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
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?YZtu, Hg2+)*vwxy[1]z{[2]|}~[3]
e345/0 l]m(ROS), €^3
‚ƒm„…, †‡e3…g(ˆ‰Š‹-ŒŽ
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NADP+”•–—5m˜™ !(rš Hg2+[e
3ˆm›‘’jk[2-5]*A AsA-GSH  +e
3[œ`[6]žŸ[7] !¡¢.£w2¤
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1.1 
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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
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 pH K 5.0~5.1(Ê 8  Í, F
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 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-’ÚÛÜ“Ú 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$Ën É! 89:AsApGSHÙ5‚úÅ, bcKLM ‚úÅ , qrsMtS , Ø89:P Hg2+EFÀ£ 5]%!      ! References [1] Zeng X-M(), Shi G-X(), Xu N( ), Xu Q-S( ), Zhang X-L( ). Effects of mercury on active oxygen metabolism and chromosome in Sagittaria trifolia. J Plant Physiol Mol Biol (), 2003, 29(3): 227−232 (in Chinese with English abstract) [2] Gu W(), Shi G-X(), Zhang C-Y(), Wang W(), Xu Q-S( ), Xu N( ), Zeng X-M(), Zhang X-L( ), Zhou H-W( ). Toxic effects of Hg2+, Cd2+ and Cu2+ on photosynthetic systems and protective enzyme systems of Potamogeton crispus. J Plant Physiol Mol Biol ( ), 2002, 28(1): 69−74 (in Chinese with English abstract) [3] Shi G X, Xu Q S, Xie K B. Physiology and ultrastructure of Azolla imbricate as affected by Hg2+ and Cd2+ toxicity. Acta Bot Sin, 2003, 45: 437−444 [4] Ken’ichi O. Glutathione-associated regulation of plant growth and stress responses. Antioxid Redox Signal, 2005, 7: 973−981 [5] Pang X(!), Wang D-H(#), Peng A($%). Effects of
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[6] Xiong F-S()*), Song P(+,), Wang F-T(-.), Gao
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