播娘蒿（Descurainia sophia）对苯磺隆的抗药性研究
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摘要
随着化学除草剂的大面积使用,农田杂草抗药性问题越来越严重。苯磺隆是ALS抑制剂类除草剂,目前是小麦田最经济、有效、安全的优秀除草剂品种之一,在我国1988年取得登记,到目前已使用20多年。在部分地区,小麦田以常规剂量喷施苯磺隆已经无法有效防除麦田主要杂草播娘蒿(Descurainia sophia)。明确播娘蒿对苯磺隆的抗药性水平及其抗药性机理,对指导抗药性播娘蒿的科学治理,延长苯磺隆的使用寿命具有重要的理论和现实意义。本研究以采自北京、天津、河北、河南、山东、山西、陕西、甘肃、青海、江苏及四川省(市)的播娘蒿为对象,研究其对苯磺隆的抗药性水平及其抗药性机理,主要结果如下。
1.整株测定结果表明,在11省(市)91个播娘蒿种群中,有42个种群对苯磺隆表现敏感,GR50值在0.10-0.12g a.i./ha之间;19个种群对苯磺隆具有较低水平的抗药性,1<抗药性指数≤10;19个种群对苯磺隆具有中等抗药性,10<抗药性指数≤100;11个种群对苯磺隆具有较高水平的抗药性,其抗药性指数在100以上。HB-8、TJ-8和GS-5种群中有个别植株抗药性很强,平均抗药性百分率分别是2.38%、1.14%和3.64%。
2. SSX-9、SSX-10、HB-8、TJ-8和GS-5种群二代对苯磺隆的抗药性检测结果表明,对苯磺隆的抗药性能够稳定遗传。
3. HB-1、SSX-8、HB-2、HB-8、SSX-4、SSX-9、SSX-10、SSX-11、SSX-12、SSX-13、TJ-8和GS-5种群ALS离体活性测定结果表明,抗药性种群的ALS对苯磺隆的敏感性降低,抗药性指数在3.17-86.33之间。
4. HB-1、HB-2、SSX-8和SSX-9种群ALS活体活性测定结果表明,抗药性种群ALS活性受到抑制后能恢复,而敏感种群则不能。
5. 4个敏感生物型和10个抗药性生物型的ALS基因克隆测序结果表明,播娘蒿ALS全长2,004bp,编码667个氨基酸,无内含子,所检测的河北、陕西、甘肃和天津的4个敏感生物型的ALS序列完全相同。
6.比较敏感和抗药性生物型的ALS序列发现,抗药性生物型在Pro197(脯氨酸)发生了突变,抗药性生物型HB-2和SSX-11 Pro197突变为Leu197(亮氨酸);SSX-9 Pro197突变为Thr197(苏氨酸);SSX-12和SSX-13 Pro197突变为Ala197(丙氨酸);TJ-8和GS-5 Pro197突变为Ser197(丝氨酸);敏感和抗药性生物型的ALS序列在GenBank上的登记号为EU520489、EU520490、EU520491和FJ715633。
7.播娘蒿种群HB-1、HB-2、SSX-8、SSX-4、SSX-9和SSX-10对ALS抑制剂包括咪唑乙烟酸(IMI类)、嘧草硫醚(PTB类)、氯酯磺草胺(TP类)、甲氧磺草胺(TP类)、双氟磺草胺(TP类)的交互抗性的测定结果表明,HB-1和SSX-8对所有供试药剂均敏感;HB-2对各供试药剂均有较高的抗药性,抗药性指数在10倍以上;SSX-4除了对咪唑乙烟酸较敏感之外,对其他药剂的抗药性指数在10倍以上;SSX-9和SSX-10对咪唑乙烟酸有低水平的抗药性,对嘧草硫醚有高水平的抗药性,对供试的三种TP类药剂都有不同程度的抗药性。SSX-4种群里个别植株对上述几种药剂抗药性很高,对嘧草硫醚、咪唑乙烟酸和双氟磺草胺的平均抗药性百分率分别为4.51%、2.71%和2.21%。
本研究首次明确了我国麦田播娘蒿种群对苯磺隆的抗药性水平与分布,及对其他5种ALS抑制剂类除草剂交互抗性水平;从酶和基因水平揭示了抗药性是由于靶标酶ALS基因突变所致。
Herbicide resistance is a worldwide phenomenon with the development of the chemical herbicides’application in agriculture, and it becomes more serious. Tribenuron-methyl is one of the most popular ALS inhibitor in wheat field with low price, high efficacy and good safety. After being applied over 20 years, some farmers complained that the flixweed (Descurainia sophia), one of the most important and widely infested broad leaved weeds in wheat fields was not sensitive to tribenuron-methyl any more which was registered in China in 1988. It is very important and necessary to confirm flixweed resistance and its mechanism to tribenuron-methyl in order to control flixweed effectively and use tribenuron-methyl in a scientific, rational way and to prolong the lifespan of tribenuron-methyl in practice. Flixweed seeds were collected from separate wheat fields that had been treated with tribenuron-methyl repeatedly over 3-15 years, and from road sides, remote hills that never received tribenuron-methyl treatment in Beijing and Tianjin Metropolitans, Hebei, Henan, Shandong, Shanxi, Shaanxi, Gansu, Qianghai, Jiangsu and Sichuan provinces with various herbicide application histories. Flixweed sensitivity to tribenuron-methyl was determined by whole plant experiments in the greenhouse, its resistant level and molecular mechanism were also conducted. The results showed as followings.
1. Results of whole plant experiments showed that 42 of the 91 flixweed populations were susceptible to tribenuron-methyl with resistance indices ranging from 0.10 to 0.12g a.i./ha, 19 of the 91 populations expressed low level resistance to tribenuron-methyl with resistance indices ranging in 1 2. The seeds of resistant populations SSX-9, SSX-10, HB-8, TJ-8 and GS-5 were harvested and planted in potted moist loam soil. Tribenuron-methyl was applied at different doses at the four-leaf growth stage of these populations. The results revealed that these populations showed high level resistance, suggesting that the resistance of these populations could be inherited.
3. The ALS activity of the populations HB-1, SSX-8, HB-2, HB-8, SSX-4, SSX-9, SSX-10, SSX-11, SSX-12, SSX-13, TJ-8 and GS-5 treated with tribenuron-methyl was detected and compared. The ALS activity of the susceptible populations SSX-8 and HB-1 was significantly inhibited. However, the activity of ALS of the resistant populations decreased slowly under tribenuron-methyl treatment, the relative I50 of the above populations ranged from 3.17 to 86.33.
4. The ALS activity-time curves of flixweed populations HB-1, HB-2, SSX-8 and SSX-9 showed that the ALS activity of the resistant populations was decreased firstly, then turned to increase slowly; the ALS activity of the susceptible populations, however, was inhibited significantly, and was irreversible.
5. The ALS genes of 4 susceptible biotypes and 10 resistant biotypes were cloned and sequenced, and the full length ALS gene being translated 667 amino acids which was approximately 2,004 nucleotides without any intron. The ALS genes of the 4 susceptible biotypes collected from Hebei, Shaanxi, Gansu and Tianjin were sequenced, and their sequences were identical. 6. Comparison of the ALS gene sequences from susceptible and resistant biotypes with Arabidopsis revealed that proline at position 197 of the ALS gene was substituted by leucine in resistant biotypes HB-2 and SSX-11, by threonine in resistant biotype SSX-9, by alanine in resistant biotypes SSX-12 and SSX-13, by serine in resistant biotypes TJ-8 and GS-5. The nucleotide sequences of susceptible and resistant biotypes were registered in the Genbank, with accession numbers EU520489, EU520490, EU520491 and FJ715633, respectively.
7. The experiments were conducted with the ALS inhibitors, tribenuron-methyl (SU type), imazethapyr (IMI type), pyrithiobac-sodium (PTB type), cloransulam-methyl (TP type), pyroxsulam (TP type) and florasulam (TP type) to detect the cross-resistance of flixweed populations HB-1, HB-2, SSX-8, SSX-4, SSX-9 and SSX-10 to the ALS inhibitors. The results showed that populations HB-1 and SSX-8 were very susceptible to all tested herbicides. Population HB-2 was resistant to all of them, with resistant indices over 10. The population SSX-4 was resistant to the herbicides tested except imazethapyr, with resistant indices over 10. The populations SSX-9 and SSX-10 were resistant to tribenuron-methyl and three TP herbicides, but susceptible to imazethapyr. Some plants of the population SSX-4 were resistant to the aboved ALS inhibitors. The average resistant percentage of the population SSX-4 to pyrithiobac-sodium was 4.51%, 2.71% to imazethapyr, and 2.21% to florasulam, respectively.
In general, the resistant levels and distributions of flixweed populations to tribenuron-methyl and its cross-resistance of these populations to five different chemical classes of ALS inhibitors were first confirmed in China. The resistant mechanism being confered to the point mutation in ALS gene were first revealed in enzyme and gene level.
1.整株测定结果表明,在11省(市)91个播娘蒿种群中,有42个种群对苯磺隆表现敏感,GR50值在0.10-0.12g a.i./ha之间;19个种群对苯磺隆具有较低水平的抗药性,1<抗药性指数≤10;19个种群对苯磺隆具有中等抗药性,10<抗药性指数≤100;11个种群对苯磺隆具有较高水平的抗药性,其抗药性指数在100以上。HB-8、TJ-8和GS-5种群中有个别植株抗药性很强,平均抗药性百分率分别是2.38%、1.14%和3.64%。
2. SSX-9、SSX-10、HB-8、TJ-8和GS-5种群二代对苯磺隆的抗药性检测结果表明,对苯磺隆的抗药性能够稳定遗传。
3. HB-1、SSX-8、HB-2、HB-8、SSX-4、SSX-9、SSX-10、SSX-11、SSX-12、SSX-13、TJ-8和GS-5种群ALS离体活性测定结果表明,抗药性种群的ALS对苯磺隆的敏感性降低,抗药性指数在3.17-86.33之间。
4. HB-1、HB-2、SSX-8和SSX-9种群ALS活体活性测定结果表明,抗药性种群ALS活性受到抑制后能恢复,而敏感种群则不能。
5. 4个敏感生物型和10个抗药性生物型的ALS基因克隆测序结果表明,播娘蒿ALS全长2,004bp,编码667个氨基酸,无内含子,所检测的河北、陕西、甘肃和天津的4个敏感生物型的ALS序列完全相同。
6.比较敏感和抗药性生物型的ALS序列发现,抗药性生物型在Pro197(脯氨酸)发生了突变,抗药性生物型HB-2和SSX-11 Pro197突变为Leu197(亮氨酸);SSX-9 Pro197突变为Thr197(苏氨酸);SSX-12和SSX-13 Pro197突变为Ala197(丙氨酸);TJ-8和GS-5 Pro197突变为Ser197(丝氨酸);敏感和抗药性生物型的ALS序列在GenBank上的登记号为EU520489、EU520490、EU520491和FJ715633。
7.播娘蒿种群HB-1、HB-2、SSX-8、SSX-4、SSX-9和SSX-10对ALS抑制剂包括咪唑乙烟酸(IMI类)、嘧草硫醚(PTB类)、氯酯磺草胺(TP类)、甲氧磺草胺(TP类)、双氟磺草胺(TP类)的交互抗性的测定结果表明,HB-1和SSX-8对所有供试药剂均敏感;HB-2对各供试药剂均有较高的抗药性,抗药性指数在10倍以上;SSX-4除了对咪唑乙烟酸较敏感之外,对其他药剂的抗药性指数在10倍以上;SSX-9和SSX-10对咪唑乙烟酸有低水平的抗药性,对嘧草硫醚有高水平的抗药性,对供试的三种TP类药剂都有不同程度的抗药性。SSX-4种群里个别植株对上述几种药剂抗药性很高,对嘧草硫醚、咪唑乙烟酸和双氟磺草胺的平均抗药性百分率分别为4.51%、2.71%和2.21%。
本研究首次明确了我国麦田播娘蒿种群对苯磺隆的抗药性水平与分布,及对其他5种ALS抑制剂类除草剂交互抗性水平;从酶和基因水平揭示了抗药性是由于靶标酶ALS基因突变所致。
Herbicide resistance is a worldwide phenomenon with the development of the chemical herbicides’application in agriculture, and it becomes more serious. Tribenuron-methyl is one of the most popular ALS inhibitor in wheat field with low price, high efficacy and good safety. After being applied over 20 years, some farmers complained that the flixweed (Descurainia sophia), one of the most important and widely infested broad leaved weeds in wheat fields was not sensitive to tribenuron-methyl any more which was registered in China in 1988. It is very important and necessary to confirm flixweed resistance and its mechanism to tribenuron-methyl in order to control flixweed effectively and use tribenuron-methyl in a scientific, rational way and to prolong the lifespan of tribenuron-methyl in practice. Flixweed seeds were collected from separate wheat fields that had been treated with tribenuron-methyl repeatedly over 3-15 years, and from road sides, remote hills that never received tribenuron-methyl treatment in Beijing and Tianjin Metropolitans, Hebei, Henan, Shandong, Shanxi, Shaanxi, Gansu, Qianghai, Jiangsu and Sichuan provinces with various herbicide application histories. Flixweed sensitivity to tribenuron-methyl was determined by whole plant experiments in the greenhouse, its resistant level and molecular mechanism were also conducted. The results showed as followings.
1. Results of whole plant experiments showed that 42 of the 91 flixweed populations were susceptible to tribenuron-methyl with resistance indices ranging from 0.10 to 0.12g a.i./ha, 19 of the 91 populations expressed low level resistance to tribenuron-methyl with resistance indices ranging in 1
3. The ALS activity of the populations HB-1, SSX-8, HB-2, HB-8, SSX-4, SSX-9, SSX-10, SSX-11, SSX-12, SSX-13, TJ-8 and GS-5 treated with tribenuron-methyl was detected and compared. The ALS activity of the susceptible populations SSX-8 and HB-1 was significantly inhibited. However, the activity of ALS of the resistant populations decreased slowly under tribenuron-methyl treatment, the relative I50 of the above populations ranged from 3.17 to 86.33.
4. The ALS activity-time curves of flixweed populations HB-1, HB-2, SSX-8 and SSX-9 showed that the ALS activity of the resistant populations was decreased firstly, then turned to increase slowly; the ALS activity of the susceptible populations, however, was inhibited significantly, and was irreversible.
5. The ALS genes of 4 susceptible biotypes and 10 resistant biotypes were cloned and sequenced, and the full length ALS gene being translated 667 amino acids which was approximately 2,004 nucleotides without any intron. The ALS genes of the 4 susceptible biotypes collected from Hebei, Shaanxi, Gansu and Tianjin were sequenced, and their sequences were identical. 6. Comparison of the ALS gene sequences from susceptible and resistant biotypes with Arabidopsis revealed that proline at position 197 of the ALS gene was substituted by leucine in resistant biotypes HB-2 and SSX-11, by threonine in resistant biotype SSX-9, by alanine in resistant biotypes SSX-12 and SSX-13, by serine in resistant biotypes TJ-8 and GS-5. The nucleotide sequences of susceptible and resistant biotypes were registered in the Genbank, with accession numbers EU520489, EU520490, EU520491 and FJ715633, respectively.
7. The experiments were conducted with the ALS inhibitors, tribenuron-methyl (SU type), imazethapyr (IMI type), pyrithiobac-sodium (PTB type), cloransulam-methyl (TP type), pyroxsulam (TP type) and florasulam (TP type) to detect the cross-resistance of flixweed populations HB-1, HB-2, SSX-8, SSX-4, SSX-9 and SSX-10 to the ALS inhibitors. The results showed that populations HB-1 and SSX-8 were very susceptible to all tested herbicides. Population HB-2 was resistant to all of them, with resistant indices over 10. The population SSX-4 was resistant to the herbicides tested except imazethapyr, with resistant indices over 10. The populations SSX-9 and SSX-10 were resistant to tribenuron-methyl and three TP herbicides, but susceptible to imazethapyr. Some plants of the population SSX-4 were resistant to the aboved ALS inhibitors. The average resistant percentage of the population SSX-4 to pyrithiobac-sodium was 4.51%, 2.71% to imazethapyr, and 2.21% to florasulam, respectively.
In general, the resistant levels and distributions of flixweed populations to tribenuron-methyl and its cross-resistance of these populations to five different chemical classes of ALS inhibitors were first confirmed in China. The resistant mechanism being confered to the point mutation in ALS gene were first revealed in enzyme and gene level.
引文
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