应用单雌系F_2代法监测田间棉铃虫种群对转Bt基因棉的抗性
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摘要
表达CrylAc毒蛋白的转Bt基因棉花在中国北方地区种植多年,虽然Bt棉因其高效和对环境低风险迅速受到种植者的青睐,但是如此商业化大面积推广使用Bt棉将会增加靶标害虫对Bt棉产生抗性的风险,室内研究已经表明棉铃虫能对CrylAc毒素和Bt棉产生高抗性。
在中国河北邱县地区,1998年开始推广种植Bt棉,1999年采用单雌系F_2代法检测到邱县地区田间棉铃虫野生种群对Bt棉初始抗性等位基因频率达到了0.0058。考虑到棉铃虫在如此高的抗性初始频率值和持续的田间选择压情况下,有可能对Bt棉产生抗性风险。2003-2007年在前人研究的基础上,我们采用单雌系F_2代法、剂量反应法、F_1代法和DNA分子生物学方法相结合的综合抗性监(检)测体系,研究长期大面积种植转Bt基因棉地区棉铃虫田间种群对Bt棉的抗性发展趋势,为设计制定合理有效的抗性治理策略提供及时、准确的科学依据,以延长转Bt基因棉在大田的使用。
2003-2005年采用Bt棉株作为筛选媒介,对河北邱县(2003-2005年)和威县(2004年)地区田间棉铃虫种群进行单雌系F_2代法抗性监测。结果表明,这两个地区3年监测的340对单雌系中有20对被确定为携带有对Bt棉抗性的等位基因,棉铃虫种群的抗性等位基因频率(E(q))范围为0.0119-0.0297。
其中邱县地区连续3年监测到的棉铃虫对Bt棉的平均抗性等位基因频率值(E(q))为0.0146(95%CI=0.0084-0.0225),明显高于1999年的监测结果(p=0.042);2003-2005年分别监测了105对、42对和131对单雌系,平均每个单雌系在Bt棉株上分别处理筛选了372.4±16.0头、328.1±9.5头和314.0±9.0头F_2代初孵幼虫,最终分别有4对、4对和7对单雌系被确定为田间亲本携带有抗性基因的阳性反应单雌系,每年的抗性等位基因频率值分别为0.0119(95%CI=0.0039-0.0243)、0.0297(95%CI=0.0099-0.0606)和0.0154(95%CI=0.0067-0.0277),监测处理单雌系的总抗性等位基因检出累计概率值(1-P_(NO))分别为96%、97%和94%;
威县地区2004年共筛选了62对单雌系,平均每个单雌系筛选了334.7±8.6头F_2代初孵幼虫,最终有5个单雌系被确定田间亲本携带有抗性基因,抗性等位基因频率值为0.0243(95%CI=0.0091-0.0471),单雌系总的抗性等位基因检出累计概率值(1-P_(NO))为95%。结果表明威县地区棉铃虫野生种群对Bt棉的抗性等位基因频率值处在一个较高的阈值,并且超过了高剂量/庇护区策略的要求。
采用Bt棉叶喂饲法处理室内敏感、抗性及两者杂交后代的初孵幼虫,通过四者在转基因棉花上5天生长发育速率的不同来加以区分。试验结果表明,敏感与抗性品系棉铃虫在Bt棉上5天的存活率与生长发育速率存在明显差异,敏感幼虫不能在转Bt基因棉叶上存活,抗性品系在转Bt基因棉叶上处理5天的存活率达到75%,正反交的幼虫存活率仅为2-6%.棉铃虫抗性品系存活幼虫中65%个体在Bt棉上发育到2龄中期、0.6mg以上;而杂合子和敏感纯合子存活幼虫均达不到此生长发育速率。因此可以确定棉铃虫幼虫在Bt棉上5天发育到2龄中期以上、体重≥0.6mg的个体为抗性个体。
2006-2007年改进单雌系F_2代法,采用Bt棉叶作为筛选媒介进行邱县田间棉铃虫抗性监测。2006年和2007年共筛选了134对和137对单雌系,平均每个单雌系分别筛选了187.9±3.4头和105.1±1.5头F_2代初孵幼虫,抗性等位基因频率值分别为0.1135(95%CI=0.0862-0.1449)和0.0745(95%CI=0.0532-0.0995),2006年和2007年单雌系总的抗性等位基因检出累计概率值(1-P_(No))分别为95%和90%。2007年,采用大田雄虫与室内抗性雌虫杂交的F_1代法监测邱县棉铃虫抗性等位基因频率,共筛选了135对单雌系,平均每个单雌系筛选了165.2±5.2头F_1代初孵幼虫,经过F_2代核查后最终确定有29对单雌系的田间亲本携带有抗性基因,抗性等位基因频率值为0.107(95%CI=0.055-0.159)。
2003-2007年,采用剂量反应法监测邱县、威县地区棉铃虫种群对CrylAc毒蛋白的抗性水平。5年的研究结果显示,田间棉铃虫对CrylAc毒蛋白的抗性倍数分别为15.3、12.9、12.5、6.7、16.5和11.5,基本达到了中等水平抗性。
2006-2007年,将单雌系F_2代法确定为携带有对Bt棉抗性等位基因的田间亲本提取基因组DNA,利用根据室内抗性和敏感棉铃虫钙粘素受体基因序列设计的三个引物对田间抗性棉铃虫进行PASA分子检测,探索性的研究这三个引物在田间抗性检测中的应用。2006年共检测了50头田间亲本成虫,2007年共检测了44头田间亲本成虫。结果显示,根据室内抗、敏棉铃虫钙粘素受体基因设计的PASA引物无法有效的应用于田间棉铃虫Bt抗性检测,田间Bt抗性分子检测技术还有待进一步的改进。
通过长达5年的田间抗性监测研究发现,邱县等大面积种植表达单一CrylAc毒蛋白Bt棉的地区,田间棉铃虫种群对Bt棉的抗性等位基因频率已经显著上升,证实了害虫抗药性发展模式中,在初始抗性等位基因频率值较高(＞0.005)地区的害虫,对Bt棉的抗性发展较快的这一推论。类似于邱县作物种植种类和结构的地区需要采取有效、灵敏的抗性检测与监测体系进行害虫的Bt抗性研究,同时结合田间生态环境,建立合理、有效的抗性管理策略,从而延长转Bt基因作物的应用。
Transgenic Bacillus thuringiensis(Bt) cotton producing Cry1Ac protein has been widely planted in northern China for a long time.The high degree of effectiveness and low environmental risk of Bt crops have encouraged Bt cotton producers to rapidly adopt this technology.The rapid and extensive commercialization of Bt cotton could increase the risk of resistance development in target insects.The studies showed that Helicoverpa armigera have been selected for high levels of resistance to Cry1Ac toxins or transgenic Bt cotton in labs.
In Qiuxian County(Hebei,China),Bt cotton has been commercially planted since 1998. Previous study using F_2 screen in 1999 found that the frequency of resistance alleles to Bt cotton in field populations of Helicoverpa armigera was 0.0058.Considering the high selection pressures in this region,we conducted another F_2 screen,high-dose bioassay, F_1screen and DNA-based method to monitor the Bt resistance in the field populations of H. armigera from 2003 to 2007 as Bt cotton acreage continued to increase over the years.The resistance monitoring program is to be used in a proactive way for resistance management, and the results of resistance monitoring will be used to ensuring the long-term durability of Bt plants
The results from the F_2 screen showed that 20 out of 340 isofemale lines during 2003 to 2005 were identified to carry resistance alleles to Bt cotton in Qiuxian County(2003 to 2005) and Weixian County(2004).The resistance allele frequency in field populations of H. armigera ranged from 0.0119 to 0.0297.
The data analysis showed an overall frequency of 0.0146 and a 95%confidence interval of 0.0084-0.0225 for the 3-year period in Qiuxian County.This value is significantly greater than the value reported from 1999(p=0.042).The F_2 screen for detecting Bt resistance alleles was successfully conducted for 105,42 and 131 lines with an average of 372.4±16.0,328.1±9.5 and 314.0±9.0 F_2 neonates were screen on Bt cotton plants, respectively from 2003 to 2005.The results showed that 4,4 and 7lines were considered to be true positive lines for Bt resistance,respectivelyand thus the resistance allele frequency for the population collected during 2003 to 2005was estimated as 0.0119 with a 95%CI of 0.0039-0.0243,0.0297 with a 95%CI of 0.0099-0.0606 and 0.0154 with a 95%CI of 0.0067-0.0277,respectively.The average detection powers of F_2 screen(1-P_(NO)) for all the lines screened were 96%,97%and 94%,respectively.
In 2004 in Weixian County,F_2 screen for detecting Bt resistance alleles was successfully conducted for 62 lines,and an average of 334.7±8.6 F_2 neonates were screened on Bt cotton plants.The results showed that 4 lines were considered to be true positive lines for Bt resistance and thus the resistance allele frequency for the population collected during 2004 was estimated as 0.0243 with a 95%CI of 0.0091-0.0471.The F_2 screen had＞80%detection power for 96%of the lines screened with an average detection power of 95.1%.
Comparing the lab strain of Bt-resistance(YCR) and Bt-susceptible(YCS) H. armigera on Bt cotton leaf for 5d,the result showed that the resistant and susceptible H. armigera had significant difference in larval surviving rate and growth rate on Bt cotton leaf.The susceptible strain(ss) was unable to survive on Bt cotton.Only 2-6% heterozygous individuals(r_1s) survived,the survivors could not reach stage of mid-2~(nd) instar and body weight of≥0.6mg.The resistant individuals(r_1r_1) had considerably higher survival rate(75%),and a majority of survivors(65%among survivors) reached≥0.6 mg and at least mid-2~(nd) instar.Therefore,the survivors reached≥0.6 mg and beyond mid-2~(nd) instar on Bt cotton leaf for 5 days were categorized as resistant individuals.
In 2006 to 2007,we used Bt cotton leaves as screen media to monitor the resistance allele frequency of field population of H.armigera in Qiuxian County.In 2006 and 2007,F_2 screen for detecting Bt resistance alleles was successfully conducted for 134 and 137lines, and an average of 187.9±3.4 and 105.1±1.5F_2 neonates were screened on Bt cotton plants, respectively.The resistance allele frequency for the population collected during 2006 and 2007 were estimated as 0.1135with a 95%CI of 0.0862-0.1449 and 0.0745 with a 95%CI of 0.0532-0.0995,respectively.The average detection powers of F_2 screen(1-P_(NO)) for all the lines screened were 95%and 90%,respectively.
In 2007,F_1 screen for detecting Bt resistance alleles was successfully conducted for 135 lines,and an average of 165.2±5.2 F_1 neonates were screened on Bt cotton plants.The results showed that 29 lines were considered to be true positive lines for Bt resistance and thus the resistance allele frequency for the population collected during 2007 was estimated as 0.107 with a 95%CI of 0.055-0.159.
Dose-response bioassay is used to monitor Bt resistance in field population of H. armigera in Hebei Province from 2003 to 2007.The results showed that the RF to Cry1Ac toxin were 15.3、12.9、12.5、6.7、16.5 and 11.5.
In 2006 to 2007,the DNA-based method was used to detect the mutation of Cadherin gene in the field resistance H.armigera individuals.The results showed that the primers designed by YCR and YCS could not be used to detect r_1 resistance gene in filed resistance individuals.
Our results suggest that significant shift in resistance allele frequency had occurred in the field populations of H.armigera in Qiuxian and Weixian County.It confirmed that after intinial resistance allele frequencies reaches 0.005;Bt resistance in field population will develop more fast.It is necessary to focuse on the development and refinement of cost-effective methods for Bt resistance decetion and monitoring.It also need to introduce Bt cotton that expressing multi-Bt toxins and integrate the Bt cotton technology with biological,chemical,and cultural practices for management of this key cotton pest.
在中国河北邱县地区,1998年开始推广种植Bt棉,1999年采用单雌系F_2代法检测到邱县地区田间棉铃虫野生种群对Bt棉初始抗性等位基因频率达到了0.0058。考虑到棉铃虫在如此高的抗性初始频率值和持续的田间选择压情况下,有可能对Bt棉产生抗性风险。2003-2007年在前人研究的基础上,我们采用单雌系F_2代法、剂量反应法、F_1代法和DNA分子生物学方法相结合的综合抗性监(检)测体系,研究长期大面积种植转Bt基因棉地区棉铃虫田间种群对Bt棉的抗性发展趋势,为设计制定合理有效的抗性治理策略提供及时、准确的科学依据,以延长转Bt基因棉在大田的使用。
2003-2005年采用Bt棉株作为筛选媒介,对河北邱县(2003-2005年)和威县(2004年)地区田间棉铃虫种群进行单雌系F_2代法抗性监测。结果表明,这两个地区3年监测的340对单雌系中有20对被确定为携带有对Bt棉抗性的等位基因,棉铃虫种群的抗性等位基因频率(E(q))范围为0.0119-0.0297。
其中邱县地区连续3年监测到的棉铃虫对Bt棉的平均抗性等位基因频率值(E(q))为0.0146(95%CI=0.0084-0.0225),明显高于1999年的监测结果(p=0.042);2003-2005年分别监测了105对、42对和131对单雌系,平均每个单雌系在Bt棉株上分别处理筛选了372.4±16.0头、328.1±9.5头和314.0±9.0头F_2代初孵幼虫,最终分别有4对、4对和7对单雌系被确定为田间亲本携带有抗性基因的阳性反应单雌系,每年的抗性等位基因频率值分别为0.0119(95%CI=0.0039-0.0243)、0.0297(95%CI=0.0099-0.0606)和0.0154(95%CI=0.0067-0.0277),监测处理单雌系的总抗性等位基因检出累计概率值(1-P_(NO))分别为96%、97%和94%;
威县地区2004年共筛选了62对单雌系,平均每个单雌系筛选了334.7±8.6头F_2代初孵幼虫,最终有5个单雌系被确定田间亲本携带有抗性基因,抗性等位基因频率值为0.0243(95%CI=0.0091-0.0471),单雌系总的抗性等位基因检出累计概率值(1-P_(NO))为95%。结果表明威县地区棉铃虫野生种群对Bt棉的抗性等位基因频率值处在一个较高的阈值,并且超过了高剂量/庇护区策略的要求。
采用Bt棉叶喂饲法处理室内敏感、抗性及两者杂交后代的初孵幼虫,通过四者在转基因棉花上5天生长发育速率的不同来加以区分。试验结果表明,敏感与抗性品系棉铃虫在Bt棉上5天的存活率与生长发育速率存在明显差异,敏感幼虫不能在转Bt基因棉叶上存活,抗性品系在转Bt基因棉叶上处理5天的存活率达到75%,正反交的幼虫存活率仅为2-6%.棉铃虫抗性品系存活幼虫中65%个体在Bt棉上发育到2龄中期、0.6mg以上;而杂合子和敏感纯合子存活幼虫均达不到此生长发育速率。因此可以确定棉铃虫幼虫在Bt棉上5天发育到2龄中期以上、体重≥0.6mg的个体为抗性个体。
2006-2007年改进单雌系F_2代法,采用Bt棉叶作为筛选媒介进行邱县田间棉铃虫抗性监测。2006年和2007年共筛选了134对和137对单雌系,平均每个单雌系分别筛选了187.9±3.4头和105.1±1.5头F_2代初孵幼虫,抗性等位基因频率值分别为0.1135(95%CI=0.0862-0.1449)和0.0745(95%CI=0.0532-0.0995),2006年和2007年单雌系总的抗性等位基因检出累计概率值(1-P_(No))分别为95%和90%。2007年,采用大田雄虫与室内抗性雌虫杂交的F_1代法监测邱县棉铃虫抗性等位基因频率,共筛选了135对单雌系,平均每个单雌系筛选了165.2±5.2头F_1代初孵幼虫,经过F_2代核查后最终确定有29对单雌系的田间亲本携带有抗性基因,抗性等位基因频率值为0.107(95%CI=0.055-0.159)。
2003-2007年,采用剂量反应法监测邱县、威县地区棉铃虫种群对CrylAc毒蛋白的抗性水平。5年的研究结果显示,田间棉铃虫对CrylAc毒蛋白的抗性倍数分别为15.3、12.9、12.5、6.7、16.5和11.5,基本达到了中等水平抗性。
2006-2007年,将单雌系F_2代法确定为携带有对Bt棉抗性等位基因的田间亲本提取基因组DNA,利用根据室内抗性和敏感棉铃虫钙粘素受体基因序列设计的三个引物对田间抗性棉铃虫进行PASA分子检测,探索性的研究这三个引物在田间抗性检测中的应用。2006年共检测了50头田间亲本成虫,2007年共检测了44头田间亲本成虫。结果显示,根据室内抗、敏棉铃虫钙粘素受体基因设计的PASA引物无法有效的应用于田间棉铃虫Bt抗性检测,田间Bt抗性分子检测技术还有待进一步的改进。
通过长达5年的田间抗性监测研究发现,邱县等大面积种植表达单一CrylAc毒蛋白Bt棉的地区,田间棉铃虫种群对Bt棉的抗性等位基因频率已经显著上升,证实了害虫抗药性发展模式中,在初始抗性等位基因频率值较高(＞0.005)地区的害虫,对Bt棉的抗性发展较快的这一推论。类似于邱县作物种植种类和结构的地区需要采取有效、灵敏的抗性检测与监测体系进行害虫的Bt抗性研究,同时结合田间生态环境,建立合理、有效的抗性管理策略,从而延长转Bt基因作物的应用。
Transgenic Bacillus thuringiensis(Bt) cotton producing Cry1Ac protein has been widely planted in northern China for a long time.The high degree of effectiveness and low environmental risk of Bt crops have encouraged Bt cotton producers to rapidly adopt this technology.The rapid and extensive commercialization of Bt cotton could increase the risk of resistance development in target insects.The studies showed that Helicoverpa armigera have been selected for high levels of resistance to Cry1Ac toxins or transgenic Bt cotton in labs.
In Qiuxian County(Hebei,China),Bt cotton has been commercially planted since 1998. Previous study using F_2 screen in 1999 found that the frequency of resistance alleles to Bt cotton in field populations of Helicoverpa armigera was 0.0058.Considering the high selection pressures in this region,we conducted another F_2 screen,high-dose bioassay, F_1screen and DNA-based method to monitor the Bt resistance in the field populations of H. armigera from 2003 to 2007 as Bt cotton acreage continued to increase over the years.The resistance monitoring program is to be used in a proactive way for resistance management, and the results of resistance monitoring will be used to ensuring the long-term durability of Bt plants
The results from the F_2 screen showed that 20 out of 340 isofemale lines during 2003 to 2005 were identified to carry resistance alleles to Bt cotton in Qiuxian County(2003 to 2005) and Weixian County(2004).The resistance allele frequency in field populations of H. armigera ranged from 0.0119 to 0.0297.
The data analysis showed an overall frequency of 0.0146 and a 95%confidence interval of 0.0084-0.0225 for the 3-year period in Qiuxian County.This value is significantly greater than the value reported from 1999(p=0.042).The F_2 screen for detecting Bt resistance alleles was successfully conducted for 105,42 and 131 lines with an average of 372.4±16.0,328.1±9.5 and 314.0±9.0 F_2 neonates were screen on Bt cotton plants, respectively from 2003 to 2005.The results showed that 4,4 and 7lines were considered to be true positive lines for Bt resistance,respectivelyand thus the resistance allele frequency for the population collected during 2003 to 2005was estimated as 0.0119 with a 95%CI of 0.0039-0.0243,0.0297 with a 95%CI of 0.0099-0.0606 and 0.0154 with a 95%CI of 0.0067-0.0277,respectively.The average detection powers of F_2 screen(1-P_(NO)) for all the lines screened were 96%,97%and 94%,respectively.
In 2004 in Weixian County,F_2 screen for detecting Bt resistance alleles was successfully conducted for 62 lines,and an average of 334.7±8.6 F_2 neonates were screened on Bt cotton plants.The results showed that 4 lines were considered to be true positive lines for Bt resistance and thus the resistance allele frequency for the population collected during 2004 was estimated as 0.0243 with a 95%CI of 0.0091-0.0471.The F_2 screen had＞80%detection power for 96%of the lines screened with an average detection power of 95.1%.
Comparing the lab strain of Bt-resistance(YCR) and Bt-susceptible(YCS) H. armigera on Bt cotton leaf for 5d,the result showed that the resistant and susceptible H. armigera had significant difference in larval surviving rate and growth rate on Bt cotton leaf.The susceptible strain(ss) was unable to survive on Bt cotton.Only 2-6% heterozygous individuals(r_1s) survived,the survivors could not reach stage of mid-2~(nd) instar and body weight of≥0.6mg.The resistant individuals(r_1r_1) had considerably higher survival rate(75%),and a majority of survivors(65%among survivors) reached≥0.6 mg and at least mid-2~(nd) instar.Therefore,the survivors reached≥0.6 mg and beyond mid-2~(nd) instar on Bt cotton leaf for 5 days were categorized as resistant individuals.
In 2006 to 2007,we used Bt cotton leaves as screen media to monitor the resistance allele frequency of field population of H.armigera in Qiuxian County.In 2006 and 2007,F_2 screen for detecting Bt resistance alleles was successfully conducted for 134 and 137lines, and an average of 187.9±3.4 and 105.1±1.5F_2 neonates were screened on Bt cotton plants, respectively.The resistance allele frequency for the population collected during 2006 and 2007 were estimated as 0.1135with a 95%CI of 0.0862-0.1449 and 0.0745 with a 95%CI of 0.0532-0.0995,respectively.The average detection powers of F_2 screen(1-P_(NO)) for all the lines screened were 95%and 90%,respectively.
In 2007,F_1 screen for detecting Bt resistance alleles was successfully conducted for 135 lines,and an average of 165.2±5.2 F_1 neonates were screened on Bt cotton plants.The results showed that 29 lines were considered to be true positive lines for Bt resistance and thus the resistance allele frequency for the population collected during 2007 was estimated as 0.107 with a 95%CI of 0.055-0.159.
Dose-response bioassay is used to monitor Bt resistance in field population of H. armigera in Hebei Province from 2003 to 2007.The results showed that the RF to Cry1Ac toxin were 15.3、12.9、12.5、6.7、16.5 and 11.5.
In 2006 to 2007,the DNA-based method was used to detect the mutation of Cadherin gene in the field resistance H.armigera individuals.The results showed that the primers designed by YCR and YCS could not be used to detect r_1 resistance gene in filed resistance individuals.
Our results suggest that significant shift in resistance allele frequency had occurred in the field populations of H.armigera in Qiuxian and Weixian County.It confirmed that after intinial resistance allele frequencies reaches 0.005;Bt resistance in field population will develop more fast.It is necessary to focuse on the development and refinement of cost-effective methods for Bt resistance decetion and monitoring.It also need to introduce Bt cotton that expressing multi-Bt toxins and integrate the Bt cotton technology with biological,chemical,and cultural practices for management of this key cotton pest.
引文
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