福建地区烟粉虱抗药性成因与控制策略
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摘要
烟粉虱Bemisia tabaci（Gennadius）是一种世界性的害虫。近20年来，由于B型生物型烟粉虱的入侵，给我国多个省市农业生产造成严重危害。上世纪末，烟粉虱入侵福建省，随后在漳州、泉州、龙岩等地市相继暴发成灾。一直以来，福建省烟粉虱防治都以化学药剂为主，特别是一些新烟碱类杀虫剂的应用，使得烟粉虱防治在短期内取得较好效果。但由于过于依赖化学药剂防治，造成化学药剂大范围、大强度的使用，几年时间就使得福建省田间烟粉虱的抗药性水平迅速提高，许多常规药剂失去防效，从而引发一系列问题。这不得不让人对以杀虫剂为主的防治策略重新定位和思考。为解决福建省烟粉虱抗药性所引起的为害问题，除对烟粉虱抗药性机制进行研究外，另一个关键问题是对烟粉虱抗药性发生成因有更深刻了解，才能制定符合福建省农业生产实际，合理、科学的烟粉虱控制策略。因此本论文首先通过田间烟粉虱抗药性监测比较，明确福建省烟粉虱田间种群对不同药剂的抗性现状和发展趋势。同时对影响烟粉虱抗药性发生成因（主要包括烟粉虱生物型、种群多样性及遗传分化、寄主和温度因子和杀虫剂亚致死效应等）等研究分析，阐明福建省烟粉虱抗药性发生的关键成因。并在此基础上构建出相应的可持续控制PUSH-PULL(推-拉)策略和生物防治措施，从而为福建省烟粉虱的抗药性治理和烟粉虱可持续控制奠定良好基础。研究结果如下：
（1）采用成虫浸叶生测法对福建省不同地区烟粉虱田间种群的抗药性水平进行了测定。结果表明，与2005年比较，福建省烟粉虱种群除对阿维菌素生物制剂仍未产生明显抗药性；对有机磷和拟除虫菊酯类杀虫剂还依旧保持高水平抗性；对新烟碱类杀虫剂的抗性发展极为迅速。其中漳州种群对吡虫啉、噻虫嗪已由2005年的中等水平抗性发展为2009年的高水平抗性，其它地区种群也由低水平抗性发展为中等水平抗性。表明福建省烟粉虱田间种群对新烟碱类杀虫剂已产生明显抗性，此类药剂在福建省烟粉虱防控上已存在很大风险，要慎重使用。
（2）采用西葫芦银叶测定和mtDNACO I序列比对方法对福建省烟粉虱不同田间种群的生物型进行鉴定，共鉴定出B型和FJ型2种生物型。除采自漳州变叶木上的烟粉虱种群为本地土著种（FJ型）外，其它10个采自大田蔬菜作物的烟粉虱种群均为B型。结果表明福建省大田未发现Q生物型入侵迹象，多年来为害福建省大田作物的烟粉虱生物型仍为B型。从而排除了福建省烟粉虱田间种群抗药性水平的迅速提高是由其它烟粉虱生物型侵入造成的可能。
（3）通过ISSR、RAPD和mtDNACOI分子标记技术对福建省B型烟粉虱不同田间种群进行了群体遗传特征研究和抗性关系分析，结果表明福建地区烟粉虱田间种群遗传多样性丰富，在抗性水平上存在一定水平的遗传分化。这种遗传分化说明在化学药剂选择压力下，不同的抗性基因在烟粉虱群体能得以快速交流，有利于其种群抗药性水平迅速提高，抵御不利的外部环境。也反应了福建省烟粉虱种群自身较强的遗传进化和适应能力。
（4）通过不同寄主和不同温度条件对烟粉虱药剂敏感性的影响研究，结果表明寄主植物和温度对烟粉虱种群的药剂敏感性有一定影响，但这种影响难以引起烟粉虱田间种群几十上百倍的抗性发展。认为吡虫啉等烟碱类杀虫剂高强度使用是B型烟粉虱田间种群对该类型药剂迅速形成抗性的重要原因。
（5）通过吡虫啉杀虫剂亚致死剂量LC_(20)对烟粉虱进行室内抗性筛选试验，结果显示连续筛选14代，吡虫啉亚致死剂量从10ppm提高到100ppm，烟粉虱抗性倍数提高8.64倍。表明杀虫剂的亚致死效应与烟粉虱抗药性有着密切关联，是烟粉虱抗药性水平迅速提高的重要因子。
（6）通过对福建省烟粉虱寄主的调查和烟粉虱与不同寄主植物适生关系研究，了解掌握对不同植物的嗜好性，为诱集物和驱避物选择奠定基础；通过对诱集植物-黄瓜和驱避物-辣椒碱的使用效果以及日本刀角瓢虫生物防治进行评价，最后集成应用诱集植物黄瓜和黄板诱集、驱避物辣椒碱驱避，同时结合在诱集物上释放捕食性天敌日本刀角瓢虫生物防治等技术，构建了适合福建省毛豆生产的可持续控制的PUSH-PULL策略和生物防治措施，为烟粉虱抗药性治理和烟粉虱可持续控制奠定基础。
The whitefly，Bemisia tabaci (Gennadius) is one of the most devastating pests worldwideand occurs in subtropical and tropical agriculture. Over the past20years, Bemisia tabaci hascaused tremendous losses in several provinces and cities in China. At the beginning of lastcentury, the outbreaks of B. tabaci populations occurred in Zhangzhou, Quanzhou, Longyanregions in Fujian province，China. In Fujian province, the control of B. tabaci has relied onchemical agent for all the times. Although, the use of chemical insecticides can cause short-term(acute) effects, especially some neonicotinoid insecticides. The irrational use of chemical agentshas brought about a series of problems, including insecticide resistance increasing quickly in B.tabaci in short time. These problems cause people to reconsider the control strategy of pesticides.In attempting to solve the problems caused by resistance of B. tabaci in Fujian province, one isto study the mechanism of insecticide resistance, another is to evaluate about effect factors on B.tabaci resistance. So we can formulate rational and scientific control strategy for B. tabaciaccording with local agriculture production. For this reason, in this paper, firstly, throughmonitoring and comparing of B. tabaci resistance, the present situation and trend of insecticideresistance in field-collected populations of B. tabaci were defined in Fujian province. Secondly,the influence factors of resistance, including biotype of B. tabaci, population diversity andgenetic differentiation, host plant species, temperature and sublethal effect of pesticides, weresurveyed and analyzed to clarify the key effect factor on B. tabaci resistance in Fujian. On thebasis of these studies, we can construct the corresponding sustainable PUSH-PULL strategy andbiological control measures, and make a good foundation for resistance management of B. tabaciin Fujian province. The results were as follows:
(1) Resistance to eight insecticides in field populations of Bemisia tabaci from six regions(Fuzhou, Zhangzhou, Longyan, Sanming, Nanping, Ningde) of Fujian Province was monitoredby adult leaf-dipping bioassay. The resistance levels to neonicotinoid insecticides developedrapidly in field-collected populations of B. tabaci. Especially, Zhangzhou population, theresistance levels to imidacloprid and thiamethoxam had already developed from medium to highlevel in2005and2009(28-102.53-fold to imidacloprid,29-227.74-fold to thiamethoxam). The populations in other regions exhibited moderate levels of resistance to neonicotinoid insecticides(19.72~44.76fold to imidacloprid,21.18~51.64-fold to thiamethoxam).
(2) The biotypes of different geographical populations of Bemisia tabaci in Fujian provincewere identified base on mtDNACOI molecular marker and squash silverleaf determination. Thebiotype B and FJ were detected. In addition to the population collected from Zhangzhou, whichwas the biotype FJ, the10field-collected populations of B. tabaci were biotype B. Due to notdetecting biotype Q, the identification of biotypes showedthat the biotype B of B. tabaci was stilldestroying agricultural and production in Fujian province over the past10years, and theresistance of B. tabaci was not caused by the recent invasion of biotype Q or others.
(3) The genetic analysis of eight field-collected populations of B. tabaci were characterizedbase on the ISSR, RAPD and mtDNACOI molecular markers in Fujian province. The resultsshowedthat the genetic diversity of field-collected B. tabaci in Fujian was abundant, and acertain level of genetic differentiation has appeared among different geographical populations.Simultaneously, The results also showedthat the Fujian populations of B. tabaci own strongability of genetic evolution and adaptability under the pesticide selection pressure. The geneticdifferentiation of B. tabaci can improve its own survivability under adverse conditions.
(4) The sensitivity of B. tabaci to different pesticides was studied under different host plantsand different temperature conditions. The results showedthat the sensitivity of B. tabaci could beaffected by host plants and temperature, but this kind of impact factors was difficult to causeinsecticide resistance to increase more than several dozens or hundreds of times. We canconclude that the most important reason of resistance increasing rapidly might be due to highstrength use of neonicotinoids.
(5) The resistance screening of laboratory population was used with sublethal doses(LC20)of imidacloprid in the laboratory in2009. After14generations, the resistance of B. tabacito imidacloprid was increased by8.64-fold (LC_(20)value from10.0ppm to100.0ppm). Therealized heritability estimate(h~2) with imidacloprid screening was0.198. The results showed thatcontinuous screening with sublethal doses of imidacloprid could further increase the resistancelevel of B. tabaci, and the sublethal effect of insecticide was closely associated with resistance ofB. tabaci.
(6) Firstly, the host plants of B. tabaci and the adaptability relationship between B. tabaciand different host plants were investigated. Secondly, the control effect of trap plants (cucumber)and repellents (capsaicin) were evaluated, including biological control (Serangini japonicum).Finally, we integrated the application of trap plants(cucumber), yellow board, repellents(capsaicin) and combined with biological control technology (Serangini japonicum), to constructPUSH-PULL strategy and biological control measures for sustainable control of B. tabaci inFujian province.
（1）采用成虫浸叶生测法对福建省不同地区烟粉虱田间种群的抗药性水平进行了测定。结果表明，与2005年比较，福建省烟粉虱种群除对阿维菌素生物制剂仍未产生明显抗药性；对有机磷和拟除虫菊酯类杀虫剂还依旧保持高水平抗性；对新烟碱类杀虫剂的抗性发展极为迅速。其中漳州种群对吡虫啉、噻虫嗪已由2005年的中等水平抗性发展为2009年的高水平抗性，其它地区种群也由低水平抗性发展为中等水平抗性。表明福建省烟粉虱田间种群对新烟碱类杀虫剂已产生明显抗性，此类药剂在福建省烟粉虱防控上已存在很大风险，要慎重使用。
（2）采用西葫芦银叶测定和mtDNACO I序列比对方法对福建省烟粉虱不同田间种群的生物型进行鉴定，共鉴定出B型和FJ型2种生物型。除采自漳州变叶木上的烟粉虱种群为本地土著种（FJ型）外，其它10个采自大田蔬菜作物的烟粉虱种群均为B型。结果表明福建省大田未发现Q生物型入侵迹象，多年来为害福建省大田作物的烟粉虱生物型仍为B型。从而排除了福建省烟粉虱田间种群抗药性水平的迅速提高是由其它烟粉虱生物型侵入造成的可能。
（3）通过ISSR、RAPD和mtDNACOI分子标记技术对福建省B型烟粉虱不同田间种群进行了群体遗传特征研究和抗性关系分析，结果表明福建地区烟粉虱田间种群遗传多样性丰富，在抗性水平上存在一定水平的遗传分化。这种遗传分化说明在化学药剂选择压力下，不同的抗性基因在烟粉虱群体能得以快速交流，有利于其种群抗药性水平迅速提高，抵御不利的外部环境。也反应了福建省烟粉虱种群自身较强的遗传进化和适应能力。
（4）通过不同寄主和不同温度条件对烟粉虱药剂敏感性的影响研究，结果表明寄主植物和温度对烟粉虱种群的药剂敏感性有一定影响，但这种影响难以引起烟粉虱田间种群几十上百倍的抗性发展。认为吡虫啉等烟碱类杀虫剂高强度使用是B型烟粉虱田间种群对该类型药剂迅速形成抗性的重要原因。
（5）通过吡虫啉杀虫剂亚致死剂量LC_(20)对烟粉虱进行室内抗性筛选试验，结果显示连续筛选14代，吡虫啉亚致死剂量从10ppm提高到100ppm，烟粉虱抗性倍数提高8.64倍。表明杀虫剂的亚致死效应与烟粉虱抗药性有着密切关联，是烟粉虱抗药性水平迅速提高的重要因子。
（6）通过对福建省烟粉虱寄主的调查和烟粉虱与不同寄主植物适生关系研究，了解掌握对不同植物的嗜好性，为诱集物和驱避物选择奠定基础；通过对诱集植物-黄瓜和驱避物-辣椒碱的使用效果以及日本刀角瓢虫生物防治进行评价，最后集成应用诱集植物黄瓜和黄板诱集、驱避物辣椒碱驱避，同时结合在诱集物上释放捕食性天敌日本刀角瓢虫生物防治等技术，构建了适合福建省毛豆生产的可持续控制的PUSH-PULL策略和生物防治措施，为烟粉虱抗药性治理和烟粉虱可持续控制奠定基础。
The whitefly，Bemisia tabaci (Gennadius) is one of the most devastating pests worldwideand occurs in subtropical and tropical agriculture. Over the past20years, Bemisia tabaci hascaused tremendous losses in several provinces and cities in China. At the beginning of lastcentury, the outbreaks of B. tabaci populations occurred in Zhangzhou, Quanzhou, Longyanregions in Fujian province，China. In Fujian province, the control of B. tabaci has relied onchemical agent for all the times. Although, the use of chemical insecticides can cause short-term(acute) effects, especially some neonicotinoid insecticides. The irrational use of chemical agentshas brought about a series of problems, including insecticide resistance increasing quickly in B.tabaci in short time. These problems cause people to reconsider the control strategy of pesticides.In attempting to solve the problems caused by resistance of B. tabaci in Fujian province, one isto study the mechanism of insecticide resistance, another is to evaluate about effect factors on B.tabaci resistance. So we can formulate rational and scientific control strategy for B. tabaciaccording with local agriculture production. For this reason, in this paper, firstly, throughmonitoring and comparing of B. tabaci resistance, the present situation and trend of insecticideresistance in field-collected populations of B. tabaci were defined in Fujian province. Secondly,the influence factors of resistance, including biotype of B. tabaci, population diversity andgenetic differentiation, host plant species, temperature and sublethal effect of pesticides, weresurveyed and analyzed to clarify the key effect factor on B. tabaci resistance in Fujian. On thebasis of these studies, we can construct the corresponding sustainable PUSH-PULL strategy andbiological control measures, and make a good foundation for resistance management of B. tabaciin Fujian province. The results were as follows:
(1) Resistance to eight insecticides in field populations of Bemisia tabaci from six regions(Fuzhou, Zhangzhou, Longyan, Sanming, Nanping, Ningde) of Fujian Province was monitoredby adult leaf-dipping bioassay. The resistance levels to neonicotinoid insecticides developedrapidly in field-collected populations of B. tabaci. Especially, Zhangzhou population, theresistance levels to imidacloprid and thiamethoxam had already developed from medium to highlevel in2005and2009(28-102.53-fold to imidacloprid,29-227.74-fold to thiamethoxam). The populations in other regions exhibited moderate levels of resistance to neonicotinoid insecticides(19.72~44.76fold to imidacloprid,21.18~51.64-fold to thiamethoxam).
(2) The biotypes of different geographical populations of Bemisia tabaci in Fujian provincewere identified base on mtDNACOI molecular marker and squash silverleaf determination. Thebiotype B and FJ were detected. In addition to the population collected from Zhangzhou, whichwas the biotype FJ, the10field-collected populations of B. tabaci were biotype B. Due to notdetecting biotype Q, the identification of biotypes showedthat the biotype B of B. tabaci was stilldestroying agricultural and production in Fujian province over the past10years, and theresistance of B. tabaci was not caused by the recent invasion of biotype Q or others.
(3) The genetic analysis of eight field-collected populations of B. tabaci were characterizedbase on the ISSR, RAPD and mtDNACOI molecular markers in Fujian province. The resultsshowedthat the genetic diversity of field-collected B. tabaci in Fujian was abundant, and acertain level of genetic differentiation has appeared among different geographical populations.Simultaneously, The results also showedthat the Fujian populations of B. tabaci own strongability of genetic evolution and adaptability under the pesticide selection pressure. The geneticdifferentiation of B. tabaci can improve its own survivability under adverse conditions.
(4) The sensitivity of B. tabaci to different pesticides was studied under different host plantsand different temperature conditions. The results showedthat the sensitivity of B. tabaci could beaffected by host plants and temperature, but this kind of impact factors was difficult to causeinsecticide resistance to increase more than several dozens or hundreds of times. We canconclude that the most important reason of resistance increasing rapidly might be due to highstrength use of neonicotinoids.
(5) The resistance screening of laboratory population was used with sublethal doses(LC20)of imidacloprid in the laboratory in2009. After14generations, the resistance of B. tabacito imidacloprid was increased by8.64-fold (LC_(20)value from10.0ppm to100.0ppm). Therealized heritability estimate(h~2) with imidacloprid screening was0.198. The results showed thatcontinuous screening with sublethal doses of imidacloprid could further increase the resistancelevel of B. tabaci, and the sublethal effect of insecticide was closely associated with resistance ofB. tabaci.
(6) Firstly, the host plants of B. tabaci and the adaptability relationship between B. tabaciand different host plants were investigated. Secondly, the control effect of trap plants (cucumber)and repellents (capsaicin) were evaluated, including biological control (Serangini japonicum).Finally, we integrated the application of trap plants(cucumber), yellow board, repellents(capsaicin) and combined with biological control technology (Serangini japonicum), to constructPUSH-PULL strategy and biological control measures for sustainable control of B. tabaci inFujian province.
引文
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