大豆蚜对吡虫啉的抗性监测及抗性机理研究
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摘要
大豆蚜Soybean aphid(Aphis glycines Matsumura)是大豆的重要害虫之一,吡虫啉自上世纪90年代推行以来,对大豆蚜良好的防治效果使其成为针对这一害虫效果最好、使用最广的当家杀虫剂,为了了解田间大豆蚜对吡虫啉抗性的发生动态,延长吡虫啉的田间使用寿命,本文通过田间抗性调查、室内抗性筛选及种群适合度研究、交互抗性研究及抗性现实遗传力研究,系统分析了大豆蚜对吡虫啉产生抗性的风险；利用室内筛选的抗性品系和敏感品系试虫为材料,分析了抗性产生的生化机制；利用分子生物学技术克隆了大豆蚜烟碱型乙酰胆碱受体基因和羧酸酯酶基因,并对羧酸酯酶基因进行了原核表达及活性分析；同时还利用荧光定量PCR技术,对大豆蚜羧酸酯酶基因mRNA水平的表达量进行了分析,取得了多项具有理论和实际意义的重要成果。
一、大豆蚜对常规杀虫剂的敏感基线构建和田间吡虫啉抗性监测
通过杀虫剂毒力测定方法建立了大豆蚜对7种常规杀虫剂的敏感基线,并以此为基准分别对2010年和2011年两年间全国大豆主要分布的5省区田间大豆蚜对吡虫啉的抗药性动态和哈尔滨地区田间大豆蚜对7种常规药剂的抗药性状况分别进行了监测,结果表明,5省(区)中除长春种群为低水平抗性外,其他4个省(区)地理种群大豆蚜抗性为中等水平抗性,7种药剂中,高效氯氟氰菊酯、吡虫啉和溴氰菊酯分别产生高、中、低等水平的抗性,其他四种药剂未产生抗药性。
二、抗吡虫啉大豆蚜品系的选育及抗性风险评估
采用浸渍法对室内连续饲养多年的的大豆蚜种群进行19代的抗性筛选,获得了抗性倍数最高为24.46倍的室内抗性品系。从抗性筛选的速度来看,大豆蚜对吡虫啉的抗性发展速度并不快,与棉蚜、桃蚜的吡虫啉抗性筛选相类似,从抗性筛选的结果来看,连续筛选的前6代抗性增长较缓慢,筛选的7到16代为快速增长期,16代后抗性基本稳定在23倍左右,进入了一个增长的平台期,总体呈现“S”型增长趋势。对室内筛选19代的大豆蚜对吡虫啉的抗性现实遗传力(h2)进行了测定,并推测出田间大豆蚜对吡虫啉的抗性提高10倍只需6-8代。这表明大豆蚜对吡虫啉的抗性增高后,持续的抗性筛选仍能使抗药性进一步上升。
三、吡虫啉抗性对大豆蚜种群适合度的影响
通过构建种群生命表,观察比较了筛选过程中吡虫啉抗性品系T9、T18和敏感品系的一系列生长发育和繁殖特征,就抗性对种群适合度的影响进行了研究。用内禀增长力来确定种群的相对适合度,结果表明,抗性品系T9和T18均表现出相对适合度下降,而且T18的相对适合度的下降幅度明显大于T9,生长发育和繁殖上也都表现出一系列的不利性。
四、大豆蚜抗吡虫啉品系的交互抗性研究
本研究以大豆蚜敏感品系为对照,测定了吡虫啉抗性品系T14和T18对6种常规杀虫剂的敏感性变化,以了解大豆蚜抗吡虫啉品系对其他药剂的交互抗性。结果表明,随着大豆蚜抗性的不断发展,抗性品系T14和T18对生产上常用的烟碱类杀虫剂和菊酯类杀虫剂均已经产生不同程度的交互抗性,但未对有机磷类杀虫剂产生交互抗性。
五、大豆蚜对吡虫啉抗性的生化机制分析
通过室内筛选获得大豆蚜的吡虫啉抗性品系,利用增效试验和解毒酶活力测定,对其抗性机理进行了研究。解毒酶活力测定表明：在抗感品系中,抗性品系的羧酸酯酶比活力明显高于敏感品系(2.1倍),而谷胱甘肽S-转移酶与抗性形成无关；增效试验结果表明：DEM在抗感品系中对吡虫啉都没有明显的增效作用,而TPP和PBO在两个品系中均有增效作用,但前者的作用更明显。综合分析认为羧酸酯酶活力增强在大豆蚜对吡虫啉的抗性形成中起重要作用。
六、大豆蚜烟碱型乙酰胆碱受体基因的克隆
昆虫烟碱型乙酰胆碱受体(nAChR)是以吡虫啉为代表的新烟碱类杀虫剂的作用靶标位点。本研究利用RT-PCR和RACE技术,从大豆蚜中成功克隆了6个烟碱型乙酰胆碱受体基因全序列,其中5个为α亚基基因,1个为β亚基基因,分别命名为：Agla1、Agla2、Agla3、 Agla4、Agla5和Aglβ1,其中Agla1亚基全长1456bp,编码427个氨基酸；Agla2亚基全长2463bp,编码592个氨基酸；Agla3亚基全长2158bp,编码537个氨基酸；Agla4亚基全长1792bp,编码532个氨基酸；Agla5亚基全长1850bp,编码549个氨基酸；Aglβ1亚基全长2406bp,编码509个氨基酸。其GenBank登录号分别为JQ690271、JF775487、JQ690273、 JN681173、JQ690274和JN681174。同时,还获得了Aglal的一个选择性剪接Aglal-2,全长为1396bp,编码406个氨基酸,GenBank登录号为JQ690272。同源性分析表明,以上亚基均具有昆虫nAChR亚基的典型特征,与其他昆虫nAChR亚基的同源性很高,说明克隆的基因确实是大豆蚜的nAChR基因。对大豆蚜nAChR亚基进行多态性分析,未发现氨基酸突变。该研究结果为今后进一步研究大豆蚜nAChR的天然亚基组成及功能,分析大豆蚜对新烟碱类杀虫剂的靶标抗性产生奠定了基础。
七、大豆蚜羧酸酯酶基因CarE的克隆与原核表达
利用RT-PCR和RACE技术,成功克隆了大豆蚜中的羧酸酯酶基因,命名为AgCarE,全长为1946bp,编码526个氨基酸,GenBank登录号为JF970181。通过序列比对和功能性分析,AgCarE具备羧酸酯酶的结构特征,如蛋白活性中心的催化三联体：Ser186, Glu313,His434,证明其属于羧酸酯酶家族。通过构建重组表达载体pET21b-AgCarE进行原核表达及Westem-blot鉴定,59.2kD的目的蛋白能够成功表达。以α-醋酸萘酯为底物的体外酶活性检测表明,其能够水解α-醋酸萘酯,说明表达产物具备羧酸酯酶活性。
八、抗感大豆蚜品系羧酸酯酶mRNA表达量研究
利用实时荧光定量PCR技术,以18SrDNA为内参基因,通过比较CT值法(即2-△△Ct法)获得了抗、感大豆蚜品系单头蚜虫羧酸酯酶mRNA的相对表达量。分析结果得出,抗性品系大豆蚜羧酸酯酶mRNA表达量平均比敏感品系大豆蚜羧酸酯酶mRNA表达量高出1.25倍,结合生化机理分析和原核表达活性分析认为,大豆蚜对吡虫啉的抗药性增加,与其体内羧酸酯酶基因mRNA水平的过量表达密切相关。
Soybean aphid(Aphis glycines Matsumura) is one of the most important pests on soybean. Imidacloprid is the key insecticide which has been widely used for control of this pest since1990s. So study on imidacloprid resistance is very important for sustainable control of A. glycines. In this paper, the resistance survey, resistance selection, insect fitness analysis, cross-resistance and realized heritability were carried out to evaluate the risk for A. glycines to develop high resistance to imidacloprid. With selected susceptible strain and resistant strains, the biochemical and molecular mechanisms for the resistance were also analyzed. Some valuable breakthroughs have been achieved.
1. Establishment of the relative susceptible baseline data to common insecticides and resistance monitoring in field populations of A. glycines
The susceptibility of A. glycines to7common insecticides was determined with laboratory susceptible strain and the baseline data were obtained. Based on these data, five field populations (Jinan, Cangzhou, Chifeng, Changchun, Harbin) collected from main soybean producing provinces of China in2010and2011were monitored for resistance to imidacloprid, and the field population of Harbin collected in2010and2011were monitored for resistance to7common insecticides. The results revealed that the aphid of Changchun had developed low level of resistance to imidacloprid, and the other four areas had developed medium level of resistance to imidacloprid. Except imidacloprid, Lambda-cyhalothrin and Deltamethrin, A. glycines had no developed resistance to other insecticides.
2. Imidacloprid resistance selected and resistance risk assessment in A. glycines
An imidacloprid-resistant strain of A. glycines was developed by continuous selection for19generations with imidacloprid in laboratory. During the selection, imidacloprid resistance was found with a phased increase. From the1st to6th generation, the resistance developed very slowly, the rapid increase occurred from7th to16th generation, and leaving17th to19th as resistance developing plate. The estimate of realized heritability of19th generation showed that resistant ratio increased10-fold was only needed6-8generations when the continuous selection was used for A. glycines with imidacloprid. It was likely that the resistance level of A. glycines to imidacloprid would increase further if the selection had continued.
3. The influence of imidacloprid resistance on the relative fitness of A. glycines
The relative fitness of resistant strain (T9and T18) had been tested by constructing life tables and taking susceptible strain as control. The results showed that two resistant strains had obvious disadvantages in their development and reproduction. The relative fitness of resistant strain was determining by intrinsic rate of increase (rm) with susceptible strain as standard. The result showed that the relative fitness of resistant strain was decreased dramatically.
4. Cross-resistance of the imidacloprid-resistant strain of A. glycines
The cross-resistance of the imidacloprid-resistant strains (T14and T18) of A. glycines to6common insecticides was measured. The results showed that A. glycines with resistance to imidacloprid had no obvious cross-resistance to organophosphorus insecticides, and significant cross-resistance existed to the neonicotinoid insecticides and pyrethroid insecticides.
5. Biochemical mechanisms for imidacloprid resistance in A. glycines
The biochemical mechanisms of imidacloprid resistance in A. glycines were studied by synergism test and detoxification enzyme analysis. The results showed that triphenyl phosphate and piperonyl butoxide had significant synergism on imidacloprid in both resistant and susceputible strains, and the synergism ratio of triphenyl phosphate was much higher than the synergism ratio of piperonyl butoxide. Diethyl mateate did not show any synergism on imidacloprid in both strains. The results of detoxification enzyme analysis in two strains showed that the activity of carboxylesterase in resistant strain was much higher than the susceptible strain (2.1-fold), however the activity of glutathione S-transferase in resistant strain had no significantly enhanced. Thus, it was concluded that the enhancement of the activity of carboxylesterase should play an important role in the imidacloprid resistance of A. glycines.
6. Cloning of nAChR genes from A. glycines
Nicotinic acetylcholine receptors (nAChRs) are the target of imidacloprid which belong to neonicotinoid insecticides. With RT-PCR and RACE technique,6different subunits of nAChR had been successfully cloned from A. glycines, include5a subunits and1β subunit. They were named as Aglal, Agla2, Agla3, Agla4, Agla5and Aglβl, and their GenBank accession number were JQ690271, JF775487, JQ690273, JN681173, JQ690274and JN681174, respectively. The Aglal was1456bp in length, and the open reading frame encoded427amino acids; the Agla2was2463bp in length, and the open reading frame encoded592amino acids; the Agla3was2158bp in length, and the open reading frame encoded537amino acids; the Agla4was1792bp in length, and the open reading frame encoded532amino acids; the Agla5was1850bp in length, and the open reading frame encoded549amino acids; the Aglβl was2406bp in length, and the open reading frame encoded509amino acids. One alternative splice was found with the clone genes. Homology analysis demonstrated that these6subunits of nAChR had high amino acid sequence identity with other insect nAChR subunits previously reported. Polymorphism analysis demonstrated that there is no point mutation exited in resistant and susceptible strains. This work facilitated the further studies on natural subunit composition, function of nAChR and the molecular mechanisms of target resistance for neonicotinoid insecticides.
7. Cloning and expression of carboxylesterase gene from A. glycines
The carboxylesterase gene was cloned by using RT-PCR and RACE methods, and it was named as AgCarE. The AgCarE was1946bp in length, and the open reading frame encoded526amino acids, the GenBank accession number was JF970181. The AgCarE possesses a catalytic triad, consisting of a Ser, a His and a Glu residue (Ser186, Glu313and His434), which is characterized by carboxylic esters (EC:3.1.1.-). The AgCarE gene had been recombined into pET21b vector. Then the HIS fusion protein was expressed by the induction of IPTG. The result of SDS-PAGE and Western blot analysis showed that the AgCarE protein in A. glycines was expressed in E. coli BL21which induced by IPTG, and its MW was found to be about59kD, similar as the predicted result. The carboxylesterase activity of AgCarE was characterized with the a-NA as a substrate and its enzyme activity of AgCarE was analyzed.
8. Relative expression of carboxylesterase mRNA in resistant and susceptible strains of A. glycines
The relative expression of carboxylesterase mRNA in resistant and susceptible strains of A. glycines were estimated by Ct comprison method. It was found that the relative expression of carboxylesterase gene in resistant strain had much higher than the expression in the susceptible strain (2.25-fold). Combination of the previous results in biochemical mechanism analysis and enzyme activity analysis of AgCarE revealed that the over-expression of carboxylesterase might associate with the high resistance to imidacloprid in A. glycines.
一、大豆蚜对常规杀虫剂的敏感基线构建和田间吡虫啉抗性监测
通过杀虫剂毒力测定方法建立了大豆蚜对7种常规杀虫剂的敏感基线,并以此为基准分别对2010年和2011年两年间全国大豆主要分布的5省区田间大豆蚜对吡虫啉的抗药性动态和哈尔滨地区田间大豆蚜对7种常规药剂的抗药性状况分别进行了监测,结果表明,5省(区)中除长春种群为低水平抗性外,其他4个省(区)地理种群大豆蚜抗性为中等水平抗性,7种药剂中,高效氯氟氰菊酯、吡虫啉和溴氰菊酯分别产生高、中、低等水平的抗性,其他四种药剂未产生抗药性。
二、抗吡虫啉大豆蚜品系的选育及抗性风险评估
采用浸渍法对室内连续饲养多年的的大豆蚜种群进行19代的抗性筛选,获得了抗性倍数最高为24.46倍的室内抗性品系。从抗性筛选的速度来看,大豆蚜对吡虫啉的抗性发展速度并不快,与棉蚜、桃蚜的吡虫啉抗性筛选相类似,从抗性筛选的结果来看,连续筛选的前6代抗性增长较缓慢,筛选的7到16代为快速增长期,16代后抗性基本稳定在23倍左右,进入了一个增长的平台期,总体呈现“S”型增长趋势。对室内筛选19代的大豆蚜对吡虫啉的抗性现实遗传力(h2)进行了测定,并推测出田间大豆蚜对吡虫啉的抗性提高10倍只需6-8代。这表明大豆蚜对吡虫啉的抗性增高后,持续的抗性筛选仍能使抗药性进一步上升。
三、吡虫啉抗性对大豆蚜种群适合度的影响
通过构建种群生命表,观察比较了筛选过程中吡虫啉抗性品系T9、T18和敏感品系的一系列生长发育和繁殖特征,就抗性对种群适合度的影响进行了研究。用内禀增长力来确定种群的相对适合度,结果表明,抗性品系T9和T18均表现出相对适合度下降,而且T18的相对适合度的下降幅度明显大于T9,生长发育和繁殖上也都表现出一系列的不利性。
四、大豆蚜抗吡虫啉品系的交互抗性研究
本研究以大豆蚜敏感品系为对照,测定了吡虫啉抗性品系T14和T18对6种常规杀虫剂的敏感性变化,以了解大豆蚜抗吡虫啉品系对其他药剂的交互抗性。结果表明,随着大豆蚜抗性的不断发展,抗性品系T14和T18对生产上常用的烟碱类杀虫剂和菊酯类杀虫剂均已经产生不同程度的交互抗性,但未对有机磷类杀虫剂产生交互抗性。
五、大豆蚜对吡虫啉抗性的生化机制分析
通过室内筛选获得大豆蚜的吡虫啉抗性品系,利用增效试验和解毒酶活力测定,对其抗性机理进行了研究。解毒酶活力测定表明：在抗感品系中,抗性品系的羧酸酯酶比活力明显高于敏感品系(2.1倍),而谷胱甘肽S-转移酶与抗性形成无关；增效试验结果表明：DEM在抗感品系中对吡虫啉都没有明显的增效作用,而TPP和PBO在两个品系中均有增效作用,但前者的作用更明显。综合分析认为羧酸酯酶活力增强在大豆蚜对吡虫啉的抗性形成中起重要作用。
六、大豆蚜烟碱型乙酰胆碱受体基因的克隆
昆虫烟碱型乙酰胆碱受体(nAChR)是以吡虫啉为代表的新烟碱类杀虫剂的作用靶标位点。本研究利用RT-PCR和RACE技术,从大豆蚜中成功克隆了6个烟碱型乙酰胆碱受体基因全序列,其中5个为α亚基基因,1个为β亚基基因,分别命名为：Agla1、Agla2、Agla3、 Agla4、Agla5和Aglβ1,其中Agla1亚基全长1456bp,编码427个氨基酸；Agla2亚基全长2463bp,编码592个氨基酸；Agla3亚基全长2158bp,编码537个氨基酸；Agla4亚基全长1792bp,编码532个氨基酸；Agla5亚基全长1850bp,编码549个氨基酸；Aglβ1亚基全长2406bp,编码509个氨基酸。其GenBank登录号分别为JQ690271、JF775487、JQ690273、 JN681173、JQ690274和JN681174。同时,还获得了Aglal的一个选择性剪接Aglal-2,全长为1396bp,编码406个氨基酸,GenBank登录号为JQ690272。同源性分析表明,以上亚基均具有昆虫nAChR亚基的典型特征,与其他昆虫nAChR亚基的同源性很高,说明克隆的基因确实是大豆蚜的nAChR基因。对大豆蚜nAChR亚基进行多态性分析,未发现氨基酸突变。该研究结果为今后进一步研究大豆蚜nAChR的天然亚基组成及功能,分析大豆蚜对新烟碱类杀虫剂的靶标抗性产生奠定了基础。
七、大豆蚜羧酸酯酶基因CarE的克隆与原核表达
利用RT-PCR和RACE技术,成功克隆了大豆蚜中的羧酸酯酶基因,命名为AgCarE,全长为1946bp,编码526个氨基酸,GenBank登录号为JF970181。通过序列比对和功能性分析,AgCarE具备羧酸酯酶的结构特征,如蛋白活性中心的催化三联体：Ser186, Glu313,His434,证明其属于羧酸酯酶家族。通过构建重组表达载体pET21b-AgCarE进行原核表达及Westem-blot鉴定,59.2kD的目的蛋白能够成功表达。以α-醋酸萘酯为底物的体外酶活性检测表明,其能够水解α-醋酸萘酯,说明表达产物具备羧酸酯酶活性。
八、抗感大豆蚜品系羧酸酯酶mRNA表达量研究
利用实时荧光定量PCR技术,以18SrDNA为内参基因,通过比较CT值法(即2-△△Ct法)获得了抗、感大豆蚜品系单头蚜虫羧酸酯酶mRNA的相对表达量。分析结果得出,抗性品系大豆蚜羧酸酯酶mRNA表达量平均比敏感品系大豆蚜羧酸酯酶mRNA表达量高出1.25倍,结合生化机理分析和原核表达活性分析认为,大豆蚜对吡虫啉的抗药性增加,与其体内羧酸酯酶基因mRNA水平的过量表达密切相关。
Soybean aphid(Aphis glycines Matsumura) is one of the most important pests on soybean. Imidacloprid is the key insecticide which has been widely used for control of this pest since1990s. So study on imidacloprid resistance is very important for sustainable control of A. glycines. In this paper, the resistance survey, resistance selection, insect fitness analysis, cross-resistance and realized heritability were carried out to evaluate the risk for A. glycines to develop high resistance to imidacloprid. With selected susceptible strain and resistant strains, the biochemical and molecular mechanisms for the resistance were also analyzed. Some valuable breakthroughs have been achieved.
1. Establishment of the relative susceptible baseline data to common insecticides and resistance monitoring in field populations of A. glycines
The susceptibility of A. glycines to7common insecticides was determined with laboratory susceptible strain and the baseline data were obtained. Based on these data, five field populations (Jinan, Cangzhou, Chifeng, Changchun, Harbin) collected from main soybean producing provinces of China in2010and2011were monitored for resistance to imidacloprid, and the field population of Harbin collected in2010and2011were monitored for resistance to7common insecticides. The results revealed that the aphid of Changchun had developed low level of resistance to imidacloprid, and the other four areas had developed medium level of resistance to imidacloprid. Except imidacloprid, Lambda-cyhalothrin and Deltamethrin, A. glycines had no developed resistance to other insecticides.
2. Imidacloprid resistance selected and resistance risk assessment in A. glycines
An imidacloprid-resistant strain of A. glycines was developed by continuous selection for19generations with imidacloprid in laboratory. During the selection, imidacloprid resistance was found with a phased increase. From the1st to6th generation, the resistance developed very slowly, the rapid increase occurred from7th to16th generation, and leaving17th to19th as resistance developing plate. The estimate of realized heritability of19th generation showed that resistant ratio increased10-fold was only needed6-8generations when the continuous selection was used for A. glycines with imidacloprid. It was likely that the resistance level of A. glycines to imidacloprid would increase further if the selection had continued.
3. The influence of imidacloprid resistance on the relative fitness of A. glycines
The relative fitness of resistant strain (T9and T18) had been tested by constructing life tables and taking susceptible strain as control. The results showed that two resistant strains had obvious disadvantages in their development and reproduction. The relative fitness of resistant strain was determining by intrinsic rate of increase (rm) with susceptible strain as standard. The result showed that the relative fitness of resistant strain was decreased dramatically.
4. Cross-resistance of the imidacloprid-resistant strain of A. glycines
The cross-resistance of the imidacloprid-resistant strains (T14and T18) of A. glycines to6common insecticides was measured. The results showed that A. glycines with resistance to imidacloprid had no obvious cross-resistance to organophosphorus insecticides, and significant cross-resistance existed to the neonicotinoid insecticides and pyrethroid insecticides.
5. Biochemical mechanisms for imidacloprid resistance in A. glycines
The biochemical mechanisms of imidacloprid resistance in A. glycines were studied by synergism test and detoxification enzyme analysis. The results showed that triphenyl phosphate and piperonyl butoxide had significant synergism on imidacloprid in both resistant and susceputible strains, and the synergism ratio of triphenyl phosphate was much higher than the synergism ratio of piperonyl butoxide. Diethyl mateate did not show any synergism on imidacloprid in both strains. The results of detoxification enzyme analysis in two strains showed that the activity of carboxylesterase in resistant strain was much higher than the susceptible strain (2.1-fold), however the activity of glutathione S-transferase in resistant strain had no significantly enhanced. Thus, it was concluded that the enhancement of the activity of carboxylesterase should play an important role in the imidacloprid resistance of A. glycines.
6. Cloning of nAChR genes from A. glycines
Nicotinic acetylcholine receptors (nAChRs) are the target of imidacloprid which belong to neonicotinoid insecticides. With RT-PCR and RACE technique,6different subunits of nAChR had been successfully cloned from A. glycines, include5a subunits and1β subunit. They were named as Aglal, Agla2, Agla3, Agla4, Agla5and Aglβl, and their GenBank accession number were JQ690271, JF775487, JQ690273, JN681173, JQ690274and JN681174, respectively. The Aglal was1456bp in length, and the open reading frame encoded427amino acids; the Agla2was2463bp in length, and the open reading frame encoded592amino acids; the Agla3was2158bp in length, and the open reading frame encoded537amino acids; the Agla4was1792bp in length, and the open reading frame encoded532amino acids; the Agla5was1850bp in length, and the open reading frame encoded549amino acids; the Aglβl was2406bp in length, and the open reading frame encoded509amino acids. One alternative splice was found with the clone genes. Homology analysis demonstrated that these6subunits of nAChR had high amino acid sequence identity with other insect nAChR subunits previously reported. Polymorphism analysis demonstrated that there is no point mutation exited in resistant and susceptible strains. This work facilitated the further studies on natural subunit composition, function of nAChR and the molecular mechanisms of target resistance for neonicotinoid insecticides.
7. Cloning and expression of carboxylesterase gene from A. glycines
The carboxylesterase gene was cloned by using RT-PCR and RACE methods, and it was named as AgCarE. The AgCarE was1946bp in length, and the open reading frame encoded526amino acids, the GenBank accession number was JF970181. The AgCarE possesses a catalytic triad, consisting of a Ser, a His and a Glu residue (Ser186, Glu313and His434), which is characterized by carboxylic esters (EC:3.1.1.-). The AgCarE gene had been recombined into pET21b vector. Then the HIS fusion protein was expressed by the induction of IPTG. The result of SDS-PAGE and Western blot analysis showed that the AgCarE protein in A. glycines was expressed in E. coli BL21which induced by IPTG, and its MW was found to be about59kD, similar as the predicted result. The carboxylesterase activity of AgCarE was characterized with the a-NA as a substrate and its enzyme activity of AgCarE was analyzed.
8. Relative expression of carboxylesterase mRNA in resistant and susceptible strains of A. glycines
The relative expression of carboxylesterase mRNA in resistant and susceptible strains of A. glycines were estimated by Ct comprison method. It was found that the relative expression of carboxylesterase gene in resistant strain had much higher than the expression in the susceptible strain (2.25-fold). Combination of the previous results in biochemical mechanism analysis and enzyme activity analysis of AgCarE revealed that the over-expression of carboxylesterase might associate with the high resistance to imidacloprid in A. glycines.
引文
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