绿盲蝽与秋季蒿类寄主的化学通讯机制
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摘要
1997年，我国开始商业化种植转Bt基因棉花，有效控制了棉铃虫Helicoverpa armigera(Hübner)等主要鳞翅目害虫的危害，但随之棉田化学农药使用量大幅度减少。棉田害虫地位发生了一系列演替，绿盲蝽Apolygus lucorum (Meyer-Dür)种群发生数量剧增，危害加重，成为了Bt棉花上一类重要的害虫，并波及了枣树、葡萄等其他多种作物。本文对秋季绿盲蝽成虫在不同植物上的种群消长动态及其与蒿类寄主植物的化学通讯机制进行了全面研究，主要研究结果如下：
秋季不同植物上绿盲蝽成虫的种群密度明显不同。在连续3年研究的119种植物中，秋季绿盲蝽成虫在艾蒿、野艾蒿、猪毛蒿、黄花蒿和藿香上的种群数量显著高于其他114种植物，此时这5种寄主植物均处于盛花期。进一步对菊科蒿类的艾蒿、野艾蒿和黄花蒿上绿盲蝽成虫在不同季节的寄主选择偏好行为研究发现，绿盲蝽成虫对三种蒿类寄主不同生育期的选择偏好性有明显差异，显著偏好花期的艾蒿、野艾蒿和黄花蒿，此时三种蒿类寄主上绿盲蝽成虫的种群密度和其对绿盲蝽成虫的相对吸引力均显著高于非花期。
室内行为测定中，花期的艾蒿、野艾蒿和黄花蒿对绿盲蝽成虫的吸引性均显著强于非花期植物和对照。通过GC-EAD测试结合GC-MS分析与鉴定，发现花期三种蒿类寄主植物挥发物中能引起绿盲蝽雌雄成虫触角电生理反应的活性物质均为顺-3-己烯醇、间二甲苯、丙烯酸丁酯、丙酸丁酯、丁酸丁酯和乙酸顺式-3-己烯酯。同时，Y型嗅觉仪研究表明间二甲苯、丙烯酸丁酯、丙酸丁酯和丁酸丁酯对绿盲蝽雌雄成虫具有显著的吸引作用。田间粘板和桶型诱捕器诱捕效果研究表明，间二甲苯、丙烯酸丁酯、丙酸丁酯和丁酸丁酯对绿盲蝽雌雄成虫的诱捕作用与对照相比均差异显著。此外，桶型诱捕器研究也表明，4种活性组分中的任何一种对绿盲蝽雌虫和雄虫的诱捕效果间不存在显著差异。两种诱捕方式对比研究表明桶型诱捕器更加适用于绿盲蝽的田间诱捕。
利用GC-EAD与GC-MS进一步比较分析发现，绿盲蝽早春寄主植物（枣树、葡萄、蚕豆和茼蒿）和夏季寄主植物（向日葵、葎草、罗勒和藿香）挥发物组分中分析鉴定出的绿盲蝽雌雄成虫具有电生理反应的活性物质与秋季三种蒿类寄主中分析鉴定出的活性物质基本相同。因此，秋季三种蒿类寄主中的活性物质在早春和夏季绿盲蝽寄主选择过程中同样起着关键的作用。
在田间评价的186种活性物质类似物中，共有17种对绿盲蝽成虫具有较好的诱捕效果，分别是α-法呢烯、α-葎草烯、α-蒎烯、丁酸乙酯、丙酸异丁酯、丙烯酸己酯、莰醇、3-环己烯-1-甲醇、(-)-4-萜品醇、壬醛、癸醛、3,4-二甲基苯乙酮、3-丙基甲苯、十四烷、左旋樟脑、(±)-樟脑和D-樟脑。
秋季绿盲蝽成虫在不同植物上种群消长动态的明确为解析多食性绿盲蝽的季节性寄主植物利用模式，以及构建绿盲蝽的环境友好型的综合治理策略等提供了重要的理论基础。同时，绿盲蝽与秋季三种蒿类寄主的化学通讯机制以及秋季三种蒿类寄主中的活性物质在早春和夏季绿盲蝽寄主选择过程中作用的研究为全面解析绿盲蝽与其寄主植物的互作关系，阐明绿盲蝽季节性寄主转换的内在机理，研发绿盲蝽的行为调控措施提供了重要的理论依据。
Since1997, wide-scale adoption of transgenic Bt (Bacillus thuringiensis) cotton has effectivelycontrolled the cotton bollworm Helicoverpa armigera (Hübner), and subsequent reduction of insecticidesprays in Bt cotton has caused Apolygus lucorum (Meyer-Dür)(Hemiptera: Miridae) to be major insectpest on Bt cotton in China. Additionally, this species has become a key pest of Chinese dates, grapesand other crops. In this study, we conduct comprehensive field trials on population abundance of A.lucorum adults on different plant species in autumn during2010-2012, and study the chemicalcommunication mechanism betwwen A. lucorum and its fall host plants of Artemisia spp. The mainresults were summarized as follows.
Population density of A. lucorum adults on different plant species shows obviously difference inautumn. During2010-2012, among119plant species tested, population density of A. lucorum adults onflowering Artemisia argyi Lévl. et Vant., Artemisia lavandulaefolia DC., Artemisia scoparia Waldst. etKit., Artemisia annua L. and Agastache rugosus (Fisch. et Meyer) O. kuntze. was markedly higher thanother many plant species in autumn. Further study on host preference of A. lucorum adults on threewide-distribution Artemisia spp.(A. argyi, A. lavandulaefolia, and A. annua) during different seasonsindicated that A. lucorum adults more prefer to three flowering Artemisia species over non-floweingperiods, and the average density of A. lucorum adults and standard attraction of three Artemisia speciesat flowering periods for this pest are extraordinary higher than non-flowering periods.
In Y-tube olfactometer bioassays, The results indicated that A. lucorum adults significantly morepreferred to the direction of the olfactometer where flowering plants of three Artemisia species werepresent compared to control or non-flowering plants. Coupled gaschromatography-electroantennography (GC-EAD) revealed the presence of totally sixelectrophysiologically active volatile compounds from flowering plants of three Artemisia species.These were identified by using gas chromatography-mass spectrometer (GC-MS) as (Z)-3-hexen-1-ol,m-xylene, butyl acrylate, butyl propionate, butyl butyrate and (Z)-3-hexenyl acetate. However, A.lucorum of both sexes only exhibited a significantly behavioral response to m-xylene, butyl acrylate,butyl propionate, butyl butyrate. Furthermore, the results obtained from sticky traps and bucket-shapedtraps showed that m-xylene, butyl acrylate, butyl propionate and butyl butyrate captured more A.lucorum adults than control in the field during2011-2012. Moreover, the bucket-shaped traps alsorevealed that the trap effect of each volatile was no difference in gender. In addition, the comparativestudy of two trap pattern found that bucket-shaped traps may be more appropriate for field attraction ofA. lucorum adults.
Additionally, by use of GC-EAD and GC-MS, further study indicated that, active compoundsidentified from early-season host plants (Ziziphus jujuba Mill., Vitis vinifera L., Vicia faba L. andChrysanthemum coronarium L.) and summer host plants (Helianthus annuus L., Humulus scandens(Lour.) Merr., Ocimum basilicum L. and Agastache rugosus (Fisch. et Meyer) O. kuntze.) that can elicit female and male A. lucorum adults electroantennogram (EAG) responses were basically the same asthose identified from three flowering Artemisia plant species. Therefore, active compounds emittedfrom autumn Artemisia spp. play an important role in the host location of A. lucorum for early-seasonand summer host plants.
Among186plant volatiles evaluated in the field in2011, there are totally17plant volatiles displaybetter trap effect to A. lucorum adults, that are α-farnesene, α-humulene, α-pinene, ethyl butyrate,propionic acid isobutyl ester, hexyl acrylate, borneol,3-cyclohexene-1-methanol,(-)-terpinen-4-ol,nonanal, decanal,3,4-dimethylacetophenone,3-propyltoluene, tetradecane,(1S)-(-)-camphor,(±)-camphor and (+)-camphor.
Our results provide groundwork for analysing the patterns of host plant use of polyphagous insectpest such as A. lucorum, and construction of integrated pest management of this pest. Additionally,research on chemical communication mechanism between A. lucorum and its fall host plants ofArtemisia spp., as well as function of active compounds emitted from autumn Artemisia spp. play in thehost location of A. lucorum for early-season and summer host plants shed light on overall analysing therelationship between A. lucorum and its hosts, clarification of host switching of A. lucorum duringdifferent seasons, and the development of environmentally sound and sustainable management tactics ofthis polyphagous mirid bug.
秋季不同植物上绿盲蝽成虫的种群密度明显不同。在连续3年研究的119种植物中，秋季绿盲蝽成虫在艾蒿、野艾蒿、猪毛蒿、黄花蒿和藿香上的种群数量显著高于其他114种植物，此时这5种寄主植物均处于盛花期。进一步对菊科蒿类的艾蒿、野艾蒿和黄花蒿上绿盲蝽成虫在不同季节的寄主选择偏好行为研究发现，绿盲蝽成虫对三种蒿类寄主不同生育期的选择偏好性有明显差异，显著偏好花期的艾蒿、野艾蒿和黄花蒿，此时三种蒿类寄主上绿盲蝽成虫的种群密度和其对绿盲蝽成虫的相对吸引力均显著高于非花期。
室内行为测定中，花期的艾蒿、野艾蒿和黄花蒿对绿盲蝽成虫的吸引性均显著强于非花期植物和对照。通过GC-EAD测试结合GC-MS分析与鉴定，发现花期三种蒿类寄主植物挥发物中能引起绿盲蝽雌雄成虫触角电生理反应的活性物质均为顺-3-己烯醇、间二甲苯、丙烯酸丁酯、丙酸丁酯、丁酸丁酯和乙酸顺式-3-己烯酯。同时，Y型嗅觉仪研究表明间二甲苯、丙烯酸丁酯、丙酸丁酯和丁酸丁酯对绿盲蝽雌雄成虫具有显著的吸引作用。田间粘板和桶型诱捕器诱捕效果研究表明，间二甲苯、丙烯酸丁酯、丙酸丁酯和丁酸丁酯对绿盲蝽雌雄成虫的诱捕作用与对照相比均差异显著。此外，桶型诱捕器研究也表明，4种活性组分中的任何一种对绿盲蝽雌虫和雄虫的诱捕效果间不存在显著差异。两种诱捕方式对比研究表明桶型诱捕器更加适用于绿盲蝽的田间诱捕。
利用GC-EAD与GC-MS进一步比较分析发现，绿盲蝽早春寄主植物（枣树、葡萄、蚕豆和茼蒿）和夏季寄主植物（向日葵、葎草、罗勒和藿香）挥发物组分中分析鉴定出的绿盲蝽雌雄成虫具有电生理反应的活性物质与秋季三种蒿类寄主中分析鉴定出的活性物质基本相同。因此，秋季三种蒿类寄主中的活性物质在早春和夏季绿盲蝽寄主选择过程中同样起着关键的作用。
在田间评价的186种活性物质类似物中，共有17种对绿盲蝽成虫具有较好的诱捕效果，分别是α-法呢烯、α-葎草烯、α-蒎烯、丁酸乙酯、丙酸异丁酯、丙烯酸己酯、莰醇、3-环己烯-1-甲醇、(-)-4-萜品醇、壬醛、癸醛、3,4-二甲基苯乙酮、3-丙基甲苯、十四烷、左旋樟脑、(±)-樟脑和D-樟脑。
秋季绿盲蝽成虫在不同植物上种群消长动态的明确为解析多食性绿盲蝽的季节性寄主植物利用模式，以及构建绿盲蝽的环境友好型的综合治理策略等提供了重要的理论基础。同时，绿盲蝽与秋季三种蒿类寄主的化学通讯机制以及秋季三种蒿类寄主中的活性物质在早春和夏季绿盲蝽寄主选择过程中作用的研究为全面解析绿盲蝽与其寄主植物的互作关系，阐明绿盲蝽季节性寄主转换的内在机理，研发绿盲蝽的行为调控措施提供了重要的理论依据。
Since1997, wide-scale adoption of transgenic Bt (Bacillus thuringiensis) cotton has effectivelycontrolled the cotton bollworm Helicoverpa armigera (Hübner), and subsequent reduction of insecticidesprays in Bt cotton has caused Apolygus lucorum (Meyer-Dür)(Hemiptera: Miridae) to be major insectpest on Bt cotton in China. Additionally, this species has become a key pest of Chinese dates, grapesand other crops. In this study, we conduct comprehensive field trials on population abundance of A.lucorum adults on different plant species in autumn during2010-2012, and study the chemicalcommunication mechanism betwwen A. lucorum and its fall host plants of Artemisia spp. The mainresults were summarized as follows.
Population density of A. lucorum adults on different plant species shows obviously difference inautumn. During2010-2012, among119plant species tested, population density of A. lucorum adults onflowering Artemisia argyi Lévl. et Vant., Artemisia lavandulaefolia DC., Artemisia scoparia Waldst. etKit., Artemisia annua L. and Agastache rugosus (Fisch. et Meyer) O. kuntze. was markedly higher thanother many plant species in autumn. Further study on host preference of A. lucorum adults on threewide-distribution Artemisia spp.(A. argyi, A. lavandulaefolia, and A. annua) during different seasonsindicated that A. lucorum adults more prefer to three flowering Artemisia species over non-floweingperiods, and the average density of A. lucorum adults and standard attraction of three Artemisia speciesat flowering periods for this pest are extraordinary higher than non-flowering periods.
In Y-tube olfactometer bioassays, The results indicated that A. lucorum adults significantly morepreferred to the direction of the olfactometer where flowering plants of three Artemisia species werepresent compared to control or non-flowering plants. Coupled gaschromatography-electroantennography (GC-EAD) revealed the presence of totally sixelectrophysiologically active volatile compounds from flowering plants of three Artemisia species.These were identified by using gas chromatography-mass spectrometer (GC-MS) as (Z)-3-hexen-1-ol,m-xylene, butyl acrylate, butyl propionate, butyl butyrate and (Z)-3-hexenyl acetate. However, A.lucorum of both sexes only exhibited a significantly behavioral response to m-xylene, butyl acrylate,butyl propionate, butyl butyrate. Furthermore, the results obtained from sticky traps and bucket-shapedtraps showed that m-xylene, butyl acrylate, butyl propionate and butyl butyrate captured more A.lucorum adults than control in the field during2011-2012. Moreover, the bucket-shaped traps alsorevealed that the trap effect of each volatile was no difference in gender. In addition, the comparativestudy of two trap pattern found that bucket-shaped traps may be more appropriate for field attraction ofA. lucorum adults.
Additionally, by use of GC-EAD and GC-MS, further study indicated that, active compoundsidentified from early-season host plants (Ziziphus jujuba Mill., Vitis vinifera L., Vicia faba L. andChrysanthemum coronarium L.) and summer host plants (Helianthus annuus L., Humulus scandens(Lour.) Merr., Ocimum basilicum L. and Agastache rugosus (Fisch. et Meyer) O. kuntze.) that can elicit female and male A. lucorum adults electroantennogram (EAG) responses were basically the same asthose identified from three flowering Artemisia plant species. Therefore, active compounds emittedfrom autumn Artemisia spp. play an important role in the host location of A. lucorum for early-seasonand summer host plants.
Among186plant volatiles evaluated in the field in2011, there are totally17plant volatiles displaybetter trap effect to A. lucorum adults, that are α-farnesene, α-humulene, α-pinene, ethyl butyrate,propionic acid isobutyl ester, hexyl acrylate, borneol,3-cyclohexene-1-methanol,(-)-terpinen-4-ol,nonanal, decanal,3,4-dimethylacetophenone,3-propyltoluene, tetradecane,(1S)-(-)-camphor,(±)-camphor and (+)-camphor.
Our results provide groundwork for analysing the patterns of host plant use of polyphagous insectpest such as A. lucorum, and construction of integrated pest management of this pest. Additionally,research on chemical communication mechanism between A. lucorum and its fall host plants ofArtemisia spp., as well as function of active compounds emitted from autumn Artemisia spp. play in thehost location of A. lucorum for early-season and summer host plants shed light on overall analysing therelationship between A. lucorum and its hosts, clarification of host switching of A. lucorum duringdifferent seasons, and the development of environmentally sound and sustainable management tactics ofthis polyphagous mirid bug.
引文
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