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- . ..

14.03.06 ,

:

..

-
2014

.
.. 5
1.
( ) ..

13

2. .. 40
2.1. 40
2.2. .. 41
2.3. . 43
2.4. 45
2.5.
47
2.6.
...

49

2.7. - ... 50
2.8. . 51
2.9. . 52
2.10. 53
2.11. ...

54

2.12. . 55
2.13. ,
56
2.14. .. 57
3. . 59
3.1. ... 59
3.1.1.
.. 59
3.1.2.
() 61

3.1.3.
. 62
3.1.4. . 64
3.1.5.
65
3.1.6.
.. 66
3.1.7. . 70
3.2.
.. 71
3.2.1. .. 71
3.2.2. ,
CCl4 ... 75
3.2.3. ,
CCl4 ... 76
3.2.4.
, CCl4 ... 82
3.2.5.
,
CCl4 .... 85
3.2.6.
, CCl4 ... 89
4. 98
5. . 120
6. 123
7. - 125
8. 126


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57

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59

3.
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60

2
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0,9% NaCl
()

4,3 /

8,6 /

13,4 /

17,02,2

6,70,9

4,70,6

3,80,5

1,00,1

24,03,1*

8,01,0

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25,03,3*

5,30,7

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61

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62

4
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0,9% NaCl
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4,3 /

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13,4 /


()


()

0,40,1

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10,21,3

6,20,8

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63

3.1.4.
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65,88,6

77,310,1

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65

6
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0,9% NaCl
18,52,4
369,518,0
2,80,4
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262,747,2*
12,11,6**
4,3 /

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353,345,9
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66

, (, , ..).

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5- , (
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67

7
(4,3-8,6-13,4 /) (1 /)

0,9% NaCl
()

4,3 /

8,6 /

13,4 /
0,9% NaCl
()

4,3 /

8,6 /

13,4 /
0,9% NaCl
()

4,3 /

8,6 /

13,4 /
0,9% NaCl
()

4,3 /

10

1-
16112
14716
(+10%)
12215
14821
(-18%)
11320*
15812
(-28%)
16018
16521
(-3%)
2-
15212
14323
(-6%)
13619*
18823
(-28%)
17221
16920
(+2%)
15713
17122
(-8%)
3-
16111
14424
(-11%)
15517
17023
(-34%)
23222*
18219
(+27%)
24422*
17619
(+39%)
4-
17111
15424
(-11%)
12126*
18217
(-34%)
19123
24618*

0,210,02

0,200,01

0,170,01

0,230,03

0,210,04

0,190,01

0,140.4

0,180,02

0,340,03

0,260,05

0,190,06

0,160,02

0,140,3

0,160,02

0,210,12

0,170,02

0,140,03

0,180,06

0,210,02

0,200,01

0,250,04

0,140,03*

0,200,09

0,180,5

0,160,03

0,180,06

0,150,02

0,210,03*

0,190,05

0,210,03

68

8,6 /

13,4 /

20923

0,9% NaCl
()

4,3 /

8,6 /

13,4 /

14222

0,9% NaCl
()
1 /

16617

4,3 / +
1 /

8,6 / +
1 /

13,4 / + 1 /
0,9% NaCl
()
1 /

4,3 / +
1 /

8,6 / +
1 /

13,4 / + 1 /

20531
21113
22821

18519

(+15%)
24119
(+15%)
5-
15617
(+10%)
17024
(-17%)
24424
(+16%)
25515
(+12%)
6-
15522
(-7%)
25321*
(+37%)

0,130,01

0,190,04*

0,130,04

0,180,03

0,220,05

0,190,04

0,160,11

0,220,01*

0,200,04

0,220,01

0,150,01

0,130,04

0,190,02

0,130,04*

21725

27519*
(+27%)

0,250,03

0,140,04*

22019

28035*
(+27%)

0,210,11

0,190,10

20121

29217*
(+45%)

0,240,06

0,120,02*

0,160,01

0,140,04

0,180,02

0,120,04*

7-
17617
16522
(-6%)
18021
24319*
(+35%)
21230

28834*
(+36%)

0,240,02

0,220,04

21122

29426*
(+39%)

0,120,04

0,230,06

22115

30319*
(+37%)

0,260,05

0,230,06

. *<0,05 .

69

. , , ,
, .
3.1.7.
- ( , , -, , , ). 4,3-8,6-13,4 / (5-7 ).
, () ( , ). (13,4 /) ( ) . () , 13,4 /. -
() .
. (4,3-8,6-13,4 /) . 13,4 /
.
. -

70

. , , ,
,
. , ,
. , (-) .

71

3.2.

3
(2,2 /, 4,3 / 11,6 /) (14 , 1
) :
, ,
-, ,
.
3.2.1.
0,4 / 50%- l4 ( 1000 / l4) 1- 4- . , , l4 40%, 50%
(50). l4 . , 2000
/ l4 .
, 16- ,
: , , ,
. , , l4, 2
( 123,114,4 248,610,4 /), 4 ( 22,89,3 88,749,7 /), - ( 1708,840,9 1930,835,4 /), - ( 2,330,83 0,980,97 /), , , .

72

( ), :
- ;
;

.
. 1 (, ) (. 12).
,
,
,
. , ,
, , .
, .
2 (, CCl4 + )
-
, 1/3 2/3
, , (. 13).
, .
2-3 -

73


().

. 12. , ()
. . 200.
. 13. , CCl4 +
()
,
(). .
200.
, CCl4 2000 /
( ),
15 . .

74

3.2.2. , CCl4
CCl4 2000
/ , , 50%
(. 8).
8
, CCl4

(%)

1. ( )
2. CCl4 + .

10

0 (0%)

14

7 (50%)

3. CCl4 + 2,2 /

12

5 (42%)

4. CCl4 + 4,3 /

14

6 (43%)

5. CCl4 + 11,6 /

14

7 (50%)

6. CCl4 + 300 /
( )
7. CCl4 + 300 /
( )

14

6 (43%)

10

10 (100%)

2,2-4,3-11,6 /
, CCl4.
42-50%.
CCl4, 2- 5- . 300 /, , . 300 /, , (100%).
7- . .

75

3.2.3. , CCl4
3.2.3.1.
,
CCl4 (2000 /) 15
(. 14). , , CCl4, ( 3 ) ( 2,5 ) ,
( 5 ) ( 2 ). .
(14 )
,
2,2 4,3 /, 11,6 /. ( ) ( ) ,
. . 300 / .
, , , ()
, CCl4. (2,2 /) (4,3 /) .

76

. - + CCl4

45

. - + CCl4

40

0.5 + CCl4

35
30
25
20
15

0.5 + CCl4

0.75 +
CCl4

1.2 + CCl4

0.75 +
CCl4

1.2 + CCl4

10

- + CCl4

5
0

+
CCl4

- + CCl4

1
0
1

. - + CCl4

10
9
8
7
6
5
4
3
2
1
0

0.5 +
CCl4

10

0.75 +
CCl4

8
0.75 +
CCl4

1.2 +
CCl4

4
1.2 + CCl4

- + CCl4

. - + CCl4

14
12

0.5 + CCl4

+
CCl4

- +
CCl4

2
0
1

+
CCl4

. - + CCl4

2,5

0.5 + CCl4

0.75 +
CCl4

1,5
1

1.2 + CCl4

0,5

- + CCl4

0
1

+
CCl4

. 14.
, CCl4
3.2.3.2.
,
. ()
() ,

77

.
() , () .
CCl4 (2000 /) 14

(. 15). , CCl4, ()
2,5 , 4 ,
10 .
. (2,2-4,3-11,6 /),
() .

. - + CCl4

120

0.5 + CCl4

100
80

0.75 +
CCl4

60

1.2 + CCl4

40

- + CCl4

20
0

+
CCl4

. - + CCl4

12

0.5 + CCl4

10
8

0.75 +
CCl4

1.2 + CCl4

- + CCl4

2
0

+
CCl4

. - + CCl4

25
20

0.5 + CCl4

15

0.75 +
CCl4

10

1.2 + CCl4

- + CCl4

0
1

+
CCl4

. 15.
, CCl4

78

, 14
.
3.2.3.3.
-,

.
CCl4 (2000 /) 15

- . (.
16). (2,2-4,3-11,6 /) 14

,
. ,
.
, -
,
,
( ), .

79

. - + CCl4

200
180
160
140
120
100
80
60
40
20
0

0.5 + CCl4

80

. - + CCl4

4,5
4
3

0.75 +
CCl4

2,5
2
1

- + CCl4

0,5
0
1

- + CCl4

20
0

+
CCl4

. - + CCl4

2,5

0.5 + CCl4

0.75 +
CCl4

1,5
1

1.2 + CCl4

0,5

- + CCl4

1.2 + CCl4

1,5

1.2 + CCl4

40

0.5 + CCl4

3,5

0.75 +
CCl4

60

- + CCl4
+
CCl4

0.5 + CCl4

100

1.2 + CCl4

. - + CCl4

120

0.75 +
CCl4

+
CCl4

+
CCl4

. 16. , CCl4
3.2.3.4. ,

.
.

.
CCl4 (2000 /) 14
4
,
(. 17). (,
)

80


. (2,2-4,3-11,6 /)
,

. 300 /
CCl4, .

. - + CCl4

120

0.5 + CCl4

100
80

0.75 +
CCl4

60

1.2 + CCl4

40

- + CCl4

20
0

+
CCl4

. - + CCl4

300

0.5 + CCl4

250
200

0.75 +
CCl4

150

1.2 + CCl4

100

- + CCl4

50
0

+
CCl4

. - + CCl4

80
70

0.5 + CCl4

60
50
40

0.75 +
CCl4

30

1.2 + CCl4

20
- + CCl4

10
0
1

+
CCl4

. 17. , CCl4
, 2,2-4,3-11,6 /
CCl4

. ,
.

81

3.2.4.
, CCl4
,
, (. 9), CCl4
( ) . , CCl4 2,2 / (p < 0,05). -,
CCl4 .
,
CCl4, ,
,
-
(p < 0,05).
,
,
(, , ).
, CCl4,
. ()
CCl4 , , -

82

9
,
CCl4,

,
/
,
/
,
/
,
/
,
/
-, /
, /
,
/
, /
, /
, /
, /
, /
, /
,
/
, /
, /
-, /

(.
)

l4 + .

l4 + 2,2
/

l4 + 4,3 /

72,452,0

70,421,47

65,182,08*

70,140,65

4,850,33

6,340,46*

4,180,44

5,400,21

1,870,11

1,660,09

1,580,04

1,870,13

45,210,95

44,341,10

43,822,47

44,081,08

8,130,27
1708,8340,89

8,140,38
1930,835,4*

8,520,43
2001,654,6*

8,840,35
1719,6144,3

22,769,32

88,7449,68*

133,6850,73** 1659,86156,75**

0,270,008

0,340,03

0,390,02*

0,430,02*

20,03,5
123,114,4
95,93,4

11,241,35*
248,6210,94*
99,928,23

7,661,49**
213,5417,81*
95,226,45

12,881,95*
198,5219,97*
119,5618,30*

14,21,5

7,170,61*

6,080,98*

9,061,13*

531,4540,15

437,863,5

656,1287,95*

604,4849,97

320,049,75

165,417,2*

159,619,55*

214,215,3*

4541,0527,9
3,110,13
0,190,04

5183,4402,80
2,620,12*
0,120,01*

5358,8354,4
2,120,07*
0,170,01

5315,8520,0
2,430,10*
0,180,02

2,330,83

0,980,97*

2,400,84

1,560,90*

. *<0,05; **p<0,01 .

83

9 ()
,
CCl4,

,
/
,
/
,
/
,
/
,
/
-, /
, /
,
/
, /
, /
, /
, /
, /
, /
,
/
, /
, /
-, /

l4 + 11,6 /

l4 + 300 /
()

(.)

l4 +
.

72,452,0

70,421,47

67,40,91

74,161,22

4,850,33

6,340,46*

4,640,72

5,580,48

1,870,11

1,660,09

1,810,06

1,910,17

45,210,95

44,341,10

44,983,17

58,5213,11

8,130,27
1708,8340,89

8,140,38
1930,835,4*

7,650,61
1989,07,97*

11,062,79
1424,2234,16

22,769,32

88,7449,68*

0,270,008

0,340,03

0,370,01*

0,370,03*

20,03,5
123,114,4
95,93,4

11,241,35*
248,6210,94*
99,928,23

6,800,58**
136,8047,79
99,385,89

14,842,21*
212,1211,68*
102,704,79

14,21,5

7,170,61*

6,060,50*

47,7625,16**

531,4540,15

437,863,5

528,8240,58

505,8103,4

320,049,75

165,417,2*

169,812,16*

271,4038,41

4541,0527,9
3,110,13
0,190,04

5183,4402,80
2,620,12*
0,120,01*

5036,80445,16
2,150,48*
0,180,01

4589,20721,59
2,580,12*
0,130,01*

2,330,83

0,980,97*

0,500,30**

1,521,11

1315,52253,22** 1838,90140,54**

. *<0,05; **p<0,01 .

84


(11,6 /), ( ), . , ,
/
,
. -
.
, (2,2-4,3
/) (300 /) -, .
3 ,
.
, , , 11,6 / 300 /
.
3.2.5.
, CCl4
(2,2-4,3-11,6 /) CCl4.
3.2.5.1. 2,2 /
, , CCl4 + 2,2 / (
3), -

85

- , 2/3 ,
, , .

. 18. , CCl4 + . ( )
,
. .
200.
. 19. , CCl4 + 2,2 / ()
,
. . 200.
. 20. , CCl4 + 4,3 / ()
,
. .
200.

86

. 3 (. 19).
3.2.5.2. 4,3 /
, CCl4 + 4,3 /
, . , -
, 1/2 3/4 .
,
.
23 (. 20).
3.2.5.3. 11,6 /
, CCl4 + 11,6 /,
, . ,
- ,
1/3 1/2 . , . 2-
(. 21).

87

. 21. , CCl4 + 11,6 / ()


, . . 200.
. 22. , CCl4 + 300 / ()
, . .
200.

3.2.5.4. 300 /
, , , CCl4 + 300 /,
. , - 1/3 , 2/3 . ,

. 1- (. 22).
, CCl4 (2000 /)
().
, ( -

88

),
().

,
11,6 /, .
3.2.6.
, CCl4
, , CCl4,
,
.
() 3 ( , ),
(), ( ) ( ) .
. 23.

89

5-

. 23.
(), (), (), 3,4-
(), (), 5- (5) 6
(),
().

. 24 ( ), . 25 () . 26
( ).
: () 3-
(3-) 3,4- () ().
: () 5--3 (5-).

90


5-

. 24.
, . 23.

5-

. 25.
, . 23.

91

5-

. 26.
, . 23.
, ( )

5-

(5-) , , (). , () / ( 3 4, =0,0009 =0,0457, ).


, CCl4 .
2,2 4,3 / ( 2 )
( , 3 4 1, < 0,01, 2, < 0,05 ). . 27.

92

DA ACB

DA

DOPAC

9
8
7
6
5
4
3
2
1
0

HVA

1.5

1.0

* *

1.0
0.5
0.5

g1

g2

g3

g4

g5

g6

0.0

0.0
g1

g2

g3

g4

g5

g6

g1

g2

DOPAC/DA

g3

g4

g5

g6

g5

g6

HVA/DA

0.25

0.125

0.20

0.100

0.15

0.075

0.10

0.050

0.05

0.025

0.00

0.000

g1

g2

g3

g4

g5

g6

g1

g2

g3

g4

. 27.
(/ );
( )

/ ( 3 4 1 2, =0,0412 =0,0390, ).
2,2 4,3 /
( < 0,05), . 28.

93

DA STR

DOPAC

DA

HVA

1.5

10.0
7.5

1.0

* *

5.0

0.5
2.5

0.0
g1

g2

g3

g4

g5

g1

g6

g2

g3

g4

g5

g6

0.0
g1

g2

DOPAC/DA

g3

g4

g5

g6

g5

g6

HVA/DA

0.2

0.2

0.1

0.1

0.0

0.0
g1

g2

g3

g4

g5

g6

g1

g2

g3

g4

. 28.
(/ );
( ) .

, , (
)
( ).
,

/ /

94

,
, /
- ( . 29).
( ),
.

. 29.
,
,
,
5- (. 30). ,

5-/

95

( 6 2, =0,0157).
( < 0,05) 5-/
, , .
( ) .
( .. ., 2006), , , .
SER Frontal cortex
SER

HIAA

2
1
1

0
g1

g2

g3

g4

g5

g6

g1

g2

g3

g4

g5

g6

HIAA/SER

0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
g1

g2

g3

g4

g5

g6

. 30.
(/ );
( ) .

96

, ,
.

97

4.


4,3-8,6-13,4 /

5-7

.
( , -,
,
)

,
( .. .,
2002, 2004, 2006, 2008; Salamone JD, Correa M., 2012).
.

, ( .., 1990; .., 2003,
2009; Curtis J.T., Wang Z., 2005; Comai S, et. al., 2012). , ()
,
(Bjrklund A., Hkfelt T., 1984;
.. ., 1996; .., 1998; Lieu C.A, Subramanian T.,
2012): 1) ( 9, ); 2) (
10, ,
, , , , , ); 3) ( 10, -

98

, 9, , , , ) 4) ( 12-14,
, , ,
). - ( .., .., 1992;
.. ., 1999, 2000, 2002; Amano T. et al,. 2012 ).

1972 . 1978 .
, ,
, ,
35- (), . , , ,
, , ,
- (Kebabian J., Calne D., 1979). . . (Kebabian J., Calne D.,
1979), , D1, , D2,
.
, , . .
, : D3, D4 D5/1b. ,

.

99

D1, D1 D5/1b , D2, D2, D3 D4 (Missale C. et al., 1998). D1b


, D1
, D5 (de Manzano . et al.,
2013).
, , (Febo M., Ferris
CF., 2007). : 1) ( ), ,
, () (Cools R., 2011) 2) -
( ),
, , , (Sagara H. et al., 2010). ,
,
().
, ,
,

,
( .., 2006; .., .., 2012;
Gderqvist S. et al., 2012). 1920- .,

100

() ,

( .., 1987, 2004;


. ., .., 1989; .. ., 2002; .., 2005;
Gderqvist S. et al., 2012). -, ,

, . -,
( ). , ,
-, ,

.
,

( .., 2007; Aragona B.J.et. al.,


2007)
,

,

. ,

, (. 31).

101

. 31.

, .

, ,
(Carlezon WA, Jr, Thomas MJ., 2009).
, ()
, ,
.

( . ., 1987; . ., . ., 1998; Morikawa H,


Paladini C.A., 2011). (, , )

(Morava E.A. et al., 2011)

, .

102

. . ).


( ., 1970;
.., 2008; Cagiano R. et al., 2008).

( . .
., 1988; . ., . ., 1998; .. ., 2004;
DeLeon K.R., 2005; Glimcher P.W. et al., 2011).

1950-


() ( , , , ). .
, (
) , , 5000 1 200 20
, , ,
( . ., 1987; . . ., 1988; Nathaniel T.I.,
2009). ,
,
,

,
, , .

103

. ,

. ,

(-)

, ,
, ,
- ,
, , ,

, - .

.

- .

, ,

,
(, ) . ,


( . . ., 1988; . ., . ., 1989; Blum K.,
2012). 1980- .



(
,

. .).

(-, )

104

. (
) (), , ,
,

. .,

. ., 1998 Blum K., 2012).


, .. (
.. ., 2002, 2004, 2006, 2008, 2011)
,

(
).

,
, , .


,
.
.

(), , , , , .

(Van den Oever M.C. et al., 2012)


( . ., . ., 1989; Lebedev A. A. et al., 1993; LeMoal M., Simon H., 1991; Panchenko G. N. et al.,
1998; McBride W. J., Murphy J. M., Ikemoto S., 1999; Koob G.F., 2011)
,
-

105

.
-

, , ,
( .. ,
2006) (. 32).

. 32.
( .. ., 2002)


D1 D2-
(Petrov E.S., Lebedev
A.A., 1997; Panchenko G.N. et al., 1998; .. ., 2002).
,

(Tritsch N.X., Sabatini B.L..,


2012).

, , ,

106

( . ., . .,
1994; Aragona B.J., Wang Z.., 2007: Guarraci F.A., 2008).
, ,

( .., 1990; Petrov E.S., Lebedev A.A., 1997).

: ,
(). ,

- ()
( .., .., 1989; Petrov E.S., Lebedev
A.A., 1997).
,
. . (1987),
,
. -,
. .. (1987)
( ),

. ,
(
) .
,

..

..

(1997)

,
, D1- SCH23390
(0,05 /) (1 /)
. SCH23390
(Petrov E.S., Lebedev A.A., 1997).

107


( .., .., 1992, .. ., 1995).
,

,
.
- ,

0,5 5 /
. 1 /
35-40%.
() ()


.
( . ., . ., 1992). ,
(1 /)
,
, , (5 /)
,
.

, :

- ,

( ..,
.., 1989; Burbassi S., Cervo L., 2008).

108


( .., 1983; .. ., 1988).
,

(Alcaro, A. et al., 2007).


,
, .
, ,
,
,

( .., .., 1989;


.., 1991; Mameli M., Lscher C., 2011; Parkes S.L.,
Westbrook R.F., 2010). -,

,

(Bardo MT, Bevins RA., 2000). ,

, c
( .., .., 1989; ..,
.., 1992; .., 2001; Mameli M., Lscher C., 2011).
,
-,
. ,
( ,

109

),


.
(-,
-,

-,

-,

( .., 2005).

(Balfour DJ., 2009; Koob G.F., 2011).
,

.

,
( .. ., 2002, 2004, 2006, 2008, 2011; ..,
2012; Tritsch N.X., Sabatini B.L., 2012; Gorwood P. et al., 2012).
, , .
()
( , ).
, ( ),
(Avena N.M., Rada P.V., 2012).
, , (13,4 /) ( ) -

110


( , ,
, ). ( )
.
( , , ,
).
, () , 13,4 /.
, ( , ,
, ).
, ( .., 2008; Dalley J.W., Roiser J.P., 2012).

,
() . 13,4 /

.
(4,3-8,6-13,4 /) . 13,4 / .
.
, ( ).

111

.
. , , ,
, .
, ()
.
,
( ), , , ( 2
, , 4,3 /), ( 8,6 13,4
/ 3-5 ). , -,
, ( , ).
, (-) (Carboni E. et
al., 2004)

(, , ), - - , - ( .., 1988).
, , - ( ..,
2008). -

112

. 0,5 / D2 ,
, ,
. 5 / ( 10 )
D1- D2-
( .. ., 2002; Afonso V.M.et al. 2009; Pare D, Duvarci S., 2012),
( , , , ). , ( , 50-100 ,
) .
.
(),
, , (Adams D.B.. 2006). ,
. (
)
1-2 / 20-25 / ( ) ,
.
, (4,3-8,6-13,4 /),

,
, .

113

. - , . ,
, , ,
, , .
(, ) . .
()

.

, , ,
, , -, , ,
,

(),

(), , , , (), ,
.
(
) , 2-
( 2000 /). CCl4 ( ) 50% , . , 2 , ,

114

, , . () , (),
- -,
, .
, 16- ,
: , , .
, , l4, 2 , 4 ,
-, -, , , .
( ), - ,

. , CCl4,
-,
1/3 2/3 , , . , .
2-3- ().

115

CCl4
,
, ,
.
.. (2006),
, ,
, 5 (1 /). .

2,2-4,3-11,6 /
300 /
14
.


. ,
, CCl4,

,

. ,
(2,2 /)
(4,3 /) ,
(11,6 /).

(), 300 / (!).
, ,
(

116

) ,
CCl4, 100%-
7- .
,
, 2,2
4,3 /, .
.
( 11,6 /)
( )
(), .
,
( ),
CCl4
.
,
(.
3.1). , (4,3-8,6-13,4 /)
, ()

).

),


.
,

CCl4.

117

, , (13,4 /) ( )
. (
) . ,
( ).
, () , 13,4 /.
, ( , ,
, ).
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