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*
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E-mail: [email protected]
زرا ﻳ ﺑﺎ ﻲ مرژ رﻮﺷ ﺶﻨﺗ ﻞﺑﺎﻘﻣ رد مﺪﻨﮔ ﻢﺳﻼﭘ ي
ﺴﺣ ﻴ ﺘﺷد ﻦ
1ﻲ
،ا* دﺎﺑآ جﺎﺗ ﺪﻤﺣ ي
2رﻮﭘ ﺴﺣ ، ﻴ ﺷ ﻦ ﻴ ﻧاﺮ
3ﻲ ﻮﻘﻧ ﺎﺿرﺪﻤﺤﻣ و
4ي
)
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هژاو ﺎﻫ ي ﻠﻛ ﻴ ﺪ ي : K(! &
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.
ﻪﻣﺪﻘﻣ
!
"
#$ %&
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/0 1$ 2+!
)
Munns et al., 2002; Munns et al., 2003
.(
&67&
8
90$
;
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)
Colmer et al., 2006
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)
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Munns et al., 2000
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?
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)
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.(
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F#
'&
)
Poustini & Siosemardah, 2004
.(
/\*R &
&\ D>d e
& P;
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>
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?g (0 MB # Y?
cH+ D&
F 90$ _^ & D >;
& %&
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%) /?
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.(
'L
; >"0=
( /? DP; 9h 90$ c! <&=
0( D )& /+
*# D')& O^&#
)
& %& )&
'L ! /, 'L
B i &
&>? D?
/? 90$ cH+
#
"+!
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Flowers & Yeo, 1995
.(
_(
f D
j\?
QQ$ %&
? bL %& )&
0 M &
0 T)
&
)
Shannon, 1984
(
*# D , Q=&
>?
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&>? JKL& # 1H& kl!
90$
Q )& D
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P6 ( 7 6 ]*
2
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(Noble & Rogers, 1992)
.
_, Q& DP;
Na+
'0^ /? ! %&
( M&(
) _ ) O7 (
90$ ?
dZ0(
&
(Gorham
et al., 1987; Munns et al., 2006)
. /+ &>??
(
1. Stress Susceptibility Index (SSI)
>+? ?
Na+
#
Cl-
O7 & ! $ F
/? DbL & >A &> % D, D<% '@; ? S
P&& 20 (
&(
/?
% &Q
>, &B O0N
m>? 7 ? %& n=? # (
(Munns et al.,
2006)
.
&>?
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&(
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[Q7 # 0 O7 & !
% ## %F& ! 20 %&
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% ?I
1@F /, &
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o! p>\ - >.
)
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.(
/?
0(& _A
_ ) O7 /? 'Z QQ$ /, /? 2
\ <&=
&>?
90$
3
9M^ P;
&
)
Munns et al., 2002; Munns et al., 2006
( j\? D
8(#qB /R?& r( (
&
'7? _ ) O7 <&6 ?
90$ 'L ?
) (ST
')& +
(Genc et
al., 2007)
. j\? S
%I 8 >g& /, & <+ (
/? _ ) O7 V"= /)#
90\*&
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>d
7? 90$ <i0(
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Dh <&=
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Pitic62
>?&>? /)
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e % >+?
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Yitpi
P6" /? & / & /, ?
\ DP#
'Z '7? 90$
( . %& 9Q P6" # &
/? # ? _(
\0N bL
<&6 \ &ST
,
')& s&6; 6 S>? kl! /,
(Genc et al.,
2007)
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F# D
&
% # t] /,
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7? 90$ /)#
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(Genc et
al., 2007)
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; '7? _
( #
&67& /N 8
K+
?
&67& TF D'7 'Z 8
K+/Na+
;
&67& n=? #
; 90$ 8
20 /?
)
Gorham, 1990
.(
H+
; 1I '\C# c
/?
*
& 9 u /X$* >( /,
>
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2. Na+ exclusion
3. Salt tolerance (ST) 4.Tissue tolerance
%&&
;
>
&$
Z
; 1I )1
(RWC
# )
&>? vY0\ # ')& <)I
H+
\C# c
; 1I '
&&
%
;
>
/? & D
* X W^# 9
60)& _ D 2
# >Z\ c!
&>?
\C# <& <+
# 1I '
; V)]
w>)& '$ <(
'
(Lafitte,
2002)
.
& x(&
%& ? bZ= 8(#qB :
)>?
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; '7? 1I &Q
( 'XA #
Na+
/? D'7?
\ <&=
%&
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(Genc et al.,
2007)
8 90$ <&6 #
) (ST
)>?
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&
F
/0 ( P] %&
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%&
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+# <&#+
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V"= # '7? 1I # 90\*&
<I /? 'Z (
.
شور و داﻮﻣ ﺎﻫ
e P] %& GH /0 <", %& P; _)KB
%#+, "+&
o>, -. %& /, '7
1.Relative water content
& GH O0F <&>
#I &
)
#F ( P; 9 #
MB&6d(
P; D (
MB&>
My &F& #
+#
? P;
. /? -Q$ &
. bL I J>
/I
&
2
,7 \.Q 9 )
Montgomery, 1991
(
zR) # ?
)
" >"*& ' &( '?^
# >L
)
> >? V0 % (
/; Ef # )
MB&6d( /;
9 MB&> /; Dt] / 9
Dt]
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My /; # t]
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t] SB (
v=0N e ] /) t
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<&;
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Q& /H; /? <(; D<)&6*# %& \? # 7 . )& ? (0 0=& &>?
>7 %&
EC × 640 × SP/100 = mg Nail / Kg soil
) ( 20 D
# /Z)$ <&; ! <%# w)& >? P\.
. /)
2. Hierarchical Design
/'0 1 2 22 '(
3 2
T. aestivum (ABBDD) T. turgidum (durum. AABB)
4 5,0
Ae. tauschii (DD)
)"!6$ )"!4
T. boeoticum (AA)
789
+
durum (106)
! T. aestivum(7654-1)
durum (23)
"#
T. aestivum(6546-3)
$%&!'
Ae. tauschii (64)
*-.
T. aestivum(2780-3)
Ae. tauschii (24)
/ T. aestivum(6851-2.2)
0!' Ae. tauschii (15)
1' T. aestivum(500-5.2)
234 Ae. tauschii (19)
*5 T. aestivum(6835-1.1)
Ae. tauschii (35)
6 T. aestivum(6548-4.1)
Ae. tauschii (4)
T. aestivum(7141-4.1)
Ae. tauschii (1)
7 T. aestivum(6979-5.1)
234 Ae. tauschii (54)
durum (51)
$%80' Ae. tauschii (17)
durum (18)
$%08' Ae. tauschii (44)
durum (27)
6' T. boeticum (3)
durum (31)
3905 T. boeticum (5)
durum (34)
234 ) -:/
( T. boeticum (48)
durum (40)
$;<
T. boeticum (14)
durum (47)
0!
T. boeticum (2)
durum (59)
. .
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8 ?
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(Genc et al., 2007; Munns,
& James, 2003)
:
3 4 "#8$ ;<=
ST =
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)>?
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/'0
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3 G '
HIJ 3 $ D$ H7$ KL2
M8N )'* MOP MQ$ RB M@S2 ;<=
3
$ 2
!TQ ( 3
$ U42# M$V QW4 HJ
*
F E(MS)
9 00=' 3&
MS/MS б2e + rб2SG(p) + rghp VS
s-1 (S)
) (
MS> /MS б2e + rб2 SG (p)+ rsб2G(p)+ rsghVP
p-1 ) (P)
>
(
MS /MS б2e + rб2 SG (p) + rsб2G(p)
p(g-1) )
(
?0'@
(G) )
(
MS/MS б2e + rб2 SG (p) +rghб2SP
(s-1)(p-1) ×
)
(
MS/MS б2e+ rб2 SG (p)
(s-1)(g-1)(P) )
( ?0'@
× ) (
б2e s gp(r-1)
A;9 BC
) (
r
p gh
! "#$ %&#
'( s
) .
*+ # , - ,*
/ ! 0+1 2,# %&#
/ 1 %, 1 $ , $
# 13, +4 25 /
:
б2G(p) = (MS3–MS 5)/ rs VP = (MS 2 – MS 3)/ rsgh
/?
#>; X
? t] ? b# /\*R # # (
/;
/? >)K, / 6N %& D(
s#
%& )& ? WARD
)^& /L7
(>u <>, &)& ? &>0(
)&
. s#
>N% >g&WARD
&
TA& /,
s#
UPGMA
(+
#>; > #
? (
9"+ &
>0, D%)
(Mohammadi
& Prasanna, 2003)
.
ﺎﺘﻧ
ﻳ ﺚﺤﺑ و ﺞ
6N
&# /
%&& bL /, & <+ V
;
>
g '$
>
/7>; &>^
&
)
#F .(
, /
>g& bL
; 9!& t] # /; D
\ / &
90$ >X %& /, & <+ # ?
/; ? (
+# <&#+ ! # < P; GH
_( # <I f
/; 9!&
t] ? D(
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F#
'&
. # /; ? 9?Q >g&
&>? [Q7 D
_ )
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V"= D>d? /, ? &
b# 90\*&
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( >u 9?Q GH ')& ! EC
. >g&
? 9?Q
&>? 6 /; 9!& t] #
bL
Na+
1I 'Z # )
(WC
\ )B /, ? &
t] b#
9?Q & (
& <+
)
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8 &67& #
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#F e (
>d S ? /,
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:DW
ZSN $ ;<=
:Na+
)'* '&
MQ$
:WC
;<= $ RB M@S2 :ST
3 %A4 [=
(
A80$@ "00=' DE )
(%CVg
"G3 *0H0 ++
*03 7
7 ) (
×
?0'@
× ) (
?0'@
BC
IJE
/
>
/
>
>ns
/ L
>ns
/ L **
/ LM**
/
>
**
/ M>
/ L DW
/
/ M LM**
/ L
>MM**
/ L L*
/ L **
/ **
/ L>M / L Na+ 1
/
>
>M / LL*
/
>ns
/
>*
/
>
**
/
>L LL**
/
>M
>
/ L WC
/
--
** -- L / L
* --
>L / L LM / ST
. 1
+6 75 %+
log(x+1)]
[
1*
/8# 9 ,**
;< =(
>
1 5 /8# ?@ ABns
.
& 'Q?R <QQ$
(Dehdari et al., 2005)
.
; 2+! <%# /? 1I 'Z D8 [ &>
8 &67& 6
')& /7 .
7 9=
6 ]*
"
H TF &
&67&
8
; WC
:
&67&
8
Na+
#
Cl-
# m>?
&67&
*I & 8
/, ) $
$ [
&67&
8
&67& n=? D?
1I 1@F 8
#
&67& WC
8 ?
(Maas & Hoffman,
1977)
.
&>
[ 8
#
"+!
D <&6
)&
?I 2 ) (ABA
&67&
8 /7
(Genc et al.,
2007)
; [) /%# < /? n=? #
>;
.
8(, p>\ D &>??
/7 ) /?
1I x>l 8(, '=
(
6)7 #
8(, 6
# ?
2+! <%# /N
8(, # /7
&67& WC
8
? .
&67&
8
WC
/? ')& "0 8(, '=
0) >g& 2+!
'
)
Tester & Davenport, 2003
(
&
[) N
&67&
% 8 ) _ ) U (%
&>
# 8 [
/N
&67& / D? 6)7 8(, 1I 1@F 8
)
#F e .(
/'0 1 ;<= HIJ KL2 )
(DW
'&
)'*
)
Na+
( $ RB M@S2 ;<=
) (WC
3 >(4 >(4 '$ ]!^ _
WC Na+
C8 DW
D>>
/ F GH>G / I
>IH / J C8 1,
K>H / J JKFG / GLI / F ) C8
KDD**
/ I KGI**
/ I GFH / I M **
217
FJ LF
FI
% O
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5 .
dZ0(
90$ c! /, & <+ bL ?
) (ST
( dZ0
\ # 'Zh
2+! <%# ? &
8 [$
& 8 <#? [$ # )
#F
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_(
&Q Df 8 [ &> ST
dZ0( D
\ # 1I 'Z # '7? _ ) &Q ? &
'& '7?
. 90$ 8(, TF D'7? _ ) 8 &67&
<&>d+(#qB > ) S ? /, >;
(Dehdari et
al., 2005; Munns et al., 2006)
& 'Q?R .
8 &67&
8 [ &> WC
90$ 8(, TF D
dZ0( &> % D >;
\ # ? &
WC
" ]*? 2+! <%# #
)
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= (r
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)
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8(, TF /, ? ST
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WCs DWs Na+s Na+n DWn ST
*
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DWs
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.
P; /, ')& s&6; O?
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w vY0\
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)
Munns et al., 2006
(
&\ & D
PN& b\*R %&
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90$
P;
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REFERENCES
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2. Colmer, T. D., Flowers, T. J. & Munns, R. (2006). Use of wild relatives to improve salt tolerance in wheat, J Exp Botany, 57, 1059-1078.
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4. Flowers, T. J. & Yeo, A. R. (1995). Breeding for salt tolerance in crop plants: Where next? Aust J Plant Physiol, 22, 875-884.
5. Genc, Y., Mcdonald, G. K. & Tester, M. (2007). Reassessment of tissue Na+ concentration as a criterion for salinity tolerance in bread wheat. Plant Cell and Environment, 30, 1486-1498.
6. Ghanem, M. E., Albacete, A., Martines-Andujar, C., Acosta, M., Romerro-randa, R., Dodd, L. C., Lutts, S. & Perez-Alfocea, F. (2008). Hormonal changes during salinity-induced leaf senescence in tomato. J Experimental Bottany, 59, 3039-3050.
7. Gorham, C., Hardy, J., WynJones, R. G., Jopa, L. R. & Law, C. N. (1987). Chromosomal location of a K/Na discrimination character in the D genome of wheat. Planta, 180, 590-597.
8. Gorham, J. (1990). Salt tolerance in the Triticeae: K/Na discrimination in synthetic hexaploid wheat. J Exp Botany, 41, 623-627.
9. Jafari-Shabestari, J., Corke, H. & Qualset, C. O. (1995). Field evaluation of tolerance to salinity stress in Iranian hexaploid wheat landrace accession. Genetic Resources and Crop Evolution, 42, 147-156.
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15. Munns, R., Husain, S., Rivelli, A. R., James, R. A., Condon, A. G., Lindsay, M. P., Lagudah, E. S., Schachtman, D. P. & Hare, R. A. (2002). Avenues for increasing salt tolerance of crops, and the role of physiologically based selection traits. Plant and Soil, 247, 93-105.
16. Munns, R. & James, R. A. (2003). Screening methods for salinity tolerance: a case study with tetraploid wheat. Plant and Soil, 253, 201-218.
17. Munns, R., James, R. A. & Lauchli, A. (2006). Approaches to increasing the salt tolerance of wheat and other cereals. J Exp Botany, 57, 1025-1043.
18. Noble, C. L. & Rogers, M. E. (1992). Arguments for the use of physiological criteria for improving the salt tolerance in crops. Plant Physiol, 146, 99-107.
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