田间棉铃虫对Bt棉花的耐性演化分析
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摘要
Bt-Cry1Ac转基因棉花于1997年在我国商业化种植以来,有效控制了棉铃虫Helicoverpa armigera的发生和为害,但人类对大规模种植Bt棉所带来的长期生态风险问题尚缺乏足够的认识。本文采用诊断剂量与DNA分子检测相结合的方法,于2006~2008年对位于我国黄河流域棉区的山东省夏津县和河北省安次县的棉铃虫种群对Bt棉耐性演化规律进行了系统监测,并根据我国不同Bt棉种植强度模式下种群发生动态,建立了种群动态模型,评价了Bt棉在不同种植模式下如何调控棉铃虫田间种群演化动态,并就目前田间棉铃虫种群对Cry1Ac和Cry2Ab两种蛋白耐性遗传相关性进行了研究。主要研究结果如下:
1在2006~2008年期间,在山东夏津和河北安次棉区分别共测定了2306和1270对单雌家系,保守的估计了夏津棉铃虫种群对Cry1Ac的抗性基因频率在2006年、2007年和2008年分别为0, 0.00022,0.00033;在安次县,三年期间均未检测到抗性家系,因此,在安次地区估计的抗性基因频率为0。该研究结果表明田间棉铃虫种群对Cry1Ac抗性基因频率在夏津和安次地区均处于很低水平(<0.0001)。在2006~2008年期间,夏津和安次县棉铃虫种群F1代幼虫在Cry1Ac饲料上的相对平均发育级别均值并没有显著性的提高,说明田间棉铃虫种群对Bt-Cry1Ac耐性这几年并没有发生明显的变化。
2根据室内已报道的与棉铃虫产生抗性相关的钙粘蛋白与APN突变位点,设计特异引物,开展田间棉铃虫抗性基因频率DNA分子检测。2006~2008年期间,我们共检测了2570头蛾子DNA,没有发现相关的突变基因在田间存在。
3两种Bt棉不同筛选强度模式下,田间棉铃虫种群动态模拟模型结果表明:棉铃虫种群数量与Bt棉在我国华北棉区种植年份呈显著的负相关,这种负相关性说明Bt棉的种植显著抑制了田间棉铃虫的种群数量。一般来说,在田间每年的6月~10月份,棉铃虫发生三代高峰期,但有趣的是,在Bt棉高密度种植区(夏津和临清棉区),田间棉铃虫种群发生代别数由抗虫棉种植前的三代演变为两代,第二代棉铃虫种群基本削弱,但这种趋势在Bt棉低密度种植区没有,且证明该代别演化趋势与Bt棉种植密度有显著的相关性。
4在2008年,夏津县和安次县分别共有572个家系和124个家系同时进行了Cry1Ac和Cry2Ab测定,测定结果表明:夏津和安次两地区棉铃虫家系的F1代对Cry1Ac相对平均发育级别与对Cry2Ab相对平均发育级别呈明显的正相关,且该相关性在F2代也存在,说明目前田间棉铃虫种群对Cry1Ac和Cry2Ab两种蛋白的耐性呈显著遗传相关性。
本研究采用诊断剂量生测与DNA分子检测技术开展抗性基因频率监测,研究结果均表明,目前田间棉铃虫种群对Bt-Cry1Ac棉的抗性基因频率还处于较低水平,不同棉区棉铃虫基因交流、Bt棉高强度种植区第二代棉铃虫种群削弱,进而阻断抗性基因的累积在延缓棉铃虫耐性发展方面发挥了积极的作用。同时,开展田间棉铃虫种群对Cry2Ab蛋白耐性反应、建立Cry2Ab敏感基线,在评价第二代Bt抗虫棉(Cry1Ac/Cry2Ab)策略来延缓棉铃虫抗性进化提供科学依据。
Genetically modified cotton expressing the Cry1Ac toxin has been commercially cultivated in China since 1997. Bt cotton provides an efficient tool for controlling cotton bollworm (CBW) Helicoverpa armigera (Hübner) in many areas of China, but little is know about their long-term ecological consequences. During 2006-2008, the sensitivities of isofemale lines of H. armigera collected from Xiajin County, Shandong Province and Anci County, Hebei Province in northern China to Cry1Ac toxin protein were monitored systematically by using both diagnostic concentration and DNA screen methods, and population dynamic models were established under different planting system in northern China. In addition, interactions between Cry1Ac and Cry2Ab in the field H. armigera populations were investigated. The main results were summarized as follows.
1. During 2006~2008, a total of 2306 isofemale lines from the Xiajin population and a total of 1270 isofemale lines from the Anci population were successfully screened on Cry1Ac diets. For each year, it was estimated that the major resistance gene frequency in Xiajin population in 2006, 2007, and 2008 was 0, 0.00022, and 0.00033, respectively. No major alleles conferring resistance to Cry1Ac were found in Anci population, the frequency of resistance alleles for Cry1Ac was 0. Based on the relative average development rates (RADR) of the H. armigera larvae in F1 tests, no substantial increase in Cry1Ac tolerance was found in either location over the 3-yr period. There were also significantly positive correlations between RADR of the lines in F1 generation and the RADR of their F2 offspring, indicating that the tolerance was genetically based.
2. We used polymerase chain reaction (RCR) primers that specifically amplify one mutant allele of a cadherin and one mutant allele of a APN gene linked with resistance to Cry1Ac in H. armigera, a major pest. We screened DNA of 2,570 insects derived from Cry1Ac cotton fields in Xiajin and Anci during 2006~2008. No resistance alleles were detected despite a decade of exposure to Bt cotton.
3. There was a negative correlation between moth densities of H. armigera and planting years of Bt cotton for the Bt cotton planting regions, which indicated that population density of H. armigera were reduced with the introduction of Bt cotton in northern China. There are three generations of moth occurred between early June and late September in the Bt cotton planting regions. Interestingly, the 2nd generation moths decreased and seemed to vanish in recent years in two intensive Bt cotton planting regions (Xiajin County and Linqing County of Shandong Province), but this tendency was not found in other two less intensive Bt cotton planting regions (Anci County of Hebei Province and Fengqiu County of Henan Province). This indicated that deployment of Bt cotton, especially in the intensive Bt cotton planting areas, have deeply suppressed the 2nd generation significantly.
4. A total of 572 isofemale families of H. armigera from Xiajin county of Shandong Province and 124 families from Anci county of Hebei Province were screened with both Cry1Ac- and Cry2Ab-containing diets in 2008. The bioassays results indicated that RADR of F1 full-sib families from field collected female moths on Cry1Ac and Cry2Ab containing diet were positively correlated. The same correlation was found in the F2 generation, indicating cross-tolerance between Cry1Ac and Cry2Ab in field populations of H. armigera in Yellow River cotton-farming region of China.
In conjunction with data from bioassays and DNA based-screen, the present results reported here indicated that the frequency for Bt resistance allele are still rare in the current field population of H. armigera. The complete control of second generation moths minimizes the impact of minor resistance genes and would be important factors contributing to the continued susceptibility to Cry1Ac in field population of H. armigera in Yellow River cotton farming region of China. Furthermore, the cross-tolerance must be considered in evaluating the utility of pyramiding Bt genes (Cry1Ac/Cry2Ab) in cotton for delaying evolution of resistance.
1在2006~2008年期间,在山东夏津和河北安次棉区分别共测定了2306和1270对单雌家系,保守的估计了夏津棉铃虫种群对Cry1Ac的抗性基因频率在2006年、2007年和2008年分别为0, 0.00022,0.00033;在安次县,三年期间均未检测到抗性家系,因此,在安次地区估计的抗性基因频率为0。该研究结果表明田间棉铃虫种群对Cry1Ac抗性基因频率在夏津和安次地区均处于很低水平(<0.0001)。在2006~2008年期间,夏津和安次县棉铃虫种群F1代幼虫在Cry1Ac饲料上的相对平均发育级别均值并没有显著性的提高,说明田间棉铃虫种群对Bt-Cry1Ac耐性这几年并没有发生明显的变化。
2根据室内已报道的与棉铃虫产生抗性相关的钙粘蛋白与APN突变位点,设计特异引物,开展田间棉铃虫抗性基因频率DNA分子检测。2006~2008年期间,我们共检测了2570头蛾子DNA,没有发现相关的突变基因在田间存在。
3两种Bt棉不同筛选强度模式下,田间棉铃虫种群动态模拟模型结果表明:棉铃虫种群数量与Bt棉在我国华北棉区种植年份呈显著的负相关,这种负相关性说明Bt棉的种植显著抑制了田间棉铃虫的种群数量。一般来说,在田间每年的6月~10月份,棉铃虫发生三代高峰期,但有趣的是,在Bt棉高密度种植区(夏津和临清棉区),田间棉铃虫种群发生代别数由抗虫棉种植前的三代演变为两代,第二代棉铃虫种群基本削弱,但这种趋势在Bt棉低密度种植区没有,且证明该代别演化趋势与Bt棉种植密度有显著的相关性。
4在2008年,夏津县和安次县分别共有572个家系和124个家系同时进行了Cry1Ac和Cry2Ab测定,测定结果表明:夏津和安次两地区棉铃虫家系的F1代对Cry1Ac相对平均发育级别与对Cry2Ab相对平均发育级别呈明显的正相关,且该相关性在F2代也存在,说明目前田间棉铃虫种群对Cry1Ac和Cry2Ab两种蛋白的耐性呈显著遗传相关性。
本研究采用诊断剂量生测与DNA分子检测技术开展抗性基因频率监测,研究结果均表明,目前田间棉铃虫种群对Bt-Cry1Ac棉的抗性基因频率还处于较低水平,不同棉区棉铃虫基因交流、Bt棉高强度种植区第二代棉铃虫种群削弱,进而阻断抗性基因的累积在延缓棉铃虫耐性发展方面发挥了积极的作用。同时,开展田间棉铃虫种群对Cry2Ab蛋白耐性反应、建立Cry2Ab敏感基线,在评价第二代Bt抗虫棉(Cry1Ac/Cry2Ab)策略来延缓棉铃虫抗性进化提供科学依据。
Genetically modified cotton expressing the Cry1Ac toxin has been commercially cultivated in China since 1997. Bt cotton provides an efficient tool for controlling cotton bollworm (CBW) Helicoverpa armigera (Hübner) in many areas of China, but little is know about their long-term ecological consequences. During 2006-2008, the sensitivities of isofemale lines of H. armigera collected from Xiajin County, Shandong Province and Anci County, Hebei Province in northern China to Cry1Ac toxin protein were monitored systematically by using both diagnostic concentration and DNA screen methods, and population dynamic models were established under different planting system in northern China. In addition, interactions between Cry1Ac and Cry2Ab in the field H. armigera populations were investigated. The main results were summarized as follows.
1. During 2006~2008, a total of 2306 isofemale lines from the Xiajin population and a total of 1270 isofemale lines from the Anci population were successfully screened on Cry1Ac diets. For each year, it was estimated that the major resistance gene frequency in Xiajin population in 2006, 2007, and 2008 was 0, 0.00022, and 0.00033, respectively. No major alleles conferring resistance to Cry1Ac were found in Anci population, the frequency of resistance alleles for Cry1Ac was 0. Based on the relative average development rates (RADR) of the H. armigera larvae in F1 tests, no substantial increase in Cry1Ac tolerance was found in either location over the 3-yr period. There were also significantly positive correlations between RADR of the lines in F1 generation and the RADR of their F2 offspring, indicating that the tolerance was genetically based.
2. We used polymerase chain reaction (RCR) primers that specifically amplify one mutant allele of a cadherin and one mutant allele of a APN gene linked with resistance to Cry1Ac in H. armigera, a major pest. We screened DNA of 2,570 insects derived from Cry1Ac cotton fields in Xiajin and Anci during 2006~2008. No resistance alleles were detected despite a decade of exposure to Bt cotton.
3. There was a negative correlation between moth densities of H. armigera and planting years of Bt cotton for the Bt cotton planting regions, which indicated that population density of H. armigera were reduced with the introduction of Bt cotton in northern China. There are three generations of moth occurred between early June and late September in the Bt cotton planting regions. Interestingly, the 2nd generation moths decreased and seemed to vanish in recent years in two intensive Bt cotton planting regions (Xiajin County and Linqing County of Shandong Province), but this tendency was not found in other two less intensive Bt cotton planting regions (Anci County of Hebei Province and Fengqiu County of Henan Province). This indicated that deployment of Bt cotton, especially in the intensive Bt cotton planting areas, have deeply suppressed the 2nd generation significantly.
4. A total of 572 isofemale families of H. armigera from Xiajin county of Shandong Province and 124 families from Anci county of Hebei Province were screened with both Cry1Ac- and Cry2Ab-containing diets in 2008. The bioassays results indicated that RADR of F1 full-sib families from field collected female moths on Cry1Ac and Cry2Ab containing diet were positively correlated. The same correlation was found in the F2 generation, indicating cross-tolerance between Cry1Ac and Cry2Ab in field populations of H. armigera in Yellow River cotton-farming region of China.
In conjunction with data from bioassays and DNA based-screen, the present results reported here indicated that the frequency for Bt resistance allele are still rare in the current field population of H. armigera. The complete control of second generation moths minimizes the impact of minor resistance genes and would be important factors contributing to the continued susceptibility to Cry1Ac in field population of H. armigera in Yellow River cotton farming region of China. Furthermore, the cross-tolerance must be considered in evaluating the utility of pyramiding Bt genes (Cry1Ac/Cry2Ab) in cotton for delaying evolution of resistance.
引文
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