抗氯氟氰菊酯甜菜夜蛾对昆虫生长调节剂的敏感性及其神经ATPase和多功能氧化酶活性变化
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摘要
甜菜夜蛾(Spodoptera exigua)属鳞翅目夜蛾科,是一种世界性分布的杂食性重要农业害虫,主要危害蔬菜、棉花和观赏性植物等。由于一直依靠化学防治,甜菜夜蛾已对多种类型农药尤其是菊酯类杀虫剂产生抗性。昆虫生长调节剂如虫酰肼和几丁质合成抑制剂类杀虫剂是近年来防治甜菜夜蛾的主要替代农药。这类杀虫剂对鳞翅目害虫都表现了高度的选择性,对哺乳动物、鸟、鱼类、其它脊椎动物和大多数天敌无毒害作用,是害虫综合防治的重要优选药剂。害虫对药剂都存在产生抗药性的风险,为了更有效地控制甜菜夜蛾的危害,掌握甜菜夜蛾对此类杀虫剂的敏感性变化十分重要。
昆虫的生命活动是通过神经系统电位传导进行调控的,神经电位是由神经膜内外离子浓度不同形成的电位差,信号传导则是神经膜电位发生连续变化而向下传递的。神经膜内外不同离子浓度的调节主要靠神经膜上的ATPase进行调控。Na-K-ATPase催化ATP末端磷酸水解,并利用其高能磷酸键释放的自由能进行Na+、K+逆电化学梯度运输,维持细胞膜内外Na+、K+离子浓度的相对恒定及渗透压的平衡。Ca2+对轴突传导、突触神经递质释放及神经膜的稳定性等起重要作用,正常情况下神经膜外Ca2+离子的浓度高于膜内浓度,Ca2+在膜内外的调节主要靠Ca-ATPase和Ca-Mg-ATPase来进行。拟除虫菊酯类杀虫剂的作用靶标主要是神经膜上的钠离子通道和维持膜内外离子浓度稳定的Na-K-ATPase、Ca-ATPase和Ca-Mg-ATPase等。目前已报道许多害虫对拟除虫菊酯类杀虫剂产生了不同程度的抗性,有些害虫的抗性甚至达到了高水平或极高水平阶段。钠离子通道变异和神经ATPase活性下降是产生抗药性的重要原因。
本文研究内容包括三部分:抗性甜菜夜蛾对昆虫生长调节剂的敏感性测定、抗性甜菜夜蛾神经ATPase活性变化和多功能氧化酶活性变化的研究。研究结果如下:
1、在室内相同条件下,分别用点滴法和浸叶法测定甜菜夜蛾抗氯氟氰菊酯品系和敏感品系3龄幼虫LD50(ug/头)或LC50值(mg/L)。用点滴法测定结果表明,甜菜夜蛾3龄幼虫对氯氟氰菊酯的抗性倍数是5163.0倍;用浸叶法测定甜菜夜蛾3龄幼虫对氯氟氰菊酯的抗药性倍数是985.2倍。浸叶法测定甜菜夜蛾抗性品系3龄幼虫对昆虫生长调节剂的敏感性结果表明,甜菜夜蛾抗性品系和敏感品系对供试药剂的敏感性均存在明显差异。甜菜夜蛾敏感品系对供试药剂的敏感性由高到低顺序为甲氧酰肼(1.866 mg/L)﹥虫酰肼(5.574 mg/L)﹥氟铃脲(13.37 mg/L)﹥抑食肼(58.61 mg/L)﹥除虫脲(178.5 mg/L);甜菜夜蛾抗性品系对供试药剂的敏感性由高到低顺序为甲氧酰肼(4.863 mg/L)﹥虫酰肼(21.45mg/L)﹥氟铃脲(64.36mg/L)﹥抑食肼(325.4mg/L)﹥除虫脲(567.3mg/L)。测定结果表明,与敏感品系相比,甜菜夜蛾抗性品系3龄幼虫对昆虫生长调节剂甲氧酰肼、虫酰肼、氟铃脲、抑食肼、除虫脲的敏感性分别下降:3.81、5.55、4.81、3.85和2.61倍,其中对部分药剂的敏感性下降倍数达到5倍以上,表现出低水平抗药性。
2、分别测定了甜菜夜蛾敏感品系和抗氯氟氰菊酯品系神经Na-K-ATPase、Ca-ATPase和Ca-Mg-ATPase的活力,结果表明,甜菜夜蛾敏感品系和抗性品系Na-K-ATPase活力差异不显著,而抗性品系Ca-ATPase和Ca-Mg-ATPase活力明显低于敏感品系。分别测定氯氟氰菊酯对敏感和抗性品系Na-K-ATPase、Ca-ATPase和Ca-Mg-ATPase活力的抑制作用,结果表明,在浓度为10-8～10-3 mol/L时,氯氟氰菊酯对敏感和抗性品系Na-K-ATPase、Ca-ATPase和Ca-Mg-ATPase的活力均有抑制作用,并且对敏感品系的抑制作用要高于对抗性品系的抑制作用。如氯氟氰菊酯浓度为10-4 mol/L时,对敏感品系Na-K-ATPase活力抑制率为29.6%,抗性品系为21.8%;对敏感品系Ca-ATPase活力抑制率为34.3%,抗性品系为21.9%;对敏感品系Ca-Mg-ATPase活力抑制率为22.3%,抗性品系为16.9%。结果表明甜菜夜蛾抗性品系上述3种ATP酶对氯氟氰菊酯的敏感性已明显下降。
3、测定了抗氯氟氰菊酯甜菜夜蛾种群在室内用人工饲料连续饲养和未用药剂筛选条件下多功能氧化酶系活性的变化规律。测定甜菜夜蛾田间种群室内F2、F20、F41代及敏感品系5龄幼虫中肠微粒体甲氧试卤灵-O-脱甲基酶、乙氧试卤灵-O-脱乙基酶、芳香基羟基化酶及艾氏剂环氧化酶活性结果表明,与敏感品系相比,田间种群甲氧试卤灵-O-脱甲基酶和艾氏剂环氧化酶的活性仅F2代差异显著,F20和F41代差异不显著;乙氧试卤灵-O-脱乙基酶和芳香基羟基化酶的活性F2、F20、F41代均差异显著。表明微粒体多功能氧化酶系不同酶在甜菜夜蛾不同抗性水平阶段的作用不同。
The beet armyworm, Spodoptera exigua (Hbüner), is a polyphagous noctuid of worldwide importance that feeds on various agricultural crops, including vegetable, cotton and omamental. The heavy dependence on insecticides for control of this pest has resulted in the emergence of resistance to a variety of insecticides. Insect growth regulators (IGRs) including tebufenozide and chitin inhibitors are the major insecticides used to control Spodoptera exigua in the field. They have selective toxicity towards different pests, and have less adverse effects on mammals, birds, fishes and other vertebtrates, as well as various beneficial insects. In order to maintain long effective controlling, we must pay more attention to sensitivity of these insecticides to beet armyworm.
The life activities of insect is controlled by its nerve system potential conduction. Nerve potential is the potential difference dues to the different ion concentration of inside and outside neurilemma. ATPase in the neurilemma mostly controls the different ion concentration of inside and outside neurilemma. Na-K-ATPase catalyses the bottom phosphoric acid of ATP to be hydrolyzed, and the high-energy phosphate bond liberatives free energy which can be used to counter-transport Na+、K+ with electrochemistry gradient, in order to maintain the relative invariablenes and balance of osmotic pressure of Na+、K+ concentration of inside and outside neurilemma. Ca2+ plays an important part in neuraxon conduction, synapse neurotransmitters release and neurilemma stability. In normal condition, Ca2+ concentration of outside neurilemma is higher than that inside neurilemma. And the modulation of Ca2+ inside and outside neurilemma mainly depends on Ca-ATPase and Ca-Mg-ATPase. Target sites of pyrethroid insecticides are sodium channel on the neurilemma and Na-K-ATPase、Ca-ATPase and Ca-Mg-ATPase which can maintain the stability of ions inside and outside neurilemma. The variation of sodium channel and the decreased activities of nerve ATPase are main causes to pyrethroid insecticide resistance.
Three parts are involved in this thesis: (1) sensitivity of insect growth regulators to resistant strain of beet armyworm, Spodoptera exigua and (2) inhibition of Lambda-cyhalothrin on Nerve Na-K-ATPase, Ca-ATPase and Ca-Mg-ATPase in the Beet Armyworm.(3)active changes of multi-function oxidases in Beet Armyworm.The results are as follows.
1. In the same condition in laboratory, LD50 or LC50 of 3rd instar larva of susceptible strain and Lambda-cyhalothrin resistant strain of beet armyworm were determined. The results detected by topical application showed that 3rd instar larva of the resistant strain had 5163.0 fold resistance to Lambda-cyhalothrin compared to the susceptible strain. The results detected by dipping method showed that 3rd instar larva of the resistant strains had 985.2 fold resistance to Lambda-cyhalothrin. Sensitivity of five insect growth regulators to resistant and susceptible stains of beet armyworm was evaluated. The order of sensitivity to 3rd instar larvae of susceptible strain detected by leaf-dipping method was methoxyfenozide﹥tebufenozide﹥hexaflumuron﹥RH-5849﹥diflubenzuron. The resistant stain showed the similar resistant orders to 5 IGRs as susceptible strain. Compared with susceptible strain, the sensitivity of resistant stain to methoxyfenozide, tebufenozide, hexaflumuron, RH-5849 and diflubenzuron decreased 3.81, 5.55, 4.81, 3.85 and 2.61 folds respectively.
2. The activities of Na-K-ATPase, Ca-ATPase and Ca-Mg-ATPase in susceptible and Lambda-cyhalothrin resistant strains of beet armyworm, Spodoptera exigua, were determined. The results indicated that there was no significant difference in Na-K-ATPase activity between the two strains, but Ca-ATPase and Ca-Mg-ATPase activities were much lower in resistant strain than in susceptible strain. Lambda-cyhalothrin at concentrations of 10-8-10-3 mol/L inhibited obviously the three above ATPase activities in two strains, and the inhibition was higher in susceptible strain than in resistant strain. For example, At concentration of 10-4 mol/L, the inhibition on Na-K-ATPase, Ca-ATPase and Ca-Mg-ATPase activities was 29.6%, 34.3% and 22.3% in susceptible strain and 21.8%, 21.9% and 16.9% in resistant strain respectively. The results showed that the sensitivity of three above ATPases in resistant strain decreased to Lambda-cyhalothrin.
3. Larvae of beet armyworm were reared on artificial diet and evaluated for resistance to Lambda-cyhalothrin for 43 generations under laboratory condition without exposure to any insecticides (for non-selection). Four indices of monoxygenase activities [methoxyresorufin O-demethylase (MROD), ethoxyresorufin O-deethylase (EROD), arylhydrocarbon hydroxylase (AHH) and aldrin epoxidase (AE)] were compared between F2, or F20, or F43 generation and the susceptible strain, respectively. Compared with the susceptible strain, the activities of MROD and AE in midguts in fifth instar larvae of F2 generation showed significant difference, those of F20 and F43 generations showed no significant difference. In the same way, the activities of MROD and AHH of F2, F20 and F43 generation also showed significant difference. This indicated that the resistance of Spodoptera exigua to lambda-cyhalothrin was closely correlated with multi-function oxidase which may play different roles at different resistant levels.
昆虫的生命活动是通过神经系统电位传导进行调控的,神经电位是由神经膜内外离子浓度不同形成的电位差,信号传导则是神经膜电位发生连续变化而向下传递的。神经膜内外不同离子浓度的调节主要靠神经膜上的ATPase进行调控。Na-K-ATPase催化ATP末端磷酸水解,并利用其高能磷酸键释放的自由能进行Na+、K+逆电化学梯度运输,维持细胞膜内外Na+、K+离子浓度的相对恒定及渗透压的平衡。Ca2+对轴突传导、突触神经递质释放及神经膜的稳定性等起重要作用,正常情况下神经膜外Ca2+离子的浓度高于膜内浓度,Ca2+在膜内外的调节主要靠Ca-ATPase和Ca-Mg-ATPase来进行。拟除虫菊酯类杀虫剂的作用靶标主要是神经膜上的钠离子通道和维持膜内外离子浓度稳定的Na-K-ATPase、Ca-ATPase和Ca-Mg-ATPase等。目前已报道许多害虫对拟除虫菊酯类杀虫剂产生了不同程度的抗性,有些害虫的抗性甚至达到了高水平或极高水平阶段。钠离子通道变异和神经ATPase活性下降是产生抗药性的重要原因。
本文研究内容包括三部分:抗性甜菜夜蛾对昆虫生长调节剂的敏感性测定、抗性甜菜夜蛾神经ATPase活性变化和多功能氧化酶活性变化的研究。研究结果如下:
1、在室内相同条件下,分别用点滴法和浸叶法测定甜菜夜蛾抗氯氟氰菊酯品系和敏感品系3龄幼虫LD50(ug/头)或LC50值(mg/L)。用点滴法测定结果表明,甜菜夜蛾3龄幼虫对氯氟氰菊酯的抗性倍数是5163.0倍;用浸叶法测定甜菜夜蛾3龄幼虫对氯氟氰菊酯的抗药性倍数是985.2倍。浸叶法测定甜菜夜蛾抗性品系3龄幼虫对昆虫生长调节剂的敏感性结果表明,甜菜夜蛾抗性品系和敏感品系对供试药剂的敏感性均存在明显差异。甜菜夜蛾敏感品系对供试药剂的敏感性由高到低顺序为甲氧酰肼(1.866 mg/L)﹥虫酰肼(5.574 mg/L)﹥氟铃脲(13.37 mg/L)﹥抑食肼(58.61 mg/L)﹥除虫脲(178.5 mg/L);甜菜夜蛾抗性品系对供试药剂的敏感性由高到低顺序为甲氧酰肼(4.863 mg/L)﹥虫酰肼(21.45mg/L)﹥氟铃脲(64.36mg/L)﹥抑食肼(325.4mg/L)﹥除虫脲(567.3mg/L)。测定结果表明,与敏感品系相比,甜菜夜蛾抗性品系3龄幼虫对昆虫生长调节剂甲氧酰肼、虫酰肼、氟铃脲、抑食肼、除虫脲的敏感性分别下降:3.81、5.55、4.81、3.85和2.61倍,其中对部分药剂的敏感性下降倍数达到5倍以上,表现出低水平抗药性。
2、分别测定了甜菜夜蛾敏感品系和抗氯氟氰菊酯品系神经Na-K-ATPase、Ca-ATPase和Ca-Mg-ATPase的活力,结果表明,甜菜夜蛾敏感品系和抗性品系Na-K-ATPase活力差异不显著,而抗性品系Ca-ATPase和Ca-Mg-ATPase活力明显低于敏感品系。分别测定氯氟氰菊酯对敏感和抗性品系Na-K-ATPase、Ca-ATPase和Ca-Mg-ATPase活力的抑制作用,结果表明,在浓度为10-8～10-3 mol/L时,氯氟氰菊酯对敏感和抗性品系Na-K-ATPase、Ca-ATPase和Ca-Mg-ATPase的活力均有抑制作用,并且对敏感品系的抑制作用要高于对抗性品系的抑制作用。如氯氟氰菊酯浓度为10-4 mol/L时,对敏感品系Na-K-ATPase活力抑制率为29.6%,抗性品系为21.8%;对敏感品系Ca-ATPase活力抑制率为34.3%,抗性品系为21.9%;对敏感品系Ca-Mg-ATPase活力抑制率为22.3%,抗性品系为16.9%。结果表明甜菜夜蛾抗性品系上述3种ATP酶对氯氟氰菊酯的敏感性已明显下降。
3、测定了抗氯氟氰菊酯甜菜夜蛾种群在室内用人工饲料连续饲养和未用药剂筛选条件下多功能氧化酶系活性的变化规律。测定甜菜夜蛾田间种群室内F2、F20、F41代及敏感品系5龄幼虫中肠微粒体甲氧试卤灵-O-脱甲基酶、乙氧试卤灵-O-脱乙基酶、芳香基羟基化酶及艾氏剂环氧化酶活性结果表明,与敏感品系相比,田间种群甲氧试卤灵-O-脱甲基酶和艾氏剂环氧化酶的活性仅F2代差异显著,F20和F41代差异不显著;乙氧试卤灵-O-脱乙基酶和芳香基羟基化酶的活性F2、F20、F41代均差异显著。表明微粒体多功能氧化酶系不同酶在甜菜夜蛾不同抗性水平阶段的作用不同。
The beet armyworm, Spodoptera exigua (Hbüner), is a polyphagous noctuid of worldwide importance that feeds on various agricultural crops, including vegetable, cotton and omamental. The heavy dependence on insecticides for control of this pest has resulted in the emergence of resistance to a variety of insecticides. Insect growth regulators (IGRs) including tebufenozide and chitin inhibitors are the major insecticides used to control Spodoptera exigua in the field. They have selective toxicity towards different pests, and have less adverse effects on mammals, birds, fishes and other vertebtrates, as well as various beneficial insects. In order to maintain long effective controlling, we must pay more attention to sensitivity of these insecticides to beet armyworm.
The life activities of insect is controlled by its nerve system potential conduction. Nerve potential is the potential difference dues to the different ion concentration of inside and outside neurilemma. ATPase in the neurilemma mostly controls the different ion concentration of inside and outside neurilemma. Na-K-ATPase catalyses the bottom phosphoric acid of ATP to be hydrolyzed, and the high-energy phosphate bond liberatives free energy which can be used to counter-transport Na+、K+ with electrochemistry gradient, in order to maintain the relative invariablenes and balance of osmotic pressure of Na+、K+ concentration of inside and outside neurilemma. Ca2+ plays an important part in neuraxon conduction, synapse neurotransmitters release and neurilemma stability. In normal condition, Ca2+ concentration of outside neurilemma is higher than that inside neurilemma. And the modulation of Ca2+ inside and outside neurilemma mainly depends on Ca-ATPase and Ca-Mg-ATPase. Target sites of pyrethroid insecticides are sodium channel on the neurilemma and Na-K-ATPase、Ca-ATPase and Ca-Mg-ATPase which can maintain the stability of ions inside and outside neurilemma. The variation of sodium channel and the decreased activities of nerve ATPase are main causes to pyrethroid insecticide resistance.
Three parts are involved in this thesis: (1) sensitivity of insect growth regulators to resistant strain of beet armyworm, Spodoptera exigua and (2) inhibition of Lambda-cyhalothrin on Nerve Na-K-ATPase, Ca-ATPase and Ca-Mg-ATPase in the Beet Armyworm.(3)active changes of multi-function oxidases in Beet Armyworm.The results are as follows.
1. In the same condition in laboratory, LD50 or LC50 of 3rd instar larva of susceptible strain and Lambda-cyhalothrin resistant strain of beet armyworm were determined. The results detected by topical application showed that 3rd instar larva of the resistant strain had 5163.0 fold resistance to Lambda-cyhalothrin compared to the susceptible strain. The results detected by dipping method showed that 3rd instar larva of the resistant strains had 985.2 fold resistance to Lambda-cyhalothrin. Sensitivity of five insect growth regulators to resistant and susceptible stains of beet armyworm was evaluated. The order of sensitivity to 3rd instar larvae of susceptible strain detected by leaf-dipping method was methoxyfenozide﹥tebufenozide﹥hexaflumuron﹥RH-5849﹥diflubenzuron. The resistant stain showed the similar resistant orders to 5 IGRs as susceptible strain. Compared with susceptible strain, the sensitivity of resistant stain to methoxyfenozide, tebufenozide, hexaflumuron, RH-5849 and diflubenzuron decreased 3.81, 5.55, 4.81, 3.85 and 2.61 folds respectively.
2. The activities of Na-K-ATPase, Ca-ATPase and Ca-Mg-ATPase in susceptible and Lambda-cyhalothrin resistant strains of beet armyworm, Spodoptera exigua, were determined. The results indicated that there was no significant difference in Na-K-ATPase activity between the two strains, but Ca-ATPase and Ca-Mg-ATPase activities were much lower in resistant strain than in susceptible strain. Lambda-cyhalothrin at concentrations of 10-8-10-3 mol/L inhibited obviously the three above ATPase activities in two strains, and the inhibition was higher in susceptible strain than in resistant strain. For example, At concentration of 10-4 mol/L, the inhibition on Na-K-ATPase, Ca-ATPase and Ca-Mg-ATPase activities was 29.6%, 34.3% and 22.3% in susceptible strain and 21.8%, 21.9% and 16.9% in resistant strain respectively. The results showed that the sensitivity of three above ATPases in resistant strain decreased to Lambda-cyhalothrin.
3. Larvae of beet armyworm were reared on artificial diet and evaluated for resistance to Lambda-cyhalothrin for 43 generations under laboratory condition without exposure to any insecticides (for non-selection). Four indices of monoxygenase activities [methoxyresorufin O-demethylase (MROD), ethoxyresorufin O-deethylase (EROD), arylhydrocarbon hydroxylase (AHH) and aldrin epoxidase (AE)] were compared between F2, or F20, or F43 generation and the susceptible strain, respectively. Compared with the susceptible strain, the activities of MROD and AE in midguts in fifth instar larvae of F2 generation showed significant difference, those of F20 and F43 generations showed no significant difference. In the same way, the activities of MROD and AHH of F2, F20 and F43 generation also showed significant difference. This indicated that the resistance of Spodoptera exigua to lambda-cyhalothrin was closely correlated with multi-function oxidase which may play different roles at different resistant levels.
引文
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