, ) (
*
:
:
E-mail: [email protected]
زﺎﺳاﺪﺟ ي
و زا ﻲﺸﺨﺑ ﻚﻴﺘﻧژﻮﻠﻴﻓ ﻲﺳرﺮﺑ نژ
زاﺮﻔﺴﻧاﺮﺗ نﻮﻴﺗﺎﺗﻮﻠﮔ
سﻼﻛ هدزﺎﻳ زا
Tauنﺎﻧ مﺪﻨﮔ ﻢﻗر ﻧاﺮﻳا
ﻲ
ﻦﺴﺤﻣ نﺎﺳﺎﺳ
1هداز يرﺎﻔﺻ ﻚﺑﺎﺑ ،* ﺖﺒﺤﻣ ﻦﺴﺣ و2
3رﺎﻛ
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FI131235 ،
FI131236 ،
FI131237 ،
FI131238 ،
FI131239 ،
FI131242 ،
FI131243 ،
FI131244 ،
FI131246 ،
FI131247 و
FI131248
, , O
, F#; B#!
(-, F0
P F%$ F Q
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+ F ) - .R , ?
# . ? );
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ﺎﻫ هژاو ي ﺪﻴﻠﻛ ي : #
<
F F B#!
F F ) .
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! " " #$ % && $' ( )*
( + )* , , !" "
, - +
. "
/$0 -$'
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748 9 ;$
-
#
<
$
$
=> ! -
#
'
$ - ,
!&? 3+ !
@$$?)' A-
, , 1 B
B 25$CD "! E
. $ "
& 1 $'
# 4F>
$ 1$* $ 1 GHI4 # % 2 $$' ! $4$ J$ @$'
KL - *&6 " " #)&
2 "
(Noctor &
Foyer, 1998; Edward & Dixon, 2005)
. ( )*
M&? && $' N" O PL+Q /"R #-
$!&>
, G$HI &* #- -
(Marrs,
1996)
. ! )$S TU -&& $' #-&4? '' 9 1? $' $0 3&6 V
! O$?6!"$- ,+ # TW 1 O$$6!&4HS
4- ,
&' . V JV ! 3Q X>
Q* " 146 %4 GA! GY 1? *&6 1? Z&4 ,S !&$? E$ 1? ! B? "!
, [B&
&'
(Dixon et al., 1998; Cho & Kong,
2007)
. /U 9 #"$? " && $'
#&4? 1+U
\"] J- G^ J-
J-&+8 J-
, ,SL -$' ! "4> J/&I0 B
) Droog, 1997
.(
& $'
&
G0& ,+ "
1+- " ! B -$' , H 26?
- ,-$' #
" U!
) McGonigle et al., 2000
. (
$' =
% ? 1_?" " ,-$' &&
*&6 " B
( 7L , 146&' "&] 15S_ " )$ -
. "
! #- ( 7L , ( )* - ? &+-
( )* -&& G H `' N #-
, ( " ,
, G$HI -
. -&& $' 1 Q*
-$' J" " ,'' #-&4? , B 2$?]
7L ,5$! 1 1? 2C
I 2!% -
, - . -&& $' #"$? &? !`L
)$ ! J$$4$ ! $ &$a ,-$' #K"- 1?
,'S* 1?
-
#
=>
, b> -
- . &? !`L
3&6!) ,Y&?
V 2$?] && $' #-
,Y&? ,S GY %4 1Q$4 " ! -^$S ,Y&? 1?
4- " " /C$&
) Listowski et al., 1988
.(
5?
-&& $' ,c
#
$$4* 1? ; ! -
GA! GY " XC0 TU ! B GV4 -
,
&' .
$$4*
,+ )*&$W #-&4? -
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. P0`_N , -&& $' $d 1?
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» #-^$S
, &4?&$W «
, e`D B
) Mueller et al., 2000
.(
1' &f -
# -$' , && $' = -
Z` "
" 748 #- .
,S 2-CB 1? 1U ?
" 56 #-0! & J,= ,- J-$ $*
Y ! ( )* 56 ^ U ! , $+$B$? /$NV
,H =S+
g " && $' Z`
B , B "4>
"CL 1 :
Alpha ،
،Mu ،Pi Theta ، Sigma ، ،Zeta Omega
9 ! , "4$
Kappa
. -$' " -&& $'
( )* -I ! 7I X>
"4> " - 26
B . ( )* ,$!&> Y $d U!
" -
" "? $48 -$' hijk
,5 B ,5_]
" U && $' B -I 1 ,
!& V TU /"R
Z 1? &* &
, $' 2a6 #&? ,L Y 1 B
T$* &?&? " $' -
7L ,B #
?
(Frear & Swanson, 1970; Board et al., 1997)
. *
14 #&? #54 #-&H+L ( )*
' " -
$ 2 B ,6&5 - .
/5S_
I ,H$4=$6 -&& $' 1
#
$> = O 146&' lI 1$S!
? J
Z` #-$ $* ,S $? 2-CB U!
#-
&4+ Pm+5 748 n
2 N"
(Dixon et al.,
2002)
.
$!&> -&& $' , -
,C &% ,SH ! ? P+L n
1 4- 4S $
0!& !
n B G$HI ,4S $
.
)*
( - /"N 1? 2 H+ - &+ !&4- &+ +
#-0! & 1? ! K* &+ !&4- 9 1 B?
, Z` O 1? o54 B
(Armstrong, 1997)
.
Z`
#-
! Tau
" ! ? -$' ,NV4Y Phi
? && $' #K` & ? 1 %
$
&4I
!&6 , 3= "
'
$
""Y&? - .
Z`< !
'
$ D V TU -&&
$ &4' 7
#
?)'
$
$
@ ' $4F> #& 1?1
$ ! B
&? O$$?)' &p 9 ! )$ " , ) 1? &$p !"
' 748 9
$ L" - ,
5& ! , +L , + .
H+- ! )$<
" "
O B o6 ٢٠٠٣
Z` && $' 14 " 1 " 3' Phi
7L T$& 1? b>
7L !
+ $?
, + ! + U B
(Cummins et al.,
2003)
. q+-
$ " &q " ! r=
skkt ?
u!" 4 2KU O$!&> #-
B ,
$!&>
7L E B %S #- " ($66
$' G$4>
Triticum tauschii
, B 1? o6
1. Xenobiotic
n 1 1? o54 1 B && $' $!&>
Z`
)Tau
g L ( J )Phi
L ( !
Lambda
) L (
?
) Zhang & Riechers, 2004
.(
" "H+- ! X<!"
hiij 1 I
1q-$' Z` && $' 1 , <C #-
1q-$' ? 1 % " " 7Lc $? ? Tau
#-
&?&? " J,I0!
2!% $$ ! & #-
I Y #&4I$?
) Roxas et al., 1997
.(
x>
Z` && $' 1U' $' " Tau
&6 ,^
B , B &+8 " B $? /"N " 1
&?&? " &+8 2!% TU
- $$ #
, &'
) Kampranis et al., 2000
.(
&^ ,I-!;> "
" "H+- ! ) E 1 skky
/"N
26&' Z` && $' ! 1 B I
" x&? $' b> " Tau
&?&?
- $$ #
" 2SY
) Moons, 2003
.(
-!;> "
3' (]"
" B 2I< ,&
& 748 oD
&?
# "? $S!
= Z` && $'
Tau
U
# O$4=$6 ,"&? ! &'
.
شور و داﻮﻣ
ﺎﻫ
3' -
#
? JS
$ 4!)? J/
d J z" J&+
J
- , J
$
"& JB!" J
# -B J ,
" ! "a 1?
4
$
? , ' = @? {SY (`> 3"= 1?
$ - , &
K |&< " }]!
$ B 1 . " -"~? "&] X>
n % -
$
&<F (
/ 1?
1%$]
2KU
L , ' " 3' 1+ &- J -
# 2I< )Q
B . 1+
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TB D J2L g
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1U"
TB # J&^$4 n
2?D" J&^$4 1U"
N"
B "&]
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#
/ E !"
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"K^ &^$4 1U"
#
B . |&84 DNA
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CTAB
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B 3Q 146 )
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0&D "a 1?
,
&>
&+
S &
, -
# @? " B 2Cp ! * 2?
= (- ,A"*
&' ! Z &?
S , -
# E &4I
3&
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#-&'W*
0&D &
, &' :
5´-AAGGGCCTGAGCTACGAG-3´
1?
L
&'W*
!&I$>
!
5´-TGCTGGCGGCTCACTTG -3´
&'W* L 1?
,4I'&?
<! &- "
µl
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1?
$ )
µl
MgCl2
"% 1?
n / ,$
J"m
dNTP
1?
$ )
,$
&'W* J"m
!&I$>
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3< &-
µl
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µl
/
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>
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&
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1$K = $ 2<&B &'
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#
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$ 1%
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$
&
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S4 10&
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1U"
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n 1U"
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1U"
&^$4 / 1?
] @
$ B 3Q 1%
. &Y* 10& "
10& @
]
$ " 1%
#
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# ! 4
3+
#+- 14B" 3+
" - B 146&' &a .
"
>
&?
#
"&?
, V J"< 2N
!" &?PCR
# =
"'*
B "6!&4 S N"
. " 1< 2 &<R G?]
= -!;>
&& $'
= }C ,
|&84
B . V {SY X> PCR
4 E #
{SY 2$<
# V 2<&B E PCR
MWG
2KU " +S*
!&I$>
5
$$
S , B .
!& J= O /$NVY )$S* ! Q4U o &D
S , , 3Q * Z&4 "
B . 3&
#-")6
,'; ! )$S* #&? ,48
= #54 #- ! -
$!&>
" -
^ >
-
# L`D , 4- 4 G?]
.
,S -!;> "
* 2? #- E DNA
#-")6 3&
ProDom ،
PROSITE ،
SMART ،
،CDD
CDART ،
BLOCKS
، وPfam BLAST
&' )$S*
.
J&^ # 1+U& #&?
,S -
&a " # 3&
")6
Transeq
B 4 . 3& @+< 1?
")6
CLC Sequence
viewer
1 &QB )$
, - u!" 1?
Neighbor Joining
) Saitou & Nei, 1987
! (
&?
2-CB Z ! -
/!
#- ,S " U ,$!&> ! #$
= ,S ! B 26 #- #-
1' ,-$' 748 #-
KU O? " U ,
= &' 1$K .
$q+- #&?
(- A"*
, ,S - (4 "^S
ClustalW
J
(Thompson et
al., 1994)
3& " U ")6
BioEdit )
Hall, 1999 (
B 4 .
ﺚﺤﺑ و ﺞﻳﺎﺘﻧ
<! 3Q X>
!0 " , -? PCR
bp
&?
# 1+
-
# 3'
"
? "6!&4 S =
"'*
N"
* 2 1?
1 "
G B -I ,
&' .
1+
V #- X> PCR
S $$5 ,
@? "
KU , =
"+B ? NCBI
-
#
&4 , {8I 1?
" 2Cp
$ !U " 1 h
/5_] D &+- 1?
$$5
S , 2 B "!* B .
3
4 5 2 3
3 0 -- .&> ?@ ; % < 8="/,#9 8. 7 6#
&-9 $A&> &>0B C00 *
D ,%E* 0 F & $% G H*
44 bp
.
8&' 7 . -A' +-* %,%
8I NCBI
90 . &; J%&* $ KLD M!-0 00
Tau
&> @
!"# $%&'%
"*+ FI131235655
-.+ FI131243640
/-# FI131236640
0'%1# FI131237456 +&2 FI131238456
3 FI131244550
304 FI131248643
5++ FI131239636
6+&% FI131246628
$7 FI131247592
$87. FI131242645
,S = +- * 2? #- '
$ &&
Z`<
&? ! ? Tau
# A
! 1 )Q $ G$
* " " @$"$?
B 3Q K .
!U ! G B
,0 $ l K4
# !N
K4
# ,S 1+U& C
1? -
$!&>
'
$ L 1? " S 3' &&
J1+
E Y&?
,
@?
,L`D #- I
, - . &?
Z ,0 K4
# !N
K4
#
C
,S 1+U&
- 1? o54
$!&>
'
$
&&
2 .
8&' 3 7 &N0 $A-. O- P%0GST
0 LI Q-*
ProDom
E value Score
Gene a ProDom Domain
Position
4e-07 129
Bronze-2 PDA0K044
19-70
4e-16 206
GstT1 PD002744
20-60
3e-08 138
Bronze-2 PD085544
20-60
1e-12 177
Gst TSI-1 PD599945
61-109
2e-17 219
Bronze-2 PD337149
137-184
a
Bronze-2
Gst TSI-1
GST
Tau
Zea mays Aegilops tauschii
.
1031 900 800 700
600
500 400
300 200 100
PROSITE 0 LI Q-* P%0C$A,- N $A,-O- &N0 7 3 6#
% S 1 , -
# ? * 2 1?
,S -
# U
1< I = @? "
,S -
# * 2 1?
2-CB N"
# ? "
-&& $' &
#
Z`<
"Tau
.
= 1< Q*
-
# '
$ &&
Z`<
" Tau
# @ 0 " !&4 ,
P`<
0
B , B?
, 4 4 ? GN0 x
Blast
2+]
!&4 ,
&? 1< " &a "
# /5_]
B 26
B J? $<
, ~0 ? ! +
S 1? "<~ 2+]
, 2 1D?& =cDNA
26
) GHB !
.(
% 1 ,S -
# 9 ? * 2 1?
Z`<
-
# ' " && $' 748
$ I -
, + 1< - ,
,S - Z`< && $'
, Tau
B?
. '
$
" 3'
# g $' 9
Z`< &&
1< 2Tau
,S -
# * 2 1?
-CB 2
# ,S 1? "
1? ?&cDNA TaGstU4
?
&4 "+B ,
AF004358
,S !
cDNA
1? ?&
TaGst28E4
&4 "+B ? ,
AF479764
I
, 4 " 1< -
$ #Q4U 1Q
BLAST
G B !
6 2Y"
$ 4=
$ GHB ,H n
, B . !`L 1?
,S -
# ? # 2-CB 3' (]"
S ,
&4B"Y $' #&? * 2 1? && $' "
) GHB n .(
.NCBI نژ ﻚﻧﺎﺑ رد ﻪﻧﻮﻤﻧ ناﻮﻨﻋ ﻪﺑ ﺪﻧﻮﻟا مﺪﻨﮔ زاﺮﻔﺴﻧاﺮﺗ نﻮﻴﺗﺎﺗﻮﻠﮔ نژ ﻲﻟاﻮﺗBlast ﻪﺠﻴﺘﻧ-3 ﻞﻜﺷ ﺺﺨﺸﻣ ﻞﻜﺷ رد ﻪﻛ نﺎﻨﭽﻤﻫ
ﻪﺑ طﻮﺑﺮﻣ ﺖﺴﺨﻧ ﻲﻟاﻮﺗ ود ﺖﺳا ﻲﻟاﻮﺗ
ﺎﻫ ﺗ ﻪﺑ طﻮﺑﺮﻣ يﺪﻌﺑ يﺎﻬﻴﻟاﻮﺗ و ﻲﻣﻮﻧژ ي ﻲﻟاﻮ
سﻼﻛ زاﺮﻔﺴﻧاﺮﺗ نﻮﻴﺗﺎﺗﻮﻠﮔ نژcDNA
رد Tau
ﺖﺳا ﻒﻠﺘﺨﻣ نﺎﻫﺎﻴﮔ .
. ,R &> ?@ M!-0 00 . $90 M 7 6#
-,> O-
J% & (- D 6R $&0SD- .
90 * 0 LI Q-* 'R $&>Tau KLD M!-0 00 V- U,-./ JR7T 6#
&; J% * $
& W)( X* JL * D YZ< &> ?@
[ * &-
Neighbor Joining
. ; ** . %,%
J% &; E,-< 6R .
$ &> 90 #<
90 * *(
& (- YZ< & J/ $ .
, &+-
$$* ,S
# * 2?
/5_]
O ) ? 3' cDNA
GN0 $!&> &
#Q4U
BLAST
,S ,5
TaGstU4
? o?_ !
"4Y 1 J"~ $!&> #"4Y 1 p &NL
* ,6&'S4 &
J2 * 2 1?
1< I
S , 3' -$' #&? B $>
#!0 4? 1N !
g Y 1 p "4Y " S* $>"
2 . & $q+-
2+]
#-
*
S 1?
, ?& ,$?$? #- B
.
= #U ! ,AB ! O$4= ,K " -
" &?" K* O$4=$6 ,"&?
. %4 )* = ( -
#
Z`< VY 1? && $' G$S 1? Tau
1 + &_Y /&p -< "
,
&B J E
_$ L ,
!
$
=> &p )
&p -
(Kilili, 2004)
J , 1'd /&p
# "
G+ "
- 748 # 4
, W !
$
&
4 ,
&?
# '
$
> "
,
B? 14B .
~S /5S_
#&4I$?
#&?
G$+
&
2+]
#- ,S - ,
&Q 1?
#U
= G<
-
# 1D?&
&' $ = %4 #&? 1<
2 ,"&? .
#-
&& $' ( )* O$4=$6
Z` 1 2 I $!&> 748 #-
&4I U O " ,A&^+- ! e& *&6 ,D
1? b>
-
* U?
) Dixon et al., 2002
.(
r$& J,!)' ,0 B Q?U J= B 7Lc #K4 ,0 r$$F ! "!*
$!&> C CST
($H 1+U
? ,A- Y 748 cL 1
"!* U? " $!&>
(Armstrong, 1998;
Wongsantichon & Ketterman, 2005)
. ,m -
" "
( )*
&& $' 1+U J2 &_
Z` &d 1H*
#- ! Tau
!&6 )U Phi
&
Z`
#- 4- CST
= $? 1H ! #-
($a 1^d $!&>
,
&' &$W 1? * !
Z`
- ,56 B 14YB # Z`
#&^ #-
&& $'
" U! -$' "
.
REFERENCES
1. Armstrong, R. N. (1997). Structure, catalytic mechanism, and evolution of the glutathione transferases.
Chemical Research in Toxicology, 10, 2-18.
2. Armstrong, R. N. (1998). Mechanistic imperatives for the evolution of glutathione transferases. Current Opinion in Plant Biology, 2(5), 618-623.
3. Board, P. G., Baker, R. T., Chelvanayagam, G. & Jermin, L. S. (1997). Zeta, a novel class of glutathione transferases in a range of species from plants to humans. Biochemical Journal, 328, 929-935.
4. Cho, H. Y. & Kong, K. H. (2007). Study on the biochemical characterization of herbicide detoxification enzyme, glutathione S-transferase. Biofactors, 30, 281-287.
5. Cummins, I., O’Hagan, D., Jablonkai, I., Cole, D. J., Hehn, A., Werck-Reichhart, D. & Edwards, R.
(2003). Cloning, characterization and regulation of a family of phi class glutathione transferase from wheat. Plant Molecular Biology, 52, 591–603.
6. Dixon, D. P., Cummins, I., Cole, D. J. & Edwards, R. (1998). Glutathione mediated detoxification systems in plants. Current Opinion in Plant Biology, 1, 258-266.
7. Dixon, D. P., Lapthorn, A. & Edwards, R. (2002). Plant glutathione transferases. Genome Biology, 3, 3004.1-3004.10.
8. Droog, F. (1997). Plant glutathione S-transferases, a tale of Theta and Tau. Journal of Plant Growth Regulation, 16, 95–107.
9. Edwards, R. & Dixon, D. P. (2005). Plant glutathione transferases. Methods in Enzymology, 401, 169–
186.
10. Frear, D. S. & Swanson, H. R. (1970). Biosynthesis of S-(4 ethylamino 6-isopropylamino 2-s-triazino) glutathione; Partial purification and properties of a glutathione S-transferase from corn. Phytochemistry, 9, 2123–2132.
11. Hall, T. A. (1999). BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium 41, 95–98.
12. Kampranis, S. C., Damianova, R., Atallah, M., Toby, G., Kondi, G., Tsichlis, P. N. & Makris, A. M.
(2000). A novel plant glutathione S-transferase/peroxidase suppresses bax lethality in yeast. Journal of
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