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2004.

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3

.
.
-
: (1)
; (2)
(, 1962; Termaat and Munns, 1986).
, , .
,
, (Marcelis and Van
Hooijdonk, 1999; Fricke, 2002).
.
(Jackson et al, 1990) ,
(, 1988),
. , ,
, , ,
:
.

(Boyer et al, 1985; Tardieu et al, 2000; Hsiao et al, 2000).
(Fricke and
Peters, 2002).
( ,
, ,
) (Cosgrove, 1993).
,
(Caderas et al., 2000).
(Zhang and Davies, 1989;
, 1996), (Cleland, 1971;
1980; ., 1984). ,
( ., 1993; Munns et al, 2002). j.

, -

_ _ _ _ ,

\ : - . - -- - ' ^'

Copyright & A K-C

4
(Munns and
Cramer, 1996).
. ,
,
(Kudoyarova et al, 1998; .,
2002) (Vysotskaya et al, 2003). ,
.
, , , -
(, 1962; Rawson et al, 1988).
, ,
.

,
.

:
1.

.
2. -
(, , ,
),

.
3. (, )
, NaCl
.
.
4. ,
,
.

Copyright & A K-C

5
. ,


. ( 10 )
, ,
.
,
.

,
.
. ,
-
, ,

.

, ,
.
.
(Creete, 2002),
(Oxford, 2003),
( 2003, ).
. 8 .
. ,
, , ,
. 94 .,
4 18 .
*tCo ,

- ...
, ..., . ..

Copyright & A K-C

6
: ... .. , .. , . , ..
. - ... ..
.

.
(Hordeum vulgare) (Triticum durum L., 139) .
2 - 3 10"5
, 24 .
10 %-, - 100 %- -I 18000 14-
. ,
, .

100 .
.
(, 1997).
.
(Thomas et al., 1999). .

() .
() : =[( -
)/( - )] * 100%. ()
: =[( - )/ ]100%.
(100-) %.
-
.
,
,
, .

Copyright & A K-C

7

80 %- .


(Veselov et al., 1992). ,
, -18, 5 80 % .
80%- ,
: : (6:1:2). ,
.
- ( 7,4)
() (Kudoyarova
et al., 1998).
Vn= , Vn- (
/ ), - (/ ), V
- ( / ) (Jackson, 1993).

1. .

. 1, NaCl

.
.
. 60 ,
30-40 %

,
, . (Thiel et al., 1988),
. . (Passioura and Munns, 2000).

Copyright & A K-C

50 -j

30

1 20
2

10

i -
10?

; [ f Ilrtg^,
10

, ,

30 40 50 60 70 80 90 100110120 130 140

-20
-30
-40
,

> ,

40 50 60 70 80 90 100110 120 130 140

. 1. ()
() .
.
: (1)
, (2) , (3)
.
1.1. .
( ) ,
( 1),
(Munns et al, 2001).
. -

Copyright & A K-C

9
.
, . ,

.
(Thiel et el., 1988; Passioura and Munns, 2000; Munns et al, 2001),
(Tardieu et al, 1998; ., 2003)
(Chazen and Neumann, 1994; , 1999).
,
, .


,
(Jeschke et al, 1993).
NaCl , ,
, , , .
,
,
(Munns et al, 2001).
,
. (Hsiao et al.,
1998) ,
.
, ,
( ) .
, ,
,
. . (Thompson et al., 1997) . (Munns, 1993)
,
. . , , .

Copyright & A K-C

10
1.2. - .
, ,
,
, . (Thomas et al., 1999).
,
.
: = (Xi - Xo)/(Lgr ) ('1 "1), -
(/)- , Xi (/)-
, Lgr () - . -
, =9,8 /106 2 (/2= ), (), - (). 1, 40
NaCl
, .
1

, (^"1)
,

20

30

40

0,890,05

0,890,06

0,710,05

2,680,12

1,790,08

1,600,07

.
, , 60
, .
(Lynch et al., 1988)
,
. ,
-
.
/. 3. .
,
-

Copyright & A K-C

11

.

. 2,

.

I5 500

600 1

S
|

400

300

200-1
| 100
S

14

| ,'11

40

NaCI, 120

800

J
600

400
? 200 4 -

. 2. ,
() ()
() (6).
(S-4100,
Hitachi) (Link, Oxford)
() ,
( ), -
(. . 2.).

Copyright & A K-C

12
2
() ,

Na+

,
20
72
7

216

65

325

6919

414

86

38114

II1124

1526

1965

19518

19414

14818

17422

142111

18213

,
.

. (Fricke, 1997),
.
, (Matsuda and Riazi, 1981)
(Thiel et al, 1988).
, .
100 NaCl ( 170 /)
0,4 ,

(170 /). 2
, 20
. , ,
60 ,
.
1.4.
.
.
,

Copyright & A K-C

13
(
). 3, 10

, ,
,
(. 1).

*-""*->>*

, 5--

*-*-^" ~ *-*-

10

20

30

40

50

60

70

80

90

100 110 120

240

10

20 30 40 50 60 70 80 90 100 120
,

3. () ()
.
. ,
, ,
(. . 4).

Copyright & A K-C

14
140

8.

13

28

43

59

73

88

105

,
4.
.
, .
,
(.3). ,
,

.
,
, -
.
(.5).
5, 10
( [100-] %)
, ,
,
.

Copyright & A K-C

15

(Sohan et al, 1999). ,
,

S?

10
30
45
75
10
75
,
. 5.
() () .
/.5.

. .
,
. ,
, .
, ,
.
, ,
. ,
.
(, 1988; Caldewell et al.2001),

(

Copyright & A K-C

16
).
, ,
, ,
. ,
,
.

(Munns, 1986; Lynch et al, 1988). ,
, ,
.
** *
,
,
: ,
, , .
,
, . , -,

. ,
: (1) , (2)
(3)
(.. ). ,
.
.
2. .

,
, . NaCl

Copyright & A K-C

17

.
2.1. .

.
( ).

(. 6).
(. 36).
250 -,

NaCI, 10

Nad, 30

Nad, 1 2 Na<3, 2 Nad, 20

20

6.
, .

(Mansfild and McAinsch, 1995).
,
, ,
(Zhang and Davies, 1991).
(Munns et a), 2001).
.

Copyright & A K-C

18
,

(17,8+1,9 19,5+2,1 )
( ) , , . ,
.
, ,
(, ), .
, ,
.
,
. ,

. ,
, ,
. ? ,
(, 1989)
(Hsiao et al, 1998).
,
,
. ,
, (Wolf et al., 1990; Zhao et al., 1991; Thomas
et al., 1992; He and Cramer, 1996; Munns and Cramer, 1996; Gomez - Caderas et al
1998). ,
.
2.2.

(Wang et al., 2001).
, , ,
, , ,
(. 7, ).

Copyright & A K-C

19
, .

- (, 1979).
,
(Blatt, 2002). (. 2),

.
. ,

-.
D

[,#111,1
10

20
30
60
15
,

50

. 7.
() ().
,
(Cleland, 1971).
,
.
,
. ,
, , .

Copyright & A K-C

20
2.3. .

( , )
. , ,
.. .. . .
( ) ,
.
,
. .
8 9,
.

. .

NaCI,

NaCI,

NaCI, 1.

. 8. ,
.
, , .
(, 1969),
( )
, .

, ?

Copyright & A K-C

21
, ,
( . 1984).

NaCI, NaCI,

NaCI, 1.

9. ,
,

, ,

(Downes and Crowell, 1998).
, (
)
, .

. ,
(Bengston et al., 1979)
(Incoll and Jewer, 1987),
. . . (Blackman
and Davies, 1983) ,

. -

Copyright & A K-C

22

, .
, .
(, 1971; Gadalah, 1999) ,
,
. , . (Gadallah, 1999),
,


K+/Na+,
.
***
, ,
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, .
: (1) , , ,
, -
; (2) ,
, ; (3)
,
.

1.

,

.

2.

.
,
.

Copyright & A K-C

23

3.

( , ) ,
.

.

4.

.
-
.
, :

1. .., .., ., ..

.
- , 24-26 2001.,
.. , 2001..13-14.
2. Kudoyarova G., Dedov A., Veselov D., Mustafina A., Sabirjanova I., Akhiyarova G. Growth response of wheat seedlings to reduced water potential of nutrient
medium. Plant under environmental stress. International symposium. 2001, Moscow,
K.A. Timiryazev Institute of Plant Physiology, 2001, p. 145-146.
3. Akhiyarova G., Veselov D., Kudoyarova G., Fricke W. The effect of salinity
on growth and water relation of barley plants. Abstracts of 13th Congress of FESPP (2-6
September, Crete, Greece, 2002), p. 504.
4. .., ..
. 6- -
- 21 , 20-24 2002 ., . . , 2002.- ..
- . 186-187.
5. D.S.Veselov, A.R.Mustafina, I.B.Sabirjanova, G.R.Akhiyarova, A.V.Dedov,
S.U. Veselov, G.R.Kudoyarova. Effect of PEG-treatment on the leaf growth response and
auxin content in shoots of wheat seedlings. Plant Growth Regulation. 2002. V. 38. P.
191-194.
6. .. , .. , .. , . , .. , .. , .. , .. , . , .. .
. . 2002. 49. 572-576.
7. .., ..
. 7- -
21 , 14-18 2003 ., . . , 2003. .150-151.
8. W.Fricke, G.Akhiyarova, D.Veselov, G.Kudoyarova. Rapid and tissue-specific
changes in ABA and in growth rate in response to salinity in barley leaves. Journal of
Experimental Botany. 2004 (in press).

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171 450054, . ., 71.
848047 42 15 1996 .
5. 100 2004 .