??????????? ??????? ??????? ??????
????? Ceratodon purpureus ? Polytrichum
juniperinum, ??????? ??????? ????? ???????? ?? 45 % ??????? ?????????.
?????????? ?????? ?????? ??????????
??? ??????-???????? ????? ??????? (???????
?????????? ???. ?????????? ? ???. ???????).
?????? ????? ?????????? ???????? ????????
???????, ??????????? ???????? ??????????????. ??????? ??????? ????????? ?????????
???? ?????? 16 ????? ?? 100 ??. ?, ??? ????????? ????? 30 ? 90 %. ????????? ???? ?? ???????, ??????????? ???? ????????? ???????
?????? ???????? ? ???????????, ? ?????
?? ?????????? ?? ???????????, ???? ????? ? ?????????????? ??????, ????? ?????,
??? Calamagrostis epigeios, Rumex acetosella,
Leucanthemum vulgare, ??????? Pinus sibirica.
??????? ??????? ???????????? 17 ????? ??
100 ??. ?, ??? ????????? ????? ???????? ??
40 ?? 100 %. ????????? ???? ??????? ???????? ?????, ?? ?? ?? ????? ????? ???????
???????????, ??? ? ?????? ??????. ? ???????
????????? ?????? ???????????? ????????
????: Rubus chamaemorus, Ledum palustre,
Eriophorum
Chamaedaphne
calyculata,
polystachion. ?? ??????? ?????????? ?????????? ?????? ? ?????????? ???????. ???????? ???????? ????? (Ceratodon purpureus ?
Polytrichum juniperinum) ????????? 90 %.
?????????? ????????? ?????? ??????????? ?? ????? ??????? ??????? ????????
?????, ? ???????????? ?. ?????????, ??? ???????????? ?????????????? ???????????? ????????????? ?????? ? ?????????????? ???,
?????, ????? ? ?????. ??? ?????????? ????-
?????????? Salix cinerea, Betula nana. ?????
????????? ?????????? ?????. ????????????
Festuca rubra, ???????? ??????? ????? ????????? 60 %. ???????????? Calamagrostis
lapponica, C. langsdorffii, Chamaenerion
angustifolium, Bromopsis inermis. ???? ??????
??????? ?????????. ???????????? ??????,
?????????????? ? ?????? ?????? ?????????,
???????? Festuca ovina, Deschampsia cespitosa
? Arctostaphylos uva-ursi. ??? (Ceratodon
purpureus ? Polytrichum juniperinum) ????????? 5 ? 60 % ??????????? ?????????.
?????? ?????? ????????? ??????? ? ??????????? ????? ????????????? ??????? ??
??????? ?????? ???. ?????????? ? ???. ??????? (????????? ?-?). ?? ???????? ?????????? ??????????? ??????????????? ??????
? ?????? ?????????? ???????? ????????????????????? ????. ?????? ????? ??????
????? ? ?? ??????? ???????? ??????. ????-
??????? ?????????? ?? ??????? ???????????? (24 ???? ?? 100 ??. ?) ? ???????????????
??????? ?? ????????? ????. ?? ????????????? ?????????? ??????????? ????? ?????, ???
Equisetum sylvaticum, E. palustre, Dactylorhiza
maculata, Orthylia secunda, Salix myrsinifolia,
???????? ?????, ?????, ???, ???? ?????. ??????? ??????? ? ??????? ????????? ?????????
??????? ????? ? ??????? Trifolium pratense,
Amoria repens, Festuca pratensis, Elytrigia
repens, Vicia cracca. ?????? ???????? ??????
????? ???????? ????????? Carex globularis
? C. aqatilis. ??? ????????? ????? 20 ? 60 %, ?
??????????? ????? ??????? ?????? (?? 40 %)
????????? ?????? Ceratodon purpureus. ????
Polytrichum juniperinum, ???????? ? ????????
???????, ??????? ???????????. ????????
??????????? Aulacomnium palustre.
?? ???????? ?????????? ? ??????? ??????? ????? ??????? ??? ?????? ?????????.
# 49 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????? ?????? ????????????? ????? ? ????????? ? ??????? ????? ???????? ??????
?????????? ?????? ?????? ??????? ? ???????? ????? ??????? ? ?? ??????? ?? ??.
??????????? ?? ?. ???????. ????????????
??????????????, ??????????? ? ??????????????, ???????????? ????????? ?????? ??
???????? ?????? ? ????????. ?? ????????
??????????? ??????????? (??? 30 ? 60 %)
?????? ? ????????? ??????? ????????????? (19 ????? ?? 100 ??. ?). ????? ???????
?????????? ????????????????? ???? ?
??????? ??????????: Equisetum arvense,
Chamaenerion angustifolium, Calamagrostis
epigeios. ??????? ??????????? ?????
Artemisia vulgaris, Hieracium umbellatum,
Dactylorhiza maculata, Salix viminalis. ???????????? ?????? ???? ?????? ???????? ??????????? ????-?????????? Crepis tectorum,
Chenopodium album, Senecio vulgaris, Erigeron
acris. ????? ????, ??????????? ? ??????????? ???????????? ???? Sonchus arvensis,
Puccinellia hauptiana, Polygonum aviculare,
Cirsium setosum. ??????? ???? ? ??????????? ?? ???????. ???????? ???????????
Aulacomnium palustre, ???????? Ceratodon
purpureus ?? ????????? 5 %.
?????????? ?????? ?????? ??????????
??? ????? ????? ????????????? ??????? ?
?? ?. ???????????? ?? ??. ???????? ?????????
?-?? ????????? ???. ?? ???? ??????? ???????????? ?????????????? ???????????? ???????? ?????? ? ????????????? ????? ? ?????.
?????????????? ?????? ????? ??????????
?????? ????????, ??? ? ???????? ???????.
????? ??????????? ??????????, ??????? ??
??????? ? ????????? ? ??????? ??????????? ????? ??? ???????? ??????, ???????
??? ??????? ??????? ????? ?? ??????????.
??-????????, ??????? ????? ? ???? ???????????? ??????? ?????? ??????, ?? ???????
??????? ??????????????, ??????????? ???????????? ???????? ? ???? ?????, ????? ?
?.?., ???????????? ?? ?????. ????????????
??????? ????????? ???????????? ??????????????? ?????????? ?????????. ? ?????????? ?????????? ????????? (??? 60 ? 100 %)
??????, ??????? ?? ??????????????? ???????
(? ??????? 46 ????? ?? 100 ??. ?). ?????????
2-3-???????. ??????? ???????? ?????????
????????????? ???????? ??????? ??????
? ????????????, ??????? ? ??????? ???????: Elytrigia repens, Phleum pratense, Festuca
pratensis, Poa palustris, P. angustifolia, Agrostis
gigantea, A. tenuis, Tanacetum vulgare. ? ??????? ????????? ????????????? ????????????
?????? ?????? ? ??????? ???????????: Vicia
cracca, Lathyrus pratensis, Achillea millefolium,
Amoria hybrida. ? ?????? ????????? ???????????? ????? Equisetum sylvaticum ? E.
arvense, ? ??????? ????????????? ??????? ? ???????? ????: Taraxacum officinale,
Stellaria graminea, Prunella vulgaris, Plantago
major, Petasites frigidus, Carex globularis ?
??. ? ??????? ????? ?????? ? ?????? ????????? ??????? ?????? ???????? Polygonum
aviculare,
Tripleurospermum
perforatum,
Lepidotheca suaveolens, Melilotus albus, M.
officinalis, Tussilago farfara. ? ?????? ?????
???????? ?????????????? ????????? ? ??????? ??????????? (Rubus idaeus, Salix caprea,
S. cinerea, S. viminalis) ? ???????? ?????????
?????: ?????? Betula pendula, ????? ? ?????.
??????????? ???? ?? ???????.
?????????????? ?????? ? ???????? ????? ????? ? ???????? ( ?? ???. ???? ?? ?.
?????????) ???????????? ??????? ???????? ????????????, ???????? ?? ??????????????? ??????? ? ???????????? ??????,
????????????????? ?? ??? ??????? ????.
????? ?????, ???????????? ??????????????
?? ???? ??????? ?????????? ????????????
????????????? (? ???????? ?????, ?????,
???) ? ???????????????? ??????, ? ? ?????????? ?-?? ?????????????? ???????? ???? ?
????? ? ????????. ??????? ?? ???????? ???-
# 50 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????? ?????? ????????????? ????? ? ????????? ? ??????? ????? ???????? ??????
???????? ?????? ?????? (Salix dasyclados, S.
pentandra, S. cinerea) ???????, ?? ??????????? ???????? ?????? ???????? ????. ???, ???
???????, ???????????????, ????? ???????
??????????? (??? 80 ? 90 %) ? 2-3-???????
????????. ??????? ???????? ????????? ????????????? ???????? ?????? (Elytrigia repens,
Phleum pratense, Festuca pratensis, Agrostis
gigantea, A. tenuis, Deschampsia cespitosa,
Calamagrostis epigeios) ? ??????? ???????????? (Cirsium setosum, Artemisia vulgaris,
Sonchus arvensis, Chamaenerion angustifolium,
Angelica sylvestris, Urtica dioica, Picris
hieracioides ? ??). ? ??????? ????????? ??????????? ???????????? ?????? ??????, ?????,
??????????????? Equisetum sylvaticum, ??????? ??????????? Lathyrus pratensis, Trifolium
medium, T. pratense, Vicia cracca, Achillea
millefolium, Pimpinella saxifraga. ? ??????
????????? ?????? Plantago major, Stellaria
graminea, Tussilago farfara, Amoria repens,
Taraxacum officinale, Ranunculus repens,
Potentilla anserina. ??????? ?????? ???????????. ????????? ?????????? ?????????? ??
??????????? ????? ???????????? ?????????
(Chenopodium album, Lepidotheca suaveolens,
Polygonum aviculare, Matricaria recutita,
Trifolium arvense), ? ????? ??????? ???????
????? ??????????? (Equisetum sylvaticum, E.
arvense, Chamaenerion angustifolium, Tussilago
farfara). ??????? ?????????? ??????? ?.?.
?????????? (2006) ?? ??????? ??????? ???
??????????? ?????. ???????????? ????????
?? ?????????????? ??????? ??????? ?? ????
??????? ?????????? ???, ????????? ?????????? ???????? ???? ?????????? ? ???????????? ?. ?????????. ??? ?????????? ???????
??????? ????????????? (? ??????? 47 ?????
?????? 56 ? ?????? ???????) ? ???????????
???? ?????, ?????? ????????????? ?? ???????? ? ?????? ????????. ??? Leucanthemum
vulgare, Prunella vulgaris, Trifolium arvense,
Veronica chamaedrys, Deschampsia cespitosa,
Potentilla anserina, Geum aleppicum, Plantago
urvillei. ????? ??????????? ????? ? ????
?????? ????????? ?? ????????.
??????????
?????????? ??????? ? ????????? ??????
??????????, ????????????? ????? ??????????? ? ???????? ?????? ???????-?????????
???????, ????????? 43 ???? ? ???????
??????????? ???????????. ????? ??? ?????????? ??? ?????? ????? ? ???????? ??
???????? ??????? ?????????? (? ??????,
?????????? ? ???????? ?????) ? ????????
?? ??????????? ? ????? ???????? (? ??????? ?????, ????? ????? ? ????????). ? ?????? ?????? ?????? ?????????? ? ???????
????????????? ??????????, ?????? ??????
???????????? ???????? ??????, ??????????????? ???????. ?? ????? ??????? ??????
??????????? ???????????? ?? ???????????
????; ???????????? ????????, ? ??? ?????
??????????? ???????????, ????????? ???????? ?????????? ?????? ?? ????? ??????? ????????????? ??????????.
? ?????? ? ?????????? ?? ???????? ??????????? ??????????? ???????-????????
?????????? ? ???????????? ?????????????????? ??????. ? ?????????????
??????? ??????????? ????? ?????? ? ??????? ???????????? ???????? ????? ? ?????????? ???????? ???????? ????? ??????????? ? ????????????. ? ???????? ? ???????
????? ???????? ???? ? ?????????? ??????
?????? ??????? ????? ? ???????, ???????
??? ??????????? ????? ??? ???????? ??????. ? ????? ????? ? ???????? ?? ????????
????? ??????????? ??????? ??????? ??????????, ???????????? ?? ??????????
??????????? ????? ?????? ????????????
???? ??????????? ????? ? ??????? ???????
???????????.
# 51 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????? ?????? ????????????? ????? ? ????????? ? ??????? ????? ???????? ??????
?????? ??????????
???????? ?.?. (2002) ????? ???????????? ????? ??????????? ???????. ???????. ????.
????. ????. ????, ???????, 14 ?.
?????? ?.?. (1994) ????????????? ?????? ??????????? ???? ???? ???????? ????? ?. ??????. ?: ????? ?.?. (???.) ?????????? ???????? ?????????????? ???????? ????. ???.: ?????,
?. 276-296
?????????? ?.?. (2006) ???????????? ?????????????? ???????????? ?????????? ??????????? ?????. ?: ????????? ?.?. (???.) ????????????? ?????????? ?????? ? ??????, ???. 2.
????????: ????? «?????», ?. 58-79
??????? ?.?. (1980) ????? ? ?????????????? ????-????????? ?????????????? ??????????
(???????? ???????). ???. ????. 9: 1174-1187
???????????? ?.?. (1979) ????????? ????????????? ????????? ????????????? ???????
?????. ???. ????. 12: 1697-1713
????? ?.?. (1971) ?????????? ???????? ?? ???????? ??????? ??????-?????? ???????????
??????. ???. ????. 3: 347-360
?????????????? ?.?. (1963) ???? ?????????????? ??????????? ? ?????? ??????????????
?????? ??????? ? ??? ????????????. ???. ????. 7: 1015-1021
????????? ?.?. (1985) ???????? ?????????????? ? ??????? ???????? ???????? ??????. ?:
?????? ?.?. (???.) ?????????? ???????? ?????? ? ???????. ?.: ?????, ?. 205-231
?????? ?.?. (2007) ?????????????? ?????????? ? ????????????? ????????????? ???????.
?????: ???-?? ???, 303 ?.
??????? ?.?., ??????? ?.?., ???????? ?.?. ? ??. (2006) ?????? ???? ????????? ??????
? ???????? ????. Arctoa. 15: 1-130
???????? ?.?., ???????? ?.?., ?????? ?.?. (1999) ??????????? ????????????? ???????
?????????????? ????????????? «????????» ? ?????????? ?????????????????? ??????????.
???. ????, ???. ????. 1: 40-48
??????? ?.?. (1974) ??????????????? ???????????? ?? ???????. 1. ?????????????? ???????? ??????? ???? ???????. ???. ????. 11: 1583-1595
????????? ?.?., ????????? ?.?. (2006) ???????????? ????????????? ??????? ?? ??????????? ???????? ????????. ?: ????????? ?.?. (???.) ????????????? ?????????? ?????? ? ??????, ???. 2. ????????: ????? «?????», ?. 47-53
??????? ?.?. (1973) ?????????????? ????????????? ?? ?????? ?????????????? ?????????. ???. ????. 11: 1581-1588
???????????? ???????????? ?? ?????????? ?????? ?????? ????????????? ????? ?????????????????? ???????? (2003). ?. ????????.
?????????? ?.?. (1975) ??????????? ???????? ?????????????? ????????????? ??????? ??
??????? ???????? ?????????? ?????? ???????? ??????. ???. ??? 1: 62-67
????????? ?? ?????????? ??????????? ????????????? ????? (1997) ?.: ??? ??, 61 ?.
??????????? ?.?. (1954) ????????? ??????. ???. ????. 1: 48-57
???????????? ?????? ???????-????????? ??????? (1985) ???????????: ?????, 251 ?.
?????? ?.?. (1979) ???????????? ?????? ?? ???????????? ??????. ???????????: ?????,
190 ?.
# 52 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????? ?????? ????????????? ????? ? ????????? ? ??????? ????? ???????? ??????
???????? ?.?. (1988) ???????????? ?????????? ?????????? ???????? ?????? ? ?????? ??????????? ????????????????? ????????????? (?????????? ?????????). ???. ????. 11: 1577-1583
????????? ?.?. (1995) ?????????? ???????? ?????? ? ???????????? ??????????. ???.: ???
? ?????-95, 991 ?.
??????? ?.?. (2005) ???????????? ?????????????? ??????. ??????: ???-?? ??? ?? ???,
158 ?.
???????? ?.?. (1964) ???????? ? ???????????. ?.: ???-?? ???, 448 ?.
????? ?.?. (1982) ????? ??? ????????? ???????. ???. ????, ???. ????. 4: 3-22
The Vegetation of Roadsides in Taiga
and Tundra Zones of West Siberia
Marina P. Tishchenko
Central Siberian Botanical Garden SB RAS,
101 Zolotodolinskaya, Novosibirsk, 630090 Russia
The natural colonization of roadsides is considered. Studied highway extends through the central and
northern regions of West Siberian plane and crosses taiga and south tundra zones. Species composition
of roadside communities includes 257 species of higher vascular plants, 8 species of mosses and
4 species of lichens. 43 species with high partial activity take the greatest part in colonization of
roadsides. At the northern segment of the road they are wide-spread pioneer species but at southern
segment ? meadow grasses. Along the whole road perennial plants of aboriginal flora are predominate.
Synanthropic plants including ruderal annuals show noticeable activity only at southern segment of
studied road. The vegetation of roadsides is represented by communities of different stages of primary
succession. At the base of floristic similarity characteristic groups of communities for different zones
(subzones) of vegetation were distinguished. Sparse grass-horsetail communities with small number
of species without moss or lichen cover are usual in tundra and forest-tundra. Communities with
well-developed moss cover and constant occurrence of northern shrubs and low shrubs are also
described at the forest-tundra. To the south from forest-tundra starting at north taiga subzone meadow
grasses and legumes increase their occurrence in roadsides colonization. The development of meadow
communities at roadsides in north taiga subzone allows considering these ecotopes as main ways of
meadow vegetation migration north from its natural limits.
Keywords: roadsides, partial activity, vegetation, West Siberian plane.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2011 4) 54-63
~~~
??? 630. 228.7:582.475.4(571.51)
Grafts of Woody Plants and the Problem
of Incompatibility Between Scion
and Rootstock (a review)
Julia A. Darikova*a,
Yulia V. Savva , Eugene A. Vaganova,
Alexi M. Gracheva and Galina V. Kuznetsovab
a
Siberian Federal University,
79 Svobodny, Krasnoyarsk, 660041 Russia
b
V.N. Sukachev Institute of Forestry of the Siberian Branch
of the Russian Academy of Sciences,
50 Akademgorodok, Krasnoyarsk, 660036 Russia 1
a
Received 4.03.2011, received in revised form 11.03.2011, accepted 18.03.2011
Grafting can occur naturally between branches or more often roots of the same species, or
artificially via asexual propagation where the tissues of two plants are encouraged to fuse with one
another. Artificial grafting is commonly used for propagation to improve qualities of trees grown
commercially via selecting for its roots (called the rootstock) and for stem, leaves, flowers or fruits
(called the scion). It is often used to reduce the time to flowering and shorten the breeding program,
improve quality and yield of fruits and seeds, improve hardiness in response to environmental
conditions and pathogens resistance, and change cultivars. Incompatibility between rootstock and
scion has been more often observed for inter-specific than intraspecific grafts, and might result in
a dieback of a graft. The objective of this paper was to review the recent literature related to graft
applications and incompatibility between scion and rootstock. Anatomical studies of incompatible
grafts demonstrated a poor vascular connection, vascular discontinuity and phloem degeneration
at the union area, which might be detected as early as few weeks after a graft establishment. These
vascular fusion problems can disturb water, nutrient and assimilate flows in the plant and might
result in a further breakdown of a union area. Incompatibility might be also a result of insufficient
plasmodesmatal coupling at an early stage of development within one of the graft partners.
Differences in abundance of phenol compounds and starch below and above the graft union, as
well as peroxidase amount at the union might serve as biochemical markers of the rootstock/scion
incompatibility. Genetically and biochemically similar scions and rootstocks could improve graft
compatibility.
Keywords: scion, rootstock, graft, compatibility.
*
1
Corresponding author E-mail address: jualeksa@rambler.ru
© Siberian Federal University. All rights reserved
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Julia A. Darikova, Yulia V. Savva? Grafts of Woody Plants and the Problem of Incompatibility Between Scion?
Introduction
Grafts are often used in agriculture to select
for and maintain desirable traits. Vegetative
propagation such as grafting is preferable to
sexual propagating to ensure reliability and
maintaining desirable quality of yield given
that the sexual propagation may genetically
transfer undesirable traits. Grafting can also be
a desirable method to propagate plants which are
hard to root such as Siberian pine (e.g., Severova,
1958; 1968; Kolegova, 1977). However, a problem
of incompatibility between grafted parts may
occur and result in a further dieback of a plant.
Thus, it is of importance to know mechanisms
of incompatibility between parts of the graft
to mitigate the potential problem even before
planting the grafts or reduce the size of the effect.
Also, recent literature has lots of information
about transfer of desirable and undesirable traits.
Summarizing of this information might be useful
for forest services. Thus, the objective of this
paper was to review the recent literature related
to graft applications and incompatibility between
scion and rootstock.
Types of grafts
Grafting occurs naturally that might involve
branches or more often roots of the same species
(inosculation) (e.g., Janick, 2009), or artificially
via asexual propagation where the tissues of two
plants are encouraged to fuse with one another.
Artificial grafting is often used for propagation to
improve qualities of trees via selecting for its roots
(called the rootstock) and for other characteristics
such as stem, leaves, flowers or fruits (called the
scion).
Natural root grafting happens when roots of
plants make physical contact with each other, while
branch or trunk grafting may occur when crossed
branches become in a close contact in a dense stand
in which bark might become injured by fusion due
to their friction under the air movements (e.g.,
winds) (e.g., Pravdin, Nekrasov, 1953). Although
natural grafting is common between tree of the
same species it has been also observed for trees
of different species but the same genus. Natural
grafting is common for holly, lime, oak, beech,
Scots pine and Jack pine (e.g., Pravdin, Nekrasov,
1953; Janick, 2009). Natural grafting might affect
ecosystems negatively by possible transmission
of tree pathogens (Epstein, 1978) or positively
through sharing water and nutrients via root grafts
(e.g., Bormann, 1966), by providing support for a
weaker tree, and an adaptation to promote fire or
wind resistance (e.g., Loehle, Jones, 1990). A study
demonstrated that root grafts can be less impacted
by hurricane damage compared to non-grafted trees
(Basnet et al., 1993). Another study demonstrated
that grafted shaded trees had significantly higher
root nonstructural carbohydrate concentrations
than non-grafted shaded trees, indicating that root
grafts partially offset the effects of shading (Fraser
et al., 2006). A recent study demonstrated that after
thinning, a growth increment was much less for
trees, which had root grafts with removed trees,
and it was linked to the support of roots and stamps
by live residual trees (Tarroux et al., 2010). Keeley
(1988) hypothesized that root grafts would be of
selective advantage to dominants regardless of the
status of the tree to which it is grafted. For example,
grafts would be of selective value in waterlogged
soils because they would enhance anchorage effect
by dispersing the tree?s compressive force over a
greater surface area.
Grafts can be classified as homografts if the
scion and rootstock are of the same species or
heterografts if they are of different species. Several
studies demonstrated that functioning and growth
of homografts is often more successful compared
to heterografts because of higher similarities in
genetic structure between the graft components
(Kovalishin, 1966; Yakovleva, 1967, 1974, 1975).
Although heterografts more often experience
incompatibility because of the differences in
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Julia A. Darikova, Yulia V. Savva? Grafts of Woody Plants and the Problem of Incompatibility Between Scion?
anatomy, morphology and physiology between the
graft components, they could be more beneficial
than homografts. Heterografts of Pinus sibirica
Du Tour demonstrated a higher growth compared
to homografts (Matveeva and Kichkildeev, 2006).
Grafting of flood-sensitive Annona species onto
A.glabra rootstock improved flood tolerance of
the species (Nuсez-Elisea et al., 1999). Grafting
Fraser fir onto Turkish fir demonstrated reduced
losses by phytophtora root rot in the southern
Appalachian Mountains of North Carolina
(Hibbert-Frey et al., 2006). Fig grafts showed an
improved growth response to nematodes, which
varied depending upon a rootstock (Hosomi et
al., 2002).
Grafts applications
Grafts are often used in agriculture to
reduce the time to flowering and shorten the
breeding program, improve quality and yield of
seeds and fruits, improve hardiness in response
to environmental conditions and pathogens
resistance and change cultivars (Severova, 1958;
Kovalishin, 1968; Browning, Watkins, 1991;
Titov, 1993; 1995; Lahav et al., 1995; Hosomi
et al., 2002; Sanou et al., 2004; Giorgi et al.,
2005; Hibbert-Frey et al., 2006; Ahmed et al.,
2006; Mitani et al., 2008; Cantin et al., 2010;
Gandev, 2009; Franzon et al., 2010; Bellefontaine
et al., 2010; Pio et al., 2010; Souza et al., 2010;
Tandonnet et al., 2010). For example, for Siberian
pine it could take two-three years after grafting
to develop reproductive organs strobills, and
about 30-40 years for a sexually-propagated
plant. Correspondingly, it could take 20 years for
grafts and 170-240 years for sexually-propagated
Siberian pines to reach the maximum seed
production of (Severova, 1958; Beh et al., 2009).
Also, Siberian pine/Scots pine heterografts had a
higher growth and height compared to Siberian
pine homografts (Matveeva, Kichkildeev, 2006).
Another study by Browning and Watkins (1991)
tested several new quince hybrid rootstocks for
pears and demonstrated that they all differed in
their influence on fruit production and quality.
A study of citrus hybrid seedlings grafted
into shiikuwasha (Citrus depressa Hayata)
demonstrated that about one third of the plants
flowered two and a half years after grafting , while
usually it takes almost 20 years from crossing
to cultivar release to the industry (Mitani et al.,
2008).
Several studies also demonstrated effect of
rootstock on branching (e.g., Jayawickrama et
al., 1991; Hirst, Ferree, 1995), crown, size and
chemical content of leaves (Jayawickrama et
al., 1991), fruit size and weight, rind thickness,
juice content, brix, total acids and brix: acid ratio
(Girogi et al., 2005; Ahmed 2006; Georgiou,
2009), and flavonoid content (Gil-Izquierdo
et al., 2004). Grafting is often used to induce
dwarfing to apple trees or other orchards because
it increases productivity by providing more fruits
per unit of land and higher quality of fruits (e.g.,
Sitarek, 2006; Prassinos et al., 2009). One of the
explanations is that when inducing dwarfing in
grafts it results in shift in resource allocation
towards reproductive organs compared to shoot
growth (Webster, 1995).
Rootstock effects on plants have not
always been positive. For example, Cardinal
et al. (2007) demonstrated that yield of Hevea
brasiliensis varied from 41% to 86% depending
upon genetic background of a rootstock. Another
study demonstrated that selection of a rootstock
or scion might suppress yield and growth of
apple trees (Univer et al., 2006). A recent study
suggested that scion might also affect root vigour
of a rootstock which was not related to carbon
supply from the shoot (Tandonnet et al., 2010).
Scion-rootstock interaction
Grafts represent two different genetic
identities functioning as two systems in one
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Julia A. Darikova, Yulia V. Savva? Grafts of Woody Plants and the Problem of Incompatibility Between Scion?
organism influencing each other. Understanding
about mechanisms of the rootstock-scion
interaction is yet limited. Several studies
demonstrated mechanisms of scion-rootstock
interaction on grafts, which were used to induce
dwarfing. One of the hypotheses is that the growth
regulator, auxin, which is transported at different
rates between grafts on different rootstocks,
affects cytokinin production in roots and causes
differences in shoot growth (Lockhard, Schneider
1981; Webster 1998). Another hypothesis assumes
that phenols accumulating at the graft union reduce
tissue viability and the rate of auxin breakdown
(Lockard, Schneider 1981, Treutter, Feucht 1991,
Errea et al., 1994). Finally, the rootstock-induced
dwarfing is caused by reduced soluble transport
across the graft union (Atkinson et al., 2003,
Basile et al., 2003). In addition, Prassinos and
coauthors (2009) showed that rootstock-induced
dwarfing might be caused by differential cessation
of terminal meristem growth and is triggered by
rootstock-specific gene regulation.
Most of studies demonstrated that there is
no horizontal gene transfer between the graft
components, scion and rootstock. However,
a recent study of grafted tobacco plants
demonstrated exchange of genetic information
via either large DNA pieces or entire plastid
genomes (Stegemann, Bock, 2009). In this
study, the analysis of the graft sites revealed
the frequent occurrence of cells harboring
both antibiotic resistances and both fluorescent
reporters. Although this research showed the
exchange of genetic material between grafted
components, the gene transfer was only
restricted to the contact zone between the grafted
components, which indicated that the changes
could become heritable only via lateral shoot
formation from the graft site. Previously, there
was a study that described hereditary changes in
grafted pepper, and suggested that these changes
might be induced by a mechanism similar to that
of transformation in higher organisms (Ohta, Van
Chuong, 1975). Yet, there are unresolved issues
of whether gene transfer depends on the cell
contacts between scion and rootstock. Although
micromolecules can go through plasmodesmata,
it is unknown if plastids or macromolecules
of DNA can go through the plasmodesmata
considering that their anatomical structure can
change due to specific proteins.
Incompatibility between scion
and rootstock
Incompatibility between rootstock and
scion has been more often observed for interspecific than intra-specific grafts, which might
further result in a dieback of a graft. Anatomical
studies of incompatible grafts demonstrated a
poor vascular connection, vascular discontinuity
and phloem degeneration at the union area,
which might be detected as early as few weeks
after a graft establishment (Pina, Errea, 2005;
Pina et al., 2009). This results in a disturbance
of water and nutrients transfer by xylem in one
direction and photoassimilates by phloem in
another direction. Plasmodesmata are important
communication pathways in plants (e.g, Pina et
al., 2009). These are channels in the plant cell wall
allowing passage of macromolecules and solutes
between the neighboring cells. A recent study
demonstrated that insufficient plasmodesmatal
coupling at an early stage of development within
one of the graft partners might result in graft
incompatibility (Pina et al., 2009).
At the graft connection area a callus forms as
a reaction to wounding. Another study of apricot
grafted onto a plum rootstock demonstrated that
only some callus differentiation into cambium
and vascular tissues occurred; while a large
portion of the callus never differentiated (Pina,
Errea, 2005). This interrupts the vascular
connections because of the lack of differentiation
that brings discontinuities in the cambium and
# 57 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Julia A. Darikova, Yulia V. Savva? Grafts of Woody Plants and the Problem of Incompatibility Between Scion?
the formation of a band of parenchymatous cells.
In a study of pear grafts typical symptoms of
graft incompatibility such as bark discontinuity,
cambial dysfunction and accumulation of
starch in the scion were observed. Little cell
necrosis was observed at the interface of
incompatible grafts at the 5-month stage of graft
development (Ermel et al., 1999). In spite of a
lack of scion/stock vascular connections, some
plants demonstrated that they could flower and
produce seeds, i.e., there were nonvascular
functional connections between the two plants,
probably of parenchyma cells (Flaishman ?t al.,
2008). Abundance of periderm rings was also
a symptom of incompatibility according to a
study by Ermel et al. (1999), which were more
abandoned in incompatible grafts. In some cases,
signs of incompatibility have been revealed a
few years after grafting and were explained by
differences in radial growth between scion and
stock that breaks tissue connectivity at the graft
union (Savva et al., 2004).
In a study of peach/plum grafts it has been
demonstrated that free amino acids per tree
decreased in rootstock of incompatible grafts
while they became stabilized in compatible grafts
between 79-89 days after grafting (Moreno et al.,
1994). Another study of peach/plum grafts by
Moing and Gaudillere (1992) demonstrated that
sorbitol was lower in roots of incompatible grafts
compared to compatible ones, and that sugars and
starch accumulated in scions of the incompatible
grafts following 78 days after grafting, but
carbon partitioning was only slightly affected
by graft incompatibility. They also showed
that nitrogen assimilation ceased 78 days after
grafting in incompatible grafts, which was not
the case in compatible grafts. This suggested
an inhibited transport of solubles from roots to
scion in rootstocks and from photosynthetic
apparatus to rootstock. A study of pear/quince
grafts demonstrated that 63kDa protein might
be responsible for incompatibility because it was
observed in compatible grafts, faintly observed
in intermediate compatible grafts and was not
detected in incompatible grafts (Gulen et al.,
2005), Absence of peroxidase A in pear/quince
grafts has been also related to graft incompatibility
(Hossein et al., 2008).
Graft incompatibility can be often
determined at an early stage of graft development.
For example, in a study of apricot grafts
accumulation of phenol compounds could serve
as an indicator of incompatibility as early as
the fi rst week after grafting (Errea et al., 2001).
Another study also indicated that accumulation
of phenol deposits at the place of graft union
might inhibit graft compatibility (Mng?omba
et al., 2008). Another study also demonstrated
that differences in phenol accumulation below
and above the graft union might serve as an
indicator of compatibility (Usenik et al., 2006).
Additionally, a study of Prunus persicu/Prunus
spp. grafts indicated that significant increase
in peroxidase activity might be observed in
incompatible unions (Zarrouk, 2010).
Conclusion
Our literature review demonstrated that
there are varieties of grafts which might inherent
desirable or undesirable properties from their
graft components. Incompatibility is a complex
problem that might occur between graft plants
and result in their further dieback. Here, we
reviewed and discussed several mechanisms
of graft incompatibility, and concluded that
compatibility might be improved by selection of
the types of rootstock and scion that might: (1)
favor callus differentiation into cambium and
vascular tissues; (2) be able to establish a close cell
contact at the graft union to maintain water and
nutrient balance between scion and rootstock; and
(3) enable synchronicity in radial growth and cell
differentiation between scion and rootstock. Graft
# 58 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Julia A. Darikova, Yulia V. Savva? Grafts of Woody Plants and the Problem of Incompatibility Between Scion?
components with similar genetic backgrounds
might be incompatible less often compared to
inter-specific grafts. Biochemical markers such
as abundance of phenol compounds and starch at
the union, below and above the union and specific
polypeptide, peroxidase activity might be helpful
to diagnose incompatibility at an earlier stage of
graft development.
Acknowledgements
The authors would like to acknowledge
the support provided to the 2010 winners of
the Russian Federation government support to
the leading scientific schools (grant ???65610.2010.4). We are also grateful to the grant
No. 53 of the Siberian Branch of the Russian
Academy of Sciences for partial support.
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???????? ????????? ????????
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?.?. ?????????, ?.?. ??????, ?.?. ????????,
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??????????????? ???????? ????? ? ?????????. ?? ???? ????? ???????? ? ????????????????
(?????????????) ? ?????????????????? (??????????) ? ????????, ????????? ? ?????????????????????? ????????????????, ???? ????? ????????? ? ?????????. ????? ?????? ??????
???????? ????? ??????????? ??????????, ????????? ? ??????????? ?????????? ? ?????????
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??????? ? ???????. ??????????? ??????????????? ?????? ? ?????? ???????? ???? ????????
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Journal of Siberian Federal University. Biology 1 (2011 4) 64-74
~~~
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????????????? ???????????
???????????? ??????? ?????????? ????????
NADH:FMN-???????????????-??????????
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?
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Received 4.03.2011, received in revised form 11.03.2011, accepted 18.03.2011
??????????? ??????? ???????????? ???????? ? ???? ?? ???????????????? ????????????
??????? ?????????? ???????? NADH:FMN-???????????????-??????????, ???????
????????????????? ? ????????????? ??????????? ???????????? ??????? ? ???????????
?????????. ?????????? ???????????? ??????? ? ???????? ?? ???????????????? ?
???????? ?????? ????????????, ????????????? ??? ???????????? ????, ? ? ????????
??????? ????????????, ????????? ? ??????????? ????, ???????????. ???????????, ???
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??????????????? ????????. ????????? ????????? ? ?????? ?????, ????????? ???????
?????????? ????????, ?? ?????? ?? ???????? ?????????????? ???????.
???????? ?????: ????????????? ??????????, NADH:FMN-???????????????, ????????????????,
???????????????? ?????????, ???????????? ?????????, ???????.
????????
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?????????: ?????????? ? NADH:FMN???????????????. ?????????? ????????????
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Corresponding author E-mail address: aebezrukih@gmail.com
© Siberian Federal University. All rights reserved
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?.?. ????????, ?.?. ??????????? ????????????? ??????????? ???????????? ??????? ?????????? ?????????
FMNH2 + RCHO + O2 ?
? FMN + RCOOH + H2O + h? .
(1)
?????, ????????? ??? ?????? ??????????????? ????? ??????????, ??????? ??????,
??? ????? ????? ????????? FMNH2, ??????? ????????????????? ?????????? ?????????? ????? ??? ?? 1 ?. (????????? ? ??., 1984).
NADH:FMN-??????????????? ????????????
???????
NADH + FMN + H+ ?
? NAD+ + FMNH2 .
(2)
? ??????????? ??????????????? ?????
???????? ?????????? ?????????????, ??? ????????? ????????? ?????????? ???????? (????????? ? ??., 1984).
? ??????? ???????? ???????????? ??????? ?????????? ???????? ????????????, ??????? ???????, ??? ?????????? ?????????????
??????? ??????? ????????? ??????????? ?
?????????? ????????? (Kricka et al., 1983;
Brolin et al., 1992; Roda et al., 2004), ? ????? ?
???????? ????-??????? ??? ??????????? ???????????? ??????????? ????????? ???? (???????? ? ??., 1987; Kudryasheva et al., 1998).
????????? ???????? ?????????? ?????????????? ??????? ? ????????????? ????????????????? ????????? ??????? ????????
?????????????? ??????, ?????????????? ?
????????? ??????? ??????????? (Kratasyuk
et al., 2001; Vetrova et al., 2007). ?????? ???????????? ???????????? ??????? ??????????
???????? ?????????????? ?? ?????????? ??????????????? ? ??????? ???????????? ???
??????????? ??????????????? ????????, ???????? ???????????, pH ? ?.?. (???????? ?
??., 1982).
????? ?? ???????? ???????? ?????????
?????????? ????????? ???????? ?? ????????????? ?? ??????? ???? ????????? (????-
??? ? ??., 1987). ????????????? ?????????
???????? ? ????????????? ????????? ??
??????-?????????? ??????????: ?????????????????, ??-????????, ????????????????
? ???????? ????? ????? ? ?.?. (?????????? ?
??., 2009). ? ????????? ??????? ??? ????????? ???????????? ????????? ?????????????
???????????????? ?????????? ?????????
(Kratasyuk et al., 2003; Esimbekova et al., 2007;
?????????? ? ??., 2009). ?????? ? ??????????? ??????? ?????????? ???????????? ?????? ?????????? ????????? ? ?????????? ??
??????????? ??? ?????????? ????????? ????????????? (??????? ? ??., 1987). ?????????
???????? ??????????? ?????? ????????????? ????????? ?, ??? ?????, ???????????
??????? ?????? ?????????? ????????? ???
?????????? ????????????? ????????? ???????? ?????????? ??????????? ?????????
(???????? ? ??., 1986; ?????????? ? ??., 2000;
?????????? ? ??., 2000).
? ????????? ????? ??????????? ????? 30
????????? ???????? ????????????? ?????????, ?????????? ?? ?????????? ??????????
(Kratasyuk et al., 2003). ???????? ?????????????? ?????????? ??? ????????? ???????????????? ?????????? ????????? ?????????? ?????????? ??? ?????? ??????????????
??????????? ???????? ??????????? ?????????????? ???? ? ???? ?? ?????? ????????
(Kratasyuk et al., 2003; Esimbekova et al., 2007;
?????????? ? ??., 2009). ?????? ????????????? ?????? ????????? ????????? ? ????
???????? ???????????? ???????????? ?????????, ????????? ?????? ???? ???????????, ???
?? ????????????? ?? ?????????? ??????????? ?????????, ?????????????? ??????????
????????? ??????????? (?????????? ? ??.,
2009).
????? ?????? ?????? ???? ????????????
??????? ????????, ????????????? ? ????????
???????? ??? ????????????? ?????????, ??
# 65 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????????? ????????????? ??????????? ???????????? ??????? ?????????? ?????????
???????????????? ???????????? ???????
NADH:FMN-???????????????-??????????,
? ????? ???????????? ????????????????? ?
???????????????? ???????????? ??????? ?
??????????? ?????????.
????????? ? ??????
? ?????? ???????????? ????????????????? ????????? ??????????????? ????????? (???????? ????????? ?????????????
????????????????, ????), ?????????????
? ??????????? ????????????????? ? ????????????? ???????????????? ????????? ????????? ?? ??? (??????????). ????
?????? ???????????????? ????????? ???????? 0,5 ?? ?????????? (L) EC 1.14.14.3
?? ??????????????? ?????? Escherichia
coli (P. leiognathi) ? 0,15 ??. ??????????
NADH:FMN-??????????????? (R) EC 1.5.1.29
(Vibrio fi scheri). ????? ? ?????? ????????????
???????? ??????????????? ????????? ?????????? ? NADH:FMN-???????????????,
?????????? ????? ?? ?????????? ????????,
??? ? ? ????????? ????. ?? ?????? ? ?????????? ????????? ??????? 9 ?? 0,05 ?
???????????????? ??????, ??????? ???????
?? ???? ? ???????????? ? ??????? 8 ? ?????
?????????????.
?? ????????? ????????? ????????:
NADH ? FMN («Serva», ????????), ????????????? («Merck», ????????), ??????? («Fluka»,
????????). ??? ????????????? ????????? ???????????? 0,05 ? ?????-????????? ?????
pH 6,8. ??????? 0,0025 %-?? ?????????????
???????? ??????????? ? 5 ?? 0,05 ? ??????
50 ??? 0,25 %-?? ?????????? ???????? ?????????.
??????????? ???? ???????? ?????????
???????. ????? ??????? ???????? ????????
???????? ????????? ? ????????? ???????? ?
??????? 30 ???. ?????, ????????? ?????????,
????????? ?? ??????????? 80 °?.
??????? ????????????? ?? ????? ????????? ??????????? ??? ????????? ???????????; ??????????? ??????????? ?????????
NADH, FMN ? ???????? ? ???? ?????? ?????????????? ??? ?????? ???????????????
?????????? VT-8 (Termex, ??????).
?????????? ???????????? ??????????? ??????? NADH:FMN-????????????????????????? ???????? ? ??????????? ????? ?????????? ???????: 500 ??? ????????
????????; 5 ??? ???????? ?????; 50 ???
0,0025 %-?? ???????? ?????????????; 10 ???
0,5 ?? ???????? FMN; 100 ??? 0,1 ?? ???????? NADH.
? ?????? ?????????????? ??????? ??????????????? ??? ?????????? ???????????
?????, ?????? ????????????, ????????
?????? ? ????????????? ? ?????????????? ???????? ???????????? ?????????????
???????? (Imax). ??? ????????? ????????? ??
?????????????? Lumat LB 9507 («Berthold
Technologies», ????????).
?????????? ???????????? ???????
??????? ? ??????????? ???????? ??????
???????????? (0,5, 1, 1,5, 5 %) ? ??? ????
(????????) ? ????????? ?????????? ?? 10 ??
50 є?. ???????????????? ???????????? ??????? ???????, ????????? ???????? ? ??????? 0,5-30 ??? ??? ?????? ???????????? ?
????????? ?? 10 ?? 50 є? ? 0,5, 1 ? 5 %-? ????????? ???????? (????????? 180 ????) ? ???
???? (????????), ? ????? ??? ?=33 є? ? 1 %-?
????????? ????????? ?????? ????????? (60
? 250 ????).
???????? ????????????????? ?????????
NADH, ????????? NADH ???????????????? ?
?????????? ?????????? ? ??? ?? ??????????
(? ??????? ????????? ????), ? ????? ????????
????????? NADH ?????????? ?????????? ?
??????????? 1 %-?? ???????? ???????? ? ???
??? ?????????? ?????????? ?? ????????? ??
??????? ?????????? ?????????, ??????????
# 66 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????????? ????????????? ??????????? ???????????? ??????? ?????????? ?????????
?? ????? ????? ????????????? ??????????
NADH (?=340 ??). ?????????? ?????????
???????? ?? ???????????????? UVIKON 943
(«Kontron Instruments», ??????). ????????
???????????? NADH ???????????? ?? ???????
?
?
?
? ? ?–
(3)
??? ? ? ?????????? ????????? ????????, ? ?
??????????? ?????????? NADH ??? ?=340
??, t ? ?????.
??? ????????? ? ?????? ???????????????? ??????????????? ??????? 350 ??? ???????? ????????, 10 ??? ???????? ????????,
50 ??? 0,0025 %-?? ???????? ?????????????,
10 ??? 0,5 ?? ???????? FMN, 50 ??? 0,8 ??
???????? NADH. ???????? NADH, FMN ? ???????? ?? ????? ???????????? ??????????? ?
?????????? ??? ??????????? 33 є?.
??????????? ?????? ?????????????????
????? ????????? ?? ????? ??? ? 3-5 ???????????? ??????????.
?????????? ? ??????????
?? ???. 1 ???????????? ???????????
????????????? ???????? ???????????? ??????? ?????????? ???????? ?? ???????????
? ??????????? ? ??????????? ????? ????????? ???????????? ????????. ??? ??????
????????????? ???????? (0,5 %) ??????????
???????????? ??????? ????????? ?? ????
????????? ????????????? ??????????. ???
?????????? ???????????? ???????? (1 % ?
?????) ??????????? ??????? ?????????????
???????? ??? ???????????, ??????? ??????????? ??????????????? ? < T?. ????????????
?????????? ????????????? ??????????? ???
???? ? ??????????? ?????? ?? ???????? ?
??????????? ?????? ?????????? ?????????
??? ?>??. ????? ????, ????????? ????????????? ???????? ??? ? ? ?? ?????????? ?????
?????????? ??? ?????????? ????????????
????????.
????????
????????????
??????
???27 °? (??????, 1980). ??????? ????????????? ???????? ??????????? ?????? ??????
???? ???????????. ????? ???????, ?????? ?????, ?????? ????????? ????????????? ????????
??????????? ?????????? ??????? ?????????
????????. ???????????? ???? ????????????????? ?????????? ? ???????? ??????????????? ????? ???? ?????? ? ????????????? ?????????????? ????????? ? ??????? ? ????????
????????????? ?????? ?????????????????
??????? ? ???????? ?????????? ????????????? ?????????? ???????? ?????. ????????
?????, ??? ????? ?????????? ????????? ?
??????????? ?????? ????????, ???????????? ??????? ???????, ?????????? ?????,
??????????????? ???????? ? ?????????????? ?? ???????????.
??? ??????????? ???? ??????????? ??????????????? ???????? ???????? ???????
????????, ????????, ???????????? ??????????? ?????????. ???????? ?????????????
???????? ? ???????? ???????? ?????? ???????????? ?? ???? ????????? ?????????????
?????????? ??????? ? ???, ??? ????????????
???????? 0,5 % ???????? ????????????? ???
??????????? ????.
??????????, ??? ???????? ???????????????? ????????????? ???????????? ???????
NADH:FMN-???????????????-??????????
??? ? ??????????? ? ??????????? ?????
0,5 %-?? ????????, ??? ? ? ??? ??????????
????? ????????????? ?????????? ????????
?? ???? ????????? ?????????? (???. 2).
??? ??????????? 30-40 є? ???????????
???????????????? ??????? ???????, ?????????? ??? ????????? ????????? ??????????? ???????????? ???????, ???????????????
????????? ???? ????? ? ??????????? ? ??????? ??????????.
# 67 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
???. 1. ??????????? ????????????? ???????? ???????????? ??????? ?? ??????????? ? ???????????
????????? ???????? ?????? ???????????? (2) ? ??? ???????? (1): ? ? 0,5 %-? ??????? ????????; ? ? 1 %-?
??????? ????????; ? ? 1,5 %-? ??????? ????????; ? ? 5 %-? ??????? ????????
???. 2. ???????????? ?????? ???????????????? ???????????? ??????? NADH:FMN-????????????????????????? ??? ?????? ????????? ??????????? (1 ? 15 є?, 2 ? 20 є?, 3 ? 25 є?, 4 ? 30 є?, 5 ? 35 є?, 6 ? 40 є?)
? ??????????? 0,5 %-?? ???????? (?) ? ??? ?????????? (?)
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????????? ????????????? ??????????? ???????????? ??????? ?????????? ?????????
?????????????
??????????
?
NADH:FMN-???????????????
????????
???????????? ?????????? ? ???????????? ????? ??????????? ? ????????? ?????????????? (Fisher et al., 1996; Koike et al.,
1998). ?????????????, ?????? ????????????? ?????????, ???????????? ???
??????????????? ???????????? ???????,
??-????????, ???????? ??????????? ????????? ?? ???????????. ???????? ??????????? ????????????? ?????? «????????»
??????? ???????????? ?????? ????????????????. ????? ????? ??????????? ?????????? ? (???) NADH:FMN-???????????????
?? ??????????? ??? ????????? ??????????? ??????? ??????????? ???????????
?????????, ??????? ????????????? ??????
«????????» ??????? ???????????? ??????
???????????????? ???????????? ???????
(???. 2).
????? ???????, ???????????????? ???????????? ??????? ?????????? ????????
?????????? ?? ??????????????? ?????????,
???????????? ??? ??????????? ???????????
??????. ???, ????????, ??????????? ??????????? ?????????? ????? ???????? ? ????
?????? ??????????? ? ??????????? ??????????? (???????? ? ??., 1986; ?????????? ? ??.,
2000; ?????????? ? ??., 2000).
??? ??????????? 15-25 є? ? ???????????
0,5 %-?? ???????? ? ? ??? ?????????? ??????????? ????????? ??????????? ???????????? ??????? ?? ??????? ???????, ????????? ?
????????? ??????????? ????????? ?? ???????????. ??????????? ??? ?????? ??????????? ? ??????????? ????????? ????????? ??
???????????.
?? ???. 2 ?????, ??? ?????? ???? ????????????????, ??????????????? ???????????
?????????, ? ???????? ?????????? ? ??????? ?????????, ??? ??? ????, ??? ???? ????????????? ????????????. ??? ???? ???????
???????? ??????????? ?? ?????? ?? ????????
???????????.
???????? ???????????????? ???????????? ??????? ? 1 %-? ???????? ???????? ????????, ??? ???????? ???????????????? ? ??????????? ? ??????????? ????? ???????? ? ???
???? ????? ?????????? ???????? ?? ?? ????
????????? ??????????, ??? ? ?????? 0,5 % ????????, ? ?????? ? ??? ??????, ????? ? > ??.
??? ??????????? 23-38 є? ???????????? ?????? ??????????? ? ???????????????????
??????????? ??????? ?? ???? «????????»
????????, ??????????????? ????????? ??????????? ? ??????????? (???. 3). ??? ? > 43 є?
??????????? ??????? ??????????? ???????
???????. ??? ??????????? ???, ??? ???????????? ?????? ???????????????? ??????????
???????????????? ????????????? ??-?? ??????? ?????????? ? ?????? ??????????? ???????? ??????????? ?????????.
??? ??????? ?????? ???????????, ?????????? ??? 5 %-?? ????????, ???? ????????, ??? ???????? ???????????????? ? ??????????? ? ??????????? ????? ???????? ?
??? ????, ??? ?? ??? ? ??? 1 %-?? ????????,
????? ??????? ??? ?????? ??? ? > ??. ???
? = 28-38 є? ??????????? ????????????????
??????? ???????, ? ??? ? > 43 є? ? ??????? ??????? (???. 4).
??? ????????????, ??????? ??????????? ??????????????? ???????? (? < ??), ? ??????????? 1 %-?? ? 5 %-?? ???????? ??????????
???????????? ????????? ???????????? ??????? ? ???????? ???????, ? ?? ????? ??? ? ?????????? ???????? ???? ????????? ???????????
?? ??????? ???????. ?????? ????? ???????????? ?????? ??? 1 %-?? ???????? ???????? ??
???. 5. ???? ????????? ????????? ???????????? ???? ???????????? ????????? ??? ?? ????????? ? ??????? ??????? ????????.
????? ????, ???? ????????, ??? ??? ????????? ???????????? ???????, ????????-
# 69 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
???. 3. ???????????? ?????? ???????????????? ???????????? ??????? NADH:FMN-????????????????????????? ??? ?????? ????????? ??????????? (1 ? 23 є?, 2 ? 28 є?, 3 ? 38 є?, 4 ? 43 є?) ? ???????????
1 %-?? ???????? (?) ? ? ??? ?????????? (?)
???. 4. ???????????? ?????? ???????????????? ???????????? ??????? NADH:FMN-????????????????????????? ??? ?????? ????????? ??????????? (1 ? 33 є?, 2 ? 38 є?, 3 ? 43 є?) ? ??????????? 5 %-??
???????? (?) ? ? ??? ?????????? (?)
???. 5. ??????????? ????????????? ???????? ???????????? ??????? ?? ??????? ???????????? ?
??????????? 1 %-?? ???????? (2) ? ??? ???? (????????) (1) ??? 10 є? ? ??????????????????? ???????????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????????? ????????????? ??????????? ???????????? ??????? ?????????? ?????????
???. 6. ??????????? ??????????? ?????????????? ???????? ???????????? ??????? ? ?????????
???????? ?????? ????????? (1 ? 60 ????, 2 ? 180 ????, 3 ? 250 ????) ? ??????????? ???????? (?????) ???
???????????? ??????? ????????? ?????????, ??? 33 o?
??? ? ??????????? ???????? ??? ???????
??????????? (35-50 є?) ? ??????? ??????????
?????????? ??????? (30 ? ? 1 ???), ? ???????
????????? ??????????? (? 20 є?) ??????????
?? ???????????: ????????, ????? 20 ??? ???????????? ??? ????????? ???????????
?????????? ???????????? ???????, ?????????????? ??????????? ? ??????? 1 ??? ???
48 є?, ?????????? ? 9 ???. ? ?????????? ???????? ????? ?????? ???? ???????????, ?????? ?
??????? ???????: ??? ??????????? ????????
? ??????? 20 ??? ???????????? ??? ????????? ??????????? ?????????? ????????????
??????? ?????????? ? 5 ???. ????? ???????,
??????? ???????????????? ????????????
??????? ???????? ???????, ??????? ???????? ???????????? ?????????????? ??????????
?????????.
????????? ???????????? ?????? ???????????????? ? ??????????? ????????? ??????
?????????, ?????????? ? ??????, ????????,
??? ?????????????? ???????? ???????????? ????????????? ???????????? ???????
? ????????? ????????? ????????? ???. ?????
????????, ???, ???????? ?? ???????? ??????????? ???????? ?????????? ????????????
??????? ? ????????? ????????, ?????????
?????????? ? ??????????? ???????? ? ?????-
????? ??? ???? ?????? ? ??????????? ???????
????????????? ????????? ??? ?????? ??????????? (???. 6).
????????? ?????? ????????????? ???????????????? ?????????? ? NADH:FMN??????????????? ? ???????? ??????????
????????, ??????????? ??????????? ?????????. ?? ?????????? ? ???????????? ????????
???????? ????????? ?????????? ????????? ??
??????? (3) ???? ?????????? ???????? ??????????????? ? ????????????????? ????????? NADH (????. 1). ???????? ??? ????????????????? ????????? NADH, ??? ? ?????????
NADH ???????????????? (? ??? ????? ? ? ??????? ????????? ????) ? ??????????? ???????? ?? ?????????? ?? ????????? ????????? ?
???????? ????????.
????? ???????, ? ?????? ????????, ??? ?????????? ???????????? ??????? NAD?:FMN???????????????-?????????? ? ???????? ??
???????????????? ? ????????, ?? ?????????? ????, (????) ? ? ???????????? ????????,
?????????? ??? ??????????? ??? ???????????? ???? ????????????? ???????? ? ??? ?????? ????????????, ???????????. ? ?? ?????
??? ????????? ????????? ? ???? ????????
???????? ? ???????? ????????????? ????????????????, ????????? ????????? ? ????-
# 71 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????????? ????????????? ??????????? ???????????? ??????? ?????????? ?????????
??????? 1. ???????? ??????????????? ? ????????????????? ????????? NADH ? ??????????? 1 %
???????? ? ??? ????
???????
V, ??????/???
?????
???????
??? ?????????
0,6 ± 0,2
0,8 ± 0,2
??????????
1,26 ± 0,3
0,93 ± 0,04
???????????????
2,6 ± 0,6
2,3 ± 0,3
????
2,5 ± 0,3
2,7 ± 0,2
??? ??????? ???????? ???????????? ?? ???????????? ? ?????????? ?????? ??????????,
??? ???? ????? ??? ???????????? ???????????
???????? ?????? ????????????????? ????????
??? ?????????????????? ??????? ?? ??????
???????????????? ? ??????????? ???? ????????? ?????????? ????????. ???????????????? ???????????? ??????? ????? ??????????
???????? ? ????????? ?? ??????????????? ??-
???????. ???????, ??????????? ? ???? ????,
???????? ?????? ???? ????????????????, ? ???????????? ???????, ????????, ????????????
????????? ???????????? ???????. ? ??????????? ???????? ???????? ??????????? ???????????? ??????? ??? ????????? ???????????.
????????? ????????? ? ?????? ?????, ????????? ??????? ?????????? ????????, ?? ??????
?? ???????? ?????????????? ???????.
?????? ????????? ??? ?????????? ????????? ???????????? ????????? ?? ????? ? ?????????? (??? «??????-?????????????? ????? ????????????? ??????», ??????????????? ???????? ? 02.740.11.0766), ???? (????? 07-04-01340-?), ???????????? ??????????? ? ?????
?????????? ????????? ? ????????????? ????? ??????????? ???????????? ? ???????? (??????
? 2.2.2.2/5309 ? RUX0-002-KR-06/BP4M02), ?????????? ???????? ???? (????????? «???????????? ? ????????? ????????»), ? ????? ?????????? ?? (????? ?? 64987.2010.4).
?????? ??????????
??????? ?.?., ?????? ?.?., ??????? ?.?. ? ??. (1987) ?????????????: ????. ??????? ???
?????: ? 8 ??. / ??? ???. ?.?. ???????, ?.?. ?????????. ?.: ?????? ?????, 159 ?. ??. 7: ???????????????? ????????.
????????? ?.?., ???????? ?.?., ????????? ?.?. ? ??. (1984) ?????????? ????????. ???????????: ?????, 278 ?.
?????????? ?.?., ????????? ?.?., ??????? ?.?. (2000) ??????????? ?????????? ????????? Luciola mingrelica ???????????????? ? ?? ???????????. ?????. ????. ??-??. ???. 2. ?????.
41: 366-370.
?????? ?.?. (1980) ?????? ???????????????? ????????. ?.: ?????, 672 ?.
?????????? ?.?., ????????? ?.?., ???????? ?.?. (2009) ????????? ???????????????? ?
??????????? ???????????? ??????? ????:???-???????????????-??????????. ????????
74: 853-859.
???????? ?.?., ????????? ?.?. (1987) ????????????? ?????????? ???????? ? ????????????????? ???????. ?????? ????????????? 21: 3-30.
# 72 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????????? ????????????? ??????????? ???????????? ??????? ?????????? ?????????
?????????? ?.?., ?????????? ?.?., ??????? ?.?. (2000) ???????? ? ???????? ???????????????? ?????????????? ?????????? ????????? Luciola mingrelica. ?????. ????. ??-??. ???.
2. ?????. 41: 362-366.
???????? ?.?., ???????? ?.?., ???????? ?.?., ???????? ?.?. (1982) ????????????????
????????????? ??????????. ???????? 47: 1773?1777.
???????? ?.?., ?????? ?.?. (1986) ?????????????? ???????????????? ????????????????.
????? ????? ? ???????. ???. ?????????????, ?. 5. ??????, ?., ?. 50-86.
Brolin S., Wettermark G. (1992) Bioluminescence Analysis. Weinheim, Germany: VCH, 151 p.
Esimbekova E.N., Kratasyuk V.A., Torgashina I.G. (2007) Disk-shaped immobilized
multicomponent reagent for bioluminescent analyses: correlation between activity and composition.
Enzyme and microbial technology 40: 343-346.
Fisher A.J., Thompson T.B., Thoden J.B., Baldwin T.O., Rayment I. (1996) The 1.5-A resolution
crystal structure of bacterial luciferase in low salt conditions. J. Biol. Chem. 271: 21956-21968.
Koike H., Sasaki H., Kobori T., Zenno S., Saigo K., Murphy M.E., Adman E.T., Tanokura M.
(1998) 1.8 A crystal structure of the major NAD(P)H:FMN oxidoreductase of bioluminescent bacterium
Vibrio fischeri: overall structure, cofactor and substrate-analog binding, and comparison with related
flavoproteins. J. Mol. Biol. 280: 259-273.
Kratasyuk, V.A., Esimbekova E.N., Gladyshev M.I., Khromichek E.B., Kuznetsov A.M., Ivanova
E.A. (2001) The use of bioluminescent biotests for study of natural and laboratory aquatic ecosystems.
Chemosphere 42: 909-915.
Kratasyuk V.A., Esimbekova E.N. (2003) Polymer Immobilized Bioluminescent Systems for
Biosensors and Bioinvestigations. In: Arshady R. (ed.) Polymeric Biomaterials, The PBM Series, vol.
1. Citus Books, London, p. 301-343.
Kricka L., Wienhausen G., Hinkley J., DeLuca M. (1983) Automated bioluminescent assay for
NADH, glucose-6-phosphate, primary bile acids, and ATP. Anal. Biochem. 129: 392?397.
Kudryasheva N.S., Kratasyuk V.A., Esimbekova E.N., Vetrova E.V., Nemtseva E.V., Kudinova I.Y.
(1998) Development of the bioluminescent bioindicators for analyses of environmental pollutions. J.
Field Analytical Chemistry and Technology 2: 277-280.
Roda A., Pasini P., Mirasoli M., Michelini E., Guardigli M. (2004) Biotechnological applications
of bioluminescence and chemiluminescence. Trends Biotechnol. 22: 295-303.
Vetrova E., Esimbekova E., Remmel N., Kotova S., Beloskov N., Kratasyuk V., Gitelson I. (2007)
A bioluminescent signal system: detection of chemical toxicants in water. Luminescence 22: 206 ?
214.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????????? ????????????? ??????????? ???????????? ??????? ?????????? ?????????
Thermoinactivation
of Coupled Enzyme System of Luminous Bacteria
NADH:FMN-Oxidoreductase-Luciferase in Gelatin
Anna E. Bezrukikha,
Elena N. Esimbekova and Valentina A. Kratasyuka,b
a
Siberian Federal University,
41 Svobodny, Krasnoyarsk, 660041 Russia
b
Institute of Biophysics,
50/50 Akademgorodok, Krasnoyarsk, 660036 Russia
b,a
Influence of gelatin solution and gel on bioluminescence of coupled enzymatic system of luminous
bacteria NADH:FMN-oxidoreductase-luciferase was analyzed, thermostability and thermoinactivation
of coupled enzyme system in gelatinous environment were studied. Activity of coupled enzyme system
and kinetics of its thermoinactivation in gelatin with low concentration, that is insufficient for gel
formation, and in gelatin with high concentration, that is sufficient for gel formation, was different. It
was determined that thermoinactivation of coupled enzyme system has nonlinear nature and occurs
with dissociative mechanism. It was showed that gel matrix of gelatin stabilizes enzymes. There was no
effect of viscous environment originating by gelatin surroundings on the rate of enzymatic reactions.
Keywords: bacterial luciferase, NADH:FMN-oxidoreductase, bioluminescence, thermoinactivation of
enzymes, stabilization of enzymes, gelatin.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2011 4) 75-85
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??? 504.064.37
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???????? ???? ??. ?.?. ???????? ?? ???,
?????? 660036, ??????????, ?????????????, 50/28 1
Received 4.03.2011, received in revised form 11.03.2011, accepted 18.03.2011
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??????????? ??????? ???????. ?????????? ?????????? ???????????? ????????? ? ????????
??????????????, ? 1960 ?. ?????????? ?? ???? ???????? ????????, ????????????? ? ????????
? ????????? ?????. ??????????? ?????? (NOAA/AVHRR and TERRA/Modis imagery) ? ??????
???????????? ???????? ?? 1996-2009 ??. ?????????????? ????????? ?????? ?????? ? ????????
???? ???????? ? ????????? ?? ????????????? ? ????????? ???????????? ????????????? ?
??????? ??????? ?????????. ????????? ?????????????? ?????? ?????? ???????-???????
?????????? ? ???????????? ????????? ???????? ????????????????? ????? ??? ???????????
???????????? ?????????? ?????? ? ??????????? ?? ???????????? ? ????????? ??????.
???????? ?????: ?????? ?????????? ????, ???????-??????? ?????????, ?????? ??????????????
????????????, ???? ??????, ???????? ??????????, ???????????? ??????????????.
????????
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?? ???????? ?????????, ? ????? ?????????? ?? ????????????????, ????????????? ?? ??????????-????????????? ??????.
???????? ?????? ???????????? ?????????
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???????????????-?????????? ???????, ???????????? ??? ???????? ?????????? ???
??????. ?????? ???? ?????????? ??????????????? ? ????????? ???????????????? ???*
1
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?????? ?????????, ??? ??????????? ??????????? ?? ????? ???????????? ???????????
??????.
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Corresponding author E-mail address: evg@ksc.krasn.ru
© Siberian Federal University. All rights reserved
# 75 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ??????????? ?????????? ?????? ?????? ????????? ????
???, ??????????? ?? ?????????? ?????????
??????????? ????????? (??????????, 1984;
Gates et al., 1965; Goward, 1989; ???????? ? ??.,
1998; Bohn et al., 2000), ? ????? ??? ????????
????????????????? ??????????? ?????? ???????? ? ??????????? ? ??? ???????????,
?????????? ? ?????????????? ???????? ??????????????, ???????? ??? ????????????,
???????????? ????????????? ??????????
(???????, 2009).
? ????????? ????? ? ??????????????
??????? ??????? ????????????????? ?????????? (SPOT-Vegetation, TERRA/Modis,
NOAA/AVHRR, Landsat-7) ??????????? ???????? ????????????????? ??????????????
????????? ??????? ?????-????????? ?????????? (???????? ? ??., 2005; ??????? ? ??.,
2007 ? ??.) ? ?????? ??????? ?????? (Kharuk
et al., 2003). ???????? ?? ????????? ?????
??????? ? ???????????? ???, ????????????
???????????????? ?????? ??????????????
????????? ?????????? ??????????? ???? ??
???????? ??????????? ??????? ? ???????????????????, ??????????????????? ? ???????
????????????????? (?????????? ? ??., 1986;
????????????????, 2006).
?????????? ??????????? ?????? ????????? ?????????? ????????? ???????????? ????????????? ????????? ??? ? ???????? ??????, ????????????? ??? ? ????????????, ??? ?
?? ???????. ??? ????? ?????? ???????? ???????????????? ????????? ?????????. ??????
?????? ?????????? ???????? ??? ????????????? ????? ?????? ???????????? ??????, ?????????? ???????? ???????? ? ??????? ?????
??????? ?????????, ? ????? ????????? ??????, ???????????? ? ???????????? ????????? (Nazimova et al., 2000; Bartalev, Belward,
2002; ??????????, ????????????, 2002).
???????? ???? ??????? ????????????
?????????? ?? ????????? ? ?????????????
???????-??????? ????????????? ??????? ??-
????? ? ?????? ???????? ???????????????
???????????? ?????????, ?????????????
??? ?????????? ????????????? ?????? ??
?????????.
????? ?????,
????????? ? ??????
???????????? ????????? ?? ????????
???????? ?????????? ????? ? ????????
«????????????», «??????» ? ????????? ????? ? ??????? «???????????» (???. 1). ?????
???????????? ??? ???????????? ???????
????????????????? ? ??? ???????-???????
????????? (???) ? ?????????? ????????????? (???????-???????????????), ????????
???????-???????? ? ?????? ? ?????-???????
????????, ???????? ??? ??????????? ?????????? ? ???????? ????????. ?????? ???
????????????? ???????????????? ?????? ?
?????????? ??????????? ??? ?????? ??????? ???????? ?????-???????? ????????? ??
???????? ??????? ? ?????????????? ???????
?????????????????.
??????????? ?????? ??? ?????????? ??
?????? ??????????? ??????????? ?????? ?
?????????? ????????? ???????????? ???????? ????????, ??????? ????? ? ???????? ?????? ??? ?????????? ? ????????? ???????????
?????????????? ??????????? ??????????.
? ?????? ?????????????? ?????? ????????????????? ?????? ??????? ????????????????? ?????????? ?? ????????? TERRA/Modis
(250 ?, 1000 ?). ???????? ?? ??????????? ????????????? ? ?????? ?????? ? ??????? ?????????? ? ???????? ?????????? ??????????,
??? ???????? ??????????? ???????? ??? ?????????? ??????????? ??????????????? ????????? ?????????. ?????????????? ??????
? ???? ???????????? ?????????? TERRA/
Modis: 0,62 ? 0,67 ???, 0,84 ? 0,87 ??? (??????
1 ? 2, ??????????????? ?? ?????? ????????????? ?????????) ? 10,78 ? 11,28 ??? (????? 31,
# 76 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ??????????? ?????????? ?????? ?????? ????????? ????
???. 1. ????? ????????????. ???????? ????????: 1 ? «????????????», 2 ? «??????», 3 ? «???????????»
??????????? ???????????????? ??????????? ???????????). ? ?????? ???????????????
?????? ?? 2005-2009 ??. ???? ???? ???????? 9
???????, ?????????????? ????? ???????????? ? ???????? ??????, 20 ??????? ???????
???? ? 8 ??????? ?? ????????-???????. ?????
(Nazimova et al., 2000; ???????? ? ??., 2005)
??????????? ???????????? ???????????
???? ?? ?????????? ?????? NOAA/AVHRR ?
1, 2 ? 5 ???????, ??????? ??????? ?????????
0,58 ? 0,68 ???, 0,725 ? 1,10 ???, 11,5 ? 12,5
??? ??????????????.
????????? ? ?????? ??????????? ??????
???? ??????????? ??????????????? ?????????, ??????? ????????????? ?????????, ????????? ?? ???? ??????, ?????????? ?????? ?
?????????????? ? ???????? ??????????????
????????. ????????????? ??????????? ?????? ???????????? ?? ????????????? ???????
? ?????????? ?????????????? ????????????? (???????, 2000) ? ?????????????? ????????????? ??????????? ????????? (ERDAS
IMAGINE).
??? ?????????????, ???????? ? ??????????? ??????????? ????? ???????????
?????, ?????????? ? ???????? SPOT-4 (???-
????????????? ?????????? 20 ?) ? ??????????
????????????????? ?????????? ????? ???? ?
???????? ???????? ????????. ????????????
?????? ????? ??????? ????????? 0,50 ? 0,59
???, 0,61 ? 0,68 ??? ? 1,58 ? 1,75 ???. ? ?????? ?? ????? ?????? ?????? ?? ?????????????
?????? (??? ? ??????) 2009 ?.
???? ?????? ?? ????????? ? ????????
????????????? ??????? ???????? ?????????
??? ??????????? ?? ?????? ???????????
?????????????? (1960 ? 2009 ??.) ????????
???????????? (?????????, ???????? 2009;
??????? ? ??., 2004). ????????? ???? ????????? ? ?????????????? ???????????? ????????????????? ?????? (???). ??? ???????? ????????? ???????? ???????????? ?? ???????
?????? ???? ??????????????, ????????????
???????? ? ???????? ????????? ?????? ??????? (????? ? ??., 2009).
?? ?????? ??????????????? ?????????
??? ?????????? ????????????? ??????????
??????????? ?????? ????????? ?????????
??????? ???? ??????????? ? ??????? ???????? ????????????. ????? ?? ?????? ???????? ??? ?????????? ? ????????? ??????????? ?????????????. ?????????? ???????
# 77 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ??????????? ?????????? ?????? ?????? ????????? ????
????????????? ???????????? ?????????? ??
????? ?????? ???????? ????????????.
?????????? ? ??????????
? ???? ????????? ??????????? ?????????? ? ????????? ?? ? ????????? ???????
???????? ??????????? ???????????? ??? ??
?????? ????????? ?? ???????? ? ????? ?????
????, ? ????? ?? ?????? ???????-????????????
????? (???) ????????? ???????. ???????
??? ????????? ???????? ????????????
??????????????? ????????? ??? ?????????
????????, ????????? ? ???????? ??? ????
??? ??????? ????????????? ????????????
????? ? ????? ??????? ?????????, ?????????
???????? ????????? ???????????? ????????????? ???????????? ? ???????? ???????
??? (????????, ????????, 1995).
?????????? ????????????? ???????
TERRA/Modis ? ???????? ????????, ???????????? ???????????????? ???????????,
???????? ?????????????????? ?????????? ?
????????? ???????. ?? ??????? ????????
«??????» (????????? ????) ??????? ????-
???????????? ?????????? (? ???????????
?????????) ? ?????-??????? ????????????
??? (???. 2). ????????? ?????????? ?????????? ??????? ????????? ? ??????? ?????? ????????? (????????????, ?????????, 2006).
????? ????, ? ???? ???????????, ????????????? ??????????? ???????????? ??????,
?????? ?????? ?????????????? ????????? (?
?????????, ???????????? ?????????? ? ??????? ???), ??????? ?? ?????????? ? ??????????????? ????? ? ??????????????????
?????????. ?????? ????????? ??????????,
??????????? ?? ??????????? ?????????????, ??????????? ? ????. 1. ? ??????? ?????? ????????? ? ???????? ???? ????? ???
??????????? ???????? ?????????????????.
?????????? ??????? ???????? ???????????
????????????? ??????????? ? ???????? ????,
? ?? ????? ??? ??????, ??????????? ?????? ?
? ????? ????, ????????? ???????? ????????
?????????? ?????? ???????????? ???????
???. ????? ?????????????? ????????????
?????????? (???????? ? ??????? ???) ? ????
??????, ??????????? ????, ?? ????????-
???. 2. ????????????? ???????-??????? ?????????? (???) ?? ?????? ????????? ?????????? ??????
TERRA/Modis. ??????? «??????». ????? ? ????????? ????? ?? ??????????? ??????????????, ?????? ?
????????? ????????????? ??????????? ? ??????????? ????????????. 1 ? ?????????-??????????? ???,
2 ? ???????????? ???, 3 ? ?????????????? ????????? (??????? ? ??????? ???)
# 78 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ??????????? ?????????? ?????? ?????? ????????? ????
??????? 1. ??????? ???????? ???????? ?????????? ???????, %
?
1
?????? ?? TERRA/Modis
???????-??????? ?????????
?????????-???????????
1
2
67,5
32,5
2
?????-???????
19,7
80,3
3
??????????????
73,5
26,5
?????? ?????????? ??????? ? ??????????
???.
??? ??????? ??????? TERRA/Modis ??
?????????? ???????? «????????????» ????????????? ?????? ????????????? ???
???? ???????????????? ??????? ? ????????????? ?????, ??????? ? ?????????? ?????????? ?????. ??????????? ????????? ?
??? ????????????-??????????????? ???????? ????? (????????). ?? ??????????? ??????????? ?????????????, ????????????? ?
??????? ??????????????, ??????? ? ???????? ?????? ?????????? ????? 70 %, ? ??????? ???????? ?????? ? ???????? ?????????
?????????? ????, ??????????????? ?? ?????? ?????????????? (???. 3). ???????????
??????? (???????????) ? ???????????????
(?????) ? ?????? ?????? ???????????? ? ?????? ? ????????? «????????? ??????????»,
??? ????? ??????? ????????? ?????? ?????????? ?????? (?????? ?????) ?? ???????????
???? ? ?????? ? ???? ????????? ?????.
????? ???????, ????????? ??????
TERRA/Modis ????????? ??????? ??????????? ????? ???????? ? ????????? ????? ?????? ?????????, ? ????? ????, ? ?????????????
????????? ?????? ????????? ???????? ??????? ??? ????? ? ???????? ? ????????? ?????????????? ?????? ??????? ?? ???? ???.
??????????? ???? ?????????? ? ?????????
????? ????????? ???? ??????????? ??????????? ??????????????? ? ?????????????? ???????? ????????? ??????? ???????.
??? ??? ??????????, ?????????? ?????????????? ?????? ?? ?????? ??????????
??????? ????? ????????? ???????. ????
?? ????????? ???????? ??? ?????????? ???????????????? ?????????? ? ?????? ?????? ? ???????????? ????????? ???????, ?
????????? ????????????? ???????? ???????
TERRA/Modis. ????????????? ?????????????? ???? ???????? ????? ?? ????????????
????????????? ????????????? ???????, ?????????????? ???????????? ????? ???? ??????????? ?????????? (???????????? ???????????, ???????????? ??????) ? ?? ?????????
(??????? ????????????, ?????????????? ?????????, ??????? ??????????? ???????? ??????). ?? ????? ???????, ? ?????????-???????
????????????? ??????? ????, ???? ????????? ??????? «???????????» (?????????? ?
??., 1986), ????? ???????? ?????? ?? ?????
??????? ??????????? ? ?????????? ?????????????? ????, ???????????? ?? ??????
?????? TERRA/Modis ? ???????? ?????????
(31 ?????). ??? ???? ???????????????? ??????????? ??????????? ??? (???????????
? ?????????) ? ??????? ????? ???????????
? ?????????????? ?????????????? ???? ?????? ???? ? ??? ??????????? ???? ??????
?????????. ????????????-???????????????
???????????? ???? ???????? ??? ????????? ? ????? ???????? ???????? ??????????????? ? ???????-???????? ?????????????????????????? ???????? ???? ?? 4 ? 5 ????
?? ???? ????? ??????? ?????? ????? ? ???????
# 79 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ??????????? ?????????? ?????? ?????? ????????? ????
???. 3. ?????? ????????????? ???????? ?????? ?? ?????? ????????? ?????? TERRA/Modis. ???????
«????????????». ???????? ??????: 1 ? ??????-???????? ???, ??????? ? ????????????? ?????; 2 ? ?????????????? ???, ?????????? ? ????????????? ???????????? (?????; ? ??????? ???); 3 ? ?????????? ???,
?????????? ? ????????????? ?????
???????????. ??????? ???????? ???????????????? ??????????? ??????????? ????? ??????? ?????????????? 40 ?? ????? ?????????? 10 ? 15 °?, ???????? ? ???????? ???? ??
2 ? 5 °?.
????????, ??? ???????-??????? ?????????????? ????? ?????????? ?? ????????????? ?????? ?????????? ?????????? ????
????????????? ???????????, ???????? ?????
?????????? ???????-???????? ??????? ??
?????? ???????. ???, ?????????? ?? ???????
(???. 4) ??????? ????? ????????????? ?????, ???????? ?? ?????????? ??????????
??????-????????? ??? (???. 4, ???????, a),
??? ??????????? ???????????? ????? ??????? ? ??????-???????? ???????????, ???????? ????????????? ? ???????????? (???. 4,
???????, b). ???????????? ????????? ??
???? ?????? ? ????????? ????????????????
?????, ????????? ??? ????? ?????? ?????????????? ???????????? ????????? ???. ???
???? ?????????????? ??????? ????????????? ??? ?? ?????????? ?????? ???????? ????????????????? ?????????? (???????? ????-
?? SPOT-4 ??????????? ?? ???. 4 ?? ???????),
??? ? ?? ??????? ? ?????? ???????????, ???????? ? ???????? ????????? ??????????
TERRA/Modis.
? ????? ???????? ???????? ???????????????? ??????????? ? ???? ???????? ??
???????? ? ????????????? ??? ?? ???????? ??????????, ? ? ??????? ??????????????
?????? ??????????? ??? ????????. ??? ??
?????? ??????????, ??? ?????????? ?????????
?????????? ??????????? ?????? ? ?????? ???????. ?????? ???????? ??????????, ???
??????? ???????, ??????? ???????? ??? ?????????????? ??? ????????? ???????? ?????????? ??????.
?? ??????? ????????????? ???????????
?????? ???????? ??????????? ??????, ??????
??????? ????? ? ???, ??? ? ?????? ???????????? ????? ?????????? ???? ??????? ???????
?? ??????????? ??????, ???? ?? ???????.
? ?????????? ????????? ??? ? ?? ???
?????? ?.?. ?????????, ?.?. ?????????????,
?.?. ????????? ? ?.?. ?????????, ?.?. ????????? ? ?????????? ? ??. ??????? ? ????????-
# 80 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ??????????? ?????????? ?????? ?????? ????????? ????
???. 4. ?????? ????????????? ??????????? ?????? ? ?????? ????????????? ??????????. ???????
«???????????». ????? ? ????????? ????? ?? ?????????? ??????????????, ?????? ? ?????????? ??
??????????? ????????? ?????? ? ???????? ?????????. ???????? ???????????: 1 ? ???????? ?????????????????????? (Pinus sylvestris, Betula pendula); 2 ? ??????????? ???????? ????: ???????????????
(Betula platyphilla, B. alba, Populus tremula), ???????? (Abies sibirica) ? ????????? ?? ????? ??????
??????? ? ???????? ??????????? ? ?????? (Pinus sibirica, Abies sibirica); 3 ? ???????????? ?????????????? (Abies sibirica, Pinus sibirica); 4 ? ????????????? ?????????? (Pinus sibirica, Abies sibirica), ????,
??????, ??????. ?? ??????? ? ???????? ??????????? SPOT-4: ? ? ??????-???????? ???; b ? ??????? ?
??????-???????? ???????????, ???????? ????????????? ? ????????????
?? ???????? ??????? ???????????? ??????????? ? ?????????????? ??? ? ???????????
?????????, ? ??????? ?????????? ????? ???????????? ????, ??????????????, ???????????? ???????. ? ????????? ???? ?????????
?????? ?? ?????? ???????? ??????? ? ????
?????????? (????? ? ??., 2002; ??????????,
2009 ? ??.). ??? ?????? ????????? ??????????? ???????????????? ??????? ? ?????????????? ????????????????? ?? ?????? ?????? ?
??????????? ?.?. ?????? ? ?????????? ???
(?????????????? ????????????..., 2007). ???????????? ??? ? ??????? ?????????????
?????? ??????? ????? ?????????????? ??????
?? ?????? ??????? Spot Vegetation (Bartalev
et al., 2003). ?????? ? ???????? ???????, ???
???????, ?? ?????????? ??????????????
?????? ?????? ?? ???????????????? ???-
??????, ??? ?? ????????? ???????? ??????
????? ?????????-????????????? ???????? ?
?????????-???????????? ???????? ?????,
?????? ??? ???? ??????????????? ? ??? ?
?????? ????? ???????????.
???????????? ????? ?????? ? ????????????? ????? ???????, ?????????? ????????? ???????? ????????? ?? ???????????? ??????? ?????????. ????? ??????????????? ???
??????????????????? ??????? ??????????
?????, ??? ??????? ????? ???????? ????????,
??????? ???????? ??????????????? ????????? ????????? ?? ???????????? ??????????? ???? ? ???????? ???????????????. ? ?????
?????? ?? ?????????? ???????-???????????
??????, ???????? ??? ?????????? ???????
?.?. ??????? ? ?.?. ????????, ? ?? ?????? ?
???????????????????? ??????????? ??????
# 81 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ??????????? ?????????? ?????? ?????? ????????? ????
?????????? ? ??????? ?????????? ???????
(Ecosystems of British Columbia, 1991 ? ??.).
???????????? ?????? ????????? ???????? ?? ??????????? ???????????? ???????
????????????????? ?????????? (????? ???
NOAA/AVHRR, TERRA/Modis) ?????? ???
??? ????????? ? ?????????? ?????: ??????????, ???????? (??????-????????), ????????????, ????????????, ??????? ????????
???????? ??????? ? ???? ????????????????
?????????? ? ?????? ???????? ??????????
???????. ??? ???????????? ????? ?????? ???????? ???????????????? ?????, ??????? ????????????? ???????? ?? ?????? ? ?? ??????
?????????? ? ??????? ???????? (1:50 000 ?
??????). ?? ???????????? ???????? ??? ????????????? ?????????????????? ????????.
?? ??????????? ??????????? ????????????
?????? ????? ? ????????????? ????????????? ?????? ? ???????? ????????? ??????? ?
????? ??????????? ???????? ??????? ?????-
?? ??????????? ?????????, ???????? ??????
?????? ??????? ?? ????????????? ?????????????.
??????????????? ?????? ? ????????????????? ? ??????????? ????????? ?????
? ?????? ????????? ?? ??????? ??????????????? ?????????????? ?????? ? ?????? ?
????????? ??????????????? ???. ??? ??????????????? ? ??????????? ??????? ????????? ?????????? ??????????? ??? ? ???????????? ?????.
??? ?????? ????????? ? ????????? ????????
???. ???? ?????? ?????????????? ? ?????????? ????????? ??? ????????????????? ????
???? ????? ????? ??????? ?????? ?? ??????????? ??????? ??????? ? ???????? ????????????????? ??????????.
??????????? ?????? ? ????????? ?????????? ??????? ????????? ??????????? ??????? ????????? ? ????????? ?????????????
??????? ?????? ??????????, ????????? ? ?????????????? ????????????? ? ????? ?????????? ?????????. ?????? ???? ?????????
????? ???? ????????????? ?? ??????? ???????? ??????????, ? ????? ? ??? ???????????
??????? ?????? ??????? ?????????????????
??????????. ????????? ??????????? ??????
??????????? ??????? ??????????????? ??
???? ?? ?????????? ??? ?? ?????? ????, ???
? ? ????????? ?????????????? ?????. ???
?????? ?????????? ??? ??????????? ???????????????? ?????? ????????? ? ?????????
?????????, ?????????????, ??????????????? ????? ? ????????? ????? ??????? ????????? ? ?????????????? ???????? ????????
?????? ??????? ?? ???? ???.
? ?????????-??????? ?????????????
??????? ???? ????? ???????? ?????? ??
????? ??????? ??????????? ? ??????????
?????????????? ????. ??? ?????????? ?????????????? ?? ??????????? ???????? ?????,
??????????? ?? ?????? ?????? TERRA/
Modis ? ???????? ????????? (31 ?????).
?????????????? ???????? ??????????????????????????? ???????????? ????? ???????? ? ???????-?????????? ?????????????????????????? ???????? ????? ????????
??????? ??????????? ? ?????? ???? (??????
?????? ??? ? ????), ??? ?????????? ????? ??????? ??????? ?????????? ?? ?????????? ???
????????????? ???????????? ??????? ???????????? ????????????? ??????? ? ????????
???????.
??????????
?? ??????? ????????????? ???????
???? ?? ??????? ??????? ????? ? ????? ????
???? ???????? ???????? ?????????? ??????? ??? ???????? ? ???????????? ??????
?????, ????????????? ? ??????????? ?????????????? ? ????????? ??????????????. ?
??????? ?????? TERRA/Modis ?????????
?????? ??????????? ????? ???????? ???-
# 82 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ??????????? ?????????? ?????? ?????? ????????? ????
?????? ????????? ??? ????????? ??????? ???? ?08-04-00-600-? ? ?08-04-00-613-?,
? 09-04-98040-?_??????_? ? ??????? ?? ??? ?27.2 ? ?????? ????????? ??? «????????????? ????????????».
?????? ??????????
????? ?.?., ??????? ?.?., ????????? ?.?., ???????? ?.?., ???????? ?.?. (2009) ???????? ????? ?????? ?? ???????????? ????? ? ?????????????? ???????? ????? ????????? ?????.
???????????? ??? ?????? ?? ??????????????? ? ????, ????????, ???????: ???. ????????.
??????-?????. ????. ???????, c. 38-43.
???????? ?.?., ????? ?.?., ????? ?.?. (1998) ?????? ?????????? ????? ?? ?????????????? ??????????? ???????????? ??????? ???????????? ???????????? ??????. ????????????
????? ?? ???????. 3: 95 ?107
????? ?.?., ?????? ?.?., ???????? ?.?. (2002) ???????????????? ???????? ?????. ? ???????: ????????????, 160 ?.
??????? ?.?. (2000) ????????????? ???????????? ? ?????????????? ??????????????
???????. ?????. 1. ?.: ??????-92, 140 ?.
??????????????? ? ???????? ?????????: ?????????????? ?????????? ? ?????????????.
(2006) ???. ?.?. ??????, ?.?. ?????, ?.?. ????????, ?.?. ???????. ???????????: ???-??
?? ???, 648 ?.
?????????? ?.?. (1984) ??????????????? ?????????? ?????????. ?.: ?????, 320 ?.
?????????????? ???????????? ??????. (2007) ?.4. ????????????? ???????????? ?????????????????. ???. ???????? ?.?. ???????????: ???, 412 ?.
??????? ?.?., ??????? ?.?., ????????? ?.?. (2004) ???????????????? ?????? ?????????
???????? ??????????????? ?????????? ? ?????????????? ???. ?????????????? ??????????.
9(42): 141-146.
???????????? ?.?., ????????? ?.?. (2006) ???? ????????????? ?????? TERRA/Modis
??? ????????? ?????????????? ?????? ????????????? ???????? ? ???????????? ????? ????????????? ????? ????. ?????. ??????. ? ??????, ???. 14, ?. 35 ? 38.
??????? ?.?., ???????? ?.?., ????? ?.?., ?????????? ?.?. (2007) ????????????? ???????????????? ??????????? ?????????????? ?????? ?????????? ?? ?????? ?????? ?????????????? ???????????? ? ???????? ?????? ???????. ?????????????? ??????????. 12 (2): 42 ? 59.
??????? ?.?. (2009) ???????? ???????? ????? ???????????????? ?????????????? ????????? ??????????? «???????-?????????????? ??????? ???????????????????? ????????». 23-25
???????? 2009 ?. ??????????, ?. 66-69.
????????? ?.?., ???????? ?.?. (2009) ???? ?????? ?? ?????????? ???????? ??? ?????? ??????????? ???????? ???????????????? ????????? ?????. ???????????? ??? ?????? ??
??????????????? ? ????, ????????, ???????: ???. ????????. ??????-?????. ????. ???????,
c. 105-111.
?????????? ?.?., ???????????? ?.?. (2002) ???? ?????????? ??????????????? ???????
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?????????? ? ??????????? ? ?????? ?????????». ?., ??????????????, c. 157-160.
# 83 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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using the SPOT ? VEGETATION instrument: new observations for climate studies. In: Proceedings of
the Use of Earth Observation data for phenological monitoring workshop,ld in Joint Research Centre,
Ispra (VA) Italy 12th-13th December 2002, pp. 41-48.
Bartalev S.A., Belward A.S., Erchov D.V., Isaev A.S. (2003) A new SPOT4-VEGETATION
derived land cover map of Northern Eurasia. Int. J. Remote Sensing. 24( 9): 1977-1982.
Bohn U., Gollub G., Hettwer C., Neuhauslova Z., Raus Th., Schluter H., Weber H., eds. (2000)
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of Forests Province of British Columbia. Victoria, B.C. Special report series, 6. 330 p.
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Opt. 4: 11-20.
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Kharuk V.I., Ranson K.J., Burenina T.A., Fedotova E.V. (2003) Mapping of Siberian forest
landscapes along the Yenisey transect with AVHRR. Int. J. Remote Sensing. 24 (1): 23-37.
Nazimova D.I., Ponomarev E.I., Fedotova E.V. (2000) Identification and mapping of altitudinal
belt classes of land cover with use of NOAA/AVHRR imagery. Remote researches and mapping of
geosystems structure and dynamics. Novosibirsk: SB RAS, p. 76-81.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ??????????? ?????????? ?????? ?????? ????????? ????
Satellite Monitoring
of Sayan Mountain Forest Ecosystems
Evgeny I. Ponomarev,
Dilshad M. Ismailova and Dina I. Nazimova
V.N. Sukachev Institute of Forest of Siberian Branch
of Russian Academy of Sciences
50/28 Akademgorodok, Krasnoyarsk, 660036 Russia
New approach to use remote sensing data of low and moderate spatial resolution is suggested for
forest cover mapping and for its seasonal monitoring. The researches of forest structure and dynamics
are conducted on three test sites of the Western and Eastern Sayan mountains since 1960. Satellite
remote data for the territory (NOAA/AVHRR and TERRA/Modis imagery) are available for 1996 ?
2009. After description of the data base the features of different forest zones and formations are shown
in context of their identification and spectral characteristics changing during phenological seasons.
Specification of mountain forest zones with different potential composition is important for forest
management and monitoring on regional and local levels.
Keywords: mountain boreal forest, altitudinal zones, remote sensing imagery, database, seasonal
monitoring, spectral characteristics.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2011 4) 86-103
~~~
??? 597.554.3: 591.9(289)
Growth, Diet and Fatty Acid Composition
of Gibel Carp Carassius Gibelio in Lake Shira,
a Brackish Water Body in Southern Siberia
Denis Yu. Rogozin a,b*,
Marina V. Pulyayevskayab, Ivan V. Zuevb,
Olesya N. Makhutovaa and Andrey G. Degermendzhia,b
a
Institute of Biophysics SB RAS
Akademgorodok, Krasnoyarsk, 660036 Russia
b
Siberian Federal University
79 Svobodny, Krasnoyarsk, 660041 Russia 1
Received 4.03.2011, received in revised form 11.03.2011, accepted 18.03.2011
For the first time the presence of gibel carp Carassius gibelio has been detected in the brackish
meromictic Lake Shira (Southern Siberia, Republic of Khakassia). Lake Shira has been continuously
studied since the late 19th century, and it has been considered so far that there is no ichthyofauna in
the lake. This work investigates the growth and population characteristics, as well as food spectra
and fatty acid composition of C. gibelio specimen at different ages that were fished out in the summers
of 2007 and 2008. The water salinity at the fishing point was 15 g l-1. The food spectrum of one- and
two-year old C. gibelio was dominated by diatomic, green and blue-green algae, while four- to sixyear old species had an increased share of copepods, beetles and chironomids in their food spectrum,
and beginning from the age of six years the amphipods Gammarus lacustris consisted significant part
of gibel carp diet. Thus, it was shown that C. gibelio is a carnivore with regard to the population of
amphipods Gammarus lacustris, which have been regarded so far as the upper link in the trophic
cascade of Lake Shira. One of the possible reasons for appearance of fish is the decreased water salinity
caused by rising of the lake level. In the recent years, the level of Lake Shira exceeded all the values
that had been registered since the late 19th century, and the lake?s salinity decreased correspondingly
to its smallest value for the whole period of observation.
Keywords: Carassius gibelio, brackish lake, food spectrum, fatty acids.
Introduction
Fish introductions are amongst the greatest
threats to freshwater biodiversity (Gherardi,
2010), and one of the most invasive freshwater
*
1
fish species is the gibel carp Carassius gibelio
Bloch (Szczerbowski, 2002; Luskova et al.,
2010) which has been recently found to have
established a population in the brackish Lake
Corresponding author E-mail address: rogozin@ibp.ru
© Siberian Federal University. All rights reserved
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Denis Yu. Rogozin, Marina V. Pulyayevskaya? Growth, Diet and Fatty Acid Composition of Gibel Carp Carassius?
Shira previously reported as a fish-less lake
(Degermendzhy, Gulati, 2002; Degermendzhy
et al., 2010). Gibel carp Carassius gibelio, being
a stenohaline species, is rather often observed
in brackish water, for example, in the littoral
zones of the Black and Baltic seas (Abramenko,
2006; Vetemaa et al., 2005), in the delta of the
Ob (Zhadin et al., 1961) etc. Lake Shira has
been studied for decades by many researchers
(Krivosheyev, Khasanov, 1990; Degermendzhy,
Gulati, 2002; and many others), but it has been
considered until now that there is no permanent
ichtyofauna in the lake (Degermendzhy, Gulati,
2002; Degermendzhy et al., 2010). In summer,
the salinity of the epilimnion water was about
15 g l-1 during last decade (Rogozin et al., 2010),
however, lower salinity was observed near
estuary of the river Son incoming to the lake.
According to the oral evidence of local residents,
gibel carp has been regularly observed in the
lake lately. However, there have been no reliable
facts proving this information so far. In 2007, we
made trial sampling that confirmed the presence
of gibel carp in the lake. Since the fish population
may provide considerable impact on previously
fishless ecosystem, it seems to be important to
study the main characteristics and ecology of this
population. At the first step, we tried to analyze
the feeding spectrum, growth characteristics
and fatty acids composition of gibel carp in the
lake. Fish species, from different ecosystems, are
known to differ in their fatty acids composition;
therefore, studies of fatty acids contents of
diverse fish from various locations are of great
importance for revealing their potential value
as sources of the essential ?3 acids for human
nutrition (Ahlgren et al., 1994; Gladyshev et al.,
2009). Polyunsaturated fatty acids (PUFAs) of the
?3 family became known as key dietary nutrients
for preventing mental, neural and, especially,
cardiovascular diseases of human (Arts et al.,
2001; Garg et al., 2006). Besides, biochemical
analysis of fatty acid composition is widely used
to trace trophic interactions between hydrobionts
and to reveal food assimilation (Desvilettes et al.,
1997; Sushchik et al., 2003). Therefore, the aims
of present work were to provide the first evidence
of presence of fish in Shira Lake, to analyze the
diet of the fish and to give some basic information
about growth characteristics and fatty acids
composition.
Materials and Methods
Lake description
Lake Shira (90.11 E, 54.30 N) is located in
the northern part of the Republic of Khakassia,
South Siberia (Russian Federation). This elliptical
(9.3 Ч 5.3 km) lake has an area of 35.9 km2 and
maximum depth of 23.8 m (2007-2010). The main
inflow is from the Son River that provides about
42 % of fresh water supply to the lake; other
sources of water are precipitation and seepage
water (Parnachev et al., 2002). The lake has no
outflow. The salinity of lake water during summer
stratification is about 14-15 g l-1 in epilimnion
(upper 5-6 m of water column) and 18-19 g l-1 in
deeper waters. The detailed description of the
lake is given elsewhere (Degermendzhy et al.,
2002; Parnachev et al., 2002; Kalacheva et al.,
2002; Rogozin et al., 2010). The surface elevation
of Lake Shira has noticeably changed over the
whole period of observation. The lake?s salinity
varied inversely with changes in water volume
(Krivosheev, Khasanov, 1990; Rogozin et al.,
2010). During the period from 1920-s to 1930-s
the lake?s level decreased by 7 m, which was
caused by the decrease in the total amount of
precipitation in the region. In this period, salinity
reached the maximum of 27 g l-1 in 1926. By the
present moment, the lake level has exceeded all
the values registered since the late 19th century,
and the salinity has dropped, respectively, to its
lowest value for the whole period of observation
(Rogozin et al., 2010).
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Denis Yu. Rogozin, Marina V. Pulyayevskaya? Growth, Diet and Fatty Acid Composition of Gibel Carp Carassius?
Sampling and water measurements
Fish sampling was carried out in 2007-2009
in Point ? (54.28.767 N, 90.14.818 E), in several
points of Area ?, and in point ? (54.29.053 N;
90.16.352 E) (Fig.1). Exact coordinates of Area
B point were not defi ned, and the location of
Area B was estimated visually on the basis of
landmarks. The dates of fishing are given on
Fig.1. Fish were captured with gill nets of various
dimensions and mesh sizes, complemented for
young specimens by drag-nets, bottle and can
traps as well as rod and line angling. In all
points the nets were set in the evening, along
macrophyte beds, at a depth of approximately
1.5 m not far from the shore; the nets were
checked early in the morning next day.
The temperature and specific conductance
of water in points ? and ? (Fig.1) were measured
with YSI 6600 (Yellow Springs, Ohio, USA)
submersible profiler. The salinity was calculated
as quadratic approximation of the dependence of
ash content upon specific conductance measured
for the Lake Shira water in laboratory:
S = -0.087 cond2 + 4.4403cond ? 37.16,
R2 = 0.95, n = 19,
where S is the salinity determined as ash content
(g l-1), Cond is specific conductance at 25°C of
the lake water samples (mS cm-1). Conductivity
sensors were calibrated against 3M KCl (YSI).
Oxygen electrodes were calibrated against
Winkler titration of the Lake Shira water with
Aquamerck kit (Merck, Germany)
Back-calculated growth,
weight?length relationship
and Fulton?s condition factor
0.01 g on electronic balances and measured to the
nearest millimeter. Scale samples for age and
growth analysis were collected from each fish.
Scales were taken above the lateral line dorsal
to the tip of depressed left pectoral fin. Growth
histories of individual fish were determined by
aging and back-calculation of total lengths (TL)
at previous ages from scales (Pierce et al., 1996).
Twelve to fifteen scales per fish were cleaned and
mounted between glass slides. All scales on slides
were viewed when ages were assigned to fish, and
a single reader did all aging. Anterior radii and
interannular distances were measured to nearest
0.01 mm by using dissecting microscope (25Ч
magnification). These radii are hereafter referred
to as scale lengths. Replicate measurements were
then averaged for each fish, providing the precise
estimates of scale lengths for back-calculations.
With the data on individual TL and mean scale
length, we then back-calculated TL at previous
ages using the Fraser-Lee formula (Pierce et al.,
1996).
Weight?length relationship and Fulton?s
condition factor were calculated from the
expressions:
(1) W = a TLb
(2) K= 100 W/SL3,
Where W is the whole body weight in grams,
TL the total length, SL is the standard length in
centimeters; a and b are the regression constants;
the factor 100 is used to bring K close to unity
(Froese, 2006).
Diet analysis
The growth history was determined for 200
fish collected from the end of July to the beginning
of August in Area ? (Fig.1). After capture all the
fish were individually weighted (wet) to the nearest
Diet was assessed on the basis of gut
contents determined by dissection of 32 fish
collected from the end of July to the beginning of
August in Area ? (Fig.1). All fish samples were
weighted and measured as described above,
and their guts were removed immediately, and
# 88 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Denis Yu. Rogozin, Marina V. Pulyayevskaya? Growth, Diet and Fatty Acid Composition of Gibel Carp Carassius?
stored in formalin (4%) until the contents were
analysed. The analysis of digestive tracts was
performed in accordance with the standard
method (Metodicheskoye rukovodstvo, 1974;
Hyslop, 1980). Animal food items were
identified to the lowest possible taxon. The diet
was analyzed for the percentage composition
by mass.
Analysis of fatty-acid composition
Detailed description of the analysis is given
elsewhere (Makhutova et al., 2003; Sushchik et al.,
2006). Briefly, lipids from samples were extracted
with chloroform:methanol (2:1, v/v) three times
simultaneously with mechanical homogenization
of the tissues with glass beads. Before extraction,
a fixed volume of an internal standard solution
(19:0) was added to the samples. Methyl esters of
fatty acids (FAMEs) were prepared in a mixture
of methanol: sulphuric acid (50:1, v/v) at 90 oC
for 2 h. FAMEs were then analysed using a GC?
MS (model GCD Plus, Hewlett Packard, USA)
equipped with a 30 m long Ч 0.32 mm internal
diameter capillary column HP-FFAP. FAMEs
peaks were identified by their mass spectra,
comparing to those in the data base (Hewlett?
Packard, USA) and to those of available authentic
standards (Sigma, USA). To determine double
bond positions in monoenoic and polyenoic acids,
GC?MS of dimethyloxazoline derivatives of FA
were used.
ever documented proof of the presence of fish
in Lake Shira. The age was determined for six
specimens of this capture (Table 1).
On August 9 2008, in Point ? the water
temperature was 23 °?, and the salinity was
14.7 g l-1. In this date, 130 two- to four-yearold specimens and 68 five- to eight-year-old
specimens were captured using nets with a mesh
size of 25 mm and 40 mm, respectively. The
specimens of this capture were analyzed only for
fatty-acid composition.
In area B (Fig.1), 200 specimens of Carassius
gibelio of different sizes and ages were captured
from the end of July to the beginning of August
2008; the water temperature and salinity were
not measured. The specimens captured in 2008
in Area ? included fish aged from 1+ to 15+ years
(Fig.2, Table 1), with the predominance of 7-yearold specimens (26.5 %). Fish aged 5+ and 7+
make up a large part of the captured total ? 15.5
and 18.0 %, respectively, of the total number of
captured specimens. There were few specimens
older than 8 years in catches. Almost in all age
groups the sex ratio was practically 1:1. It was
only in the 4+ and 7+ age groups that males
dominated. The body length of the fish in this
sample was from 54 mm to 460 mm, and the body
mass was from 2.47 to 1358 g.
In May 2009, no specimens were caught in
points A and C (Fig.1). The water temperature at
that time varied within the range of 10.7-13.2 ??,
and the salinity was 13.7 ? 14.5 g l-1.
Results
Fishing results
On July 24 2007, in the Point ? (Fig. 1)
the water temperature was 24 °?, and salinity
was 15.1 g l-1. The salinity in the central part
of the lake at this period was 15 g l-1, i.e. it was
practically the same as in the fishing point. The
captured fish included 33 specimens of 2- and
3-year-old Carassius gibelio, and one 7-year-old
specimen of the same species. This was the first
Weight?length relationship
and Fulton?s condition factor
The length-weight relationship calculated
for total length data (Table 1), was
W = 0.012 TL3.134, (R 2 = 0.995, n= 124) for
females,
W = 0.015 TL3.068, (R 2 = 0.978, n= 76) for males.
The relationship between TL and SL was
SL = 0.786 TL + 0.405, (R 2 = 0.998, n= 206).
# 89 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
N
S
1
C
A
B
Son River
1 km
Fig. 1. Fish sampling sites at Shira Lake in 2007-2009: (A) ? 24 July 2007, 9 August 2008 and 30 May 2009;
Area (B) ? 21-30 July and 3-5 August 2008; (C) ? 30 May 2009; (1) ? Town Zhemchouzhnyi and spa resort ?Lake
Shira?
Table 1. Length and mass of Carassius gibelio from Lake Shira collected on July, 2007, and July-August, 2008.
M ? total mass, m ? mass without entrails
Age
TL±SE (cm)
SL±SE (cm)
M±SE (g)
m±SE (g)
n
2007
2+
14.1 ± 0.3
10.2 ± 0.5
42.7 ± 4.6
35.3 ± 3.5
3
3+
14.8 ± 0.6
11.8 ± 0.3
61.8 ± 2.1
48.9 ± 1.6
2
7+
24.0
17.3
177
133
1
1+
7.3 ± 1.7
5.5 ± 1.3
6.4 ± 0.4
5.1 ± 0.3
34
2008
2+
-
-
-
-
0
3+
14.9 ± 1.1
12.0 ± 1.0
54.4 ± 1.4
47.0 ± 1.3
21
4+
16.5 ± 1.8
13.5 ± 2.0
83.1 ± 3.0
68.8 ± 2.8
10
5+
18.4 ± 0.8
14.8 ± 1.2
114.1 ± 2.5
92.0 ± 2.1
31
6+
19.9 ± 0.7
16.1 ± 0.9
149.2 ± 2.0
122.7 ± 2.0
54
7+
21.8 ± 0.8
17.6 ± 0.8
182.3 ± 4.0
147.3 ± 3.8
36
8+
23.0 ± 1.3
18.6 ± 1.8
210.7 ± 5.3
170.4 ± 5.2
10
9+
25.2 ± 1.5
20.8 ± 2.0
308.5 ± 1.8
265.3 ± 3.3
2
10+
26.0
21.2
301
264
1
15+
46.0
36.0
1358
1108
1
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Denis Yu. Rogozin, Marina V. Pulyayevskaya? Growth, Diet and Fatty Acid Composition of Gibel Carp Carassius?
Table 2. Fulton?s condition index (Kf ) of Carassius gibelio collected from Lake Shira in July and August, 2008
Males
Females
Age
Kf
n
Kf
n
1+
3.59 ± 0.03
30
3.79 ± 0.11
4
2+
-
0
-
0
3+
3.13 ± 0.05
13
3.22 ± 0.12
8
4+
-
0
3.41 ± 0.11
10
5+
3.33 ± 0.08
19
3.67 ± 0.08
12
6+
3.51 ± 0.04
33
3.51 ± 0.08
21
7+
3.37 ± 0.11
18
3.29 ± 0.10
18
8+
3.15 ± 0.09
8
3.6 ± 0.01
2
9+
3.43 ± 0.08
2
-
0
10+
-
0
3.16
1
15+
2.91
1
-
0
The Fulton?s condition index (Kf ) undergoes
minor changes in specimens belonging to
different age groups. The (Kf ) varied from 2.91 to
3.79 (Table 2). The (Kf ) of males and females was
almost equal, except the 5+ and 8+ age groups,
in which males were noticeably better-fed than
females (Table 2).
Growth history
Generally, the relative length increment
estimated from scales decreased with the age (Fig.3,
Table 3). The back-calculated patterns of length at
the age were similar among six generations, and
the year-to-year differences among the fish of equal
age-class were not significant, except youngest
group of 3+ (Fig. 3, Table 3). The relative length
increment of 3+ fish in 2005 was significantly
higher than for other generations at this age (Fig. 3).
However, this may be explained by shortcoming of
back-calculation method too (Pierce et al., 1996;
Martinson et al., 2000).
Analysis of diet
The diet of Carassius gibelio in Lake Shira
is rather diverse; it includes 4 divisions of algae,
9 groups of water invertebrates, remains of
terrestrial insects and higher aquatic vegetation
(Table 4, Fig. 4). In addition, the guts of fish in all
age groups contained sand which was obviously
consumed with food.
The largest number of diatoms was found at
the inflow of the River Son. These are diatoms
that are preferred by juvenile fish. The algal flora
of the lake typically includes algae of Pyrrophyta
division, but they were not present in the guts of
the caught gibel carps. The cyanobacteria of the
genera Spirulina, Oscillatoria, and Anabaena
dominating in the littoral zone of the lake were
not present in gut contents.
The zooplankton of Lake Shira is
characterized by a relatively small diversity of
species. The dominating species are Copepoda
and Rotatoria (Degermendzhy, Gulati, 2002)
which are present in the diet of gibel carp. Several
species of Cladocera are found in the south-eastern
part of the lake (at the inflow of the River Son).
Though their population size and biomass are
low, these animals are the most favorite delicacy
of gibel carp among all crustaceans. Benthic
animals also display little diversity, but they
constitute a considerable part of the C. gibelio?s
diet (Table 4, Fig 4).
# 91 #
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35
30
Fish number
25
20
15
10
5
0
1+
2+
3+
4+
5+
6+
7+
8+
9+
10+ 11+ 12+ 13+ 14+ 15+
Age, years
Fig. 2 Age and sex distribution of Carassius gibelio from Shira Lake collected on July-August, 2008. males;
males
females;
50
8+
Annual length
increment, %
40
3+
7+
6+
5+
4+
30
20
10
0
2000
2001
2002
2003
2004
2005
Years
Fig. 3. Back-calculated relative increments of total length of Carassius gibelio from Shira Lake collected on
August 2008. Numbers 8+?..3+ indicate age at capture
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Table 3. The year of hatching, mean total length at capture, mean back-calculated lengths-at-age, standard errors,
and mean annual growth increments of Carassius gibelio from Shira Lake collected in August, 2008. Ten fishes
of each age class were analyzed
Back-calculated lengths at age (mm)
Year
TL at capture
1+
2+
3+
4+
5+
6+
7+
8+
Class
TL
SE
TL
SE
TL
SE
TL
SE
TL
SE
TL
SE
TL
SE
TL
SE
TL
SE
149.0
1.2
165.2
1.8
183.7
1.6
196.6
1.6
218.0
1.6
230.3
1.3
57.2
1.6
58.7
2.1
66.4
2.2
61.1
2.9
53.0
1.5
45.3
1.5
35.0
108.2
1.9
95.2
2.5
97.6
2.8
95.1
3.1
81.1
2.0
74.3
2.2
32.2
149.0
1.2
129.8
2.1
131.1
2.4
124.8
3.0
109.8
3.2
100.1
2.4
28.4
165.2
1.8
155.5
2.3
151.7
2.7
139.9
2.8
125.1
2.8
27.9
183.7
1.6
173.8
2.4
171.0
0.6
155.0
1.9
24.2
196.6
1.6
194.1
1.4
181.7
1.7
25.4
218.0
1.6
208.7
1.5
21.6
2005
2004
2003
2002
2001
2000
Mean TL increment (mm)
230.3
1.3
Table 4. Food components of Carassius gibelio from Lake Shira collected on July and August, 2008. Numbers
indicate weight percentages of a species in a food component
Fish age
Food component
1+
3+
4+
5+
6+
7+
1
2
3
4
5
6
7
Bacillariophyta
1. Navicula sp.
2. Cymbella sp.
39
22
70
48
73
20
80
57
3. Sinedra ulna
8.8
20
18
-
-
20
4. Sinedra acus
6.9
-
14
7
-
23
5. Melosira sp.
1.0
-
-
-
-
-
6. Pinnularia sp.
1.0
-
10
-
-
-
7. Gomphonema olivaceum
0.7
-
-
-
-
-
8. Cocconeis placentula
2.0
-
-
-
-
-
9. Achnanthes sp.
0.3
-
-
-
-
-
10. Eunotia sp.
3.0
-
-
-
-
-
11. Amphora ovalis
2.0
-
-
-
-
-
12. Rhopalodia gibba
2.3
-
5
-
-
-
13. Cyclotella sp.
11
10
5
-
20
-
Chlorophyta
14. Staurastrum sp.
15. Ankistrodesmus sp.
8
50
-
-
-
-
20
-
-
-
-
-
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Table 4. (Continue)
1
2
3
4
5
6
7
-
-
-
-
16. Tetraedon minimum
2
-
17. Scenedesmus quadricauda
10
50
-
-
100
-
18. Scenedesmus acumenatus
5
-
25
100
-
-
19. Pediastrum angulosum
7
-
25
-
-
-
20. Pediastrum tetras
10
-
25
-
-
-
21. Cosmarium sp.
10
-
25
-
-
-
22. Pandorina morum
10
-
-
-
-
-
23. Oedogonium sp.
10
-
-
-
-
-
24. Dictyosphaerium sp.
8
-
-
-
-
-
Cyanophyta
25. Merismopedia tenuis
26. Gomphospheria sp.
79
-
-
-
-
-
21
-
-
-
-
-
100
-
-
-
-
-
10
-
-
-
-
-
90
-
100
-
-
-
100
-
100
-
100
100
-
100
-
100
100
100
-
100
-
100
100
100
10
3
10
10
5
12
10
5
10
10
5
10
50
75
80
50
-
-
20
10
-
30
-
-
10
7
-
-
-
-
-
-
-
-
90
78
Euglenophyta
27. Phacus
Nemathelmintes
Rotatoria
Family Asplanchnidae
28. Asplanchna sp.
Family Branchionidae
29. Keratella sp.
Type Annelida
Class Oligochaeta
30. Setae of Oligochaeta
Type Mollusca
Sub type Conchifera
31. Class Gastropoda
Type Arthropoda
Class Arachnida
Gen. Acarina
32. Family Hydrocarinae
Class Crustacea
Sub class Branchiopoda
Gen. Cladocera
Family Daphniidae
33. Daphnia sp.
Family Bosminidae
34. Bosmina sp.
Family Chydoridae
35. Chydorus sp.
Sub class Copepoda
Gen. Cyclopoida
36. Family Cyclopoidae
37. Gen. Calanoida
Class Malacostraca
Gen. Amphipoda
Family Gammaridae
38. Gammarus lacustris
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Table 4. (Continue)
1
2
Class Insecta
Gen. Diptera
Larvae of Chironomidae
39. Endochironomus sp.
40. Chironomus sp.
3
4
5
6
7
-
10
11
20
15
8
-
80
69
70
60
64
18
41. Polypedilum sp.
-
10
10
-
15
42. Cricotopus sp.
100
-
10
10
10
Gen. Coleoptera
43. Larvae of beetles
100
100
-
100
100
100
44. Remains of terrestrial insects
-
-
100
100
100
100
45. Remains of macrophytes
-
100
100
100
100
100
46. Indeterminable substance
100
100
100
100
100
100
47. Sand
100
100
100
100
100
100
100
90
80
G u tc o n te n ts ,%
70
60
50
40
30
20
10
0
1+
3+
4+
5+
6+
7+
F is ha ge
Bacillariophyta
Chlorophyta
Cyanophyta
Euglen?phyta
Rotatoria
Oligochaeta
Cladocera
Copepoda
Mollusca
Hydrocarinae
Beetles
Chironomidae
Macrophytes
Gammarus
Fig. 4 Mass percentage of gut contents components of Carassius gibelio from Lake Shira collected on JulyAugust 2008. Species compositions of components are given in table 4
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Denis Yu. Rogozin, Marina V. Pulyayevskaya? Growth, Diet and Fatty Acid Composition of Gibel Carp Carassius?
The guts collected from Lake Shira were
relatively evenly distributed by age groups, which
allowed us to reveal differences in the diets of
specimens of different ages (Fig. 4). Obviously,
the diet is not the same throughout the whole
life, the percentage of diet components varies
depending on age (Fig. 4). The guts of 2-yearolds contain mostly algae, with the predominance
of Bacillariophyta; there are also Rotatoria,
Oligochaeta and equal percentage of Cladocera
and Copepoda. At an age of 3+ , the percentage
of small crustaceans and insects increases, while
Gastropoda, Hydrocarina and the remains of
higher aquatic vegetation appear in the diet of
gibel carp. The percentage of algae is small,
while worms are not absent. Chironomidae and
Coleoptera are the dominant food category of
5- and 6-year-old specimens. Small crustaceans,
gastropods, aquatic mites and worms are also
common food for fish of this age group. Older
C. gibelio prefer Amphipoda, Chironomida and
Coleoptera. The percentage of small crustaceans
and worms is insignificant; there are remains of
higher aquatic vegetation and sporadic traces of
algae (Fig. 4).
Fatty acid composition
More than sixty fatty acid (FA) species
were identified in all muscle samples of C.
gibelio from the Shira Lake, and percentage of
43 prominent acids are given in Table 5. The
average level of saturated fatty acids (SFAs)
was 27.8 %. Among them 16:0, 18:0 and 14:0
dominated. The percentage of markers of
bacterial food, odd and branched acids was
4.7 %, among them ai17:0 and i17:0 dominated.
Monounsaturated fatty acids (MUFAs) were in
average 29.8 % (Table 5). Monoenoic acids were
primarily represented by 18:1?9, 16:1?7, 18:1?7,
20:1?9. Polyunsaturated fatty acids (PUFAs)
were in average 39.9 %. Docosahexaenoic acid
(DHA, 22:6?3), eicosapentaenoic acid (EPA,
20:5?3), arachidonic acid (ARA, 20:4?6) and
linoleic acid (LIN, 18:2?6) were the dominant
species among PUFAs. The ratio of ?3/?6 was
2.8. It was shown that value of ?3/?6 depends
on trophic positions of fish e.g., the optimal
?3/?6 ratio in the carnivorous-benthivorous
freshwater fish is 3.8, for carnivorouspiscivorous fish the ratio is 2.6, and for
herbivorous?omnivorous fish is 2.0 (Ahlgren
et al., 2009). Thus, ?3/?6 ratio of Carassius
gibelio from the Shira Lake corresponds with
the ratio for carnivorous-piscivorous fish. The
total FA concentration in muscle samples of C.
gibelio from the Shira Lake was 7.2±0.7 mg/g
wet weight.
We compared FA content and composition
of muscle samples of C. gibelio from saline Lake
Shira and from the freshwater reservoir Bugach
(a small reservoir situated 56°03 N and 92°43 E
in the Bugach river, secondary tributary of the
Yenisei river). FA composition of the freshwater
and saltwater C. gibelio differed significantly
(Table 5). C. gibelio from the Shira Lake had
higher level of MUFA and BFA, but lower levels
of SFA and PUFA in comparison with C. gibelio
from Bugach Reservoir (Table 5). Among SFA
the levels of dominant FAs, 16:0, 18:0 and 17:0
were significantly higher in freshwater fish.
The levels of most fatty acids of MUFA were
significantly higher in saltwater population.
These differences may be the result of several
causes. Firstly, desaturation of 18:0 to 18:1?9
might be more effective in saltwater population.
This result is in good agreement with our previous
data on freshwater and saltwater populations of
Gammarus lacustris (Makhutova et al., 2003).
Secondly, differences in nutrition spectrum
can take place. The acid 16:1?7 is a marker of
diatoms (Claustre et al., 1988/1989; Leveille et
al., 1997; Shin et al., 2000). Fatty acids 20:1?11,
20:1?9, 20:1?7, as well as 22:1?11, 22:1?9 and
22:1?7 which obtained in saltwater fish in traces
# 96 #
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Table 5. Average levels (M, %) of prominent fatty acids in muscles of Carassius gibelio from Shira Lake (n=9)
and Bugach Reservoir (n=6), t ? Student?s test, * significant difference. The total, mg/g wet weight.
12:0
Fatty acids
Shira
M±SE
Bugach
M±SE
t
1
2
3
4
0.11
±
0.02
0.14
±
0.02
1.06
14:0
1.52
±
0.19
1.19
±
0.10
1.54
15:0
0.82
±
0.21
1.17
±
0.07
1.58
16:0
17.69
±
0.31
22.81
±
0.79
6.03*
17:0
0.74
±
0.10
1.57
±
0.11
5.58*
18:0
6.50
±
0.42
8.26
±
0.19
3.82*
20:0
0.23
±
0.03
0.19
±
0.01
1.26
ai15:0
0.22
±
0.03
0.18
±
0.02
1.11
i15:0
0.14
±
0.03
0.01
±
0.003
4.31*
i16:0
0.22
±
0.01
0.17
±
0.01
3.54*
ai17:0
0.79
±
0.04
1.52
±
0.11
6.24*
i17:0
0.30
±
0.01
0.23
±
0.01
4.95*
16:1?9
0.18
±
0.05
0.75
±
0.19
2.90*
16:1?7
6.93
±
0.71
3.24
±
0.15
5.08*
16:1?5
0.38
±
0.02
0.33
±
0.03
1.39
17:1
0.80
±
0.16
0.37
±
0.04
2.61*
18:1?9
13.09
±
0.99
6.94
±
1.30
3.76*
18:1?7
5.68
±
0.17
3.73
±
0.24
6.63*
18:1?5
0.29
±
0.03
0.05
±
0.01
7.59*
20:1?11
0.23
±
0.04
0.09
±
0.01
3.40*
20:1?9
1.07
±
0.07
0.58
±
0.03
6.43*
20:1?7
0.44
±
0.03
0.14
±
0.01
9.49*
16:2?4
0.43
±
0.07
0.04
±
0.003
5.57*
16:3?4
0.26
±
0.05
tr.
16:3?3
0.04
±
0.01
0.01
±
0.004
2.79*
16:4?3
0.06
±
0.01
0.03
±
0.005
2.68*
16:4?1
0.02
±
0.01
tr.
18:2?6
3.43
±
0.40
3.90
±
0.86
0.50
18:2?4
0.17
±
0.01
0.02
±
0.01
10.61*
18:3?6
0.26
±
0.03
0.10
±
0.02
4.44*
18:3?3
1.52
±
0.14
1.58
±
0.06
0.39
18:4?3
0.62
±
0.10
0.27
±
0.02
3.43*
20:2?6
0.34
±
0.03
0.41
±
0.09
0.74
20:3?6
0.48
±
0.03
0.39
±
0.1
0.86
20:4?6
4.19
±
0.27
5.46
±
0.34
2.93*
20:3?3
0.24
±
0.02
0.37
±
0.03
3.61*
20:4?3
0.86
±
0.06
0.56
±
0.02
4.74*
20:5?3
7.87
±
0.32
6.26
±
0.49
2.75*
22:4?6
0.29
±
0.04
0.09
±
0.02
4.47*
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Denis Yu. Rogozin, Marina V. Pulyayevskaya? Growth, Diet and Fatty Acid Composition of Gibel Carp Carassius?
Table 5. (Continue)
1
22:5?6
2
3
0.98
±
0.08
22:4?3
0.02
±
0.01
1.95
22:5?3
2.50
±
0.16
2.38
22:6?3
13.80
±
1.20
21.38
SFA
27.76
±
0.70
35.44
BFA
MUFA
0.26
±
4
0.07
6.77*
±
0.17
-11.33*
±
0.33
0.33
±
1.66
-3.70*
±
1.13
-5.78*
4.69
±
0.49
2.30
±
0.1
4.78*
29.75
±
1.80
16.59
±
1.63
5.42*
PUFA
39.91
±
1.28
45.68
±
1.49
-2.94*
?18-22 ?3 PUFA
27.52
±
1.27
34.76
±
2.11
-2.94*
?18-22 ?6 PUFA
9.97
±
0.58
10.59
±
1.03
-0.52
7.2
±
0.72
4.54
±
0.23
3.52*
Total
are the markers of marine copepods (Mayzaud
et al., 1999; Falk-Peterson et al., 2001; Nelson et
al., 2001; Scott et al., 2002). Fatty acid 18:1?7 is
considered as marker of bacteria. Thus, perhaps
C. gibelio from the Shira Lake fed diatoms and
copepods more selectively or in a large quantity.
Moreover, the level of PUFA markers of diatoms
(16:2?4, 16:3?4, 16:4?1 and 20:5?3) were
significantly higher in saltwater population.
Among C18-22 PUFA the levels of 18:2?4,
18:3?6, 18:4?3, 20:4?3, 20:5?3, 22:4?6 and
22:5?6 were significantly higher in saltwater
fish. A high level of long-chained fatty acids
may be the evidence of prevalence animals in
nutrition of C. gibelio from the Shira Lake. It
should be emphasized that the level of essential,
physiologically important acid, 22:6?3 was in
two times higher in freshwater population.
The total FA concentration in muscles of
C. gibelio from the Shira Lake was in one and
a half times higher than in the fish from the
Bugach Reservoir. A fattier animal contains a lot
of triacylglycerols (TAG), which is characterized
by a high level of SFA, though in our case fat
less population of C. gibelio from the Bugach
Reservoir had significantly higher levels of SFA
and PUFA (Table 5).
Discussion
This paper for the first time refers to reliable
facts indicating presence of ichthyo-fauna in
Shira Lake. Although there is oral evidence that
fishermen caught C. gibelio at the inflow of the
Son River as early as in the 1980-s, we do not
have any evidence referring to an earlier date.
Total absence of C. gibelio in our catches of
May 2009 can be explained by the low water
temperature, whereas the salinity at this period
had the lowest values as compared to the other
dates of sampling.
The length-weight relationship indicates
that population of C. gibelio in Shira Lake is
relatively well-fed. For example, in femaledominated population of C. gibelio from
Chimaditis Lake (Greece) the exponent b was
2.81 for TL (Leonardos et al., 2008) (compare
to range, 3.13-3.07, in our study). As it was
shown by Tsoumani with co-authors (2006), the
exponent b may increase with trophic state of
the lake. In eutrophic lakes these b values were
significantly lower than those in oligotrophic or
mesotrophic lakes (Tsoumani et al., 2006). In
this respect, the value of b in Shira Lake is close
rather to mesotrophic and eutrophic lakes than
to oligotrophic ones. According to coefficient b
# 98 #
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Denis Yu. Rogozin, Marina V. Pulyayevskaya? Growth, Diet and Fatty Acid Composition of Gibel Carp Carassius?
Elevation, m
356
(a)
354
352
350
348
Salinity, ppt
30
(b)
25
20
15
10
1890
1910
1930
1950
1970
1990
2010
Fig. 5. Surface elevation and salinity of Lake Shira (delivered from Rogozin et al., 2010): (a) surface elevation
ASL: ( ) m ) measured by meteoservice (1936-2007); ( rev) ? previously reported (Krivosheev, Khasanov, 1990 );
(b) salinity: ( e sal ) average salinity calculated according to estimates of volume change; ( ly m) ? previously
measured and reported (Krivosheev, Khasanov, 1990; Kalacheva et al., 2002; Parnachev et al., 2002; Rogozin et
al., 2010); ( ) ? upper ) ? upper value of Carassius gibelio tolerance according to literature
values, the females were more ?succulent? than
males (see Results).
Obviously, absence of other fish species
makes gynogenetic reproduction of C.gibelio
in Shira Lake impossible. Almost equal sex
distribution (Fig. 2) indicates sexual reproduction
of C.gibelio in the lake.
We suggest that elevated salinity level has
been keeping gibel carp out of the lake. The
salt concentration in the lake varied inversely
to the lake level (Fig. 5) (Krivosheyev,
Khasanov, 1990; Rogozin et al., 2010). In the
last two decades the trend has been towards
level increase and salinity decrease; in 2007
the historical level maximum occurred, and,
therefore, the historical salinity minimum was
reached, about 14-15 g l-1, measured in the
epilimnion in summer (Fig. 5).
Therefore, we can assume that the fish range
could have expanded throughout the lake in the
recent years, and this trend can continue with the
lake level increase (and salinity decrease) which
might be caused both by increase of climate
humidity and inflow of man-made discharge into
the lake (Krivosheev, Khasanov, 1990). Apart
from the years-long changes, the water salinity
in the upper layers of Shira changes during
a year with minimum in May and early June
(Rogozin et al., 2010). The upper limit of the C.
gibelio?s salinity tolerance is about 15 g l-1; this
species cannot survive for a long time at higher
values (Nico, Schofield, 2006). In steppe lakes of
Western Siberia the Carassius gibelio is known
to withstand the increases of salinity up to 11-14
g l-1 (Yadrenkina, Interesova, 2010). However, the
salt composition of Shira Lake water is dominated
by sulphate-ion (Kalacheva et al., 2002), that
makes difficult to compare the salinity tolerance
of fish with sea water and other reservoirs where
chloride-ion dominates.
Despite many information is available
on occurrence of Carassius gibelio and other
cyprinids in brackish and mildly saline waters
of seas and lakes (Ishitobi et al., 2000; Vetemaa
et al., 2005; Abramenko, 2006; Kanaya et al.,
2009; Yadrenkina et al., 2010), we could not
# 99 #
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Denis Yu. Rogozin, Marina V. Pulyayevskaya? Growth, Diet and Fatty Acid Composition of Gibel Carp Carassius?
fi nd in literature any study of invasion the fishfree saline lake by fish. Obviously, the invasion
of fish-free saline lake is not common event.
Therefore, the opportunity to monitor the early
point of the invasion stage in a saline lake seems
to be very rare. It is not known, however, when
exactly the gibel carp appeared in the lake.
Apparently, the fish could always be present at
the inflow of the river, where the water salinity
is minimal.
However, even the minimal salinity values
registered by us in the lake are not suitable for
normal development of eggs and juveniles of this
species (Nico and Schofield, 2006). Thus, the
spawning place of the gibel carp living in Lake
Shira is not known. Neither is known from what
water bodies the gibel carp can enter the Shira
Lake, what total biomass this species has in the
lake, and what seasonal dynamics this biomass
has, and therefore, to what degree C. gibelio
influences the biomasses of organisms being
its diet components. To answer these questions,
further studies are necessary.
Acknowledgements
We thank Mr. Vladimir Zykov, Mrs.
Anastasia Kharuk, Dr. Yury Barkhatov
(Institute of Biophysics SD RAS), Mr. Maxim
Ousenko and officers of Khakassian National
Park for assistance in field work. We are also very
grateful to Prof Dr. Anatoly Vyshegorodtsev,
Prof. Sergey Chuprov (Siberian Federal
University) and two anonymous reviewers
for valuable comments and advice on article
preparation. This work was partly supported
by the Russian Foundation for Basic Research
and Krasnoyarsk Regional Science Foundation,
Grant No. 09-04-98036, by Siberian Branch
of Russian Academy of Sciences, Integrative
Project No. 95.
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????, ???????, ? ?????????????? ??????
??????????? ?????? Carassius gibelio
? ??????????? ????? ????, ????? ??????
?.?. ????????,?, ?.?. ???????????,
?.?. ?????, ?.?. ?????????, ?.?. ????????????,?
?
???????? ????????? ?? ???
?????? 660036, ??????????, ?????????????
?
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ?????????, 79
? ??????????? ??????????????? ??. ???? (????? ??????, ?????????? ???????)
???????????????? ??????????? ??????????? ?????? Carassius gibelio. ????? ????
?????????? ????????? ? ????? XIX ?., ? ?? ??? ??? ?????????, ??? ?????????? ? ??????
????? ???????????. ? ????????? ?????? ??????????? ??????????????? ??????????????,
?????? ??????? ? ?????????????? ?????? ?????? Carassius gibelio ?????????? ????????,
??????????? ? ??. ???? ? ?????? ????? 2007 ? 2008 ??. ????????? ???? ? ?????? ?????? ????
????? 15 ?/?. ????? ??????? ??????????? ?????? ???????? 1-2 ??? ??????????? ??????????,
??????? ? ????-??????? ?????????. ? ?????? ?? ??????? ?? ???? ??? ? ???? ?????????????
???? ???????, ????? ? ?????????. ??????? ? ???????????? ???????? ???????????? ???? ?
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??????? ?????? ???????????? ??????? ? ?????????? ??. ????. ???? ?? ????????? ??????
????????? ?????? ? ????? ? ???????? ?????????, ????????????? ?????????? ?????? ?????. ?
????????? ??????????? ??????? ??. ???? ???????? ??? ????????, ????????? ??? ??????? ?????
? ????? XIX ?. ??????????????, ????????? ???? ? ????? ????????? ?? ???????????? ????????
?? ???? ?????????????????? ?????.
???????? ?????: ?????????? ??????, ??????????? ?????, ?????? ???????, ?????? ???????.
??? ????????? ???????? ?? ??????????? ???????????? ???????
????????????????? ?????????? (????? ???
NOAA/AVHRR, TERRA/Modis) ?????? ???
??? ????????? ? ?????????? ?????: ??????????, ???????? (??????-????????), ????????????, ????????????, ??????? ????????
???????? ??????? ? ???? ????????????????
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?????????? ? ??????? ???????? (1:50 000 ?
??????). ?? ???????????? ???????? ??? ????????????? ?????????????????? ????????.
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????????? ??????????????? ???. ??? ??????????????? ? ??????????? ??????? ????????? ?????????? ??????????? ??? ? ???????????? ?????.
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???? ????? ????? ??????? ?????? ?? ??????????? ??????? ??????? ? ???????? ????????????????? ??????????.
??????????? ?????? ? ????????? ?????????? ??????? ????????? ??????????? ??????? ????????? ? ????????? ?????????????
??????? ?????? ??????????, ????????? ? ?????????????? ????????????? ? ????? ?????????? ?????????. ?????? ???? ?????????
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Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ?????????? ??????????? ?????????? ?????? ?????? ????????? ????
Satellite Monitoring
of Sayan Mountain Forest Ecosystems
Evgeny I. Ponomarev,
Dilshad M. Ismailova and Dina I. Nazimova
V.N. Sukachev Institute of Forest of Siberian Branch
of Russian Academy of Sciences
50/28 Akademgorodok, Krasnoyarsk, 660036 Russia
New approach to use remote sensing data of low and moderate spatial resolution is suggested for
forest cover mapping and for its seasonal monitoring. The researches of forest structure and dynamics
are conducted on three test sites of the Western and Eastern Sayan mountains since 1960. Satellite
remote data for the territory (NOAA/AVHRR and TERRA/Modis imagery) are available for 1996 ?
2009. After description of the data base the features of different forest zones and formations are shown
in context of their identification and spectral characteristics changing during phenological seasons.
Specification of mountain forest zones with different potential composition is important for forest
management and monitoring on regional and local levels.
Keywords: mountain boreal forest, altitudinal zones, remote sensing imagery, database, seasonal
monitoring, spectral characteristics.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2011 4) 86-103
~~~
??? 597.554.3: 591.9(289)
Growth, Diet and Fatty Acid Composition
of Gibel Carp Carassius Gibelio in Lake Shira,
a Brackish Water Body in Southern Siberia
Denis Yu. Rogozin a,b*,
Marina V. Pulyayevskayab, Ivan V. Zuevb,
Olesya N. Makhutovaa and Andrey G. Degermendzhia,b
a
Institute of Biophysics SB RAS
Akademgorodok, Krasnoyarsk, 660036 Russia
b
Siberian Federal University
79 Svobodny, Krasnoyarsk, 660041 Russia 1
Received 4.03.2011, received in revised form 11.03.2011, accepted 18.03.2011
For the first time the presence of gibel carp Carassius gibelio has been detected in the brackish
meromictic Lake Shira (Southern Siberia, Republic of Khakassia). Lake Shira has been continuously
studied since the late 19th century, and it has been considered so far that there is no ichthyofauna in
the lake. This work investigates the growth and population characteristics, as well as food spectra
and fatty acid composition of C. gibelio specimen at different ages that were fished out in the summers
of 2007 and 2008. The water salinity at the fishing point was 15 g l-1. The food spectrum of one- and
two-year old C. gibelio was dominated by diatomic, green and blue-green algae, while four- to sixyear old species had an increased share of copepods, beetles and chironomids in their food spectrum,
and beginning from the age of six years the amphipods Gammarus lacustris consisted significant part
of gibel carp diet. Thus, it was shown that C. gibelio is a carnivore with regard to the population of
amphipods Gammarus lacustris, which have been regarded so far as the upper link in the trophic
cascade of Lake Shira. One of the possible reasons for appearance of fish is the decreased water salinity
caused by rising of the lake level. In the recent years, the level of Lake Shira exceeded all the values
that had been registered since the late 19th century, and the lake?s salinity decreased correspondingly
to its smallest value for the whole period of observation.
Keywords: Carassius gibelio, brackish lake, food spectrum, fatty acids.
Introduction
Fish introductions are amongst the greatest
threats to freshwater biodiversity (Gherardi,
2010), and one of the most invasive freshwater
*
1
fish species is the gibel carp Carassius gibelio
Bloch (Szczerbowski, 2002; Luskova et al.,
2010) which has been recently found to have
established a population in the brackish Lake
Corresponding author E-mail address: rogozin@ibp.ru
© Siberian Federal University. All rights reserved
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Denis Yu. Rogozin, Marina V. Pulyayevskaya? Growth, Diet and Fatty Acid Composition of Gibel Carp Carassius?
Shira previously reported as a fish-less lake
(Degermendzhy, Gulati, 2002; Degermendzhy
et al., 2010). Gibel carp Carassius gibelio, being
a stenohaline species, is rather often observed
in brackish water, for example, in the littoral
zones of the Black and Baltic seas (Abramenko,
2006; Vetemaa et al., 2005), in the delta of the
Ob (Zhadin et al., 1961) etc. Lake Shira has
been studied for decades by many researchers
(Krivosheyev, Khasanov, 1990; Degermendzhy,
Gulati, 2002; and many others), but it has been
considered until now that there is no permanent
ichtyofauna in the lake (Degermendzhy, Gulati,
2002; Degermendzhy et al., 2010). In summer,
the salinity of the epilimnion water was about
15 g l-1 during last decade (Rogozin et al., 2010),
however, lower salinity was observed near
estuary of the river Son incoming to the lake.
According to the oral evidence of local residents,
gibel carp has been regularly observed in the
lake lately. However, there have been no reliable
facts proving this information so far. In 2007, we
made trial sampling that confirmed the presence
of gibel carp in the lake. Since the fish population
may provide considerable impact on previously
fishless ecosystem, it seems to be important to
study the main characteristics and ecology of this
population. At the first step, we tried to analyze
the feeding spectrum, growth characteristics
and fatty acids composition of gibel carp in the
lake. Fish species, from different ecosystems, are
known to differ in their fatty acids composition;
therefore, studies of fatty acids contents of
diverse fish from various locations are of great
importance for revealing their potential value
as sources of the essential ?3 acids for human
nutrition (Ahlgren et al., 1994; Gladyshev et al.,
2009). Polyunsaturated fatty acids (PUFAs) of the
?3 family became known as key dietary nutrients
for preventing mental, neural and, especially,
cardiovascular diseases of human (Arts et al.,
2001; Garg et al., 2006). Besides, biochemical
analysis of fatty acid composition is widely used
to trace trophic interactions between hydrobionts
and to reveal food assimilation (Desvilettes et al.,
1997; Sushchik et al., 2003). Therefore, the aims
of present work were to provide the first evidence
of presence of fish in Shira Lake, to analyze the
diet of the fish and to give some basic information
about growth characteristics and fatty acids
composition.
Materials and Methods
Lake description
Lake Shira (90.11 E, 54.30 N) is located in
the northern part of the Republic of Khakassia,
South Siberia (Russian Federation). This elliptical
(9.3 Ч 5.3 km) lake has an area of 35.9 km2 and
maximum depth of 23.8 m (2007-2010). The main
inflow is from the Son River that provides about
42 % of fresh water supply to the lake; other
sources of water are precipitation and seepage
water (Parnachev et al., 2002). The lake has no
outflow. The salinity of lake water during summer
stratification is about 14-15 g l-1 in epilimnion
(upper 5-6 m of water column) and 18-19 g l-1 in
deeper waters. The detailed description of the
lake is given elsewhere (Degermendzhy et al.,
2002; Parnachev et al., 2002; Kalacheva et al.,
2002; Rogozin et al., 2010). The surface elevation
of Lake Shira has noticeably changed over the
whole period of observation. The lake?s salinity
varied inversely with changes in water volume
(Krivosheev, Khasanov, 1990; Rogozin et al.,
2010). During the period from 1920-s to 1930-s
the lake?s level decreased by 7 m, which was
caused by the decrease in the total amount of
precipitation in the region. In this period, salinity
reached the maximum of 27 g l-1 in 1926. By the
present moment, the lake level has exceeded all
the values registered since the late 19th century,
and the salinity has dropped, respectively, to its
lowest value for the whole period of observation
(Rogozin et al., 2010).
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Denis Yu. Rogozin, Marina V. Pulyayevskaya? Growth, Diet and Fatty Acid Composition of Gibel Carp Carassius?
Sampling and water measurements
Fish sampling was carried out in 2007-2009
in Point ? (54.28.767 N, 90.14.818 E), in several
points of Area ?, and in point ? (54.29.053 N;
90.16.352 E) (Fig.1). Exact coordinates of Area
B point were not defi ned, and the location of
Area B was estimated visually on the basis of
landmarks. The dates of fishing are given on
Fig.1. Fish were captured with gill nets of various
dimensions and mesh sizes, complemented for
young specimens by drag-nets, bottle and can
traps as well as rod and line angling. In all
points the nets were set in the evening, along
macrophyte beds, at a depth of approximately
1.5 m not far from the shore; the nets were
checked early in the morning next day.
The temperature and specific conductance
of water in points ? and ? (Fig.1) were measured
with YSI 6600 (Yellow Springs, Ohio, USA)
submersible profiler. The salinity was calculated
as quadratic approximation of the dependence of
ash content upon specific conductance measured
for the Lake Shira water in laboratory:
S = -0.087 cond2 + 4.4403cond ? 37.16,
R2 = 0.95, n = 19,
where S is the salinity determined as ash content
(g l-1), Cond is specific conductance at 25°C of
the lake water samples (mS cm-1). Conductivity
sensors were calibrated against 3M KCl (YSI).
Oxygen electrodes were calibrated against
Winkler titration of the Lake Shira water with
Aquamerck kit (Merck, Germany)
Back-calculated growth,
weight?length relationship
and Fulton?s condition factor
0.01 g on electronic balances and measured to the
nearest millimeter. Scale samples for age and
growth analysis were collected from each fish.
Scales were taken above the lateral line dorsal
to the tip of depressed left pectoral fin. Growth
histories of individual fish were determined by
aging and back-calculation of total lengths (TL)
at previous ages from scales (Pierce et al., 1996).
Twelve to fifteen scales per fish were cleaned and
mounted between glass slides. All scales on slides
were viewed when ages were assigned to fish, and
a single reader did all aging. Anterior radii and
interannular distances were measured to nearest
0.01 mm by using dissecting microscope (25Ч
magnification). These radii are hereafter referred
to as scale lengths. Replicate measurements were
then averaged for each fish, providing the precise
estimates of scale lengths for back-calculations.
With the data on individual TL and mean scale
length, we then back-calculated TL at previous
ages using the Fraser-Lee formula (Pierce et al.,
1996).
Weight?length relationship and Fulton?s
condition factor were calculated from the
expressions:
(1) W = a TLb
(2) K= 100 W/SL3,
Where W is the whole body weight in grams,
TL the total length, SL is the standard length in
centimeters; a and b are the regression constants;
the factor 100 is used to bring K close to unity
(Froese, 2006).
Diet analysis
The growth history was determined for 200
fish collected from the end of July to the beginning
of August in Area ? (Fig.1). After capture all the
fish were individually weighted (wet) to the nearest
Diet was assessed on the basis of gut
contents determined by dissection of 32 fish
collected from the end of July to the beginning of
August in Area ? (Fig.1). All fish samples were
weighted and measured as described above,
and their guts were removed immediately, and
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Denis Yu. Rogozin, Marina V. Pulyayevskaya? Growth, Diet and Fatty Acid Composition of Gibel Carp Carassius?
stored in formalin (4%) until the contents were
analysed. The analysis of digestive tracts was
performed in accordance with the standard
method (Metodicheskoye rukovodstvo, 1974;
Hyslop, 1980). Animal food items were
identified to the lowest possible taxon. The diet
was analyzed for the percentage composition
by mass.
Analysis of fatty-acid composition
Detailed description of the analysis is given
elsewhere (Makhutova et al., 2003; Sushchik et al.,
2006). Briefly, lipids from samples were extracted
with chloroform:methanol (2:1, v/v) three times
simultaneously with mechanical homogenization
of the tissues with glass beads. Before extraction,
a fixed volume of an internal standard solution
(19:0) was added to the samples. Methyl esters of
fatty acids (FAMEs) were prepared in a mixture
of methanol: sulphuric acid (50:1, v/v) at 90 oC
for 2 h. FAMEs were then analysed using a GC?
MS (model GCD Plus, Hewlett Packard, USA)
equipped with a 30 m long Ч 0.32 mm internal
diameter capillary column HP-FFAP. FAMEs
peaks were identified by their mass spectra,
comparing to those in the data base (Hewlett?
Packard, USA) and to those of available authentic
standards (Sigma, USA). To determine double
bond positions in monoenoic and polyenoic acids,
GC?MS of dimethyloxazoline derivatives of FA
were used.
ever documented proof of the presence of fish
in Lake Shira. The age was determined for six
specimens of this capture (Table 1).
On August 9 2008, in Point ? the water
temperature was 23 °?, and the salinity was
14.7 g l-1. In this date, 130 two- to four-yearold specimens and 68 five- to eight-year-old
specimens were captured using nets with a mesh
size of 25 mm and 40 mm, respectively. The
specimens of this capture were analyzed only for
fatty-acid composition.
In area B (Fig.1), 200 specimens of Carassius
gibelio of different sizes and ages were captured
from the end of July to the beginning of August
2008; the water temperature and salinity were
not measured. The specimens captured in 2008
in Area ? included fish aged from 1+ to 15+ years
(Fig.2, Table 1), with the predominance of 7-yearold specimens (26.5 %). Fish aged 5+ and 7+
make up a large part of the captured total ? 15.5
and 18.0 %, respectively, of the total number of
captured specimens. There were few specimens
older than 8 years in catches. Almost in all age
groups the sex ratio was practically 1:1. It was
only in the 4+ and 7+ age groups that males
dominated. The body length of the fish in this
sample was from 54 mm to 460 mm, and the body
mass was from 2.47 to 1358 g.
In May 2009, no specimens were caught in
points A and C (Fig.1). The water temperature at
that time varied within the range of 10.7-13.2 ??,
and the salinity was 13.7 ? 14.5 g l-1.
Results
Fishing results
On July 24 2007, in the Point ? (Fig. 1)
the water temperature was 24 °?, and salinity
was 15.1 g l-1. The salinity in the central part
of the lake at this period was 15 g l-1, i.e. it was
practically the same as in the fishing point. The
captured fish included 33 specimens of 2- and
3-year-old Carassius gibelio, and one 7-year-old
specimen of the same species. This was the first
Weight?length relationship
and Fulton?s condition factor
The length-weight relationship calculated
for total length data (Table 1), was
W = 0.012 TL3.134, (R 2 = 0.995, n= 124) for
females,
W = 0.015 TL3.068, (R 2 = 0.978, n= 76) for males.
The relationship between TL and SL was
SL = 0.786 TL + 0.405, (R 2 = 0.998, n= 206).
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N
S
1
C
A
B
Son River
1 km
Fig. 1. Fish sampling sites at Shira Lake in 2007-2009: (A) ? 24 July 2007, 9 August 2008 and 30 May 2009;
Area (B) ? 21-30 July and 3-5 August 2008; (C) ? 30 May 2009; (1) ? Town Zhemchouzhnyi and spa resort ?Lake
Shira?
Table 1. Length and mass of Carassius gibelio from Lake Shira collected on July, 2007, and July-August, 2008.
M ? total mass, m ? mass without entrails
Age
TL±SE (cm)
SL±SE (cm)
M±SE (g)
m±SE (g)
n
2007
2+
14.1 ± 0.3
10.2 ± 0.5
42.7 ± 4.6
35.3 ± 3.5
3
3+
14.8 ± 0.6
11.8 ± 0.3
61.8 ± 2.1
48.9 ± 1.6
2
7+
24.0
17.3
177
133
1
1+
7.3 ± 1.7
5.5 ± 1.3
6.4 ± 0.4
5.1 ± 0.3
34
2008
2+
-
-
-
-
0
3+
14.9 ± 1.1
12.0 ± 1.0
54.4 ± 1.4
47.0 ± 1.3
21
4+
16.5 ± 1.8
13.5 ± 2.0
83.1 ± 3.0
68.8 ± 2.8
10
5+
18.4 ± 0.8
14.8 ± 1.2
114.1 ± 2.5
92.0 ± 2.1
31
6+
19.9 ± 0.7
16.1 ± 0.9
149.2 ± 2.0
122.7 ± 2.0
54
7+
21.8 ± 0.8
17.6 ± 0.8
182.3 ± 4.0
147.3 ± 3.8
36
8+
23.0 ± 1.3
18.6 ± 1.8
210.7 ± 5.3
170.4 ± 5.2
10
9+
25.2 ± 1.5
20.8 ± 2.0
308.5 ± 1.8
265.3 ± 3.3
2
10+
26.0
21.2
301
264
1
15+
46.0
36.0
1358
1108
1
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Denis Yu. Rogozin, Marina V. Pulyayevskaya? Growth, Diet and Fatty Acid Composition of Gibel Carp Carassius?
Table 2. Fulton?s condition index (Kf ) of Carassius gibelio collected from Lake Shira in July and August, 2008
Males
Females
Age
Kf
n
Kf
n
1+
3.59 ± 0.03
30
3.79 ± 0.11
4
2+
-
0
-
0
3+
3.13 ± 0.05
13
3.22 ± 0.12
8
4+
-
0
3.41 ± 0.11
10
5+
3.33 ± 0.08
19
3.67 ± 0.08
12
6+
3.51 ± 0.04
33
3.51 ± 0.08
21
7+
3.37 ± 0.11
18
3.29 ± 0.10
18
8+
3.15 ± 0.09
8
3.6 ± 0.01
2
9+
3.43 ± 0.08
2
-
0
10+
-
0
3.16
1
15+
2.91
1
-
0
The Fulton?s condition index (Kf ) undergoes
minor changes in specimens belonging to
different age groups. The (Kf ) varied from 2.91 to
3.79 (Table 2). The (Kf ) of males and females was
almost equal, except the 5+ and 8+ age groups,
in which males were noticeably better-fed than
females (Table 2).
Growth history
Generally, the relative length increment
estimated from scales decreased with the age (Fig.3,
Table 3). The back-calculated patterns of length at
the age were similar among six generations, and
the year-to-year differences among the fish of equal
age-class were not significant, except youngest
group of 3+ (Fig. 3, Table 3). The relative length
increment of 3+ fish in 2005 was significantly
higher than for other generations at this age (Fig. 3).
However, this may be explained by shortcoming of
back-calculation method too (Pierce et al., 1996;
Martinson et al., 2000).
Analysis of diet
The diet of Carassius gibelio in Lake Shira
is rather diverse; it includes 4 divisions of algae,
9 groups of water invertebrates, remains of
terrestrial insects and higher aquatic vegetation
(Table 4, Fig. 4). In addition, the guts of fish in all
age groups contained sand which was obviously
consumed with food.
The largest number of diatoms was found at
the inflow of the River Son. These are diatoms
that are preferred by juvenile fish. The algal flora
of the lake typically includes algae of Pyrrophyta
division, but they were not present in the guts of
the caught gibel carps. The cyanobacteria of the
genera Spirulina, Oscillatoria, and Anabaena
dominating in the littoral zone of the lake were
not present in gut contents.
The zooplankton of Lake Shira is
characterized by a relatively small diversity of
species. The dominating species are Copepoda
and Rotatoria (Degermendzhy, Gulati, 2002)
which are present in the diet of gibel carp. Several
species of Cladocera are found in the south-eastern
part of the lake (at the inflow of the River Son).
Though their population size and biomass are
low, these animals are the most favorite delicacy
of gibel carp among all crustaceans. Benthic
animals also display little diversity, but they
constitute a considerable part of the C. gibelio?s
diet (Table 4, Fig 4).
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35
30
Fish number
25
20
15
10
5
0
1+
2+
3+
4+
5+
6+
7+
8+
9+
10+ 11+ 12+ 13+ 14+ 15+
Age, years
Fig. 2 Age and sex distribution of Carassius gibelio from Shira Lake collected on July-August, 2008. males;
males
females;
50
8+
Annual length
increment, %
40
3+
7+
6+
5+
4+
30
20
10
0
2000
2001
2002
2003
2004
2005
Years
Fig. 3. Back-calculated relative increments of total length of Carassius gibelio from Shira Lake collected on
August 2008. Numbers 8+?..3+ indicate age at capture
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Table 3. The year of hatching, mean total length at capture, mean back-calculated lengths-at-age, standard errors,
and mean annual growth increments of Carassius gibelio from Shira Lake collected in August, 2008. Ten fishes
of each age class were analyzed
Back-calculated lengths at age (mm)
Year
TL at capture
1+
2+
3+
4+
5+
6+
7+
8+
Class
TL
SE
TL
SE
TL
SE
TL
SE
TL
SE
TL
SE
TL
SE
TL
SE
TL
SE
149.0
1.2
165.2
1.8
183.7
1.6
196.6
1.6
218.0
1.6
230.3
1.3
57.2
1.6
58.7
2.1
66.4
2.2
61.1
2.9
53.0
1.5
45.3
1.5
35.0
108.2
1.9
95.2
2.5
97.6
2.8
95.1
3.1
81.1
2.0
74.3
2.2
32.2
149.0
1.2
129.8
2.1
131.1
2.4
124.8
3.0
109.8
3.2
100.1
2.4
28.4
165.2
1.8
155.5
2.3
151.7
2.7
139.9
2.8
125.1
2.8
27.9
183.7
1.6
173.8
2.4
171.0
0.6
155.0
1.9
24.2
196.6
1.6
194.1
1.4
181.7
1.7
25.4
218.0
1.6
208.7
1.5
21.6
2005
2004
2003
2002
2001
2000
Mean TL increment (mm)
230.3
1.3
Table 4. Food components of Carassius gibelio from Lake Shira collected on July and August, 2008. Numbers
indicate weight percentage
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