全 文 ： ACTA AGRONOMICA SINICA 2008, 34(5): 823−830 http://www.chinacrops.org/zwxb/
ISSN 0496-3490; CODEN TSHPA9 E-mail: xbzw@chinajournal.net.cn

:

  (2006BAK02A18);  (Z306300);  (973
)(2002CB10804)
 :

!(1983–), , #$%&, ()*, +,-./012*345
*
6789(Corresponding author): :;4Tel: 0571-88206481; E-mail: pzhch@zju.edu.cn
Received(<=>?): 2007-09-23; Accepted(@A>?B: 2007-12-22.
DOI: 10.3724/SP.J.1006.2008.00823
AsA-GSH  
  1  1   1,*  2
(1   ,  310058; 2 ,
 310006)
 :  11  ,  0.4 mmol L−1 Hg2+  
 !-#$%&(AsA-GSH)( )*+,-., Hg2+,  /0 H2O2123 2O− 45678
MDA123GSSG123DHA129:;<=5>;  /0 GSH/GSSG8 AsA/DHA?@AB<=5>, C
D E8F Hg2+12.9B<=5>*G.  H AsA-GSH ( Hg2+IJKLMB
NOP78QRJS,  TUVWX*
: Hg2+; ;  ; ;  !-#$%&(
Effect of AsA-GSH Cycle on Hg2+-Tolerance in Rice Mutant
ZENG Bin1, WANG Fei-Juan1, ZHU Cheng1,*, and SUN Zong-Xiu2
(1 State Key Laboratory of Plant Physiology & Biochemistry, College of Life Science, Zhejiang University, Hangzhou 310058, Zhejiang; 2 State Key
Laboratory of Rice Biology, China National Rice Research Institute, Hangzhou 310006, Zhejiang, China)
Abstract: Mercury (Hg) toxicity is an in rice growth problem throughout the world. In the present study, Zhonghua 11 and a
Hg2+-tolerant rice mutant (MT) were used in a solution culture to investigate the effect of 0.4 mmol L−1 Hg2+ treatment on reactive
oxygen species (ROS) metabolism and dynamic change of ascorbate-glutathione (AsA-GSH) cycle. The results indicated that the
H2O2 content, 2O
−

evolution rate, MDA content, GSSG and DHAcontents of the leaves were higher in the wild type than in the
mutant; both the ratio of GSH to GSSG and the ratio of AsA to DHA were higher in the mutant than in the wild type, while the
accumulation of Hg2+ in roots and stems of the mutant was more than that of the wild type. The results reveal that AsA-GSH cycle
was less inhibited in the mutant than in the wild type, thus the mutant was able to scavenge ROS more. An effective AsA-GSH
cycle is important for the mercury resistance of mutant.
Keywords: Hg2+; Rice; Mutant; Reactive oxygen species; AsA-GSH cycle
  “”
 Hg2+, ! Hg2+
#$%&(Hg2+)*+,-./01+234
567, 89:3;<=>?4, &@A?B
CD(EFGHIJKLMNOP:23QR ,
SGTUV Hg2+23QR(WX, YZEF[
Hg2+\]^^_`abOcd(Hg2+[e
3fg2hijkl]mnopqbr(s
?YZtu, Hg2+)*vwxy[1]z{[2]|}~[3]
e345/0 l]m(ROS), €^3
‚ƒm„…, †‡e3…g(ˆ‰Š‹-ŒŽ
 (AsA-GSH  )Ge345Oˆm‘
’ [4], “89 AsA/DHAGSH/GSSGNADPH/
NADP+”•–—5m˜™ !(rš Hg2+[e
3ˆm›‘’jk[2-5]*A AsA-GSH  +e
3[œ`[6]žŸ[7] !¡¢.£w2¤
b¥¦§¨,© AsA-GSH  hEF[ Hg2+ª«]r‘˜¬­§¨(YZtu, ®¯Be3
*A®¯e3®°¯[±²ª«\³_`
´µ¶®[8-9], WX·¶®¸¹º»¼½4
824      34

¾YZ¿bcd[10](ÀÁYZ Hg2+ª«Ã¼½4
Ä^Å\]ÆÇh AsA-GSH  ¢È_`
¶Ér‘, *ÊËÌ Hg2+[EF…g_*AE
F[ Hg2+\]_`/0YZÍ(
1 
1.1 
*.IEFYZ£EF^3ÎIÏÐÑÒÓ
ÔÕEF.Ö 11 (Oryza sativa L. subsp. japonica,
cv. Zhonghua 11, WT)ʳ4×ؼ½4Ù.‚Ú
ÛÜRÝ Hg2+-\]¼½4 (Hg2+-tolerant mutant,
MT)[10], Þ PCR ßàtuX¼½4‘á…âãäå
9æ.^4–—ç]‘¼½4(Hg2+è·éê
Ê 0.4 mmol L−1, ë 7 dìÄ^ż½4íîu
ïÆÇ(ð 1)(ñòónäå1ô?·õ, ö÷
îø¼½4[ Hg2+\])*ùú¸¹(*ø¼
½4 T6nÊûÍ=üYZ(



 1 0.4 mmol L−1 Hg2+  7 d  
Fig. 1 Phenotypic difference between wild type (WT) and
mutant (MT) under 0.4 mmol L−1 Hg2+ for 7 d
A: ; B: Hg2+ 
A: Water culture without Hg2+; B: Water culture with Hg2+.

EF¯ƒýþš
., EäåK 2 
ìK 6 L å  25 cm Í, 
( å IEFYZ£[11], 
 pH K 5.0~5.1(Ê 8  Í, F
!ú+5, # 2 $, # 16 $, äåK 6
% HgC12 ë(&()ÑÒô*, ëéê
+Ê 0.4 mmol L−1(
,-./-0(BSO)G γ-Œ1/21‹
‡›(γ-ECS)`3, YZtu, 2 mmol L−1 BSO
ë)*ï456e3–—5 GSH [12](17
8(3-amino-1,2,4-triazole, AT)G9m9›(CAT)
) `3[13], ëì)€e345H2O2
/0(äåK 6 EF:‚Ê 3 â, ;
RâÊ[<%ë, ;=â 2 mmol L−1 ATë
2 d(AT>%K å ., Eä), ;â? 2 mmol
L−1 BSO(ë 12 h, @íìAB&CD 2 mmol
L−1 ATë 2 d(
1.2 
@ Hg2+ë 123 4 dEF&E
F°àú Hg2+éê(? 20 mmol L−1 EDTA-Na2
G HI 15 min*JKEFtLUM Hg2+, D
KڃEAB(£bF°+ 70NÃOžKP, 
QRST_U–, 9 60 Vè, @SW 0.1~0.2 g,
X H2SO4-HNO3Y‹(1Z1)[, AFS-930 ™ƒ
\œœ]^àú[ . Hg2+éê(
@ Hg2+ëì 0123 4 dEF; 2_
´µ¾ (^`aùúbant] ), <
Arnon[14]càúde ; fcàúH2O2 ; fcàú ^no ; f=t(MDA) ; uvœ[18]càúˆ‰Š
‹(AsA)w9ˆ‰Š‹(DHA)*A˜™ÅŒŽ
(GSH)mŌŽ(GSSG) (
£bÑÒxy 3 z, #Ýë 3 Ýpü(
X SPSS 10.0’{|}=üƂ~(
2 
2.1 Hg2+  
ð 1 )­, Hg2+-\]¼½4Ä^Å+€
‚}Ã^ƒ7ÀR, Þ 0.4 mmol L−1 Hg2+ë 7 d
ì=„tÅÆÇï4, Ä^Å_…†‡; ¼½
4 C‚_xˆ•^l‰(¼½4[ Hg2+ª«
\]uï šÄ^Å(
2.2 Hg2+  
Hg2+
t 1)­, 0.4 mmol L−1 Hg2+ª«Ã, ë
Š‹Œƒ, Ä^ż½4&E.
Hg2+éꍎ%; ¼½4&E. Hg2+éêx
uÄ^Å, <‘„_. Hg2+éê’bï
4ÆÇ(t 1)(
 5  : AsA-GSH   825


 1 0.4 mmol L−1 Hg2+    Hg2+
Table 1 Hg2+ concentration in roots, stems, and leaves of wild type (WT) and mutant (MT) under 0.4 mmol L−1 Hg2+ for different
exposing time
 Root (mg kg−1 DW)

 Stem (mg kg−1 DW)

 Leaf (mg kg−1 DW)

Hg2+
Hg2+ exposing time WT MT WT MT WT MT
0 d 37.01±1.50 e 34.65±1.38 e 6.74±0.24 e 7.43±0.41 e 8.66±0.17 e 9.26±0.42 e
1 d 2051.9±7.3 d 2342.5±6.0 d** 529.7±2.6 d 608.2±1.0 d** 83.2±3.1 d 81.9±1.7 d
2 d 3372.8±2.4 c 3505.9±3.5 c** 1086.3±3.5 c 1159.6±2.9 c** 113.1±1.7 c 105.8±2.8 c
3 d 3427.3±3.3 b 5632.1±2.3 b** 1128.5±3.5 b 1214.3±1.1 b** 145.1±2.9 b 143.4±2.1 b
4 d 3727.3±2.6 a 7054.1±1.9 a** 1156.8±2.7 a 1541.4±1.1 a** 374.3±1.4 a 383.5±2.1 a
   P<0.05*** !#$%&(P<0.05)(P<0.01)
Values within the same column followed by a different letter are significantly different at the 0.05 probability level. * and ** represent
significantly different at P<0.05 and P<0.01 between wild type and mutant, respectively.

2.3 Hg2+  
 0.4 mmol L−1 Hg2+,  
; 
 ,  !#$%&Hg2+
() 4 d, * 0 d +,-.,  
 /01 39.8% 16.8%(2 2)



 2 0.4 mmol L−1 Hg2+  

Fig. 2 Chlorophyll content in leaves of wild type (WT) and mu-
tant (MT) under 0.4 mmol L−1 Hg2+ for different exposing time

2.4 Hg2+   
 H2O2MDA
H2O234567 89:2 3 ;
 <= >?@ H2O2 A
 Hg2+()  BCDE , F 
2O
−
>?@ H2O2GH0.4 mmol L−1 Hg2+
() 1I2I3 4 d, −
>?@
/0.K 10.4%I12.4%I15.1% 19.3%;
<= H2O2 /0.K 31.7%I
40.7%I53.6% 43.3%, LMANOP
MDA3QRS:T >U, VW1XYQ Z
[\]2 4^_,  Hg2+()  , 
 <= MDA `DE, F
CKa0.4 mmol L−1 Hg2+() 1 d
,   MDAbcLM, F
() 2I3 4 d , <= MDA /
0. %d1 40.2%I39.6% 35.9%, 4eL
MNfOPgC Hg2+()  ,
Qhijf S:T[kGH
2.5 Hg2+   GSH
GSSGAsA DHA
GSH AsA3lUJmn o:Tp, 3q
rso:Thi 67tuv/0.4 mmol L−1 Hg2+
(),  <= GSHAwD
E Hg2+() 2 d,  
<= GSH ACDExNfy&z, /0
. 0 d E{ 40.7% 40.2%; F()
 , J GSH |}n 0.4
mmol L−1 Hg2+() 4 d,   GSH
/0. 0 d  68.9% 35.8%, 
\C{a ~()S\=,
<= GSHA{aHg2+(
),  = GSSG A€DE
Hg2+() 1I2I3 4 d, <=
GSSG/0. 0 d %d 25.8%I91.1%I
128.7% 170.5%; <= GSSG‚/0%d
6.9%I18.4%I35.8% 44.7%;  %!
{a  GSH/GSSG.z 
B#$; ‚wDE , n-ƒ(
)  GSH/GSSG.zA{a
(„ 2)
826      34




 3 0.4 mmol L−1 Hg2+    H2O2
Fig. 3 Effect of Hg2+ treatment on generation rate and H2O2 content in leaves of wild type (WT) and mutant (MT)




 4 0.4 mmol L−1 Hg2+  
MDA
Fig. 4 Effect of Hg2+ treatment on

MDA content in leaves of
wild type (WT) and mutant (MT)
0.4 mmol L−1 Hg2+(), <= AsA
  , wD
E ;  = DHAA
E{ („ 3) <= AsA 
n()…†‡?DE, n 2 dNfy&z, 
Hg2+() 1I2I3 4 d,  DHA
/0. 0 d %d 9.1%I53.3%I60.2%
77.7%; ‚. 0 d %d 7.4%I28.4%I
42.1% 51.6%, %!CHa* GSH/GSSG
T-ˆ, <= AsA/DHA .z#$
 , n =‚wDE 
, x‰n-ƒ()4eLM, 
AsA/DHA .zC{a+,„C, 
JG{ GSH/GSSG.z AsA/DHA.zcŠa‹
€Œ :TŽ‘, ’“ Hg2+”• :T()

 2 0.4 mmol L−1 Hg2+   GSH GSSG
Table 2 GSH and GSSG contents in leaves of wild type (WT) and mutant (MT) under 0.4 mmol L-1 Hg2+ for different exposing time

GSH (nmol mg−1 protein)


GSSG (nmol mg−1 protein)

/ 
GSH/GSSG

Hg2+ 
Hg2+ exposing time
WT MT WT MT WT MT
0 d 127.13±1.25 b 138.52±1.12 c* 36.61±1.37 e 37.92±1.51 d 3.47±0.17 a 3.65±0.12 c
1 d 133.62±1.59 b 167.27±1.19 b** 46.05±1.32 d 40.53±1.07 d** 2.90±0.08 b 4.13±0.05 b**
2 d 161.03±1.32 a 193.07±0.96 a** 69.98±1.09 c 44.87±2.56 c** 2.30±0.04 c 4.30±0.05 a**
3 d 104.54±1.87 c 136.93±1.63 c** 83.73±1.11 b 51.49±2.16 b** 1.25±0.07 d 2.66±0.08 d**
4 d 50.10±2.38 d 101.47±1.42 d** 99.05±2.16 a 54.85±1.32 a** 0.51±0.04 e 1.85±0.06 e**
 P<0.05  ! * ** #$%&()*+ (P<0.05),
(P<0.01)!
Values within the same column followed by a different letter are significantly different at the 0.05 probability level. * and ** represent
significantly different at P<0.05 and P<0.01 between wild type and mutant, respectively.

 5  : AsA-GSH   827


 3 0.4 mmol L−1 Hg2+   AsA DHA
Table 3 AsA and DHA contents in leaves of wild type (WT) and mutant (MT) under 0.4 mmol L−1 Hg2+ for different exposing time
-./0
AsA (µmol mg−1 DW)

12-./0
DHA (µmol mg−1 DW)

-./0/12-./0
AsA/DHA

Hg2+ 
Hg2+ exposing time
WT MT WT MT WT MT
0 d 3.66±0.63 a 3.84±0.76 c** 3.19±0.85 d 1.85±0.57 e ** 1.15±0.02 a 2.07±0.04 a**
1 d 3.28±1.04 b 4.51±0.85 b** 3.48±0.97 c 3.06±0.98 d** 0.94±0.03 b 1.47±0.07 b**
2 d 2.97±0.72 c 4.76±0.94 a** 4.89±0.69 b 3.66±0.87 c** 0.61±0.05 c 1.30±0.06 c**
3 d 2.65±0.91 d 3.72±1.14 cd 5.11±1.12 b 4.05±1.09 b** 0.52±0.04 d 0.92±0.04 d**
4 d 1.97±0.84 e 3.59±1.03 d** 5.67±0.91 a 4.52±1.23 a** 0.35±0.05 e 0.79±0.05 e**
 P<0.05 !* ** #$%&()*+ (P<0.05), (P<0.01)!
Values within the same column followed by a different letter are significantly different at the 0.05 probability level. * and ** represent
significantly different at P<0.05 and P<0.01 between wild type and mutant, respectively.

2.6 (BSO)  (AT)
 H2O2 !
2 mmol L−1 2 d,  
 GSH ,   GSH 
 ;   H2O2
 !,   H2O2# 
$ 52.3%, %&()BSO *+
,,   GSH-./0
1 0 d 28.1% 24.6%, $234
5678;   H2O2-
.#, ! 79.1% 180.3%, 9
:;4<78!=;   GSH
 H2O2>((? 4))@ AsA-GSH
ABCDEFGH, IJKL Hg2+MNGO
PQRST; GSHU/0VW AsA-GSHAB
XY, Z[FGHU)
2.7 Hg2+# $%&()*
\? 5]^_, $4`a H2O2b c
debMDA bDHA fghi, ja
AsA/DHAbGSH/GSSGfkhi)AsA/DHAb
GSH/GSSGa L Hg2+lmnIMGop
hi)

 4 2 mmol L−1(AT) 2 d   GSH H2O2 BSO
Table 4 GSH and H2O2 contents in leaves of wild type (WT) and mutant (MT) under 2 mmol L−1 AT for 2 d and effect of BSO pre-
treatment
 GSH (nmol mg−1 protein)

3 2 H2O2 (mmol g−1 FW)

4
Treatment group WT MT WT MT
56 Control 126.03±1.69 b 137.70±2.32 b* 1.77±0.32 c 1.65±0.41 c
AT 154.75±1.87 a 191.17±1.58 a** 3.34±0.35 b 2.39±0.44 b**
BSO + AT 32.11±1.57 c 33.84±1.88 c 6.52±0.41 a 6.70±0.36 a*
 P<0.05 !* ** #$%&()*+ (P<0.05), (P<0.01)!
Values within the same column followed by a different letter are significantly different at the 0.05 probability level. * and ** represent
significantly different at P<0.05 and P<0.01 between wild type and mutant, respectively.

 5 Hg2+  !#$%
Table 5 Correlation among several physiological indexes of mutant (MT) under Hg2+ stress

3 2
H2O2
7 89:
2O
−


;<=
MDA


GSSG
12
-./0
DHA
-./0
/12-./0
AsA/DHA

/ 
GSH/GSSG
>?@ Chlorophyll
−0.634* −0.672* −0.577* −0.358 −0.545* 0.595* 0.556*
3 2 H2O2 0.516 0.935* 0.537* 0.784* −0.645* −0.563*
7 89: 2O
−


0.668* 0.866* 0.887* −0.863* −0.610*
;<=MDA
0.537* 0.911* −0.760* −0.549*
 GSSG
0.785* −0.946* −0.951**
12-./0 DHA
−0.999** −0.813*
-./0/12-./0
AsA/DHA
0.913*
* ** #$&A BC, BC!
*
and ** denote significantly different at P<0.05 and P<0.01, respectively.
828     D 34E

3 
  
[19],   !
#$% Hg2+&()*+ Hg2+&,
-./0123, 456789:;<=
> , $%?@ABC=D Hg2+EFG
H!IJ<KLMNOP:QRS, T
UVWXYZ[\]^$_[20]!O`
 a Hg2+EF,, KLMbcde5fgh
ij , KLM>kl , mn5>WXoSp
qrsMt!u#vw$%, Hg2+EFZx456
p89: H2O2 >lkyz{ 2O
−
e5BC!4
56 H2O2 p 2O
−
 e5BC01|}89:;
Hg2+EF(89:~ MDA I, qr
oS€<, ‚d89:ƒ|„KLMbc
…†R‡!
:ƒˆMt…†‰WXˆMt
(GSHŠAsAŠ5‹ŒŠVit E)pˆMtŽ(Mt
tŽ SODŠsMtŽ PODŠsMt‘Ž CATŠ
’“”•–—Ž GR)˜™!GSH:ƒ
š›œˆMt, ždŸ:ƒ ¡Mt–—¢
£!¤¥¦dKLM¢£, §žd AsA-GSH
¨©bc H2O2[21]!ª«ƒ GSH p
GSH/GSSH 7¬­® AsA-GSH ¨©¯°„C|
I‡±^[22]!u²³$%, › 0.4 mmol L−1 Hg2+
&(,456p89:~ GSH ´µ
¶·|(¸¹ , O›EF(º , 456
~ GSHB<%0{}89:!»¼½EF
¾ºGSH·|‚PEF¿¡À£
L¢£, ÁÂsz{ GSHÙ, ÄňÆL;
EF(º456ƒGSHÇÈ‚
^ª«ƒ> GSH É}Ê¡Ë s€, Ì
PC ÙÍ, ΈMtžd AsA-GSH ¨
©ŠbcKLMω]!›ÐÑÒÓpÔÕE
Fs€~, 89: GSH Ö×|}456, Ø
‚?:ƒždGSHÙpÙ5 γ-’ÚÛܓÚ
Ýَ(γ-ECS)Š’“”•Ã™Ž(GSHS)Š’“”
•–—Ž(GR)KL|Þx!GSHp AsAMt–
—ßàdPÆáÀ£âãä‡, AsA å‚
æçd GSHè£pMt–—ßàéâä‡[23]!›
Hg2+EF¾º456:ƒ GSH êRÞë·,
O GSSG ë·B<s GSH, GSH/GSSG Ö
×ìí¸¹; 89: GSSG Ä;}
456, GSH/GSSGµ¶ë·(¸¹!89
:| GSH/GSSG, ¿îïRð}ñíª«ƒ
–—©á, ò¿îï§Rð}ó|AsA-GSH¨©
„C!
(AT)   , 
 Hg2+ GSH ,  GSH 
 !#, γ-ECS  GSH $%
&()*+,-./γ-ECS 
BSO0, 1234567GSH89:;
<=>?@A, AsA-GSH BCDEF(G
HI0, 123456JK GSH 
H2O2DLMN/, 456GSH89*LO
123P H2O2*L?123; BSO
Q=HI0, 123456JK GSH89
D*L:;<=R@A, STUV !/
W, H2O2 89*LXYZ[LM\], ^R_`
aRb)* AsA-GSH BCcdefg
h#(
GSHcô›bcKLMs€~æç Éõ,
–öÕ(PC)ÙÍ, dP 
÷Lâãä‡[24-25], PCÂsd øXöÃ
ÄÅùˆEF‚ú[26]!Hg2+Pû]5
WXRüÅýpú , ^þ89:ƒ|
GSH ‚n
¡É, ¿îïó| AsA-GSH ¨©
„C; ò¿îïd Hg2+¥¦öÃÎÂsÙ PC
d Hg2+-¦öÃ!u#$%, Hg2+EF()*:ƒ
 Hg2+œkl›, 89:~ Hg2+J<
%0|}456, Ø‚^89:Ù>
 PC,  Hg2+öÃt ®, 2 Hg2+m
nMtEF!
AsA APXXè:t H2O2–—, y
hMt™ ‘ˆjÝ(DHA); ( ‘ˆj
ݖ—Ž(DHAR)t–—[27]!AsA p DHA ä
Pfg› AsA-GSH¨©~n É!456p
89: DHA´+ Hg2+EF,-./µ·|
¸¹, O456Ä;>}89:!¿7
AsA/DHA 7¬æ, ›äEF€<89
: AsA/DHA 7¬´01|}456, %89:
ƒ| AsA pÙ5‚úd?|bcKL
MEF„Câãä‡!²³$%, AsAPqr
sMtRÉ [28]!ÂsP456p89:
 5  : AsA-GSH   829


MDAŠAsAŠAsA/DHA7p89:
~ MDAŠDHA AsA/DHAä‡L
W$%, DHA d MDA µ01Ðä‡,
AsA/DHA d MDA µ01ä‡!AsA 
AsA-GSH ¨© ™2 ‰ Hg2+mns
MtS!Ød?²³vw¿x[29-30]!
u²³$%, › Hg2+EF, ÷LÅ89
:‚ñí| GSHŠAsA  GSH/GSSGŠ
AsA/DHA)f, 456 GSHŠAsA%0I
 GSSGŠDHA%0·|, AsA-GSH¨©„C
, N‚R„bcª«ƒKLM!89:
GSHp AsA›ÔÕEFs€´|}456, Ø
‚d89:ƒ GRŠDHARKL|R‡, §‚
 GSHŠAsAَKLÄÅ !!
4 
#ô¿Õ)* Hg2+-÷L89:!› Hg2+EF
, )* AsA-GSH¨©PEF$Ën É!
89:AsApGSHÙ5‚úÅ, bcKLM
‚úÅ , qrsMtS , Ø89:P
Hg2+EFÀ£ 5]%!

   
!
References
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[2] Gu W(), Shi G-X(), Zhang C-Y(), Wang
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), 2002, 28(1): 69−74 (in Chinese with
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[3] Shi G X, Xu Q S, Xie K B. Physiology and ultrastructure of
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