东亚飞蝗对有机磷杀虫剂的抗性及其机理研究
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摘要
东亚飞蝗Locusta migratoria manilensis(Meyen)是为害严重的农业害虫,有机磷杀虫剂是防治这一害虫使用较广的杀虫剂,为了深入探讨东亚飞蝗对有机磷杀虫剂的抗性机制,针对东亚飞蝗不同种群制定相应的治理策略,更有效地进行抗性治理,本文通过东亚飞蝗田间种群对有机磷农药的敏感性分析,探讨了东亚飞蝗对有机磷农药产生抗性的现状;同时利用现代分子生物学技术,研究分析了抗性产生的生化和分子生物学机理,主要研究内容如下:
一、东亚飞蝗对有机磷农药的敏感性分析和增效作用测定
本研究针对我国东亚飞蝗发生规模、分布特点、农药使用背景差异和种群遗传特征,采集了河北黄骅、海南黄流和山东无棣田间种群,选择有代表性的有机磷农药进行敏感性分析。结果发现,东亚飞蝗无棣种群对马拉硫磷最为敏感,黄骅种群对毒死蜱和辛硫磷仅有5.4和2.9倍的抗药性,而黄骅和黄流两种群对马拉硫磷已产生了57.5和14.8倍的抗药性。可以看出,由于东亚飞蝗不同种群马拉硫磷选择压力的差异已经造成不同田间种群对其抗性程度产生了变化,而且不同的有机磷农药对同一种群的选择压力已产生不同的抗药性水平。由此说明,研究东亚飞蝗对常用杀虫剂抗性发生现状,建立有效的抗性治理策略,是十分必要的。
通过增效剂磷酸三苯酯(TPP)、顺丁烯二酸二乙酯(DEM)及氧化胡椒基丁醚(PBO)在河北黄骅田间种群和敏感品系中对马拉硫磷的增效试验,结果发现,无论在黄骅种群还是敏感品系中,PBO对马拉硫磷都没有增效作用(增效比分别为0.9和1.1),但在黄骅种群中TPP和DEM对马拉硫磷表现出显著的增效作用(增效比分别为16.2和3.3),而在敏感品系中TPP和DEM没有明显的增效作用(增效比分别为1.4和1.2);由此推测,酯酶和谷胱甘肽S-转移酶与黄骅种群马拉硫磷抗性相关,而细胞色素P450与马拉硫磷抗性无关。
二、东亚飞蝗有机磷抗性代谢解毒酶生化机制研究
本研究分析了东亚飞蝗代谢解毒酶系(酯酶、谷胱甘肽S-转移酶和细胞色素P450氧化酶)的生化特性。东亚飞蝗河北黄骅种群和敏感品系代谢解毒酶活性表现出不同程度的差异,其中黄骅种群酯酶对α-NA、α-NB及β-NA三种底物的水解活性和谷胱甘肽S-转移酶对底物CDNB和DCNB水解活性显著高于敏感品系,分别是敏感品系的2.1-3.2倍和1.2-2.0倍。而对于细胞色素P450活性,两者之间没有显著差异。酯酶聚丙烯酰胺凝胶电泳结果显示,黄骅种群和敏感品系电泳图谱中条带带型没有差异,但黄骅种群条带染色明显深于敏感品系。据此推测,黄骅种群与敏感品系之间的酯酶活性和谷胱甘肽S-转移酶差异与河北黄骅东亚飞蝗的抗药性有关。
用对氧磷、马拉氧磷两种有机磷抑制剂对东亚飞蝗海南黄流和山东无棣种群的羧酸酯酶进行体外抑制,实验结果表明,黄流和无棣种群的pI_(50)值有显著差异,黄流种群分别是山东种群的1.16和1.27倍,说明黄流种群对对氧磷和马拉氧磷的敏感性较低。然而黄流种群和无棣种群的羧酸酯酶活性测定结果表明,以α-NA和β-NA为底物时,无棣种群羧酸酯酶活性分别是黄流种群1.75和1.50倍,具有显著差异。动力学研究表明,无棣种群和黄流种群的羧酸酯酶K_和V_(max)值没有显著差异。据分析黄流种群和无棣种群羧酸酯酶活性的差异可能是由于生态环境的不同所导致的种群分化引起的。根据目前的研究结果尚难以断定对马拉硫磷生物测定中LD_(50)值较低的无棣种群对其它农药的敏感性如何。基于无棣种群较高的羧酸酯酶活性,我们推测,东亚飞蝗无棣种群可能对其它的农药具有较高的抗药性。
三、东亚飞蝗有机磷抗性靶标酶生化机制研究
本研究分析了东亚飞蝗有机磷靶标乙酰胆碱酯酶(acetylcholinesterase,AChE)的生化特性。对东亚飞蝗黄骅种群、黄流种群、无棣种群和敏感品系AChE活性和动力学比较结果表明,三个东亚飞蝗田间种群AChE活性与敏感品系相比有显著差异,黄骅种群、黄流种群和无棣种群AChE活性分别是敏感品系的4.04、9.55和1.99倍,并且三个田间种群的K_m值与敏感品系相比也有显著差异;根据有机磷杀虫剂对AChE的体外抑制结果,河北黄骅种群AChE对马拉氧磷、氧化毒死蜱及辛硫磷的敏感性与敏感品系相比分别下降了3.2、2.2和1.1倍。而海南黄流种群对马拉氧磷和氧化毒死蜱的敏感性分别下降了11.3和19.2倍。同时发现,对马拉硫磷生物测定中仅有1.8倍抗药性的山东无棣种群,其AChE对马拉氧磷和氧化毒死蜱的敏感性也分别下降了4.8和2.4倍。
结合东亚飞蝗解毒酶的生化机制研究,本文认为黄骅种群对马拉硫磷的抗性机理主要包括酯酶、谷胱甘肽S-转移酶活性升高和乙酰胆碱酯酶的活性升高以及敏感性下降。黄流种群产生抗性主要是由于AChE的活性升高和敏感性降低引起的。而无棣种群可能对其它的农药有较高的抗药性,导致其AChE敏感性的显著下降。据此认为,东亚飞蝗不同种群抗性产生机制是不尽相同的,有可能包括多种不同的抗性机制。
四、东亚飞蝗有机磷抗性分子机制研究
本研究利用简并性引物和降落PCR技术从东亚飞蝗中克隆获得了3个东亚飞蝗羧酸酯酶(carboxylesterases,CarEs)基因的cDNA片段,分别命名为CarE1,CarE2和CarE3。根据昆虫羧酸酯酶和谷胱甘肽S-转移酶的保守区(motif),采用生物信息学方法对东亚飞蝗表达序列标签(expressing sequence tags,ESTs)数据库进行了全面搜索,并对其进行了羧酸酯酶和谷胱甘肽S-转移酶ESTs拼接、注释,获得了32条羧酸酯酶ESTs和12条谷胱甘肽S-转移酶ESTs,与克隆获得的3条羧酸酯酶基因片段同源比对和拼接后,共得到34条羧酸酯酶基因片段。采用实时定量PCR技术对上述羧酸酯酶和谷胱甘肽S-转移酶基因在黄骅种群和敏感品系中的表达量进行了比较,结果发现,羧酸酯酶基因CarE1、CarE4、CarE6、CarE12、CarE15、CarE19、CarE22、CarE23、CarE26、CarE27、CarE30和CarE32在黄骅种群中的表达量分别是敏感品系的3.29、1.13、8.78、3.16、3.17、11.84、2.24、4.92、6.24、4.66、4.47和4.73倍;谷胱甘肽S-转移酶基因GST1、GST2、GST3和GST12的表达量分别是敏感品系的1.25、1.33、2.47和1.61倍。据此认为这些羧酸酯酶基因和谷胱甘肽S-转移酶基因表达提高与东亚飞蝗的抗药性密切相关。
对东亚飞蝗AChE基因克隆和Western blot分析结果认为直翅目昆虫AChE基因可能有其特殊性,尚有待进一步研究证实。
本文通过东亚飞蝗不同种群对有机磷农药的敏感性分析,揭示了东亚飞蝗的田间抗性现状。通过生化和分子生物学方法分析了东亚飞蝗对有机磷的抗性机理,发现东亚飞蝗不同种群在产生抗性过程中的机制是不尽相同的,有可能包括多种不同的抗性机制。这些研究结果对于有效地进行抗性治理,延缓东亚飞蝗的抗性发展均具有重要的实践意义。
本文首次借助现有的东亚飞蝗的ESTs数据库对抗性相关解毒酶基因进行了系统研究,用real-time PCR对代谢解毒酶羧酸酯酶基因和谷胱甘肽S-转移酶基因在抗性和敏感种群中差异表达进行了定量分析,确定了抗性形成过程中的关键代谢解毒酶基因,这将极大地推动东亚飞蝗抗性机制的系统深入研究。
综上所述,本文研究结果对东亚飞蝗抗性机制和东亚飞蝗的抗性治理均具有重要的理论和实践意义。
Locusta migratoria manilensis(Meyen) is one of the seriously endangering agricultural pests.Organophosphate(OP)insecticides have been used extensively to control the locust.In order to delaying organophosphorus resistance and carrying out effective management strategies of different populations of L.migratoria manilensis,the objectives of this study were to evaluate the resistance spectrum of the field locust populations using the bioassay to OPs and investigate major biochemical and molecular mechanisms of resistance to OPs by the modem molecular biology technology.Elucidation of the resistance spectrum and resistance mechanisms in L.migratoria manilensis will be significantly important in theory and practice.
1.Resistance spectrum to selected OP insecticides and synergism of TPP, PBO and DEM
Based on the scale,distribution,history of insecticides control and population characteristics of L.migratoria manilensis,the field populations of the locust were collected from Huanghua County of Hebei Province, Huangliu County of Hainan Province and Wudi County of Shandong Province.The sensitivity analysis or resistance spectrums to the representative OP insecticides were evaluated.The results showed that, compared with the susceptible strain(SS),the population of Wudi population (WD) was the most sensitive to malathion and Huanghua population(HH) showed 2.9-fold to phoxim,5.4-fold to chlorpyrifos of resistance levels. However,HH and Huangliu population(HL) developed 57.5-fold and 14.8-fold resistant to malathion.In conclusion,As a consequence of the various malathion selective pressures,there has been developed different resistance to malathion in different locusts populations and various resistant levels in the same locust population has been developed by the different OPs. Consequently,it is significantly necessary to evaluate the resistance spectrum and detect the effective resistance management strategy of the field locust populations.
The malathion resistance of the HH was significantly diminished by TPP (synergism ratio:16.2) and DEM(3.3),but was unchanged by PBO(0.9).In contrast,none of these synergists significantly affected the toxicity of malathion in the SS.These results suggested that both ESTs and GSTs were involved in OP resistance in the HH,whereas P450s were unlikely to be involved in the resistance simply because PBO did not synergize the toxicity of malathion by the synergism of triphenyl phosphate(TPP),diethyl meleate (DEM) and piperonyl butoxide(PBO) on malathion in the HH and SS of the locust.
2.Biochemical mechanisms of organophosphate resistance mediated by metabolic detoxification enzymes in L.migratoria manilensis
The metabolic detoxification enzymes(cytochrome P450 monooxygenases(P450s),general esterases(ESTs) and glutathione S-transferases(GSTs)) were biochemical characterized.The results showed that the mean of the ESTs activities in the HH was 2.1-3.2-fold significantly higher than that of the SS whenα-NA,α-NB andβ-NA were used as substrates,respectively.The GST activity was 1.2-2.0- fold significantly higher in the HH than in the LS when CDNB and DCNB were used as substrates,respectively.For P450 activity assay,there was no significant difference between the HH and SS of the locust.The non-denaturing PAGE analysis of ESTs in the HH displayed the similar bands but increased staining intensity compared with the LS.Therefore,ESTs and GSTs played major roles in conferring malathion resistance in the HH.
Two organophosphorus inhibitors such as paraoxon and malaoxon were used to compare the sensitivity levels of carboxylesterases(CarEs) between the HL and WD.Significant differences were observed in pI_(50) values between HL and WD for all the two OP inhibitors.Specifically,CarEs from the HL were 1.16- and 1.27- fold less sensitive to inhibition by paraoxon and malaoxon,respectively,than that from the WD based on the differences in the pI_(50) values between the two populations.However,there were significant differences in CarEs specific activities between the WD and HL of the locust. CarEs specific activities in the WD were 1.75- and 1.50-fold higher than those in the HL whenα-NA andβ-NA were used as substrates,respectively. The increased CarEs specific activities in the WD appeared to be associated with a 1.8-fold decreased susceptibility to malathion.The differences of CarEs activities may attribute to the locust differentiation between HL and WD due to the different locust breeding environment.It is not known whether or not the WD can be resistant to other OPs,the WD might be resistant to other OPs due to the increased Care activity.
3.Bichemical mechanisms of organophosphate resistance mediated by target enzyme in L.migratoria manilensis
The biochemical characteristics of acetylcholinesterase(AChE,EC 1.1.1.7) were compared between the three populations and the susceptible strain of L. migratoria manilensis.AChE from the HH,HL and WD showed 4.04-,9.55-, and 1.99-fold significantly higher activity than that from the SS.Kinetic studies indicated that the K_m values of AChE from the three populations presented significantly difference compared with the SS.Inhibition kinetics revealed that AChE from the HH was 3.2-,2.2- and 1.1-fold less sensitive than that from the SS to malaoxon,chlorpyrfos oxon and phoxim respectively, AChE from the HL was 11.3- and 19.2-fold less sensitive to inhibition by malaoxon and chlopyrifos oxon,respectively,than that from the SS. Meanwhile,AChE from the WD was 4.8- and 2.4-fold less sensitive to inhibition by malaoxon and chlopyrifos oxon,respectively,in despite of 1.8-fold marginal resistance to malathion.
Combined with the biochemical mechanisms of organophosphate resistance mediated by metabolic detoxification enzymes in L.migratoria manilensis,it was indicated that the malathion resistance in the HH was conferred by multiple mechanisms,including increased detoxification by ESTs and GSTs,and increased activity and reduced sensitivity of AChE to OP inhibition;the mild resistance to malathion in HL was associated with reduced sensitivity and increased catalytic activity of ACHE;the WD might be resistant to other OPs due to the decreased AChE sensitivity.Consequently, it was proposed that L.migratoria manilensis from different populations utilized different mechanisms,even including multiple mechanisms,to adapt to the OP insecticide selective pressure.
4.Molecular mechanisms of organophosphate resistance in L.migratoria manilensis
Three CarEs gene cDNA fragments of CarE1,CarE2 and CarE3 were cloned using the degenerate primers designed from conserved motifs in CarEs gene family and touchdown PCR program.According to the conservative regions(motifs) of CarEs and GSTs in insects,the database of expressing sequence tags(ESTs) in L.migratoria manilensis was searched, spliced,32 ESTs of CarEs and 12 ESTs of GSTs were obtained by bioinformatics methods.Total 34 CarEs gene fragments were produced by analyzing 32 ESTs of CarEs and 3 CarEs gene cDNA fragments cloned.The expression levels of these CarEs and GSTs gene were compared between the HH and SS using regular RT-PCR,then differential expression genes were further analyzed by real-time quantitative PCR.It was showed that CarE1, CarE4,CarE6,CarE12,CarE15,CarE19,CarE22,CarE23,CarE26,CarE27, CarE30 and CarE32 in the HH were expressed 3.29,1.13,8.78,3.16,3.17, 11.84,2.24,4.92,6.24,4.66,4.47 and 4.73- fold higher than that in the SS respectively;GST1,GST2,GST3 and GST12 in the HH were expressed 1.25, 1.33,2.47 and 1.61- fold higher than that in the SS respectively.These results indicated that higher expression level of CarEs and GSTs may contribute to the insecticides resistance of L.migratoria manilensis in HH.
The gene cloning and Western blotting of acetylcholinesterase(ACHE) suggested that the AChE gene of insects from Orthoptera may be different from other insects.It will be necessary to be further confirmed.
Elucidation of the resistance spectrum and resistance mechanisms in L. migratoria rnanilensis will help us to make knowledge-based selections of insecticides for locust control and resistance management in the field.
In our study,we searched all of the CarEs and GSTs genes from the EST database of L.migratoria manilensis using the genome informatics(for example EST),screened out the CarEs and GSTs genes correlated with insecticide resistance and characterized these genes systematicly.
Consequently,these findings were significant to help researchers detect and monitor insecticide resistance in L.migratoria manilensis,and to devise effective chemical control methods for this important insect pest in the field.
一、东亚飞蝗对有机磷农药的敏感性分析和增效作用测定
本研究针对我国东亚飞蝗发生规模、分布特点、农药使用背景差异和种群遗传特征,采集了河北黄骅、海南黄流和山东无棣田间种群,选择有代表性的有机磷农药进行敏感性分析。结果发现,东亚飞蝗无棣种群对马拉硫磷最为敏感,黄骅种群对毒死蜱和辛硫磷仅有5.4和2.9倍的抗药性,而黄骅和黄流两种群对马拉硫磷已产生了57.5和14.8倍的抗药性。可以看出,由于东亚飞蝗不同种群马拉硫磷选择压力的差异已经造成不同田间种群对其抗性程度产生了变化,而且不同的有机磷农药对同一种群的选择压力已产生不同的抗药性水平。由此说明,研究东亚飞蝗对常用杀虫剂抗性发生现状,建立有效的抗性治理策略,是十分必要的。
通过增效剂磷酸三苯酯(TPP)、顺丁烯二酸二乙酯(DEM)及氧化胡椒基丁醚(PBO)在河北黄骅田间种群和敏感品系中对马拉硫磷的增效试验,结果发现,无论在黄骅种群还是敏感品系中,PBO对马拉硫磷都没有增效作用(增效比分别为0.9和1.1),但在黄骅种群中TPP和DEM对马拉硫磷表现出显著的增效作用(增效比分别为16.2和3.3),而在敏感品系中TPP和DEM没有明显的增效作用(增效比分别为1.4和1.2);由此推测,酯酶和谷胱甘肽S-转移酶与黄骅种群马拉硫磷抗性相关,而细胞色素P450与马拉硫磷抗性无关。
二、东亚飞蝗有机磷抗性代谢解毒酶生化机制研究
本研究分析了东亚飞蝗代谢解毒酶系(酯酶、谷胱甘肽S-转移酶和细胞色素P450氧化酶)的生化特性。东亚飞蝗河北黄骅种群和敏感品系代谢解毒酶活性表现出不同程度的差异,其中黄骅种群酯酶对α-NA、α-NB及β-NA三种底物的水解活性和谷胱甘肽S-转移酶对底物CDNB和DCNB水解活性显著高于敏感品系,分别是敏感品系的2.1-3.2倍和1.2-2.0倍。而对于细胞色素P450活性,两者之间没有显著差异。酯酶聚丙烯酰胺凝胶电泳结果显示,黄骅种群和敏感品系电泳图谱中条带带型没有差异,但黄骅种群条带染色明显深于敏感品系。据此推测,黄骅种群与敏感品系之间的酯酶活性和谷胱甘肽S-转移酶差异与河北黄骅东亚飞蝗的抗药性有关。
用对氧磷、马拉氧磷两种有机磷抑制剂对东亚飞蝗海南黄流和山东无棣种群的羧酸酯酶进行体外抑制,实验结果表明,黄流和无棣种群的pI_(50)值有显著差异,黄流种群分别是山东种群的1.16和1.27倍,说明黄流种群对对氧磷和马拉氧磷的敏感性较低。然而黄流种群和无棣种群的羧酸酯酶活性测定结果表明,以α-NA和β-NA为底物时,无棣种群羧酸酯酶活性分别是黄流种群1.75和1.50倍,具有显著差异。动力学研究表明,无棣种群和黄流种群的羧酸酯酶K_和V_(max)值没有显著差异。据分析黄流种群和无棣种群羧酸酯酶活性的差异可能是由于生态环境的不同所导致的种群分化引起的。根据目前的研究结果尚难以断定对马拉硫磷生物测定中LD_(50)值较低的无棣种群对其它农药的敏感性如何。基于无棣种群较高的羧酸酯酶活性,我们推测,东亚飞蝗无棣种群可能对其它的农药具有较高的抗药性。
三、东亚飞蝗有机磷抗性靶标酶生化机制研究
本研究分析了东亚飞蝗有机磷靶标乙酰胆碱酯酶(acetylcholinesterase,AChE)的生化特性。对东亚飞蝗黄骅种群、黄流种群、无棣种群和敏感品系AChE活性和动力学比较结果表明,三个东亚飞蝗田间种群AChE活性与敏感品系相比有显著差异,黄骅种群、黄流种群和无棣种群AChE活性分别是敏感品系的4.04、9.55和1.99倍,并且三个田间种群的K_m值与敏感品系相比也有显著差异;根据有机磷杀虫剂对AChE的体外抑制结果,河北黄骅种群AChE对马拉氧磷、氧化毒死蜱及辛硫磷的敏感性与敏感品系相比分别下降了3.2、2.2和1.1倍。而海南黄流种群对马拉氧磷和氧化毒死蜱的敏感性分别下降了11.3和19.2倍。同时发现,对马拉硫磷生物测定中仅有1.8倍抗药性的山东无棣种群,其AChE对马拉氧磷和氧化毒死蜱的敏感性也分别下降了4.8和2.4倍。
结合东亚飞蝗解毒酶的生化机制研究,本文认为黄骅种群对马拉硫磷的抗性机理主要包括酯酶、谷胱甘肽S-转移酶活性升高和乙酰胆碱酯酶的活性升高以及敏感性下降。黄流种群产生抗性主要是由于AChE的活性升高和敏感性降低引起的。而无棣种群可能对其它的农药有较高的抗药性,导致其AChE敏感性的显著下降。据此认为,东亚飞蝗不同种群抗性产生机制是不尽相同的,有可能包括多种不同的抗性机制。
四、东亚飞蝗有机磷抗性分子机制研究
本研究利用简并性引物和降落PCR技术从东亚飞蝗中克隆获得了3个东亚飞蝗羧酸酯酶(carboxylesterases,CarEs)基因的cDNA片段,分别命名为CarE1,CarE2和CarE3。根据昆虫羧酸酯酶和谷胱甘肽S-转移酶的保守区(motif),采用生物信息学方法对东亚飞蝗表达序列标签(expressing sequence tags,ESTs)数据库进行了全面搜索,并对其进行了羧酸酯酶和谷胱甘肽S-转移酶ESTs拼接、注释,获得了32条羧酸酯酶ESTs和12条谷胱甘肽S-转移酶ESTs,与克隆获得的3条羧酸酯酶基因片段同源比对和拼接后,共得到34条羧酸酯酶基因片段。采用实时定量PCR技术对上述羧酸酯酶和谷胱甘肽S-转移酶基因在黄骅种群和敏感品系中的表达量进行了比较,结果发现,羧酸酯酶基因CarE1、CarE4、CarE6、CarE12、CarE15、CarE19、CarE22、CarE23、CarE26、CarE27、CarE30和CarE32在黄骅种群中的表达量分别是敏感品系的3.29、1.13、8.78、3.16、3.17、11.84、2.24、4.92、6.24、4.66、4.47和4.73倍;谷胱甘肽S-转移酶基因GST1、GST2、GST3和GST12的表达量分别是敏感品系的1.25、1.33、2.47和1.61倍。据此认为这些羧酸酯酶基因和谷胱甘肽S-转移酶基因表达提高与东亚飞蝗的抗药性密切相关。
对东亚飞蝗AChE基因克隆和Western blot分析结果认为直翅目昆虫AChE基因可能有其特殊性,尚有待进一步研究证实。
本文通过东亚飞蝗不同种群对有机磷农药的敏感性分析,揭示了东亚飞蝗的田间抗性现状。通过生化和分子生物学方法分析了东亚飞蝗对有机磷的抗性机理,发现东亚飞蝗不同种群在产生抗性过程中的机制是不尽相同的,有可能包括多种不同的抗性机制。这些研究结果对于有效地进行抗性治理,延缓东亚飞蝗的抗性发展均具有重要的实践意义。
本文首次借助现有的东亚飞蝗的ESTs数据库对抗性相关解毒酶基因进行了系统研究,用real-time PCR对代谢解毒酶羧酸酯酶基因和谷胱甘肽S-转移酶基因在抗性和敏感种群中差异表达进行了定量分析,确定了抗性形成过程中的关键代谢解毒酶基因,这将极大地推动东亚飞蝗抗性机制的系统深入研究。
综上所述,本文研究结果对东亚飞蝗抗性机制和东亚飞蝗的抗性治理均具有重要的理论和实践意义。
Locusta migratoria manilensis(Meyen) is one of the seriously endangering agricultural pests.Organophosphate(OP)insecticides have been used extensively to control the locust.In order to delaying organophosphorus resistance and carrying out effective management strategies of different populations of L.migratoria manilensis,the objectives of this study were to evaluate the resistance spectrum of the field locust populations using the bioassay to OPs and investigate major biochemical and molecular mechanisms of resistance to OPs by the modem molecular biology technology.Elucidation of the resistance spectrum and resistance mechanisms in L.migratoria manilensis will be significantly important in theory and practice.
1.Resistance spectrum to selected OP insecticides and synergism of TPP, PBO and DEM
Based on the scale,distribution,history of insecticides control and population characteristics of L.migratoria manilensis,the field populations of the locust were collected from Huanghua County of Hebei Province, Huangliu County of Hainan Province and Wudi County of Shandong Province.The sensitivity analysis or resistance spectrums to the representative OP insecticides were evaluated.The results showed that, compared with the susceptible strain(SS),the population of Wudi population (WD) was the most sensitive to malathion and Huanghua population(HH) showed 2.9-fold to phoxim,5.4-fold to chlorpyrifos of resistance levels. However,HH and Huangliu population(HL) developed 57.5-fold and 14.8-fold resistant to malathion.In conclusion,As a consequence of the various malathion selective pressures,there has been developed different resistance to malathion in different locusts populations and various resistant levels in the same locust population has been developed by the different OPs. Consequently,it is significantly necessary to evaluate the resistance spectrum and detect the effective resistance management strategy of the field locust populations.
The malathion resistance of the HH was significantly diminished by TPP (synergism ratio:16.2) and DEM(3.3),but was unchanged by PBO(0.9).In contrast,none of these synergists significantly affected the toxicity of malathion in the SS.These results suggested that both ESTs and GSTs were involved in OP resistance in the HH,whereas P450s were unlikely to be involved in the resistance simply because PBO did not synergize the toxicity of malathion by the synergism of triphenyl phosphate(TPP),diethyl meleate (DEM) and piperonyl butoxide(PBO) on malathion in the HH and SS of the locust.
2.Biochemical mechanisms of organophosphate resistance mediated by metabolic detoxification enzymes in L.migratoria manilensis
The metabolic detoxification enzymes(cytochrome P450 monooxygenases(P450s),general esterases(ESTs) and glutathione S-transferases(GSTs)) were biochemical characterized.The results showed that the mean of the ESTs activities in the HH was 2.1-3.2-fold significantly higher than that of the SS whenα-NA,α-NB andβ-NA were used as substrates,respectively.The GST activity was 1.2-2.0- fold significantly higher in the HH than in the LS when CDNB and DCNB were used as substrates,respectively.For P450 activity assay,there was no significant difference between the HH and SS of the locust.The non-denaturing PAGE analysis of ESTs in the HH displayed the similar bands but increased staining intensity compared with the LS.Therefore,ESTs and GSTs played major roles in conferring malathion resistance in the HH.
Two organophosphorus inhibitors such as paraoxon and malaoxon were used to compare the sensitivity levels of carboxylesterases(CarEs) between the HL and WD.Significant differences were observed in pI_(50) values between HL and WD for all the two OP inhibitors.Specifically,CarEs from the HL were 1.16- and 1.27- fold less sensitive to inhibition by paraoxon and malaoxon,respectively,than that from the WD based on the differences in the pI_(50) values between the two populations.However,there were significant differences in CarEs specific activities between the WD and HL of the locust. CarEs specific activities in the WD were 1.75- and 1.50-fold higher than those in the HL whenα-NA andβ-NA were used as substrates,respectively. The increased CarEs specific activities in the WD appeared to be associated with a 1.8-fold decreased susceptibility to malathion.The differences of CarEs activities may attribute to the locust differentiation between HL and WD due to the different locust breeding environment.It is not known whether or not the WD can be resistant to other OPs,the WD might be resistant to other OPs due to the increased Care activity.
3.Bichemical mechanisms of organophosphate resistance mediated by target enzyme in L.migratoria manilensis
The biochemical characteristics of acetylcholinesterase(AChE,EC 1.1.1.7) were compared between the three populations and the susceptible strain of L. migratoria manilensis.AChE from the HH,HL and WD showed 4.04-,9.55-, and 1.99-fold significantly higher activity than that from the SS.Kinetic studies indicated that the K_m values of AChE from the three populations presented significantly difference compared with the SS.Inhibition kinetics revealed that AChE from the HH was 3.2-,2.2- and 1.1-fold less sensitive than that from the SS to malaoxon,chlorpyrfos oxon and phoxim respectively, AChE from the HL was 11.3- and 19.2-fold less sensitive to inhibition by malaoxon and chlopyrifos oxon,respectively,than that from the SS. Meanwhile,AChE from the WD was 4.8- and 2.4-fold less sensitive to inhibition by malaoxon and chlopyrifos oxon,respectively,in despite of 1.8-fold marginal resistance to malathion.
Combined with the biochemical mechanisms of organophosphate resistance mediated by metabolic detoxification enzymes in L.migratoria manilensis,it was indicated that the malathion resistance in the HH was conferred by multiple mechanisms,including increased detoxification by ESTs and GSTs,and increased activity and reduced sensitivity of AChE to OP inhibition;the mild resistance to malathion in HL was associated with reduced sensitivity and increased catalytic activity of ACHE;the WD might be resistant to other OPs due to the decreased AChE sensitivity.Consequently, it was proposed that L.migratoria manilensis from different populations utilized different mechanisms,even including multiple mechanisms,to adapt to the OP insecticide selective pressure.
4.Molecular mechanisms of organophosphate resistance in L.migratoria manilensis
Three CarEs gene cDNA fragments of CarE1,CarE2 and CarE3 were cloned using the degenerate primers designed from conserved motifs in CarEs gene family and touchdown PCR program.According to the conservative regions(motifs) of CarEs and GSTs in insects,the database of expressing sequence tags(ESTs) in L.migratoria manilensis was searched, spliced,32 ESTs of CarEs and 12 ESTs of GSTs were obtained by bioinformatics methods.Total 34 CarEs gene fragments were produced by analyzing 32 ESTs of CarEs and 3 CarEs gene cDNA fragments cloned.The expression levels of these CarEs and GSTs gene were compared between the HH and SS using regular RT-PCR,then differential expression genes were further analyzed by real-time quantitative PCR.It was showed that CarE1, CarE4,CarE6,CarE12,CarE15,CarE19,CarE22,CarE23,CarE26,CarE27, CarE30 and CarE32 in the HH were expressed 3.29,1.13,8.78,3.16,3.17, 11.84,2.24,4.92,6.24,4.66,4.47 and 4.73- fold higher than that in the SS respectively;GST1,GST2,GST3 and GST12 in the HH were expressed 1.25, 1.33,2.47 and 1.61- fold higher than that in the SS respectively.These results indicated that higher expression level of CarEs and GSTs may contribute to the insecticides resistance of L.migratoria manilensis in HH.
The gene cloning and Western blotting of acetylcholinesterase(ACHE) suggested that the AChE gene of insects from Orthoptera may be different from other insects.It will be necessary to be further confirmed.
Elucidation of the resistance spectrum and resistance mechanisms in L. migratoria rnanilensis will help us to make knowledge-based selections of insecticides for locust control and resistance management in the field.
In our study,we searched all of the CarEs and GSTs genes from the EST database of L.migratoria manilensis using the genome informatics(for example EST),screened out the CarEs and GSTs genes correlated with insecticide resistance and characterized these genes systematicly.
Consequently,these findings were significant to help researchers detect and monitor insecticide resistance in L.migratoria manilensis,and to devise effective chemical control methods for this important insect pest in the field.
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