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149.Журнал Сибирского федерального университета. Сер. Биология №2 2013

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Журнал Сибирского федерального университета
2013
Journal of Siberian Federal University
6 (2)
Биология
Biology
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Editorial Advisory Board
Chairman:
Eugene A. Vaganov
Members:
Josef J. Gitelzon
Vasily F. Shabanov
Andrey G. Degermendzhy
Valery L. Mironov
Gennady L. Pashkov
Vladimir V. Shaidurov
Vladimir V. Zuev
Editorial Board:
Editor-in-Chief:
Mikhail I. Gladyshev
Founding Editor:
Vladimir I. Kolmakov
Managing Editor:
Olga F. Alexandrova
Executive Editor for Biology:
Elena S. Kravchuk
CONTENTS / СОДЕРЖАНИЕ
Vera Yu. Kovaleva,
Vadim M. Efimov and Yuri N. Litvinov
Directional Asymmetry of Morphological Traits During Postnatal
Ontogeny in Root Vole Microtus oeconomus Pall. (Rodentia,
Cricetidae)
? 115 ?
Е.П. Турпаева, А.К. Райский
l%!?*,? C=3*, !%?= Colossendeis (Colossendeidae,
Pycnogonida) ?%! r.?????= , C!,???=?,. =*"=2%!,L
? 130 ?
А.А. Савченко, К.П. Базарин
q%?2% ?,? =*2,"?%?2, m`d- , m`dt-?=",?,?/.
???,?!%???=? " ??L2!%-,???/. ?!=?3?%?,2=. 3 ?C%!2????%"
" ?,?=?,*? 2!??,!%"%??%?% ?,*?=
? 151 ?
В.С. Бондарь, А.П. Пузырь,
А.В. Горева, А.Е. Буров
j%??2!3,!%"=?,? *%?C%?,?,%??/. ?=2?!,=?%" ?= %??%"?
?=?%=??=?%" , !=?!3?=??/. K,%C%?,??!%"
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П.А. Шестерня, А.С. Сергеева,
С.Ю. Никулина, В.А.Шульман
k%*3? 9!21.3 $ ????2,???*,L C!??,*2%! 2 ???2, *%!%?=!?%?%
=2?!%?*??!%?= 3 ?3??,?
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Editorial board for Biology:
Elena Kravchuk ? Series Editor, Institute of Biophysics,
Russian Academy of Sciences, Siberian Branch,
Russia
Nadezhda Sushchik ? Institute of Biophysics, Russian
Academy of Sciences, Siberian Branch, Russia
Sergey Bartsev ? Institute of Biophysics, Russian
Academy of Sciences, Siberian Branch, Russia
Alexander Bolsunovsky ? Institute of Biophysics,
Russian Academy of Sciences, Siberian Branch,
Russia
Marc d?Alarcao ? Tufts University, USA
Nicolai Gaevsky ? Siberian Federal University, Russia
Woodland J. Hastings ? Harvard University, USA
Malcolm Hughes ? the University of Arizona, USA
Takayoshi Koike ? Hokkaido University, Japan
Valentina Kratasyuk ? Siberian Federal University;
Institute of Biophysics, Russian Academy of
Sciences, Siberian Branch, Russia
Elena Muratova ? Institute of Forest, Russian Academy
of Sciences, Siberian Branch, Russia
Akira Osawa ? Kyoto University, Japan
Frank B. Salisbury ? NASA, USA
Ernst-Detlef Shulze ? Max Planck Institute of
Biogeochemistry, Germany
Tatyana Volova ? Siberian Federal University; Institute
of Biophysics, Russian Academy of Sciences,
Siberian Branch, Russia
Eugene Vysotsky ? Institute of Biophysics, Russian
Academy of Sciences, Siberian Branch, Russia
Egor Zadereev ? Institute of Biophysics, Russian
Academy of Sciences, Siberian Branch, Russia
Liliana Zalizniak ? RMIT University, Australia
Ramesh Gulati ? NIOO/Centre of Limnology, The
Netherlands
Mikhail Karpinsky ? Russian Federal Research Institute
of Fisheries and Oceanography, Russia
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Г.Т. Омурова, В.В. Баринов,
О.В. Кардаш, В.С. Мыглан
r?2=?%"???,? "!????, ?2!%,2????2"= (C?!??2!%L*,)
m=?/??*%?% ?%!%?*=: ????!%.!%?%?%?,???*,L
=?C?*2
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Е.Г. Крылова
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(Asteraceae) * ??L?2",? ?3??-=2= ?,?*= ?=
?=?=???/. .2=C=. %?2%?????=
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Т.Т. Ефремова, С.П. Ефремов
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K?!??? *%"
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Journal of Siberian Federal University. Biology 2 (2013 6) 115-129
~~~
??? 574.3, 591.3
Directional Asymmetry of Morphological
Traits During Postnatal Ontogeny
in Root Vole Microtus oeconomus Pall.
(Rodentia, Cricetidae)
Vera Yu. Kovalevaa*,
Vadim M. Efimova,b,c and Yuri N. Litvinova
a
Institute of Systematics and Ecology of Animals SB RAS,
11 Frunze Str., Novosibirsk, Russia 630091
b
Institute of Cytology and Genetics SB RAS,
10 Pr. Lavrentyeva, Novosibirsk, Russia 630090
c
Tomsk State University,
36 Pr. Lenina, Tomsk, Russia 634050
Received 05.11.2012, received in revised form 22.11.2012, accepted 19.05.2013
We analyzed Microtus oeconomus material collected for 25 years (1982?1992, 1994?2005,
2009, 2010). Directional asymmetry (DA) was demonstrated for teeth morphotypes, adrenal and
kidney weights, and for length of the hind paw as well. DA may be formed both at early and
later development stages and is not dependent on environmental factors and population density
dynamics. Although each trait has its own dynamics DA in ontogeny, usually DA increases in
the period of sexual maturation. Uncoordinated development of the sides in this period seems
to associate with an increase in intensity of developmental processes. Our study argues that
sexual dimorphism of continuous traits appears at the later developmental stages. This finding
is consistent with Geodakian?s evolutionary theory of sex. Evolutionary significance of the DA
is to increase both short-term individual fitness and long-term population adaptive capacity.
Developmental approach to the study of morphological and functional asymmetries in animals
opens new possibilities for the study of evolutionary processes.
Keywords: Microtus oeconomus, directional asymmetry, teeth morphotypes, adrenal and kidney
weights, paw lengths, environmental conditions, population density dynamics.
*
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: vkova@ngs.ru
# 115 #
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Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
Introduction
In recent decades, the study of asymmetry
in zoology has been reduced to fluctuating
asymmetry as an indicator of stability of
individual development and/or quality of the
natural environment (Zakharov, Graham 1992;
Parsons 1992; Palmer, Strobeck, 1992, 2003;
Clarke, 1993; Leung et al., 2000; Zakharov, 2003;
Van Dongen, 2006). Directional asymmetry (DA)
and antisymmetry were more often regarded
as nuisance safer to omit from estimation of
fluctuating asymmetry (Graham et al., 1998;
Stige et al., 2006). Nevertheless, DA of bilateral
traits is widespread in nature, most likely inherent
in most animal species (Levin, Palmer, 2007;
Palmer, 2009).
The impetus for rapid research in the field
of DA came from the finding of asymmetrically
expressing genes in early development, such as
Shh and Nodal in chicken (Levin et al., 1995) and
Nodal and Lefty1 in mice (Collignon et al., 1996;
Lowe et al., 1996; Meno et al., 1998). It is known
that numerous genes are involved in patterning
left-right visceral asymmetry (Shiratori,
Hamada, 2006). That DA has a genetic basis is a
fact beyond question. It is of relevance that early
ontogeny only, when the primordial structures
are laid down, have been so far studied (LopezGracia, Ros, 2007).
It is known that there are genes and their
aggregates that are differentially expressed at
particular developmental stages (DS) (Zahn et
al., 2007). Therefore, we can assume that it is true
for DA also, i.e. leading to the DA corresponding
gene network can work at each late DS. In wild
populations, such networks must pass through
natural selection and their effects should be
manifested at the phenotypic level. Hence, the
aim of our research is to study both discrete and
continuous traits bilateral DA depending on sex,
DS, environmental conditions and population
density dynamics in wild populations of animals.
Materials and Methods
Data collection
and morphometric measurements
This study was performed in Gorno-Altai
region, at the northern shore of the lake Teletskoye,
in the site where the Institute?s biological station
is located (51°47?34? N, 87°16?33? E). Voles
were wild-caught in the vicinities of the station,
in terraces around the lake. Many years of
observations on the dynamics of a community
of wild mouselike rodents inhabiting near the
station showed that the root vole (Microtus
oeconomus) and the northern red-backed vole
(Myodes rutilus = Clethrionomys rutilus) prevail
among vole species (Litvinov et al., 2006).
Here we analyzed the root vole material
collected for 25 years (1982?1992, 1994?2005,
2009, 2010). Root vole populations fluctuate in
cycles with a 3?4 year periodicity (Fig. 1A). In
addition, there is a long-term trend of population
density, possibly associated with an increase in
average annual temperature (Fig. 1B).
Specimens were analyzed for bilateral
morphotypic variation in first lower molar M1
(N=707) and the third upper molar M3 (N=709)
teeth. To discriminate between different
morphotypes, we used the number of closed
triangles on chewing surface and the shape
complexity of the labial and lingual sides of the
anterior unpaired loop of M1 and talon of M3
(Pozdnyakov, 1993; Kovaleva et al., 2002). On
this basis, morphotype complexity (MC) was
calculated for each of the morphotypes (Fig. 2).
In addition, morphotype complexity asymmetry
dR-L was calculated for each of the specimens as
follow (Pozdnyakov, 2010):
dR-L = MC(right) ? MC(left)
Measurements of metric traits were taken
exclusively by one of the authors (VYuK). The
excised left and right kidneys and adrenals were
# 116 #
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Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
Fig. 1. A) Microtus oeconomus relative abundance dynamic (per 100 trap-nights, lg); B) Average annual
temperature dynamic (Ongudai meteorological station), gray curve ? distance weighted least squares fitting
L1
L2
L3
L4
L5
L6
L7
L8
24
24
25
25
26
26
26
27
M1
L1
L2
L3
L4
L5
L6
L7
L8
L9
L10
L11
11
12
12
12
13
13
13
14
14
14
15
M3
Fig. 2. Microtus oeconomus morphotypic variation in chewing surface of the first lower (M1) and the third upper
molar (M3). Morphotype complexity values are given under each morphotype designation. (Outlines are made
by A.A. Pozdnyakov)
weighed with a torsion balance to a precision of
1 mg. It is very important to reduce measurement
errors in asymmetry studies, since the difference
between the bilateral traits is usually small
(Palmer, Strobeck, 1986; Palmer, 1994; Merilд,
Bjцrklund, 1995). For this reason, the lengths
of the external morphometric traits, the front
and hind paw, were measured 2-3 times with a
caliper 0.1 mm to reduce measurement errors. If
the difference between the measures was larger
than 0.3 mm, measurements were repeated. The
median, i.e. the non-parametric point estimate,
was chosen.
The relative age of voles was estimated
according to the degree of physiological
maturation and the data on body length and mass
by the standard methods (Bashenina, 1953; Larina,
Lapshov, 1974). According to this, all voles were
divided into four age groups: immature, juveniles
and subadults, and mature, adults and senescents.
# 117 #
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Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
Males and females were analyzed separately.
Morphotypes variability showed no particular
relation to age and sex, therefore we considered
all sets as a pooled sample containing more than
700 molar pairs.
Statistical analysis
The correlation coefficient between dR-L for
M1 and M3 was calculated. Then, we calculated
contingency tables between right-side and leftside both M1 and M3 morphotypes. Morphotypes
observed fewer than 8 times, were excluded from
further analysis.
All metric traits were log transformed
(Graham et al., 2003; Palmer, Strobeck, 2003).
The significance of the presence of DA in every
sample was tested using Student?s paired t-test
(Palmer, 1994).
The following values were assigned to
each specimen to calculate the correlation with
DS, environmental conditions and population
parameters: Year ? trapping year; Trapping
date ? number of calendar days from the 1th May
of the trapping year to the trapping day; Sexual
maturity ? 0 for immature voles, 1 for mature
ones; Abundance dynamic (AD): AD ? number
of captures per 100 trap-nights in trapping year,
lg; ADdev ? AD deviation from regression line;
ADabs(dev) ? absolute deviation from regression
line; TMay, TJun, TJul, TAug ? average monthly
temperatures in May, June, July and August,
accordingly. We included the ADdev and ADabs(dev)
parameters in the calculation in order to take into
account the possible dependence between AD
deviations from population norm in the direction
of higher or lower density and DA. The population
norm was itself increased during the period of
our studies (Fig. 1).
DA, although it is an individual property,
can only be estimated for a group of individuals,
therefore either age-independent qualitative
traits or homogeneous samples composed of
size-age matched individuals were used. In the
case of quantitative traits, samples collected from
natural populations are, as a rule, heterogeneous.
Clearly, then, the difference between left and
right sides of traits depends on size, which,
as a rule, increases in the course of individual
development. It is a convention to reduce sizeage heterogeneity by transformation of metric
traits to natural logarithms (Graham et al., 1998;
Graham et al., 2003; Palmer, Strobeck, 2003). A
Box-Whiskers plot was applied to estimate DA
variability within each age group for continuous
traits. Kruskal-Wallis test and ANOVA (F
test) were used to evaluate significance of DA
dynamic.
Results
First, we investigated the bilateral asymmetry
of discrete traits. Asymmetry of shape of M1 and
M3 was the most obvious among discrete traits.
No correlation between dR-L for M1 and M3 was
found (r=0.0253; n=711).
As expected, most of morphotype pairs
turned out to be symmetrical, with asymmetrical
pairs accounting for 29 % and 25 % of the total
number of pairs of M1 and M3, respectively
(Table 1, 2). Absolute value of dR-L, |dR-L|, did not
usually exceed 1, sometimes reaching 2 (n=10 for
M1 and n=13 for M3) (Fig. 3A, B).
The morphotypes in the contingency tables
are ordered according to their complexity.
More complex morphotypes of the M1 and M3
significantly developed more often on the right
side (Fig. 3A, B). Consequently, there was a DA
in morphotype complexity.
Three of the four investigated continuous
traits showed a DA (Table 3), which did not depend
on environmental temperatures, population
dynamics and trapping seasons for both sexes
unlike an individual size (Table 4). Nevertheless,
asymmetry can depend not only on trait size as
such, but also on the DS of an individual.
# 118 #
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Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
Table 1. Contingency table between right- and left-sides morphotypes M1 of Microtus oeconomus voles in
Teletskaya population. The morphotypes (1?8) are ordered according to morphotype complexity (MC).
Frequencies of symmetrical morphotype pairs are along the diagonal (n=502)
MC (M1)
24
24
25
25
26
26
26
27
L1
L2
L3
L4
L5
L6
L7
L8
Total
R1
190
0
3
33
0
0
1
0
227
R2
0
3
0
0
0
0
0
0
3
R3
12
0
18
2
3
0
0
0
35
R4
85
2
1
244
1
0
4
0
337
R5
4
0
9
17
25
0
3
1
59
R6
2
0
0
5
0
4
0
0
11
R7
2
0
0
11
1
0
16
0
30
R8
0
0
0
1
1
0
1
2
5
Total
295
5
31
313
31
4
25
3
707
Table 2. Contingency table between right- and left-sides morphotypes M3 of Microtus oeconomus voles in
Teletskaya population. The morphotypes (1?11) are ordered according to morphotype complexity (MC) (11?15).
Frequencies of symmetrical morphotype pairs are along the diagonal (n=530)
MC
(M3)
11
12
12
12
13
13
13
14
14
14
15
L1
L2
L3
L4
L5
L6
L7
L8
L9
L10
L11
Total
R1
14
1
2
5
0
0
0
0
0
0
0
22
R2
0
4
0
0
0
1
1
0
0
0
0
6
R3
1
0
4
1
1
0
0
0
0
0
0
7
R4
7
0
0
312
0
8
24
4
1
1
0
357
R5
0
1
0
0
4
0
0
1
0
0
0
6
R6
0
1
0
16
0
38
2
11
0
0
0
68
R7
1
0
0
48
1
3
98
4
2
0
0
157
R8
0
0
0
5
0
10
8
45
0
0
2
70
R9
0
0
0
0
0
1
1
0
6
0
0
8
R10
0
0
0
0
0
0
1
0
0
3
0
4
R11
0
0
0
0
0
0
1
1
0
0
2
4
Total
23
7
6
387
6
61
136
66
9
4
4
709
Asymmetry for adrenal weight steadily
increased in the course of development both
in males and females. The average adrenal
weight and maximum difference between
the left and right sides of this trait reached
greater values in females compared to males
(Fig. 4).
The right-side DA for kidney weight was
significant in males and in females, and it
also depended on DS. There was a maximum
for dextral kidney asymmetry before sexual
maturation. The end stages of population
ontogeny were characterized by symmetry of
these paired organs (Fig. 5).
# 119 #
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Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
Fig. 3. Histogram of the distribution of morphotypes complexity asymmetry d R-L among Microtus oeconomus
specimens in Teletskaya population; A) M1; B) M3. Zero value of d R-L (the biggest columns) can appear n for
specimens with symmetrical morphotypes, and also for ones with asymmetric morphotypes, but with the same
complexity (n=7 for each tooth). Specimens with nonzero values of dR-L anytime have asymmetric morphotypes.
Mean of morphotypes complexity asymmetry dR-L is significantly greater than zero for both teeth (M1: t=7.874;
p<0.001; M3: t=2.419; p=0.015). Abscissa ? d R-L, ordinate ? the number of specimens
Table 3. Statistical parameters of DA of morphological traits in samples of Microtus oeconomus voles in Teletskaya
population
623
471
Student?s paired
t-test
12.08
9.71
<1*10 -29
<1*10 -21
Males
Females
524
384
9.81
4.84
<1*10 -8
<1*10 -5
Hind paw length
Males
Females
228
182
6.40
5.76
<1*10 -9
<1*10 -7
Front paw length
Males
Females
227
182
1.93
0.98
>0.05
>0.05
Trait
Sex
N
Adrenal weight
Males
Females
Kidney weight
Dextral DA for the length of the hind paw
was significant only in males. The right hind paw
was longer than the left in young and adult voles
(Fig. 6).
Discussion
It is believed that the progressive evolution
of the animal kingdom is accompanied by a
p-value
natural loss of symmetry (Beklemishev, 1969;
Geodakian, 1991; Palmer, 2009). MCA of voles is
an example of trait with incomplete evolutionary
transition from symmetric to asymmetric state.
The existence of asymmetry is well known
and has repeatedly been observed in different
rodent species (Angermann, 1973; Kovaleva
et al., 2002; Vasil?ev et al., 2004; Pozdnyakov,
# 120 #
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Table 4. Correlation coefficients of DA (R-L) and trait size ((L+R)/2) of Microtus oeconomus voles in Teletskaya
population with environmental temperatures, population dynamics and trapping date (*p<0.05; **p<0.01;
***
p<0.001; with Bonferroni correction), see text (Statistical analysis) for details
Adrenal weight
Females (N=485)
Males (N=632)
DA
Size
DA
Size
-0.08
-0.28***
0.05
-0.38***
-0.21***
0.80***
-0.18***
0.77***
AD
-0.11
0.06
0.04
-0.14*
ADdev
-0.08
0.13
0.01
-0.01
0.02
0.15
0.03
0.00
Year
-0.09
-0.13
0.08
-0.29***
TMay
0.01
-0.07
0.07
-0.15*
TJun
-0.07
-0.10
-0.04
-0.10
TJul
0.03
-0.02
-0.05
-0.02
TAug
0.01
-0.13
0.12
-0.08
Trapping date
Sexual maturity
ADabs(dev)
Kidney weight
Females (N=402)
Males (N=552)
DA
Size
DA
Size
0.01
-0.22***
0.05
-0.33***
Sexual maturity
-0.12
0.82***
-0.17**
0.88***
AD
-0.05
0.17**
ADdev
ADabs(dev)
Trapping date
0.01
-0.12
-0.03
***
0.21
0.01
-0.01
-0.01
0.20***
0.05
0.02
Year
-0.04
-0.03
0.00
-0.27***
TMay
-0.04
0.01
-0.02
-0.13
TJun
-0.08
-0.10
-0.03
-0.22***
TJul
0.03
-0.07
0.02
0.01
TAug
-0.02
-0.13
-0.08
-0.08
Hind paw length
Females (N=205)
Males (N=257)
DA
Size
DA
Size
Trapping date
0.03
-0.01
-0.04
-0.12
Sexual maturity
0.05
0.44***
0.22
0.41***
AD
0.15
0.12
0.11
0.08
0.10
0.09
0.05
0.11
-0.08
-0.14
-0.16
-0.28**
Year
0.16
0.11
0.14
-0.03
TMay
-0.05
-0.12
-0.20
-0.26*
TJun
-0.02
0.06
0.07
0.20
TJul
-0.02
0.00
-0.01
-0.13
TAug
0.09
-0.11
0.00
-0.01
ADdev
ADabs(dev)
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
Fig 4. Age-depended dynamics of adrenal weight asymmetry of Microtus oeconomus voles in Teletskaya
population. Developmental stages: juv ? juveniles, sad ? subadults, adu ? adults, sen ? senescents. The numbers
under the developmental stages are age group numbers. ? ? median; ? ? 25 %?75 %; ? non-outlier range; ? ?
outliers. F ? Fisher F-test. KW-H ? Kruskal-Wallis H-test
Fig 5. Age-depended dynamics of kidney weight asymmetry of Microtus oeconomus voles in Teletskaya
population. For other designations, see Fig 4
2010). It is known that dental traits are highly
heritable (Bader, 1965; Berry, 1975; Hlusko
et al., 2011; Polly et al., 2011). In addition, the
different species of the voles have a common
set of morphotypes which are distinct only in
frequency of occurrence (Angermann, 1973;
Pozdnyakov, 2005; Pozdnyakov, 2011).
The development paths of these traits
which are formed in the ancestral lines of living
Arvicolinae, are highly conservative. This is
apparently based on the evolutionary stability
of the trait?s basic mechanisms of development
which canalize all deviations (Waddington,
1942; Vasil?eva, 1999; Siegal, Bergman, 2002;
Vasil?ev, 2009). Perhaps, this suggestion explains
the fact that the dR-L in our study was small, its
absolute value did not exceed 2, and it had no
developmental dynamics.
# 122 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
Fig 6. Age-depended dynamics of hind paw length asymmetry of Microtus oeconomus voles in Teletskaya
population. For other designations, see Fig 4
According to our data, the left adrenal
weighed more than the right in both sexes. The
occurrence of this phenomenon among wild
populations has been, to our knowledge, first
described for the water vole Arvicola amphibius
from Western Siberia (Galaktionov, Efimov
2003). There is a large body of laboratory data
supporting this observation for other mammalian
species (Idelman, 1978; Abramov et al., 1996;
Coleman et al., 1998; Droste et al., 2003; Trut et
al., 2002).
The founded prevalence of right asymmetry
of kidney weight for the root vole is consistent
with the previous reports. For instance, 7 natural
populations of the house mouse caught in Great
Britain, the Pacific Ocean islands, and the
Subantarctica showed a statistically significant
greater weight of the right kidney compared to the
left one, with difference decreasing with increasing
kidney size and age (Berry, 1978; Mittwoch, 1979,
2008). In our view, the evolutionary implications
for population ontogeny are clear: the organism
strives to minimize energy expenditures under
peak loads. For the growing organism, it is more
advantageous to partition over time the same
phases of organ growth and development at the
right and the left than to go through the phases
synchronously.
Asymmetry study of human limbs takes
back to the nineteenth century (Arnold, 1844;
Gennadis, 1858; Tarkhanov, 1879). Nonetheless,
the issue of the effects of environmental and
genetic factors remains very unclear. It is
known that handedness is strongly correlated
with hemispheric brain asymmetry, which has
a considerable genetic component (Geschwind
et al., 2002). The handedness?brain asymmetry
relationship has been extended to primates,
mammals and to some extent vertebrates as
a whole (Singh, 1971; Pande, Singh 1971;
Bianki, 1993). Postnatal measurements on
foxes demonstrated that the DA of limb bones
increases after birth and continues to change
(increasing or decreasing) during postnatal
growth (Kharlamova et al., 2010). Our current
study revealed that most voles of the examined
population have the hind version (?footedness?)
of this trait, rather than the front one
(?handedness?). Perhaps this is due to the spatial
and motor asymmetry which has been shown
for many species of vertebrates (Bianki, 1993,
Agulova et al., 2010; Agulova et al., 2012).
# 123 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
The prevalent view of most contemporary
papers is that the asymmetries in the early
development stages are largely due to the
contribution of internal cytogenetic factors. This
is supported by the finding of specific genes
responsible for establishing the body left-right
axes of the vertebrate embryo. They are key
components of the signaling cascade which is
responsible for the formation of this axis, and
also the morphological asymmetry of visceral
organs (Fujunaga, 1996; Wood, 1997; Burdine,
Schier, 2000). A. Palmer (1996) has suggested
that the asymmetries appeared in the later
stages of ontogeny are mostly due to external
environmental factors.
Our studies do not support this hypothesis.
Developmental dynamics of the DA of three
bilateral traits did not depend on external or
intrapopulation factors, despite the fact that the
absolute sizes of traits are affected by a number
of factors (Table 4). Similar results were obtained
early (Kovaleva, Faleev, 1994).
The DA trajectories for all the set of traits
before sexual maturation start are similar in
males and females. This similarity allowed us
to put forward the hypothesis that the two sexes
have a common genetic program during its
establishment in ontogeny. In the period of sexual
maturation asymmetry increases and then DA
trajectories of these organs are different in males
and females.
Reasons for the divergence of developmental
trajectories of the investigated traits are not
accidental, and they are in good agreement with
the Geodakian?s evolutionary theory of sex
(Geodakian, 1991). A trait changes only in males
over many generations for diclinous forms due to
replacement from stabilizing selection to moving
one. Females keep the old value of trait. The trait
evolution trajectory bifurcates into male and
female branches. This is a divergent phase of trait
evolution. Genetic sexual dimorphism arises in
such a way.
Conclusion
DA of morphological traits is perhaps more
widespread than it is generally thought. It may be
formed both at the early and later development
stages and, contrary to the Palmer? hypothesis
may be not dependent on environmental factors.
Developmental approach to the study of
the DA formation reveals its dynamic character.
Although each trait has its own dynamics DA in
ontogeny, usually DA increases in the period of
sexual maturation. Uncoordinated development
of the sides in this period seems to be associated
with an increase in intensity of developmental
processes. Our study argues that sexual
dimorphism appears at the later developmental
stages. Its manifestation is consistent with
Geodakian?s evolutionary theory of sex.
Evolutionary significance of the DA is
to increase both short-term individual fitness
and long-term population adaptive capacity.
Developmental approach to the study of
morphological and functional asymmetries in
animals opens new possibilities for the study of
evolutionary processes.
Acknowledgments
We thank Anna Fadeeva for translation of the
manuscript from Russian into English. The work
was supported by RFBR grant No.11-04-00141-?,
project for Basic Research SB RAS and UB RAS
70, integration project SB RAS 18.13, the Russian
President grant SS-5278.2012.4 and RAS program
A.II.6. We also thank two anonymous reviewers
for helping in improvement of the manuscript.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
References
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
Abramov V.V., Karmatskikh O.L., Kozlov V.A., Os?kina I.N. (1996) Functional asymmetry of
adrenal glands in (CBA x C57BL/6)F1 mice. Dokl. Akad. Nauk 347(6): 831?833.
Agulova L.P., Andreevskikh A.V., Bolshakova N.P., Kravchenko L.B., Suchkova N.G., Moskvitina
N.S. (2010) Dynamics of motor asymmetry profiles in striped field mouse (Apodemus agrarius
Pall.) populations. Rus. J. Ecol. 41(6): 506?512. (in Russian)
Agulova L.P., Bolshakova N.P., Andreevskikh A.V., Suchkova N.G., Kravchenko L.B., Moskvitina
N.S. (2012) Functional motor asymmetry in three species of mouse-like rodents from natural
populations. Neurosci. Behave. Physiol. 42(7): 718?724.
Angermann R. (1973) Homologous variability of molars in voles (Microtinae). In: Vorontsov N.N.
(ed.) Problems of evolution, vol. 3. Novosibirsk: Nauka, p. 104?118 (in Russian)
Arnold F. (1844) Handbuch der Anatomie des Menschen. Freiburg: Herder?sche.
Bader R.S. (1965) Heritability of dental characters in the house mouse. Evolution 19(3): 378?
384.
Bashenina N.V. (1953) On age determination of common vole (Microtus arvalis Pall). Rus. J. Zool.
32 (4): 730?743. (in Russian)
Beklemishev V.N. (1969) Principles of comparative anatomy of invertebrates. Chicago: University
of Chicago Press, 1058 p.
Berry A.C. (1975) Factors affecting the incidence of non-metrical skeletal variants. J. Anat. 120:
519?535.
Berry R.J. (1978) Genetic variation in wild house mice: where natural selection and history meet.
Am. Sci. 66: 52?60.
Bianki V.L. (1993) The mechanisms of brain lateralization. New York?London: Gordon Breach,
290 p.
Burdine R.D., Schier A.F. (2000) Conserved and divergent mechanisms in left?right axis
formation. Genes Dev. 14: 763?776.
Clarke G.M. (1993) Fluctuating asymmetry of invertebrate populations as a biological indicator of
environmental quality. Environ. Pollut. 82: 207?211.
Coleman M.A., Garland J.T., Marler C.A., Newton S.S., Swallow J.G., Carter P.A. (1998)
Glucocorticoid response to forced exercise in laboratory house mice (Mus domesticus). Physiol.
Behav. 63: 279?285.
Collignon J., Varlet I., Robertson E.J. (1996) Relationship between asymmetric nodal expression
and the direction of embryonic turning. Nature 381: 155?158.
Droste S.K., Gesing A., Ulbricht S., Mцller M.B., Linthorst A.C.E., Reul J.M.H.M. (2003) Effects
of long-term voluntary exercise on the mouse hypothalamic-pituitary-adrenocortical axis.
Endocrinology 144(7): 3012?3023.
Fujinaga M. (1997) Development of sidedness of asymmetric body structures in vertebrates. Int.
J. Dev. Biol. 41: 153?186.
Galaktionov Y.K., Efimov V.M. (2003) Variability of morpho-physiological indicators and
bilateral asymmetry indices depending on phase of population density dynamics. In: Ravkin Y.S.
(ed.) Realization of morphological diversity in natural mammalian populations. Novosibirsk: SB
RAS, p. 125?130. (in Russian)
# 125 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
19. Gennadis G. (1858) Textbook of Regional Anatomy. 1st Edn., Athens: Antoniadis Publisher,
p. 270?271.
20. Geodakian V.A. (1991) The evolutionary theory of sex. Priroda (8): 60?69 (in Russian)
21. Geschwind D.H., Miller B.L., DeCarli C., Carmelli D. (2002) Heritability of lobar brain volumes
in twins supports genetic models of cerebral laterality and handedness. PNAS 99(5): 3176?3181.
22. Graham J.H., Freeman D.C., Emlen J.M., Leame L.J., Kiesler J.A. (1998) Directional asymmetry
and the measurement of developmental instability. Biol. J. Linn. Soc. 64: 1?16.
23. Graham J.H., Shimizu K., Emlen J.M., Freeman D.C., Merkel J. (2003) Growth model and the
expected distribution of fluctuating asymmetry. Biol. J. Linn. Soc. 80: 57?65.
24. Hlusko L.J, Sage R.D., Mahaney M.C. (2011). Modularity in the mammalian dentition: Mice and
monkeys share a common dental genetic architecture. J. Exp. Zool. B Mol. Dev. Evol. 316(1):
21?49.
25. Idelman S. (1978) The structure of the mammalian adrenal cortex. In: Jones I.C., Henderson I.W.
(eds.) General, comparative and clinical endocrinology of the adrenal cortex. London: Academic
Press, p. 1?199.
26. Kharlamova A.V., Trut L.N., Chase K., Kukekova A.V., Lark K.G. (2010) Directional asymmetry
in the limbs, skull and pelvis of the silver fox (V. vulpes). J. Morph. 271(12): 1501?1508.
27. Kovaleva V.Y., Faleev V.I. (1994) Morphological variability of Microtus oeconomus (Rodentia,
Cricetidae) in different temperature conditions. Zool. Zh. 73(9): 139?145. (in Russian)
28. Kovaleva V.Y., Pozdnyakov A.A., Efimov V.M. (2002) Investigation of variability structure of
root vole (Microtus oeconomus Pallas) molar morphotypes using bilateral asymmetry. Zool. Zh.
81(1): 111?117. (in Russian)
29. Larina N.I., Lapshov V.A. (1972) On methods for distinguishing age groups in some voles. In:
Physiological and population ecology of animals, vol. 2(4). Saratov: Saratov Univ., p. 92?97. (in
Russian)
30. Leung B., Forbes M.R., Houle D. (2000) Fluctuating asymmetry as a bioindicator of stress:
comparing efficacy of analyses involving multiple traits. American Naturalist 155(1): 101?115.
31. Levin M., Johnson R.L., Stern C.D., Kuehn M. Tabin C. (1995) A molecular pathway determining
left-right asymmetry in chick embryogenesis. Cell 82: 803?814.
32. Levin M., Palmer A.R. (2007) Left-right patterning from the inside out: widespread evidence for
intracellular control. BioEssays 29: 271?287.
33. Litvinov Y.N., Abramov S.A., Kovaleva V.Y., Krivopalov A.V., Novikov E.A., Chechulin A.I.
(2007) Structural-temporal organization of a rodent community in the Teletskaya taiga, Altai
Mountains. Rus. J. Ecol. 38(6): 444?449.
34. Lopez-Gracia M.L., Ros M.A. (2007) Left-right asymmetry in vertebrate development. Adv.
Anat. Embryol. Cell Biol. 188: 1?121.
35. Lowe L.A., Supp D.M., Sampath K., Yokoyama T., Wright C.V., Potter S.S., Overbeek P., Kuehn
M.R. (1996) Conserved left-right asymmetry of nodal expression and alterations in murine situs
inversus. Nature 381: 158?161.
36. Meno C., Shimono A., Saijoh Y., Yashiro K., Mochida K., Ohishi S., Noji S., Kondoh H., Hamada
H. (1998) Lefty-1 is required for left-right determination as a regulator of lefty-2 and nodal. Cell
94: 287?297.
# 126 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
37. Merilд J., Bjцrklund M. (1995) Fluctuating asymmetry and measurement error. Systematic Biol.
44(11): 97?101.
38. Mittwoch U. (1979) Lateral asymmetry of kidney weights in different populations of wild mice.
Biol. J. Linn. Soc. 11: 295?300.
39. Mittwoch U. (2008) Different gene expressions on the left and the right: a genotype/phenotype
mismatch in need. Ann. Hum. Genet. 72(1): 2?9.
40. Palmer A.R. (1994) Fluctuating asymmetry analysis: a primer. In: Markow T., ed. Developmental
instability: its origins and evolutionary implications. Dordrecht: Kluwer Academic Publishers,
p. 335?364.
41. Palmer A.R. (1996) From symmetry to asymmetry: phylogenetic patterns of asymmetry variation
in animals and their evolutionary significance. PNAS 93: 14279?14286.
42. Palmer A.R. (2009) Animal asymmetry. Curr. Biol. 19(12): 473?477.
43. Palmer A.R. Strobeck C. (1986) Fluctuating asymmetry: measurement, analysis, patterns. Annu.
Rev. Ecol. Syst. 17: 391?421.
44. Palmer A.R., Strobeck C. (1992) Fluctuating asymmetry as a measure of developmental stability:
implications of non-normal distributions and power of statistical tests. Acta Zool. Fenn. 191: 55?70.
45. Palmer A.R., Strobeck C. (2003) Fluctuating asymmetry studies revisited. In: Polak M., ed.
Developmental instability: causes and consequences. Oxford: Oxford University Press, p. 279?319.
46. Pande B.S., Singh I. (1971) One-sided dominance in the upper limbs of human fetuses as evidenced
by asymmetry in muscle and bone weight. J. Anat. 109: 457?459.
47. Parsons P.A. (1992) Fluctuating asymmetry: a biological monitor of environmental and genomic
stress. Heredity 68: 361?364.
48. Polly P.D., Killick L., Ruddy M. (2011) Using left-right asymmetry to estimate non-genetic variation
in vole teeth (Arvicolinae, Muridae, Rodentia). Palaeontologia electronica 14(3, 41A): 1?12.
49. Pozdnyakov A.A. (1993) Morphotypical variability of molars in meadow voles of the ?maximoviczi?
group (Rodentia, Arvicolidae, Microtus): an attempt of quantitative statistical analysis. Zool. Zh.
72(11): 114?125. (in Russian).
50. Pozdnyakov A.A. (2005) The structure of the grey voles M3 morphological variability (Microtus
Schrank, 1798). In: Abramson N.I., Averianov A.O., eds. Systematics, paleontology and phylogeny
of rodents. Proc. Zool. Inst. RAS, vol. 306. St. Petersburg: Zool. Inst. RAS, p. 102?115. (in
Russian)
51. Pozdnyakov A.A. (2010) Morphological Diversity: characteristics, structure, analysis. In:
Bolshakov V.N., ed. Animal communities and populations: ecological and morphological
analysis. Proc. Inst. Syst. Ecol. SB RAS, vol. 46. Novosibirsk?Moscow: KMK Press, p. 133?157.
(in Russian)
52. Pozdnyakov A.A. (2011) The structure of morphological variability (with the masticatory surface
morphotypes of the lower first molar in the voles as an example). Biol. Bull. Rev. 1(5): 471?481 (in
Russian)
53. Shiratori H., Hamada H. (2006) The left-right axis in the mouse: from origin to morphology.
Development 133: 2095?2104.
54. Siegal M., Bergman A. (2002) Waddington?s canalization revisited: Developmental stability and
evolution. PNAS 99: 10528?10532.
# 127 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
55. Singh I. (1971) One-sided dominance in the limbs of rabbits and frogs, as evidenced by asymmetry
in bone weight. J. Anat. 109: 271?275.
56. Stige L.C., David B., Alibert P. (2006) On hidden heterogeneity in directional asymmetry ? can
systematic bias be avoided? J. Evol. Biol. 19(2): 492?499.
57. Tarkhanov I.R. (1879) Psychomotor centres and their development in human and animals.
St. Petersburg: L.F.Panteleev, 168 p. (in Russian)
58. Trut L.N., Prasolova L.A., Kharlamova A.V., Plyusnina I.Z. (2002) Directional left-sided
asymmetry of adrenals in experimentally domesticated animals. B. Exp. Biol. Med. 133(5): 506?
509. (in Russian)
59. Van Dongen S. (2006) Fluctuating asymmetry and developmental instability in evolutionary
biology: past, present and future. J Evol. 19(6): 1727?1743
60. Vasil?ev A.G. (2009) Phenogenetic variability and population meronomy. Zh. Obsh. Biol. 70(3):
195?209 (in Russian)
61. Vasil?ev A.G., Faleev V.I., Galaktionov Yu.K., et al. (2004) Realization of the morphological
diversity in natural populations of mammals. Novosibirsk: SB RAS, 232 p. (in Russian)
62. Vasil?eva I. (1999) Epigenetic divergence of Asian high-mountain voles of the subgenus Aschizomys
from southern and north-eastern Siberia. Folia Zool. 48(Suppl. 1): 105?114.
63. Waddington C.H. (1942) Canalization of development and the inheritance of acquired characters.
Nature 150: 563?565.
64. Wood W.B. (1997) Left-right asymmetry in animal development. Ann. Rev. Cell Dev. Biol. 13:
53?82.
65. Zahn J.M., Poosala S., Owen A.B., et al. (2007) AGEMAP: A gene expression database for aging
in mice. PLoS Genet. 3: 201.
66. Zakharov V.M. (2003) Linking developmental stability and environmental stress: a whole organism
approach. In: Polak M., ed. Developmental instability: causes and consequences. Oxford: Oxford
University Press, p. 402?414.
67. Zakharov V.M., Graham J.H. (1992) Developmental stability in natural populations. Acta Zool.
Fenn. 191: 7?30.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Vera Yu. Kovaleva, Vadim M. Efimov? Directional Asymmetry of Morphological Traits During Postnatal Ontogeny?
???????????? ??????????
??????????????? ?????????
? ??????? ?????????????? ??????????
? ???????-???????? Microtus oeconomus Pall.
(Rodentia, Cricetidae)
?.?. ????????a,
?.?. ??????a,?,?, ?.?. ????????a
a
???????? ??????????? ? ???????? ???????? ?? ???,
?????? 630091, ???????????, ??. ??????, 11
?
???????? ????????? ? ???????? ?? ???,
?????? 630090, ???????????, ???????????, 10
?
??????? ??????????????? ???????????,
??????, 634050, ?????, ??. ??????, 36
??????????????? ???????? ?? ???????????? ????????????? ??????????????? ?????????
???????-???????? Microtus oeconomus, ????????? ? ??????? 25 ??? (1982?1992, 1994?2005,
2009, 2010). ???????? ???????????? ?????????? (??) ?????????? ???????? ?????, ????
????????????? ? ?????, ? ????? ????? ?????? ????. ????????, ??? ?? ????? ?????????????
??? ?? ??????, ??? ? ?? ??????? ??????? ?????????? ?, ??-????????, ????? ??????
???????????? ????????????. ????? ????, ?? ??????? ???????????? ???????? ????? ????
??????????? ???????????????? ??????????, ??????? ?? ??????? ?? ???????? ????? ?
???????? ??????????? ?????????. ? ?????? ???????? ?????????? ?????????? ?????????? ??
???? ??????????? ?????????, ???????????????? ????????? ? ??????????? ?????????????
????????? ????????. ????????, ??? ??????? ????????? ?? ??????????? ?? ???????
??????? ??????????, ??? ?????? ??????????? ? ???????????? ??????? ???? ?.?. ?????????.
???????????? ???????? ?? ??????????? ? ????????? ?????????? ???????????? ???
?????????? ?????????, ??? ? ???? ?????????. ???????????????? ?????? ? ???????????? ??????????????????? ?????????? ? ???????? ????????? ????? ??????????? ??? ????????????
???????????? ?????????.
???????? ?????: Microtus oeconomus, ???????????? ??????????, ????????? ?????, ??? ?????
? ?????????????, ????? ???, ??????? ?????, ???????? ???????????.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2013 6) 130-149
~~~
??? 595.315
??????? ????? ???? Colossendeis
(Colossendeidae, Pycnogonida) ???? ????????
? ?????????? ?????????
?.?. ????????, ?.?. ???????*
???????? ??????????? ??. ?.?. ??????? ???,
?????? 117997, ??????, ??????????? ????????, 36
Received 12.12.2012, received in revised form 18.01.2013, accepted 04.06.2013
? 1996 ? 2000 ??. ???????????? R/V «Polarstern» ANT XIII/3 ? ANT XVII/3 ???? ??????? 17 ?????
Colossendeis. ???????? ????? ? ???? ?????? ?????????? ?????????????? ??? ?. megalonyx
Hoek, 1881, ?????????????? ????? ?????????. ???????? ???????? ???????? ?. megalonyx
megalonyx. ???????????????? ???????????? ????????? C. megalonyx megalonyx. ?????????
???? ??? ??????????? ???????? ?. megalonyx. ????????????? ????????????, ? ?????
??????????????? ????? ? ???????? ???? Colossendeis ????????? ????????????, ??? ? ????
???????? ? ????? ?? ?????????? ??????????????? ?????????? ? ??????? ????????????? ???????
?????????? ??????????????? ?????? ???? ? ?????????? ??????????? ??? ?????????????? ?
?????????????, ????? ? ????????? ??????.
???????? ?????: Pycnogonida, ???? ??? ??????????? ???????? Colossendeis megalonyx,
??????????????? ? ???? ????????.
????????
??? ???????????? ?????? ????? ?????????? ? ???????? ?????? ????? ?????? ??????????? ??????? ??????? ?????, ?????????? ????????? ???????? Colossendeis Jarzinsky, 1870.
???????? ? ??????????? ????? ????? ???? ???
?? ???? ???????????? ????????? ? ?????????.
? ???? ????? ?????? ????????? ?????? ???????? ?????????? ??????? ????????? ????? ? ?????????? ?? ?????? ????? (? 1996 ? 2000 ??.)
????? ??????????, ???????? ?? ??????? ????
???????? ?????????????? ????????????
*
?? ????? «Polarstern». ?????? ????? ?????
???? ? ???????????? ???????? Colossendeis
megalonyx Hoek, 1881 ???????? ? ?????? ?
???, ??? ???? ???????? ???????? ????? ??
??????? ??????????????? ?????????????
?????? ????? ?????? ?????????.
????????? ? ??????
???????? ??? ????????? ?????? ???
?????? ? ???? ???????? ? ??????????? R/V
«Polarstern» ? 1996 ?. (???? ANT XIII/3) ?
2000 ?. (???? ANT XVII/3) (????. 1).
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: alex_rayskiy@mail.ru
# 130 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
??????? 1. ???? Colossendeis, ????????? ???????????? R/V Polarstern ? ???? ???????? ? 1996 ? 2000 ??.
? ?????? ????? ????????? ? ???? ????? ????? ?????????? ????? (?????????) ? ????? ??????????
????????? ??????????? (???????????)
?????? ?????
???? Colossendeis
??? ????????
1996 ? 2000 ??.
??? ???????
1996 ?.
????? ???????????
?-?? 2000 ?.
C. australis Hodgson
4/5
?
?
C. avidus Pushkin
1/3
2/2
1/1
C. brevirostris Child
1/1
?
?
C. drakei Calman
4/5
1/1
2/3
C. fragilis Pushkin
2/2
1/2
?
C. glacialis Hodgson
3/8
?
?
C. korotkewitschi Pushkin
1/2
?
?
10/25
1/1
?
C. lilliei Calman
C. longirostris Gordon
?
1/1
?
C. megalonyx megalonix Hoek
16/59
4/10
7/34
C. megalonyx arcanus Turpaeva.
4/11
?
?
C. megalonyx frigida Hodgson
3/5
?
1/1
C. megalonyx rugosa Hodgson
2/7
2/2
?
C. megalonyx weddellensis Turpaeva
2/3
?
?
C. potentis Turpaeva
1/1
?
?
C. pseudochelata Pushkin
1/1
?
?
C. robusta Hoek
?
?
1/5
C. scotti Calman
4/9
?
?
C. smirnovi Pushkin
1/3
?
?
C. tortipalpis Gordon
2/2
3/7
2/4
C. wilsoni Calman
3/5
?
?
???????? ??????? ???? ???????? ?????? ?????, ? ??? ?????? ?? ???????? ???
??????????? ?????? ????. ??????? ?????
?????? ????? ????????? ? ???? ???????????? ????????? ???????, ?????? ???????,
??????, ?????????? ???????????????? ????????. ????? ? ???? ????????, ???????????? ???????????? 70-72? ?.?. ? 10-15? ?.?.
? ????????? ?? ??????? ??????????? ??
700 ?, ??? ?????????? ? 1996 ? 2000 ??. ?????, ????????????? ????? ???? ???????
(Vestkapp) ????? 71-74? ?.?. ? 21-23? ?.?.
?? ???????? 200-2000 ?, ??? ?????????? ?
1996 ?. ????? ???????? ????? ???????????
???????? ????? ???????????? 61-63? ?.?. ?
60-61? ?.?. ? ????????? 200-1000 ?, ??? ?????????? ? 2000 ?. ????? ????, ? ?????? ???????????? ????????? ?? ????????? ????????? ???????????, ????????? ????????????
??? «???????? ????????» ? 1967 ?., ???
«???????? ????????» ? 1971 ?. ? ??? «??????» ? 1988 ?.
? ????? ?? ???????? ???? ???????????? ??? ?????? ?? ?????????????? ???????????? (??????, 1993; Child, 1995), ? ??????? ????????? ?????? ?????????????? ?????
# 131 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
Colossendeis. ?????? ?.?. ??????? ???????
?? 26 ?????, ?? ??????? 5 ????? ??? ?????,
? ??? ????? ???? ??????? ? ???? ????????
(Fry, Hedgpeth, 1969). ?????? ?????? (Child,
1995) ??????? ?? 17 ?????, ?? ??????? ?????
5 ????? ????? ??? ?????. ?????????? ?????
?????????????? Colossendeis ??????? ? ?????
???????. ? ????????? ?? ???? ????????, ????????? ? 1996 ? 2000 ??., ????????? 17 ?????
Colossendeis, ?????? ???? ???, C. megalonyx,
??????????? ????? ?????????. ?????? ???????????? ????? Colossendeis ???????? ?
????. 1.
??????????
??????????? ???????????? ? ????
???????? ????? Colossendeis ??????? ?
?????????? ???????????? ?????. ???????
???? ?????????? ????????, ???????????
???????? ????????? ????, ? ??????????
????????? ??????????????? ???? C. megalonyx.
Colossendeis avidus Pushkin, 1970
Colossendeis avidus ??????, 1970, ?. 14881490, ???. 1; ? 1993. ?. 86-88, ???. 66; ????????,
1998, ?. 65; Child, 1995, p. 74, fig. 1 f-g.
????????.
1?,
R/V
«Polarstern»,
ANT XVII/3, ??????? 13/GSN, 14.II 1996,
73?36ґ3 ?.?., 22?19? ?.?., ??????? 620 ?; 1?,
ANT XVII/3, 2.V 2000, ??????? 12/GSN,
61?58ґ5 ?.?., 60?18ґ7 ?.?., ??????? 804-930 ?.
??????? (?????) ????? ????????????? ???? ??? ?????? ? 1957 ?. ???????????
?? ????? «???» ? ???? ??????????? ?? ??????? 270 ? (??????, 1993). ? 1963 ?. ??????????? ?? R/V «Eltanin» ? ??????-????????
????? ???? ???????? ? ?????????? ????????
??? ?????? ??? ???? ????????? C. avidus,
???? ?? ????????, ????? (Child, 1995). ? ??????? ?? ?????? ????? Colossendeis, C. avidus
???????? ?????? ??????? ???????. ????????
??????? ???? ??????????? ?????? ?? ????????, ?????????? ????? ??????? ?????????,
?? ?????? ? ?? ???? ???????????. ????? ???-
???. 1. Colossendeis avidus (1-5), Colossendeis megalonyx megalonyx (????????? ?? ?????? ???? ????????)
(6-9). 1, 7 ? ??????? ?????, 2 ? ????? ?? ??????? ???????, 3 ? ?????????? ????? 10-?? ??????? III ??????????,
4 ? ???????? ???? 1-?? ????, 5, 9 ? 7-? ? 8-? ??????? ????, 6 ? ???? ?? ??????? ???????, 8 ? 10-? ??????
III ??????????. ??????? 5 ??
# 132 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
??? ????, ????????????? ????????, ??????
????????? ??????????? ????, ??????? ? ??????? ?????. ???? ?????????? ????? ? ?????????? ?????? (Child, 1995) ? ????????? ?????????? ?? ????????. ? ???????? ????? ?????
????? ????????? ??????? ? ??????? ?????,
????? ??????? ??????? ????? ? ??????????????? ????????? ? ????????????????? ???????, ????? ??????? ????? ????????? ???.
????????? ??????? ?????, ???? ??????????
??????? ? ??????????? ????? ?? ??????? 13/
GSN.
???????? ? 14.2, ????? ? 15.1, ??????? ?
2.8;
II ?????????? ? 25.2 (1-? ?????? ? 0.5,
2-? ? 7.9, 3-? ? 0.8, 4-? ? 5.1, 5-? ? 2.3, 6-? ? 2.6,
7-? ? 2.1, 8-? ? 2.0, 9-? ? 1.9);
III ?????????? ? 40.2 (1-? ?????? ? 1.0,
2-? ? 1.2, 3-? ? 1.5, 4-? ? 9.6, 5-? ? 5.4, 6-? ?
11.2, 7-? ? 3.1, 8-? ? 2.6, 9-? ? 2.5, 10-? ? 2.1),
??????? ? 0.81;
???? III ???? ? 84.0 (1-? ?????? ? 2.0,
2-? ? 2.5, 3-? ? 2.1, 4-? ? 25.2, 5-? ? 22.0, 6-? ?
18.5, 7-? ? 6.4, 8-? ? 5.3), ?????? ? 4.6.
?????? ???, ???????? ?? ??????????????
? ?????????????? ???????? ??????????????
???.
Colossendeis potentis Turpaeva, 2008
Colossendeis pontetis ????????, 2008,
?. 886-888, ???. 2.
????????. 1 ?, R/V «Polarstern» ANT
XVII/3, ??????? 111-5, 5.IV 2000, 71?07ґ5 ?.?.,
11?27ґ7 ?.?., ??????? 66 ?.
??????? ????????? (????? ????????
22.9 ??, ????? ?????? 32.3 ??, ????? ???? III
???? 119.5 ??), ???????? ?????? ? C. glacialis
Hodgson, 1907, C. leniensis Pushkin, 1993 ?
C. elephantis Child, 1995, ?? ?????????? ??
??? ????? ???????? ?????????, ????? ???????? ????????? ? ???????, ?????? ???????? ?????, ?????? ???????????? ????? ??-
???. 2 ???????????? ????? ? ??????????? Colossendeis megalonyx megalonyx. 1-4 ? ??????? ?????,
5-10 ? 7-? ? 8-? ??????? II ???????????, 11-15 ? 10-? ??????? III ??????????? (1, 5, 7, 11, 12 ? ??????????
?? ?????? ???? ????????; 2, 6, 13, 14, 15 ? ?????????? ?? ?????? ????? ??????????? ????????;
3, 8 ? ?????????? ? ????????????? ??????; 4, 9, 10 ? ?????????? ?? ????-?????????? ??????). ???????
1 ??
# 133 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
????????? III ???? ? ??????????? ????????
???.
Colossendeis megalonyx Hoek, 1881
Colossendeis megalonyx Hoek, 1881, p. 6769, figs. 1-3; Fry, Hedgpeth, 1969, p. 29-30, figs. 7,
8, 10-23 (sin.); ??????, 1993, ?. 52-54, ???. 33,
34; Child, 1995, p. 86 (sin.); ? 1998, p. 58-59.
?????? ???????????????? ? ?????????????? ????? ?????????????? ???. ???
???????? ????????: ???????? ??????????,
?????????? ????? ???????? ??????????
?????????????? ????? ?????? ?????????,
?????? ???? ???????, ??????? ????? ???????, ?????????? ??? ??????????????????. ?
??????????? II ???? 2-? ?????? ????? ???????, 5-? ?????? ?????? 10 % ????? ???? ?????, 6-? ?????? ????? ??? ? 3 ???? ??????? ????? ??????, 7-? ?????? ??????????? ??????
8-??. ????????? ???? ???????, ?? 4-? ??????
????? ???????, 6-? ?????? 5-??, 7-? ???????
8-??, ?????? ?????? ??????? ??????? ???????? 8-?? ???????.
?? ?????????????? ? ????????????????? ??? ??????? ????????? ????, ???????
? C. megalonyx. ????? ?? ??? ????, ??????
? ????? (Fry, Hedgpeth, 1969; Child, 1995)
??????? ?????????: C. megalonyx megalonyx
Hoek, 1881, C. megalonyx frigida Hodgson,
1907, C. megalonyx or?adens? Hodgson,
1908, C. megalonyx scoresbii Gordon, 1932,
C. megalonyx arundorostris Fry, Hedgpeth,
1969, C. megalonyx rugosa Hodgson, 1907.
??????? (????????, 1975) ? ???? ??????? ???
?????? ??????-????????????? ??? C. tenera,
????????? ???????? (Hilton, 1943) ? ????????????? ???????? (Hedgpeth, 1943), ?????? ?????? ??????? ???????? C. tenera Hilton,
1943 c C. megalonyx. ? ????????????? ??????
???? ?????????? ??? ??? ??????? ? C. megalonyx arcanus Turpaeva, 2008 ? C. megalonyx
weddellensis Turpaeva, 2008 (????????, 2008).
???? ??????? ????????? ? ????? ????????? 6 ???????? C. megalonyx, ??????????? ???????????? C. megalonyx megalonyx ?
????????? ???? ??? ??????????? ????????
????? ????.
Colossendeis megalonyx
megalonyx Hoek, 1881
Colossendeis megalonyx megalonyx Fry,
Hedgpeth, 1969, p. 30-32, figs. 7, 8, 11-16, 23;
????????, 1998, ?. 66; Child, 1995, p. 86-87.
Colossendeis
megalonyx
????????,
1975, ?. 236-239, ???. 5, 6; ??????, 1993, ?. 5254, ???. 33-34.
Colossendeis
tenuipedis
??????,
1993, ?. 56-58, ???. 37, 38.
????????. 1?, R/V «Polarstern», ANT
XIII/3, ??????? 1, 5.II 1996, 71?02ґ1 ?.?.,
11?19ґ3 ?.?., ??????? 462-481 ?; 1 ???????, ANT
XVII/3, ??????? 65-1, 31.III 2000, 71?06ґ?.?.,13?
48ґ ?.?., ??????? 615-648 ?; 1?, ??????? 102-1,
3.IV 2000, 71?11ґ9 ?.?., 12? 21ґ7 ?.?., ??????? 323-312 ?; 2 ???., ??????? 109-1, 4.IV
2000, 71?11ґ3 ?.?., 12?18ґ5 ?.?., ??????? 311316 ?; 1?, ??????? 111-9, 5.IV 2000, 71?07ґ5 ?.?.,
11?27ґ8 ?.?., ??????? 62-145 ?; 4?, 3 ???????, ??????? 111-16, 5.IV 2000, 71?07:5 ?.?.,
11?27ґ6 ?.?., ??????? 80 ?; 1?, ??????? 135-1,
10.IV 2000, 70?50ґ2 ?.?., 10?34ґ7 ?.?., ???????
274-273 ?; 26 ???. (4?, 8?, 8 ???????, ? ????? ??????????? ??? ?? ?????????), ???????
136-1, 10.IV 2000, 70?50ґ2 ?.?., 10?35ґ4 ?.?., ??????? 271-251 ?; 12 ???., ??????? 173-1, 30.IV
2000, 63?01ґ2 ?.?., 61?08ґ7 ?.?., ??????? 352379 ?; 1?, ??????? 177-1, 1.V 2000, 62?49ґ5 ?.?.,
60?49ґ3 ?.?., ??????? 202-200 ?; 1?, 2 ???????, ??????? 178-2, 2.V 2000, 61?58ґ5 ?.?.,
60?18ґ7 ?.?., ??????? 804-930 ?; 3?, 3?, 3 ???????, ??????? 183-1, 3.V 2000, 62?06ґ7 ?.?.,
60?21ґ6 ?.?., ??????? 204-200 ?; 7 ???., ???????
184-1, 3.V 2000, 62?00ґ9 ?.?., 60?20ґ7 ?.?., ??????? 374-338 ?; 10 ???., ??? «???????? ???-
# 134 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
?????», ??????? 135, 1965 ?., 60?37ґ ?.?.,
44?39ґ ?.??., ??????? 330-450 ?; 7 ???., ???????
1035, 1967 ?., 45?06ґ, ?.?., 60?11ґ ?.?., ???????
130-124 ?; 9 ???., ??? «???????? ????????»,
??????? 882, 1970 ?., 57?09ґ ?.?., 26?40ґ ?.?., ??????? 1687-1837 ?; 11 ???., ??????? 888, 1970 ?.,
57?06ґ ?.?., 26?45ґ ?.?., ??????? 318 ?.
?????
??????
????????????????
???????????????????? ?????? C. megalonyx. ??????? ?? ???????? 60-4795 ?. ???
???????? ????????: ???????? ???????, ??????????, ?????????? ????? ???????? ?????????? ????? ????? ???????? ?????????;
?????????? ??????? ? ???????, ?????? ??
?????????? ???????? ??????????? II ????
????? ?????? ??????? ? ?? ??????? 8-?
???????? ??? ????????? ??? ???????? ???????; ??????? ????? ???????; ????? ???????
??????? ????????, ?????????????? ? ????????????? ????? ? ??????????? ? ???????
? ?????????? ??????. ?????????? II ???? ?
C. megalonyx megalonyx ? 3-4 ???? ???????
????????; ?? 7-? ?????? ????? ????????.
????????? ???? ? 10-15 ??? ??????? ????????, ?? ????? ??????? ?????? 4-?, 6-?
?????? ?????? 5-??, 7-? ?????? ??????? 8-??,
????? ????? 7-?? ? 8-?? ???????? ????? ????? ????? 6-?? ???????, ?????? ????? ?????
?????? 8-?? ???????.
???? ?????? ?????? ?? ???????????
?? ???? ?????. ??????? ??? ????????? ??
???????? ????? ??????? ? ??????????? ????? ?? ???? ???????? ?? ??????? 177-1.
???????? ? 10.1, ????? ? 20.6, ??????? ?
1.4;
II ?????????? ? 28.9 (1-? ?????? ? 0.8,
2-? ? 10.2, 3-? ? 0.8, 4-? ? 7.1, 5-? ? 2.1, 6-? ?
3.0, 7-? ? 1.0, 8-? ? 1.7, 9-? ? 2.2);
III ?????????? ? 44.9 (1-? ?????? ? 1.0,
2-? ? 0.8, 3-? ? 1.0, 4-? ? 14.0, 5-? ? 7.1, 6-? ?
13.7, 7-? ? 1.1, 8-? ? 1.9, 9-? ? 2.2, 10-? ? 2.1),
??????? ? 1.35;
???? III ???? ? 102.2 (1-? ?????? ? 1.5,
2-? ? 3.0, 3-? ? 2.8, 4-? ? 28.1, 5-? ? 25.2, 6-? ?
20.5, 7-? ? 11.6, 8-? ? 9.5), ?????? ? 5.5.
??? ????????? ??????????? C. megalonyx megalonyx, ????????? ????????????
R/V «Polarstern» ? ???? ????????, ? ????????? ?? ? ???????????? ????? ???????, ?????????? ???????????? ??? «????????
????????» ? ????-??????????? ?????? ?
??? «???????? ????????» ?? ???????????? ?????? (????????, 1974, 1975), ????????? ???????? ???????????? ?? ?????????.
??? ?? ?????? ? 67 ??????????? C. megalonyx
megalonyx ?? ??????? ??????? (????????????
?????, ????-????????? ?????, ????? ????? ??????????? ???????? ? ????? ????
????????) ???? ???????? ????? ????????
(?? ??????? ???? ?? ????????? ????????),
????? ?????? ? ??? ?????? ? ??????? ?????,
????? ?????????? ???????? II ??????????,
????? ???? ?????????? ???????? ? ????? ????????? ???. ?? ??????????? ???? ?????????
?????????? ??????????? ????????? ??????
???? ????????????? ???????????, ?? ??????? ????????, ???????????? ?????????? ??
??????? ? ??????????? ???????? (????. 2).
? ???????? ??????????? ??????? ???????
???????? ?? ???? ??????? ???? ??????, ??
???????? ????????????? ?????????? ? ???????? ???????????? ???????? ????????? ??
??????????? ??????? ???????????? ????????
?????? ? ???? ???????? ?? ????????? ? ?????? ???????? ?????????????? ??????.
??????? ???????? ????????????? ?????
?????? ? ?????? ??????? ?????????? ?????????????. ?????? ? ??????????? ?? ??????????
??????? ????????????? ????? ?????? ???? ????? ??????????, ??? ? ???????????, ????????? ? ???????? ????? ????? ?????????. ?
????????? ?????? ?? ?????????? ???????
????????????? ????? ?????? ????????? ??????????? ?????? ??????? ???????. ???, ? ?????
# 135 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
??????? 2. ?????????????? ????????? ??????????? ????????? ???????????? ??????????? C. megalonyx
megalonyx ?? ?????? ??????? ( x ? ??????? ????????, ? ? ???????????? ??????????, v ? ???????????
????????, n ? ????? ??????????? ? ??????)
????????????
?????. ???
«????????
????????»,
??????? 135 ? 1035
????-?????????
?????. ???
«????????
????????»,
??????? 882 ? 888
x
10.40
10.64
8.98
9.48
?
±0.712
±0.748
±1.846
±1.933
v
0.068
0.069
0.206
0.204
n
17
20
11
14
x
1.97
2.07
2.004
2.13
?
±0.212
±0.105
±0.150
±0.249
v
0.108
0.053
0.075
0.103
n
17
20
11
14
x
6.64
6.18
7.06
6.49
?
±0.590
±0.358
±1.407
±0.583
v
0.089
0.058
0.199
0.089
n
19
19
11
14
x
0.50
0.53
0.43
0.53
?
±0.058
±0.054
±0.104
±0.098
v
0.116
0.101
0.242
0.183
n
16
19
11
14
x
0.98
1.00
0.96
1.03
?
±0.129
±0.043
±0.086
±0.063
v
0.132
0.043
0.090
0.062
n
16
19
11
16
x
0.76
0.59
0.64
0.55
?
±0.122
±0.055
±0.057
±0.054
v
0.160
0.094
0.090
0.099
n
16
20
11
16
?????????
?????
????????, ??
????? ??????/
????? ????????
?????: ?????/
??????
II ??????????:
????? 7-??
???????/????? 8-??
???????
????:
????? ????? 7-??
? 8-?? ????????/
????? 6-?? ???????
????:
????? ?????/?????
8-?? ???????
???????? 8.5 ??, ?????????? ? ???? ????????, ????? ??? ? 2.6 ???? ??????? ????????, ?
? ????? ???????? ? 11.0 ??, ?????????? ????? ???? ???????, ? ? 2.7 ???? ??????? ????????, ?????? ? ????? ??????? ????? ??????
???? ??????????? ??? ???????.
??? ??? ???? ????????, ????? ? C. megalonyx megalonyx ???????? ? ??????? ?????.
??????? ????? ?????????? ? ?????? ??????
????? ?????
????? ????
???????????
????????. R/V
????????. R/V
Polarstern, ???????
Polarstern, ???????
111-9, 111-16, 135-1
173-1 ? 184-1
???????????. ???????? ?????????? ? ???????? ??????????? ???? ?????? ? ???????????
?? ????-?????????? ??????. ? ?????????
????? ?????? ????? ??? ? 6-6.8 ???? ???????
????? ??????; ? ????? ?? ???? ?????? ?????? ???? ??? ????? ??????????: ?????????
?? ????? ? ?????? ?????????? ? ????????
5.6-6.3 ????. ? ??????????? ?? ??????????
??????? ?????? ????????? ????? ?????? ?
# 136 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
???????? ??????????? ?? ????? ? ??????
????? ???????????. ????? ????? ????????
????? ? ??????? ????? (? ????????? ??????
10.5 ??) ?? ??????? 184-1, ?? ??? ???????
????? ?????? ? 10.8 ????.
???????????? ?????????? ? ? ????? ???????? ?????. ? ??????? ????? ????????????? ??????????? ??????? ????? ?????????? ? ????????? ?????????, ?? ?????
?????????; ??? ????? ?????? ?????? ??????? ?????? ?????? ? ?????????, ??????
????????????? ????? ???????? ????? ??????
????????? ????? (???. 2, 1-4). ?????? ? ????????? ??????????? ?? ???? ???????? ????????????? ????? ???????, ???????????????.
????? ???????, ?? ????? ??????? ????????????? ????? ? ????????????? ???????????
?? ????-?????????? ??????, ? ? ??????????? ? ????????????? ?????? ?????????????
????? ???????? ????? ??????, ??????????????.
??-????????, ? ?????? ??????? ?????????????? ??? ? C. megalonyx megalonyx ???????
? ????? ???????? ????? ????? ???? ?????.
? ?????????? ?? ???? ???????????, ?????????? ???????? (1993) ??? C. tenuipedis
Pushkin, 1993, ??????? ????? ??????????????????, ? ???????? ????????????? ??????.
????? ? ???? ??????????? ??????? ? ? ??????????? ????????????????. ? ???????????,
????????? ?????? 300 ?, ??????? ???????
?????? ?? ????????? ?????.
??????????? ????????? C. megalonyx
????????? ??????????? ????? 7-?? ? 8-??
???????? II ??????????. ??? ???????? ???????? ??? ???????, ??? 7-? ?????? ???? ?????
????? ?????????. ?? ??? ??????? 7-? ??????
??????, ??????????????, ????? ?????? 8-??
(Hoek, 1881, tabl. IX, fig. 3). ???? ? ??????
(Fry, Hedgpeth, 1969, p. 32) ??????? ???????
???????? II ??????????? ? ???? ???????????
?. megalonyx megalonyx ?? ???? ?????. ????
?? ?? ??????????, ??????? 7-? ???????? ??
?????? ? ????? ????? ???? ?????????????,
????????? ? ????????? ??????? ????? ?? 1/3
????? (??????????? ????? 7-?? ? 8-?? ???????? ?????????? ? ???? ??????????? ? ????????
0.47-0.8). ?????? (1993), ???????? C. megalonyx ?? ?????? ??????? ????????, ???????,
??? 7-? ?????? II ?????????? ? ??? ?????????? ?????????????, ??????? ?????? ?????
?????? ? ????? ??? ????? 57 % ????? 8-??
???????. ? ????????????? ???? ???????????
????????????? 7-? ???????? ??????????? II
???? ?? ????. ? ????? ????????? ??? ??????, ???????? ? ????????????? ?????, ?????????????? ??? ????????????? ?????????, ? 1.5-2 ???? ??????? ????? ?????? (???. 2,
5-10). ? ????????? ??????????? ????? ? ?????? ???? ???????? ???? ???????????. ????
?? ???????? ???????????? ????????, ???????? ????????? 7-? ??????? II ??????????? ?
C. megalonyx megalonyx ?? ????-??????????
??????, ? ???????? ????????? ? ? ?????? ????? ??????? ?? ?????? ????? ???????????
????????. ?????? ?? ????? 7-? ??????? II ??????????? ?????? ?? ?????? ????????? ?????
?? ?????????? ?? ???? ???????? ? ???????????, ????????? ??????????? ????????, ?
???? ??????? ??????? ??? ??????? ??????
???????? 8-? ????????.
???????????? ????????? ? ??? ???????????? ?????????? ??????????? ???????
?????????? ???????? ??????????? III ????.
?????? ? ???????? ???????? ??? ?????????? ??????? ?? ?????? ???? ??????? ???????????? ?????, ????????????? ??? ????
???????????? ???????? ??????????? ?????
? ???????? ? ????? ?????????? ??????,
?????????????? ? ????????? ???????? ?????, ???????????? ???? ?? ????????? ?
??? ???????. ?????, ??? ??????? ??????????? ?? ?????????????, ???? ???? ????????,
??? ????? ????? ?????? ????? ? ????????
? ????????? ????????????? (????????, 1975).
# 137 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
??????????? ?????????? ???? ???????? ?
??? ??????? ?????????????? ???????????.
? ??????? ?????? ?? ???????? ???? ?? 2-4
???? ?????? ?????, ? ??????? ???????????? ? ?? 6-8 ?????. ? ?????? ???????????
??????????? ????? ??????? ? ????? ??????? ?? 10-? ??????? III ??????????. ? ??????
?? ?????? ???? ???????? ??? ??????? ????
??????? ? ? 1.5-2 ???? ?????? 10-? ????????;
? ?????? ?? ?????? ????? ???????????
???????? ???????? ??????? ???? ????? ????
?????????? ? ? 3 ???? ?????? 10-? ????????.
?????? ? ???? ???????????, ????????? ??
???????? 135-1 ? 177-1, ???????? ??????? 10-?
???????? ???? ???????? (?????? ?? 20-22 %
?????? ????????), ?????????, ? ?????????
??????? ?? ?????????? ????. ????? ????,
?? ???? ???????? ????????? ??? ??????? ????????? ????, ????????????? ??????? ???????? ???????????? ?????. ??? ???? ????
??????? ??????? ???????????? ????? 1-??
???? ? ?????? ?????? ?? ???????? (???. 2,
11, 14). ????????? ???????? ? ???? ??????????? ???? ???????????? ??? ??????? C. megalonyx megalonyx.
? ?????????? ?????? (1993) ??????
????? ??? C. tenuipedis, ??????? ? C. megalonyx. ???????? ?????????, ??????????
C. tenuipedis ?? ?????? ????? Colossendeis,
?????? ??????? ????????? ????????? ????? 7-?? ? 8-?? ???????? ??? ? ????? 6-?? ???????. ? ???????????, ?????????? ????????, ??? ????????? ? ????? ???? ????? 1.09,
? ????? ? 1.06. ? ????????????? ???? ??????????? ??????? ???????? ????? ???????????
?????? ? ??????? (0.96-1.03). ? ?????? ???????? ±? ???????? ????? ??????????? ? ?????????? ??????? ????????? ??? ????????
? ??????????? ?? ???? ??????????? (???????? ???????). ???? ?? ???????? ???????
(1993, ?. 56-57), ?????? ??????? C. tenuipedis
?? C. megalonix megalonix ???. ???????
C. tenuipedis ?? ??????? ????????? C. megalonyx megalonyx (?? Fry, Hedgpeth, 1969).
???????? ?????????? ??? ???????????
????? ?????????? ???????? ??? ? ????? ????????? ???? ? ???????????, ????????? ??
???????????? ??????. ? ????? ????? ?? ????? ?????? ????????? ????? ???? ??????????
???????? ??? ???? ???? ? 1.3 ???? ??????
????? 6-?? ???????, ? ? ?????? ????? ? ?????? ?? 30 %. ??????? ????????, ??? ? ?????? ?
???? ?? ?????????? ???????? ????? ??????????? ?? ?????? ????? ???????????. ???, ????????, ? ????? ?? ?????? ????? ???????????
???????? ?? ???? I ???? ??? ???? ????? 0.97,
?? ???? II ???? ? 1.13, ?? ???? III ???? ? 1.03;
? ????? ?? ?????? ???? ???????? ? ?????????????? 1.0, 1.07, 1.05.
??????????? ????????? ? ???????? ????????? ????? ????????? ????? ? ????? 8-??
??????? ????. ???????? ???????? ?????
???? ? ??????, ????????? ?? ????????????
??????. ? ????????? ??????????? ?? ?????
?????? ????? ????? ? ????? 8-?? ??????? ???
???? ????? ???????????. ? ?????? ??????
????? ???? ?? 25-30 % ?????? 8-? ????????.
? ?????? ????? ??????? ?? ?????? ???????, ?
????? ? ?? ???? ??????????? ???????? ????? ????????? ??? ??????? ??????? ????????
8-? ????????.
?????? ???????????? ???????? ????????? ? C. megalonyx megalonyx ?? ?????? ????
????????. ?????? ??????? ?????????? ??
?. megalonyx megalonyx ???? ???????????,
???? ???????????, ?? ????????????? ?????
???????.
Colossendeis megalonyx arcanus
Turpaeva, 2008
Colossendeis
arcanus
????????,
2008, ?. 288-289, ???. 3.
????????. R/V «Polarstern», ANT
XIII/3,1 ?, c?????? 7, 7.II 1996, 71?24ґ3 ?.?.,
# 138 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
???. 3. Colossendeis megalonyx arcanus. ????? ?? ??????? 7 ? 1-9, ????? ?? ??????? 29 ? 10-12. 1 ?
???? ?? ??????? ???????, 2, 10 ? ??????? ?????, 3, 12 ? ?????????? ??????? II ??????????, 4 ? 10-? ??????
III ??????????, 5-7 ? ???????????? ????: 1-?? ???? (5), 2-?? ???? (6), ???? ???????? (7), 8 ? ?????
???????????? ???????????? ????? ?? ?????????? ??????? III ??????????, 9 ? 8-? ?????? ???? ? ??????,
11 ? ????? ?????. ??????? 5 ??
13?44ґ4 ?.?., ??????? 212 ?; 4?, 1?, 1 ???????, ??????? 29, 28.II 1996, 71?30ґ3 ?.?.,
12?27ґ8 ?.?., ??????? 504-529 ?; 2 ??????? ??????????, ??????? 9, 26.II 1996, 71?34ґ8 ?.?.,
12?25ґ9 ?.?., ??????? 560-571 ?; 1 ???????, ??????? 26, 24.II 1996, 71?29ґ2 ?.?., 14?18ґ4 ?.?., ??????? 216-222 ?; ??? «???????? ????????»,
11 ????, 1?, ??????? 882, ????-?????????
?????, 02.XII 1971, 53?17ґ ?.?., 24?54ґ ?.?., ??????? 1729-1879 ?; 1 ??????? ?????????, ??????? 927, 17.XII 1971, 52?40ґ ?.?., 56?08ґ ?.?., ??????? 1640-1664 ?.
C. megalonyx arcanus ?????????? ??
?. megalonyx megalonyx ???????? ? ?????? 7-?? ??????? II ??????????, ????? ??????? ??????? ?????? ? ???????????????
????????????? ??????, ????? ?????????
?????? ?????? ?? ????????? ??????????
???????? III ???????????, ????? ????????
???????? ???????? ????????? ???. ?? ????
?????? ???????? C. megalonyx arcanus ???????? ?????? ? C. megalonyx arundorostris,
?????????? ?? ???? ?????. ?????? ? C. megalonyx arundorostris ????? ??????, ?????? ???
?????????? ? ??????? ?????, 7-? ?????? II ?????????? ????? ????????, ? 5 ??? ?????? 8-??
???????, ?? ?????????? ???????? III ?????????? ?? 4 ???? ?????, ? ??????? ? 3.5 ????
?????? 10-?? ???????.
Colossendeis megalonyx frigida
Hodgson, 1907
Colossendeis frigida Hodgson, 1907, p. 6364, pl. IX, X, figs. 3, 5, 6; Fry, Hedgpeth, 1969, p.
29; ??????, 1993, ?. 62-64, ???. 43, 44 (??????????).
????????. R/V «Polarstern», 1?, ???????
173-1, 30.IV 2000, 63?01ґ2 ?.?., 61?02ґ9 ?.?., ??????? 352-379 ?; ??? «???????? ????????»,
1?, ??????? 217?, ?????????????? ???? 1967 ?.,
# 139 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
???. 4. Colossendeis megalonyx orcadens? ? 1-6; C. megalonyx rugosa ? 7, 8. 1 ? ???? ?? ??????? ???????,
2 ? ??????? ?????, 3 ? 5-9-? ??????? II ??????????, 4 ? 7-10-? ??????? III ??????????, 5 ? ???????? ????
1-?? ????, 6 ? 7-8-? ??????? ????, 7, 8 ? ???????????? ???? ??????? ???????? ????. ??????? 2 ??
49?03ґ ?.?., 60?14ґ ?.?., ??????? 240-300 ?; 1?,
2?, ??????? 756, ?????????????? ???? 1978 ?.,
61?37ґ ?.?., 53?41ґ ?.?., ??????? 400-435 ?.
???????? ???????, ?????????? ?????
???????? ?????????? ??? ?????? ?????????. ????? ??????, ? 2-2.3 ???? ??????? ????????, ?????? ???????? ? ??????? ?????.
7-? ?????? II ?????????? ???????? ? ????????????? ?????, ? 2.5 ???? ?????? 8-?? ???????, 9-? ?????? ???? ????? ??????? ???????
8-??. ?? ?????????? ???????? III ??????????
4 ???? ???????? ?????. ?? ??????? 8-?? ??????? ????????? ??? ?????? ??? ????? ???????? ???????. ??????? ?????? ??????, ??????
?????????, ? 2.4-1.8 ???? ?????? 8-?? ???????
???.
??????, ?????? ???????????????? ?
?????????????? ?????. ??????? ? ????? ?????, ??????, ?ґ??????, ???????, ???????,
???????????, ?????-???????, ? ?????????
????? ???? ????????, ? ????? ????? ?????-
????????? ?????? ? ???????? ????? ???????????, ????? ??????????, ????? ??????????, ? ????? ???????, ? ????????????
????????, ?? ?????? ? ??????? ????????? ? ?
??????? ????????. ??????? ???????? 3-830 ?
(??????, 1993).
Colossendeis megalonyx orcadense
Hodgson, 1908
Colossendeis orcadense Hodgson, 1908,
p.184; Helfer, Schlottke, 1935, p. 294.
Colossendeis orcadensis Stock, 1963, p. 334,
fig. 7.
Colossendeis megalonyx Fage, 1956, pp. 177178, figs. 18-20.
Colossendeis megalonyx orcadense Fry,
Hedgpeth, 1969, pp. 34-35, figs. 7, 8, 11-14, 22,
23; ????????, 1991, ?. 42.
????????. 1?, ??? «??????», ???????
2629, 23.XI 1988, 25?12ґ2 ?.?., 35?41ґ8 ?.?., ??????? 1530-1200 ?.
# 140 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
???? ????? ???? ?????? ????? C. megalonyx ???? ??????? ??????????? ?????????
??????????? ? ????? ????? ?????????????
??????? ?????? ?????????? ?????? ??????.
???? ????????? ?? ????????, ??? ??? ? ?????????? ??????? ?????? ??????? ????????
??????.
???????? ?????????? ??????????????????. ?????? ??????????? ????? ???????? ?????????? ????? ?????? ?????????.
???????? ????? ??????? ???????? ?????????
??? ?????? ??????? ????????? I ????. ?????? ???? ????????. ??????? ????? ?????????, ??????????????, ???? ???. ????? ? 1.5
???? ??????? ???????? ? ? 4.5 ???? ???????
????? ?????????? ??????, ?????????????? ? ????????????? ?????, ???????? ?
??????? ????? ? ? ?????; ????? ?????? ??????
??????? ????. ??????? ??????, ??????????
?????? ???? ??????? ?? ???????? 1-? ???????? ??? IV ????. ?????????? II ???? ? 2.5
???? ??????? ????????, ???????????????,
??????, ???????; ???? ?? ????????? ???????? ??????? ????? ????????? ?????? ???????. ????? ??????? ?????? II ??????????
2-?, 4-? ?? ј ?????? 2-??, 5-? ?????? ?? ј
?????? 6-??, 7-? ??????, ?????? ???????? ?
????????????? ?????, ?? ????? ????? ???????? 8-?? ???????, 9-? ?????? ??????? ?????? 8-??. ?????????? III ???? ? 4 ???? ???????
????????; ?? ????? ??????? ??????? 4-? ?
6-?, ????? ?????????? ???????? ??????????? ? ????? ?????. ?? ??????? ???????
?????????? ???????? 2 ???? ???????? ????? ? ???????? ?????? ? ????????????????
???????? ?????? ?????. ??????? ??????
?????????, ?????????, ??????? ?????? 10?? ???????. ????????? ???? ????? ? 10 ???
??????? ????????. ????? ??????? ??????
??? 4-?, 5-? ????? ? 1.5 ???? ?????? 4-??, 6-?
??????? ?????? 5-??; 7-? ?????? ???????
8-??, ? ?? ????? ????? ?????? ????? 6-??
???????. ?????? ??????, ????? ??????, ????
?? 12 % ?????? 8-?? ???????.
??????? ????? ? ???????????.
???????? ? 8.0, ????? ? 11.8, ??????? ?
1.9;
II ?????????? ? 19.5 (1-? ?????? ? 0.3,
2-? ? 5.6, 3-? ? 0.9, 4-? ? 4.2, 5-? ? 2.1, 6-? ? 2.8,
7-? ? 0.9, 8-? ? 1.4, 9-? ? 1.3);
III ?????????? ? 33.9 (1-? ?????? ? 0.6,
2-? ? 0.7, 3-? ? 0.9, 4-? ? 9.6, 5-? ? 5.3, 6-? ?
11.0, 7-? ? 1.8, 8-? ? 1.4, 9-? ? 1.3, 10-? ? 1.1),
??????? ? 0.6;
???? III ???? ? 77.0 (1-? ?????? ? 1.5, 2-? ?
1.7. 3-? ? 1.5, 4-? ? 24.3, 5-? ? 18.3, 6-? ? 15.2,
7-? ? 8.2, 8-? ? 6.3), ?????? ? 5.6.
C. megalonyx orcadens? ?????????? ?? ?????? ???????? C. megalonyx ????????? ?????????? ?????????. 7-? ?????? II ??????????
? ??? ??????????? ??????? ?????? ????????
8-?? ???????, ? 9-? ?????? ??????? ?????? ?
?????? 8-?? ???????. ??????? ????? ???????
????, ??? ? ?????? ????????; ????? ???? ?????????? ??????? ? ??????????????????,
???? ?? ??????? ??????. ????? ???????????? ???????? ? ????????? (? ?????????????? ???? ?????????? ????? ?????? ? 4.5 ????
??????? ?????? ????????????? ????????), ?
????? ????????? ????? ????????? ??? ?????
????? ????? ?? 8-?? ???????.
C. megalonyx orcadens? ? ????????????? ??????. ??? ??????? ??? ?????? ? ??????? ??????, ? ?????? ????? ??????????
???????? (??????? ?? ???????). ??????? ??
??? ????????? ? ??????????? ? ???????? ?????? ??????? ?????? ?? ???????? ????? 4 ????? ?. ?????? ????? ???????? C. megalonyx
orcadense ? ????? ????? ??????. ? ????????????? ?????? (?? ?????? ?????????) ??
??? ????????? ?? ???????? 2740-3260 ? ? ?
???????????? ???????: ? ???-???????? ????? ?? ???????1530-1200 ? ? ??????-?????????
????? ?? ??????? 3410 ?.
# 141 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
Colossendeis megalonyx rugosa
Hodgson, 1907
Colossendeis megalonyx weddellensis
Turpaeva, 2008
?olossendeis rugosa Hodgson,1907, p. 6466, pl. IX, fig. 7; Calman, 1915, p. 16-17; Gordon,
1938, p. 11; ??????, 1993, ?. 48-50, ???. 29, 30;
????????, 1998, ?. 66;.
M???????. R/V «Polarstern», ANT XIII/3,
1?, ??????? 11/BT, 13.II 1996, 73?22ґ6 ?.?.,
21?10ґ6 ?.?., ??????? 338-333 ?; 1?, ??????? 6/
AGT, 25.II 1996, 71?32ґ3 ?.?., 13?42ґ8 ?.?., ??????? 354-362 ?; 1?, 3?, 2 ???????, ??????? 29/
BPN, 29.II 1996, 71?30ґ3 ?.?., 12?27ґ8 ?.?., ??????? 504-528 ?.
????? ??????????? ??????? ????? ???-
Colossendeis weddellensis ????????, 2008,
?. 885-886, ???. 1.
????????. R/V «Polarstern», ANTXVII/3.
1 ?, ??????? 111-19, 5.IV 2000, 71?17ґ5 ?.?.,
11?28ґ3 ?.?., ??????? 112 ?; 1?, 1 ??????? ?????????, ??????? 111-16, 5.IV 2000, 71?07ґ5 ?.?.,
11?27ґ6 ?.?., ??????? 80 ?.
?? C. megalonyx megalonyx ? ?????? ???????? C. megalonyx ???? ?????? ??????????
????????? ?????????? ?????????: ???????? ??????? ??????? ??????, ???????????
??????? ??????? ?????????? ???????? III
?????????? (???????? ??????? ????? ???????? ?????, ?? ??????? ??????? ????????
???????, ? ??????????? ???????? ??????
?????), ???????? ??????? ????????? ??
??????? 7-?? ? 8-?? ???????? ?????????
??? ? ????? ???????? ???????? 8-?? ???????
???? ? ???????????? ?????????, ???????????? ????, ???????? ???????? ?? ???????
???????? ???. ?? ?????????? C. megalonyx
rugosa ????????? ?????????? ?? C. megalonyx
megalonyx. ??????? ????? ? ???? ? ??????????????? ?????????, ????? ????? ??????????????. ? ?????????????? ???? ????? ?? ??????? 29/BPN ????? ????? ??????????????,
? 1.4 ???? ??????? ???????? ? ? 6.4 ????
??????? ????? ??????. ?????????? II ????
???????????????, ?? 6-? ?????? ? 3.4 ????
??????? ????? ??????, 7-? ?????? ?????
????????, ? 3 ???? ?????? 8-??, 9-? ??????
??????? ??????? 8-?? ???????. ?? ?????????? ???????? III ??????????? 1 ??? ???????
???????? ?????, 2-? ??? ?????? ????????
????? ? ????? ???????? ? ????? ?????????
?????? ????? ????????? ????? ?????. ????????? ???? ???????. ?? 6-? ?????? ??????? ????? ?????????? 7-?? ? 8-?? ????????,
?????? ?????? (7-? ?????? ?? 8-10 % ???????
8-??). ?????? ?? ????? ????? ????? 8-?? ???????.
???? ?????? ??? ???????? ?? ?????????????? ??????? ?????????????? ??? ? ? ????
??????? ? ????????? ??? ?? ???? ?????.
? ???? ???????? ????????? ???????. ??????? ?? ???????? 229-650 ?.
???.
??? ????? ?? ???????????? ???? ??????
? ????????, ??????? C. megalonyx, ????????? ? ???? ????????, ?? ?????????? ???????? ? ??????????? ?????? ????? ????? ?
?????????? ???? ?? ????? ????? ?????-?????
??????????. ??? ????????? ??? ?????????
???? ??? ??????????? ???????? C. megalonyx, ??????? ? ???? ????? ??????? ?? ?????? ??????? ??????.
???? ??? ???????????
???????????? ???????????
???????? Colossendeis megalonyx
1(2) ????? ??????, ??????????? ? ??????? ? ?????????? ??????, ? 6-7 ??? ???????
????? ?????? ? ? ??????????? ???????????
? 1.7-2.2 ???? ??????? ????????. 7-? ?????? II
?????????? ??????, ???????? ? ????????????? ?????, ? 1.7-2.5 ???? ?????? 8-?? ???????.
?? ?????????? ???????? III ?????????? 2
???? ???????? ????? ? ???????? ? ??????-
# 142 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
???? ??????, ?????????????? ? 7-8 ?????
?????. ?? ????????? ????? ????? ??????, ????
???????? ???????, ??????? 8-? ???????? ???
??? ??????? ? ?????, ?????? ??? ?????? 8-??
??????? ?? 50-20 %..................................................
................................?. megalonyx megalonyx Hoek
2(1) ????? ????????? ????
3(8) ?? ???????? ????????? ??? ???????
???? ??? ???????.
4(5) ?? ???????????, ???????? ?? ???????
???????? ???, ??????????? ????????? ???????????? ???? ??????????...?.?..
???????C. megalonyx rugosa Hodgson
5(4) ????? ?? ??????????? ???, ????????? ???? ????????? ?????????.
6(7) ?? ?????????? ???????? ??? ??????????? ???? ???????? ? ????? ??????? ???????, ?? ??????? 8-? ???????? ?????? ????
?????? ???????, ????? ????? ??????? ?????????????................???????????
.????.???.C. megalonyx frigida Hodgson
7(6) ?? ??????? ?????????? ????????
??? ?????? ??????? ?????? ?? ???????? ?????? ???????, ????? ? 1.3 ???? ??????? ???????? ??????????????..................
???.?...C. megalonyx weddellensis Turpaeva
8(3) ??????? ??????????? ???????, ???
?????, ???????, ???? ????, ?? ???? ???????,
??????.
9(10) ????? ?? ????? ????? ????? ????????, ?????? ????????? ??? ??????? 8-?? ??????? ? 1.5-1.3 ????????????....?????
..........??...??.C. megalonyx scoresbii Gordon
10(9) ????? ??????? ????????.
11(14) ????? ? 2.5 ???? ??????? ????????, ?????? ????????? ??? ?????????????? ??
40 % ?????? 8-?? ???????.
12(13) ????? ? 8 ??? ??????? ????? ??????; 7-? ?????? II ?????????? ??????, ???
??????? ?? ??????, ? 2.5-3 ???? ?????? 8-??
???????. ?? ??????? ??????? ??????????
???????? III ?????????? ?? 2 ???? ????????
????? ? ???????? ? ??????? ??????, ?????????????? ? 6-8 ???????? ????????.?
.....??..??.?.C. megalonyx arcanus Turpaeva
13(12) ????? ? 11-12 ??? ??????? ?????
??????; 7-? ?????? II ?????????? ????????,
? 4 ???? ?????? 8-??. ?? ?????????? ????????
III ?????????? 4 ???? ???????? ????? ??....
?......C. megalonyx arundorostris Fry, Hedgpeth
14(11) ????? ? 1.5 ??? ??????? ????????.
15(16) 7-? ?????? II ?????????? ??????
Ѕ 8-?? ???????, ?????? ????????? ??? ??????? ?? 8-?? ???????, ??????? ????? ???????,
???????????????, ????? ???????. ?????.
..........??..????.C. megalonyx tenera Hilton
16(15) 7-? ?????? II ?????????? ???????
Ѕ 8-?? ???????, ?????? ????????? ??? ??????
?? 8-?? ???????, ??????? ????? ??????, ??????????????, ????? ??????????? ??? ?? ???. ...
???...??..C. megalonyx orcadense Hodgson
??????????
?????????????? ???? Colossendeis ?
????????????? ??????????? ?????. ??????????? ?? ??? ??????? ?? ?????? ? ????
?????????????? ??????????????? ?????????????? ???, ????? ???????? (-0.8-0.0°?),
?????????? ?????????? (6.9-9.5 ??/?), ??????? ????????? 34.7?, ? ? ?????????????
??????? ? ?? 35? (??????????? ? ??., 2000).
????? ?????????? ????????? ?????????????? ?????????????? ????????, ??????????????, ??-????????, ?????????? ????
????? ???????? 60° ?.?. ? ??????????? ????
?????????? ? ?????? ???????. ?????? ? ????? ????? ?????????????? ?????? ??????????? ???? ????????? ????? ?? 30° ?.?.
????? ????????? ? ??????? ????????
???????? ?????? ??? ???? ???????? ? ????????? ????? ?????? ??? ???????????????
??????????????? ???????, ????????? ????
????? ?????? ??????. ?????? ??? ???? ????? ????????? ????? ???????. ?????????
# 143 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
???? ???????? ????? ?????????? ? ?????????????? ????????, ????????? ????????,
????????????? ?????????, ??? ???????
??????? ??? ????????????? ? ???????????
? ????????????? ????? (??????????? ? ??.,
2000).
????????????? ??????????, ????????? ???????????? R/V «Polarstern» ? 1996
? 2000 ??., ? ????? ?????????????? ???????? (??????,1993; Child, 1995) ?????????
??????? ???????? ??????????????? ???????????? ? ???? ???????? ????? Colossendeis.
??????? ????? ?? ??? ?????? ?????????????? ? ?????????????? ?????, ?????? ????????? (????) ????? ??????? ??????????. ???,
?????? ? ???? ???????? ????????? C. potentis,
????????? ? ?????? ???? ????????. ? ????
???????? ? ? ????? ????? ??????????????
?????? ???????? C. pseudochelata Pushkin,
1993 ? C. smirnovi Pushkin, 1988. ???? ???,
C. fragilis Pushkin, 1993, ??????? ? ???? ????????, ? ????? ????? ?????????????? ?????? ? ? ????????????? ??????? ?????????????? ???. ???? ???, C. wilsoni Calman, 1915,
??????? ? ???? ???????? ? ? ???? ?????. ?
???????? ????? ?????????? ? ? ???? ???????? ? ????????????? ??????? ??????????????
??? ? ??????????? ?. avidus Pushkin, 1970,
C. brevirostris Child, 1995, C. korotrevitschi
Pushkin, 1984 ? C. longirostris Gordon, 1938.
????????? ???? ????????????????????.
?? ??? C. drakei Calman, 1915, C. glacialis
Hodgson, 1907, C. lilliei Calman, 1915, C. scotti
Calman, 1915 ? C. tortipalpis Gordon, 1932
???????? ?????? ?? ?????????????? ???,
C. australis Hodgson, 1907 ? C. robusta Hoek,
1881 ??????????? ????? ? ????? ????? ?????????????? ??????.
????? ?????? ???????????????? ????? ???????? C. megalonyx. ?? ??? ?????? ???????? ???? ??????? ? ???? ???????? (???. 5).
?? ??? ???????? ????? ??????????? C. megalonyx megalonyx ? ????????????????????
??????, ??????????? ?? ????? ? ? ???????
????? ????? ?????????????? ??????. ????? ??????????????????? ???????????????
? ???????? ??????? ??????? ? C. megalonyx
frigida, ?????????????? ? ????? ?????????
???. 5. ??????????????? ???????? ?. megalonyx ? ??????? ??????: 1 ? C. megalonyx megalonyx,
2 ? C. megalonyx orcadense, 3 ? C. megalonyx tenera, 4 ? C. megalonyx arcanus, 5 ? C. megalonyx frigida,
6 ? C.megalonyx scoresbii, 7 ? C. megalonyx weddellensis, 8 ? C. megalonyx arundorostris, 9 ? C. megalonyx
rugosa. ????????? ???????? ?????????????? ???? ??????????
# 144 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
?? ??????????? ??????. ????????? ???????
C. megalonyx, ??????????? ? ???? ????????, ?????????????? ????? ??????. ?? ???
C. megalonyx weddellensis ???????? ???? ?????? ? ???? ????. C. megalonyx arcanus ??????
? ???? ???????? ? ? ????? ????? ?????????????? ?????? (? ????-??????????? ?????? ? ?? ?????? ???????????? ?????????).
?. megalonyx rugosa, ????? ???? ????????,
???????? ?? ???? ????? ? ?????????????? ??????? ?????????????? ???.
????????? ?????? ??????? ? ???? ???????? ? ? ????????? ? ???? ??????? ?? ?????????. C. megalonyx s?oresbii Gordon, 1932 ??????? ?? ??????????? ?????? ? ? ????????????
????????, C. megalonyx arundorostris ????????
?????? ?? ???? ?????, C. megalonyx orcadensis
??????? ? ????? ????? ?????????????? ?????? ? ???-???????? ????? ?????????? ??????,
?. megalonyx tenera ?????? ? ???????? ?????
?????? ??????.
??????????? ????? Colossendeis ????
???????? ? ?????????????? ?????. ??????
????????? ???? ? ??????? ????????? ?
????? ???????? ???? ??????. ????????????
????????????? ????? ? ???????? ?????????? ???????????? ?? ???. 6 ? 7. ?? ????????
????????????? ????????? ? ????????????
??? ?? ????? ?????? ???? Colossendeis ????
???????? ????? ????????? ?? 3 ??????. ?
1-? ?????? ????????? ???? ? ????????????
?????????? 250-300 ?: C. scotti, C. wilsoni,
C. robusta ? ?. pseudochelata, ??????? ???????
????????????? ????????? ??????? ??????????? ?? ??????? 50-150 ?, ? ????? ????? ????????????? C. brevirostris ? C. megalonyx
rugosa, ????????? ? ?????? 300-600 ? ? 300650 ? ??????????????. ?? 2-? ?????? ?? ??????? ????, ???????????? ???????? ???????
???????????? ?? ??????? ????? 1000 ? ? ??
??????? ??????????? ?? ??????????? ????????? ? ???????, ??????? ?? ??????????????
???. 6. ???????????? ????????????? ????? Colossendeis, ??????????? ? ???? ????????: 1 ? C. australis,
2 ? C. drakei, 3 ? C. glacialis, 4 ? C. fragilis, 5 ? C. potentis, 6 ? C. lilliei, 7 ? C. korotkevitschi, 8 ? C. scotti,
9 ? C. wilsoni, 10 ? C. robusta, 11 ? C. pseudochelata, 12 ? C. brevirostris, 13 ? C. avidus, 14 ? C. smirnovi,
15 ? C. tortipalpis, 16 ? C. lonirostris
# 145 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
?????? ?????????? ?? ??????? 800-1200 ?
(??????, 1985; ??????, ????????, 1998; ???????????, ?????? ?????????). ? ???? ??????
????????? C. australis, C. drakei, C. glacialis,
C. fragilis, C. lilliei, C. korotkevitschi, C. megalonyx frigida, C. megalonyx scoresbii, ? ?????
????? ????????????? ?. avidus, C. smirnovi
? C. tortipalpis. ? 3-? ?????? ??????? ??????? ???? ? ????? ??????? ????????????
?????????? ? 2-5 ?????? ?. ?? ?????????
? ???? ???????? Colossendeis ? ???? ?????? ????????: ?. longirostris ? C. megalonyx
megalonyx, ????????????? ?? ??????? ??????????? ?? ??????????? ?????? (???. 7),
?
?????
?????????????-??????????
?. megalonyx ar?anus. ????? ????, ?? ???????? ? ??? ?????? C. megalonyx orcadens? ?
C. megalonyx tenera, ?? ??????????? ? ????
????????, ?? ?????????????? ??????? ? ??????????????? C. megalonyx megalonyx. ???
????? ???? ?????? ????????? ? ???????????? ???????: C. longirostris ? ? ?????????
?????, C. megalonyx megalonyx ? C. megalonyx
arcanus ? ? ?????????????, C. megalonyx
orcadens? ? ? ????????????? ? ?????????,
C. megalonyx tenera ? ? ????? ?????.
??? Colossendeis ? ????? ?? ??????????
????? ???????? ?????? ??????????? ? ???????? ? ???????? ???????? ????? ?????? ????????? (???. 8), ?? ????, ?? ????????????
?. ?????? (Ekman, 1953), ? ??????????????
???????? ?????, ??? ?????????? ????? ???????? ????? ????? ????. ? ??????????? ?
?????????????? ????? ????????? ? ??????
????????? ??????? 27 % ????? ????? ????,
? ???????? ?????? ?????????????? ? ?????? ??????? ? ?? 13 %, ? ? ??????????? ?????
??????????? ????? 2 %. ?????? ?????????????
???? Colossendeis ????? ???????? ???????
??????????????? ? ??????? ??????, ????????
???? ?? ?????? ? ?????? ????????? ?????????? ?????? (???????, ????????, 2006). ?????
???.
7.
????????????
?????????????
????????
Colossendeis
megalonyx
Hoek:
1 ? C. megalonyx megalonyx, 2 ? C. megalonyx orcadens?,
3 ? C. megalonyx tenera. 4 ? C. megalonyx arcanus,
5 ? C. megalonyx frigida, 6 ? C. megalonyx scoresbii,
7 ? C. megalonyx weddellensis, 8 ? C. megalonyx
arundorostris, 9 ? C. megalonyx rugosa
???? ??????????? ??????????? ????????????? ????????????? ????? ??????? ?????? ??
?????????????? ???? (????????, ????????,
1997). ?? ???? ???????????, ??????????????
??? C. megalonyx ???? ???????????? ????? ?
???? ???????.
???????? ?????????? ????? ? ???????? ???? Colossendeis ???????????? ?????????? ??????????????? ????????? ??????????????? ???????? ??? ? ???? ???????? ?
?????? ??????????????? ??????????? (???????????, 2003).
# 146 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
???. 8. ?????????? ??????????? ??????????????? ????????????? ????? ???? Colossendeis ? ???????
??????: 1 ? ????, ????????? ? ???????, 2 ? ? ???????? ????? ?????????????? ??????, 3 ? ? ???????? ?????
?????? ??????, 4 ? ? ??????????-?????????????? ????? ? ??????????? ????????? ? ?????? ?????????,
5 ? ? ?????????? ? ?????????????? ?????
??? C. megalonyx ??? ???? ? ????? ?????
??????????????? (???. 5) ?????? ???????????
????????? ??????????????? ??????????????? ??????? ? ??????????? ?????????????? ???
? ??????? ?????? ? ?????????? ??????? (????????, ????????, 1997). ??? ???? ????????????? ???? ???? ???????? ? ??? ???????? ?????
????????? ?? ??????-???????? ????? ??????????????? ???????????, ???????????, ? ?????????, ????????????? ??? ????????? ??????????????? ????? (???????????, 2003, ?. 72).
??????????
??????? ????????? ? ?????? ??????????
????????, ? ????? ???????????? ?????????????
???????????? ? ??????????? ???????????????
????? ? ???????? ???? Colossendeis ????????? ????????????, ??? ? ???? ???????? ? ????? ?? ?????????? ??????????????? ??????????
? ??????? ????????????? ??????? ??????????
??????????????? ?????? ???? ? ??????????
??????????? ??? ?????????????? ? ?????????????, ????? ? ????????? ??????.
?????? ??????????
1.
2.
3.
4.
5.
6.
7.
??????????? ?.?., ?????? ?.?., ???????? ?.?. (2000) ????????????? ?????? ? ???????
?????? ??????: ????????????? ?????????, ?????? ????? ? ??????? ?? ????????. ??-? ??????????? ??. ?.?.??????? ???. ???. ????, ?., 105 ?.
?????? ?.?. (1985) ????????. ???????? ????????? ? ????? ??????. ?????, ?., 501 ?.
?????? ?.?., ???????? ?.?. (1998) ??????????? ?????? ????? ?? ?????? ? ??????? ??????????. ??????????? ??????? ?? ???? ??????. ??????????. ????? ?? ???????? ??????????
???. ?????, ?., ?. 140-147.
???????? ?.?., ???????? ?.?. (1997) ?????????? ? ????????????? ??????? ????? ????????? (Pycnogonida). ???????? ??, ????? ????????????? ?3: 306-314.
??????????? ?.?. (2003) ????????????? ????????? ? ??????? ?????????? ??????????.
???-?? ?????, ?., 296 ?.
?????? ?.?. (1970) ????? ???? Colossendeis (Pantopoda). ????. ????. 49 (10): 1488-1496.
?????? ?.?. (1993) ????? ??????????????? (Pycnogonida) ?????? ??????. ????????????
????? ?????. XX (XXX). ?????????? ????????????? ???????????? ????????? ?????????????? ??????????. 8. ?-??????????????????? (???????), 397 ?.
# 147 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
8.
??????? ?.?., ???????? ?.?. (2006) ????????????? ??????? ????? (Pycnogonida) ????????
????????? ? ?? ??????????????? ? ??????? ??????. ??????????? 46 (1): 63-58.
9. ???????? ?.?. (1974) ??????????????? (Pycnogonida) ???? ?????? ? ?????????? ???.
????? ??-?? ??????????? 98: 277-305.
10. ???????? ?.?. (1975) ????????? ????????????? ???? ??????????????? (Pycnogonida), ????????? ? ??????-???????? ? ???-????????? ??????? ?????? ??????. ????? ??-?? ??????-
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
????? 103: 230-246.
???????? ?.?. (1991) ????????? (Pycnogonida), ????????? ?? ?????? ???-????????? ?????? ? ? ?????????? ?????. ????. ????. 70 (12): 33-43.
???????? ?.?. (1998) ?????????????? ?????? ???????? (Pycnogonida) ?? ????????? ????? ???? ????????. ?: ???????? ?.?., ?????? ?.?. (???. ???.) ?????? ?????????????? ???????. ?? ???, ?., ?. 56-68.
???????? ?.?. (2008) ????? ???? ??????? ?????? ???? Colossendeis (Colossendeidae,
Pycnogonida) ?? ???????? ????? ???? ????????. ????. ????. 87 (7): 885-890.
Calman W.T. (1915) Pycnogonida. British Antarctic (Terra Nova) Expedition, 1910. Zoology 3
(1): 1-74.
Child C.A. (1995) Antarctic and subantarctic Pycnogonida: Nymphonidae, Colossendeidae,
Rhynchothoraxidae, Pycnogonidae, Endeidae, and Callipallenidae. In: Cairns S. (ed.)
Antarctic Research series, vol. 69. Biology of the Antarctic Seas. XXIV, Washington, D.C.,
p. 1-165.
Child C.A. (1998) The marine fauna of New Zealand: Pycnogonida (Sea Spiders). Department of
Invertebrate Zoology National Museum of Natural History Smithsonian Institution, Washington,
D.C. 20560. USA. NIWA. Biodiversity Memoir. 109: 1-71.
Ekman S. (1953) Zoogeography of the sea. Sidgwick and Jackson, London, 417 p.
Fage L. (1956) Les Pycnogonides (exel. le genre Nymphon). Galathea Reports 2: 167-182.
Fry W.G., Hedgpeth J.W. (1969) Pycnogonida. 1. Colossendeidae, Pycnogonidae, Endeidae,
Ammotheidae. Fauna of the Ross Sea. Part 7. Memoirs of the New Zealand Oceanographic
Institute 40: 1-139.
Gordon I. (1938) Pycnogonida. Scientific Reports of the Australasien Antarctic Expedition,
Zoology and Botany 2 (8): 1-40.
Hedgpeth J.W. (1943) On a species of pycnogonid from the North Pacific. Journal of the Washington
Academy of Sciences 33 (7): 223-224.
Helfer H., Schlottke E. (1935) Pantopoda. In: Dr. H.G. Bronns (ed.) Klassen and Ordnungen des
Tierreichs. vol. 3 (4). Akademische Verlagsgesellschaft m. b. H., Leipzig, p. 1-314.
Hilton W.A. (1943) Pycnogonids from the Pacific. Family Colossendeidae. Journal of Entomology
and Zoology of Pomona College 35 (1): 2-4.
Hodgson T.V. (1907) Pycnogonida National Antarctic Expedition 1901-1904. Report of the
National Antarctic Expedition 1901-1904. Natural History 3: 1-72.
Hodgson T.V. (1908) The Pycnogonida of the Scottish National Expedition. Transaction of the
Royal Society of Edinburgh 46 (1), Part 6: 159-188.
Hoek P.P.C. (1881) Report on the Pycnogonida dredged by HMS Challenger during the years
1873-1876. Reports of the Scientific Results Exploring Voyage of HMS Challenger 3 (10): 1-167.
# 148 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ??????? ????? ???? Colossendeis (Colossendeidae, Pycnogonida) ???? ?????????
27. Stock J.H. (1963) South African deep-sea Pycnogonida, with description of five new species.
Annals of the South African Museum 46, Part 12: 321-340.
Sea Spiders of the Genus Colossendeis
(Colossendeidae, Pycnogonida)
from the Weddell Sea and Adjacent Waters
Elena P. Turpaeva and Alexey C. Rajsky
P.P. Shirshov Institute of Oceanology RAS
36, Nahimovski prospect, Moscow, 117997 Russia
Seventeen species and five subspecies of Colossendeis were sampled during the expeditions of R/V
Polarstern ANT XIII/3 and ANT XVII/3 in 1996 and 2000 in the Weddell Sea. Polytypical species
C. megalonyx Hoek, 1881 was frequently met in this collection. Subspecies C. megalonyx megalonyx
was numerous; its traits variability is analyzed. An identification key for subspecies of C. megalonyx
has been proposed. Based on the distribution of species and subspecies of Colossendeis as well as
intraspecific variability, it is assumed that it is in the Weddell Sea, with its complicated hydrology and
relief features, that the speciation of the genus takes place. Representatives of Colossendeis from the
Weddell Sea are found now in the Atlantic, the Pacific and the Indian Oceans.
Keywords: Pycnogonida, identification key for the subspecies of Colossendeis megalonyx, speciation
in the Weddell Sea.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
????? ???????? ???????? ? ????????? ????????? ????????? ??????????? ??. ?.?. ???????. ????? ????????? ????????????? ????? ???????????????
?????????? ??????? ???????? ?????????????? ???
??? ?????? ? ??????????????? ????? ???????? ???????????, ???? ? 1947 ?. ????? ?? ?????? ??????????.
???????? ??? ??????? ???? ?????? ???????? ????????????, ?????????? ?????????????? ????????, ?????
???????? ???????? ?????? ?????? ??????????? ? ??????? ???????? ? ???????????? ??????????? ?????????
? ??????????? ??????????? ???????? ???????. ?? ??-??
?????? ?.?. ???????? ????? ???????? ???????? ????
???????????? ??????????? ? 1950 ?. ??? ? ????? ?????????????, ???????? ????????? ?????????. ??????
??? ??? ???????? ? 1959 ?. ????? ???????? ???????? ????????????? ?????????? ? ????? ?????: ????????? ???????? ???????, ? ????? ??????????, ?????????? ? ??????? ?? ??? ?????
? ???????????????? ??????????. ?? ????????? ?????????? ?????????? ?????? ?????????
???? ??????????? ?????? ???????????????? ?????????? ??????????, ? ??? ????? ? ?? ???????????? ????????, ??? ????? ????? ?????????? «????????????? ?????? ?????????? ??????????» (1987) ? ????????? ?????? ?????????? ???????????.
????????????, ?????? ??????????? ??????? ???????? ??????????????, ????? ????????
?????????? ? ??????? ??????????? ? ???????? ??????? ??????, ??????????????? (Pycnogonida),
? ? ????????? ???? ??? ???? ????? ???????? ? ?? ??????? ?????????????. ??? ??????????????
?? ????? ???????? ????????????? ?? ????????????, ?????????? ? ??? ??????? ???????????,
??????????? ??????????. ???????? ?? ???? ?????????? ???????, ????? ???????? ??????????
??????? ???????? ? ????? ??????????, ???????? ???? ?????? ??? ??????.
????????? ????? ???????, ???????? ?????????????, ?.?. ???????? ????? ? ????? ??????
???????????? ????????????? ????????????? ? ????????.
???????, ???????????? ????????? ???????? ???????, ??????????, ?????????????, ?????????? ????????? ? ??????????? ??????????????????? ???????? ??????????? ????? ???????? ???????? ? 90-??????!
?????? ????????????? ????,
??????? ??????? ????????? ?????,
???????????? ??????????? ????????? ???
?.?. ??????????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2013 6) 151-162
~~~
??? 616.155.34
????????? ?????????? ???- ?
????-????????? ????????????
? ????????????? ???????????? ? ???????????
? ???????? ?????????????? ?????
?.?. ?????????,?, ?.?. ????????*
?
??? ??????????? ??????? ?????? ?? ????,
?????? 660022, ??????????, ??. ????????? ?????????, 3?
?
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ?????????, 79
Received 04.11.2012, received in revised form 03.12.2012, accepted 27.12.2012
??????????? ?????????????? ??????? ????????????? ???????????? ????? ???????????,
?????????????? ????????? ????? ?????? ? ???????????????? ??????????? ????????????
? ???????? ? ??????? ????, ? ????????? ????? ???????? ?????????????? ??????????.
? ????? ????????????????? ??????? ? ????????????? ???????????? ????????????
???????? ?????????????? ????????: ?????????? ? ???????? ??????????, ?????????????????? ??????????????? ???????, ? ????? ???????????? ???????, ????????????
??????? ??????? ?????????????????? ??????? ? ???????????? ??????. ?? ???? ?????????
? ????? ????????????????? ??????? ? ??????????? ??????????? ?????????? ????????
?????????? ??????????? ???- ? ????-????????? ????????????, ??? ?????????? ????????
????????????????? ?????????, ??????? ????????????? ??????, ?????????? ???????????
????????. ? ?????? ?????? ??????? ?????????????? ?????????????? ????????? ?
????????????? ???????????? ? ??????????? ?? ??????????.
???????? ?????: ?????, ???????? ???????, ????????????? ???????????, ??????????
?????????????????, ?????????????? ?????????????????.
????????
???????? ??????? ???????? ????? ??
???????? ?????????????? ???????????, ?????????? ??????? ????????? ?? ??????-
*
????? ????????? ???????? ??????? ?????,
? ??? ????? ? ?????????-???????????????.
?????????? ????????????, ? ??????? ?????????? ???????? ?????????? ? ??????????
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: kpbazarin@gmail.com
# 151 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ????????? ?????????? ???- ? ????-????????? ???????????? ? ??????????????
????????????? ???????????, ?????? ???????? ???????????? ????????? ???????????????? ?????????, ????????? ????? ???
«?????????? ?????????????». ? ?????????
10-15 ??? ?????????? ??????????? (exercise
immunology, sport immunology) ???????? ? ??????????????? ??????? ??????????. ? ????
?????? PubMed ?????????? ????? ???? ?????
??????????, ??????????? ???????? ??????? ?????????? ???????????? ?? ????????
???????. ????? ??????????? ??????? ? ?????
????? ??????? ?????? ? ???????????????
???????, ??????????????? ???????????
?????????? ?? ????????? ? ?????????? ?????????????? ??????????? ??????? ?????????????? ??????????, ?????????? ? ?????????? ???????????? ????????? (Friman,
Wesslen, 2000; Shephard, 2000; Koch et al.,
2007).
????????, ??? ??? ?????????? ????????
?????????? ?? ?????? ???????????? ???? ?????????? ??????????? ???????? ???????????? ???????????, ?? ???????? ? ????????
?????????? ???????????, ??? ??? ?????????
(?????????????, ???????, 2003), ? ???????????????? ?????? ????????????? ???????????
?????????????? ? 40 % ???????????, ???????
????????????? ???????????? ????? ? ????????????? ?????????? ??????????? ??????? ?
????????? (?????????, 2009).
??????? ?????????? ???????? ????????
????????? ????????? ???????? ???????, ????????????, ? ?????????, ? ???????? ?????? «?????????????? ??????» ???????????,
???????????? ??????????, ???????? ??????????????? ?????????? NK-?????? (natural
killer), ??? ?????? ?????? ?? 3 ?? 24 ? ?????
????????? ???????? ? ??????????? ?? ?? ???????????? ? ?????????????. ?????? ????????? ????????? ???? ????? ?????????? ? ???
??????, ???? ???????????? ???????? ????????? 1,5 ?, ? ????????????? ?????????? 5575 % ?? ?????? ????????????? ???????????
????????? (???). ?????? ??????????? ?????????? ????? ????? ? ???????? ???????????? ????????? ??????????, ????? ????????????????? ??????? ??????????? ????? ?????
????? ????????? ????????, ?? ???????????-
? ???????? ??????? ????? ????????? ??????
?? ??????? ????????, ???????????? ????????????????? (??????, ???????, 1987). ???????? ??????? ????? ????? ? ?????????????
???????? ????????? ??????????. ??? ???????
? ???????? ?? ? ???????? ????????? ???????,
???????????? ? ?????????? ??????????????
????????????-????????????????? ?????????
(?????????, 2009).
??????? ???????? ????????? ???????????????? ????????? ? ??????????????????????? ???????????, ?? ?????? (??????
? ??., 2002), ?????????? 50-60 %, ??? ??????????? ? ???????? ???????? ????????? ?
??????? ??????????? ? ???????? ?????????
?????. ? ???????????-?????????????? ?????????? ????? ??????? ??????? ??????????????
??????????? (????????, 2007). ?? ??????
?????? ??????? ??????? ?? ?????? ??????
(Gleeson, 2004).
??? ?? ?????, ???????? ?? ????????????
?????????? ???????????? ? ??????? ?????????? ???????????, ???????? ?????????????? ?????????????? ???????????? ?????????
? ???????? ???????, ??????????? ??? ??????? ?????????? ????????, ? ????? ??????????????? ????????? ????? ?????????. ?
?????????, ??????? ???????????? ???????????? ?????????? ???????? ??????????????
?????????? ????????????? ????????????
????? ? ???????????, ?????????????? ??????? ??????? ?? ?????????? ????????? ???????? ? ????? ?????????? ?????? (Bazarin,
Savchenko, 2011). ?????? ?????????? ???- ?
????-????????? ???????????? ??????? ?
???????? ??????????? ???????????????? ??-
# 152 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ????????? ?????????? ???- ? ????-????????? ???????????? ? ??????????????
????????? ?? ??????? ?????????????????
??????? ??????? ?????? ????????? ? ??????????? ?????????????? ?????????? ??????????? ?????? ???????? ??????? (???????, ????????, 1998; ????????, ???????, 2011; Rutter
et al., 2010; Youngster et al., 2010; Sirover, 2011).
????????? ? ??????
? ???????????? ??????? ??????? ?????????? ???????? ????, ????????????? ????????? ????? ?????? ? ????????????????
??????????? ????????????. ??????????
??????????? ????????? 51 ???????. ???????
22,1±3,7 ???. ?????????? ????????????: 1-?
?????? ? 5 ???????, ???????? ? ??????? ?????? ? 20 ???????, ?????? ?????? ? 18 ???????,
?????? ?????? ?????????????? ?????? ? 8
???????. ? ???????? ??????????? ?????? ??????????? 64 ??????? ???????? ???? ???????? ????, ??????? 20,6±1,8 ???, ?? ???????????? ?????????? ??????? ??????????
????????.
??????? ???????? ????? ? ??????? ??
??????????? ???????? ?????? ? ???????
???? ? ? ????? ????????????????? (????????
??? ?????), ????????????????? (?????????
???????????? ?????????) ? ??????????? (?????) ????????; ? ??? ?? ??????????? ?????? ? ??????????. ??????? ????? ?? ????????
???? ?????, ???????, ? ????????? ?????, ???
??????? ????? 12 ? ????? ????????? ?????????? ????????. ???????? ????? (10 ??) ??
???????? ???? ???????? ? ????????????
????????, ?????? ???????????? ? 80 ??
????????. ????????? ????????????? ???????????? ???????????? ? ??????? ?????????
????????? ??????-?????????? (p=1,077 ?/??3
??? ????????? ?????????????? ?????? ?
?=1,119 ?/??3 ??? ????????? ?????????????
????????????). ???? ????????????? ???????????? ????????? ?????????? ? ???????????? ???????? ? ?????? ???????? ? ?????-
?????????? ???????? ??? 400 g ? ??????? 10
???. ?? ????????? ???????? ????????????????? ??????????? ???????, ? ??????????
?????? ???????????? ? 2 ?? ???????? ??????.
??????? ??????????? ? ?????? ???????. ?????? ??????? ?????????? ????????????? ???????????? (??? ????? ?????? ?????????? ??
????????????-?????) ?????????? ??????????
?????????? ? ???????????. ????? ????????
?????????? ?????? ? ????????? ???????? ????????? ?????? ?? 1 ??? ??/??.
??????????? ?????????? ???- ? ????????????? ???????????? ? ?????????????
???????????? ????????? ????????????????? ??????? (????????, 1989). ?????? ??????? ???????????? ?????????? ?????????
?????????: ???????-6-???????????????????
(?6???), ????????-3-???????????????????
(?3???), ?????-???????? (???????),
???- ? ????-????????? ??????? ?????????????????? (??? ? ????-???), ???- ?
????-????????? ??????? ????????????????? (??? ? ????-???), ????- ? ?????????????? ??????? ????????????????????? (??????? ? ?????-???), ???- ?
????-????????? ??????? ????????????????????? (?????? ? ????-???), ???- ?
????-?????????
?????????????????????
(??????? ? ????????) ? ?????????????????? (??). ??? ????? ???????????? ?????????????? ????????????????? ????????
???(?)?:??????????????????-??????????
(??????????? ?? ????????? ???????? ???????????? ????????????? ? ????-??????????
?????????? ?? Photobacterium leiognathi ? ??????????????? ?? Vibrio fischeri ? ?????????
????????? ?? ???, ?. ??????????). ????????? ????????? ?? ?????????????????????
??????????? «CL-3606» (???? «?????», ??????????). ?????????? ???????????? ????????
? ?????????????? ???????? (1 ?=1 ??????/
???) ?? 104 ?????? (???????, 1998).
# 153 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ????????? ?????????? ???- ? ????-????????? ???????????? ? ??????????????
????????????? ???????? ??????? ??????? ??????????? ?????????? ????????? ? ???????????????? ???????, ??????????? ?
??????????? ??????, ????????? ?? ?????????????????? U-???????? ?????-?????. ?????????????? ?????? ???????????? ? ??????
???????? Statistica 7.0 (StatSoft Inc.).
?????????? ? ??????????
??? ???????????? ??????? ??????????
????-????????? ???????????? ? ????????????? ???????????? ? ??????????? ? ???????? ?????????????? ????? ??????????, ???
?? ???? ????? ??????????? ???????? ?????????? ?6??? (???. 1?). ?? ?????? ?????????
?????????? ????????? ? ??????? ???? ?????
??????? ? ????????????? ?????????? ? ?????? ??????. ?????????? ?? ? ??????????? ?
??????????? ????? ????????????? ??????????
???? ???????? ???????????? ???????????
?????? ? ????? ????????????????? ??????? (???. 1?). ?? ???? ???????????? ????????? (????????? ????????????????? ???????)
?????????? ??????? ???????? ?????????, ?
?????? ?????? ??????????? ?? ??????????,
?????????, ??????, ????????????? ??????????
????, ??? ? ??? ??????????? ??????. ??????? ?????????? ???????? ? ??????????? ? ??????????? ?? ???? ????? ? 5,2 ????
???????? ?????????? ??????????? ??????
(???. 1?). ? ?????? ????????? ?????????? ???????? ??????????? ????????? ? ??????????
?????? ? ?????? ??????. ?????????? ????????????? ??????? ????????????????????? ?
??????????? ? ??????????? ?? ???? ???????
?????????????? ????? ???? ?????????????
?????????? ???????? ???????????? ??????????? ?????? (???. 1?). ? ?? ?? ????? ?????????? ?????-????????? ??????? ??? ?
??????????? ?? ???? ????? ???????????????
???????????? ??????, ????? ??? ????????????? ?????????? ????????? ?? ???? ?????????
(???. 1?). ?? ????? ?????? (?????????? ??????) ?????????? ????? ???????? ??????????
????, ??? ? ??????????? ??????. ??????????
??????? ? ????????????? ????????????
??????????? ?? ???? ??????? ??????????????
????? ??????????????? ??????????? ??????.
???????????? ??????? ?????????? ???????????? ???????????? ? ?????????????
???????????? ??????????? ? ???????? ?????????????? ????? ????????? ?????????? ??????????? ??????????????. ???, ?????????? ???
?? ???? ????? ???? ????????????? ?????????? ???????? ???????????? ????????????
?????????, ????? ????????? ? ????? ????????????????? ??????? ? ?? ????????????????? ?
?????????? ??????? (???. 2?). ?????????? ?????????? ?????? ?????????? ???, ??????
? ??????? ? ????????????? ????????????
? ???????????: ????????????? ?????????? ?????????? ? ????? ????????????????? ???????
(???????? ??? ?????), ?????????? ?????????
????? ????????? ?? ???? ????????? (????????? ????????????????? ???????) ? ????????????? ?????? ?????????????? ? ?????? ??????
(???. 2?-?). ? ?? ?? ????? ?????????? ?3??? ?
????????????? ???????????? ? ???????????
? ????? ????????????????? ??????? ?? ?????????? ?? ??????? ? ??????????? ??????, ??
????????????? ?????????? ????????? ? ?????
????????????????? ??????? ? ?????????? ?
?????????? ?????? (???. 2?).
????????????
???????
??????????
????-????????? ??????? ???, ??? ? ???
? ????????????? ???????????? ? ??????????? ? ???????? ???????? ??????????????
????? ????????? ?????????? ????????? ??????????????: ? ????? ?????????????????
??????? (???????? ??? ?????) ??????????
???? ??????????? ???????? ????????????
??????????? ?????? (???. 3?-?); ? ????? ????????????????? ??????? ??????????? ??????
???????? ?????????? ? ????????? ?? ??????-
# 154 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ????????? ?????????? ???- ? ????-????????? ???????????? ? ??????????????
???. 1. ??????? ?????????? ?????????? ????-????????? ???????????? ? ????????????? ????????????
? ??????????? ?? ?????? ??????? ?????????????? ?????; ? ? ????????; ? ? ????????? ?????????????????
??????? (???????? ??? ?????); ? ? ????????? ????????????????? ??????? (?????????); ? ? ?????????
??????????? ??????? (?????); ? ? ????????????? ???????? ??????????? ??????? ??????? ? ???????????
??????
???????? ? ?????????? ??????, ?????? ?? ??????????? ??????, ??????????? ?? ????????
???? ?????.
??????????? ???????? ???????? ???????? ??????? ?? ?????? ??????????????
????? ??????, ??????? ? ????? ????????????
?? ??????????????? ? ?????????????? ???????????. ???, ????? ?? ?????????? ????-
??????????? ???????????? ?????????????
???????????? ?????? ????????????? (???
???????????) ?????, ?????????? ???????? ?????????????? ????? ????????? ????????????????? ????? (Braga et al., 2009;
Bazarin, Savchenko, 2011; Seth et al., 2011).
?6??? ? ???????? ? ????????????????
??????? ????????????????? ????? (????-
# 155 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ????????? ?????????? ???- ? ????-????????? ???????????? ? ??????????????
???. 2. ??????? ?????????? ?????????? ???-????????? ???????????? ? ????????????? ???????????? ?
??????????? ?? ?????? ??????? ?????????????? ?????. ??????????? ???????? ? ????????????? ????????
??? ?? ???. 1
???, ????????, 1998; ????????? ? ??., 2006).
????????? ?????????? ??????? ???????? ?
????????????? ???????????? ? ???????????
?? ???? ????? ?????????? ??????? ???????
?????????????? ?????????? ?????? ?? ????
??????? ????? ? ???????????????? ?????,
??????? ???????????? ????-????????? ???
??????? ??????????-????????. ? ?? ?? ?????
?? ???? ????????? ??????????? ??????????
???????? ?????????? ??????? ????????, ???,
? ???? ???????, ?????????? ????????? ?????? ??????? ???????? ???? ????????? ?, ??????????????, ?????????????? ??????????
????????????? ????????????. ? ?????? ?????? ?????????? ???????? ??????????, ?? ??
????????????????? ?????????.
??? ??????????? ???????????????? ??????? ?? ???? ?????????????? ?????? ? ???-
# 156 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ????????? ?????????? ???- ? ????-????????? ???????????? ? ??????????????
???. 3. ??????? ?????????? ?????????? ????-????????? ??????? ? ????????????? ???????????? ?
??????????? ?? ?????? ??????? ?????????????? ????? (??????????? ???????? ? ????????????? ????????
??? ?? ???. 1)
?????????? ???????????? ?????? ?????????????? ?????????-????????? ???????????????
??????? (Yuan et al., 2009; Harwood et al., 2011).
?????????????, ? ??????????? ? ?????????
????????? ???? ????? ??????????? ????????? ?????????? ??, ??????? ??????????
??????? ???????? ? ????? ?????????????????
??????? ???? ?????? ??????????? ?????? ?
?? ???????????????? ? ?????? ??????. ?????????????, ?????????? ?????????-?????????
??????????????? ??????? ? ??? ??????? ?
??????????? ???????, ??? ????? ???????????? ???????? ?? ????????? ????????????????
??????? ??????? ???? ??????. ??????????
????????, ??? ????????????? ???????????
?????????????? ?????? ????? ????????
?????????? ?6??? ? ?? ? ?????????????
???????????? ? ??? ??????????? ??????
? ??????????? ? ???????? ??????????????
????? ?? ??????????. ??? ??????? ? ???, ???
?????, ???????????? ?? ????????????????????????????? ?????? ????????????????? ?????, ??????????? ?? ?????? ??? ?????????????? ??????????? ??????????, ?? ? ?
?????-?????????? ???????.
?????????? ????????? ??????? ????????
???????? ??????????????????? ????????? ? ????????????? ???????????? (Schuster
et al., 2007; van Raam et al., 2008). ??? ????
?6??? ? ???????? ????????? ????????? ??
????????. ??? ?? ????? ???????? ?????????
?????????? ?????????? ??????? ??? ? ????????????? ???????????? ??????????? ??
????????? ?????? ?????????????? ????? ????????? ????????? ?????????? ?6???. ?????
?? ?????????? ???????????? ??????????????
??????? ????????? ?????? ?????? ?????????
????????? ??????????? ????? ?3??? (???????, ????????, 1998). ??????????, ??? ?
??????????? ? ????????????? ????????????
# 157 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ????????? ?????????? ???- ? ????-????????? ???????????? ? ??????????????
?????????? ??????? ???????? ???? ???????
?? ???? ????? ? ? ?????? ?????????, ?? ?????????? ? ?????? ??????. ??-????????,
??????????? ????? ?? ????????? ? ?????????
????????? ???? ????? ? ?? ???? ?????????
??????????????? ???????????? ???????
?????????? ??????? ????? ?????????????????? ???????? ??????, ????? ??? ?? ?????
?????? ??????? ??????????? ??????????
??????? ????????? ?, ??????????????, ????????????? ?????????? ?????????? ?3???.
? ????????? ???? ?????????? ??????????? ???????????? ???? ??????????? ?
?????????? ? ? ?????????? ??????????????
??????? ????????????? ???????????? (???????? ? ??., 2003; ????? ? ??., 2009; Aulik
et al., 2011; Geering, Simon, 2011). ???????????
??????????? ?????????? ?? ??????????? ????????? ???????? ??????????? ? ?????????
???? ???????????????? ????????? ??? ?????????????? ????????????. ??? ??????? ?????????? ?????????????? ?????????????????
???????????? ????????????? ????????????
? ??????????? ? ???????? ??????????????
????? ???? ??????????, ??? ?????? ?????????? ??? ? ??????? (???????? ?????
????????????? ??????) ???????? ? ?????
????????????????? ???????, ??? ???????? ?
???????????? ???????????? ?????? ?? ?????
?????? ?, ??????????????, ???????? ??????
????????? ???????. ? ?? ?? ????? ? ?????
????????????????? ? ??????????? ????????
?????? ?????????? ?????? ????????? ??????????? ???????, ??? ?????????? ?????????
?????? ????????? ???????. ???????? ??????????????? ??? ??????????????? ???????
????? ????????????? ??????, ??????????
??????? ?????????? ???????? ??? ????????
??????? ???????????? ???? ? ???????????????? ????????????. ????? ????, ????????
?????????? ???????? ? ????????????????
???????????? ????? ????????? ? ??????-
???? ????? ???????? ???????? ?????? (Kim,
Park, 2010; Jung, Park, 2011; Kil et al., 2011). ???
???? ???????? ?????????? ??????? ????????
????? ????????????? ?????????????? ?????
??????. ?????????????, ?????? ??????????
???????? ? ????????????? ????????????
? ??????????? ????????????? ?? ?????? ?????? ??????????????? ??????? ????????????
?????? ?? ????? ????????????? ??????, ?? ?
?????????? ???? ???????? ???????? ???????
???? ??????.
???? ????????????? ?????? ????????
??????????????, ????? ?????? ? ?????????
??????????????? ?????? ????? ???(?)- ?
????(?)-????????? ?????????????????????
(???????, ????????, 1998). ? ?????????????
???????????? ??????????? ???????? ??????
????-?????????? ?????? ?????????? ? ?????
?????? ?? ??????? ??????????????? ??????
? ???? ?????????????? ????? ?????????????
???????? ?????????? ??????????? ????????? ????????? ?????: ??????? ??????? ? ????????? ????????? ???? ?????, ???????? ??
???? ????????? ? ????????????? ?????????????? ?? ????? ??????. ?????-????????? ????? ??????????? ????? ?????? ??? ????????: ?
????? ????????????????? ??????? ?? ????????????? ?????? ??????????? ??????, ??????????? ???????????? ???????? ?? ???? ????????? (? ????? ????????????????? ???????) ?
????????? ????????? ? ?????????? ??????.
?????? ?????????? ?? ???? ?????????????
?????? ????? ?????? ????? ?????????????
??????????? ???????? ????????? ????????????????? ???????????? ? ???? ???????? ??????????: ??????? ??????? ?? ???????? ????
?????, ???????? ?? ????? ????????????????? ???????, ????????????? ??????????????
? ?????? ??????. ?????????? ??????? ?
????????????? ???????????? ? ???????????
???? ??????? ?? ???? ?????????? ????????
??????????. ?????????????, ????? ???????-
# 158 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ????????? ?????????? ???- ? ????-????????? ???????????? ? ??????????????
??? ? ????? ????????????? ?????? ?? ???????
??????????????? ?????? ?????????????? ????? ??????????, ??? ??????. ??? ???? ???????????? ??????????? ????? ?????????? ?
??????????? ???????? ?????????? ????????? ? ???? ???????????????-??????????????
??????????. ?????? ? ?????? ?????? ? ????????????? ???????????? ??????????? ??????????? ????????? ?????-??????????
?????? ?????????? ? ????????? ????? ??
??????? ??????????????? ??????. ? ?????
????? ?????????, ??? ??????? ????????????
?????????? ????? ?????? ?????? ? ????????? ??????????????? ?????? ? ???????????
??????????? ????????????? ????? ??????????
? ?????????????? ????????? ?? ???????? ?????.
??????????
? ????? ????????????????? ??????? ?
??????????? ? ????????????? ???????????? ???????????? ???????? ??????????????
????????: ?????????? ? ???????? ??????????,
?????????-????????? ??????????????? ???????, ? ????? ???????????? ???????, ???????????? ??????? ??????? ??????????????????
??????? ? ???????????? ??????.
?? ???? ????????? (? ????? ????????????????? ???????) ? ??????????? ???????????
?????????? ???????? ?????????? ??????????? ???- ? ????-????????? ????????????,
??? ?????????? ???????? ?????????????????
?????????, ??????? ????????????? ??????,
?????????? ??????????? ????????.
? ?????? ?????? ??????? ??????????????
?????????????? ????????? ? ?????????????
???????????? ? ??????????? ?? ??????????.
? ???? ?????? ??????????? ????????? ????
?????????? ??????????, ? ??? ????? ?? ????
????????? ??????? ?????????? ? ???????
????????? ???????????, ? ????? ??????????????? ?????? ?????????? ? ????? ????????????? ?????? ?? ??????? ??????????????? ??????.
???????????? ????????? ??? ?????????? ????????? ????, ?????? ?12-0431145\12 ???_?.
?????? ??????????
1.
2.
3.
4.
5.
6.
7.
?????? ?.?., ??????? ?.?. (1987) ???????????????? ???????????? ??? ????????? ??????? ?????????? ????????. ?.: ??????????? ? ?????, 210 ?.
??????? ?.?., ???????? ?.?. (1998) ????????????? ?????. ?.: ????????, 704 ?.
????????? ?.?. (2009) ??????????? ????????????? ? ???????????. ?.: ????????? ?????,
192 ?.
??????? ?.?. (2011) ????? ??? ?????????: ??????-?????????? ???????. ???????? ?????
? ???????????? ????????. 3: 1-6.
????? ?.?., ??????? ?.?., ????? ?.?., ??????? ?.?., ????????? ?.?. (2009) ?????????????
?????????????? ??????????? ?????????????????? ??????????? ? ???????? ???????????
??? ????????? ????????????????? ????????????. ???????????. 5: 267-269.
????????? ?.?., ?????? ?.?., ?????????? A.B. (2006) ???????????? ???????? ???????-6??????????????????? ?? ?????? ???? ? ???????? ?????????????? ????????????????????
????????? ??? ??????????? ????????. ?????????????? ?????. 52(3): 278-286.
???????? ?.?., ???? ?., ??????? ?. (2003) ???????-??????????? ???? ????????? ??????
??????????? ????????, ?????????????? TNF?. ???????? ? ??????????. 2(1): 29-35.
# 159 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ????????? ?????????? ???- ? ????-????????? ???????????? ? ??????????????
8.
?????? ?.?., ?????? ?.?., ??????? ?.?. (2002) ??????? ???????? ??????? ?? ??????????
????????. Russian Journal of Immunology. 7(1): 1-24.
9. ???????? ?.?. (2007) ??????????? ? ???????????? ???????????. ????????? ??????. ?.:
??? «???», 184 ?.
10. ???????? ?.?., ????????? ?.?., ??????? ?.?., ???????? ?.?. (2011) ???????? ??? ??????
???????????????????? ???????. ??????????, 213 ?.
11. ???????? ?.?., ??????? ?.?. (1989) ???????????????????? ??????????? ?????????? ????-
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
???????? ? ?????????? ?????????????? ????? ????????????????? ???????. ???????????? ????. 11: 23-5.
????????????? ?.?., ??????? ?.?. (2003) ????? ??????? ? ????????? ?????????? ??????????? ???????????????. ?????? ? ???????? ?????????? ????????. 1: 18-23.
Amulic B., Hayes G. (2011) Neutrophil extracellular traps. Curr Biol. 21(9): 297-298.
Aulik N.A., Hellenbrand K.M., Kisiela D., Czuprynski C.J. (2011) Mannheimia haemolytica leukotoxin
binds cyclophilin D on bovine neutrophil mitochondria. Microb Pathog. 50(3-4): 168-178.
Bazarin K.P., Savchenko A.A. (2011) Special features of the state of functional activity of
neutrophil granulocytes in blood during the high level of physical exercises. Journal of Siberian
Federal University. Humanities & Social Sciences. 4 (9): 1251-1259.
Braga P.C., Dal Sasso M., Culici M., Spallino A., Falchi M., Bertelli A., Morelli R.,
Lo Scalzo R. (2009) Antioxidant activity of Calendula officinalis extract: inhibitory effects
on chemiluminescence of human neutrophil bursts and electron paramagnetic resonance
spectroscopy. Pharmacology. 83(6): 348-355.
Cуrdova A., Sureda A., Tur J.A., Pons A. (2010) Immune response to exercise in elite sportsmen
during the competitive season. J. Physiol. Biochem. 66(1): 1-6.
Friman G., Wesslen L. (2000) Special feature for the Olympics: effects of exercise on the immune
system: infections and exercise in high-performance athletes. Immunol. Cell. Biol. 78(5): 510-522.
Geering B., Simon H.U. (2011) Peculiarities of cell death mechanisms in neutrophils. Cell Death
Differ. 18(9): 1457-1469.
Gleeson M. (2004) Immune function and exercise. European Journal of Sport Science. 4(3):
52-66.
Hдger M., Cowland J.B., Borregaard N. (2010) Neutrophil granules in health and disease. J. Intern.
Med. 268(1): 25-34.
Harwood D.T., Darlow B.A., Cheah F.C., McNeill N., Graham P., Winterbourn C.C. (2011)
Biomarkers of neutrophil-mediated glutathione and protein oxidation in tracheal aspirates from
preterm infants: association with bacterial infection. Pediatr. Res. 69(1): 28-33.
23. Hawley J.A., Burke L.M. (2010) Carbohydrate availability and training adaptation: effects on cell
metabolism. Exerc. Sport Sci. Rev. 38(4): 152-160.
24. Jung K.H., Park J.W. (2011) Suppression of mitochondrial NADP(+)-dependent isocitrate
dehydrogenase activity enhances curcumin-induced apoptosis in HCT116 cells. Free Radic.
Res. 45(4): 431-438.
25. Kil I.S., Jung K.H., Nam W.S., Park J.W. (2011) Attenuated mitochondrial NADP+-dependent
isocitrate dehydrogenase activity enhances EGCG-induced apoptosis. Biochimie. 93(10):
1808-1815.
# 160 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ????????? ?????????? ???- ? ????-????????? ???????????? ? ??????????????
26. Kim S.Y., Park J.W. (2010) Modulation of hypoxia-inducible factor-1? expression by mitochondrial
NADP+-dependent isocitrate dehydrogenase. Biochimie. 92(12): 1908-1913.
27. Koch A.J., Wherry A.D., Petersen M.C. (2007) Salivary immunoglobulin A response to a collegiate
rugby game. J. Strength. Cond. Res. 21(1): 86-90.
28. Mason E.F., Rathmell J.C. (2011) Cell metabolism: an essential link between cell growth and
apoptosis. Biochim. Biophys. Acta. 1813(4): 645-654.
29. Mulukutla B.C., Khan S., Lange A., Hu W.S. (2010) Glucose metabolism in mammalian cell
culture: new insights for tweaking vintage pathways. Trends Biotechnol. 28(9): 476-484.
30. Nuutila J. (2010) The novel applications of the quantitative analysis of neutrophil cell surface
FcgammaRI (CD64) to the diagnosis of infectious and inflammatory diseases. Curr. Opin. Infect.
Dis. 23(3): 268-274.
31. Rutter J., Winge D.R., Schiffman J.D. (2010) Succinate dehydrogenase ? assembly, regulation and
role in human disease. Mitochondrion. 10(4): 393-401.
32. Schuster D.P., Brody S.L., Zhou Z., Bernstein M., Arch R., Link D., Mueckler M. (2007) Regulation
of lipopolysaccharide-induced increases in neutrophil glucose uptake. Am. J. Physiol. Lung Cell
Mol. Physiol. 292(4): 845-851.
33. Seth R., Ribeiro M., Romaschin A., Scott J.A., Manno M., Scott J.A., Liss G.M., Tarlo S.M. (2011)
Occupational endotoxin exposure and a novel luminol-enhanced chemiluminescence assay of
nasal lavage neutrophil activation. J. Allergy Clin. Immunol. 127(1): 272-275.
34. Shephard R.J. (2000) Special feature for the Olympics: effects of exercise on the immune system:
overview of the epidemiology of exercise immunology. Immunol. Cell. Biol. 78(5): 485-495.
35. Sirover M.A. (2011) On the functional diversity of glyceraldehyde-3-phosphate dehydrogenase:
biochemical mechanisms and regulatory control. Biochim. Biophys. Acta. 1810(8): 741-751.
36. Summers C., Rankin S.M., Condliffe A.M., Singh N., Peters A.M., Chilvers E.R. (2010) Neutrophil
kinetics in health and disease. Trends Immunol. 31(8): 318-324.
37. van Raam B.J., Sluiter W., de Wit E., Roos D., Verhoeven A.J., Kuijpers T.W. (2008) Mitochondrial
membrane potential in human neutrophils is maintained by complex III activity in the absence of
supercomplex organization. PLoS One. 23 (3-4): e2013.
38. Witko-Sarsat V., Pederzoli-Ribeil M., Hirsch E., Sozzani S., Cassatella M.A. (2011) Regulating
neutrophil apoptosis: new players enter the game. Trends Immunol. 32(3): 117-124.
39. Yeo W.K., Carey A.L., Burke L., Spriet L.L., Hawley J.A. (2011) Fat adaptation in well-trained
athletes: effects on cell metabolism. Appl. Physiol. Nutr. Metab. 36(1): 12-22.
40. Youngster I., Arcavi L., Schechmaster R., Akayzen Y., Popliski H., Shimonov J., Beig S., Berkovitch
M. (2010) Medications and glucose-6-phosphate dehydrogenase deficiency: an evidence-based
review. Drug Saf. 33(9): 713-726.
41. Yuan W., Wang Y., Heinecke J.W., Fu X. (2009) Hypochlorous acid converts the gamma-glutamyl
group of glutathione disulfide to 5-hydroxybutyrolactam, a potential marker for neutrophil
activation. J. Biol. Chem. 284(39): 26908-26917.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????. ????????? ?????????? ???- ? ????-????????? ???????????? ? ??????????????
Dynamics of Neutrophilic Granulocyte?s NADand NADP-Dependent
Dehydrogenases Activity in Athletes
Andrei A. Savchenkoa,b and Kirill P. Bazarinb
Research Institute of Medical Problems of the North
of Siberian Branch of Russian Academy of Medical Sciences,
3g Partizana Geleznyaka, Krasnoyarsk, 660022 Russia
b
Siberian Federal University,
79 Svobodny, Krasnoyarsk, 660041 Russia
a
The present research is focused on metabolic reactions of neutrophilic granulocytes in the blood of
athletes, representatives of various sports who showed endurance in dynamics (in the course of a year),
at different phases of an annual training macrocycle. It was found that the main metabolic processes
(anaerobic and aerobic energy production systems, glutathione-depended antioxidant system and
plastic reactions defining the level of macromolecular synthesis and cells reactivity) are activated at
the end of preparing period. The metabolic reactions of neutrophilic granulocytes are significantly
disturbed at the end competitive period. This reduces their functional and regulatory capacity. At the
end of recovery period disturbed reactions were not sufficiently recovered.
Keywords: sport, immune system, neutrophils, spontaneous chemiluminescence, induced
chemiluminescence.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2013 6) 163-174
~~~
??? 620.22:577.11
??????????????? ?????????????? ??????????
?? ?????? ???????????
? ??????????? ????????????
?.?. ????????,?*, ?.?. ???????,?,
?.?. ???????,?, ?.?. ??????,?
?
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ?????????, 79
?
???????? ????????? ?? ???,
?????? 660036, ??????????, ?????????????, 50, ???.50
?
???? «?????», ??? ?? ???,
?????? 660049, ??????????, ??. ????, 53
Received 17.03.2012, received in revised form 13.02.2013, accepted 26.02.2013
??????????? ??????? ??????????????? ? ?????????? ?????????????? ??????????
?? ?????? ??????????? ? ???????????????? ??????????????????????. ?? ???????
????????????? ????????? ?? 3-???????????????????? (3-???) ? ????????????????
??????????? (???) ????????? ??????? ?????????????????? ??????????? ?????????
???????????????? ????????? ???????? ?????????????? ????????????? ?? ???????????????
????????? Ad N2. ????????, ??? ??? 20-???????? ????????? ? ?????? ???????? ?? 1 ??
????????? ?? ??????????? ????? ???? ?????????????? ?? 70 ?? 150 ??? ?????????? ?
??????????? ?? ???????? ??????????? ???? ??????????? ? ???????.
???????? ?????: ?????????????????????, 3-???????????????????, ????????????????
??????????, ??????????, ?????????????? ?????????.
????????
??????????? ????????????? ?????????
???????? ????????????? ???????????, ???????????? ??? ???????????? ??????? ???????? ??????? ???????????? ?????. ? ?????????, ?????????????? ??????????????
??????????? ???????? ????????????????????? (???) ? ????????? ?????? ??????
*
?????????????? ????????????? (?????? ?
??., 2006). ??? ???????? ????? ?????? ?????????????????? ???????, ? ??????: ????????????????? ??? ??????????? ?????????
?????????, ??????? ?????????????????, ???????????? ????????? ???????????? ???????
??????? ? ???????. ??? ????????? ???????????? ?? ? ???????????? ??????? ?????????
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: bondvs@mail.ru
# 163 #
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?.?. ???????, ?.?. ??????? ??????????????? ?????????????? ?????????? ?? ?????? ????????????
?????, ?????????? ? ?????????? ??? ?????????????? ?????????? ? ????????, ????????????, ??????? ??????????????, ????????
????????? ? ?.?. (?????? ? ??., 2008; ?????? ?
??., 2011; ???????? ? ??., 2011; Shishatskaya,
2008). ???????? ?????? ????????, ???????? ? ????? ????? ???????????, ????????
?????????? ??????? ????????? ??? ??????? ??????? ? ??????-?????????? ??????? ??
???? ??????????????????? ??????? ? ??????
???????? ??????????? ???????????? ??? ???????? ?????? ?????? ??????? ? ???????????
????????????? ?????????? ? ??????, ???????,
????????, ?????, ?????????? ? ?.?.
????????????? ? ??????? ???????????????? ??? ????????? ????????????? ?? ???
????????????? ????????? ???????????????
??????????. ????????, ? ????????? ??????
????? ??????? ??? ?????????? ???????????
????????????? ??? ? ???????????? ???- ?
?????????? ??????????? ??? ?????????????
??????? (??????? ????????????? ?????????)
??? ?? ????????? ? ???????????????? ????????; ?????? ?????, ??????? ? ????????? ???
?????????????, ???????????? ? ????????????
???????? (????????? ? ??., 2011; Shishatskaya
et al., 2006; Shishatskaya, 2008; Shishatskaya et
al., 2008; Shishatskaya et al., 2011). ????????,
??? ?????????? ????? ?????????? ? ????????? ?????? ???????????? ???????? ?????
?????????????? ?? ?????? ?? ????? ???????????? ??????? ? ????????? ????????????? ???????? ??????????? ? ?????.
???????? ?? ????? ??????????? ???, ??????? ????????, ??? ?????????? ?????????? ?
??????????? ???????? ???? ???????????? ?
???????????? ??????? ?????? ?????? ?? ????????????? ?????????????? ??????????,
????????, ???????????? ??????????? ?????????? ? ???????? ????? ?????? ???????? ???????? ???????? ? ????? ??????? ?????????
? ???????????. ????? ?? ????????? ???????
???? ???????? ???????? ??????????????? ??
?????? ??? ?????????????? ??????????.
??????????????, ??? ????????? ???????????, ?????????? ???????????? ??????????
(??? ?????????), ???????? ???????????? ?? ?
???????? ????????? ??? ?????????? ??????????????? ??????? ? ????????????? ??????????, ????? ???????? ??????-??????????
? ??????????? ?????????????? ??????????
?????????? ?, ??? ?????????, ????????? ??????????? ??????????? ??????? ?? ??? (????????, ??? ?????????????, ???????????? ?
???????????? ????????).
? ???????? ???????????? ?????????? ??? ???????? ?????????? ?? ?????? ???
?????????????? ????????????? ????????????? ???????????????? ??????????? (???)
????????? ??????? (Bondar, Puzyr, 2004) ?? ?????????? ???? ??????. ??????-??????????
???????? ??? (????????? ??????, ????????? ???????????, ???????? ???????????
? ??????? ?????????? ???????????? ? ????????????? ??????) ?????????? ?? ???????
????????????? ???????? ? ????????? ??????????? ????????????? ? ???????????????
???????, ??? ????????? ????????? ??? ??????????? ??? ?????????? ????? ??????????
? ??????? ??????????????? ??????????. ????????, ???????? ??????????? ????????????? ??? ? ???????? ???????????????????
?????????? ??? ???????????? ????????? ?
??????? ?????? (Bondar et al., 2004; Puzyr et
al., 2007a; Purtov et al., 2008; ?????? ? ??.,
2010), ???????? ??? ?????????? ? ????????????? ???????? (Puzyr et al., 2007b; Mogilnaya et
al., 2010a; Puzyr et al., 2010), ?????? (????????)
??? ??????????????? ????? ?????? ?????????, ????????????? ??????????? ? ????????
???????? ????????????? ?????????? (Puzyr et
al., 2004; Bondar et al., 2008; Purtov et al., 2010;
Purtov et al., 2011; Mamaeva et al., 2011). ?????? ???????? ??? ?????? ??? ???????? ?
# 164 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ??????? ??????????????? ?????????????? ?????????? ?? ?????? ????????????
??, ??? ?????????? ????????????? ????????????? in vivo ???????? ??????? ???????????????? ???? ?????????? ? ???????????
????????????????? ???????? (?????? ? ??.,
2004?; ?????? ? ??., 2004?; ?????? ? ??., 2005;
Puzyr et al., 2007a; Mogilnaya et al., 2010b).
????? ???????, ????? ?????? ????????
???????? ??????????? ????????? ?????????????? ?????????? ?? ?????? ??? ? ???????????????? ??????????? ?????????
???????. ? ???????? ?????????? ??????? ???
???????????? ???? ??????? ?????????????
??????????, ??? ???? ?? ???????? ???????
????????? ??????? ??? ????????????? ? ???????? ??????? (????????, ??????, 2006).
????????? ? ??????
? ???????????? ???????????? ?????????? ?? 3-???????????????????? (3-???)
? ????????? ? ????????? 1?20 ??. ???????????? ??????????? ????????? ????????? ?
?? ??????-?????????? ???????? ???????? ???????? ? ??????? (????????, ??????, 2006;
?????? ? ??., 2012). ?????????? ????????
????????? ???????. ?? ????????? ???????
?????????? ???????? ?????????????????? (Centrifuge 5415R, Eppendorf, Germany)
??? 9300 g ? ??????? 5 ??? ??? ???????????
15 °?. ?????????? ?????? ?????? ????????? ?????????????? ????? (Milli-Q system,
Millipore, USA) ??? ???????? ?????????
???????? ????????????, ??????? ??? ?????????????? ? ??????? ????????? ????? ?? ?????????. ?????????? ?????? ??? ???????????????? ? ?????? ?????? ??????????????
???? ? ???????? ?????????????????? ???
????????? ???? ????????. ??????? ?????????? ???????????????? ? ??????????????
???? ? ???????????? ??? ????????????.
??? ????????? ?????????????? ?????????? ????????? ???, ?????????? ???????
?????????? ????????????? ? ?????????????
?????? ? ??????? ??????? ????????? ? ????????? 30 ? 125 ?? (??????????? ???????? ????????? ?? ??????????? Coulter #5, BeckmanCoulter, USA). ?????????? ????????? ??? ?
?? ??????-?????????? ?????????????? ???????? ???????? ???? ????? (Bondar, Puzyr,
2004; ??????, ???????, 2005; Puzyr et al.,
2005; Puzyr et al., 2007?). ??? ?????????????
???????????? ????????? ? ?????????????
??? 10,0 ?/?, ??????? ???????? ???????????
?????????????? ???? ? ??????? ??????? ??????????.
???????? ??????????? ????????? ?????????? 3-??? ?????????? ? ??? ? ?????????????? ???? ????????? ????????? ???????. ??????? ???????? ?????? ????????? ???
?????? ??????? ???????????? ???????????
(?????????? ? ???) ? ???????????? ??
??? ????????? ??????????? ? ??????? 1 ? ???
?????????? ????????????? ?? ?????????
200 rpm (?????? OS-10, BIOSAN, ??????).
????????? ???????? ?????? ????????? ?????????, ??????? ???????????? ??? ??? ??
???????? ??? ?????????? ???. ????? ?????
?????????? ???????? ??????????????????
??? 2300 g ? ??????? 5 ??? ??? ???????????
15 °?. ? ?????????? ????????????? ?????????? ?????????? ???, ?? ??????????? ? ????????????, ?? ???????? ?????????? ?????????
?? ????? ????? 400 ?? (UV/VIS ??????????????? UVIKON 943, Kontron Instruments, ??????). ??? ?????????????? ?????? ???????????? ????????????? ???????????, ??????????
???????????????? ?? ?????????? ? ?????????
????????????? ??? (???. 1). ?????? ??????
???????? ?????????????? ????? ??? ???????? ???????? ???, ?? ???????????????? ?
????????????. ??? ???? ?????????? ?????? ??? ???????????????? ? ?????? ??????
????, ???????? ?????????????????? ???
????????? ???? ???????? ? ???????? ???????????? ??? ??????????? ?????????? ???.
# 165 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ??????? ??????????????? ?????????????? ?????????? ?? ?????? ????????????
???. 1. ??????????? ?????????? ????????? ?????????? ??? ?? ????? ????? 400 ?? ?? ????????????
??????????
????????? ??????????????? ?????????
? ??????? ??????? ?????????????? ????????????? ????????? ?? ????????? ?????.
??????? ? ?????????????? ???? ????????
????????? ?????????? : ???, ??? ??????
??????? ??????????? ??????????? ? 9:1, 5:1
? 2:1. ? ???????? ???????? ???????????? ????????? ????????? ??? ?????????? ???. ?
?????????? ????????? ????????? ???????
??????? ?? ???????????? 5 ?? ? ???????????? ?? ? ??????? 20 ??? ??? ????????? ??????????? ? ?????????? ????????????? ??
????????? 200 rpm. ????? ????? ??????????
???????? ?????????????????? ??? ????????, ????????? ? ?????????? ???????. ?????? ????????? ??????????? ???????? (3 ????
?????????????? ?????, 1 ??? 1? NaCl, 3 ????
?????????????? ?????) ??? ???????? ???????? ???, ?? ??????????? ? ????????????.
?????????? ?????? ??? ????????????????
? ??????????? ?????? ? ???????? ?????????????????? ??? ????????? ???? ????????. ?????????? ???????????? ???????? ???
??????????? ?????????? ????????? ???, ??
?????????? ? ??????? ? ????????????.
????????????? ???????????? ??????????????? ???????????? ? ??????????? ??????? ?????????? ??????????? ????????? ?
??????? ?????????, ??????????? ????????? ???, ????????? ? ??????? ?????????
??????????? ??????????? (?????????? ??
1000 ? S-5500, Hitachi, ??????).
?????????? ? ??????????
?? ????????? ????? ???????????? ????
????????? ???????? ??????????? ?????????
??????????????? ????????? ???-??? ???
????????? ?????? ??? ? ???????????? ??
3-??? ? ??????????? ????? (??????????????
????). ? ?????????? ??????????? ????????????? ???? ???????????, ??? ? ???? ???????
??? ?? ??????????????? ? ????????????.
?? ?????? ????????????? ??????? ???????,
??? ????? ??????????? ????????????? ????????? ????????? ? ??? (??? ?????? ??????? ???????????? ???????????) ??? ????-
# 166 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ??????? ??????????????? ?????????????? ?????????? ?? ?????? ????????????
1
2
???. 2. ??????? ??? ??????? ?????????: 1 ? ???????????, 2 ? ??????? ????? ????????? ? ?????????
??? ? ??????????? ????? ? ??????????? ??????? ?????????????? ?????
??????? ?????????????? ? ?????????????,
?????????? ?? ??????? ?????????????????
????????? ? ??????????? ??????? ?????????. ? ?????? ????, ??? ? ???????? ??????????? ????? ?? ?????????? ?????????? ???
? ???????????? ?? 3-???, ???????????????
????? ?????????? ????????? ????? ????????? ????? ?? ????????? ? ??? ? ??????????? ??????? (???. 2).
?????????? ?????? ???????????? ?????????? ?????????? ????????????? ?????????? ?? ??? ? ????????????? ?????????
?? ?? ?????? ?????????????? ?????????? ?
????????? ??? ? ??????????? ?????? ?????.
?? ??????? ????, ??? ?????????????????? ??
??????? ????????? ?? 3-???.
??? ??? ???????????? ? ???????????
???????? ?? ???? ?????????????? ??????????
? ?? ????????? ???????? ??????????????
???????? ?? ??? ? ???, ???? ???????????
??????????? ????????? ?????????? ?????????? 3-??? ? ??? ? ??????? ???????
????????????? ?? ????????? AdN2. ?????
????????????????? ?????? ??????????? ??
????????? ??????. ????????????, ??? ???
???????? ??????????? ?????? ?????? ?
???????? ? ????? ????????? ????????????
(?=O) ?????? (????????, ?????? ? ??., 2006).
????????, ??? ?? ??????????? ??? ???????
???????????? ?????? ??????????????????
????? (?????????????, ?????????????, ????????????) (Gibson et al., 2009). ?????? ??
?????, ??????????? ??? ? ??????? ???
????? ???????? ?? ?????????????? ??????????? ??????? ?????????????? ?????????????. ???????? ?????? (????, 1988) ???????
?????????????? ????????????? ?? ????????
????????????????? ??????????????? ????????? AdN2 (????????????? ?? ????? ?=O)
???????? ? ??? ?????? (???. 3): ?????????
????????????? ?????????? ?? ??????? ?????
? ???????????? ??????????, ??????? ?????
?????? ????????? ?????????????? ????????.
????????? ???? ???????? ???????????? ?????? ??????? ?????? ????????????, ??? ???
?????????? ??????? ????????? ??????????
??????? ? ???????? ????? ?????????????. ???????? ?????, ??? ??????? ? ????????????
???????????? ?????????????? ????????? ??
???? ?????????????? ????? ????????? ????????? ? ???????????? ????????????????????
????????????.
# 167 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ??????? ??????????????? ?????????????? ?????????? ?? ?????? ????????????
R
R
O
Nu-
R
R
O
H+
R
Nu-
O
Nu-
H+
R
???. 3. ????? ??????? ?????????????? ????????????? ?? ???????? ????????????????? ??????????????
??????????????? ????????? AdN2, ??? ??????????? (Nu-) ????? ????????: OH, R 2O, NH2, R 2NH, R 2CH
? ?.?.
?
?
1
2
3
4
1
2
3
4
???. 4. ??????? ??? ????????? (?) ? ??????? (?) ??????????? ????????? (1) ? ??????? ?????????
(2-4) ????? ????????????? ??? ???????? ?????????????? ????????????? ??? ??????? ????????????
??????????: ??????????? ? 9:1, 5:1 ? 2:1 ??????????????
? ???? ???????????? ???? ???????????
??????????? ????????? ?????????? ?????????? 3-??? ? ??????? ??? ? ??????? ??????? ?????????????? ????????????? ??? ??
?????????? ? ?????? ????????. ? ?????? ?????
??????????????? ????????? ????? ????????? ?
????? ?????????? ??????? ? ?????? ???????? ? ??????????? ??????? ?????????? (? ??
?????????) ????? ???????? (?????) ????, ?
??????? ???????? ? ?????????? ???? ?? ????
??????? ????????? ?????? ??? (???. 4).
?????? ??????????? ???????, ?????????? ??? ??????????? ??????????? ????????
??????????? ? ??????? ?????????, ?????
??????????????? ? ?????????? ?????? ???
? ???????????? ? ?????????? ??????? ????????????? (???. 5). ??? ????? ?? ??????????????
??????, ? ??????? ???????? ??????????????
??????? ?????????????? ????????? ? Na, Si
? Fe ?? ????????? ? ???????????? ???????-
?? ?????????. ????????? ????????, ??? ??
??????????? ?????? ??? ??????? ???????
????? ????????, ??? Na, Si ? Fe (Bondar, Puzyr,
2004; Puzyr et al., 2012), ?????????? ????????,
??? ?????????????? ????????, ???????????? ? ??????? ???????? ?????????, ???????
? ???????? ? ??? ?????? ???.
??? ????????? ?????????????????
???????? (?????????????? ????, ???????????? ???????, ????? ?????????, ??????????? ?
???????? ?????????????) ??????????? ??????? ???????????? 3-??? ?????????? : ???
??? ????????? ??????????????? ?????????
???????? ??????????? 2:1, ????????? ??? ???
??????????? ?????????? ???????? ?????????? ?????????? ? ????????????. ?? ????????,
?????????? ????? ?????????? ??????? ???
?????? ??????? ???????????? ???????????
? ??????????? ??????? ?????????, ???????,
??? ?? 1 ?? ????????? ?? 3-??? ????? ????
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1
2
???. 5. ?????????? ?????? ????????? ?? ????????? ??????????? 3-??? (1) ? ??????????? ?????????
3-??? ? ??? (2), ?????????? ???????? ?????????????? ?????????????
??????? 1. ?????????? ?????? ???, ???????????????? ?? ?????????? ?? 3-??? ????????
?????????????? ?????????????, ? ??????????? ?? ???????? ??????????? ??????????? ? ????????
?????????
??????? ??????????? ???????????
(?????????? : ???)
?????????? ???????????????? ??????????
(??? ??? / ?? ?????????)
9:1
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149
?????????? ?????????????? ?? 70 ?? 150 ???
?????? ??? (????. 1). ?? ????????, ??? ?????? ???????????? ??????????? ???????????
(? ?????????, ?????????? ????????? ? ?????
?????????????? ?? ???????? ? ?????? ?????????? ???????? ?????????????? ?????????)
? ??????????? ??????? ?????????? ???????
?????????????? ????????????? ????? ???????? ?????????? ????????? ??? ? ??????????. ???????? ????? ??????? ???????? ????????? ?????????? ????????????.
?????????? ????????? ??????????? ??????????? ??????????? ? ??????? ????????
????????? ?? ??????? ???????-?????? ???????????? ????????? ?? ??????? ? ?????
?????????? (???. 6). ??? ?????? ????? ????????????????? ? ?????? ????, ??? ? ???????
20 ??? ????????? ????????? ? ?????? ???????? (???????????? HCl ? 5 ??, ?? ????? ?
2-3) ? ????????????? ??????????? ?????????????? ? ????????????? ???????????
?? 3-??? ?? ?????????? ???????????? ????????????? ?????????. ??? ????????????
??????? ???????? ???? ???????? (???. 7),
??? ??????? ??? ????????????? ?? ??????????? ????????? ? ???? ????????? ????????? (?????????). ??????? ???????? ?????,
??? ? ???? ??????????-????????????????
???????????? ???? ???????? ????????????
????????? ??????-?????????? ????????????? ??????????? ??????????????? ?????????. ???, ??? ???????????? ????????????
???? ???????????? ?????????? ?? ??????????? «?????????» ??????????? ????????? 3-???-??? ? ??????? ?? ???????????
?????????, ??? ??????? ???? ?????? ??? ???????????????.
?? ????????, ??? ?????????? ????????
????? ????? ?????????? ? ?????? ??????
???????????? ????????. ??????? ???????????? ?????????????? ???, ???????? ? ?????? ?????????, ????????? ??????????????
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?.?. ???????, ?.?. ??????? ??????????????? ?????????????? ?????????? ?? ?????? ????????????
?
?
???. 6. ???-??????????? ???????? ????????? ?? 3-???: ? ? ????????, ? ? ????? ????????????? ???
???????? ?????????????? ?????????????. ?????? ? 100 ?m
???. 7. ??????? ??? ????????? ??? (???????? ?????????), ???????????????? ?? ???????????
?????????? ?? 3-???. ??? ??????????? ???????? ??????????, ?????? ? 1 ?m
??? ????????????? ? ???????? ???????, ??????????? ? ?????????????, ???????????? ?
???????????? ???????? ??? ??????????????
??????? ??????. ????????, ? ???????? ?????? (Zhang et al., 2012) ???? ???????? ??????????? ?????????? ??? ???? ????? ???????????? ????????? ?? ?????? ???????????
? ????????????????? ??????????? ???? ?????????? ???????. ????? ????, ??????????
???????? ???????? ????????????? ???????
??? ??????????????? ??????? ???????? ????????????? ??????????, ??????? ???????
????????????????? ????????. ??????????????, ??? ???????? ? ?????? ????????? ???????
??? ?? ???? ?? ?????????????? ? ?????????
???????? ??????????? ????? ????????????
??????? ????????????? ???????? ???????
????????? ? ????????? ???????????. ???-
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?.?. ???????, ?.?. ??????? ??????????????? ?????????????? ?????????? ?? ?????? ????????????
???? ????????, ??? ??? ?????????? ??????
???????? ???????? ? ?????????????? ?????????? ????? ???????? ?? ????????????? ???
?????????? ?? ????????? ????? ??????????
??????????????? ??????? ? ???????? in vivo.
? ?????? ???????????? ???????????????
????????? ???-??? ??? ???? ????? ??????????????? ?????????? ???????? ????????????, ? ??????? ???? ????????, ??? ?????????? ????? ?????????? ?? ????????? ????????
? ??????? ????? ???? ?????????? ???? (Chow
et al., 2011).
?????? ?? ?????, ????????????? ???????????? ??????-?????????? ? ????????????
??????? ???????? ????????? ?? 3-??? ? ??
?????????? ? ????????? ???, ???????????
??????? ????????? ?????????????? ??????????, ? ????? ???????? ?????????????
??????? ?????????????? ?????????? ?????? ? ?????? ????????? ??????????? ???????? ????????????? ???????? ?????????? ????????????.
??????????
????? ???????, ? ?????? ???????????
??????????? ???????? ??????????????
?????????? ?? ???????????????? ??????-
???????????????? ? ????????????? ???????????. ????????, ??? ? ??????? ??????? ?????????????? ?????????????, ?????????? ?
?????? ???????? ?????, ?? ??????????? 1 ??
????????? ?? 3-??? ?? 20 ??? ????? ????
?????????? ?????????????? ?? 70 ?? 150
??? ?????? ???. ????????? ??????????
?????????????? ??????? ??????????? ??????? ? ????????? ??????????? ???????????
????????????????? ????????, ? ????? ??????????? ????????????. ??? ?????????
????????????????? ???????? ??? ?????????
?????????? ???????? ??????????? ???????
???????????? ??????????? ?????????? :
??? ???????? ??????????? 2:1, ?????????
??? ????? ??????????? ??????????? ?????????? ????????????? ????????????? ?????????? ?? ??????????? ? ?? 150 ??? ???
?? 1 ?? ?????????. ?? ?????? ?????????
??????????? ??????????? ???????, ??? ??????? ??? ????????????? ?? ???????????
????????? ??????????? ? ???? ?????????
(?????????). ?????????? ???????????? ?????
?????????? ?? ??????????? ??????? ????????? ??????????? ?????????? ?? ??????
??? ? ??? ? ???????? ??????????? ??
?????????? ? ??????????? ?????.
?????? ???????????? ?.?. ??????? (???? ?? ???, ?. ??????????) ?? ?????????? ???????????? ?????????? ? ??????????? ??????? ????????? ??????? ??????????? ???????????. ?????? ?????????? ?????????? ????????????? ?? «? ????? ?? ???????????
??????? ?????? ? ??????? ????????? ??????», ????????????? ????????????? ?? ? 220
(?????? «????????????? ????? ?????????????»), ?????????? ??? ? 24 (?????? 57).
?????? ??????????
1.
2.
3.
?????? ?.?., ??????????? ?.?., ???????? ?.?. (2006) ????????????????? (???) ? ?????????????? ???????? ??? ????????. ??????????: ???????, 288 ?.
?????? ?.?., ??????? ?.?., ???????? ?.?., ?????????? ?.?. (2008) ??????????? ????????????? ????????????? ?????????? ??? ??????????????? ?????????? ???? ??????????.
???. 418 (3): 1-4.
?????? ?.?., ??????? ?.?, ???????? ?.?., ?????? ?.?., ?????????? ?.?., ?????????? ?.?.,
?????? ?.?., ???? ??? ??, ???? ?? ????, ????????? ?.?. (2011). ????????????? ??????# 171 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ??????? ??????????????? ?????????????? ?????????? ?? ?????? ????????????
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
???????????????? (???) ? ????????? ???? ? ????????????? ???-????????????? ????????. ?????????????. 80 (2): 266?274.
?????? ?.?., ???????? ?.?., ?????? ?.?., ?????? ?. ??. (2012) ?????????????? ?????????? ??????????? ?? ?????? ????????????? ?????????? ????????? ?????? ? ????????
????????? ??? ????????????? ? ???????? ??????????. ?????????????????? ??????????.
54(2): 224-236.
???? ?. (1988) ???????????? ?????: ???????, ????????? ? ?????????, ?.3. ??????: ???,
456 ?.
????????? ?.?., ???????? ?.?., ??????? ?.?., ?????? ?.?., ?????? ?.??. (2011) ????????????? ???????????? ????????? ?????????, ?????????? ?? ????????????? ?????????????????????? ?????????? ??????????? ???????. ????????? ???????????????? ? ????????
?????????. 6(4): 63-67.
?????? ?.?., ??????? ?.?. (2005) ?????? ????????? ??????????? ????????? ??????? ? ?????????? ?????????? ?????????????. ?????? ?? ?2252192, ?????. 20.05.2005
???. ?14.
?????? ?.?., ??????? ?.?., ??????????? ?.?., ?????????? ?.?., ????????? ?.?. (2004?)
?????????? ???????????? ?????????? ?????????? ????????????? ??????????? ? ???????? ??????????????. ????????? ??????????? ?????????. 2-3 (31-32): 25-28.
?????? ?.?., ??????? ?.?., ??????????? ?.?., ??? ?.?., ????????? ?.?., ?????????? ?.?.
(2004?) ???????? ????????? ??????????????? ??????????? ???????????? ????? ???
?????????? ??????????? ???????? ????????? ???????????. ????????? ??????????? ?????????. 4 (33): 19-23.
?????? ?.?., ????????? ?.?., ???????? ?.?., ??? ?.?, ??????????? ?.?., ??????? ?.?.,
??????? ?.?. (2005) ? ??????????? ????????????? ???????? ?????????? ????? ???????????????? ???????????. ????????? ??????????? ?????????. 1 (34): 20-24.
?????? ?.?., ????? ?.?., ?????? ?.?., ??????? ?.?. (2010) ?????????? ? ?????????????: ?????????? ??? ????????? ?????? ? ???????? ???????????? ????-??????. Journal of
Siberian Federal University. Biology. 3 (4): 418-433.
???????? ?.?., ?????? ?.?. (2006) ???????????? ?? ??????????????? ???????????????? ?
???????? ???????? ??? ????????????? ??????????. ????????????? ?????????. 4: 65-70.
???????? ?.?., ?????? ?.?., ???????? ?.?., ?????? ?.?. (2009) ????????????? ? ?????????
?????????? ??????????? ? ?????? ?????????? ??????? ???????????? ???????? ??? ???????????? ????????. ????????? ????????????????? ???????? ? ????????. 11: 542-546.
Bondar V.S., Puzyr A.P. (2004) Nanodiamonds for biological investigations. Phys. Solid State. 46:
716-719.
Bondar V.S., Pozdnyakova I.O., Puzyr A.P. (2004) Applications of nanodiamonds for separation
and purification of proteins. Phys. Solid State. 46: 758-760.
Bondar V.S., Purtov K.V., Puzyr A.P, Baron A.V., Gitelzon I.I. (2008) Catalytic activity of
nanodiamond particles in organic reactions. Dokl. Biochem. Biophys. 418: 11-13.
Chow E.K., Zhang X.Q., Chen M., Lam R., Robinson E., Huang H., Schaffer D., Osawa E., Goga
A., Ho D. (2011) Nanodiamond therapeutic delivery agents mediate enhanced chemoresistant
tumor treatment. Sci. Transl. Med. 3: 73ra21.
# 172 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ??????? ??????????????? ?????????????? ?????????? ?? ?????? ????????????
18. Gibson N., Shenderova O., Luo T.J.M., Moseenkov S., Bondar V., Puzyr A., Purtov K.,
Fitzgerald Z., Brenner D.W. (2009) Colloidal stability of modified nanodiamond particles.
Diam. Relat. Mater. 18: 620-626.
19. Mamaeva E.S., Baron A.V., Puzyr A.P., Burov A.E., Bondar V.S. (2011) Modified nanodiamonds
obtained by detonation synthesis in constructing biochemical indication systems (as exemplified
by the glucose determination system). Dokl. Biochem. Biophys. 439: 182-184.
20. Mogilnaya O.A., Puzyr A.P., Bondar V.S. (2010a) Growth and bioluminescence of luminous
bacteria under the action of aflatoxin B1 before and after its treatment with nanodiamonds.
Appl. Biochem. Microbiol. 46: 33-37.
21. Mogilnaya O.A., Puzyr A.P., Baron A.V., Bondar V.S. (2010b) Hematological parameters and
the state of liver cells of rats after oral administration of aflatoxin B1 alone and together with
nanodiamonds. Nanoscale Res. Lett. 5: 908-912.
22. Puzyr A.P., Baron A.V., Purtov K.V., Bortnikov E.V., Skobelev N.N., Mogilnaya O.A., Bondar
V.S. (2007a) Nanodiamonds with novel properties: A biological study. Diam. Relat. Mater. 16:
2124-2128.
23. Puzyr A.P., Purtov K.V., Shenderova O.A., Luo M., Brenner D.W., Bondar V.S. (2007b) The
adsorption of aflatoxin B1 by detonation-synthesis nanodiamonds. Dokl. Biochem. Biophys.
417: 299-301.
24. Puzyr A.P., Burov A.E., Bondar V.S., Trusov Yu.N. (2010) Neutralization of aflatoxin B1 by
ozone treatment and adsorption by nanodiamonds. Nanotechnologies in Russia. 5: 137-141.
25. Puzyr A.P., Pozdnyakova I.O., Bondar V.S. (2004) Design of a luminescent biochip with
nanodiamonds and bacterial luciferase. Phys. Solid State. 46: 761?763.
26. Puzyr A.P., Bondar V.S., Bukayemsky A.A., Selyutin G.E., Kargin V.F. (2005) Physical and
chemical properties of modified nanodiamonds. NATO Science Series. II. Mathematics,
Physics and Chemistry. 192: 261-270.
27. Puzyr A.P., Burov A.E., Selyutin G.E., Voroshilov V.A., Bondar V.S. (2012) Modified nanodiamonds
as antiwear additives to commercial oils. Tribology Transactions. 55: 149-154.
28. Purtov K.V., Puzyr A.P., Bondar V.S. (2008) Nanodiamond sorbents: new carriers for column
chromatography of proteins. Dokl. Biochem. Biophys. 419: 72-74.
29. Purtov K.V., Petunin A.I., Burov A.E., Puzyr A.P., Bondar V.S. (2010) Nanodiamonds as
carriers for address delivery of biologically active substances. Nanoscale Res. Lett. 5: 631636.
30. Purtov K.V., Petunin A.I., Puzyr A.P., Burov A.E., Bondar V.S. (2011) Model drug delivery
system based on nanodiamonds. Nanotechnologies in Russia. 6: 256-264.
31. Shishatskaya E.I. (2008) Biomedical investigation of biodegradable PHA. Macromol. Symposia.
269: 65-81.
32. Shishatskaya E.I., Chlusov I.A., Volova T.G. (2006) A hybrid PHA-hydroxyapatite composite
for biomedical application: production and investigation. J. Biomater. Sci. Polymer. Edn. 17:
481-498.
33. Shishatskaya E.I., Voinova O.N., Goreva A.V., Mogilnaya O.A., Volova T.G. (2008)
Biocompatability of polyhydroxybutyrate microspheres: in vitro and in vivo evaluation.
J. Mater. Sci: Mater. Med. 9: 2493-2502.
# 173 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ??????? ??????????????? ?????????????? ?????????? ?? ?????? ????????????
34. Shishatskaya E.I., Goreva A.V., Kalacheva G.S., Volova T.G. (2011) Biocompatability and
resorption of intravenously administered polymer microparticles in tissue of internal organs of
laboratory animals. J. Biomater. Sci. Polym. Ed. 22: 2185 ? 2203.
35. Zhang Q., Mochalin V.N., Neitzel I., Hazeli K., Niu J., Kontsos A., Zhou J.G., Lelkes P.I., Gogotsi
Y. (2012) Mechanical properties and biomineralization of multifunctional nanodiamond-PLLA
composites for bone tissue engineering. Biomaterials. 33: 5067-5075.
The Engineering of Composite Materials Based
on Nanodiamonds and Degradable Biopolymers
Vladimir S. Bondara,b,*, ?lexey P. Puzyra,b,
Anastasiya V. Gorevaa,b and ?ndrey E. Burova,c
a
Siberian Federal University,
79 Svobodny, Krasnoyarsk, 660041 Russia
b
Institute of Biophysics SB RAS,
50/50 Akademgorodok, Krasnoyarsk, 660036 Russia
c
SDTB ?Nauka? KSC SB RAS,
53 Mira, Krasnoyarsk, 660049 Russia
Issues related to engineering and application of composites based on nanodiamonds and degradable
polyhydroxyalkanoates are discussed. The ability of composite engineering is demonstrated on the
3-polyhydroxybutyrate microspheres ? modified detonation nanodiamonds composite fabricated
through the bimolecular nucleophilic addition (AdN2) reaction. It was shown that during 20-min
incubation at acid pH from 70 to 150 ?g of the nanoparticles can be immobilized on 1 mg of the
microspheres depending on the components weight ratio.
Keywords: polyhydroxyalkanoates, 3-polyhydroxybutyrate, modified nanodiamond, composite
materials.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2013 6) 175-184
~~~
??? 616.13-004.6-055.1:575
????? 9?21.3 ? ???????????? ?????????
??????? ??????????? ????????????? ? ??????
?.?. ????????*, ?.?. ????????,
?.?. ????????, ?.?. ???????
???????????? ??????????????? ??????????? ???????????
??. ?.?. ?????-??????????
?????? 660022, ??????????, ??. ????????? ?????????, 1
Received 27.02.2013, received in revised form 03.03.2013, accepted 07.07.2013
???? ???????????? ? ??????? ??????????? ?????? 9?21.3 ? ???????? ???????????
????????????? ? ????????? ?????????. ? ???????????? ???????? 255 ??????? (211 ??????,
44 ???????) ? ????????? ???????? (??) ???????????? ???? ? ???????? ? 65 ???
(52,56±7,98 ???). ???????? ??? ???????? ?? ???????? ????? ?????-???????????? ???????.
??????????? ????? ??????????? ? ??????? ???????????? ?????? ??????? (???) ? ??????
????????? ??????? (????? TaqMan, Applied Biosystems 7900HT). ? ???????????? ?????????
??? ???????????????? ???????????? (???), ????????????? ? ?????? 9p21.3 ? rs10757278
? rs1333049. ?????? ??????? ??????? ????????? ??????????? ????? ????????????? ?
???????????? ?????? ? ???? ??????????? ???????????????? ?????????????? (Philips Allura
Xper FD10), ??????????? ?????????? ?????????? ?????????, ???????????? ??????? Gensini
? SYNTAX. ??????? ? ????????? ????????? ? ?????? ???????? ??????????? ????????? GG
rs10757278 ? ?? rs1333049 ? ???????? ??????????? ?????????????.
???????? ?????: ??????? ????????, ???????????????? ???????????, rs10757278, rs1333049,
????? 9?21.3, ????????????.
????????
? ??????? ??????????? ??????? ????????
????? ???????????? ??????????, ??????????? ?????????????? ??????????????????? ?
??????????? ??????? ?????? (???) ? ???????? ???????? (??), ????????? ???????????.
????????? ? ???????? ???????? ??? ????????? ?????????????? ??????? ?????? ? ????????
*
???????? ???????????????? ?????????????
(???) ????????, ????? ????????? ?????????? ??????? ????????????? ????????????? ???????????? (Genome-Wide Association
Study ? GWAS). ??????? ? 2007 ????, ?????
??????? ???? ???????????????? ???? ?????? 9?21.3 ? ???????? ?? ? ??????????, ???
?????? ???? ???????????? ? ?????????
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: shesternya75@mail.ru
# 175 #
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?.?. ????????, ?.?. ????????? ????? 9?21.3 ? ???????????? ????????? ??????? ??????????? ??????????????
??????????, ??????? ? ?????? (????????
? ??., 2012; ???????? ? ??., 2012). ? ??????????? ???? ???????? ????????? ????????
????? ???, ??????????????? ? ?????????
??? ? ?? (Deloukas et al., 2012; Prins et al.,
2012). ?????? ???????? ??????? ? ??????????? ???????????? ??????????? ??? ? ??,
????????? ? ?????????? ???????, ????????
????????? ??????? ?????? (???????? ~ 58kb),
????????????? ?? ???????? ????? 9-? ????????? (????? 9?21.3). ?????????? ? ???? ??????? ?????, ????????????? ?? ?????? ?????????
??????, ??????????? ?????????? ????????????? ??????????? ?????? 9?21.3 ? ???. ???????????, ??? ? ?????? 9?21.3 ?????????? ???????????? ???????????? ???????????????
??????? (????? ANRIL). ??????? ???????
?????????? ANRIL ??????? ? ?????? ? ???????, ?????????? ?????????????? (Broadbent
et al., 2006). ?????????? ?????????????, ???
9?21.3 ?? ?????? ?? ??????? ????????????,
?? ? ?? ????? ???????????????? ??????????? ????????????? (Patel et al., 2010; Adrissino
et al., 2011; Chan et al., 2011). ????????????
???? ???????? ? ??????? ??????????????
?????????? ???????????, ????????????
????? 20 ????????????, ???????????? ??
???????? ???? ?????? 9?21.3. ??? ???????
??????????? ? ???????????????? ??????
???????? ?????? ????? ?? ?????????? ?????? 9?21.3 ?????? ? ???????? ???????????
?????????????, ? ?? ? ????????? ?? (Chan et
al., 2013). ? ????? ?????? ??????????????
?????, ??????????? ???????? ????? ???????, ???? ?? ???????.
???? ?????? ?????? ? ??????? ??????????? ??? rs10757278 ? rs1333049 ??????
9?21.3 ?? ???????? ???????????? ??????????? ????????????? ? ??????? ??.
?????? ??????? ?????? ?????? ?? ?????????
???????????? ? ????? ????????? ? ?????????????? ??? (Schunkert et al., 2010). ??????????? ????? ?????? ? ??????? ?????? 9?21.3
?? ??????????? ??????????????????? ??????
(Buysschaert et al., 2010). ?????? ???????? ?????????? ???????? ??????? ??????? ??????
???????? ??????????? ?? ??? ???.
??????? ????? ??? ????????? ????? ??
????????? ?????????? ??????????? ????????? ?????? 9?21.3 ? ???????? ? ??????????
???????????????????? ????????? ??????????? ?????. ??????? ?????? ? ???????????
?????? ????? G rs10757278 ? ??????????????? ? ????????????? ????? ?????????
?????????? ??????? (??), ???????? ??????
????? ?? (Patel et al., 2010). ? ??????? ????????? ???????????? Ottawa Heart Genomics
Study ?????????? ? ???????? ????????? ??
???? ???????? ??? ??????? ??? ??????? ?????? rs1333049 (Dandona et al., 2010). ?? ?????? ???? ???????, ???????? ??????? ??????
????????? ? ??????
? ???????????? ???????? 255 ???????,
??????????????????? ? ????????? ?? ?
??????? ???????????????? ?????? ?. ???????????: «????????? ??????????? ????????
?20 ??. ?.?. ???????» ? «??????? ??????????? ????????» ? 01.01.2009 ?? 30.06.2010 ??.
? ??????????????? ????????? ?????????.
?????????? ????????? ????????: 1) ???????????????? ??????? ??; 2) ??????? ? ??????? ? ???????? ? 65 ???; 3) ????????????
????; 4) ??????? ???????????? ???????????????? ???????? ????????. ??????? ???????????? ???????? ????????? ????????? ????????????? ???????????????? ????????????
????????????. ??????? ??????? ????????? ??
22 ?? 65 ??? (??????? ??????? 52,56±7,98 ???;
???????? 25, 50, 75 % = 47,0/53,0/59,0). ? ??????????? ?????? ??????????? ??????? ? 211
??????? (82,7 %), ?????? ???? 44 (17,3 %).
??????????? ? ??????????????? ?????????????? ??????? ????????? ? ????. 1.
# 176 #
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?.?. ????????, ?.?. ????????? ????? 9?21.3 ? ???????????? ????????? ??????? ??????????? ??????????????
??????? 1. ??????????? ? ??????????????? ?????????????? ??????? ??
?????????? ???????, ???.
(% ?? ?????? ?????)
??????????????
??????????????? ??????, ??????? ?????
???????, ???, M±?
52,56±7,98
??? (???????/???????)
211 (82,7) / 44 (17,3)
???????? ?????? ? ????????
25 (9,8)
???????????? ??????????? ? ????????
170 (66,7)
??????????????????? (????? 5,0 ?????/?)
162 (63,5)
???????
152 (59,6)
?????????? ????? ???? (??? ?25)*
167 (65,5)
???????? (??? ?30)*
63 (24,7)
?????????????? ??
?? ? ???????? ???????? ST
199 (78,0)
?? ??? ??????? ???????? ST
56 (22,0)
???????? ??????????? ??
112 (43,9)
?????????? ?? (??????? ??????, ???????, ??????)
118 (46,3)
??????????? ??
25 (9,8)
Q-???????? ??
180 (70,6)
???????????????? ??????
????????? ?????????? ???????:
- ??????????????
96 (37,7)
- ??????????????
84 (32,9)
- ??????????????
57 (22,4)
- ?????????? ???????????????????? ?????????
18 (7)
?????????? ??????????????? ???????? ????????, M±?
2,20±1,38
?????? GENSINI, M±?
48,27±39,55
?????? SYNTAX, M±?
13,90±8,27
* ??? ? ?????? ????? ???? ?? Quetelet (?????(??)/????(?)2).
???????????-???????????? ????????????.
???????? ??? ???????? ?? ?????????? ???????? ????? ?????-???????????? ???????.
??????????? ????? ??????????? ? ???????
??? ? ?????? ????????? ??????? ? ???????????? ? ?????????? ?????-????????????? ??
??????? AB 7900HT (????? TaqMan, Applied
Biosystems, ???). ? ???????????? ???????
??? ???, ????????????? ? ?????? 9p21.3 ?
rs10757278 ? rs1333049.
??????????? ?????????? ??????? ??????????? ?? ???????? Allura Xper FD10 (Philips,
?????????). ?????? ???????????? ????????? ???????????????? ?????????????? ????
??????? ? ???????????? ?????? ??? ??????
??????? ??????? ????????? ??????????? ????? ??????????? ??????? ?????????? ?????????? ????????? ?? ? ???????????? ???????
Gensini ? SYNTAX.
??????? ?????????? ???????? ??????????? ?? ???????????? ??????????????? ???????? (>50 % ????????) ???????? ? ?????? ????????? ??????????? ????? (????? ????? ??,
?????????????, ??????? ? ?????????? ???-
# 177 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ????????? ????? 9?21.3 ? ???????????? ????????? ??????? ??????????? ??????????????
????? ?????? ?? ????????????? ??????? ?
????????? ?????, ???????? ??????????????? ????? ????? ?? ? ?????? ??).
? ??????? Gensini ????????? ??????????
????????? ??????????? ?????, ? ????????????????? ????????? ??????????, ??????
? ?????? ???????????? ????? ???????? ?????????? ???????, ????? ?????? ????, ??????????????? ???????. ???????????? ????????????? ??????? Gensini ???????? ????
?????????????? ?????????? ??????? ???????. ??????? ???????? ?? 25 % ??????????? 1
????, 25-50 % ? 2, 50-75 % ? 4, 75-90 % ? 8, 9099 % ? 16, ?????? ???????? ? 32 ?????. ?????
?????????? ???????? ???????? ?? ??????????? ?? 0,5 ?? 5 ? ??????????? ?? ???????????
??????? ???????. ??? ??????? ??????????
???????? ?????????? ???????? ??? ???????
?? ??? ??????????? (Montorsi et al., 2006).
??? ??????? ??????? SYNTAX ?????????????? ?? ?????? ???????? ? ???????????, ??
? ????????????? ???????, ??????? ?????????? ??, ???????? ???????? ? ??. ?????? ????????????? ???????????? ????? SYNTAX
???????? ???? ?????? ?????????????? ???????? ??? ??????? ??????????????????? ????????? ? ??????. ????? ?????? ?22 ????????
???????? ???????, 23-32 ????? ? ??????? ?
?33 ?????? ? ??????? ????????? ???????????
????? (Sianos et al., 2005).
?????????????? ????????? ??????????
?????? ??????????? ??? ?????? ?????????
SPSS, ?????? 19.0. ???????????? ??????????
??????????? ???????????? ???????????? ???
???????????? ????????? ? ???? ??????????
?????, ??? ?????????????? ? ? ???? ???????
?????????????? (?) ? ??????????? ?????????? (?). ?????????? ???????? ????????????
????????? ? ??????? ?????????? ?????????
??? ?????? ?????????????????? ???????? ?2
??????? ? ????????? ?? ?????????????. ???
??????? ????????????? ???????? 5 ? ????? ??-
??????????? ?????? ???????? ??????. ???
?????? ?????????? ?????????????? ????????
?? ?????????????? ????????? ?????????
????????? ????? ?? ?????????????????? ????????? ???????? ?????-?????. ???????? ??
???? ??????? ????????? ??? ?????????????
???????? ??? ?<0,05. ??? ?????? ???????
????? ?? ??????????? ?????? ??? ????????
??????????? ?????? ????????? ?????? (??)
? ???????? ????????????? 2?2 ? ???????? ????????????? ?????????? (??).
?????????? ? ??????????
?? ??????? ????????? ??????????? ????? ???????? ???? ????????? ?? ???????
????-, ????- ? ?????????????? ?????????
??, ? 10 (3,9 %) ????????? ???? ????????
????????? ??, ? 8 (3,1 %) ??????? ???????
?????????????? ???????? ??? ??????????????????? ????????? ? ?? (????. 1). ????????
??? ??????????????????? ????????? ?? ????
????????? ?? ??????????? ???????????????
?????. ??? ??????? ?????????? ??????????
???????????? ? ???????? (14 (5,5 %) ???????)
?????? ???????? ?? ???????????.
?????????? ????? ??????????? ??? ???????? ??????? ?? rs1333049 ? ??????? GG
rs10757278. ???? ?? ???????? ?????????????
???????? ?????????? ????? ?????????? ????????? ?????? 9?21.3 ? ??????????? ?????????? ???????. ??? ?? ????? ????? ???????
? ?????????????? ????? ????????? ???????
GG rs10757278 ??????? ? 32,0 ?????? 21,0 %
????? ??????? ? ????- ? ?????????????? ?????????? ? 33,3 ?????? 23,3 % ??? ???????? ?? rs1333049 ??????????????. ?????????
?????????? ???????? ? ????????? ?? ??????
????? rs1333049 ? rs10757278 ???? ???? ?
????????? ? ??????? ??????????, ?????? ??
????? ?????????????? ??????????. ? ???????? ??????? ?????? ??????? ? ????????
????? ???? ???????? ?????????? ???????????
# 178 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ????????? ????? 9?21.3 ? ???????????? ????????? ??????? ??????????? ??????????????
???????? ? ?????????? ?? rs1333049 (42,2 %
?????? 21,5 ? ?????? 1-3 ????????, ?=0,005) ?
GG rs10757278 (36,6, % ?????? 20,7 ? ??????
1-3 ????????, ?=0,005). ???????? ???????? ??
rs1333049 ? ???????? GG rs10757278 ?????
????? ??????? ??????????? ??????? ?????
???? ????????. ?????????? ???????? ??????????? ?????????? ??? ?????? 9?21.3 ? ???????? ??????? ?????????? ?? ? ????????
?????? ????? ??. ???????? ????????? ?????
?? rs1333049 ? GG rs10757278 ????? ?????
????? ??????? ???? ???????? ??????? ??????
????? ?? (????. 2).
???????????? ??????????? ??????? ????????? ??????????? ????? ? ?????? ??????
????? ??????? ?????? Gensini, ???????????
?? ?????? ??????????, ?? ? ???????????????? ?????????? ????????? ????????. ???????? ????????? ?? rs1333049 ? GG rs10757278
????? ?????????? ????? ??????? ????????
??????? Gensini ? ????????? ? ??????????,
???????? ? ???????? ???? ???? ?????? ????? ??? ?? ???????? ??? ?????. ????????
??????? SYNTAX ???? ???? ? ????????? ????????? ????? ????? ????????? ???, ??????
???????? ?? ???? ????????????. ?????????
???????? ????? ???????? ????????? ???????
???????? ??? ??????????????????? ?????????
?? ??? ??????? ??????? SYNTAX, ? ?? ?????
??? ?????? Gensini ???????????? ?????? ? ????????? ? ?????????????? ??. ? ?????????
?????????????, ?????????? ????????? ??
???????????? ???????? ? ????????????? ?????? ????????? rs10757278 ? rs1333049 ???? ??
???????? (????. 2).
??????????? ??????????? ?????, ?????????? ??????????? ?????? 9?21.3 ? ?????????? ??????????????, ????????? ? ??????????? ? ???????????? ??????????; ? ??
?? ????? ???????????? ????????? ??????
? ????? ?? ??????? ???????? ??????????
(Hinohara et al., 2008; Shen et al., 2008). ????
??????? ? ?????? ???????????? ??????, ?????????? ? ???? ?????????????? ???????, ??
?????????? ??? ?????? 9?21.3 ? ????????
???????????????????? ????????? ??????????? ????? ? ??????? ??. ?????? ?????????????? ???????? ??????????????, ???? ??????
??????? ??????????? ?????????? ? ????????? ?????????.
? ???????????????? ?????????????, ???????????, ??? ???????, ? ?????? ?????????????? ?????????, ??????????? ?????? ??????????????, ??????????? ??? ?? ??????????
?????????? ? ??????? ? ?????? ??????????
?????????, ??? ? ??? ?????????? ????????
?????????? ?????????? ????????????. ????? ?? ?????????? ??????????? ? ???????????? ?? ??????????? ?????????? (Dandona
et al., 2010; Patel et al., 2010). ???? ????? ??
??? ???? ????????? ?? ???????? ????????? ?
???? ???????, ??????????? ? ????????? ?????????? ?????? ? ????????? ?? (Adrissino et
al., 2011). ???????? ??????? ??????????? ???
?????? ???????????, ????? ? ????????????
????????????, ??? ????????? ???????, ?????????? ?? ? ? ??????????? ??????????
? ???????????? ????????????, ? ??????? ??
??? ??????????? ? ????????? ?????????? ?????? ????? ?????? ??????????. ?????? ???????????????, ????????? ??????? ???????
?? ??????????, ???????? ????????????? ??????? ????????? ?? ????????? ?????????/
??????????. ? ?????????, ??????? ????????????? ?????????, ?????? ????? ?????????
???????. ???, ? Ottawa Heart Genomics Study
????????, ??????? ???????? ??????, ??????????? ?? ???????????? (Dandona et al., 2010).
? ????? ?????? ??????? ?? ???????? ? ???????????? ????? ??? ??????????? ? ???????, ????????????? ???????? ??????? ?? ????????????? ????????? ?? ????????????.
???????? ???? ????????????? ?????????? ? ????????? ????????????? ???????
# 179 #
10 (24,4 %)
??????? ????? ???? ????????, n ( %)
13 (20,3 %)
???????? ??????? Gensini ? 70
??????, n ( %)
9 (17,0 %)
2 (14,3 %)
????????? ????????? ??, ??? ( %)
????????????? ??,
n ( %)
?
0,002*
GG
?? 11 (50,0 %)
??? 44 (21,1 %)
0,031*
?? 15 (36,6 %)
??? 36 (20,7 %)
?? 21 (32,8 %)
??? 31 (20,3 %)
59,40±45,07
11 (78,6 %)
32 (60,4 %)
26 (51,8 %)
13,73±7,90
?? 16 (32,0 %)
??? 35 (21,0 %)
?? 12 (22,6 %)
??? 43 (24,2 %)
?? 1 (7,1 %)
??? 54 (25,2 %)
15,53±9,08
1,38 (1,11-2,42)
30 (46,9 %)
45,02±34,67
0,125*
0,820*
0,111*
0,161
0,048*
0,031
0,238
2,38±1,50
2,21 (1,06-4,61)
16 (39,0 %)
2,19±1,24
3,75 (1,53-9,22)
5 (22,7 %)
AG
2 (14,3 %)
9 (15,5 %)
8 (14,8 %)
13,45±8,78
10 (15,4 %)
45,98±43,69
9 (20,0 %)
2,00±1,41
5 (23,8 %)
GG
CC
?? 11 (50,0 %)
??? 51 (23,3 %)
?? 19 (42,2 %)
??? 38 (21,6 %)
2,44±1,55
0,006 #
?? 25 (38,5 %)
??? 33 (20,9 %)
11 (78,6 %)
36 (62,1 %)
28 (51,9 %)
13,38±7,86
?? 18 (33,3 %)
??? 39 (23,2 %)
?? 13 (22,4 %)
??? 49 (26,9 %)
?? 1 (7,1 %)
??? 61 (27,4 %)
15,44±9,15
0,095#
0,494#
0,138#
0,100
0,019
0,005 #
0,105
0,004 #
?
58,13±45,24
1,78 (1,19-2,65)
30 (46,1 %)
43,84±34,74
2,10 (1,26-3,50)
17 (37,8 %)
2,17±1,27
3,16 (1,41-7,07)
6 (27,3 %)
CG
rs 1333049
&
n ( %)- ?????????? ??????? ( % ?? ?????? ?????).
* ?????????????? ???????? ?? ???????????? ????????? ????? ?????????? ???????? GG rs1075728.
# ?????????????? ???????? ?? ???????????? ????????? ????? ?????????? ???????? ?? rs1333049.
3. ??? ?????????????? ????????? (????????? ?????????? ????????, ?????? Gensini, ?????? SYNTAX) ??????????? ???????? ???? ????? ?????????? ?????? ????????? (AA, AG
? GG ??? rs1075728 ???? GG, GC ? CC ??? rs 1333049).
4. C???????????? ??????????? ???????? p ? ????????????? ????????? ???????? ?????? ???????.
8 (16,2 %)
?????????????? ????????? ??, n
( %)
1.
2.
13,68±8,59
?????? SYNTAX, M± ?
?? (95 % ??)
48,94±43,89
?????? Gensini, M± ?
?? (95 % ??)
2,08±1,34
6 (27,3 %)
????????? ?????????? ????????,
M± ?
?? (95 % ??)
????????? ?????? ????? ??, n ( %)&
AA
rs 1075728
??????? 2. ?????????? ??????????? ????????? ???????? rs1075728 ? rs1333049 ? ????????????????? ???????????????? ??????????? ?????????????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?
69,25±29,26
0,068
5 (71,4 %)
43,87±42,65
0,687* 21 (61,8 %)
?? 2
(28,6 %)
??? 6
(21,4 %)
?????? G
46,68±36,62 57,72±47,27
p
0,570
0,271
?? 4 (44,4 %)
??? 6
0,176#
(21,4 %)
2,36±1,80
0,013#
?? 15
(41,7 %)
??? 32
(21,6 %)
44,83±35,96 57,96±46,90 0,040
2,08 (1,17-3,70)
0,128
0,002#
p
2,45±1,51
3,67 (1,51-8,69)
21 (58,3 %)
2,11±1,37
CC
?? 10
(52,6 %)
??? 41
(22,7 %)
???????
?????? G
rs1333049
?? 1 (33,3 %)
??? 10
0,792# 9 (47,4 %)
(26,3 %)
CC
42,30±41,34 58,91±38,53
0,029* 5 (55,6 %)
?? 13
(38,2 %)
??? 30
(20,5 %)
0,097
0,213
2,07±1,34
0,002* 2 (66,7 %)
p
???????
2,45±1,53
2,39 (1,08-5,33)
1,000
2,00±1,03
4,11 (1,56-10,86)
2,00±1,34
2,16±1,29
GG
?? 10
(52,6 %)
??? 37
(21,3 %)
???????
?? 1 (33,3 %)
??? 7
0,587* 9 (47,4 %)
(20,0 %)
GG
p
rs1075728
&
n ( %)- ?????????? ??????? ( % ?? ?????? ?????).
* ?????????????? ???????? ?? ???????????? ????????? ????? ?????????? ???????? GG rs1075728
# ?????????????? ???????? ?? ???????????? ????????? ????? ?????????? ???????? ?? rs1333049.
3. ??? ?????????????? ????????? (????????? ?????????? ????????, ?????? Gensini) ??????????? ???????? ???? ????? ?????????? ?????? ????????? (AA, AG, GG ??? rs1075728
???? GG, GC, CC ??? rs 1333049).
4. ????????????? ??????????? ???????? p ? ????????????? ????????? ???????? ?????? ???????.
1.
2.
?????? Gensini,
M±?
?? (95 % ??)
?????????
??????????
????????,
M±?
??????? ?????
???? ????????,
n ( %)
?? (95 % ??)
?????????
?????? ????? ??,
2 (66,7 %)
n ( %)&
?????? ?
???????
??????? 3. ????????? ?????? ?????????? ??????????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ????????? ????? 9?21.3 ? ???????????? ????????? ??????? ??????????? ??????????????
?????????, ?? ??????? ?????????? ????????? ?????? ????????????? ??????????.
?????????? ?????? ???????? ???????????
????????? ????? ??? ?????? 9?21.3 ? ?????
??????? ??????????????????? ??????????
?? ????????????? ? ??????. ???????? ???????? ?? rs1333049 ? ???????? GG rs10757278
????? ????? ??????? ??????????? ???????
????? ???? ???????? ??, ? ?????? ???? ????
???? ????????? ?????? ????? ??. ??? ????
???????????? ?????? Gensini ??? ????????????? ??????? ???? ?????? ????? ?????? ?
????????? ?? rs1333049 (????. 3).
?????????????? ? ????. 2 ? 3 ??????
???????? ??????????? ?????? ??????????
? ????????????? ????????? ????????? ????? (???????? GG rs1075728 ? ???????? ??
rs1333049) ??? ?? ???????, ??? ? ?? ????????? ???????????????????? ????????? ??????????? ?????. ????????? ??? rs1333049 ?
rs10757278 ??????????? ?????????? ?????? ?
?? ?????????? 1025 ???????????? ???. ???
???????? ????? GG rs1075728 ? ?? rs1333049
???????? ???????????????????, ? ????????
???????????? (r=0,949). ????????, ??? ???
rs1075728 ? rs1333049 ? ????? ????????? ?????? ? ???? ???? ?????????, ??? ??????????? ?
?????????? ???????. ?? ?????? ???????????? INTERHEART, ??????????? ?????????? ????? ????? ????? ??? ?????? 9?21.3
????????? ?? 0,92 ?? 0,98 ? ??????????? ??????? ????????? (Do et al., 2011).
??????????
??????????? ????????????? ????????
?????????? ??? rs1075728 ? rs1333049 ?????? 9?21.3 ? ???????? ????????????????????
????????? ??????????? ????? ? ??????,
??????? ??. ? ??????????? ????? ??????
???????? ??????? ????????????? ????????
?????????? ???????? ??????????????? ????? ? ????????? ????????????, ???, ? ????
???????, ??? ???????????? ???????????
??????? ?????. ?????????? ????????????
??????????? ? ????????? ????? ??????
????? ? ?????????? ???????-????????????
??????????? ????????? ??????????????
????????????? ? ??????????? ??????????????? ????? ???????? ???????????. ?????????? ? ????? ?????? ??????????? ??????????? ????????? ?????? 9?21.3 ? ????????
??????????? ????????????? ????? ???? ???????????? ??? ??????????? ??? ?????????
???????????? ??.
?????? ??????????
1.
2.
3.
???????? ?.?., ??????? ?.?., ????? ?.?. ??????? ?.?., ???????? ?.?., ????????? ?.?.,
??????? ?.?. (2012) ???????? ??????????? ????? ??????? ????????????????? ?????????????? ? ????????? ???????? ?? ????????? ?????????. ??????? ???? 5: 24-29.
???????? ?.?., ??????? ?.?., ???????? ?.?. ????????? ?. ?., ??????? ?. ?., ????? ?.
?., ???????? ?. ?., ??????? ?. ?. (2012) ???????????? ???? ????????????? ?????????
9?21.3 ? ?? ??????????? ? ??????????? ????????????????? ? ???????? ???????? ????????.
?????????? ???????????????? ?????? 6(98): 14-18.
Adrissino D., Berzuini C., Merlini P.A., Mannucci P.M., Surti A., Burtt N., Voight B., Tubaro M.,
Peyvandi F., Spreafico M., Celli P., Lina D., Notarangelo M.F., Ferrario M., Fetiveau R., Casari
G., Galli M., Ribichini F., Rossi M.L., Bernardi F., Marziliano N., Zonzin P., Mauri F., Piazza A.,
Foco L., Bernardinelli L., Altshuler D., Kathiresan S. (2011) Influence of 9p21.3 genetic variants
on clinical and angiographic outcomes in early-onset myocardial infarction. J. Am. Coll. Cardiol.
58(4): 426-434.
# 182 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ????????? ????? 9?21.3 ? ???????????? ????????? ??????? ??????????? ??????????????
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
Broadbent H.M., Peden J.F., Lorkowski S., Goel A., Ongen H., Green F., Clarke R., Collins R.,
Franzosi M.G., Tognoni G., Seedorf U., Rust S., Eriksson P., Hamstein A., Farrall M., Watkins H.
(2008) Susceptibility to coronary artery disease and diabetes is encoded by distinct, tightly linked
SNPs in the ANRIL locus on chromosome 9p. Hum. Mol. Genet. 17: 806-814.
Buysschaert I., Carruthers K.F., Dunbar D.R., Peuteman G., Rietzschel E., Belmans A., Hedley A.,
Meyer T.D., Budaj A., Van de Werf F., Lambrechts D., Fox K.A.A. (2010) A variant at chromosome
9p21 is associated with recurrent myocardial infarction and cardiac death after acute coronary
syndrome: The GRACE Genetics Study. Eur. Heart J. 31: 1132?1141.
Chan K., Motterle A., Laxton R.C., Ye S. (2011) Common variant on chromosome 9p21 predict
severity of coronary artery disease. J. Am. Coll. Cardiol. 57: 1497?1498.
Chan K., Patel R.S., Newcombe P., Nelson C.P., Qasim A., Epstein S.E., Burnett S., Vaccarino
V.L., Zafari A.M., Shah S.H., Anderson J.L., Carlquist J.F., Hartiala J., Allayee H., Hinohara K.,
Lee B.S., Erl A., Ellis K. L., Goel A., Schaefer A.S., Mokhtari N.E.E., Goldstein B.A., Hlatky
M.A., Go A.S., Shen G.Q., Gong Y., Pepine C., Laxton R.C., Whittaker J.C., Tang W.W.H.,
Johnson J.A., Wang Q.K., Assimes T.L., Nцthlings U., Farrall M., Watkins H., Richards A.M.,
Cameron V.A., Muendlein A., Drexel H., Koch W., Park J. E., Kimura A., Shen W., Simpson I.A.,
Hazen S.L., Horne B.D., Hauser E.R., Quyyumi A.A., Reilly M.P., Samani N.J., Ye S. (2013)
Association between the chromosome 9p21 locus and angiographic coronary artery disease
burden. A collaborative meta-analysis. J. Am. Coll. Cardiol. 61(4): 957-970.
Dandona S., Stewart A.F.R., Chen L., Williams K., So D., O?Brien E., Glover C., LeMay M.,
Assogba O., Vo L., Wang Y. Q., Labinaz M., Wells G.A., McPherson R., Roberts R. (2010) Gene
dosage of the common variant 9p21 predicts severity of coronary artery disease. J. Am. Coll.
Cardiol. 56: 479-486.
Deloukas P., Kanoni S., Willenborg C. et al. (2012) Large-scale association analysis identifies new
risk loci for coronary artery disease. Nature Genetics 45(1): 25-33.
Do R., Xie C., Zhang X. et al. (2011) The Effect of chromosome 9p21 variants on cardiovascular
disease may be modified by dietary intake: evidence from a case/control and a prospective study.
PlosMed 8(10): e1001106.
Hinohara K., Nakajima T., Takahashi M., Hohda S., Sasaoka T., Nakahara K., Chida K., Sawabe
M., Arimura T., Sato A., Lee B.S., Ban J.M., Yasunami M., Park J.E., Izumi T., Kimura A. (2008)
Replication of the association between a chromosome 9p21 polymorphism and coronary artery
disease in Japanese and Korean populations. J Hum Genet 53: 357-359.
Montorsi P., Ravagnani P.M., Galli S., Rotatori F., Veglia F., Briganti A., Salonia A., Deho F.,
Rigatti P., Montorsi F., Fiorentini C. (2006) Association between erectile dysfunction and coronary
artery disease. Role of coronary clinical presentation and extent of coronary vessels involvement:
the COBRA trial. Eur. Heart J. 27: 2632?2639.
Patel R.S., Su S., Neeland I.J., Ahuja A., Veledar E., Zhao J., Helgadottir A., Holm H., Gulcher
J.R., Stefansson K., Waddy S., Vaccarino V., Zafari A.M., Quyyumi A.A. (2010) The chromosome
9p21 risk locus is associated with angiographic and progression of coronary artery disease. Eur
Heart J. 31: 3017-3023.
Prins B.P., Lagou V., Asselbergs F.W., Fu J. (2012) Genetics of coronary artery disease: genomewide association studies and beyond. Atherosclerosis 225(1): 1-10.
# 183 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ????????? ????? 9?21.3 ? ???????????? ????????? ??????? ??????????? ??????????????
15. Schunkert H., Erdmann J., Samani N.J. (2010) Genetics of myocardial infarction: a progress
report. Eur. Heart J. 31(8): 918-925.
16. Shen G.Q., Li L., Rao S., Abdullah K.G., Ban J.M., Lee B.S., Park J.E., Wang Q.K. (2008) Four
SNPs on chromosome 9p21 in a South Korean population implicate a genetic locus that confers
high cross-race risk for development of coronary artery disease. Arterioscler Thromb Vasc Biol
28: 360-365.
17. Sianos G., Morel M.A., Kappetein A.P., Morice M.C., Colombo A., Dawkins K., Van den Brand M.,
Van Dyck N., Russell M.E., Mohr F.W., Serruys P.W. (2005) The SYNTAX score: an angiographic
tool grading the complexity of coronary artery disease. EuroIntervention 1: 219?227.
Locus 9p21.3 ? Genetic Predictor
of Coronary Atherosclerosis Severity in Siberians
Pavel A. Shesternya, Anna S. Sergeeva,
Svetlana Yu. Nikulina and Vladimir A. Shulman
Krasnoyarsk State Medical University
named after ?rof. V.F. Voino-Yasenetsky,
1 Partizan Zheleznyak Str., Krasnoyarsk, 660022 Russia
The purpose of this study was to investigate association between 9p21.3 locus single nucleotide
polymorphisms (SNP) and severity of coronary atheromatous burden. A total of 255 Caucasians (211
male, 44 female) with myocardial infarction younger 65 years old (mean 52,56±7,98 years) were enrolled.
Genome DNA was extracted from venous blood using the phenol-chloroform extraction method. Two
SNPs rs10757278 and rs1333049 (locus 9p21.3) were tested using real-time polymerase chain reaction
(PCR) according to protocol (probes TaqMan, Applied Biosystems, 7900HT). The coronary angiograms
were reviewed by independent angiographers who were blinded to the results of the genotype (Philips
Allura Xper FD10). The total number of lesions, Gensini scores and SYNTAX score were derived. For
the first time in males of Siberian population, genotype CC rs1333049 and genotype GG rs10757278
demonstrated a direct strong association with severity of coronary atheromatous burden.
Keywords: myocardial infarction, single nucleotide polymorphisms, rs10757278, rs1333049, locus
9p21.3, atherosclerosis.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2013 6) 185-195
~~~
??? 57.045
???????????? ??????? ????????????? (???????????)
?????????? ???????:
????????????????????? ??????
?.?. ????????*, ?.?. ????????,
?.?. ???????, ?.?. ???????
?
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ?????????, 79
?
??? «???????? ?????????? ??????»,
?????? 628310, ????????? ???., ?????-?????????? ?? ?
????, ???????????, ??. ????, 1
Received 18.07.2012, received in revised form 05.10.2012, accepted 22.04.2013
?????? ????????? ???????? ?????????? ?????????????????????? ?????? ??? ????????????
??????? ?????????? ? ??????????? ???????????????? ????????? ? ?????????? ???????.
??????????? ???????????? ???????, ????????? ? ??????????? ????????-????????? ??????????
?? ?????? ????????? ???????, ?? ???????????? ????????? ? ??????????? ????????. ??????????
?????????? ????????? ?????????? ??????? ?????????? (???????????) ??????????????
?????????: ????????????? ??????????? XV ?., ???????? ???????? ? ?????? ????????
??????????? XV ?., ????? XV ? ?????? XVI ????????, ??????-????????? ??????????? XVI ?.,
?????? ???????? XVII ?. ? ? ????????? ?????????? ????????-????????? ?????????? «?????»,
?????????? ??????????? ???????????? ????????? ???????? ????????? ?????????????? ?
??????? ? ?????? ??????? ?. ?????.
???????? ?????: ???????????????, ????????-????????? ??????????, ??????????????? ?????????,
??????, ????????? ???????.
????????
????????? ??????? ? ?????????? ??????????????????? ???????? ???????? ???????????? ????????? ?????????????? ????
??????-???????? ?????? (???. 1). ??????????
???????????? ???????????? ???????????????? ????????? ????? ??????? ??? ????????
*
???????????? ????????, ?????????, ?????????? ????????? ? ??????????? ??????? ???????????? ????????? ??????-???????? ??????
(??????, 2009). ?? ??? ??? ?????????????
???????? ??????????????? ??????, ?????????
?? ???????? ??? ????????? ? ????????????????. ? ?????????, ?????????? ?????????
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: gulzar03522@mail.ru
# 185 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ???????? ???????????? ??????? ????????????? (???????????) ?????????? ????????
???. 1. ????? ? ?????-????? ???????????? ?????????? ??????? ? ???????????? ?????????? «????????
????», ?????? ? ??? ?? ????????? ??????? ? ???-??????
????? ???????? ???? ????????? ???? ???????
??????? ??????? ? ?????? XVI?XVII ??., ?.?.
?????? ????????????? ???????? ?????? ?
???????? ??????????? (??????, 2009). ??????, ???????? ?????????? ??????????? ?????????? ??????????? ???????? ??????????
???????, ???????? ??????????? ??? ???????????? ???????????? ??????? ??????????????
?????????? ?????????????????????? ???????. ?????? ????? ????????? ????? ? ??????? ?????? ???? ??????????????? ????????
?? ????????????? (?????????) ??? ?????????? ????????? ????????? ?? ??????????????
?????? ??????? ???????????? ??????? ??
?????????? ?????????? ???????? ???????
(??????, 1980; ??????, 1996; ??????, ????????, 1997; ??????, ??????????, 2000; ??????? ? ??., 2002; ?????? ? ??., 2009; ?????? ?
??., 2010; Grabnera et al., 2007; Eckstein, 2007;
Saskia et al., 2009 ? ??.).
?????? ????????????????????? ???????????? ?????????? ??????? ???? ????????? ?.?. ????????? (??????? ? ??., 2002). ?????? ?? ????????? ???? ??? ??????? ?????
??????????????? ???????? ? ????? ??????
????? ?????????, ??????? ????????? ????????? ????? ???????????? ?.?. ?????????
??????? ????????????? ?????????. ?????
???????, ???? ??????? ???????????? ? ?????????? ???????????? ????????? ?????????? ???????.
????????? ? ??????
????????? ??????? ????????? ?? ??????
?????-????????? ??????????? ?????? ????????? ???????, ? ?????? ?. ?????, ? 60 ?? ?
??????-?????? ?? ?. ?????, ?? ???????. ??????
????????? ??? ??????? ???? ????????? (??
30 ?) ??????, ??????????? ??????? ????????? ???????? ???? ? ????? ????????? 80 ? 25
?. ???????????? ?????? ????????? ?????????? ?????? ? ?????????? ????? ??????????
4.5 ? (???. 1). ? ????????? ????? ?????????
??????? ???????????? ????? ?????????????
??? ????????? ???????????? ????????? ???????? ???????? ???????? ????? 650 ??. ?.
(??????, 2009).
??????? ??? ??????????????????????
??????? ????????? 78 ???????? ??????????????? ?????????, ?????????? ? 2011 ?.
# 186 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ???????? ???????????? ??????? ????????????? (???????????) ?????????? ????????
? ???????? ? 2 (?????? ???????? 5.30-5.70;
4.90-5.00; 4.98-4.72) ? ? 12. ????? ??????
???????? ????????? ???, ??? ?? ????? ??????
????????????? ???? ????????? ?????? ???
??????????? ???????? ??????????????? ???????????? ??????? ????????????? ? ??????????? ???????. ?? ?????? ?????? ????????
????????? ??? ???? ?????????, ??????????
????, ?? ??????? ?????, ?????? ???? ??????????, ? ????? ????????? ?????????? ????????? ?????????? ???????. ??? ??????????
??????? ???????? ?????? ???????? ?????????
?????? ????????, ??????????? ???????????
????????? ?????????? ???????????? ?????.
? ?????? ??????? ???? ? ????? ???????? ?????
????????? ?? ????? ???????? ???????????, ?
?????????? ?????? ??????? ??????????? ?
?????? ??????????? ????? ??????????? ??????. ????????, ??? ? ????????? ????? ?????
? ???????? ??????? ?????? ?? ????????????, ??? ???????? ???????? ???????????????
????????? ? ??????????? ????? ??????? ??
????? ???????? ?????? ?. ?????, ?? 40 ??
???? ?? ??????? ?? ???????????? ???????,
???? ???????? ??? ??????? ???????, ???
???? ???????? ????? ??????????? ????????? (Larix sibirica Ledeb.). ?????? ??????
????????? ????????? ???????? ??????????
??????????.
????????? ?????? ???????? ?????
???? ????????? ?? ??????????????????
????????? «LINTAB» (? ????????? 0,01 ??).
????????? ?????????? ????? ???? ???????????? ??????????? ????????? ???????????
???????????? ????????? (Douglass, 1919) ?
?????-??????????????? ??????? ? ?????? ?????????????????? ???????? ??? ????????????????????? ???????????? ? DPL (Holmes,
1984) ? «TSAP V3.5» (Rinn, 1996). ? ??????
?????????????? ????????-????????? ?????????? ???? ????????? ???????? ? 2/3 ?????
?????? ????? ? ????????? ARSTAN (Cook,
Kairiukstis, 1990). ??? ??????????? ????????
?????????? ????????-????????? ??????????
?? ?????????? ??????? ???? ????????????
?????????? «???????? ????», ??????????? ?.
????????????? ?? ??????????? ?????????, ?
778 ?? 1892 ??. (F. Schweingruber, International
Tree Ring Data Bank (ITRDB) http://www1.ncdc.
noaa.gov/pub/data/paleo/treering/chronologies/
asia/ russ176w_crns.crn).
??????????
?????????? ????? ???????????? ????????? ???????????? ??????? ???? ????????????:
?????????? ??????????? ????????-?????????
?????????? ?? ?????? ???????????? ? «?????????» ????????-????????? ????, ??????????? ?? ??????????????? ?????????. ? ???? ????? ?? ?????????? ?????? ???????????????
????????? ???? ????????? ? ????????????????? ????????-????????? ?????????? ?? ?????? ????????? ??????: ????? Pinus sibirica Du
Tour ? «????», ??????????? L. sibirica ? «???????????» ? ??? Picea obovata Ledeb.? «???».
??????????? ???????????? ?????? ???????,
??? ??????? ????????-????????? ?????????? ?? ???????????, ????? ? ??? ????? ??????
????? ?????, ??? ????????? ?????????? ??
? ?????????? ?????????? «?????», ???????????? ?????? ? 1010 ?? 1616 ??. (???. 2).
?????? ???????? ????????????? ??????????
????? ?????????? ????????-????????? ??????????? «?????» ? ????????????????????
???????????? ?? ?????? ????????? ??????
??????, ??? ?? ???? ?????????? ? ?????????
??????????? ?????? ??? ?????? ???, ????????? ? ?????????? ??????? ????????????? ?
?????????????????? ????????? ?????? ????????? ????? ?? ????????? ??????? ???????
?????. ????? ???????, ???????? ????? ??????, ??????????????? ??????? ?????????
?????????????? ?? ????????? ??????????
????? ? ???????? ?. ????? (????. 1).
# 187 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ???????? ???????????? ??????? ????????????? (???????????) ?????????? ????????
???. 2. ??????????? ???????????? ????????? ??????????????? ?????????? ?? ????????? ??????? («???»,
«???????????», «????») ? ?? ?? ?????? ?????????? ?????????? ?????????? «?????» ?? ??????????
???????
??????? 1. ???????????? ?????????? ????? ???????????????????? ????????-??????????
???????????? «?????», «????», «???????????», «???», «???????? ????» (?????? ? 1250 ?? 1450 ??.,
???????????????? ??? ????????????? ????????-????????? ?????). ??? ???????? ?????????? ??????????
??? p < 0.05
«????»
«???????????»
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??? ???????? ? ??????????? ????? ??????? ?????????? «?????» ???? ????????????
????????-????????? ?????????? «????????
????», ????????????? ? 300 ?? ? ?????? ??
?????????? ???????. ??????????? ??????????? ???????? ???????????? ????????? ????????? ?? ???. 3. ????? ???????, ? ??????????
??????????? ???????????? ????????? ????
???????????? ???????? ? ??????????? ?????
??????? ?????????????? ????? ?? ?????????? ? 2 ? 12 (????. 2, 3).
????????? 2. ?? ?????????? 55 ????????
?????????? 48, ?? ??????? ????????? ?????
«?????»
0.41
??? ?????????????? ?????????? ??????????:
??? ? 17, ???? ? 16, ??????????? ? 15 (????. 2).
??????? ???????? 5.30?5.70. ? ????? ????????? ???? ???????? 15 ???????? ??????????????? ?????????, ?? ??????? ???????????? 11 (????. 2). ??????? ????????????
?????????? ?????????? ????????: ??? ? 5,
???? ? 5, ??????????? ? 1.
?????? ???????????? ???????? ???????? ???????? ??? ??????? ?????????????
(????. 2). ?????? ?????? ???????????? ????????? ng070 ? ng071, ? ??????? ???? ???????????? ??????????? ?????? ?????????? ??
# 188 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
???. 3. ???????????? ????????? ????????-????????? ?????????? «?????» (?????? ?????) ? «????????
????» (??????? ?????)
??????? 2. ?????????? ???????????? ????????? ???????? ????????? ? 2 ?????????? ???????: r ?
????????????? ??????????? ?????????? ??? ??????????????????? ????? ????????; m ? ???????????
????????????????; SD ? ??????????? ??????????; ??????? ?????? ???????? ???????, ??????? ??????????
?? ?????? ? 1464 ?? 1470 ??.; ??????? ?????? ? ? ????? XV ?. ?? ?????? XVI ?.; ????? ?????? ? ??????
??????????? XVI ?.; ?????? ?????? ? ?????? ???????? XVII ?.
??????? ???????? 5.70 ? 5.30
?
1
??????? ?????. ?????.
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3
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6
7
8
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ng144
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42
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???
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??????? ???????? 4.72 ? 4.98 (?? ?????)
43
ng131
-
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1271
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1256
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0.54
0.28
0.29
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1355
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0.88
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48
ng128
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??????????? ?????????
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49
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0.45
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????????? ????. 2
50
ng120
1297
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0.31
0.37
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1283
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0.48
?????.
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1360
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0.26
0.28
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1483
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1422
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0.31
0.33
?????.
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??????????: ?????. ? ?????? ????????????? ??????; ?????. ? ????????? ????????????? ??????.
??????? 3. ?????????? ???????????? ????????? ???????? ????????? ? 12 ?????????? ???????:
r ? ????????????? ??????????? ?????????? ??? ??????????????????? ????? ????????; m ? ???????????
????????????????; SD ? ??????????? ??????????; ????? ?????? ???????? ???????, ??????? ??????????
?? ??????-?????? ??????????? XV ?.; ??????? ?????? ? ? 1464 ?? 1470 ??. XV ?.
?
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?????????? ???? ???????????, ?????
8
ng329
1181
1422
0.47
0.21
0.42
???
?????????? ????
9
ng327
1320
1423
0.55
0.35
0.38
???
?????????? ???? ???????????
10
ng328
1204
1426
0.57
0.30
0.37
???
11
ng326
1273
1429
0.63
0.27
0.36
?????.
?????????? ????, ?????
??????????? ?????????
?????????? ???? ???????????, ?????
12
ng324
1187
1431
0.58
0.52
0.48
?????.
?????????? ???? ???????????, ?????
13
ng323
1274
1432
0.57
0.35
0.41
?????.
?????????? ???? ???????????, ?????
14
ng320
1275
1435
0.54
0.26
0.39
?????.
?????????? ???? ???????????
15
ng325
1237
1435
0.58
0.34
0.44
?????.
?????????? ???? ???????????, ?????
16
ng319
1194
1437
0.50
0.32
0.40
?????.
???????? ?????????
17
ng316
1014
1447
0.49
0.21
0.32
????
?????????? ???? ???????????, ?????
18
ng331
1084
1448
0.58
0.23
0.39
????
?????????? ????
19
ng318
1093
1453
0.46
0.24
0.39
????
???????? ?????????
20
ng312
1090
1455
0.62
0.21
0.29
????
?????????? ???? ???????????, ?????
21
ng315
1120
1456
0.48
0.25
0.35
???
?????????? ???? ???????????, ?????
22
ng317
1122
1459
0.49
0.24
0.44
????
?????????? ???? ???????????, ?????
23
ng313
1098
1464
0.54
0.20
0.29
????
?????????? ????, ?????
??????????: ?????. ? ?????? ????????????? ??????; ?????. ? ????????? ????????????? ??????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ???????? ???????????? ??????? ????????????? (???????????) ?????????? ????????
1464 ? 1470 ??. ???? ?? ????? ??????? ??????? ng085b, ng086b, ng078, ? ??????? ??????????? ????? ??????? ?????, ? ?????????
???????????? ?????? ????????? ? ?????????
? 1464 ?? 1470 ??. ????? ???????, ?????? ??
?????????? ?????? ????? ????????????,
??? ????? ??????????????? ?????????????
???????? ?? ???????? ???????? ? ??????
???????? ??????????? XV ?. ?????? ?????? ????????? ??????? ng082, ng077, ng062,
ng064, ????? ??????????? ???????????? ????? ? ??????? ?????????? ?? ?????? ??????????? XVI ?. ? ???? ?? ?????? ????? ???????
??????? ng074 (1505 ?.), ? ???????? ?? ???????
??????? ??????? ?????????????? ????? ?????, ??????????? ?? ?????????? ????? ?????
?????????? ?????????? ??????? ????? ?????.
????????, ??? ?? ? ?????? ??????? ?? ????
?????? ?? ??????????? ?????????? ??????,
????? ??????????, ??? ??????????? ???????????? ??????? ???? ????????? ?? ?????
1519 ?., ? ?????? ?????, ? ??????? ? ?????????
??????????? XVI ?. ?? ??????? ???????? ??????? ??????????? ????????? ??????? ng064,
????? ???????????? ?????????? ????????????? ?????? ???????? ????????? ???????? ?
???, ??? ??????????? ?????????????? ?? ????? 1616 ?., ? ?????? ?????, ?? ?????? ? ???????
??????????? XVII ?.
??????? ???????? 4.90-5.00. ?? ????
?????? ???????? ???? ???????? 27 ????????, ?? ??????? ???????????? 25 (????. 2), ??
??????? ????????? ????? ??? ?????????????? ????????? ???????: ??? ? 7, ???? ? 10,
??????????? ? 8. ?????? ?????? ???????
???????? ????????? ???????? ??? ???????
???????????. ? ?????? ?????? ????? ??????? ??????? ng148, ng149, ng151, ng153, ng154,
ng156, ng167, ? ??????? ???? ????????????
???????????? ????? ?????????? ?? 1445 ?
1469 ??. ?????? ?? ?????????? ? ???????? ?????????? ?????, ????? ????????????, ???
?????? ??????? ??????????? ? ??????????
??? ?????? ???????? 5.30 ? 5.70 ?????? ??????? ? ????? ?????????? ?? ???????? ???????? ? ?????? ???????? ??????????? XV ?. ?????? ?????? ???????????? ????????? ng144,
ng150, ng155, ng157, ng159, ng160, ng161, ng164,
ng165, ???????????? ?????????? ????, ????
???????????? ???????? ????????? ? ????????
?????????? ????????? 1490 ? 1506 ??. ???? ??
????? ??????? ??????? ng145, ng146, ng166,
ng168, ng170, ? ??????? ??????????? ????? ???????????? ?????. ? ???? ?????? ????? ????????, ??? ????? ???? ??????????? ? ?????
XV ? ?????? XVI ????????.
??????? ???????? 4.98-4.72. ? ?????????????? ????? ?????? ????????? ???? ?????
13 ????????, ?? ??? ???????????? 12 (????. 2),
?? ??????? ????????? ????? ??? ?????????????? ????????? ???????: ??? ? 5, ????
?1, ??????????? ? 6. ? ????? ?????? ?????
??????? ??????? ng120, ng123, ng124, ng125,
ng128, ng129, ng130 ? ng127, ? ??????? ?????????? ?????????? ???? ? ????? ???????????? 1465 ?. ? ???? ?????? ????? ????????,
??? ?? ???? ???? ??????? ???????? ?????
??????????????? ?????? ???????????, ???????????? ?? ???????? ???????? ? ?????? ???????? ??????????? XV ?. ??????? ng119, ng121,
ng122, ? ??????? ?? ?????????? ??????????
????, ??????????, ??? ????? 1484 ?., ????????,
???? ??? ???? ???????????. ???? ???????????
? ???????, ??????????? ??? ?????? ???????? 4.90-5.00, ?? ??????? ????????????, ???
? ????? ??????? ?????????? ???? ???????????, ???????????? ?? ????? XV ? ?????? XVI
????????.
????? ???????, ?????? ?????? ?? ????????? 2 ???????? ???????? ?????????
??????? ????????????? (???????????): ???????? ???????? ? ?????? ???????? ???????????
XV ?., ????? XV ? ?????? XVI ?., ?????? ? ????????? ??????????? XVI ?., ?????? ? ??????
# 192 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ???????? ???????????? ??????? ????????????? (???????????) ?????????? ????????
??????????? XVII ?. ???????, ??? ??????,
?????????? ??? ?????? ??????? ????????,
?? ???????????? ? ????????? ???? ?????.
????????? 12. ? ?????????? ??????????? ???? ????? 23 ???????, ?? ??????? ??????????? 21 ??????? (????. 3). ????????? ?????? ???? ???????????? ????????? ???????:
??? ? 4, ???? ? 9, ??????????? ? 8.
?????? ??????? ???????? ???????? ???????? ??? ??????? ????????????? ???????.
? ??????? ????? ??????? ??????? ng329
? ng328, ???? ???????????? ???????????
?????? ??????? ?????????? ?? 1422, 1426 ??.
???????? ? ????? ?? ?????????? ????????? ????????? ??????? ng310, ng314, ng321,
ng322, ng327, ng332, ? ??????? ???????????
????? ???????? ?????. ? ???? ?????? ????? ????????????, ??? ????? ??????????
????????? ? 12 ???????? ?? ?????? ??????????? XV ?. ?? ??????? ??????? ????? ??????? ??????? ng313 (? ???????? ??????????
?????????? ????), ng312, ng315, ng317, ng318,
? ??????? ??????????? ????? ????????????
?????. ? ??? ?? ?????? ????? ???????? ??????? ng324 ? ng326, ?? ??????? ??????????? ??????? ??????????? ????? ???????
??????????, ??-?? ???? ??????????? ???????????? ??????. ??? ? ? ????????? 2, ????
?????? ??????????? ???????? ?? ???????
??????????? XV ?.
????? ???????, ???????? ??????, ?????????? ?? ?????????? ? 2 ? ? 12, ??????? ?????????? ? ??????????? ??????????
??????? ?????????? ?? ?????? ???????????
XV ?., ???????? ???????? ? ?????? ????????
??????????? XV ?., ????? XV ? ?????? XVI
????????, ??????-????????? ???????????
XVI ?. ? ?????? ???????? XVII ?.
??????????
????????????? ????????? ????????
?????????? ??????? ???????? ????? ?? ???-
??? ???????? ????????????. ????? ? ????????? ???????? ??????? ?????? (????, ???,
???????????) ?? ????????????, ? ??????????, ??? ?????????? ????????????? ????????? ????????? ?????? ???????, ????????????? ????? ?? ??????? ?. ?????, ????????
?? 15-20 ??. ? ????????? ?????? ??????
??????????, ? ????? ???? ??? ????????? ?????????????? ????????????? ?????????, ?
? ????? ? ???????, ??????????? ????????
????. ???? ?????????? ????????? ????????
?? ????? ????????, ? ?????? ????????????
? ????????? ??????????, ?????? ?? ????
??? ?????, ????? ????? ???????? ? ???????????????? ???????? ????????? ?????????
??? ?????????????. ?????????????? ?????????? ? ?????? ????????????? ? ?????????
????????? ?? ????????? ??????? (????????
? ??????? 30 ??) ???????? ??? ????, ??? ????
??????? ????????? ? ???? ?????? ????????? ???????, ??? ?????? ??????????? ?????????? ????? ?????, ?.?. ??????? ????????
????????? ?????????? ?? ????????? ??????? ???????.
??? ??????????? ???????????? ???????? ??????? ??? ?????????? ?????????? ????? ???????? ????????? ???, ?????
??? ????????? ???????????? ??????????
??????? ?????. ??? ????????? ??????????? ???? ?? ???????? ? ????? ????????????
????????? ???????????? ????? ? ??????????? ????????? ?????? ?? ?????????
???????? ????????????? (???????????)
?????????. ???, ????????, ?? 78 ?????????? ??? ???????????? ???????? ???? ????
?????????? 85,5 %, ? ?? ????? ??? ? ?????
?????????????? ??????? (??????, 2009) ??
1350 ???????? ???????????? ???? 340, ???
????????? 25,2 %.
?????????? ??????? ???????????? ????????????? ?????????? ???? ????????
????????????? (???????????) ??????????
# 193 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ???????? ???????????? ??????? ????????????? (???????????) ?????????? ????????
??????? ? ???????, ??????????? ????? ?.?.
????????? ? ??. (2002). ????????? ??????????? ????????, ??? ???????????? ????? ?????
????????? ??????? ???????????? ????? ???
?? ??????? ???? ? ?? ????????????? ??????????? XV ?. ? ???????????? ?.?. ????????
? ????????? (2002) ??????????? ??????????
???? ??????? ???????? ???????? ? ??????
???????? ??????????? XV ?., ????? XV ? ?????? XVI ????????, ???, ????????, ??????? ?
??????????? ? ??? ????????? ???????? ?? ????? ?????? ??????????. ???????? ?????????
??????????? ??????? ? ???????? 1590?1595
??., ??????? ?? ?????????????? ?? ???????????? ???? ??????????. ????????? ????? ??????????? ????????? (1620-1630 ??.)
?????????, ? ????? ?????????? ??? ???????
(1650-1655, 1690-1695 ??.), ??????? ?? ?????????????? ? ????? ????????????. ?????????????? ???? ??????????? ?????????? ??
1725-1730 ??. ? ???????? ????? ??? ????? ????????????.
??????????
?????????? ?????? ????????, ??? ? ????????????? ???????? ?????? ???????? ?????? ????????-????????? ??????????, ??????????? ?? ?????? ????????? ???????
? ?????? ???????????????? ?????????, ?????????? ?????? ??????????? ? ??????????
?????????? ???? ? ??????. ?????????? ??
?? ?????? ?????????? ????????-?????????
?????????? «?????» ????????? ???????????
??????????? ???????? ??????????????? ????????? ?? ????????-????????? ??????????
«???????? ????». ? ?????????? ???? ???????? ???????????? ?????????? ? ???????????
?????????? ???????, ??? ????????? ??????????? ??? ?????????? ?????????? ?????????
?????????????????????? ????????? ? ??????? ???????. ??????????? ????????-?????????
?????????? «?????» ????? ???? ???????????
???????? ???????????? ??????? ????????
?????? ????????? ????? ?? ????? ????????????? ??????.
?????? ??????????
1.
2.
3.
4.
5.
6.
7.
8.
??????? ?.?., ???????? ?.?., ?????? ?.?. (2002) ?????????? «?????????? ????????» ?
??????? ????????-?????????? ???????. ?????????? ? ???????????? ??????????????? ?????????? ???????? ???????? ?????? ? ???????????? ??????????. ??????: ???-?? ????????? ??????? ???????? ?????? ?? ???, ?. 22?24.
?????? ?.?. (2009) ????????? ??????? ? ????? XVI ? ?????? ????? XVIII ??. ??????? ?
???????????? ????????. ????????????-???????????: ????????, 360 ?.
?????? ?.?., ??????? ?.?., ????????? ?.?. (2010) ????????: ????????-?????????????
??????. ??????????: ???-?? ???, 152 ?.
?????? ?.?., ?????????? ?.?., ????????? ?.?. (2009) ???????? ??????? ?? ??????????: ????????????????????? ??????. ??????????, ?????????? ? ???????????? ??????? 3: 103?110.
?????? ?.?. (1996) ???????????????? ? ??????????. ?.: NOX, 216 c.
?????? ?.?., ???????? ?.?. (1997) ???????????????? ????????????? ?????????? ?? ?????? ?????? (??????? ???????? ????? ? ?????). ?????????? ?????????? 1: 109?124.
?????? ?.?. (1980) ????????? ?????????? ?????????? ???????? ?????????????????????
???????. ?: ???????????? ??????? ???????? ???. ?. 1. ?.: ???????????????, ?. 93?107.
?????? ?.?., ?????????? ?.?. (2000) ????????????????????? ????????? ????????? ??????????? ?? ???????????????? ????????? ???? VI ?? ??????????? ????. ?: ?????????
?????. ???. 1. ????????????-????????: ??? ???, ?. 112?121.
# 194 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ???????? ???????????? ??????? ????????????? (???????????) ?????????? ????????
9.
10.
11.
12.
13.
14.
15.
Cook E.R., Kairiukstis L. (1990) Methods of dendrochronology: applications in environmental
sciences. Dordrecht; Boston; L.: Kluwer Acad. Publ., 394 p.
Douglass A.E. (1919) A study of the annual rings of trees in relation to climate and solar activity.
In: Climatic cycles and tree-growth, v. 1. Washington: Carnegie Inst., p.127.
Eckstein D. (2007) Human time in tree rings. Dendrochronologia 24: 53?60.
Grabnera M., Kleina A., Geihofera D., Reschreiterb H., Barthb F.E., Sormazc T., Wimmera V.
(2007) Bronze age dating of timber from the salt-mine at Hallstatt, Austria. Dendrochronologia
24: 61?68.
Holmes R.L. (1983) Computer-assisted quality control in tree-ring dating and measurement. TreeRing Bulletin 43: 69?75.
Rinn F. (1996) TSAP V3.5. Computer program for tree-ring analysis and presentation. Frank Rinn
Distribution, Heidelberg, 269 p.
Van de Gevel S.L., Hart J.L., Grissino-Mayer H.D., Robinson K.W. (2009) Tree-ring dating of oldgrowth longleaf pine (Pinus palustris Mill.) logs from an exposed timber crib dam, Hope Mills,
North Carolina, USA. Tree-Ring Research 65(1): 69?80.
Periods of Construction (Reconstruction)
of Nadymsky Gorodok: Dendrochronological Aspect
Gulzar T. Omurovaa, Valentin V. Barinova,
Oleg V. Kardashb and Vladimir S. Myglana
a
Siberian Federal University,
Russia 660041, Krasnoyarsk, Svobodny, 79
b
A?NO ?Institute of Archeology of the North?,
Russia 628310, Tyumen region, Hanty ?
Mansiisk autonomous area ?
Ugra, Nefteugansk s., st. Mira,1
This work is devoted to the application of dendrochronological methods to establish the time of
construction and reconstruction of the archaeological monument named Nadymsky gorodok where
located 60 km to the north-north-west from the town of Nadym, Yamal-Nenets autonomous area,
Tyumen region. We discussed aspects related to the building of tree-ring chronologies of different tree
species and their cross-dating and relation with calendar periods. The next periods of construction
(reconstruction) of Nadymsky gorodok were distinguished: the third decade of the XV century, middle
of seventh ? beginning of eighth decade of the XV century, end of XV ? beginning of XVI century, the
third or fourth decade of the XVI century, the first quarter of the XVII century. Generalized tree-ring
chronology of Nadymsky gorodok was built and covered the period from 1010 to 1616 AD.
Keywords: dendroindication, tree ? ring chronology, archeological wood, Siberia, Nadymsky
gorodok.
# 195 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2013 6) 196-204
~~~
??? 504.45.054-034
???????????? ?????????????? ????
Bidens (Asteraceae) ? ???????? ???????? ?????
?? ????????? ?????? ??????????
?.?. ???????*
???????? ???????? ?????????? ???
??. ?.?. ???????? ?????????? ???????? ????,
?????? 152742, ??????????? ???????, ???. ?????
Received 07.08.2012, received in revised form 13.09.2012, accepted 22.04.2013
??????? ??????? ???????? ????? (0.1 ? 50 ??/?) ?? ??????????? ????? ? ????????? ?????
???????? ?????????? ???? ????? ???? Bidens L. ???????? ???????? ????? ?? ???? ?????????
????????? ???????????? ?? ??????? ????????????? ??????? ?? ???????????? ?????????
????? B. tripartita L. ??? 50 ??/? ???????? ??????????? ???????? ???????????? ?????????
????? B. frondosa L. ???? (50 ??/?) ?????? ??????????? ?????? ?? ???? ? ???????? ????????
??????? ?????????? ???? ????????????? ?????, ? ?????? ? B. cernua ? B. frondosa ????????????
??????? ??????, ? B. tripartita ?????????? ????????? ??? ?????. ????????? ????? ??????????
?????? ??????????? ?? ???????? ?????????? ???????????? ????? ? ?????. ???????????
???? ????????????? ? ???? ? ??????? ?????????? ???????????? ? ???????? ???????? ?????:
B. tripartita > B. frondosa > B. cernua.
???????? ?????: ??????? ?????, ??????????? ?????, ???????? ??????????, Bidens tripartita,
B. cernua, B. frondosa.
????????
??????? ??????? (??) ????????????
????????? ?????? ??? ???????? ? ????????
?????????? ?? ?????? ??????????? ? ???????????? ?????????? ? ?????????? ????? ??
??????? ???????, ????????? ?? ??????????
???????????? ??????? ? ??? ??????????????
????? (?????????, 2006). ??????????? ????? ????????????? ?? ? ????????? ?????? ?
???????????? ????????? ????????????. ??-
*
?????? ? ?????????? ?????????, ??? ?????
???????????? ????????? ???????????, ???
?? ???????????? ?? ?????? ????? ??? ????????? ???.
???? ????????? ? ????? ???????????
?????????????? (????????????? ?????????),
???????? ?? ???? ? ?????????? ???????? ????????, ?????? ? ?????? ???????? ??????? ?????? ????????? ? ????????? ?????????????
??????????????? ??????? (?????????????,
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: panova@ibiw.yaroslavl.ru
# 196 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????????????? ???? Bidens (Asteraceae) ? ???????? ???????? ????? ?? ??????????
1991). ?????? ? ???? ??????? ????????????
????? ????????? ???? ? ???????? ????????,
?????????? ?????????? ???????? ????????? ? ????????? ???????, ???????? ??????
??????? ??????? ? ???????? ??????????,
???????, ????????????, ??????? ????????
???? ? ?????? (????? ? ??., 1985; ????????
? ??., 2005; Khudsar et al., 2004; Wang et al.,
2009). ????????????? ?????????? ? ??????????????????? ??????????? ????? ???????,
??????????? ?????????, ?? ?? ????????????
(????????, 1988).
??????????? ????? ??? ????? ? ?????????? ?????????? ?? ???????. ???????
????????? ??????, ?????????? ??????????, ?????? ?????? ?? ??????????? ???????
???????????? ??????? (???????, ????????,
2009). ? ????? ? ??????????????? ???????
????? ?????? ???? ?????? ???????? ????????? ???????????? ???????? ????? ?? ??????????? ?????, ???? ? ???????? ??????????
???? ????? ???? Bidens L.: ?????? ?????????????? (Bidens tripartita L.), ?????? ???????? (B. cernua L.) ? ?????? ???????????
(B. frondosa L.). B. tripartita ? B. cernua ? ??????????? ??? ??????? ?????? ???????????
????? ?????? ?????????, ????????????? ?? ??????? ?????, ? ???????-??????? ?
?????????-?????? ???????????. B. frondosa ? ?????????? ???, ???????? ?? ???????
???????? ? ??????????? ??????????. B. frondosa ? ????????? ????? ?????? ???????????
? ????????? ?? ????????? ????????? B. tripartita ? B. cernua (???????????, 2003; ??????????? ? ??., 2010).
????????? ? ??????
?????? B. tripartita ? B. cernua ????????
? ??????????? ??????? ? ???????????? ????
??????, B. frondosa ? ? ???????????? ?. ????????, ? ???????? 2009 ?. ????? ????????
??????? ????????????? ??? ???????????
4?8 °? ? ??????? 4?5 ??????? ?????? ??????????? ? ??????????? ? ?????? ????? ???
??????????? 20?25 °? ?? ?????????????? ??????, ????????? ?????????? ZnSO4· 7H2O, ?
?????? 15 ?? ?? ?????? ?????. ???????????
???????????? ???? 0.1, 1, 10, 25, 50 ??/? ??
??????? ?? ??? ????? Zn2+. ???????? ????????
?? ???????????????? ????. ???????????? ???
???????? ????????: ???????????????? ????
? ?????????? ????????????? ????. ?????
????????? ? ???? ???????????? ??? ???????????? 3200 ????, ??????????? 9 ? / 15 ? (???? :
???????), ????????????????? ?????? 15 ???.
?????????? ????????? ??????????: ???????????? ????????? ? ???? ????????? ????? ?
????? ???????????? ?? ?? ?????? ????? ? %;
???-????? ? ????? (???) ????? ??????? ???????????? ? ??????? ???????????; ?????? ??????????? (???), ? ????? ????????? ?????????? ?? ????????? ??????????. ?? ?????????
????? ? 10 ?????????? ?? ??????? ????????
???????? ????? ???????? ?????, ??????????,
?????????, 1-?? ?????????? ?????, ??????
?????????, ? ????? ????? ??????????? ??????.
?????? ???????????? ? ??????????????
???????? Excel, Statistica (StatSoft Inc., Tulsa,
USA).
?????????? ? ??????????
??????? ???????? ?????
?? ??????????? ?????
????????????? ??????????? ????????
???????????? ????????? ??????????? ?????
? ???????????????? ? ?????????? ????????????? ???? ?? ??????????. ???? ??????????
??? ??????? ? ???? ????????????? ?????
(????. 1). ?????? B. tripartita ? B. cernua ???????? ?????????? ?? 2-? ????? ?????, B. frondosa ? ?? 3-? ?????, ?????? ??????????? ???
???? ????? ???????? 7?8 ????. ??????? ??????????? ????? ??????????????? ?? ?? ???-
# 197 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????????????? ???? Bidens (Asteraceae) ? ???????? ???????? ????? ?? ??????????
??????? 1. ???????????? ????????? ????? Bidens tripartita, B. cernua ? B. frondosa ??? ???????? ????????
?????, % ????????? ????? ? ????? ???????????? ?? ?? ?????? ?????
???
????????
???????????? ?????, ??/?
0.1
1
10
25
50
9.7±5.8
96.7±5.8
86.7±10.4
90.0±8.7
85.0±1.0
86.7±14.4
B. cernua
83.3±1.7
95.0±5.0
96.7±5.8
80.0±5.0
95.0±5.0
68.3±18.9
B. frondosa
96.6±3.3
88.3±7.6
93.0±8.7
96.7±2.9
93.3±7.6
68.3±16.1*
B. tripartita
* ??????????? ???????? ????? ??????????? ? ??????? ?????????? (t-???????? ?????????, p < 0.05).
??????? ?????????? ? ??????????? ? ???????????.
???????? ???????? ????? ?? ???????
????????????? ??????? ?? ????????????
????????? ????? B. tripartita, ?? ???? ????????? ????? ??????????? ???????? ?? ????? ?????????? ????????????? (????. 1). ???
????? B. cernua ???????? ??????????????
?????????, ? ??? ????? B. frondosa ??????????? ?????????? ????? ?????????? ???
50 ??/?.
?????? ?? ???????? ??????????? ?????????????, ??? ???????? ????? ????? ?????????? ?? 3?4-? ????? ??? B. cernua ? ?? 5?6-?
????? ??? B. tripartita ? B. frondosa (???.). ?
?????????, ?????????? ???? ?????, ??????
B. tripartita ? B. frondosa ?????????? ???????
?????????, ??? ? ????????. ??????? ????????
?????????? ??? ??????????? ???????? ???????????? 50 ??/? ? ?????????? ????. ?? ???????? ??????????? ????? B. cernua ???????
????? ?????? ????????? ???????: ???????????? 0.1 ? 1 ??/? ????????????? ???????????,
? ???????????? 50 ??/? ??????????? ????????
???? ???????.
???????? ?????????? ??? ????????????? ?????? ???????????? ????? ???? ???????? ?.?. ????????? ? ?.?. ?????????
(2009) ??? ?????????? ?????????????????
Batrachium trichophyllum (Chaix) Bosch., ? ????? ???????? ????????????? ????????????
????????? ? ?????? ???????????. ?.?. ???-
???? ? ?????????? (2009) ? ????????????
?????? ????????, ??? ???????????? ?????
10 -6 ?10 -3 ? ?? ?????? ?? ??????? ??????????? ? ????????? ????? ????????? ????????
Setaria viridis L.
??-????????, ???? ????? ?? ????????? ????? ???????? ????????, ???????????
???????? ????? ??????? ?????????? ????? ????????? ?? ??????????? ? ?????? ????????????? ???????, ??? ???????? ? ? ??????????
(Rengel, 2000).
????? ???????, ???????? ??????????? ?
???????? ????????? ???????????? ????????
????? ????????? ?????? B. tripartita. ??? ??
?????????, ???? ??? ????? ????? ???????? ?
???????? ????????? ?????? ? ???? (??????? ? ??., 2011). ???????? ?????? ?????????
????? B. tripartita ? ????????????? (???????????????) ???????? (??????????? ? ??.,
2008). ???? ????? ???????????? ??? ??????????????.
??????? ???????? ?????
?? ???????? ??????????
????????????? ??????????? ????????
???????????????? ??????????? ? ??????????? ?????????? ? ???????????????? ? ?????????? ????????????? ???? ?? ??????????. ?
???????? ?????????? ????? ???????? ?????
???? ???????? ? ?????????? B. frondosa, ?????????? ? B. cernua, ? ?? ????? ??? ?????
?????????? ???? ???????????? ? ???????-
# 198 #
0
20
40
60
80
100
0
0
3
2
3
4
5
5
??? ?????, ???
?
??? ?????, ???.
4
?
6
6
7
7
50 ??/?
25 ??/?
10 ??/?
1 ??/?
0.1 ??/?
????????
8
8
?????????, %
0
20
40
60
80
100
120
0
2
3
4
5
??? ?????, ???.
?
6
7
8
???. ???????? ??????????? ????? Bidens tripartita (?), B. cernua (?) ? B. frondosa (?), ?? ??? ??????? ? ?????????, % ????????? ????? ? ????? ???????????? ??
?? ?????? ?????
0
20
40
60
80
100
120
?????????, %
?????????, %
120
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????????????? ???? Bidens (Asteraceae) ? ???????? ???????? ????? ?? ??????????
??????? 2. ???????????????? ?????????? ?????????? Bidens tripartita, B. cernua ? B. frondosa,
?????????? ??? ????????? ????????????? ????? ????? ? ?????
??????????
????? ????????
?????, ??
??????????
???????????
??????, ??
???????????? ?????, ??/?
???
????????
0.1
1
10
25
50
Bt
47.6±5.2
33.2±12.8
42.8±8.4
34.2±25.2
18.7±11.23
3.1±0.8*
Bc
23.2±1.7
30.1±5.7
29.6±6.3
22.2±6.8
0*
0*
Bf
55.7±5.6
45.6±11.6
64.2±28.6
70.5±12.3
26.1±8.9
0*
Bt
4.8±0.5
4.5±1.5
4.3±1.9
5.6±2.5
7.0±2.4
4.5±1.9
Bc
2.0±0.3
1.6±0.5
2.0±0.2
2.2±0.7
2.9±0.9
0.5±0.3
Bf
4.0±0.3
3.7±1.3
3.9±1.1
4.2±1.2
6.8±2.2
5.6±0.9
Bt
34.8±2.5
39.8±7.4
36.7±5.9
39.6±5.3
35.8±6.7
39.4±7.6
?????
??????????, ??
Bc
22.9±1.5
23.7±3.6
22.2±4.2
22.7±4.6
20.1±1.7
9.8±5.5
Bf
26.7±1.6
25.5±2.0
25.8±5.9
27.4±4.8
25.7±3.5
26.7±3.3
Bt
8.1±0.3
8.9±1.4
8.9±1.3
8.9±1.6
8.2±0.9
8.4±0.7
?????
????????, ??
Bc
5.3±0.2
5.1±0.6
5.5±0.7
5.6±0.6
5.3±0.5
5.1±0.8
Bf
8.8±0.3
8.4±0.8
8.8±1.0
8.8±0.9
9.8±1.4
8.7±1.4
Bt
3.3±0.1
4.1±0.4
3.5±0.5
3.7±0.5
3.2±0.3
3.0±0.3
??????
????????, ??
Bc
2.7±0.1
2.8±0.4
2.9±0.4
2.9±0.5
2.6±0.3
2.8±0.4
Bf
2.4±0.1
2.1±0.3
2.5±0.3
2.5±0.3
2.4±0.2
2.2±0.2
Bt
1.7±0.3
1.5 ±1.3
1.3±0.5
1.6±0.8
1.3±0.8
1.0±0.4
????? 1-??
?????, ??
Bc
0.7±0.1
0.7±0.2
1.3±0.6
1.3±0.5
0.9±0.3
0.5±0.4
Bf
0.9±0.1
0.7±0.4
0.7±0.5
1.2±0.5
0.9±0.2
0.7±0.2
Bt ? Bidens tripartita, Bc ? B. cernua, Bf ? B. frondosa; * ? ??????????? ???????? ????? ??????????? ? ???????
?????????? (t-???????? ?????????, p < 0.05).
??? B. tripartita, ? ??????????? ? ? B. cernua
(????. 2).
???????? ???????? ????? ?? ???? ? ???????? ?????????? ??????????? ????? ??????????? ? ??????????? ?? ??? ????????????
? ?????. ?????????? ??????????? ??????
???????? ????? ??? ??????? ? B. cernua.
??? ???????????? ????? 25 ??/? ? ???????? ??????? ?????? ? ?????????? ????? ????
????????????, ?? ?????????? ???????????
?????? (??????????? ? ?????????? ????????? ??? 25 ? 50 ??/?) ? ??????????? ???????????????? ??????????? ????????? ?????
?????????? ?????????? ?? ?????????? ??
???????? ??? ???? ?????????????. ??????? ?? ???????? ??????????? ??? ?????-
??????? ???????????? ???? ? ??????????
(????. 2).
B. frondosa ???????? ????? ??????????
? ???????? ???????? ????? ????? ?? ????????? ? B. cernua. ? ????? ???? ????? ????????
????? ?????????? ????????? ??? ???????????? ????? ? ????? 25 ??/?, ? ??? 50 ??/? ?????? ????????????. ?????????? ???????????
?????? ?? ?????????, ? ????????????????
?????????? ????????? ????? ?????????? ?????????? ?? ?????????? ?? ???????? ??? ????
????????? ????????????? (????. 2). ????????????? ????? ??????, ??? ????????, ???? ??
???? ????????? ?????????? ?? ?????? (Ruano
et al., 1988). ?????? ? ??? ??????? ?????????? ???????????? ?????????? ???? ????? ??
# 200 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????????????? ???? Bidens (Asteraceae) ? ???????? ???????? ????? ?? ??????????
???? ?? ??????????? ???????????? ????? ??
???????.
???????? ??????????? ? ???????? ???????? ????? ????????? ????????? B. tripartita.
?????????? ?????? ???? ????? ?. Bidens
??????????? ?????? ? ??????? ???? ????????? ?? ????? ? ???????? ???????? ??????? ??????????. ?????? ??????????? ???????? ????? ???????? ????? ?? ????????? ? ????????
?????????? ???? ??? 50 ??/?, ? ??????????
??????????? ?????? ??????????? ?? ?????????? (????. 2). ?????????? ????? ????????? ????? ?????????? ?? ???? ?? ??? ?????
????????????. ??????? ???????? ?????, ???
??????? ????? ???????????? ???????? ????????? ? B. tripartita ??? ???????????? 1 ??/?, ?
B. cernua ? ??? 1 ? 10 ??/?. ???????? ???????
?????????? ?? ?? ?? ???? ?????????? ? ??????? ???? (??????? ? ??., 2011).
????? ?.?. ????????? (2008) ???? ???????? ??????? ???????????? B. tripartita ?
???????? ??????? ?????? (? ?????????????
0.01?25 ??/?) ?? ?????? ?????? ??????????.
??? ???? ?? ????????? ???????????? ?????????, ??? ? ? ????? ?????, ? ? ??????????
??????????? ???? ????? ???????? ????? ?
??????? ?????????? ?????. ???????? ?????
?????? ??? ???????? ???????????? Zn 21 ??/?
? ????? ?????????? Myriophyllum spicatum L.
? ??? ?????????? ???????? ?? ???????? ?????? ? ?????????? ?????????????????, ?? ???
??????? ????????????? (0.01?0.1 ??/?) ??????????? ? ? ?????? ??????? (???????, ????????, 2009; Stoyanova et al., 1997).
????????? ?????? ?????????? ? ????????? ?????????? ? ?????????? B. cernua ???
???????????? ????? 50 ??/?, ? ?? ????? ??? ?
?????????? B. tripartita ? B. frondosa ???????
?? ?????????. ??????? ????????? ? ??????? ?
?????????? ???? ??????????? ????? ?? ???????????, ???, ??-????????, ??????????????? ??
?????????? ???????? ?????????? ???????-
?????????? ?????????. ?.?. ??????? ? ?????????? (2009) ????????, ??? ? Setaria viridis
????? ?????????? ???????????, ????????????? ?????? ??????????? II ???? ??? ??????? ????????????? ????? (160 ? 320 ??/??
?????) ?? ??????????, ? ?????? ????????????
???????? ?????????? ??????? ?????????.
?????? ???? ??????, ?????????????? ??????????? ?????? ????? ??? ???????? ????????,
????????????? ? ??????? ??????? (?? ???? ????????? ??????? ??????????), ??????????? ?????????, ? ????? ? ???????? ????????
??????????? (Singh, 1988; Ruano et al., 1988).
??-????????, ????????? ????? ?????????
? ???????? ???????? ????? ?? ????????? ?
?????????? ? ??????? ???????? ????????
????????.
????? ???????, ???????? ?????????????? ? ?????????? ????? ?. Bidens ? ????????
???????? ????? ????????? ???????? ???????.
?????? ??????????? ?????? ? ???????????
????? ?????????? ??-???????: ????????? ?????? ???????? ????? ?????????? ??????????
?????? ? B. tripartita ??? ????????????? 25 ?
50 ??/?. ?????? ??????????? ?????? ???????
????? ? B. tripartita ??? ???????????? ?????
50 ??/?, ????? ??? ? B. cernua ? B. frondosa ???
???? ???????????? ??????????? ????? ?????????? ? ?????????? ?????????.
? ????? ??????? ????? ????? ????????
??? ????????? ?????? ?????????? ?????????????? ?. Biden, ??? ???????? ?????? ? ????.
???? ?????? ???????????? ??????????
???????????? ?.?. ???????? ? ??????????
(2010) ?? ??????? ?????, ??????? ????????,
??? ????????????? ??????????? ????? ?????? ?????????? ??? ????????????, ? 30 ???
??????????? ?????????? ????????????
????, ? 50 %-??? ????????????? ??????????? ????? ? ????????? ?????? ???? ??????? ? ???????????? ? 20 ??? ???????, ???
????.
# 201 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????????????? ???? Bidens (Asteraceae) ? ???????? ???????? ????? ?? ??????????
??????????
?????? ?????? ?????????????? (B. tripartita), ?????? ???????? (B. cernua) ? ??????
??????????? (B. frondosa) ???????? ???????
?????????????????, ? ???????? ??? ?????
???????????? ????????? 83?96 %. ????????
???????? ????? ?? ??????? ?????????????
??????? ?? ???????????? ????????? ?????
B. tripartita. ? B. frondosa ??????????? ??????????? ?????????? ???????????? ????????? ??? ???????????? 50 ??/?. ????????????? ????? ? ???????? ?????????? ????????
?? ??????? ?????????????? ? ????????????
????? ? ?????. ?????????? ??????????? ?????? ???? ????? ??????? ?? ???? ? ????????
???????? ??????? ?????????? ???? ??????????? ?????. ? B. cernua ? B. frondosa ???
???????????? 50 ??/? ??????? ?????? ????????????, ? B. tripartita ?????????? ?????????
??? ?????. ?? ??????? ?????????? ????? ?
???????? ?????????? ??????????? ???? ????????????? ? ????????? ????: B. tripartita
> B. frondosa > B. cernua. ??????? ?????????? ???????????? ?????????? ???? ?????
?? ???? ?? ??????????? ???????????? ????? ?? ???????.
?????????????
????? ???????? ???????? ??????????????? ???????? ???????? ?????????? ????????????? ????????? ??. ?.?. ????????, ??????? ????????????? ???? ?.?. ?????????? ??
???????????? ???????????? ? ?????? ? ?????????????? ????????? ????????? ? ????????
?????????? ????????? ???????? ??????????
??? ??. ?.?. ???????? ???, ????????? ????????????? ???? ?.?. ?????????? ?? ??????
? ?????????? ??????.
?????? ??????????
1.
2.
3.
4.
5.
6.
7.
8.
??????????? ?.?. (2003) ????????????????? ???????? ???????????? ??????? (?? ??????? ???? Bidens) ?: ???????? ???????? ??????????? ? ???????????? ????? ? ????????
???. ????????? ?????. ????. ????, ?. 31?33.
??????????? ?.?., ????? ?.?. (2008) ??????????????? ???????????? ??????????? ? ???????????? ????????? ????? ???????????? ? ???????????? ?????? ?: ???????????????:
???????? ? ??????????? ??????????. ????????? ????. ?????. ????., ??????. 135-?????
?.?. ?????????, ?. 1. ?????, ?. 17?19.
??????????? ?.?., ??????? ?.?., ????? ?.?. (2010) ?????? ????? ????? ??????? ??????:
?????????? ???? ???????? ? ??????????? ??????? ??????. ?.: ????, 512 ?.
??????? ?????????? ????????. ?????????????? ?????????? I-IV ?????: ?????. ???. (1988)
??? ???. ?.?. ?????? ? ??. ?.: ?????, ?. 18-34.
??????? ?.?., ???????? ?.?., ???????? ?.?. (2010) ????????????? ??????? ??????? ???????????? ????? ???? ? ????? ? ???????? ?? ?????????????? ? ????????? ?????. ?????????? ???????? 57(6): 864-873.
????? ?.?., ?????? ?.?., ?????? ?.?. (1985) ??????? ??????? ???????? ?? ????, ????????
? ??????????? ???????????????????? ???????. ????????? 6: 90-100.
??????? ?.?., ????? ?.?., ???????? ?.?., ??????? ?.?. (2009) ???????????? ?????????
???????? ? ?????????? ????????????? ?????. ???????? ???. ????? ????????????? 6:
677-684.
??????? ?.?., ????????? ?.?. (2011) ???????? ???????? ?????? ?? ????????? ????? ?????????? ???????? ???? ????? ???? Bidens (Asteraceae). ???????????? ??????? 1: 65-71.
# 202 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????????????? ???? Bidens (Asteraceae) ? ???????? ???????? ????? ?? ??????????
9.
10.
11.
12.
13.
??????? ?.?., ????????? ?.?. (2011) ??????????? ????? ? ???????? ?????????? ?????????????? ???? Bidens (Asteraceae) ? ????????? ???????? ????. ??????? ???????? ???????????????? ???????????? 352: 207-210.
??????? ?.?. (2008) ??????? ????????? ??????? ???????? ?? ??????????? ????? ? ???????? ?????????? Alisma plantago-aquatica (Alismataceae) ? Bidens tripartita (Asteraceae). ????.
??????? 4: 98?106.
??????? ?.?., ???????? ?.?. (2009) ??????? ???????????? ????? ????????? ???????
???????? ?? ????????? ????? ?????????? ?????????? ????????????????? (Batrachium
tricjophyllum) (Chaix) Bosch.). ??????? ???????? ???????????????? ???????????? 323:
364-369.
????????? ?.?. (2006) ???????? ???????????? ??????????? ????? ??????? ???????? ??????? ?????????? ? ???? ??????? ? ????????? ? ???. ???. ?????. ?? ??? 8(1): 332-340.
???????????? ? ??????? ???????? ???????????? ????? (1991) ??? ???. ?.?. ??????????-
???????. ?.: ?????, 214 ?.
14. ???????? ?.?., ??????? ?.?., ???????? ??.?. (2005) ????????? ? ??????? ?????????????
????? ???? ? ????? ???????? ??????????? ?????? ? ??????????? ?? ????????????? ? ????? ??????????????. ?????????? ???????? 52(6): 848-858.
15. Khudsar T., Mahmooduzzafar N., Jqbal M., Sairam R.K. (2004) Zinc-induced changes in morpho-physiological and biochemical parameters in Artemisia annua. Biol. Plant. 48: 255-260.
16. Rengel Z. (2000) Ecotypes of Holcus lanatus tolerant to zinc toxicity also tolerate zinc deficiency.
Ann. Bot. 86: 1119-1126.
17. Ruano A., Poschenrieder Ch., Boacelo J. (1988) Growth and biomass partitioning in zinc-toxic
bush beans. J. Plant Nutr. 11(5): 577-588.
18. Singh M.V. (1988) Response of rice (Oryza sativa) and wheat (Triticum aestivum) to phosphorus
and zinc fertilization on sodic soils. Indian J. Agr. Sci. 58(11): 823-826.
19. Stoyanova D.P., Tchakolova E.S. (1997) Cadmium-induced ultrastructural changes in chloroplasts
of the leaves and stems parenchyma in Myriophyllum spicatum. Physiol. Plant. 34: 241-248.
20. Wang C., Zhang S.H., Wang P.F., Hou J., Zhang W.J., Li W., Lin Z.P. (2009) The Effect of excess
Zn on mineral nutrition and antioxidative response in rapeseed seedling. Chemosphere 75: 14681476.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ?????????????? ???? Bidens (Asteraceae) ? ???????? ???????? ????? ?? ??????????
Stability of the Genus Bidens (Asteraceae)
to Action of Zinc Sulfate
in the Early Stages of Ontogeny
Elena G. Krylova
Institute for Biology of Inland Waters RAS,
Borok, Yaroslavl region, 152742 Russia
The effect of zinc sulfate (0.1 ? 50 mg/L) on seed germination and early stages of development of seedlings
of three species of the genus Bidens L. Zinc sulfate solution in the whole range of concentrations
indicated no significant effect on laboratory germination B. tripartita L. A significant decrease of
laboratory germination B. frondosa L. was found at 50 mg/L. Zinc (50 mg/L) had a toxic effect on the
growth and root development of seedlings of all species studied: namely, B. cernua and B. frondosa
missed the main root, while B. tripartita had the root which was significantly reduced in length. The
above-ground portion of seedlings was less responsive to the elevated concentrations of zinc in the
medium. Investigated species are located in the following series in order of decreasing resistance to
zinc sulfate: B. tripartita> B. frondosa> B. cernua.
Keywords: zinc sulfate, seed germination, seedling growth, Bidens tripartita, B. cernua, B. frondosa.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2013 6) 205-219
~~~
??? 630.114.351
??????? ????????? ??????? ?????????
???????? ??????????
?.?. ????????*, ?.?. ???????
???????? ???? ??. ?.?. ???????? ?? ???
?????? 660036, ??????????, ????????????? 50, ???. 28
Received 28.06.2012, received in revised form 14.09.2012, accepted 19.12.2012
? ???????? ?????????????? ??????? ?? ????????? ?? ????????????? ?? ??????????????????????? ????? ???????? ?????????? ???????????? ??????????????? ????
????????? (???????????????????, ???????????????????, ???????????, ???????????,
???????????? ? ????????), ??????? ??????????????? ?????? ? ???????????? ????????
?????. ???????? ?????????????-????????????? ?????? ??????????? ????? ?????????????
??????????? ?????????, ??????????-?????????? ??????? ????????? ???????, ?????????,
??????? ???????, ?????? ? ????????. ???????? ???????? ?????????????? ????? ???????????
? ??????????????? 3-? ???????, ??? ???? ????????? ????????????? ??? ?????????? ? ?????
?????????????????. ????????? ????????????? ????? ??????????? ????????? ? ??????????,
???????? ??????? ? ?????? ???????? ?? ????????, ?????????????? ???????, ? ???????
?????????? ???????? ????? ???????. ? ?????????? ????????? ???????????? ??????????
??????? ???? ????????? ??????????? ???????????? ??????, ???????????????? ????????
???????? ?????????? ????????? ????????? ????? ???? ??? ???????? ?????????? ???????
??????????. ????????? ???? ????????? ?????? ??????? ???. ????? ???????, ?????????
???????, ???????????? ? ???????? ?????? ???????????? ??????? ???????? ??????????,
????????????? ?????? ????????????? ???????????? ????????? ??????? ?????????.
???????? ?????: ???????????, ?????????, ???????? ?????????, ????????????? ??????.
????????
????????? ? ???? ?? ??????? ?????????????????? ??????????? ?????? ?????????. ???????? ????????? ??????? ?????? ??
???????????? ????????????? ????????????
?????? ????, ????????????? ?????????????
??????????? ? ?????? ???????????? ???-
*
???????, ?????????? ?? ???????????????? ????????? ? ??????? ?????????? ??? ????????????? ????????????? ????????? ??????????.
????????? ???????? ????? ????????????
????????????? ??????????? ??????? ????????????? ??????? ????????????. ???? ?????????? ? ?????????-???????????? ?????-
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: efr2@ksc.krasn.ru
# 205 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
???, ??????????? ? ???????????? ???????
?????????, ???????????? ????????????? ?????????? ????????????? ????????? ? ???????
???? ???????? ?? ????????? ???????? ?????????, ???????????? ????????? ????????????
???????????????? ?? ?????? ??????? ? ??????????. ? ??? ??? ???? ??????? ??? ???????
????? ????????? ? ???? ?????????? (????????, 1973; ??????????, 1983; ?????????,1983;
Luo, Christie, 2001; Kakei, Clifford, 2002; Batty,
Younger, 2007 ? ??.). ?? ??????? ??????????? ??????????? ? ???????????? ???? ?????
???????? ????????? ???????? ??????, ????
??????? ? ???????????? ????????? ??????
???????????? ?????????? ??????? ???????
?????????? 27,4 % (??????? ? ??., 2005; ???????? ? ??., 2009, 2010?, 2010?; Sekretenko et
al., 2011).
?????? ????????????? ???????????
???????? ????????? ?????? ?? ???????????????? ????????, ??????? ? ???????????? ???? ????????????? ?????????????????? ? ?????????????? ??????????
????????????? ?????????, ??????? ??????????
?????? ? ?????? ???????? ????????, ??????????? ?????????? ?????????, ?? ???????????
?????????, ????????????-?????????????????
????????? ? ?????? ???????? ?????, ?????????
? ???????? ????? ?????????? ?????? ?????????????. ?????????????, ???, ????????????
??????????? ?????????, ?????????? ????????? ?????????? ? ????????????? ????????????? ? ??????????????? ? ????????.
?????????? ??????????? (??) ???? ????????????? ?? ?????????? ????? ?+ ? ?????? ????? ??????????? ??????, ??????????????? ?
???????? ??????? ? ???????? ?????? ? ???????? ?????????? ????????? ???????, ?????????? ?????????? ??????????. ??????????????? ??????????? ?????? ? ????????????
???????? ???????? ? ????? ?????? ?????????
???????????, ??????? ???????? ? ????????-
??? ? ????????????? ??? ?????????????? ??
????????? ??????????. ??????? ??????????????? ???????????, ???????????? ? ??????? ???????? ???????? ????????? (?? 8,3),
???????????? ????????????? ??? ??????????
????? ????????????? ???????????, ??????????????? ????????? ?????????? ???? ????????? ??????????? (?????????, ?????????,
2006).
???? ?????? ?????? ? ????????????????
??????????? ????????? ???????? ??????????
? ???????? ??????????????? ????, ??????????? ?? ????????? ???????, ??????? ???????
??????? ???????????? ????????? ???????,
?????????? ??????? ??????????? ?????????? ? ??????????-?????????? ??????? ????????????? ???????? ??? ????????? ???????????????? ????????????? ?????????????
?????, ?????????? ?????????. ????? ??????
???????? ? ????????? ??????? ???????? ??????????? ?? ????????? ?????? ?? ?????????
??????? ????????? ? ???? ????????? ???????? ?????????? ? ?????? ?? ?????????????
????????.
????? ?????,
????????? ? ??????
???????? ???????????? ????????? ????????????? ????? ?????????? ??????, ?????????????? ? ??????????? ??????? ????????
??????. ?? ?????????? ??? ? ???? ??? ????
?? ????? ??????? ???????? ???????? ?????
???????? 2,3 ???. ??. ? ?????????? ???????
????? ?????????????? ???????? ???????????? ?????????????? ???????????????? ????,
?????????? ????????? ??????? ???????? ????? ?. ?????? (56°23?710???.?., 84°34?043???.?.) ?
??????? ????????????? ??????? ? 100?140 ?
?? ?????, ?? ???????? ??????????? ????????
?????? ?????????????? ?????? ????? ???.
?????????? ????????????? ????????????
??????? ?????????????? ????? ?????????-
# 206 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
??? ????? ?????? ? ?????????? ??????? ??????? ????????? ? ?????????? ???????????.
???????? ???????????? ????? ??????????
?????? ???????? (Betula pubescens Ehrh.),
??????????? ? ?????. ?? ???? ????????
?? ????????? ????? ?????????? ???????
????????-????????? ???, ????????? ?? ???????????, ????????????? ????????? ????
(???. 1), ??? ????????????? ???????????????
????? ???????????? ???????????? ???????????. ????? ???????? ???? ??????????????
?????? ???????? ??????????, ????????? ????????????? ??????? ?? ????? ??????????????
????? ?????? ????????? ??????? (270Ч60 ?). ?
??? ???????? ???????? ????????? ???? ????:
0?30 ? ?? ????? ? ?????????????-???????????????????? ????????, ?? ??????????
30-100 ? ? ??????????????????????, 100140 ? ? ?????????-???????? ?????????????,
140-230 ? ? ???????-???????-?????????, 230270 ? ? ???????-????????-???????????????.
??????????? ????????? ???????????? ??
????????????? ? ???????? ?? ???????????
??????????? ? ??????? ???????? ??????????????????? ?????????? (????????????, 1977;
??????????, 1984; ????????? ? ??., 2004). ?
??????????? ???????? ?????????? ?????
?????????? ???????????????? ???? ????????
????? ????? ?????? ?????????: ???????????????????, ???????????????????, ?????????????????? (???????????), ???????????,
???????????? ? ????????. ????????? ????? ???????? ?? 3,6 ?? 6,2 ?? ? ??????????
?????? ?????????? 0,047?0,079 ?/??3. ????
????????? ??????????, ??? ???????, ? ???????????? ??????? ??????????????? ???????,
????????? ? ????????? ??????????. ??????????????????? ??????????????? ?????????
(? ?????????????? ? L, F, H) ???????????? ??
????????, ??????????????? ??????????? ?
????? ?????????????? ????? (0?30 ?). ??????
????????? ??????? ???????? ????????? ??????????. ??????????????????? ?????????
???????? ???????????????? ???????????
(?) ???? ????????????. ???????????, ???????????, ???????????? ??????? ??????????????? ???????? ? ??????????????
??????????????, ???????? ? ?????? ????????? ????. ????????? ??????????? ? ?? ???????
???????????? ???????????. ? ??????- ? ??????????????????? ?????????? ????????????? ?????? ????? ???? (0?100 ?? ?? ?????)
?????????
??????????
40
40
30
30
20
20
10
10
0
??????? ??????????, ??
???????? ?????????, %
50
0
0-30
30-70
70-100 100-140 140-190 190-230 230-270
????????? ?? ????? ?????, ?
???. 1. ??????? ?????????? ? ????????? ????????? ???????? ?????????? ? ???????? ??????????????
??????? (??????? ???????? ?? ?????? ?????? 2005?2007 ??.)
# 207 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
226
242
217
215
235
0-30
30-70
70-100
100-140
140-190
190-230
230-270
??
??
??
??????? ???
219
???????
25
?????
0
230-270
236
50
190-230
86
50
????????
75
140-190
97
77
100
100-140
100
125
0-30
150
0
150
2
206
200
???????? ?????, ?/?
?????????? ?? ????? ?????????????? ?????, ?
?????????? ?? ????? ?????????????? ?????, ?
??
250
?????? ???????????, ??/??
0,93
??????
8,28
0,75
?????????
0,71
?????
??
?????????
4,47
5,02
0,65
???????????
4,47
0,28
??????? ???
235
205
7,77
4,56
196
200
????
???
150
120
100
50
46
30,0
32,7
26,2
0,25
3,72
3
50
38
5,0
5,4
4,8
0
6
9
12
0-30
0
15
??????????? ??????????, %
70-100
????????? ???
17,6
100-140
????????
230-270
12,24
1,1
190-230
1,72
???????
140-190
??
30-70
????? ?????, ?/?
2
175
????????? ???
250
70-100
269
30-70
280
277
??
????????? ????
???????-??????? ????
300
?
??????????
?
?????????? ?? ????? ?????????????? ?????, ?
???. 2. ????? ?????????? ????? ? ?????? ???????-???????? ??????? ? ????????? ??????? ??????????????
??????? ???????? ?????????? (?), ???????? ??????????? ????? ???????? ? ??????? ????????????
??????? (?), ?????????? ??????? ? ????????? ? ??????????? ????? ???????? (?), ????????????????
??????????? ???? ? ?????? ??????? ? ????????? ? ??????????? ???????????? ???????? (?)
?????????? ???????? ????, ??????????????? ?????? ???????, ?????????, ???????????
(???. 2?, ?). ? ??????????? ? ?????????????????? ????????? (100?140 ?) ??????? ??????
? ???????????? ????????????? ??????? (???????, ????) ?????????????? ????????????.
??????????? ? ???????????? ????????? ??????? ??????????????? ?????????? ?????????. ????? ????????? ?????????-????????????
?????????????? ????? ????????? ????????? ?
?????? (???????? ? ??., 2010?).
???????????? ??????? ????????? ? ???????? ?????????????? ???? ??????????? ??
?????????? 2004?2007 ??. ??? ?????? ??-
?????? ?????????? 10 ????? ??????????? ??
5 ?????? ? ????? ?? ????????? ????? ? ???????? ??????? ???? ????. ????????? ???? ?????????? ?? 70 ??????????????? (100Ч100 ??),
????????????? ????? 10 ? ???? ?? ????? ??
????? ?????? ???????? ?????????. ????????
???????????? ????????????? ??????? ???????????? ?????????? ? ???????? 50Ч50 ?? ?? 20
????????? ? ?????? ???? ??????????.
??????-?????????? ???????? ????????? ???????????? ????????????? ? ???????????? ???????? (?????????????? ??????
???????????? ????, 1975). ???????? ?????
????????? (??? ??????? ?? ???????) ??????-
# 208 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
?????? ? ??????? ???????? ??????? ??????,
????????? ? ????? ???????????? ?????????
? ?????? ??? 450 °? ? ??????? 6 ?. ????????
?? ?????? ??????? ???????? ??????????????????. ??????????????? (????? ?????????????) ???????????? ?? ?????? ???????,
???????? ??????????? ? ?? ?????? ?????????. ?????????? ???????? ???????????????
??????????? ???????? ?? 1,75 ? ???????
???????? ??????????? ?? ????????? ?????????? ?+. ????? ???????? ????????? (Ca2+
+ Mg2+) ???????????? ?? ?????? ????????
??????????, ????? (???????) ??????????
??? ? ?????????????????? ??????? ? ??????????? ?????????? ????? ??????? ????????
????? ?????? ?????? ? ??????? ??????. ?????????? ????????? ?????????? ? %, ?.?. ??
100 ? ????????? ??? ????????, ??????????
?? ??????????? ???? ??? 105 °?. ?????????
? ??????????? ?????? ????????????? ???????? ???????? ?? ???????? ?.?. ????????-
??? ?????????? ?? ??????? ???? ? ???????
6 ?????. ? ????????? ??????? ?????? ?? ???????? ??????? ???????? ????????? ??????? ??????????? ?????????????? ?????????? 1N H2SO4. ????????????? (??-1, ??-2
? ??-3) ????????? ?? ???????? ????? ????????? ??????????????? ???????? ???????
? ????????? ??????. ??????????? ???????
???????????? (??-1?) ???????????? ??????????????? ? 0,1 N ???????? H2SO4. ???????
???, ?.?. ?????????? (1980). ????????????
????? ???????? ? ????? ?????????? ?? ?????? ??????? ? ?? ????????? ??????????????.
? ?????? ????????? ??????? ???????? ???
???????? ??????: ????????? ??????? (??)
? ????????????? (??), ??????? ???????????
??????????????, ?????????????, ?????????? ????????, ???????? ???????? ????????
? ??. ??????????. ?????? ?? ????? ????????
????????? ????????? ???????, ??????????
????????? ????? ????????? ??????? ? ???????????? ??????????? ??????, ?/??? ???????
???????????? ????????? ??????? ? ??????
??????? ???????????? ?????????????? ????????????? ????????? (???????? ????????,
?????????). ? ???? ??????? ?????? ???????
????????? ?????? (??-1 + ??-1) ???????????
??????????????? ????????? 0,1 N NaOH ??
??????, ?????? ??????? (??-2+??-2) ? ????????? 0,1 N NaOH ????? ???????????????,
?????? ??????? (??-3 + ??-3) ? 0,02 N NaOH
?????????? ? ??????????
???????? ???????, ????????? ?????? ?
???????????? ??-1? ?????????? ??????????? ??????? ?? ??????. ?????????? ?????
? ??????? ????????????? ???????? ????????
? ????????? ?? ????? ?????? ?????????????
???????? ? ?????????.
????????????????? ???????? ????????? ???????? ??????????????? ? ??????????????? ?????????????? ??????? ? ?????? ???????? STATISTICA 6.0.
?????????-???????????? ???? ????????? ???????? ?????????? ???????????? ?????
?????????????? ????? ??????????? ????????? ?????? ????????????? ????????: 4,5 ??/?2
(0?30 ?) > 3,7 (30?100 ?) > 2,1 (100?140 ?) < 2,5
(140?230 ?) < 3,1 ??/?2 (230?270 ?). ?????????
?????????? ??????????????? ? ???????????????? (????. 1). ?????????? ???????????
????????? ???????????: Fe2O3 ? 0,53?1,64 %,
Al2O3 ? 0,70?1,57, ??? ? 0,30-1,44 %, ????????
????????? (Ca+Mg) 33,54?86,6 ??-???./100 ?
???????. ?????????????? ??????? ?????????????? ?????????? ?????. ????????? ?????
???????????????? ??????? (100?140 ?) ?????? ???????????? ???????, ????????????
???????? ????????? ? ???????? ????????????? ???????? ?????????? ????? ????????? ??????????????. ? ??????????? ?????
?????????????? ????? ????????????????
???????? ????????? ??????????: ??????????
# 209 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
??????? 1. ??????? ???????? ?????????? ??????? ? ??????????-????????? ?????? ?????????????
???????? ????????? ???????? ??????????
??????????
????????????????? ???? ????????? ? ??????????
?? ????? ?????????????? ?????, ?
????????????
?????????????????????????????- ??????????????,
??????,
??????,
???,
?????,
?????,
100-140
140-190
190-230
230-270
0-30
30-100
?????????? ????????, % ?? ????? ?????????
?????????
15,73
???
1,44
Fe2O3
1,64
Al2O3
1,57
??+?g, ??-???./100 ?
????? ??????????
14,86
10,26
8,03
7,27
8,52
1,15
0,67
1,24
0,93
0,46
0,32
0,30
0,92
0,66
0,53
1,34
0,77
0,70
0,89
1,10
86,62
30
71,26
54,09
44,68
44,3
33,54
30
30
15
15
30
????????? ? ??????????? ?????? ????????????? ????????, % ?? ?????? ???????? ?????????
????????? C/N
18,3
22,37
27
30,22
32,05
40,24
15,77
11,89
10,26
11,85
8,75
6,33
????????? ???????:
??-1
??-2
???
???
???
???
???
???
??-3
9,07
8,36
8,03
8,04
7,7
7,42
????? ??
24,84
20,25
18,28
19,89
16,45
13,75
??-1?
3,011
2,93
3,51
2,75
3,27
2,97
??-1
13,43
13,41
12,45
10
12,59
11,78
??-2
3,0
2,65
3,22
3,44
3,64
2,97
?????????????
??-3
5,12
5,23
5,51
6,03
5,78
6,02
????? ??
24,66
24,22
24,69
22,24
25,28
23,74
??-1+??-1
29,2
25,3
22,71
21,85
21,34
18,11
???+???
49,5
1
44,47
0,84
42,96
0,74
42,13
0,89
41,73
0,65
37,49
0,58
5
5
5
5
5
5
??? /???
????? ??????????
????????? ????????, ?????????, ??????????
??????? ????????????? ????????, ????????
?????????. ?????? ????????? ??????? ??? ?????????? ????????? ? ???????? ????? ??????????.
??????? ??????? ??????????????? ?
???????? ?????????????? ?????????????
???????? ?????????. ? ??????? ???????????? ???????????? ???????? ??????????????? ??????? ?? ??????????? (???+???) ?
37,5?49,5 % ? ????????? ?????? ???????? ?
?????? ????? (C/N), ??????? ? ??????????
??????????? ??????????? ?? ???? ??????????? ? ????? ???? ? 40,24 ?? 18,3. ? ???????
????????? ?????? ???????????? ???????
??-1, ?????????? ??????? ???? ??????????
? ??????????????????? ????????? (15,77 %),
?????????? ? ? ???????? (6,33 %). ????????? ? ?????????? ??-1 (10,0?13,43 %) ?????
????????, ???? ? ????????? ??????????
# 210 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
???????????? ??-1 ?????????. ?????????
??????? ??-3 ????????????????? ??????????????? ? ??????????????????? ?????????,
????????? ? ???????? ?????????????? ??????? ?? 7,42 ?? 9,07 %, ????????????? ??-3
? ?????????? 5,12?6,03 % ?????????????
? ??????? ?????????. ????????? ???????
??????? ??-2, ????????? ? ????????, ?????????????, ??? ?????????? ??? ?????? ?????????????. ????????????? ? ??-2 ? ??????????? ??????? ??-1? ?????????? ?? ???? ?????
????????? ??????????? ? ?????????? ?????????? (????? 3 %) ?, ?????????????, ?? ?????
??????? ?????????????? ????????. ?????
???????????????????, ??????????????????? ? ???????????? ????????? ??????????????? ???????? ?????: ????????? ????????
????????? ?????? ? ???????? ????????????
(??? / ???) ? 0,74?1, ????? ??????????? ?
???????? ????????? ? ?????????-????????
?????: ??? / ??? 0,58?0,65 (?????? ??: ????,
????????, 1974).
????????? ???????? ?????????? ????????????????? ?????? ? ????????????
????????, ??????????? ?? ??????? ?????????????? ??????? ?? 3,7 ?? 5,5 ??, ??????? ????????????? ????????????: ?????
(???????????????) 75,9?174,4 ? ????????
3,7?25,8 ?????(+)/100 ? (????. 2). ?? ???? ???????? ?? ????? ?????????????? ????? ?????????? ??????????????? ????????? ? ??????????????? ? ???????? ?????? (Cv ? 10 %) ?
??????? ???????????????.
??? ???? ???????????, ???????? ?????????????? ??????? (R2, ?-??????? ??????????), ????? ? ??????? ????? ???????????
??????? 2. ?????????????? ????????? ????? ??????????? ????????? ???????? ?????????? ??
?????????????? ???????
230?
270
190?
230
140?
190
100?
140
70?
100
30?70
?????????? ?? ????? ?????????????? ?????, ?
0?30
??????????????
??????
?? ??????????????
???????
? ?????
????? ????????????? (???????????????) ???????????, ?????(+) / 100 ?
???????
91,1
91,4
95,4
118,4
138,3
149, 9
151,9
119,8
???????
75,9
77,9
76,8
106,2
116,5
122,6
139,3
75,9
????????
104,0
99,2
108,2
132,0
150,5
165,3
174,3
174,4
Cv
7,1
6,3
7,5
6,2
8,3
9,5
5,4
22,3
n
30
15
15
30
15
15
30
150
???????
5,7
5,3
6,0
10,2
13,9
15,8
16,2
10,5
???????
4,0
3,7
4,1
7,2
9,8
11,7
11,5
3,7
????????
8,5
7,3
8,5
13,5
19,1
25,8
22,0
25,8
Cv
22,2
22,3
23,1
17,5
23,1
24,4
?????????? ??????????? (?? ??????)
19,2
48,1
???????
5,0
5,0
5,0
4,6
4,3
4,1
4,0
4,6
???????
4,7
4,8
4,8
4,4
4,0
3,8
3,7
3,7
????????
5,5
5,2
5,3
4,9
4,5
4,3
4,2
5,5
Cv
3,5
2,2
3,1
3,3
4,5
4,1
3,6
10,1
???????? ???????????, ?????(+) / 100 ?
Cv ? ??????????? ????????, n ? ????? ????????? ??? ???? ????? ???????????.
# 211 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
??
25
y = -1 0 , 5 6 x + 5 8 , 8 1
R 2 = 0,88,
p < 0,001
20
????? ?????????????
30
15
10
5
0
3 ,5
4
4 ,5
5
??
160
y = -5 7 , 7 x + 3 8 3 , 8
R 2 = 0,95,
p < 0,001
140
120
100
80
3 ,5
4
4 ,5
5
5 ,5
??
?
30
?
25
?????(+)/100?
180
60
5 ,5
??
???????? ???????????,
???????????, ?????(+)/100?
?????(+)/100?
???????? ???????????,
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
y = 0,17x - 10,41
R 2 = 0,85,
p < 0,001
20
15
10
5
0
60
90
120
150
180
? ? ? ? ? ? ? ?? ? ?? ? ?? ? ? ?
? ? ? ?? ?? ? ? ?? , ? ? ? ?? (+ )/ 1 0 0 ?
???. 3. ?????? ????????????? ?????? ?????: ???????? ??????????? ? ???????? ?? (?), ?????
????????????? ??????????? ? ???????? ?? (?), ???????? ? ????? ????????????? ??????????? (?)
????????????? (???. 3). ????????????? ?????
??????????? ??????????? ? ? ?????????? ??????????? ????, ?? ???, ??? ???????, ??????????????? ?????? ??????????? (?????, 1985;
?????????, ?????????, 2006). ???????, ????????????? ???????? ?????????????? ????? ????? ??????????? ? ????????? ????????
?????????? ???? ????????? ?? ?????????????
?? ? ???????? ??????????????? ?????????
??? ???????? ?? ?? ????????? ???????, ???
?????? ???????? ?????? ?.?. ?????? (1985).
??????? ????? ????? ??????????? ?????? ????? ????????????? ???????????, ??????? ?????? ? ????????? ??????????? ????????
????????? (Ca+Mg) ?????????? ? ????????
???????????????? ???????? ????????????
?????????? ????????? ???????? ??????????
(???????? ? ??., 2010?).
????????????? ??????????? ?????????
????? ?????????? ?? ?????? ??????????. ????
?? ???????? ?????? ????? ? ?????????????
????????? ??????????? ?????? ???????????
???????????? ??????? ? ???????? ????????????????? ???????????????, ??????, ??????
? ?????? ????????, ? ????? ?????????????
????????? ???????, ???????????? ? ?????????? ??????? ????????????? ?????????????
???????????? ????????. ???????? ???????????????? ????????? ? ???????? ?????????
??????? ????????? ???????? ?????????? ???????? ?????? ??????????? ???????.
????????
??????????????
???????
(????. 3) ??????? ????????????? ???????? ???????????? ??????????? ????? ????? ????????????? ???????????? ? ????????????,
???????????????? ??????? ?????????????
# 212 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
??1+??-1
??1+??-3
-0,83
-0,90
-0,84
-0,64
0,94
-0,91
0,80
0,75
0,86
0,77
0,71
-0,92
0,87
Fe2O3
-0,94
0,91
???
0,71
0,98
-0,95
0,96
0,96
0,99
0,96
0,51
-0,86
0,98
Al2O3
-0,70
0,86
0,71
???
0,78
-0,56
0,61
0,51
0,67
0,54
0,74
-0,74
0,68
???
-0,92
0,92
0,98
0,78
???
-0,91
0,94
0,91
0,98
0,92
0,58
-0,88
0,96
C/N
0,93
-0,84
-0,95
-0,56
-0,91
???
-0,96
-0,93
-0,95
-0,93
-0,56
0,90
-0,97
??**
-0,86
0,80
0,96
0,61
0,94
-0,96
???
0,96
0,98
0,96
0,54
-0,86
0,99
??-1
-0,83
0,75
0,96
0,51
0,91
-0,93
0,96
???
0,97
1,00
0,30
-0,72
0,95
??-3
-0,90
0,86
0,99
0,67
0,98
-0,95
0,98
0,97
???
0,98
0,48
-0,84
0,99
??-1+?-?3 -0,84
0,77
0,96
0,54
0,92
-0,93
0,96
1,00
0,98
???
0,32
-0,74
0,96
??-3
-0,86
-0,84
??-1
0,93
0,92
??-3
-0,92
0,86
??-1
C/N
-0,70
0,91
??**
CaO
-0,94
???
???
Fe2O3
-0,96
????????? -0,96
?*
?*
Al2O3
??????????
?????????
??????? 3. ???????????? ?????????? ??????? ?????????? ? ????????????? ??????????? ?????????
???????? ?????????? (????????????? ??????????? r > 081 ? ???????????? 95 %)
??-1
-0,64
0,71
0,51
0,74
0,58
-0,56
0,54
0,30
0,48
0,32
???
-0,85
0,59
??-3
0,94
-0,92
-0,86
-0,74
-0,88
0,90
-0,86
-0,72
-0,84
-0,74
-0,85
???
-0,90
??1+??-1
-0,91
0,87
0,98
0,68
0,96
-0,97
0,99
0,95
0,99
0,96
0,59
-0,90
???
?* ? ????? ????????????? ???????????, ??** ? ??????? ??????????? (???+???).
????????????? ?????????: ?????????????
?? ???????? ??????????? (??? + ???) ? ????????????? ? ?????????? C / N. ??? ????
??????? ????? ?????? ????????? ???????
????? ???????. ????????????? ?????? ??
????????? ???????? ????????? ????????????: ? ???????? ??-3 ??????????? ??????? ????????????? ?????, ? ??-1 ? ?????????????.
????? ?????????? ????? ???????? ?????????? ?????????? ? ???????????? ??????? ????????????? ???????? ? ???????? ???????????
?????????, ????????? ????????? ?????????????? ?????????????? ???????. ????????
???????? ????????????? ? ?????????????
???????????? ??????????: ??-3, ??-3, ?????? ????????? ?????? (??-1+??-3), ?????????
??????? 1-? ??????? (??-1+??-1) ? ????? ????????? ?????? (???+???). ??? ?????? ? ?????? ?????????? ?????? ??????????? ? ?????
?????. ??????? ? ?????? ????????????? ????????? ???????????? ????????? ?????????.
?? ????????? ?? ??????????? ?????????? ??-
??????? ????????????????? ???????????????
????????? ?? ??? ????????, ??? ??? ?? ????
?????????, ??????? ??????? ??????????,
???????? ???????? ?????? ????, ??????? ?
??????? ???? ?????? ?? ????????? ?????????? (???????, 2003)
?? ??????????? ?????????? ??????????? ????????? ???????? ???? ????? ???????
? ?????? ????????? ??????? 1-? ??????? ?
??-3 ? ???????? ???????? ???????? ??????????????? ??????????? ????? ????????????? ??????????? ????????? (R2 ? 0,84?0,89)
(????. 4). ??? ????????? ?????????? ????????? ??????? ???????????, ??? ???????,
?? ?????? ??? ???????????????. ????????????? ??????????? ????? ?????????????
??????????? ? ??????????? ????????????
3-? ??????? ???????????, ????????, ???, ???
??? ?????????? ?? ????????? ????? ??????????? (????????, 1992). ?????? ???????????? ??????? ???????????, ? ????????? ?.?.
????????????? (1980), ???????????? ?????
# 213 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
??????? 4. ?????????????? ?????? ????????? ????????????? ???????? ????????? ????? ?????
????????????? ???????????, ?????????? ??????? ??????? ? ?????? ? ??????????-?????????? ???????
????????????? ???????? ? ????????? ???????? ??????????
??????????
?????????????????
??????????? ? ??? ??????
?
???????? ???????? ??????
p
R2
F-????????
p
??????????? ????????
Ca?
Fe2O3
-0,956
0,003
0,91
???????? ?? ??????
42,90
< 0,003
0,89
31,19
< 0,005
0,89
31,19
< 0,005
0,84
20,55
< 0,010
0,89
31,19
< 0,005
0,84
20,55
< 0,010
0,91
42,90
< 0,003
0,91
42,90
< 0,003
0,85
23,1
<0,009
????????? ????????
??-3
??-3
0,941
??-3
??-1 + ??-1
0,941
??-3
??-1 + ??-1
0,005
???????? ?? ??????
0,005
???????? ?? ??????
???????? ?? ??????
-0,915
0,010
??-3
??? + ???
0,941
0,005
??-1 + ??-1
??? + ???
-0,915
???????? ?? ??????
0,010
???????? ?? ??????
????????? + ??????????? ????????
Ca?
??-3
-0,956
Ca?
??-1 + ??-1
-0,956
Ca?
??-1 + ??-3
-0,923
0,003
???????? ?? ??????
0,003
???????? ?? ??????
0,009
???????? ?? ??????
????????????? ??????????????????, ? ???????? ???????? ??????? ??????????? ?????? ?????????????? ??????. ??????????
??????? ????????? ?????? ???????????? ?
?????????? ?????????? ? ??????? ????????? ?????????????. ????????????? ???????
?????????? ??????????? ????????? ? ??????
(??-1+??-1) ???????????, ??-????????, ????? ?????? ???????? ????????? ?????? 1-?
???????. ????????? ?????, ??? ????????????
????? ??????????-?????????????? ???? ?
???????? ????????? ????? ????????? ?????????????????? ????????? ??????????????
????????? ?????? ? ??????????? ?????? ?????????. ???????? ?.?. ????????????? (1980),
????????? ???????, ? ????????? ?????? ?
????????, ??? ?????????????? ? ?????????? ????????? ???????? ??????????? ????,
? ??????? ?????? ?????? ? ?????? ????????
????? ???????? ? ?? ???????? ? ????????
????????. ??? ???? ??????????? ?????????
????????? ????? ????????????? ?????, ??????? ????? ?????????? ????????? ????????
? ???????-????????? ?????????. ??? ?????????? ???????? ? ??????? ???? ?? ?????
?????? ??????????? ?, ?? ??????? ???????????? ? ????????, ????????? ????????????
??????? ?? ????????????? ?????? ?????. ?????????? ?????????? ?????? ???????? ????
??????????????. ???, ??????- ? ?????????-
# 214 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
?????????? ?????????, ? ??????? ?????????
?????? ??????? ?????????? 1-? ???????, ??????????????? ? ????? ?????? ????????????
?? ????????? ? ??????? ??????.
????????????? ??????????? ???????????? ? ?????????? ??????? ????? ??????????? ????????? ? ???????????? ????????????. ???????????, ??? ????? ?????????????
??????????? ????? ???????????? ??????? ?
?????????? ?????????, ???????? ???????,
?????? ? ?? ??????? ? ????????? ?????? ? ??????? ???????? (????. 3). ????? ????????,
????? ?? ??????? ?????????, Fe2O3 ??? CaO,
?????? ????????????? ????? ?????, ????????? ?????????? ??????? ?? ???????? ?????
????????????? ???????????, ????????? ???????? ????????? ????????? ?????????????? ?????????????? ???????. ? ??????????
?????????? Fe2O3 ? CaO ? ???????? ??????????? ? ????????? ????????????? ?????????
??? ??????? ?????? ????? ???????, ????????? ? ???????????? ????? ?????????????
??????????? ?? 91 % (????. 4). ????? ??????
???????? ?? ?????? ??? ???????????????.
????????, ??????, ??????? ??????????? ????
? ?????????? ?????????, ?????? ? ??????
???????? ????? ???????? ? ????? ?????? ??
????????? ???????? ?????????, ??? ??? ??
???????? ? ???????? ????????.
?? ????? ??????????? ????????????
? ??????????? ???????????, ???????? ?????
????????? ? ????????????, ????? ?????????????? ?????????? ??????? ?????? ??????????? ?????????. ??????????? ?????
???????? (????????????) ? ????????????
??????? (R2 0,85?0,91) ? ????????????? ????????. ?????? ??? ?? ????????? ? ???????, ?? ??????????? ? ?????? ??????????,
? ?????????? ??????? (??? ??? ??????????
?? ????? ?????? ????? ?????????) ?????? ??
????????? ???????. ?? ????? ???????, ??? ?
????????????? ????????? ??????? ???????-
?? ???????? ?????????? ?????? ???? ??????????? ??????? ??? ?????? ?? ????????
????????? ?? ???????????? ???????.
??????????? (???????) ???????? ????????? ????????? ? ????????? ????????
?????????? ? ????????-?????????? ??????,
???????????? ????????, ???????? ? ???????? ?????????? ???, ?? ??????? ??????? ?
????????????. ??????? ??????? ??????? ??????? ? ??????????? ???????????? ??????? ?
????????? ????? ?? ???????? ????? ????????????? ??????????? ??????????? ? ???????
???????????? ?????? ?????????????? ?????????????? ???????. ???????? ??????? ???????????? ? ?????? (R2 0,93), ????????????
??????? ???????? ????????????, ?????????
????? ??? ???????????? ????? ????????????? ??????????? ????????? ???????? ?????????? (????. 5). ???????? ?????????????????? ????????????, ??????????? ??????????
?????? ??????? ??????????, ???????????????
? ???, ??? ??????? ??????? ???? ????????????
??????? ?? ??????????? ????????? ??????????. ????? ??????????? ??????? ??????????
????? ? ????????????? ?? ??????. ????????????? ??????? ?????? ?????? ?????????????? ?? ????????? ???????, ???????? ?? ?
???????????? ?????? ??????????? ???? ???????? ????? ? ??????? (???????????? ? ??.,
1980; ????????, 1991; ?????? ? ??., 2006, 2008;
? ??.).
??????????? ???????????? ????????????? ??????? ??????? ?? ?????? ? ????????? ????? ????????????? ????????, ???????????? ? ????????? ?? ????????? ??????
??????????????, ?? ? ? ????????????? ???
??????????? ???????. ????????????????
?????????? ?????????? ????? ? ????????
?????????????? ??????? ?????????? ????? ?
?????????? 221,2 ?/? 2 (???. 2?). ???????? ???????????? ????????????? ???????, ????????, ????????? ? ??????? ????????, 77?280
# 215 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
??????? 5. ?????????????? ?????? ????????? ????????????? ????????? ????? ?????????????
??????????? ?????????, ??????? ???? ? ????????? ????? ? ???????-??????? ??????????? ???????
??????????
???????????? ????????? ? ?? ??????
?????????????????
?????????????
t-????????
?
???????? ???????? ??????: R = 0,93 F-???????? = 28,4 ? < 0,004
2
?????????
???????????
??????
????????? ????
167,78
4,338
0,012
?0,843
?2,216
?3,772
0,019
?0,147
?2,715
?0,656
0,547
?/? 2, ??? ??????????? ??????? ??????????????????????? ?????????? ??????????????
???? ???????? ??????????. ??? ???? ??????????? ???? ???????? ???????????? ??????? ? ??????? ??? ??????? ???? ??????????? ??????????? ???? ? ??????? (???. 2?).
? ?????????, ?????? ????????????? ? ???????? ? ?????????? ????? ???????, ????????, ??????. ?????????? ????????????,
??????????? ??????? ??????? ?? ???????
0?100 ? ???????????? ?????, ?????????? ??
196 ?? 235 ??/?? ??????? ??????? ? 26,2?30
??/?? ???????, ??? ? 5-6 ??? ??????, ???
???????-???????????? ? ??????? ??????????, ????????? ?? ???????????? ???????
?????????????? ????? (???. 2?).
????? ??????????????, ??? ? ????????
????????? ???????? ?????????? ??????????
???????????????? ???????? ????????????
??????? ?? ??????????? ??????????? ??????????, ? ??? ?????, ???????-?????????
?????????, ????????? ? ???????? ????????????? ????????? ????? ????????. ? ?????????????? ???? ???????? ??????????
??? ?????? ?????????? ??? ? ?????????,
??? ???? ????????? ????? ?????????????
???????????. ??????????? ???????????????? ????????? ????????????? ??????????:
y = 164,34 ? 55,49 x, ??? y ? ????? ????????????? ???????????, ?????(+) / 100 ?, x ?
??? %. ?????????? ??????? ???????? ????-
??: R2 = 0,95, F-???????? = 76,48, ?-???????
< 0,001. ?? ????????? ???????: ??? ??????????? ???????? ?????? ??????? ?? ??????? (?
???????? 0,3?1,44 %) ????? ?????????????
??????????? ????????? ????????? ? ???????
?? 55,49 ?????(+) / 100 ?.
?????????? ????????? ??????????????? ? ???, ??? ??????????? ????????? ??????????????? ???? ???????? ?????????? ??
??????? ????? ???????????? ????????????
?????????? ???????????? ????????????? ???????, ????????????? ?????????? ???????.
??????????? ??????????? ???????-????????
????? ? ??? ?????????????? ? ???????????????? ????????????? ??????????? ??? ??????????? ????? ???????????? ????????,
??? ? ????? ?????????? ??????????? ????????? (? ??? ????? ???????), ???????????
? ?????????. ????????? ???????, ??????????
??????? ? ?????????????? ???? ?? ?????????? ? ????????, ??????? ? ????????? ?????????? ?????????? ????????? ?????, ??
???? ??????? ??????????? ??????????????
??????????? ???????? ???????????? ???????? ?????? ??????.
??????????
? ????????????? ???? ???????? ?????????? ?? ????????????? ?? ??????????????????????? ????? ???? ????????????
????????? ???? ????????? (???????????-
# 216 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
????????, ???????????????????, ???????????, ???????????, ???????????? ? ????????),
??????? ??????????????? ?????? ? ???????????? ???????? ?????. ????????????
???????? ???? ???????? ????? ????????
????? ???? ???????? ??????????. ????? ????????????????? ??????? ???????? ???????
???? ???????????? 3-? ??????? ? ?????????
?????????? ????? ??????????? ???????. ?
????? ????????, ???????????? (????????????????) ??????????? ?????, ???????????
???????????????? ???? ???????. ????????,
??? ?????????? ????? ? ?????????? ????????? ???????? ??????????, ? ??? ????? ????????, ????????? ??????????? ????????????
??????, ???????????????? ???????? ????????
?????????? ????????? ????????? ????? ????.
? ???????? ????? ?????? ????????? ????????? ?????, ????? ?? ??????? ?????? ??????
?????? ??????????????, ????????????? ?????? ????????????? ???????????? ????????? ???????? ?????????. ????????? ????
???????? ?????????? ????????? ?????? ??????? ???.
?????? ????????? ??? ?????????? ????????? ????????? ?????????? ??? ? 30 (?????? ?? ??? ? 30.11).
?????? ??????????
1.
2.
3.
?????????????? ?????? ???????????? ???? (1975). ?.: ?????, 656 ?.
???????????? ?.?. (1980) ???????????? ???????? ????? ? ???????? ??? ????????????.
?.: ?????, 288 ?.
???????? ?.?. (1991) ??????? ??????????? ???????? ?? ???????? ????? ? ?????? ???. ??-
?????????? 10: 131-143.
???? ?.?., ???????? ?.?. (1974) ??????? ?????? ? ???????????? ???????? ? ??????????????????? ???????? ???? ?????? ?????. ???????: ??????????, 113 ?.
5. ????????? ?.?., ????? ?.?., ??????? ?.?., ??????????? ?.?. (2004) ? ????????? ????????????? ???????? ???????????? ?????? ?????????. ??????????? 4: 17?30.
6. ????????? ?.?., ????????? ?.?. (2006) ????????????? ???????????. ??????? ? ??????????. ???????????? 4: 421?431.
7. ??????? ?.?., ???????? ?.?., ??????? ?. (2005) ????????????? ?????????????? ? ??????????
??? ????????? ?????? ????? ???????? ??????. ????????? ????????????? ?????? 1: 29-44.
8. ???????? ?.?. (1973) ???????????? ?????? ????????? ??? ???????????? ????????? ????????? ?????. ?: ??????????? ?????? ????? ? ???????????? ????? ? ????? ? ?? ???????????. ???????????: ?????, ?.142?162.
9. ???????? ?.?. (1992) ???????????????? ? ??????????? ???????? ???? ???????? ?????????. ???????????? 12: 25?35.
10. ???????? ?.?., ?????? ?.?., ??????? ?.?., ?????????? ?.?. (2009) ??????????? ????????????? ????????????? ???????? ????????? ???????? ??????????. ???????????? 10: 1203-1212.
11. ???????? ?.?., ??????? ?.?., ?????? ?.?. (2010?) ? ????????????? ?????????????????
????? ????????? ? ?? ?????????? ? ???????? ??????????. ???????????? 8: 920-928.
12. ???????? ?.?., ??????? ?.?., ?????? ?.?., ?????????? ?.?. (2010?) ??????????? ?????????? ????????? ???????? ??????????: ??????? ?????????? ? ????? ? ???????????????????
??????????. ?????? ?????????? ???????????? ????????????, ???????? 3(2): 211?223.
4.
# 217 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
13. ???????????? ?.?. (1977) ???????? ?????????? ??????? ? ?????? ????????????. ?.: ???-??
???, 312 ?.
14. ???????????? ?.?., ???????? ?.?., ?????????? ?.?. (1980) ? ???????? ???????? ?????????? ??????? ? ?????? ?????????????. ???????????? 5: 40?49.
15. ?????????? ?.?. (1983) ???????? ?????????? ? ????????? ??????? ???????? ?????. ?:
???? ????????? ? ?????? ?????????????. ?.: ?????, ?. 89?90.
16. ?????? ?.?., ??????? ?.?., ?????? ?.?. (2006) ??????????? ? ??????????? ????? ????
?????? ?????. ?: ???????? ?????? ????: ????????????????, ????????? ? ???????. ???:
?????, ?. 215-254.
17. ?????? ?.?., ????????? ?.?., ?????? ?.?. (2008) ??????????? ????? ???? ????????????? ?????. ?.: ?????, 342 ?.
18. ????? ?.?. (1985) ????? ????. ?.: ???-?? ???, 576 ?.
19. ?????????? ?.?., ????????? ?.?. (1980) ????? ? ????????????????. ?.: ?????, 222 ?.
20. ????????? ?.?. (1983) ?????? ????????? ??? ?????? ????????????????. ?: ???? ?????????
? ?????? ?????????????. ?.: ?????, ?. 168.
21. ??????? ?.?. (2003) ?????????????? ?????? ??????????? ??????. ?????????? ?????? ?????????? ???????? STATISTICA. ?.: ??????????, 312 ?.
22. ?????????? ?.?. (1984) ?????? ????????? ? ????????????, ????????????? ? ??????????.
???????????? 5: 96?105.
23. Batty L.C., Younger P.L. (2007) The effect of pH on plant litter decomposition and metal cycling
in wetland mesocosms supplied with mine drainage. Chemosphere 66(1): 158?164.
24. Kakei M., Clifford P.E. (2002) Long-term effects of liming on needles, soil properties, and soil
water in a Sitka spruce stand on deep peat. Forestry 75(5): 553?567.
25. Luo Y.M., Christie P. (2001) Short-term effects of alkaline biosolids on pH and trace metals
growth of Picea sitchensis. Forestry 74(2): 145-159.
26. Sekretenko O.P., Efremova T.T., Avrova A.F., Efremov S.P. (2011) Factors influencing the spatial
pattern of the ash content of bog birch forest litter. Procedia Environmental Sciences 3: 99?104.
Nature of Acidic Properties
of Litter in Bog Birch Forests
Tamara T. Efremova and Stanislav P. Efremov
V.N. Sukachev Institute of Forest SB RAS,
50/28 Akademgorodok, Krasnoyarsk, 660036 Russia
Different litter types (strongly decomposed, average decomposed, rhizomatous, peaty, peatified and
peaty) characterized by acid and heavily acid reaction of environment, have been formed in the
ecological series of bog birch forests beginning from the tall herb to the sphagnum- dead cover forest
types. The key factor of differences among sections of this series is moistening. Relationships between
total potential acidity of litter and humic fractional and group composition and ash substances,
# 218 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
including calcium and iron oxides, were studied using correlation and regression analyses. Fulvoacids
of the third fraction, i.e. products of the initial section of detritus humification, were significantly
correlated with the total acidity, hence, these compounds may be considered as acidic-forming. ?mong
factors which regulate (neutralize) the acidity of environment, the preferential role of calcium has
been revealed. It was shown that the most contribution of ash substances, including calcium, to litter
enrichment is made by ground vegetation which biochemical selection takes place due to forest type
succession. Finally, it is the soil moisture, having an impact on species composition of lower vegetation
layers, that is indirectly a prime cause of formation of acidic litter properties. Plant debris of birch
forests makes only a basic background.
Keywords: acidity, litter, bog birch forests, regression analysis.
? ?????????????????? ??????????? ?????? ?????????. ???????? ????????? ??????? ?????? ??
???????????? ????????????? ????????????
?????? ????, ????????????? ?????????????
??????????? ? ?????? ???????????? ???-
*
???????, ?????????? ?? ???????????????? ????????? ? ??????? ?????????? ??? ????????????? ????????????? ????????? ??????????.
????????? ???????? ????? ????????????
????????????? ??????????? ??????? ????????????? ??????? ????????????. ???? ?????????? ? ?????????-???????????? ?????-
© Siberian Federal University. All rights reserved
Corresponding author E-mail address: efr2@ksc.krasn.ru
# 205 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
???, ??????????? ? ???????????? ???????
?????????, ???????????? ????????????? ?????????? ????????????? ????????? ? ???????
???? ???????? ?? ????????? ???????? ?????????, ???????????? ????????? ????????????
???????????????? ?? ?????? ??????? ? ??????????. ? ??? ??? ???? ??????? ??? ???????
????? ????????? ? ???? ?????????? (????????, 1973; ??????????, 1983; ?????????,1983;
Luo, Christie, 2001; Kakei, Clifford, 2002; Batty,
Younger, 2007 ? ??.). ?? ??????? ??????????? ??????????? ? ???????????? ???? ?????
???????? ????????? ???????? ??????, ????
??????? ? ???????????? ????????? ??????
???????????? ?????????? ??????? ???????
?????????? 27,4 % (??????? ? ??., 2005; ???????? ? ??., 2009, 2010?, 2010?; Sekretenko et
al., 2011).
?????? ????????????? ???????????
???????? ????????? ?????? ?? ???????????????? ????????, ??????? ? ???????????? ???? ????????????? ?????????????????? ? ?????????????? ??????????
????????????? ?????????, ??????? ??????????
?????? ? ?????? ???????? ????????, ??????????? ?????????? ?????????, ?? ???????????
?????????, ????????????-?????????????????
????????? ? ?????? ???????? ?????, ?????????
? ???????? ????? ?????????? ?????? ?????????????. ?????????????, ???, ????????????
??????????? ?????????, ?????????? ????????? ?????????? ? ????????????? ????????????? ? ??????????????? ? ????????.
?????????? ??????????? (??) ???? ????????????? ?? ?????????? ????? ?+ ? ?????? ????? ??????????? ??????, ??????????????? ?
???????? ??????? ? ???????? ?????? ? ???????? ?????????? ????????? ???????, ?????????? ?????????? ??????????. ??????????????? ??????????? ?????? ? ????????????
???????? ???????? ? ????? ?????? ?????????
???????????, ??????? ???????? ? ????????-
??? ? ????????????? ??? ?????????????? ??
????????? ??????????. ??????? ??????????????? ???????????, ???????????? ? ??????? ???????? ???????? ????????? (?? 8,3),
???????????? ????????????? ??? ??????????
????? ????????????? ???????????, ??????????????? ????????? ?????????? ???? ????????? ??????????? (?????????, ?????????,
2006).
???? ?????? ?????? ? ????????????????
??????????? ????????? ???????? ??????????
? ???????? ??????????????? ????, ??????????? ?? ????????? ???????, ??????? ???????
??????? ???????????? ????????? ???????,
?????????? ??????? ??????????? ?????????? ? ??????????-?????????? ??????? ????????????? ???????? ??? ????????? ???????????????? ????????????? ?????????????
?????, ?????????? ?????????. ????? ??????
???????? ? ????????? ??????? ???????? ??????????? ?? ????????? ?????? ?? ?????????
??????? ????????? ? ???? ????????? ???????? ?????????? ? ?????? ?? ?????????????
????????.
????? ?????,
????????? ? ??????
???????? ???????????? ????????? ????????????? ????? ?????????? ??????, ?????????????? ? ??????????? ??????? ????????
??????. ?? ?????????? ??? ? ???? ??? ????
?? ????? ??????? ???????? ???????? ?????
???????? 2,3 ???. ??. ? ?????????? ???????
????? ?????????????? ???????? ???????????? ?????????????? ???????????????? ????,
?????????? ????????? ??????? ???????? ????? ?. ?????? (56°23?710???.?., 84°34?043???.?.) ?
??????? ????????????? ??????? ? 100?140 ?
?? ?????, ?? ???????? ??????????? ????????
?????? ?????????????? ?????? ????? ???.
?????????? ????????????? ????????????
??????? ?????????????? ????? ?????????-
# 206 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
??? ????? ?????? ? ?????????? ??????? ??????? ????????? ? ?????????? ???????????.
???????? ???????????? ????? ??????????
?????? ???????? (Betula pubescens Ehrh.),
??????????? ? ?????. ?? ???? ????????
?? ????????? ????? ?????????? ???????
????????-????????? ???, ????????? ?? ???????????, ????????????? ????????? ????
(???. 1), ??? ????????????? ???????????????
????? ???????????? ???????????? ???????????. ????? ???????? ???? ??????????????
?????? ???????? ??????????, ????????? ????????????? ??????? ?? ????? ??????????????
????? ?????? ????????? ??????? (270Ч60 ?). ?
??? ???????? ???????? ????????? ???? ????:
0?30 ? ?? ????? ? ?????????????-???????????????????? ????????, ?? ??????????
30-100 ? ? ??????????????????????, 100140 ? ? ?????????-???????? ?????????????,
140-230 ? ? ???????-???????-?????????, 230270 ? ? ???????-????????-???????????????.
??????????? ????????? ???????????? ??
????????????? ? ???????? ?? ???????????
??????????? ? ??????? ???????? ??????????????????? ?????????? (????????????, 1977;
??????????, 1984; ????????? ? ??., 2004). ?
??????????? ???????? ?????????? ?????
?????????? ???????????????? ???? ????????
????? ????? ?????? ?????????: ???????????????????, ???????????????????, ?????????????????? (???????????), ???????????,
???????????? ? ????????. ????????? ????? ???????? ?? 3,6 ?? 6,2 ?? ? ??????????
?????? ?????????? 0,047?0,079 ?/??3. ????
????????? ??????????, ??? ???????, ? ???????????? ??????? ??????????????? ???????,
????????? ? ????????? ??????????. ??????????????????? ??????????????? ?????????
(? ?????????????? ? L, F, H) ???????????? ??
????????, ??????????????? ??????????? ?
????? ?????????????? ????? (0?30 ?). ??????
????????? ??????? ???????? ????????? ??????????. ??????????????????? ?????????
???????? ???????????????? ???????????
(?) ???? ????????????. ???????????, ???????????, ???????????? ??????? ??????????????? ???????? ? ??????????????
??????????????, ???????? ? ?????? ????????? ????. ????????? ??????????? ? ?? ???????
???????????? ???????????. ? ??????- ? ??????????????????? ?????????? ????????????? ?????? ????? ???? (0?100 ?? ?? ?????)
?????????
??????????
40
40
30
30
20
20
10
10
0
??????? ??????????, ??
???????? ?????????, %
50
0
0-30
30-70
70-100 100-140 140-190 190-230 230-270
????????? ?? ????? ?????, ?
???. 1. ??????? ?????????? ? ????????? ????????? ???????? ?????????? ? ???????? ??????????????
??????? (??????? ???????? ?? ?????? ?????? 2005?2007 ??.)
# 207 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
226
242
217
215
235
0-30
30-70
70-100
100-140
140-190
190-230
230-270
??
??
??
??????? ???
219
???????
25
?????
0
230-270
236
50
190-230
86
50
????????
75
140-190
97
77
100
100-140
100
125
0-30
150
0
150
2
206
200
???????? ?????, ?/?
?????????? ?? ????? ?????????????? ?????, ?
?????????? ?? ????? ?????????????? ?????, ?
??
250
?????? ???????????, ??/??
0,93
??????
8,28
0,75
?????????
0,71
?????
??
?????????
4,47
5,02
0,65
???????????
4,47
0,28
??????? ???
235
205
7,77
4,56
196
200
????
???
150
120
100
50
46
30,0
32,7
26,2
0,25
3,72
3
50
38
5,0
5,4
4,8
0
6
9
12
0-30
0
15
??????????? ??????????, %
70-100
????????? ???
17,6
100-140
????????
230-270
12,24
1,1
190-230
1,72
???????
140-190
??
30-70
????? ?????, ?/?
2
175
????????? ???
250
70-100
269
30-70
280
277
??
????????? ????
???????-??????? ????
300
?
??????????
?
?????????? ?? ????? ?????????????? ?????, ?
???. 2. ????? ?????????? ????? ? ?????? ???????-???????? ??????? ? ????????? ??????? ??????????????
??????? ???????? ?????????? (?), ???????? ??????????? ????? ???????? ? ??????? ????????????
??????? (?), ?????????? ??????? ? ????????? ? ??????????? ????? ???????? (?), ????????????????
??????????? ???? ? ?????? ??????? ? ????????? ? ??????????? ???????????? ???????? (?)
?????????? ???????? ????, ??????????????? ?????? ???????, ?????????, ???????????
(???. 2?, ?). ? ??????????? ? ?????????????????? ????????? (100?140 ?) ??????? ??????
? ???????????? ????????????? ??????? (???????, ????) ?????????????? ????????????.
??????????? ? ???????????? ????????? ??????? ??????????????? ?????????? ?????????. ????? ????????? ?????????-????????????
?????????????? ????? ????????? ????????? ?
?????? (???????? ? ??., 2010?).
???????????? ??????? ????????? ? ???????? ?????????????? ???? ??????????? ??
?????????? 2004?2007 ??. ??? ?????? ??-
?????? ?????????? 10 ????? ??????????? ??
5 ?????? ? ????? ?? ????????? ????? ? ???????? ??????? ???? ????. ????????? ???? ?????????? ?? 70 ??????????????? (100Ч100 ??),
????????????? ????? 10 ? ???? ?? ????? ??
????? ?????? ???????? ?????????. ????????
???????????? ????????????? ??????? ???????????? ?????????? ? ???????? 50Ч50 ?? ?? 20
????????? ? ?????? ???? ??????????.
??????-?????????? ???????? ????????? ???????????? ????????????? ? ???????????? ???????? (?????????????? ??????
???????????? ????, 1975). ???????? ?????
????????? (??? ??????? ?? ???????) ??????-
# 208 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
?????? ? ??????? ???????? ??????? ??????,
????????? ? ????? ???????????? ?????????
? ?????? ??? 450 °? ? ??????? 6 ?. ????????
?? ?????? ??????? ???????? ??????????????????. ??????????????? (????? ?????????????) ???????????? ?? ?????? ???????,
???????? ??????????? ? ?? ?????? ?????????. ?????????? ???????? ???????????????
??????????? ???????? ?? 1,75 ? ???????
???????? ??????????? ?? ????????? ?????????? ?+. ????? ???????? ????????? (Ca2+
+ Mg2+) ???????????? ?? ?????? ????????
??????????, ????? (???????) ??????????
??? ? ?????????????????? ??????? ? ??????????? ?????????? ????? ??????? ????????
????? ?????? ?????? ? ??????? ??????. ?????????? ????????? ?????????? ? %, ?.?. ??
100 ? ????????? ??? ????????, ??????????
?? ??????????? ???? ??? 105 °?. ?????????
? ??????????? ?????? ????????????? ???????? ???????? ?? ???????? ?.?. ????????-
??? ?????????? ?? ??????? ???? ? ???????
6 ?????. ? ????????? ??????? ?????? ?? ???????? ??????? ???????? ????????? ??????? ??????????? ?????????????? ?????????? 1N H2SO4. ????????????? (??-1, ??-2
? ??-3) ????????? ?? ???????? ????? ????????? ??????????????? ???????? ???????
? ????????? ??????. ??????????? ???????
???????????? (??-1?) ???????????? ??????????????? ? 0,1 N ???????? H2SO4. ???????
???, ?.?. ?????????? (1980). ????????????
????? ???????? ? ????? ?????????? ?? ?????? ??????? ? ?? ????????? ??????????????.
? ?????? ????????? ??????? ???????? ???
???????? ??????: ????????? ??????? (??)
? ????????????? (??), ??????? ???????????
??????????????, ?????????????, ?????????? ????????, ???????? ???????? ????????
? ??. ??????????. ?????? ?? ????? ????????
????????? ????????? ???????, ??????????
????????? ????? ????????? ??????? ? ???????????? ??????????? ??????, ?/??? ???????
???????????? ????????? ??????? ? ??????
??????? ???????????? ?????????????? ????????????? ????????? (???????? ????????,
?????????). ? ???? ??????? ?????? ???????
????????? ?????? (??-1 + ??-1) ???????????
??????????????? ????????? 0,1 N NaOH ??
??????, ?????? ??????? (??-2+??-2) ? ????????? 0,1 N NaOH ????? ???????????????,
?????? ??????? (??-3 + ??-3) ? 0,02 N NaOH
?????????? ? ??????????
???????? ???????, ????????? ?????? ?
???????????? ??-1? ?????????? ??????????? ??????? ?? ??????. ?????????? ?????
? ??????? ????????????? ???????? ????????
? ????????? ?? ????? ?????? ?????????????
???????? ? ?????????.
????????????????? ???????? ????????? ???????? ??????????????? ? ??????????????? ?????????????? ??????? ? ?????? ???????? STATISTICA 6.0.
?????????-???????????? ???? ????????? ???????? ?????????? ???????????? ?????
?????????????? ????? ??????????? ????????? ?????? ????????????? ????????: 4,5 ??/?2
(0?30 ?) > 3,7 (30?100 ?) > 2,1 (100?140 ?) < 2,5
(140?230 ?) < 3,1 ??/?2 (230?270 ?). ?????????
?????????? ??????????????? ? ???????????????? (????. 1). ?????????? ???????????
????????? ???????????: Fe2O3 ? 0,53?1,64 %,
Al2O3 ? 0,70?1,57, ??? ? 0,30-1,44 %, ????????
????????? (Ca+Mg) 33,54?86,6 ??-???./100 ?
???????. ?????????????? ??????? ?????????????? ?????????? ?????. ????????? ?????
???????????????? ??????? (100?140 ?) ?????? ???????????? ???????, ????????????
???????? ????????? ? ???????? ????????????? ???????? ?????????? ????? ????????? ??????????????. ? ??????????? ?????
?????????????? ????? ????????????????
???????? ????????? ??????????: ??????????
# 209 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ???????? ??????? ????????? ??????? ????????? ???????? ??????????
??????? 1. ??????? ???????? ?????????? ??????? ? ??????????-????????? ?????? ?????????????
???????? ????????? ???????? ??????????
??????????
????????????????? ???? ????????? ? ??????????
?? ????? ?????????????? ?????, ?
????????????
?????????????????????????????- ??????????????,
??????,
??????,
???,
?????,
?????,
100-140
140-190
190-230
230-270
0-30
30-100
?????????? ????????, % ?? ????? ?????????
?????????
15,73
???
1,44
Fe2O3
1,64
Al2O3
1,57
??+?g, ??-???./100 ?
????? ??????????
14,86
10,26
8,03
7,27
8,52
1,15
0,67
1,24
0,93
0,46
0,32
0,30
0,92
0,66
0,53
1,34
0,77
0,70
0,89
1,10
86,62
30
71,26
54,09
44,68
44,3
33,54
30
30
15
15
30
????????? ? ??????????? ?????? ????????????? ????????, % ?? ?????? ???????? ?????????
????????? C/N
18,3
22,37
27
30,22
32,05
40,24
15,77
11,89
10,26
11,85
8,75
6,33
????????? ???????:
??-1
??-2
???
???
???
???
???
???
??-3
9,07
8,36
8,03
8,04
7,7
7,42
????? ??
24,84
20,25
18,28
19,89
16,45
13,75
??-1?
3,011
2,93
3,51
2,75
3,27
2,97
??-1
13,43
13,41
12,45
10
12,59
11,78
??-2
3,0
2,65
3,22
3,44
3,64
2,97
?????????????
??-3
5,12
5,23
5,51
6,03
5,78
6,02
????? ??
24,66
24,22
24,69
22,24
25,28
23,74
??-1+??-1
29,2
25,3
22,71
21,85
21,34
18,11
???+???
49,5
1
44,47
0,84
42,96
0,74
42,13
0,89
41,73
0,65
37,49
0,58
5
5
5
5
5
5
??? /???
????? ??????????
????????? ????????, ?????????, ??????????
??????? ????????????? ????????, ????????
?????????. ?????? ????????? ??????? ??? ?????????? ????????? ? ???????? ????? ??????????.
??????? ??????? ??????????????? ?
???????? ?????????????? ?????????????
???????? ?????????. ? ??????? ???????????? ???????????? ???????? ??????????????? ??????? ?? ??????????? (???+???) ?
37,5?49,5 % ? ????????? ?????? ???????? ?
?????? ????? (C/N), ??????? ? ??????????
??????????? ??????????? ?? ???? ??????????? ? ????? ???? ? 40,24 ?? 18,3. ? ???????
????????? ?????? ???????????? ???????
??-1, ?????????? ??????? ???? ??????????
? ??
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