稻田稗草对丁草胺的抗性测定及机理初步研究
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摘要
本论文测定了我国部分稻田稗草对丁草胺的抗药性，并对其产生机理从代
谢角度进行了初步探讨。
在湖北，黑龙江等地调查发现田间用丁草胺防除稗草的效果明显不如以
前。本文以华中地区三种群岳阳（未使用过丁草胺）、湖北漳湖垸农场3队（使用
丁草胺14年）、湖北漳湖垸农场4队（使用丁草胺18年），东北地区三种群牡丹
江（未使用过丁草胺）、民主（使用丁草胺10年）、阿城（使用丁草胺14年）为研究
对象，运用室内盆栽生物测定方法研究了其抗药性现状。试验结果表明，LC_(50)
值以岳阳种群最小，牡丹江种群最大，分别为54.73g a.i./ha和1271.6g a.i./ha；
敏感性由高至低依次为：岳阳、阿城、3队、民主、4队、牡丹江。以岳阳种
群为参照，阿城、3队、民主、4队稗草种群的抗药性系数分别为5.21、5.85、
8.00、12.01，表明长期用药使其产生了对丁草胺的抗药性。而牡丹江种群种子
粒大，千粒重较其它种群显著高，加上可能由于其它除稗剂禾大壮、二氯喹啉
酸等的使用，使其具有获得抗性（或耐药性）的可能。
本文还应用生物化学方法，选择了三个苗龄的稗草来测定各种群幼苗中
GSTs酶活和GSH、Cysteine等自由巯基的含量，初步研究了它们分别与LC_(50)
值之间的相关性。结果表明：（1）当用5ppm丁草胺处理稗草幼苗48小时时，
酶活有不同程度的提高。GSH、半胱氨酸含量差异显著，其中，南方处理种群
GSH含量与LC_(50)值之间正相关，北方处理种群Cysteine含量与LC_(50)值之间呈
负相关。（2）而用5ppm丁草胺处理9天的稗草，处理种群与未处理种群比较，
其酶活有不同程度地提高（阿城种群例外），但各种群酶活与其生测的致死中量
LC_(50)值之间未表现出线性相关性。各未处理种群之间GSH含量差异极其显著，
它们与LC_(50)值之间的线性拟合方程为y=0.0321x+25.029（R~2=0.8382）。（3）种植
14天的稗苗，未用药剂处理，酶活表现出一定的规律性，即：酶活与LC_(50)值
的关系，华中种群为正相关，北方种群为负相关。而南方种群幼苗GSH含量
与LC_(50)值有正相关趋势，北方种群幼苗GSH含量与LC_(50)值有负相关趋势。（4）
测定GST专一性酶活结果表明，所采用的方法未检测到GS-丁草胺，可能是
与GST酶对甲草胺、乙草胺、异丙甲草胺的专一性酶活相比，对丁草胺的酶
活较低，这与Scarponi（1991年）报道的结果一致，其专一性酶活的测定方法需
稻山稗草对丁芋胺的抗性测定及机理初步Uf究
进一步探索。而体外轭合实验表明，在一定条件下非酶结合是可以实现的，这
己通过TLC和质谱获得证实。
总之，田间调查和室内生物测定结果表明我国部分稻田稗草对丁草胺已产
生不同程度的抗药性。通过GST酶活的测定以及簿层层析和质谱分析表明GST
酶代谢丁草胺解毒是其抗或耐药性的机制之一。抗或耐丁草胺的稗草种群GSH
等自由疏基含量和酶活有显著提高，酶活或GSH含量同LC。。值己表现出一定
的相关性，但南方种群与北方种群相关性表现不一致，可能与气候、土壤等环
境条件、作物种植管理水平不同、丁草胺的特殊结构及抗性水平有关：稗草对
丁草胺的抗药性机制还可从其它途径进行研究。
The present status of butachlor resistance of Echinochlor crus-
galli(L.)Beauv.(CE) occurring in paddy rice in north and south of China was studied,
and the resistant mechanism from the point of metabolism was studied. With
continuing application of butachlor in the fields of Hubei and I-Ieilongjiang
Provinces, it was noticed that the herbicidal efficacy for controlling EC was reduced.
Seeds of different EC populations collected from 6 sites (Yueyang, no butachlor
applied, Team III on Zhanghuyuan Farm with 14-year application of butachlor, Team
IV on Zhanghuyuan Farm with 18-year application of butachlor in the south;
Mudanjiang, no butachlor applied, Minzhu with 10-year application of butachlor,
Acheng with 1 4-year application of butachlor in the north)were tested to evaluate
butachlor resistant status in the areas. It was revealed that LC~0 for controlling of
Yueyang population was the lowest, that of Mudanjiang population was the highest
among the six populations, 54.73 g a.i./ha and 1271.6 g a.i.Iha respectively. The
order of sensitivity of the populations from high to low ranged: Yuyang, Acheng,
Team III, Minzhu, Team IV, Mudanjiang. Using Yuyang population as the control,
RI indexes of other populations excluding Mudanjiang population were Acheng 5.21,
Team III 5.85, Minzhu 8.00, Team IV 12.01, which indicated that the four
populations had evolved resistance to bubachlor.
Three seedling stages was selected to test glutathione S-transferase(GST)
activity and the amount of GSH, cysteine, and whether it was correlative with LC50
values were studied. Biochemical tests showed that (1) when the seedlings of EC
were treated with butachlor (5 ppm) for 48 hours, GSTs activities increased, the
amount of GSH, cysteine show significant difference. The amount of GSH of treated
populations was positively correlative to LC0 for controlling of the populations in
the south, the amount of Cysteine of treated populations was negatively correlative
to LC50 in the north. (2) The seedlings of EC were treated with butachlor (Sppm) for
9 days, compared to the untreated populations, the GSTs (CDNB) activities of each
treated population were higher with exception of Acheng population, but the amount
of GSH varied significantly according to the different populations. There was linear
trend between the GSH and LC, i.e. y=0.032l+25.029 (R2=0.8382). (3) GSTs
activity and the amount of GSH of the unbeated seedlings grown for l4 days showed
positive correlation to LC,, for controlling of the POpulations in the south
respectively, but negative correlation to LC,, of those in the north. Although GS-
butachlor was not found in vivo study, it was confirmed in vitro conjugating tests by
TLC and mass spectrum techniques. The free thiols and GSTs activities to CDNB in
EC populations indicatCd that butachlor resistance in EC was increased notably.
GSTs activity or the amoun of GSH showed direct correlation with the LC,, fOr
controlling of different poPulations to some extent. It is suggested that the analysis
method needs to be improved, studying of other possible mechanism is needed.
谢角度进行了初步探讨。
在湖北，黑龙江等地调查发现田间用丁草胺防除稗草的效果明显不如以
前。本文以华中地区三种群岳阳（未使用过丁草胺）、湖北漳湖垸农场3队（使用
丁草胺14年）、湖北漳湖垸农场4队（使用丁草胺18年），东北地区三种群牡丹
江（未使用过丁草胺）、民主（使用丁草胺10年）、阿城（使用丁草胺14年）为研究
对象，运用室内盆栽生物测定方法研究了其抗药性现状。试验结果表明，LC_(50)
值以岳阳种群最小，牡丹江种群最大，分别为54.73g a.i./ha和1271.6g a.i./ha；
敏感性由高至低依次为：岳阳、阿城、3队、民主、4队、牡丹江。以岳阳种
群为参照，阿城、3队、民主、4队稗草种群的抗药性系数分别为5.21、5.85、
8.00、12.01，表明长期用药使其产生了对丁草胺的抗药性。而牡丹江种群种子
粒大，千粒重较其它种群显著高，加上可能由于其它除稗剂禾大壮、二氯喹啉
酸等的使用，使其具有获得抗性（或耐药性）的可能。
本文还应用生物化学方法，选择了三个苗龄的稗草来测定各种群幼苗中
GSTs酶活和GSH、Cysteine等自由巯基的含量，初步研究了它们分别与LC_(50)
值之间的相关性。结果表明：（1）当用5ppm丁草胺处理稗草幼苗48小时时，
酶活有不同程度的提高。GSH、半胱氨酸含量差异显著，其中，南方处理种群
GSH含量与LC_(50)值之间正相关，北方处理种群Cysteine含量与LC_(50)值之间呈
负相关。（2）而用5ppm丁草胺处理9天的稗草，处理种群与未处理种群比较，
其酶活有不同程度地提高（阿城种群例外），但各种群酶活与其生测的致死中量
LC_(50)值之间未表现出线性相关性。各未处理种群之间GSH含量差异极其显著，
它们与LC_(50)值之间的线性拟合方程为y=0.0321x+25.029（R~2=0.8382）。（3）种植
14天的稗苗，未用药剂处理，酶活表现出一定的规律性，即：酶活与LC_(50)值
的关系，华中种群为正相关，北方种群为负相关。而南方种群幼苗GSH含量
与LC_(50)值有正相关趋势，北方种群幼苗GSH含量与LC_(50)值有负相关趋势。（4）
测定GST专一性酶活结果表明，所采用的方法未检测到GS-丁草胺，可能是
与GST酶对甲草胺、乙草胺、异丙甲草胺的专一性酶活相比，对丁草胺的酶
活较低，这与Scarponi（1991年）报道的结果一致，其专一性酶活的测定方法需
稻山稗草对丁芋胺的抗性测定及机理初步Uf究
进一步探索。而体外轭合实验表明，在一定条件下非酶结合是可以实现的，这
己通过TLC和质谱获得证实。
总之，田间调查和室内生物测定结果表明我国部分稻田稗草对丁草胺已产
生不同程度的抗药性。通过GST酶活的测定以及簿层层析和质谱分析表明GST
酶代谢丁草胺解毒是其抗或耐药性的机制之一。抗或耐丁草胺的稗草种群GSH
等自由疏基含量和酶活有显著提高，酶活或GSH含量同LC。。值己表现出一定
的相关性，但南方种群与北方种群相关性表现不一致，可能与气候、土壤等环
境条件、作物种植管理水平不同、丁草胺的特殊结构及抗性水平有关：稗草对
丁草胺的抗药性机制还可从其它途径进行研究。
The present status of butachlor resistance of Echinochlor crus-
galli(L.)Beauv.(CE) occurring in paddy rice in north and south of China was studied,
and the resistant mechanism from the point of metabolism was studied. With
continuing application of butachlor in the fields of Hubei and I-Ieilongjiang
Provinces, it was noticed that the herbicidal efficacy for controlling EC was reduced.
Seeds of different EC populations collected from 6 sites (Yueyang, no butachlor
applied, Team III on Zhanghuyuan Farm with 14-year application of butachlor, Team
IV on Zhanghuyuan Farm with 18-year application of butachlor in the south;
Mudanjiang, no butachlor applied, Minzhu with 10-year application of butachlor,
Acheng with 1 4-year application of butachlor in the north)were tested to evaluate
butachlor resistant status in the areas. It was revealed that LC~0 for controlling of
Yueyang population was the lowest, that of Mudanjiang population was the highest
among the six populations, 54.73 g a.i./ha and 1271.6 g a.i.Iha respectively. The
order of sensitivity of the populations from high to low ranged: Yuyang, Acheng,
Team III, Minzhu, Team IV, Mudanjiang. Using Yuyang population as the control,
RI indexes of other populations excluding Mudanjiang population were Acheng 5.21,
Team III 5.85, Minzhu 8.00, Team IV 12.01, which indicated that the four
populations had evolved resistance to bubachlor.
Three seedling stages was selected to test glutathione S-transferase(GST)
activity and the amount of GSH, cysteine, and whether it was correlative with LC50
values were studied. Biochemical tests showed that (1) when the seedlings of EC
were treated with butachlor (5 ppm) for 48 hours, GSTs activities increased, the
amount of GSH, cysteine show significant difference. The amount of GSH of treated
populations was positively correlative to LC0 for controlling of the populations in
the south, the amount of Cysteine of treated populations was negatively correlative
to LC50 in the north. (2) The seedlings of EC were treated with butachlor (Sppm) for
9 days, compared to the untreated populations, the GSTs (CDNB) activities of each
treated population were higher with exception of Acheng population, but the amount
of GSH varied significantly according to the different populations. There was linear
trend between the GSH and LC, i.e. y=0.032l+25.029 (R2=0.8382). (3) GSTs
activity and the amount of GSH of the unbeated seedlings grown for l4 days showed
positive correlation to LC,, for controlling of the POpulations in the south
respectively, but negative correlation to LC,, of those in the north. Although GS-
butachlor was not found in vivo study, it was confirmed in vitro conjugating tests by
TLC and mass spectrum techniques. The free thiols and GSTs activities to CDNB in
EC populations indicatCd that butachlor resistance in EC was increased notably.
GSTs activity or the amoun of GSH showed direct correlation with the LC,, fOr
controlling of different poPulations to some extent. It is suggested that the analysis
method needs to be improved, studying of other possible mechanism is needed.
引文
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49. Hatzios, Kriton K. et al. (1993) In vitro Conjugation of chloroacetanilide Herbicides and Atrazine with Thids and Contribution of Nonenzymatic Conjugation to Their Glutathione-Mediated Metabolism in Corn. J. Agricultural Food Chemistry. vol.41.No.10:1736-1742
50. Heap, Ian M. (1997) The occurrence of herbicide-resistant weeds world-wide. Pestic. Sci. 51,235-243
51. Heap, Ian M. Identification and documentation of herbicide resistance.(1998)Personal communication
52. Herbicide Resistance Action Committee (1997) Guideline to the Management of Herbicide Resistance
53. Jablonkai, Istvan, et al.(1999) Factors Affecting the phytotoxicity of the Herbicide Acetochlor to Monocotyledonous and Dicotyledonous Weeds. Pesticide Science 533:95-397
54. Jakoby, William B.(1985) Glutathione Transferases: An overview. In Methods in Enzymology Vol. 113.
55. Kosower, Edward et al.(1978) 1,5-Diazabicyclo[3,3,0]octadienediones (9,10-Dioxabimanes).Strongly Fluorescent Syn Isomers. Journal of the American chemical society/100:20/september 27,1978.6516-6518.
56. Lebaron, Homer M. (1991) Distribution and seriousness of herbicide-resistant weed infestations worldwide. In: Herbicide Resistance in Weeds and Crops, J.C. Caseley, et al. (eds.). 1991,27-43
57. Matthews, J M (1994) Management of herbicide resistant weed populations. In: Herbicide Resistance in Plants: Biology and Biochemistry. S. B. Powls and J. A. M. Holtum (eds.), Boca Raton FL,CRC Press Inc. 317-335
58. Maxwell, Bruce D. and A. Martin Mortiner (1994) Selection for herbicide resistance. In: Herbicide Resistance in Plants: Biology and Biochemistry. S. B. Powls and J. A. M. Holtum (eds.), Boca Raton, FL, CRC Press Inc.
59. McGonigle, Brian et al.(1998) Homoglutathione Selectivity by Soybean Glutathione S-Transferases. Pesticide Biochemistry and Physidogy. 62:5-25
60. Menendez, J. M.; R. DE Prado; M D Devine(1997) Chlorsulfuron crossresistance in a chlorotoluron-resistant biotype of Alopecurus myosuroides. Proceedings 1997 of Brighton Crop Protection Conference-Weeds 1,319-324
61. Mortimer, M. IRRI(1998) Herbicide resistance in weeds. MEETING AND TRAINING ON HERBICIDE RESISTANCE 29 june-3 July, 1998. Origanised by FAO and Kyungpook National University, Republic of Korea
62. Moss, S. R.(1995) Techniques for determining herbicide resistance. In: BRIGHTON CROP PROTECTION CONFERENCE-WEED-1995.547-556
63. Murray, Bruce G. et al.(1996). A seed Bioassay to Identify Acetyl-CoA carboxylase Inhibitor Resistant Wild Oat (Avena fatua) Populations. Weed Technology. Volume 10:85-89
64. O'connell, K.; M. Breaux, E. J. & Fraley, R. T.,(1988) Different rates of metabolism of two chloroacetanilide herbicides in pioneer 3320 corn. Plant physiol. 86:359-363
65. Ohkawa, Hideo; Hisae Tsujii and Yasunobu Ohkawa(1999) The use of Cytochrome P450 genes to introduce herbicide tolerance in crops a review. Pesticide Science 55:867-874
66. Oudejans, Jan H(1991) Integrated control of weds. In: Agro-pesticides: Properties and Functions in Integrated Crop Protection. ESCAP,Bangkok. 1991,263-290
67. Paisance, Kathryn L. and John W. Gronwald (1999) Enhanced Catalytic Constant for Glutathione S-Transferase(Atrazine) Activity in an Atrazine-Resistant Abutilon theophrasti Biotype. Pesticide Biochemistry and physiology 63:34-49
68. Personal communication (2000) from E-mail: lanHeap@WeedSmart.com Web: http://WeedScience.com
69. Preston, Christopher(1994) Resistance to Photosystem Disrupting Herbicides. In: Herbicide Resistance in Plants: Biology and Biochemistry. S. B. Powls and J. A. M. Holtum (eds.), Boca Raton, FL, CRC Press, Inc.61-81
70. Reade, John PH and Andrew H Cobb (1999) Purification Characterization and comparison of glutathione S-transferases from Black-grass (Alopecurus myosuroides Huds)biotypes Pesticide Science 55:993-999
71. Reade, John PH and Andrew H Cobb(1999) Glutathione S-Transferases in black-grass(Alopecurus myosuroides Huds): properties and involvement in herbicide resistance. Pestic. Sci. 55:349-351
72. Riechers DE, Irxyk GP, Jones S. S. and Fuerst E. P. (1997) Partial characterisation of glutathione S-transferases from wheat (Triticum spp) and purification of a safener-induced glutathione S-transferase form Triticum tauschii. Plant Physiol. 114:1461-1470
73. Robert, I.; C Leavitt and donald Penner(1979) In Vitro Conjugation of Glutathione and Other Thiols with Acetanilide Herbicides and EPTC Sulfoxide and the Action of the Herbicide Antidote R-25788. J. Agric. Food Chem. Vol.27No.3:533-536
74. Roberts, Terry R(1998) Metabolic Pathways of Agrochemicals Part One：Herbicides and Plant Growth Pegulators(ISBNO-85404-494-9) The Royal Society of Chemistry
75. Rossini et al.(1998) Alachlor Regulation of Maize Gluta-thione S-Transferase genes. Pesticide Biochemistry and physiology 60:205-211
76. Rossini, Laura; Carla frova，et al.(1998) Alachlor Regulation of Maize Glutathione S-Transferase Genes. Pesticide Biochemistry and physiology. 60,205-211
77. Scarponi, Luciano, Piero Perucci, and Luca Martinetti (1991). Conjugation of 2-chloroacetanilide Herbicides with Glutathione: Role of Molecular Structures and of Glutathione S-transferase Enzymes. J. Agric. Food Chem. 39:2010-2013
78. Shaner D. L.(1996) Herbicide-resistant crops: a new tool. In: Herbicide-resistant weed management. In: Second International Weed Control Congress Copenhagen. 1996. 421-425
79. Tal A, Romano M. L., Stephenson G. R., Schwan A. L. and Hall J. C.(1993) Glutathione conjugation:a detoxification pathway for fenoxaprop-ethyl in barley, oat and wheat. Pestic Biochem Physiol. 46:190-199
80. Thomas J. Mozer et al. (1983) Purification and characterization of Corn Glutathione S-Transferase. Biochemistry 22:1068-1072
81. Timmerman, Kurt P.(1989) Molecular characterization of corn glutathione Stransferase isozymes involved in herbicide detoxication. Physiologia plantarum. (Copenhagen) 77:465-471
82. Tiwari, Narendra K., et al.(1991) Purification and Characterization of Glutathione Stransferases of Eoreuma Cofiini and Diatraea saccharalis(Lepidoptera: pyralidae). Journal of Economic Entomdogy. Vol.84,No.5:1424-1431
83. Toennies, Gerrit and Joseph J. Kolb(1951) Techniques and Reagents for Paper Chromatography. Analytical Chemistry Vol. 23 No.6:823-826
84. Ulmasov, Tim et al.(1995) The soybean GH2/4 Gene That Encodes a Glutathione S-Transferase Has a promoter That Is Activated by a Wide Range of Chemical Agents. Plant physiol. 108:919-727
85. WRAG(1997) Revised Guidelines for Preventing and Managing Herbicideresistant Grass.
86. Zajc, Alois et al.(1999) Herbicide detoxification By glutathione S-transferases as implicated From X-ray Structures. Pesticede Science 55:248-252
87. ZHANG, Chaoxian (1998) Current herbicide use in weed management system in China. MEETING AND TRAINING ON HERBICIDE RESISTANCE 29 june-3 July, 1998 Organized by FAO and Kyungpook National University, Republic of Korea
88. Zopes, Heinrich; Sigrid Klapheck and Ludwig Bergmann (1993) The Function of Homoglutathione and Hydroxymethylglutathione for the Scavenging of Hydrogen Peroxide. Plant Cell Physiol. 34(4): 515-521
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47. Hatton, Pamela J.; David J. Cole and Robert Edwards (1996) Influence of Plant age on Glutathione levels and Glutathione Transferases Involved in Herbicide Detoxification in Corn (Zea mays )and Giant Foxtail (Setaria faberi Herrm) Pesticide Biochemistry and Physiology 54:199-209
48. Hatton; P.; J. Edwards, R. & Cole, D. J.(1993) Glutathione transferases in major weed species. Pestic. Sci. 43:173-175
49. Hatzios, Kriton K. et al. (1993) In vitro Conjugation of chloroacetanilide Herbicides and Atrazine with Thids and Contribution of Nonenzymatic Conjugation to Their Glutathione-Mediated Metabolism in Corn. J. Agricultural Food Chemistry. vol.41.No.10:1736-1742
50. Heap, Ian M. (1997) The occurrence of herbicide-resistant weeds world-wide. Pestic. Sci. 51,235-243
51. Heap, Ian M. Identification and documentation of herbicide resistance.(1998)Personal communication
52. Herbicide Resistance Action Committee (1997) Guideline to the Management of Herbicide Resistance
53. Jablonkai, Istvan, et al.(1999) Factors Affecting the phytotoxicity of the Herbicide Acetochlor to Monocotyledonous and Dicotyledonous Weeds. Pesticide Science 533:95-397
54. Jakoby, William B.(1985) Glutathione Transferases: An overview. In Methods in Enzymology Vol. 113.
55. Kosower, Edward et al.(1978) 1,5-Diazabicyclo[3,3,0]octadienediones (9,10-Dioxabimanes).Strongly Fluorescent Syn Isomers. Journal of the American chemical society/100:20/september 27,1978.6516-6518.
56. Lebaron, Homer M. (1991) Distribution and seriousness of herbicide-resistant weed infestations worldwide. In: Herbicide Resistance in Weeds and Crops, J.C. Caseley, et al. (eds.). 1991,27-43
57. Matthews, J M (1994) Management of herbicide resistant weed populations. In: Herbicide Resistance in Plants: Biology and Biochemistry. S. B. Powls and J. A. M. Holtum (eds.), Boca Raton FL,CRC Press Inc. 317-335
58. Maxwell, Bruce D. and A. Martin Mortiner (1994) Selection for herbicide resistance. In: Herbicide Resistance in Plants: Biology and Biochemistry. S. B. Powls and J. A. M. Holtum (eds.), Boca Raton, FL, CRC Press Inc.
59. McGonigle, Brian et al.(1998) Homoglutathione Selectivity by Soybean Glutathione S-Transferases. Pesticide Biochemistry and Physidogy. 62:5-25
60. Menendez, J. M.; R. DE Prado; M D Devine(1997) Chlorsulfuron crossresistance in a chlorotoluron-resistant biotype of Alopecurus myosuroides. Proceedings 1997 of Brighton Crop Protection Conference-Weeds 1,319-324
61. Mortimer, M. IRRI(1998) Herbicide resistance in weeds. MEETING AND TRAINING ON HERBICIDE RESISTANCE 29 june-3 July, 1998. Origanised by FAO and Kyungpook National University, Republic of Korea
62. Moss, S. R.(1995) Techniques for determining herbicide resistance. In: BRIGHTON CROP PROTECTION CONFERENCE-WEED-1995.547-556
63. Murray, Bruce G. et al.(1996). A seed Bioassay to Identify Acetyl-CoA carboxylase Inhibitor Resistant Wild Oat (Avena fatua) Populations. Weed Technology. Volume 10:85-89
64. O'connell, K.; M. Breaux, E. J. & Fraley, R. T.,(1988) Different rates of metabolism of two chloroacetanilide herbicides in pioneer 3320 corn. Plant physiol. 86:359-363
65. Ohkawa, Hideo; Hisae Tsujii and Yasunobu Ohkawa(1999) The use of Cytochrome P450 genes to introduce herbicide tolerance in crops a review. Pesticide Science 55:867-874
66. Oudejans, Jan H(1991) Integrated control of weds. In: Agro-pesticides: Properties and Functions in Integrated Crop Protection. ESCAP,Bangkok. 1991,263-290
67. Paisance, Kathryn L. and John W. Gronwald (1999) Enhanced Catalytic Constant for Glutathione S-Transferase(Atrazine) Activity in an Atrazine-Resistant Abutilon theophrasti Biotype. Pesticide Biochemistry and physiology 63:34-49
68. Personal communication (2000) from E-mail: lanHeap@WeedSmart.com Web: http://WeedScience.com
69. Preston, Christopher(1994) Resistance to Photosystem Disrupting Herbicides. In: Herbicide Resistance in Plants: Biology and Biochemistry. S. B. Powls and J. A. M. Holtum (eds.), Boca Raton, FL, CRC Press, Inc.61-81
70. Reade, John PH and Andrew H Cobb (1999) Purification Characterization and comparison of glutathione S-transferases from Black-grass (Alopecurus myosuroides Huds)biotypes Pesticide Science 55:993-999
71. Reade, John PH and Andrew H Cobb(1999) Glutathione S-Transferases in black-grass(Alopecurus myosuroides Huds): properties and involvement in herbicide resistance. Pestic. Sci. 55:349-351
72. Riechers DE, Irxyk GP, Jones S. S. and Fuerst E. P. (1997) Partial characterisation of glutathione S-transferases from wheat (Triticum spp) and purification of a safener-induced glutathione S-transferase form Triticum tauschii. Plant Physiol. 114:1461-1470
73. Robert, I.; C Leavitt and donald Penner(1979) In Vitro Conjugation of Glutathione and Other Thiols with Acetanilide Herbicides and EPTC Sulfoxide and the Action of the Herbicide Antidote R-25788. J. Agric. Food Chem. Vol.27No.3:533-536
74. Roberts, Terry R(1998) Metabolic Pathways of Agrochemicals Part One：Herbicides and Plant Growth Pegulators(ISBNO-85404-494-9) The Royal Society of Chemistry
75. Rossini et al.(1998) Alachlor Regulation of Maize Gluta-thione S-Transferase genes. Pesticide Biochemistry and physiology 60:205-211
76. Rossini, Laura; Carla frova，et al.(1998) Alachlor Regulation of Maize Glutathione S-Transferase Genes. Pesticide Biochemistry and physiology. 60,205-211
77. Scarponi, Luciano, Piero Perucci, and Luca Martinetti (1991). Conjugation of 2-chloroacetanilide Herbicides with Glutathione: Role of Molecular Structures and of Glutathione S-transferase Enzymes. J. Agric. Food Chem. 39:2010-2013
78. Shaner D. L.(1996) Herbicide-resistant crops: a new tool. In: Herbicide-resistant weed management. In: Second International Weed Control Congress Copenhagen. 1996. 421-425
79. Tal A, Romano M. L., Stephenson G. R., Schwan A. L. and Hall J. C.(1993) Glutathione conjugation:a detoxification pathway for fenoxaprop-ethyl in barley, oat and wheat. Pestic Biochem Physiol. 46:190-199
80. Thomas J. Mozer et al. (1983) Purification and characterization of Corn Glutathione S-Transferase. Biochemistry 22:1068-1072
81. Timmerman, Kurt P.(1989) Molecular characterization of corn glutathione Stransferase isozymes involved in herbicide detoxication. Physiologia plantarum. (Copenhagen) 77:465-471
82. Tiwari, Narendra K., et al.(1991) Purification and Characterization of Glutathione Stransferases of Eoreuma Cofiini and Diatraea saccharalis(Lepidoptera: pyralidae). Journal of Economic Entomdogy. Vol.84,No.5:1424-1431
83. Toennies, Gerrit and Joseph J. Kolb(1951) Techniques and Reagents for Paper Chromatography. Analytical Chemistry Vol. 23 No.6:823-826
84. Ulmasov, Tim et al.(1995) The soybean GH2/4 Gene That Encodes a Glutathione S-Transferase Has a promoter That Is Activated by a Wide Range of Chemical Agents. Plant physiol. 108:919-727
85. WRAG(1997) Revised Guidelines for Preventing and Managing Herbicideresistant Grass.
86. Zajc, Alois et al.(1999) Herbicide detoxification By glutathione S-transferases as implicated From X-ray Structures. Pesticede Science 55:248-252
87. ZHANG, Chaoxian (1998) Current herbicide use in weed management system in China. MEETING AND TRAINING ON HERBICIDE RESISTANCE 29 june-3 July, 1998 Organized by FAO and Kyungpook National University, Republic of Korea
88. Zopes, Heinrich; Sigrid Klapheck and Ludwig Bergmann (1993) The Function of Homoglutathione and Hydroxymethylglutathione for the Scavenging of Hydrogen Peroxide. Plant Cell Physiol. 34(4): 515-521