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桔小实蝇及其幼虫寄生蜂的嗅觉行为研究
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摘要
桔小实蝇Bactrocera dorsalis (Hendel)是一种毁灭性害虫,可为害果树、蔬菜和花卉等46科的250多种作物的果实。本研究采用“Y”型嗅觉仪测定了桔小实蝇及其寄生蜂---切割潜蝇茧蜂Psytallia incise (Silvestri)对不同寄主植物挥发性物质的行为反应。采用气质联用仪鉴定、对比了不同寄主植物挥发物的组成成份。测定了寄主挥发物主要成份对桔小实蝇的触角电位反应(EAG)。同时,对桔小实蝇的3种寄生蜂,即切割潜蝇茧蜂、布氏潜蝇茧蜂Fopius vandenboschi (Fullaway)和长尾潜蝇茧蜂Diachasmimorpha longicaudata (Ashmead)对主要挥发物的触角电位反应及触角显微结构进行了研究。以期为研制桔小实蝇及其寄生蜂的引诱剂,将害虫及其天敌吸引到相对小的空间,提高生物防治效果提供参考。
     (1)嗅觉仪测定结果表明,10种水果果肉对桔小实蝇两性成虫均能产生显著的引诱效果,强弱顺序为:香蕉、芭乐、木瓜、杨桃、芒果、桃、龙眼、草莓、梨和橙。对于同种水果,果肉对桔小实蝇的引诱效果明显高于同种水果叶片;成熟度高的果实引诱雌虫的数量显著高于成熟度低的果实;被桔小实蝇取食过的果肉、鲜果肉和产卵过的果肉,其引诱桔小实蝇的效果依次减弱。此外,对于同种水果,雌虫比雄虫反应强烈,未交配成虫比已交配的成虫反应强烈。
     (2)嗅觉仪测定结果也表明,切割潜蝇茧蜂雌、雄成虫对8种水果果肉均表现出行为反应,8种水果引诱效果强弱顺序为:香蕉、龙眼、芭乐、木瓜、杨桃、芒果、橙和桃。切割潜蝇茧蜂对桔小实蝇各虫态的趋性强弱顺序是:桔小实蝇幼虫、卵、蛹、成虫。该茧蜂雌虫对实蝇幼虫的趋性比雄蜂强,但对桔小实蝇其他虫态的趋性,雌雄蜂之间差异不显著。桔小实蝇取食会影响切割潜蝇茧蜂对水果的趋性,趋性强弱顺序如下:桔小实蝇幼虫取食过的水果、鲜水果、桔小实蝇成虫产过卵的水果、桔小实蝇成虫取食过的水果。从性别上看,雌雄虫反应差异不明显。未交配成虫和已交配成虫的反应差异也不显著。
     (3)本研究分析了芭乐、草莓、梨、芒果、木瓜、橙、杨桃、龙眼和桃等10种桔小实蝇寄主植物的果实和叶片的挥发性物质组成。萜烯类物质是主要的挥发性物质成份,不仅在种类上多而且含量上也高。烷烃类物质很少,酮、醛、醇、酯和酸的量都比较少。不同的寄主植物中有的还有苯、腈、胺和杂环类物质中一种或多种。
     (4)桔小实蝇对其寄主植物挥发物成份反-2-癸烯醛、香橙烯、α-法呢烯、橙花叔醇、朱栾倍半萜、紫罗兰酮、大根香叶烯、柠檬烯、里那醇和石竹烯(浓度为10μg/μL)均产生触角电位反应。雌雄虫触角最高EAG反应值均由反-2-癸烯醛引发,最低EAG反应值均由石竹烯引发。对此10种挥发物成份,桔小实蝇雄成虫交配前的触角电位反应值均显著大于交配后的。雌成虫对柠檬烯和石竹烯的触角电位反应值,则是交配前的小于交配后的;对其余8种成份,则交配前的触角电位反应值大于交配后的。交配前桔小实蝇是雄虫比雌虫的触角电位反应值大;而交配后的雌虫对香橙烯的触角电位反应值小于雄虫,而对其余9种成分,雌虫的触角电位反应值均比雄虫大。
     (5)三种寄生蜂成虫对测试的9种植物挥发物标准化合物(浓度为10μg/μL),均可引诱产生触角电位反应,且寄生蜂性别、交配前后均影响它们对各化合物的触角电位反应值。切割潜蝇茧蜂,交配前的雌雄虫对反-2-癸烯醛和香橙烯反应最强烈;交配后,雌雄虫触角最高EAG反应值均由里那醇引发,雌虫除了里那醇在交配前后差异显著外,其余的化合物在雌虫的交配前后差异不显著。切割潜蝇茧蜂雄虫除了里那醇和α-法呢烯在交配前后差异显著外,其余的化合物在雄虫的交配前后差异不显著。布氏潜蝇茧蜂交配后的雌雄成虫均对朱栾倍半萜反应最大,均对石竹烯反应最小。而交配后的长尾潜蝇茧蜂雌虫对α-法呢烯反应最大,其次是反-2-癸烯醛;雄虫对反-2-癸烯醛反应最大,其次是香橙烯和α-法呢烯。
     (6)切割潜蝇茧蜂的雌虫有4种感器,为毛形感器、板状感器、腔锥形感器和蒲姆氏鬃;雄虫除了有以上4种外,还多了种腔形感器,同时雄虫的腔锥形感觉器比雌虫的数量多。布氏潜蝇茧蜂的雌虫有5种感器,为毛形感器、板状感器、腔锥形感器、蒲姆氏鬃和腔形感器,而雄虫没有腔锥形感器,有其余4种感器。长尾潜蝇茧蜂雌雄虫的触角感器一样,是毛形感器、板状感器、腔锥形感器、蒲姆氏鬃4种。
The oriental fruit fly, Bactrocera dorsalis (Hendel), is a serious pest insect,attacking more than250species from46families of fruits and vegetables. In presentresearch, the behavior responses of the fly and its parasitoid, Psytallia incise(Silvestri), to their host volatiles were tested with Y-tube olfactometers. Volatiles fromthe tested host plants were analyzed with GC-MS for the component compounds.Afterward, the individual compounds were determined for their elicitedelectroantennogram detection (EAG) responses to antennae of B. dorsalis and its3parasitoid species, i.e. P. incise, Fopius vandenboschi (Fullaway) andDiachasmimorpha longicaudata (Ashmead). At last, the antennal sensilla wereobserved under scanning electronic microscopy (SEM) for P. incise and another twoparasitoid species of the fly, F. vandenboschi (Fullaway) and D. longicaudata(Ashmead).
     (1) Olfactometer bioassays showed that fruits of all10tested host plant speciesexhibited attraction to both sexes of B. dorsalis adults. The attraction the tested plantsreduces in the order: Musa paradisiacal, Psidium guajava, Chaenomelessinensiskoehne, Averrhoa carambola, Mangifera indica, Amygdalus persica, Dimocarpuslonggana, Fragaria ananassa, Pyrus spp and Citrus sinensis. Fruits attractedsignificantly more B. dorsalis adults than leaves. The attraction of fruits increased asthey ripening. The flies preferred the fly-eaten fruits over the fresh ones which,however, showed higher attraction than the fruits with the fly eggs oviposited in. Tothe same fruits, female adults of the fly presented higher behavior response than themales.
     (2) Olfactometer bioassays revealed that female and male adults of P. incisechose volatiles of tested fruits of8host plant species. The attraction of fruits to theparasitoids reduced in the order: Musa paradisiaca, Dimocarpus longgana, Psidiumguajava, Chaenomelessinensis Koehne, Averrhoa carambola, Mangifera indica,Citrus sinensis and Amygdalus persica. The preference of P. incise to the flyvolatiles reduced in the order of the fly life stages: lavae, eggs, pupae and adults.The female parasitoids showed stronger orientation to the fly lavae volatilesthan the males. But to the volatiles of eggs, pupae and adults of the oriental fruit fly, both sexes of the parasitoids exhibited similar preference. The attraction ofhost plants to P. incise was affected by the fly feeding, and reduced in the order:fruits eaten by the fly larvae, fresh fruits, fruits with the eggs ovipositted by the fly,and fruits eaten by the fly adults. Mated and unmated adults of the parasitoidsexhibited similar behavior reposes to the same host volatiles.
     (3) Fruit and leaf volatiles of10host plant species, i.e. Psidium guajava,Fragaria ananassa, Pyrus sp, Mangifera indica, Chaenomelessinensis koehne, Citrussinensis, Averrhoa carambola, Dimocarpus longgana, and Amygdalus persica, wereanalyzed for their component compounds. The volatiles components were dominatedby terpenes with high content and diversity. The volatiles from all tested plant speciescontained a little of alkyl compounds as well as ketones, aldehydes, ethylations,esters and acids. In volatiles of some host plant species, there are a little of benzene,cyanogens, amines and one or several heterocyclic compounds.
     (4) The oriental fruit fly produced EAG responses to10μg/μL of each(E)-2-decenal, aromadendrene, alpha-farnesene, nerolidol, valencene, ionone,germacrene, dl-Limonene, linalool and caryophyllene. The strongest EAG responsesby adult females and males of the oriental fruit fly were elicited by (E)-2-decenal, andthe lowest EAG response by Caryophyllene. To each of the tested10compounds,EAG response by unmated adult males of the fly was significantly stronger than thatby the mated males. The unmated adult females of the fly were also produced strongerEAG response than the mated females to the tested compounds, except dl-Limoneneand Caryophyllene to which the unmated females produced weaker EAG responsethan the mated. To same compounds, unmated adult males of the fly producedstronger EAG responds than the unmated females. In contrast, the mated femaleadults of the fly presented stronger EAG response than the mated males, except thatthe EAG response to Aromadendrene was stronger by mated males than by matedfemales.
     (5) The tested parasitoids of the oriental fruit fly, i.e. P. incise, F. vandenboschi,and D. longicaudata, presented EAG responses to all9tested component compoundsof the volatiles of the fly host plants. The responses were different between femaleand male adults of the parasitoids, as well as between the unmated and mated adults.For P. incise, Both of the unmated females and males produced significantly strongerEAG responses to (E)-2-decenal and aromadendrene than to other tested compounds.Mated adults of both sexed presented higher EAG responses to linalool than to other tested compounds. No significant difference of EAG responses was tested betweenthe unmated and mated females when they were exposed to the same testedcompounds, except for the EAG response to linalool. The significant difference ofEAG responses between unmated and mated males of P. incise was only tested withalpha-farnesene and linalool. For F. vandenboschi, both of the mated and unmatedadults presented strongest EAG responses to valencene among the9testedcompounds, and weakest responses to caryophyllene. Mated D. longicaudata femalesproduced the strongest EAG response to alpha-farnesene, and the second strongest to(E)-2-decenal among the tested compounds; the strongest EAG response of the maleswas elicited by (E)-2-decenal, and the second strongest response by aromadendreneand alpha-farnesene.
     (6) SEM observation of antennal sensilla revealed that P. incis possessed sensillatrichodea, sensilla placodea, sensilla ampullaceous and Bohm bristles with additionalsensilla coeloconica and more sensilla ampullaceous for the males. Parasitoid F.vandenboschi females has5types of antennal senilla, i.e. sensilla trichodea, sensillaplacodea, sensilla ampullaceous, Bohm bristles and sensilla ampullaceous. Male F.vandenboschi adults have the same senilla types as the females except that the maleslack sensilla ampullaceous. D. longicaudata adults of both sexes have4types ofantennal sensilla such as sensilla trichodea, sensilla placodea, sensilla ampullaceousand Bohm bristles.
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