,1, 李慧敏,1,2, 操海群2, 王桂荣,1Mechanisms and Applications of Plant-Herbivore-Natural Enemy Tritrophic Interactions Mediated by Volatile Organic Compounds
WANG Bing,1, LI HuiMin,1,2, CAO HaiQun2, WANG GuiRong,1通讯作者:
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收稿日期:2020-05-27接受日期:2020-07-6网络出版日期:2021-04-16
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Received:2020-05-27Accepted:2020-07-6Online:2021-04-16
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王冰, 李慧敏, 操海群, 王桂荣. 挥发性化合物介导的植物-植食性昆虫-天敌三级营养级互作机制及应用[J]. 中国农业科学, 2021, 54(8): 1653-1672 doi:10.3864/j.issn.0578-1752.2021.08.007
WANG Bing, LI HuiMin, CAO HaiQun, WANG GuiRong.
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1980年,PRICE等[1]提出了植物-植食性昆虫-天敌昆虫三级营养级互作的理论,揭示了植物对植食性昆虫和天敌昆虫之间具有直接或间接、积极或消极等多方面影响。在错综复杂的信息网中,化学信息调节三者之间的相互作用以及相互影响,是昆虫与昆虫、昆虫与植物、甚至植物与植物之间特殊的“语言”,是农田生态系统的重要组成部分[2,3,4]。近40年来,随着传统化学生态学研究的不断深入,特别是化学分析手段和灵敏度的不断提高以及电生理研究技术的广泛渗入,新的研究理念、研究手段快速形成与发展[5,6],使得三级营养级关系之间的化学“语言”逐渐被破译,如利马豆（Phaseolus lunatus）-二斑叶螨（Tetranychus urticae）-智利小植绥螨（Phytoseiulus persimilis）[7,8];番茄（Solanum lycopersicon）-美洲棉铃虫（Heliothis zea）-短管赤眼蜂（Trichogramma pretiosum）[9]等一些研究进展极大地推动了三级营养级关系在化学生态领域中的认知。本文对挥发性化合物介导的植食性昆虫、天敌昆虫与传粉昆虫行为的影响进行了综述,对挥发物在三级营养级之间的互作机制、研究现状及其应用进行了归纳与总结,以期为昆虫行为调控产品的研发以及害虫的绿色防控提供一定的借鉴作用和理论指导。
1 化学线索调控昆虫行为选择
在健康植物的生长发育过程中,植物能够持续不断地释放大量的挥发物,植食性昆虫能够以植物挥发物为化学线索,远距离识别和定位寄主,这不仅满足了植食性昆虫自身的营养需求,而且为寻找合适的产卵场所提供了必要条件[10,11,12]。当植物受到植食性昆虫或病原菌危害和侵袭后,会形成一定的防御机制来应对各种伤害。虫害诱导的植物挥发物（herbivore- induced plant volatile,HIPV）是一种特殊的植物“语言”,可以向植食性昆虫传递“警告”信号,亦可以向天敌昆虫发出“求救”信号,同时“告诫”临近植株危险的到来[2,13-14]。HIPV主要包括萜烯类化合物和绿叶挥发物（green leaf volatile,GLV）,还包括少量的含氮、含硫化合物[15,16,17]。此外,昆虫种间和种内信息素对于同种或异种昆虫行为选择有一定的影响,尤其是植食性昆虫挥发物作为重要的化学线索影响着天敌昆虫的寄主定位。这些化学信号参与调控植物-植食性昆虫-天敌昆虫三级营养级关系之间的生态平衡。1.1 植物挥发物对植食性昆虫的影响
植物挥发物对于植食性昆虫搜寻和定位寄主、产卵以及寻找配偶等生命活动具有重要意义[10,18]。在鳞翅目昆虫中已开展了系统的研究。例如,苹果蠹蛾（Cydia pomonella）幼虫能识别成熟的西洋梨（Pyrus communis）释放的挥发物（反式）-α-法尼烯、（反式）-2-（顺式）-4-癸二烯酸甲酯、（反式）-2-（顺式）-4-癸二烯酸乙酯,并在行为上表现出明显的偏好性[19]。油菜（Brassica campestris）的挥发物异硫氰酸烯丙酯、2, 5-己二醇、异戊酸叶醇酯、（顺式）-3-甲基丁酸-3-己烯酯不仅能强烈地激活小菜蛾（Plutella xylostella）触角的EAG反应,还能引起小菜蛾的定向飞行和着陆行为[20]。有研究显示,（顺式）-茉莉酮和1-戊醇是吸引棉铃虫（Helicoverpa armigera）幼虫的关键成分[21]。“Y”型嗅觉仪试验表明,烟草植物挥发物乙酸香叶酯对烟青虫（Helicoverpa assulta）雌蛾有显著的吸引作用,橙花叔醇能显著地吸引雌、雄蛾[22]。由此可见,寄主植物挥发物在鳞翅目昆虫定位寄主的过程中发挥着关键作用。另外,一些特定寄主植物挥发物对鞘翅目、半翅目和双翅目等昆虫也具有引诱作用。例如蓝桉树（Eucalyptus globulus）植物挥发物莰烯、（+）-α-蒎烯和苯乙醇能够显著地吸引鞘翅目昆虫普来特象鼻虫（Gonipterus platensis）[23]。芸香科植物挥发物α-石竹烯、β-石竹烯和（1R）-（+）-α-蒎烯能显著地吸引半翅目昆虫柑橘木虱（Diaphorina citri）成虫[24,25]。葱属（Allium）植物和侧耳属（Pleurotus）植物共有的挥发物柠檬烯对双翅目昆虫韭菜迟眼蕈蚊（Bradysia odoriphaga）的3龄幼虫以及雌成虫具有显著的引诱作用[26]。
不同植食性昆虫对同种植物挥发物的行为反应存在着差异。以1-辛烯-3-醇对多种双翅目昆虫的作用为例,研究显示1-辛烯-3-醇对橘小实蝇（Bactrocera dorsalis）雄蝇具有驱避作用,但是对已交配的雌虫却具有显著的引诱作用[27]。另外,1-辛烯-3-醇能增加斑翅果蝇（Drosophila suzukii）的搜寻时间并降低雌蝇产卵量[28]。来源于蘑菇、三叶草以及牛呼吸气流中的1-辛烯-3-醇能够吸引按蚊属（Anopheles）和伊蚊属（Aedes）的蚊类,但是对致倦库蚊（Culex quinquefasciatus）却有显著的驱避作用[29,30]。寄主植物挥发物己醛、甲基丙基二硫醚、1-辛烯-3-醇对韭菜迟眼蕈蚊的3龄幼虫以及雌成虫具有驱避作用[26]。
在农田生态系统中,植食性昆虫往往通过对植物“化学语言”不同编码方式的识别而选择和定位寄主植物。多种化合物在环境空间中的组成决定了昆虫对植物的偏好性选择程度。例如,（顺式）-3-己烯醇、正己醇、苯甲醛、（反式）-β-法尼烯、水杨酸甲酯等15种蚕豆（Vicia faba）植株挥发物的混合物能够显著吸引蚕豆蚜（Aphis fabae）[31]。另外,具有特定比例的混合物比单一组分更能引起昆虫强烈的定向行为。研究显示成熟芒果（Mangifera indica）挥发物组分乙醇、醋酸和苯乙醇以1﹕22﹕5的比例混合比单一组分对黑腹果蝇（Drosophila melanogaster）的引诱作用更强[32]。以100﹕78﹕9比例混合的葡萄（Vitis vinifera）挥发物（反式）-β-石竹烯、4, 8-二甲基-1, 3, 7-壬三烯（DMNT）和（反式）-β-法尼烯能强烈地吸引葡萄花翅小卷蛾（Lobesia botrana）,缺少其中的任何一种都会失去引诱效果[33,34]。麦麸发酵物棕榈酸乙酯、亚油酸乙酯、亚油酸甲酯和亚油酸以10﹕24﹕6﹕0.2的比例混合对家蝇（Musca domestica）的引诱效果显著高于单一组分,并显著提高产卵量[35]。因此,具有特定比例的混合物对于鳞翅目、鞘翅目、双翅目等多种植食性昆虫的搜寻、定位、产卵等生命活动具有重要意义,可为开发害虫引诱剂提供一定的理论参考。
1.2 虫害诱导的植物挥发物（HIPV）对昆虫的影响
1.2.1 植物的防御机制以及对植食性昆虫的影响 在植物的生长周期中,经常受到大量植食性昆虫的危害。因此,植物在长期的进化过程中形成了多种高效的防御途径[36,37,38,39],包括水杨酸（salicylic acid,SA）、茉莉酸（jasmonic acid,JA）、乙烯（ethene,ETH）等植物激素调节的信号转导途径[40,41,42,43,44]。植食性昆虫取食以及产卵等行为会诱导植物产生HIPV,是植物应对外来侵害的一种防御策略[2,13-14,45]。研究表明HIPV能够增强植物对植食性昆虫的防御反应。例如,草地贪夜蛾（Spodoptera frugiperda）取食水稻（Oryza sativa）释放挥发物吲哚能诱导未被取食的组织中丝裂原活化蛋白激酶OsMPK3的转运、积累和活化,增强OsWRKY70以及多个茉莉酸合成基因的转录,而茉莉酸信号传导有利于植物防御,该研究解析了吲哚增强水稻对草地贪夜蛾防御反应的作用模式[17]。类似地,甜菜夜蛾（Spodoptera exigua）危害玉米植株释放的绿叶挥发物（顺式）-3-己烯乙酸酯也能增加玉米植株茉莉酸的合成,从而启动植物对植食性昆虫的防御[16]。另外,昆虫的挥发物也能影响植物的防御反应。例如,双翅目秋麒麟瘿蝇（Eurosta solidaginis）雄虫释放的主要挥发物混合成分螺结缩长时间处理北美一枝黄花（Solidago altissima）,诱导黄花关键防御激素茉莉酸水平的增加,从而增强植物防御反应,降低对怀卵雌虫的吸引[46]。地下环境中生存的昆虫病原线虫挥发物苯甲醛、苯甲醇、苯乙酮、壬醛和吲哚能够促进植物的防御反应并且影响植食性昆虫的行为。研究显示昆虫病原线虫挥发物能够减少地上部植食性昆虫马铃薯甲虫（Leptinotarsa decemlineata）幼虫对马铃薯叶片的取食,并且抑制马铃薯甲虫雌虫的产卵行为,这主要是由于马铃薯植株经过昆虫病原线虫挥发物的诱导后能产生更高水平的水杨酸和茉莉酸,从而使植株产生防御反应[47]。有些植食性昆虫利用HIPV操纵植物的防御反应,更适于自身的繁衍。研究显示烟粉虱（Bemisia tabaci）诱导的马铃薯植物挥发物β-月桂烯或β-石竹烯能抑制临近植株的水杨酸防御途径,使临近植株更适于烟粉虱的生长发育[48]。地下害虫红胸律点跳甲（Bikasha collaris）幼虫取食乌桕（Triadica sebifera）,释放2-乙基己醇和壬醛能吸引同种成虫取食地上部分的叶片,根部酚类次生防御物质含量降低,从而有利于幼虫的生长发育[49]。植食性昆虫识别HIPV后,“推断”寄主植物的适生情况以及是否存在竞争者,从而表现出一定的行为反馈[50]。为了提高同种后代的存活率,降低后代生存竞争,植食性昆虫往往会避免在已被危害的植物上定殖或产卵。例如,烟芽夜蛾（Heliothis virescens）雌蛾通过识别烟草（Nicotiana tabacum）在夜间释放的（顺式）-3-己烯丁酸、（顺式）-3-己烯乙酸酯等多种绿叶挥发物的混合物从而产生驱避行为,并且避免在植株上产卵[51]。类似地,海灰翅夜蛾（Spodoptera littoralis）取食棉花后,诱导的挥发物DMNT能抑制同种害虫对性信息素的选择,从而影响交配和产卵行为[52]。
VEYRAT等[53]研究表明,虫害诱导的玉米植株挥发物吲哚能提高海灰翅夜蛾幼虫的死亡率并降低其取食量,驱避雌成虫产卵,从而达到直接防御的作用。此外,虫害诱导的植物挥发物法尼烯的同分异构体（反式,反式）-α-法尼烯（49%）、（反式）-β-法尼烯（26%）、（顺式）-β-法尼烯（18%）以及（顺式,反式）-α-法尼烯（7%）能显著地抑制烟青虫雌虫的产卵行为[54]。虫害诱导的玉米植株挥发物（反式）-β-法尼烯对玉米蚜（Rhopalosiphum maidis）具有显著的驱避作用[55]。麦长管蚜（Sitobion avenae）危害小麦释放的3种诱导挥发物6-甲基-5-庚烯-2-酮、6-甲基-5-庚烯-2-醇和水杨酸甲酯对有翅蚜具有较好的驱避效果[56]。茶尺蠖（Ectropis obliqu）幼虫取食后诱导的茶树挥发物苄腈和芳樟醇对同种未交配的雌、雄成虫具有显著的驱避作用[57]。HIPV对同一生态位的异种植食性昆虫也会产生一定的影响。例如,烟芽夜蛾危害烟草植株能诱导释放大量的烟碱,对同一植株上的西花蓟马（Frankliniella occidentalis）产生忌避作用[58]。HIPV对某些植食性昆虫具有驱避作用是植物在长期的进化过程中产生的自身防御反应,以减少植食性昆虫的危害。
1.2.2 HIPV对天敌昆虫行为的影响 植株被植食性昆虫危害后能产生一系列自身防御反应,释放多种诱导挥发物作为利他素以吸引植食性昆虫的自然天敌,从而间接地调控生态系统的组成结构[2,59-61]。最早报道的有关植物利用HIPV吸引天敌的证据显示,二斑叶螨危害利马豆释放的DMNT能吸引天敌智利小植绥螨[60,62],从而形成间接防御。随着研究的深入,越来越多证据表明HIPV能够远距离地吸引植食性昆虫的天敌,尤其是寄生性天敌昆虫,研究较多的是鳞翅目昆虫危害玉米产生的HIPV对寄生蜂行为的影响。例如,甜菜夜蛾幼虫危害玉米植株释放的（顺式）-3-己烯乙酸酯、（顺式）-3-己烯醇和芳樟醇等11种虫害诱导的混合挥发物,对幼虫寄生蜂缘腹盘绒茧蜂（Cotesia marginiventris）有显著的引诱作用[63]。草地贪夜蛾取食玉米植株1—2 h后,释放的绿叶挥发物（顺式）-3-己烯醛、（反式）-2-己烯醛、（反式）-3-己烯醇和（顺式）-3-己烯乙酸酯与少量单萜芳樟醇和β-月桂烯混合显著地吸引卵寄生蜂短管赤眼蜂,而危害玉米12 h后的植株却失去了对短管赤眼蜂的引诱作用,表明该寄生蜂更易被新鲜伤口产生的挥发物所吸引[64]。此外,草地贪夜蛾危害玉米产生的α-蒎烯和α-古巴烯的混合物对卵寄生蜂岛甲腹茧蜂（Chelonus insularis）有极显著的引诱作用,是主要的活性成分[65]。由此可见,虫害诱导的玉米植株挥发物通过不同活性成分的组合模式能吸引多种寄生蜂前来定位猎物的幼虫或卵,从而发挥生物控害的效果。
HIPV对害虫的间接防御作用具有普遍性。“Y”型嗅觉仪试验显示,棉铃虫幼虫危害反枝苋（Amaranthus retroflexus）诱导产生的植物挥发物6-甲基-5-庚烯-2-酮和β-榄香烯可以显著地吸引雌性中红侧沟茧蜂（Microplitis mediator）,有利于棉铃虫绿色防控产品的开发与应用[66]。半翅目昆虫红圆蚧（Aonidiella aurantii）危害柑橘后诱导产生的D-柠檬烯和β-罗勒烯是吸引天敌昆虫印巴黄蚜小蜂（Aphytis melinus）的主要化合物,在红圆蚧的生物防治中发挥着重要作用[67]。已有的研究表明植物防御过程中产生的萜烯类化合物DMNT和TMTT是天敌昆虫重要的利他素,利用转基因的方法使水稻释放DMNT和TMTT能显著地吸引水稻害虫的天敌二化螟绒茧蜂（Cotesia chilonis）,起到对害虫的间接防御作用,为培育新型抗虫品种提供了新思路[68]。
捕食性天敌昆虫同样能够识别环境中的多种HIPV,从而快速定位猎物。有研究显示,麦长管蚜危害小麦诱导产生的（顺式）-3-己烯乙酸酯对黑带食蚜蝇（Episyrphus balteatus）具有显著的引诱作用,水杨酸甲酯能够特异性地吸引异色瓢虫（Harmonia axyridis）,这些化合物在调控麦长管蚜的种群消长中发挥着重要作用[69]。单一或混合组分的HIPV对不同种天敌昆虫的引诱效果存在差异。例如,田间试验结果表明虫害诱导挥发物罗勒烯对四条小食蚜蝇（Paragus quadrifasciatus）具有明显的引诱作用[70],嗅觉行为试验显示低浓度的单一组分水杨酸甲酯能够显著地吸引对多异瓢虫（Hippodamia variegate）[71]。单一组分（顺式）-3-己烯乙酸酯对七星瓢虫（Coccinella septempunctata）具有引诱作用[70]。而（顺式）-3-己烯乙酸酯和水杨酸甲酯的混合物比单一组分对食螨瓢虫（Stethorus punctum picipes）的引诱作用更强[72]。杂食性天敌昆虫矮小长脊盲蝽（Macrolophus pygmaeus）能够识别桃蚜（Myzus persicae）取食茄子叶片产生的（反式）-β-法尼烯和TMTT等HIPV的混合物,从而偏好性地选择有蚜虫定殖的茄子植株[73]。
植物地上部分HIPV可以吸引捕食性或寄生性天敌昆虫寻找猎物,植物地下根系分泌的化学信息素也能吸引天敌,从而定位寄主。有研究显示昆虫病原线虫能识别玉米根萤叶甲（Diabrotica virgifera virgifera）幼虫危害玉米根部释放的（反式）-β-石竹烯,对这种化学线索识别的特异性使其精准定位猎物[74]。
1.3 花的挥发物参与昆虫行为调控
植物与传粉昆虫之间的相互作用在维持大多数陆地生态系统功能完整性中发挥着重要作用[75]。大约75%的农作物种类以及88%的开花植物都需要传粉昆虫授粉从而增加作物产量[76,77,78]。反之,植物为传粉昆虫提供花粉或花蜜,传粉昆虫可以通过识别花中挥发物寻找食物来源,在帮助植物传粉的同时有利于传粉昆虫自身的生长发育与繁殖[79,80,81]。花的挥发物是吸引传粉昆虫的重要化学信号。为了研究花香的演变与昆虫化学交流的关联性,SCHIESTL[82]分析了来源于96种植物和87种昆虫中的71种最常见的花香挥发性有机化合物的发生、共性和进化模式,研究显示植物和昆虫产生的挥发性有机化合物有87%的重叠,其中芳香族化合物在被子植物中进化出信号功能,主要用于吸引传粉媒介。传粉昆虫的嗅觉系统在识别花挥发物的过程中发挥着重要作用[83,84]。花的挥发物由花的不同结构和部位散发,传粉昆虫可以通过识别挥发物差异定位花的不同部位。在无花蜜的植物中,花药是植物提供给访花昆虫的主要报偿。SOLíS-MONTERO等[81]研究表明,刺萼龙葵（Solanum rostratum）传粉型花药挥发物丁香酚、α-古巴烯和甲基丁香酚,以及取食型花药挥发物苯甲酸甲酯、苯甲酸乙酯、（反式）-α-法尼烯、γ-癸内酯和（反式）-金合欢醇均能显著地吸引东方熊蜂（Bombus impatiens）前来访花,选择行为试验表明东方熊峰更加偏好选择取食型花药。另外,研究发现刺萼龙葵取食型花药和传粉型花药的挥发物均能引起一种麦蜂（Melipona solani）的触角电位反应[81]。有研究表明菊科植物丝路蓟（Cirsium arvense）花的挥发物苯乙醛、水杨酸甲酯、2-甲氧基苯甲酸甲酯是引起黑带食蚜蝇触角电位反应的主要成分[83]。田间调查试验表明,神后鸢尾（Iris planifolia）主要的传粉昆虫为熊蜂（Bombus ruderatus）和意大利蜜蜂（Apis mellifera ligustica）,其次为黑带食蚜蝇。神后鸢尾花的挥发物以1, 4-对苯二甲醚、苯乙醇和2-甲氧基苯甲醛等芳香化合物为主,推测是吸引传粉昆虫前来取食和访花的主要化学线索[85]。此外,蜜蜂和食蚜蝇等传粉昆虫在柠檬烯释放量较高的甜叶菊（Stevia rebaudiana）上访问数量最多[86]。牡丹草属植物Gymnospermium scipetarum花的挥发物主要为烃类化合物、脂肪酸、长链的醇、醛类和少量芳香族化合物,能够吸引双翅目、膜翅目和鞘翅目昆虫[77]。另外,花挥发物也能吸引夜行性访花昆虫。比如,含有芳樟醇和（反式）-β-罗勒烯等巴西香可可（Paullinia cupana）花挥发物的混合物能吸引夜行性传粉昆虫隧蜂（Megalopta genalis）[87]。夜行性盲蝽（Neella floridula）能识别花挥发物（顺式）-茉莉酮,从而定位寄主万年青属植物Dieffenbachia aurantiaca,并将其作为取食和交配场所[88]。另外,花挥发物也能引起一些植食性昆虫的电生理反应或行为反应。例如,大叶醉鱼草（Buddleja davidii）的花挥发物苯甲醛、茶香酮、2, 2, 6-三甲基-1, 4-环己二酮、氧化异佛尔酮、（反式,反式）-α-法尼烯等能引起甘蓝尺蠖（Trichoplusia ni）的触角电位反应[89]。诱捕试验表明,花挥发的混合物β-月桂烯与苯乙醛能显著地吸引大豆尺蠖（Pseudoplusia includens）[90]。
一些植物可以通过模拟并释放昆虫的信息素组分吸引传粉昆虫,从而增加植物自身的传粉。疏花火烧兰（Epipactis veratrifolia）可以挥发一些类似于蚜虫报警信息素组分的萜烯类化合物,例如α-蒎烯、β-蒎烯、β-月桂烯和β-水芹烯等利他素,从而吸引黑带食蚜蝇前来产卵,有助于为其传粉[91,92,93]。澳大利亚兰花（Chiloglottis trapeziformis）可以释放一种与雌性黄蜂（Neozeleboria cryptoides）性信息素组分2-乙基-5-丙基-1, 3-环己二酮相同的化合物以吸引传粉昆虫授粉[94,95]。在长期进化过程中,植物发展的这种“欺骗”行为有助于自身的生存和繁衍。
开花植物释放的化合物具有动态节律性[96,97]。花的个体发育阶段以及花性别差异导致其释放的挥发物对传粉昆虫和植食性访花昆虫的引诱作用也存在一定的分化。THEIS等研究发现雌雄异株的丝路蓟雄蕊和雌蕊花的挥发物在开花初期和花期均以芳香族化合物为主,但是雄蕊花大多吸引鞘翅目金龟甲和半翅目盲蝽等植食性访花昆虫,而雌蕊花大多吸引蜜蜂和集蜂科的传粉昆虫。而雌雄同株的美洲近缘种蓟属植物Cirsium repandum同样在开花初期和花期大量释放芳香族化合物,对植食性访花昆虫和传粉昆虫均具有引诱作用,其中对凤蝶等传粉昆虫的引诱效果最好。无论是雌雄异株的丝路蓟还是雌雄同株的C. repandum在开花末期单萜的释放量显著提高,但对访花昆虫的引诱作用明显下降[98]。ZHOU等揭示了渐狭叶烟草（Nicotiana attenuata）的花在夜间释放的挥发物（反式）-α-香柠檬烯吸引烟草天蛾（Manduca sexta）成虫进行传粉,而日间烟草的叶片释放的同一化合物作为一种HIPV用于吸引烟草天蛾幼虫的天敌,这种植物挥发物释放的时空变化策略有助于解决同为传粉昆虫和植食性害虫的困境,同时为传粉昆虫和植食性昆虫对花香信号与植物防御之间形成的协同进化机制提供了遗传证据[99]。植物繁殖方式不同也影响植物对传粉昆虫的吸引。HABER等研究表明,远亲繁殖的玄参科植物猴面花（Mimulus guttatus）挥发物中的一个关键组分（反式）-β-香柑油烯的释放量显著高于近亲繁殖的量,传粉昆虫东方熊蜂能够识别这种释放量的差异对远亲繁殖的种类完成授粉过程[100]。因此,花挥发物的来源、植物繁殖类型、花的个体发育阶段与花性别差异在访花昆虫与植物互作中都发挥着重要作用。
1.4 植食性昆虫与天敌之间的化学联系
天敌昆虫可以通过植食性昆虫挥发物以及性信息素组分等化学线索寻找和定位猎物[101]。具有代表性的案例是蚜虫天敌对蚜虫蜜露挥发物和蚜虫报警信息素等化学线索的精准识别。LEROY等研究表明,巢菜修尾蚜（Megoura viciae）蜜露挥发物3-羟基-2-丁酮、3-甲基-2-丁烯醛、3-甲基-1-丁醇和柠檬烯可以显著地吸引七星瓢虫[102],这些化合物的鉴定为巢菜修尾蚜的生物防治提供了重要的化学线索。另一研究显示蚜虫报警信息素（反式）-β-法尼烯能被黑带食蚜蝇识别并诱导产卵行为[103,104]。周氏啮小蜂（Chouioia cunea）是一种寄生在美国白蛾（Hyphantria cunea）蛹中的内寄生蜂,“Y”型嗅觉仪试验表明美国白蛾蛹的挥发物1-十二烯能引起已交配的周氏啮小蜂强烈的定向选择,这一结果为揭示蛹寄生机制提供了分子依据[105]。另外,植食性昆虫释放的性信息素可以作为天敌昆虫定位寄主的重要化学线索[106,107]。例如雌性麦蛾茧蜂（Bracon hebetor）能识别雄性寄主大蜡螟（Galleria mellonella）释放的性信息素混合物壬醛和十一醛,并以此作为寻找产卵场所的直接线索[108]。ZHU等研究表明豌豆蚜（Acyrthosiphon pisum）性信息素组分（1R, 4aS, 7S, 7aR）-荆芥醇能够显著吸引金眼草蛉（Chrysopa oculata）成虫,有利于其迅速定位猎物[109,110]。由此可见,天敌昆虫对于植食性昆虫释放的化学线索的识别具有普遍性和特异性,然而目前对于识别的分子机制并不十分清楚。
2 昆虫化学感受基因参与化学线索的识别
随着转录组和基因组等测序技术的飞速发展,研究人员已经鉴定出大量的昆虫化学感受基因。主要的嗅觉基因包括气味结合蛋白（odorant binding protein,OBP）、化学感受蛋白（chemosensory protein,CSP）、气味受体（odorant receptor,OR）、离子型受体（ionotropic receptor,IR）、感觉神经元膜蛋白（sensory neuron membrane protein,SNMP）等,参与了外周嗅觉系统信号的转导[111,112]。外部气味分子通过触角极孔进入感器淋巴液,通过气味结合蛋白将其运输至神经元树突膜后,释放气味分子的同时激活气味受体,将化学信号转变为电信号并传递到更高级的神经中枢,从而指导昆虫行为[113,114]。在这个过程中,气味结合蛋白和气味受体的功能获得了广泛的研究,为解析昆虫触角外周神经系统嗅觉识别的分子机制奠定了基础。2.1 气味结合蛋白参与化学线索的识别
气味结合蛋白联系着气味受体与气味分子之间的相互作用。目前,通过转录组或者基因组测序已经鉴定了包括鳞翅目、双翅目、鞘翅目、半翅目[115,116,117,118,119,120,121]等许多种植食性昆虫的气味结合蛋白基因。其中,对在鳞翅目昆虫中较为保守的性信息素结合蛋白（pheromone binding protein,PBP）和普通气味结合蛋白（general odorant binding protein,GOBP）的功能已经开展了广泛的研究[122,123,124,125,126,127,128]。例如桃小食心虫（Carposina sasakii）的性信息素结合蛋白CsasPBP3能特异性结合两种性信息素成分（顺式）-7-二十烯-11-酮和（顺式）-7-十九烯-11-酮,而普通气味结合蛋白CsasGOBP1对这两种信息素组分结合力最强,同时也能结合植物挥发物,CsasGOBP2偏好结合植物挥发物,表明CsasPBP和CsasGOBP分别在识别性信息素和寄主植物挥发物的过程中发挥着不同的作用[129]。有研究显示,经典的气味结合蛋白也能结合性信息素组分,例如甜菜夜蛾的SexiOBP7除了可以结合苯乙酮和金合欢醇等寄主植物挥发物以外,对甜菜夜蛾的主要性信息素组分（顺式,反式）-9, 12-十四碳二烯醇醋酸酯的结合能力最好,推测SexiOBP7可能参与性信息素识别或相关行为[130]。现有的研究显示大多数昆虫气味结合蛋白均能结合寄主植物挥发物,例如鳞翅目昆虫二化螟（Chilo suppressalis）的CsupOBP8能高度结合植物挥发物β-紫罗酮、橙花叔醇、金合欢醇和2-己酮[131];半翅目昆虫柑橘木虱的DcitOBP1对β-石竹烯、α-石竹烯、α-水芹烯和（1R）-（+）-α-蒎烯等寄主植物挥发物具有高亲和力,并且对这些化合物具有强烈的行为趋向性[25];双翅目韭菜迟眼蕈蚊的BodoOBP1和BodoOBP2均能结合寄主植物挥发物二丙基三硫醚[132];鞘翅目华北大黑鳃金龟（Holotrichia oblita）的HoblOBP13和HoblOBP9分别对（反式）-2-己烯醇和苯乙醇具有高亲和力[133]。有关天敌昆虫嗅觉编码机制的研究还比较匮乏。尽管通过组学分析已经鉴定出了许多寄生性天敌以及捕食性天敌的化学感受基因,但是对于基因的功能则研究较少[3,134-142]。对鳞翅目昆虫的寄生性天敌中红侧沟茧蜂的研究发现,7个气味结合蛋白能够识别具有不同化学构象和官能团的配体,其中MmedOBP2、MmedOBP4和MmedOBP6能结合萜烯类化合物,并且后两者对含有十五碳的萜烯类化合物有较好的亲和性,推断这些气味结合蛋白对中红侧沟茧蜂搜寻和定位寄主具有一定的作用[143]。对捕食性天敌中华通草蛉（Chrysoperla sinica）气味结合蛋白功能的研究结果显示,CsinOBP1和CsinOBP10结合谱较广并且是结合萜烯类化合物的两个关键气味结合蛋白,CsinOBP1不仅能结合植物挥发物金合欢醇、己酸、（顺式）-3-己烯酯、香叶基丙酮、β-紫罗兰酮、2-十三烷酮和（反式）-橙花叔醇,还能结合蚜虫报警信息素成分（反式）-β-法尼烯,这为寄主搜寻和定位提供了直接证据[144]。
2.2 气味受体功能研究进展
气味受体是外周神经系统中接收嗅觉识别信号的关键因素,其功能鉴定对于解析嗅觉编码机制十分重要[145,146]。目前,研究人员对于鳞翅目夜蛾科昆虫性信息素识别机制的研究较为广泛,夜蛾科昆虫的性信息素受体（pheromone receptor,PR）在进化上较为保守,一些研究揭示了性信息素受体的功能在蛾类求偶交配等生理活动中发挥着重要作用。例如,棉铃虫性信息素受体HarmOR13能特异性识别性信息素主要成分（顺式）-11-十六碳烯醛,HarmOR6识别性信息素组分（顺式）-9-十六碳烯醛和（顺式）-9-十四碳烯醛,HarmOR16主要被次要组分（顺式）-11-十六碳烯醇激活[147]。随后,利用CRISPR/Cas9技术敲除雄性棉铃虫气味受体HarmOR16,发现OR16-/-突变体提前与未成熟的雌性交配,明确了棉铃虫次要性信息素成分作为性信息素拮抗剂参与调控棉铃虫的最优交配时机[148]。最近的研究显示鳞翅目近缘种棉铃虫和烟青虫的同源性信息素受体OR14b跨膜结构域上两个氨基酸位点分化决定了功能差异,阐明了性信息素受体结构与功能之间的关系[149]。这些研究为解析棉铃虫识别性信息素的外周编码和物种形成机制提供了证据。此外,对烟青虫[150,151]、烟芽夜蛾[152]、斜纹夜蛾（Spodoptera litura）[153]、甜菜夜蛾[154]以及海灰翅夜蛾[155,156]等许多夜蛾科昆虫的性信息素受体功能开展了研究并取得了一定的进展。植食性昆虫普通气味受体的激活是对寄主植物挥发物最初的信号识别与接收,参与调控昆虫的行为选择。体外功能研究显示一些寄主植物挥发物能够激活植食性昆虫的气味受体。例如斜纹夜蛾的气味受体SlituOR12能专一性识别植物挥发物（顺式）-3-己烯乙酸酯[157]。（顺式）-3-己烯乙酸酯能够激活绿盲蝽（Apolygus lucorum）气味受体AlucOR28,可能参与调控绿盲蝽趋花行为[158]。对雌性绿盲蝽具有引诱作用的植物挥发物（反式）-2-己烯醛能够激活绿盲蝽雌性触角高表达的气味受体AlucOR46[159]。另外,研究显示引起棉铃虫成虫触角EAG反应的6种结构相似的植物挥发物香叶醇、β-香茅醇、3, 7-二甲基-3-辛醇、（-）-芳樟醇、芳樟醇和（反式）-2-己烯乙酸酯均能激活鳞翅目夜蛾科3个近缘种棉铃虫、烟青虫和烟芽夜蛾的同源气味受体HarmOR12、HassOR12和HvirOR12[160]。一些研究证据显示单一气味受体的激活与植食性昆虫行为选择相关联。体内和体外功能研究表明豌豆蚜气味受体ApisOR5能被蚜虫报警信息素组分（反式）-β-法尼烯及其类似物乙酸香叶酯激活,通过RNAi技术基因沉默该受体后,豌豆蚜的跌落行为下降,证实ApisOR5通道的激活与蚜虫驱避行为相关,为筛选蚜虫驱避剂提供了理论依据[161]。对烟青虫气味受体和神经元的功能研究显示HassOR40以及短毛形感器中的B神经元能特异性识别3种结构类似的烟草挥发物乙酸香叶酯、香叶醇和橙花叔醇,并且橙花叔醇能显著地引起雌、雄虫的吸引行为,这种利用反向化学生态学理念以气味受体和神经元的功能鉴定为基础的高通量筛选可以极大地提高害虫行为调控产品的开发效率[22]。另外,鳞翅目昆虫的幼虫是主要的植食者,同样利用灵敏的嗅觉系统感受寄主植物挥发物。研究显示月桂烯、（顺式）-茉莉酮、苯乙醛和1-戊醇的混合物对棉铃虫1龄幼虫具有显著的引诱作用,其中（顺式）-茉莉酮和1-戊醇是混合物中的必要组分,可以激活在棉铃虫幼虫的同一嗅觉神经元上表达的OR41和OR52,初步阐明了棉铃虫幼虫寄主识别的神经和分子机制[21]。2020年,最新的研究发现烟青虫气味受体HassOR31在雌虫产卵器中高表达,能够感受顺-3-己烯基丁酯等12种寄主植物气味分子,明确了HassOR31参与产卵选择行为[162]。
目前,在天敌昆虫,尤其是鳞翅目幼虫寄生性天敌昆虫中有少数气味受体的功能得到鉴定。例如（顺式）-茉莉酮作为一种HIPV能特异性地激活齿唇姬蜂（Campoletis chlorideae）雌虫触角高表达的气味受体CchlOR62,选择行为试验表明该化合物能够显著地吸引交配后的齿唇姬蜂雌虫及其猎物棉铃虫幼虫,并且（顺式）-茉莉酮能显著地提高齿唇姬蜂雌虫的寄生效率,该研究初步解析了寄生蜂识别HIPV的嗅觉机制,为发展寄生蜂生物防控策略提供了理论依据[163]。另有研究显示,寄主挥发物β-石竹烯、十一烷、（反式）-α-金合欢烯、（+）-香橙烯和（顺式）-3-己烯醇能够显著引诱平腹小蜂（Anastatus japonicus）产卵。利用RNAi技术沉默气味受体AjapOR35后,平腹小蜂对β-石竹烯和（反式）-α-金合欢烯的EAG反应显著降低,并且β-石竹烯和（反式）-α-金合欢烯对平腹小峰的产卵引诱效果消失,表明AjapOR35可能参与调控平腹小蜂产卵行为[164]。天敌昆虫化学感受基因对于HIPV和猎物挥发物的识别尤为重要,是调控天敌昆虫寄主定位和产卵选择行为的重要靶标。目前对于天敌昆虫化学感受基因的功能研究还比较匮乏,是今后研究的一个重要方向。
3 应用
挥发性次生代谢物质调节植物与其他生物之间的作用一直被认为是可持续农作物保护的机遇。近年来,挥发物调节三级营养级关系已经成功地应用于农业生产[165]。目前,常见的田间防控策略包括气味诱芯的释放以调控昆虫的行为、诱导改变植物的引诱效果、培育新品种以促进挥发物的释放、间作套种（即推拉策略）等。但总的来说主要是利用各种化学信息素调节昆虫的行为。这些应用策略并不是独立起作用的,而是通过各种手段的综合应用以达到最优效果[2]。3.1 化学信息素调控昆虫行为
田间直接施用合成的挥发性昆虫行为调控产品在诱杀、驱避害虫或吸引天敌方面已经取得了成功[166,167,168]。已商品化的昆虫行为调控产品大部分是针对植食性昆虫研制而成,其中引诱产品中以鳞翅目昆虫的性信息素及其类似物居多。利用挥发性化合物吸引天敌的应用已经有了成功案例,例如含有水杨酸甲酯的混合组分可以吸引草蛉等捕食性天敌昆虫,利用水杨酸甲酯和其他引诱成分复配的引诱剂已经得到商业化推广[169]。目前,国内外对于天敌昆虫行为调控产品的开发与应用仍然具有极大的发展潜力。而驱避剂产品在市场上的占比较低,主要用于防治蚊类和蚜虫等。越来越多的证据表明,植物内源激素是植物诱导防御的重要信号物质,参与直接和间接的防御过程。例如,外施茉莉酸可以诱导番茄或水稻产生HIPV引起间接防御作用,提高对寄生蜂的吸引力,从而达到自然控害的效果[170,171]。然而这一结果在田间并没有得到广泛地证实。（顺式）-茉莉酮是一种挥发性植物激素,已在田间开展了应用研究,成功用于诱导大豆植物间接防御反应从而吸引卵寄生蜂[172]。化学遗传学筛选显示,阔叶杂草的除草剂2, 4-二氯苯氧基乙酸作为一种生长素的同系物可以极大地促进水稻对优势种卵寄生蜂的吸引[173]。
3.2 转基因作物应用
了解挥发性化合物介导的防御信号传导过程不仅可以使用标记辅助分子育种,而且可以通过识别信号的生物合成功能基因和调控基因促进育种程序。当这些信号是次生代谢物或与之相关的化合物时,转基因明显是一条更直接的途径。培育转基因新品种以驱避害虫或吸引天敌可能是进行害虫生物防治简单有效的方法之一[174,175]。蚜虫报警信息素对蚜虫的驱避效果显而易见,如果利用转基因技术从农作物中释放蚜虫报警信息素,将极大地降低蚜虫危害[176,177]。BEALE等[178]研究证实了拟南芥对桃蚜抑制以及提高寄生蜂Diaeretiella rapae搜寻能力的原理,并在十字花科植物中进行了验证。BRUCE等[179]在实验室利用转基因方法在小麦中共表达（反式）-β-法尼烯和法呢基二磷酸前体的合成酶基因与质体靶向的氨基酸序列,证实了该小麦品种显著驱蚜并且提高寄生蜂的搜寻效率,但是在两年的田间试验中却没有达到相似的结果。作物差异导致的HIPV种类变异使得对天敌昆虫的引诱作用造成分化。例如,不同玉米品系释放的HIPV存在明显的差异,导致天敌对植食性昆虫的寄生率产生一定的影响[180,181]。因此,在选择转基因植物合成靶标基因时应考虑到合成挥发性化合物是否具有特异性和共性,以提高害虫综合治理的效率。
3.3 种植方式的改变——推-拉策略
自从1990年MILLER等提出推-拉策略的理论,并将此策略应用于防治洋葱蝇（Delia antiqua）以来[182],推-拉策略得到了广泛地关注,该理论充分地将特殊植物挥发物和植物种植方式结合起来,利用间作套种的方式防治农业害虫,已在田间得到成功应用[165,183-184]。在非洲将玉米和糖蜜草（Melinis minutiflora）间作,可以有效地驱避（推）鳞翅目蛀茎类害虫,并对寄生蜂Cotesia sesamiae具有引诱作用（拉）,使得对蛀茎害虫幼虫的寄生率显著提高[185]。推-拉策略也已成功扩展到其他谷类作物[186],但需要进一步鉴定、繁殖、培育有引诱作用或者驱避作用的植物,并且植物之间地下部分的相互作用不容忽视。此外,这种策略还可以通过灵活地应用抗聚集信息素、报警信息素、产卵忌避素、拒食剂等驱避剂以及聚集信息素、性信息素、产卵刺激素等引诱剂调控昆虫行为,以达到对植食性昆虫进行绿色、高效防控的目的。另一种方法是间作对病原微生物或植食性昆虫易感的特定植物品种,诱导释放HIPV并对临近植物释放防御信号,同时能强烈地吸引有益的节肢动物。易感植物可作为有效生物防治因子的繁殖地,也可用于早期害虫的监测。4 展望
挥发性化合物在植物、植食性昆虫与天敌昆虫三级营养级关系的相互作用中具有重要的作用。本文综述了挥发物介导的三营养级间的互作机制以及国内外****在该研究领域中的贡献和重要进展,概括了部分已鉴定的重要植物挥发物对植食性昆虫、天敌昆虫以及传粉昆虫的生态调控作用（表1）。值得一提的是,我国****在昆虫化学感受基因的鉴定与功能研究方面开展了大量的工作,阐释了一些重要农业昆虫感受化学信号从而产生求偶、交配、寄主定位以及产卵等重要生理行为的生化机制,我国昆虫化学生态学研究的整体水平从研究技术与手段到研究平台的建设与利用等方面均获得了极大的提升,这为进一步开发具有自主知识产权的昆虫行为调控产品打下了基础。Table 1
表1
表1挥发性化合物在三级营养级关系中的生态调控作用
Table 1
| 化合物名称 Chemical name | 来源Source | 物种及调控作用 Species and regulation effect | 参考文献Reference | ||||
|---|---|---|---|---|---|---|---|
| 普通植物 挥发物 Plant volatile | 虫害诱导的 植物挥发物 Herbivore- induced plant volatile | 花挥发物 Flower volatile | 植食性昆虫 Herbivore | 天敌昆虫 Natural enemy | 传粉昆虫 Pollinator | ||
| (反式)-α-法尼烯、(反式)-2-(顺式)-4-癸二烯酸乙酯 (E)-α-farnesene, ethyl (2E, 4Z)-decadienoate | 西洋梨 Pyrus communis | 苹果蠹蛾 Cydia pomonella◆ | [19] | ||||
| 2, 5-己二醇、异戊酸叶醇酯 2, 5-Hexanediol, (Z)-3-hexenyl isovalerate | 油菜 Brassica campestris | 小菜蛾 Plutella xylostella◆● | [20] | ||||
| 莰烯、苯乙醇、(+)-α-蒎烯 Camphene, 2-phenylethanol, (+)-α-pinene | 蓝桉树 Eucalyptus globulus | 普来特象鼻虫 Gonipterus platensis◆ | [23] | ||||
| (1R)-(+)-α-蒎烯、α-石竹烯、β-石竹烯(1R)-(+)-α-pinene, α-caryophyllene, β- caryophyllene | 芸香科植物 Rutaceae | 柑橘木虱 Diaphorina citri◆ | [24-25] | ||||
| β-石竹烯β-caryophyllene | 马铃薯 Solanum tuberosum | 烟粉虱 Bemisia tabaci◆ | [48] | ||||
| 己醛、甲基丙基二硫醚、1-辛烯-3-醇Hexanal, methyl propyl disulfide, 1-octen- 3-ol | 葱属和侧耳属植物Allium and Pleurotus | 韭菜迟眼蕈蚊3龄幼虫和雌成虫 3rd instar larva and female adult of Bradysia odoriphaga★ | [26] | ||||
| (顺式)-3-己烯醇、正己醇、苯甲醛、(反式)-β-法尼烯、水杨酸甲酯等混合物 Mixture of (Z)-3-hexen-1-ol, 1-hexanol, benzaldehyde, (E)-β-farnesene and methyl salicylate, et al. | 蚕豆植株 Vicia faba | 蚕豆蚜 Aphis fabae◆ | [31] | ||||
| (顺式)-茉莉酮 (Z)-jasmone | 桑叶 Mulberry leaves | 棉铃虫 Helicoverpa armigera◆ | [21] | ||||
| 乙醇﹕醋酸﹕苯乙醇 Ethanol﹕acetic acid﹕2-phenylethanol (1﹕22﹕5) | 芒果 Mangifera indica | 黑腹果蝇 Drosophila melanogaster◆ | [32] | ||||
| (反式)-β-石竹烯﹕4, 8-二甲基-1, 3, 7壬三烯﹕(反式)-β-法尼烯(E)-β-caryophyllene﹕DMNT﹕(E)-β-farnesene (100﹕78﹕9) | 葡萄 Vitis vinifera | 葡萄花翅小卷蛾 Lobesia botrana◆ | [33-34] | ||||
| 化合物名称 Chemical name | 来源Source | 物种及调控作用 Species and regulation effect | 参考文献Reference | ||||
| 普通植物 挥发物 Plant volatile | 虫害诱导的 植物挥发物 Herbivore- induced plant volatile | 花挥发物 Flower volatile | 植食性昆虫 Herbivore | 天敌昆虫 Natural enemy | 传粉昆虫 Pollinator | ||
| 棕榈酸乙酯﹕亚油酸乙酯﹕亚油酸甲酯﹕亚油酸Ethyl palmitate﹕ethyl linoleate﹕methyl linoleate﹕linoleic acid (10﹕24﹕6﹕0.2) | 麦麸发酵物 Wheat bran fermentation | 家蝇 Musca domestica◆ | [35] | ||||
| (反式, 反式)-α-法尼烯、(反式)-β-法尼烯、(顺式)-β-法尼烯、(顺式, 反式)-α-法尼烯 (Z, E)-α-farnesene (49%)、(E)-β-farnesene (26%) , (Z)-β-farnesene (18%), (Z, E)-α-farnesene (7%) | 烟草 Nicotiana tabacum | 烟青虫 Helicoverpa assulta★ | [54] | ||||
| 1-辛烯-3-醇 1-Octen-3-ol | 蘑菇、三叶草等Matsutake pine mushroom, shamrock, et al. | 按蚊属和伊蚊属 Anopheles and Aedes◆;致倦库蚊Culex quinquefasciatus★ | [29-30] | ||||
| 2-乙基己醇、壬醛 2-Ethyl-1-hexanol, nonanal | 乌桕植物 Triadica sebifera | 红胸律点跳甲 Bikasha collaris ◆ | [49] | ||||
| 吲哚 Indole | 水稻 Oryza sativa | 草地贪夜蛾 Spodoptera frugiperda★ | [17] | ||||
| 玉米 Zea mays | 海灰翅夜蛾 Spodoptera littoralis★ | [53] | |||||
| (顺式)-3-己烯乙酸酯 (Z)-3-hexenyl acetate | 玉米 Z. mays | 甜菜夜蛾 Spodoptera exigua★ | [16] | ||||
| (反式)-β-法尼烯 (E)-β-farnesene | 玉米 Z. mays | 玉米蚜 Rhopalosiphum maidis★ | [55] | ||||
| 烟碱 Nicotine | 烟草植物 Nicotiana | 西花蓟马 Frankliniella occidentalis★ | [58] | ||||
| (顺式)-3-己烯丁酸、(顺式)-3-己烯乙酸酯(Z)-3-hexenyl butyrate, (Z)-3-hexenyl acetate | 烟草植物 Nicotiana | 烟芽夜蛾 Heliothis virescens★ | [51] | ||||
| (3E)-4, 8-二甲基-1, 3, 7-壬三烯 DMNT | 棉花Gossypium hirsutum | 海灰翅夜蛾 S. littoralis★ | [52] | ||||
| α-蒎烯、α-古巴烯 α-pinene, α-copaene | 玉米 Z. mays | 岛甲腹茧蜂 Chelonus insularis◆ | [65] | ||||
| (反式)-β-石竹烯 (E)-β-caryophyllene | 玉米Z. mays | 昆虫病原线虫 Heterorhabditis bacteriophora◆ | [74] | ||||
| D-柠檬烯、β-罗勒烯 D-limonene, β-ocimene | 柑橘Citrus | 印巴黄蚜小蜂 Aphytis melinus◆ | [67] | ||||
| (反式)-β-法尼烯、(E, E)-4, 8, 12-三甲基-1, 3, 7, 11-十三碳四烯 (E)-β-farnesene, TMTT | 茄子Solanum melongena | 矮小长脊盲蝽Macrolophus pygmaeus◆ | [73] | ||||
| (3E)-4, 8-二甲基-1, 3, 7-壬三烯 DMNT | 利马豆Phaseolus lunatus | 智利小植绥螨Phytoseiulus persimilis◆ | [60,62] | ||||
| (顺式)-3-己烯乙酸酯、(顺式)-3-己烯醇和芳樟醇等混合物 Mixture of (Z)-3-hexenyl acetate, (Z)-3- hexen-1-ol and linalool, et al. | 玉米 Z. mays | 缘腹盘绒茧蜂 Cotesia marginiventris◆ | [63] | ||||
| 化合物名称 Chemical name | 来源Source | 物种及调控作用 Species and regulation effect | 参考文献Reference | ||||
| 普通植物 挥发物 Plant volatile | 虫害诱导的 植物挥发物 Herbivore- induced plant volatile | 花挥发物 Flower volatile | 植食性昆虫 Herbivore | 天敌昆虫 Natural enemy | 传粉昆虫 Pollinator | ||
| (顺式)-3-己烯醛、(反式)-2-己烯醛、(反式)-3-己烯醇和(顺式)-3-己烯乙酸酯与少量单萜芳樟醇和β-月桂烯的混合物Mixture of (Z)-3-hexenal, (E)-2-hexenal, (E)-3-hexen-1-ol and (Z)-3-hexenyl acetate with a small amount of linalool and β-myrcene | 玉米 Z. mays | 短管赤眼蜂 Trichogramma pretiosum◆ | [64] | ||||
| 6-甲基-5-庚烯-2-酮、β-榄香烯 6-Methyl-5-hepten-2-one, β-elemene | 反枝苋Amaranthus retroflexus | 中红侧沟茧蜂 Microplitis mediator◆ | [66] | ||||
| (顺式)-3-己烯乙酸酯、水杨酸甲酯(Z)-3-hexenyl acetate, methyl salicylate | 小麦等植物Triticum aestivum, et al. | 黑带食蚜蝇、异色瓢虫、多异瓢虫、七星瓢虫、食螨瓢虫Episyrphus balteatus, Harmonia axyridis, Hippodamia variegate, Coccinella septempunctata, Stethorus punctum picipes◆ | [69-72] | ||||
| (反式, 反式)-α-法尼烯、苯甲醛、茶香酮、2, 2, 6-三甲基-1, 4-环己二酮、氧化异佛尔酮 (E, E)-α-farnesene, benzaldehyde, 4- oxoisophorone, 2, 2, 6-trimethylcyclohexane- 1,4-dione, oxoisophorone oxide | 大叶醉鱼草 Buddleja davidii | 甘蓝尺蠖Trichoplusia ni● | [89] | ||||
| 1, 4-对苯二甲醚、苯乙醇、2-甲氧基苯甲醛等混合物 Mixture of 1,4-dimethoxybenzene, 2- phenylethanol and 2-methoxybenzaldehyde, et al. | 神后鸢尾 Iris planifolia | 熊蜂、意大利蜜蜂、黑带食蚜蝇 Bombus ruderatus, Apis mellifera ligustica, E. balteatus◆ | [85] | ||||
| (顺式)-茉莉酮(Z)-jasmone | 万年青属植物 Dieffenbachia aurantiaca | 夜行性盲蝽Neella floridula◆ | [88] | ||||
| α-蒎烯、β-蒎烯、β-月桂烯和β-水芹烯等混合物Mixture of α-pinene, β-pinene, β-myrcene and β-phellandrene, et al. | 疏花火烧兰 Epipactis veratrifolia | 黑带食蚜蝇 E. balteatus◆ | [91-93] | ||||
| 柠檬烯Limonene | 甜叶菊 Stevia rebaudiana | 蜜蜂、黑带食蚜蝇等 A. mellifera, E. balteatus, et al.◆ | [86] | ||||
| (反式)-β-罗勒烯和芳樟醇等混合物Mixture of (E)-β-ocimene and linalool, et al. | 巴西香可可 Paullinia cupana | 隧蜂Megalopta genalis◆ | [87] | ||||
| 丁香酚、α-古巴烯、甲基丁香酚的混合物Mixture of eugenol, α-copaene and methyleugenol;苯甲酸甲酯、苯甲酸乙酯、(反式)-α-法尼烯、γ-癸内酯和(反式)-金合欢醇的混合物 Mixture of methyl benzoate, ethyl benzoate, (E)-α-farnesene, γ-decalactone and (E)-farnesol | 刺萼龙葵 Solanum rostratum | 东方熊蜂Bombus impatiens◆ | [81] | ||||
| 苯乙醛、水杨酸甲酯、2-甲氧基苯甲酸甲酯Phenylacetaldehyde, methyl salicylate, dimethyl salicylate | 丝路蓟 Cirsium arvense | 黑带食蚜蝇 E. balteatus● | [83] | ||||
| 2-乙基-5-丙基-1, 3-环己二酮 2-Ethyl-5-propylcyclohexan-1, 3- dione | 澳大利亚兰花 Chiloglottis trapeziformis | 黄蜂 Neozeleboria cryptoides◆ | [94-95] | ||||
| (反式)-β-香柑油烯 (E)-β-bergamotene | 猴面花 Mimulus guttatus | 东方熊蜂 B. impatiens● | [100] | ||||
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虽然挥发物介导植物与昆虫之间互作关系的研究有了一些突破性进展,但是目前在农业生产的应用方面还存在着一定的局限性,例如,有大量的研究表明这些化合物具有引诱或驱避植食性昆虫、吸引天敌昆虫等调节昆虫行为的作用,但如何把这些化学信息素进行整合加工进而商业化生产,并科学地应用于实际的害虫综合治理过程中还有待于进一步的系统性研究。另外,在三级营养级关系中的许多作用机理还不清楚,尤其是植物与昆虫之间的互作机制、植物挥发物与昆虫激素协同作用的分子机制、天敌昆虫化学感受分子机制等相关研究还比较匮乏。因此,需要利用多种研究手段和策略进行深入地探索与实践,以期为发展绿色、安全和高效的昆虫行为调控技术提供理论参考。
参考文献 原文顺序
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被引期刊影响因子
[本文引用: 1]
DOI:10.1146/annurev-ento-020117-043507URLPMID:29324043 [本文引用: 5]
Tritrophic interactions between plants, herbivores, and their natural enemies are an integral part of all terrestrial ecosystems. Herbivore-induced plant volatiles (HIPVs) play a key role in these interactions, as they can attract predators and parasitoids to herbivore-attacked plants. Thirty years after this discovery, the ecological importance of the phenomena is widely recognized. However, the primary function of HIPVs is still subject to much debate, as is the possibility of using these plant-produced cues in crop protection. In this review, we summarize the current knowledge on the role of HIPVs in tritrophic interactions from an ecological as well as a mechanistic perspective. This overview focuses on the main gaps in our knowledge of tritrophic interactions, and we argue that filling these gaps will greatly facilitate efforts to exploit HIPVs for pest control.
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DOI:10.1016/j.tplants.2005.04.003URLPMID:15949760 [本文引用: 2]
Locating a host plant is crucial for a phytophagous (herbivorous) insect to fulfill its nutritional requirements and to find suitable oviposition sites. Insects can locate their hosts even though the host plants are often hidden among an array of other plants. Plant volatiles play an important role in this host-location process. The recognition of a host plant by these olfactory signals could occur by using either species-specific compounds or specific ratios of ubiquitous compounds. Currently, most studies favor the second scenario, with strong evidence that plant discrimination is due to central processing of olfactory signals by the insect, rather than their initial detection. Furthermore, paired or clustered olfactory receptor neurons might enable fine-scale spatio-temporal resolution of the complex signals encountered when ubiquitous compounds are used.
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URLPMID:22616005 [本文引用: 2]
DOI:10.1186/1471-2229-14-173URLPMID:24947749 [本文引用: 2]
BACKGROUND: The induction of plant defenses in response to herbivory is well documented. In addition, many plants prime their anti-herbivore defenses following exposure to environmental cues associated with increased risk of subsequent attack, including induced volatile emissions from herbivore-damaged plant tissues. Recently, we showed in both field and laboratory settings that tall goldenrod plants (Solidago altissima) exposed to the putative sex attractant of a specialist gall-inducing fly (Eurosta solidaginis) experienced less herbivory than unexposed plants. Furthermore, we observed stronger induction of the defense phytohormone jasmonic acid in exposed plants compared to controls. These findings document a novel class of plant-insect interactions mediated by the direct perception, by plants, of insect-derived olfactory cues. However, our previous study did not exclude the possibility that the fly emission (or its residue) might also deter insect feeding via direct effects on the herbivores. RESULTS: Here we show that the E. solidaginis emission does not (directly) deter herbivore feeding on Cucurbita pepo or Symphyotrichum lateriflorum plants--which have no co-evolutionary relationship with E. solidaginis and thus are not expected to exhibit priming responses to the fly emission. We also document stronger induction of herbivore-induced plant volatiles (HIPV) in S. altissima plants given previous exposure to the fly emission relative to unexposed controls. No similar effect was observed in maize plants (Zea mays), which have no co-evolutionary relationship with E. solidaginis. CONCLUSIONS: Together with our previous findings, these results provide compelling evidence that reduced herbivory on S. altissima plants exposed to the emission of male E. solidaginis reflects an evolved plant response to olfactory cues associated with its specialist herbivore and does not involve direct effects of the fly emission on herbivore feeding behavior. We further discuss mechanisms by which the priming of HIPV responses documented here might contribute to enhanced S. altissima defense against galling.
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DOI:10.1105/tpc.18.00569URLPMID:30760558 [本文引用: 2]
Herbivore-induced plant volatiles prime plant defenses and resistance, but how they are integrated into early defense signaling and whether a causal relationship exists between volatile defense priming and herbivore resistance is unclear. Here, we investigated the impact of indole, a common herbivore-induced plant volatile and modulator of many physiological processes in plants, bacteria, and animals, on early defense signaling and herbivore resistance in rice (Oryza sativa). Rice plants infested by fall armyworm (Spodoptera frugiperda) caterpillars release indole at a rate of up to 25 ng*h(-1) Exposure to equal doses of exogenous indole enhances rice resistance to S. frugiperda Screening of early signaling components revealed that indole pre-exposure directly enhances the expression of the leucine-rich repeat-receptor-like kinase OsLRR-RLK1 Pre-exposure to indole followed by simulated herbivory increases (i.e. primes) the transcription, accumulation, and activation of the mitogen-activated protein kinase OsMPK3 and the expression of the downstream WRKY transcription factor gene OsWRKY70 as well as several jasmonate biosynthesis genes, resulting in higher jasmonic acid (JA) accumulation. Analysis of transgenic plants defective in early signaling showed that OsMPK3 is required and that OsMPK6 and OsWRKY70 contribute to indole-mediated defense priming of JA-dependent herbivore resistance. Therefore, herbivore-induced plant volatiles increase plant resistance to herbivores by positively regulating early defense signaling components.
URLPMID:29229187 [本文引用: 1]
DOI:10.1007/s001140100244URLPMID:11572015 [本文引用: 1]
The alkyl ethyl and methyl esters of (2E,4Z)-2,4-decadienoic acid found in head-space samples of ripe Bartlett pear (Pyrus communis L.) stimulated a response from neonate larvae of the codling moth (CM), Cydia pomonella (L.), in both static-air Petri-plate and in upwind Y-tube and straight-tube olfactometer bioassays. In comparison with the known CM neonate attractant, (E,E)-alpha-farnesene, ethyl (2E,4Z)-2,4-decadienoate was attractive at 10-fold and 1,000-fold lower threshold dosages in the Petri-plate and in the Y-tube bioassays, respectively. Methyl (2E,4Z)-2,4-decadienoate was attractive to CM neonates in these bioassays at much higher doses than ethyl (2E,4Z)-2,4-decadienoate. Other principal head-space volatiles from ripe pear fruit and pear leaves, including butyl acetate, hexyl acetate, (Z)-3-hexenyl acetate, and (E)-beta-ocimene, were not attractive to CM neonates. The potential uses of these pear kairomones for monitoring and control of CM in walnuts and apple are discussed.
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[本文引用: 2]
DOI:10.1016/j.ibmb.2018.04.006URLPMID:29698698 [本文引用: 2]
Plant volatiles are vital cues in the location of hosts for feeding and oviposition for Lepidoptera moths. The noctuid Helicoverpa assulta is a typical polyphagous moth, regarded as a good model for studying the olfactory reception of plant volatiles. In this study, four full-length genes encoding odorant receptors HassOR24, HassOR40, HassOR41, and HassOR55 expressed in antenna in H. assulta were functionally characterized. The highly expressed HassOR40 was narrowly tuned to a few structurally-related plant volatiles: geranyl acetate, geraniol and nerolidol. By systematically analyzing responses of single neuron in both trichoid sensilla and basiconic sensilla using single sensillum recording, the specific neuron B in one type of short trichoid sensilla was found to be mainly activated by the same chemicals as HassOR40 with high sensitivity, and with no significant difference between male and female neurons. Thus, a clear
[本文引用: 1]
DOI:10.1146/annurev-ento-120811-153542URLPMID:23317046 [本文引用: 1]
The Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Psyllidae), is the most important pest of citrus worldwide because it serves as a vector of
[本文引用: 2]
[本文引用: 2]
DOI:10.1016/j.ibmb.2018.07.002URLPMID:30026095 [本文引用: 1]
The Oriental fruit fly, Bactrocera dorsalis, is a highly destructive pest of various fruits. The reproductive and host-finding behaviors of this species are affected by several plant semiochemicals that are perceived through chemosensory receptors. However, the chemosensory mechanisms by which this perception occurs have not been fully elucidated. We conducted RNA sequencing analysis of the chemosensory organs of B. dorsalis to identify the genes coding for chemosensory receptors. We identified 60 olfactory receptors (ORs), 17 gustatory receptors and 23 ionotropic receptors-including their homologs and variants-from the transcriptome of male antennae and proboscises. We functionally analyzed ten ORs co-expressed with the obligatory co-receptor ORCO in Xenopus oocytes to identify their ligands. We tested 24 compounds including attractants for several Bactrocera species and volatiles from the host fruits of B. dorsalis. We found that BdorOR13a co-expressed with ORCO responded robustly to 1-octen-3-ol. BdorOR82a co-expressed with ORCO responded significantly to geranyl acetate, but responded weakly to farnesenes (a mixture of isomers) and linalyl acetate. These four compounds were subsequently subjected to behavioral bioassays. When each of the aforementioned compound was presented in combination with a sphere model as a visual cue to adult flies, 1-octen-3-ol, geranyl acetate, and farnesenes significantly enhanced landing behavior in mated females, but not in unmated females or males. These results suggest that the ORs characterized in the present study are involved in the perception of plant volatiles that affect host-finding behavior in B. dorsalis.
DOI:10.1093/ee/nvx125URLPMID:28981656 [本文引用: 1]
In agricultural settings, examples of effective control strategies using repellent chemicals in integrated pest management (IPM) are relatively scarce compared to those using attractants. This may be partly due to a poor understanding of how repellents affect insect behavior once they are deployed. Here we attempt to identify potential hallmarks of repellent stimuli that are robust enough for practical use in the field. We explore the literature for success stories using repellents in IPM and we investigate the mechanisms of repellency for two chemical oviposition deterrents for controlling Drosophila suzukii Matsumura, a serious pest of small fruit crops. Drosophila suzukii causes injury by laying her eggs in ripening fruit and resulting larvae make fruit unmarketable. In caged choice tests, reduced oviposition was observed in red raspberry fruit treated with volatile 1-octen-3-ol and geosmin at two initial concentrations (10% and 1%) compared to untreated controls. We used video monitoring to observe fly behavior in these caged choice tests and investigate the mode of action for deterrence through the entire behavioral repertoire leading to oviposition. We observed fewer visitors and more time elapsed before flies first landed on 1-octen-3-ol-treated fruits than control fruits and concluded that this odor primarily inhibits behaviors that occur before D. suzukii comes in contact with a potential oviposition substrate (precontact). We observed some qualitative differences in precontact behavior of flies around geosmin-treated fruits; however, we concluded that this odor primarily inhibits behaviors that occur after D. suzukii comes in contact with treated fruits (postcontact). Field trials found reduced oviposition in red raspberry treated with 1-octen-3-ol and a combination of 1-octen-3-ol and geosmin, but no effect of geosmin alone. Recommendations for further study of repellents for practical use in the field are discussed.
URLPMID:18092970 [本文引用: 1]
DOI:10.12688/f1000research.6646.1URLPMID:26543554 [本文引用: 1]
Since the discovery in the early 1980s that 1-octen-3-ol, isolated from oxen breath, attracts tsetse fly, there has been growing interest in exploring the use of this semiochemical as a possible generic lure for trapping host-seeking mosquitoes. Intriguingly, traps baited with 1-octen-3-ol captured significantly more females of the malaria mosquito, Anopheles gambiae, and the yellow fever mosquito, Aedes aegypti, than control traps, but failed to attract the southern house mosquito, Culex quinquefasciatus. Additionally, it has been demonstrated that this attractant is detected with enantioselective odorant receptors (ORs) expressed only in maxillary palps. On the basis of indoor behavioral assays it has even been suggested that 1-octen-3-ol might be a repellent to the southern house mosquito. Our approach was two-prong, i.e., to isolate 1-octen-3-ol-sensitive ORs expressed in maxillary palps and antennae of southern house female mosquito, and test the hypothesis that this semiochemical is a repellent. An OR with high transcript levels in maxillary palps, CquiOR118b, showed remarkable selectivity towards ( R)-1-octen-3-ol, whereas an OR expressed in antennae, CquiOR114b, showed higher preference for ( S)-1-octen-3-ol than its antipode. Repellency by a surface landing and feeding assay showed that not only racemic, but enantiopure ( R)- and ( S)-1-octen-3-ol are repellents at 1% dose thus suggesting the occurrence of other ( S)-1-octen-3-ol-sensitive OR(s). Female mosquitoes with ablated maxillary palps were repelled by 1-octen-3-ol, which implies that in addition to OR(s) in the maxillary palps, antennal OR(s) are essential for repellency activity.
URLPMID:18584254 [本文引用: 1]
DOI:10.1023/a:1022931816351URLPMID:12775150 [本文引用: 1]
Bioassays with a variety of overripe fruits, including mango, plum, pear, and grape, and their extracts showed that odors from overripe mango were most attractive to adult vinegar flies, Drosophila melanogaster. Combined gas chromatography-electroantennographic detection (GC-EAD) analyses of solid-phase microextraction (SPME) and Tenax extracts of overripe mango odors showed that several volatile compounds, including ethanol, acetic acid, amyl acetate, 2-phenylethanol, and phenylethyl acetate elicited significant EAG responses from antennae of female flies. Most of the volatile compounds in the extracts were identified by mass spectral and retention index comparisons with synthetic standards. In cage bioassays, lures with a blend of ethanol, acetic acid, and 2-phenylethanol in a ratio of 1:22:5 attracted six times more flies than any single EAG-active compound. This blend also attracted four times more flies than traps baited with overripe mango or unripe mango. However, in field trials, the blend was not as attractive as suggested by the laboratory bioassay.
DOI:10.1016/j.phytochem.2011.04.011URLPMID:21596403 [本文引用: 1]
Volatile plant secondary metabolites are detected by the highly sensitive olfactory system employed by insects to locate suitable plants as hosts and to avoid unsuitable hosts. Perception of these compounds depends on olfactory receptor neurones (ORNs) in sensillae, mostly on the insect antennae, which can recognise individual molecular structures. Perception of blends of plant volatiles plays a pivotal role in host recognition, non-host avoidance and ensuing behavioural responses as different responses can occur to a whole blend compared to individual components. There are emergent properties of blend perception because components of the host blend may not be recognised as host when perceived outside the context of that blend. Often there is redundancy in the composition of blends recognised as host because certain compounds can be substituted by others. Fine spatio-temporal resolution of the synchronous firing of ORNs tuned to specific compounds enables insects to pick out relevant host odour cues against high background noise and with ephemeral exposure to the volatiles at varying concentrations. This task is challenging as they usually rely on ubiquitous plant volatiles and not those taxonomically characteristic of host plants. However, such an odour coding system has the advantage of providing flexibility; it allows for adaptation to changing environments by alterations in signal processing while maintaining the same peripheral olfactory receptors.
DOI:10.1007/s00114-005-0077-7URLPMID:16450082 [本文引用: 1]
Host plant odours attract gravid insect females for oviposition. The identification of these plant volatile compounds is essential for our understanding of plant-insect relationships and contributes to plant breeding for improved resistance against insects. Chemical analysis of grape headspace and subsequent behavioural studies in the wind tunnel show that host finding in grapevine moth Lobesia botrana is encoded by a ratio-specific blend of three ubiquitous plant volatiles. The odour signal that attracts mated females to grape consists of the terpenoids (E)-beta-caryophyllene, (E)-beta-farnesene and (E)-4,8-dimethyl-1,3,7-nonatriene. These compounds represent only a fraction of the volatiles released by grapes, and they are widespread compounds known throughout the plant kingdom. Specificity may be achieved by the blend ratio, which was 100:78:9 in grape headspace. This blend elicited anemotactic behaviour in moths at remarkably small amounts. Females were attracted at release rates of only a few nanograms per minute, at levels nearly as low as those known for the attraction of male moths to the female sex pheromones.
URLPMID:27667397 [本文引用: 1]
DOI:10.1038/ni1253URLPMID:16177805 [本文引用: 1]
Although adaptive immunity is unique to vertebrates, the innate immune response seems to have ancient origins. Common features of innate immunity in vertebrates, invertebrate animals and plants include defined receptors for microbe-associated molecules, conserved mitogen-associated protein kinase signaling cascades and the production of antimicrobial peptides. It is commonly reported that these similarities in innate immunity represent a process of divergent evolution from an ancient unicellular eukaryote that pre-dated the divergence of the plant and animal kingdoms. However, at present, data suggest that the seemingly analogous regulatory modules used in plant and animal innate immunity are a consequence of convergent evolution and reflect inherent constraints on how an innate immune system can be constructed.
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DOI:10.1093/jisesa/ieu075URLPMID:25502025 [本文引用: 1]
Recent study showed that exogenous abscisic acid (ABA) acts as a regulator of plant resistance. This study investigated average injury scale and callose contents of rice, and vitellogenin (Nlvg) mRNA expression in Nilaparvata lugens (Stal) (Hemiptera: Delphacidae) adult females after third instar nymphs fed on exogenous ABA-treated susceptible [Taichung Native one (TN1)] and moderately resistant (IR42) rice cultivars. The results showed that exogenous ABA significantly decreased average injury scale of rice and Nlvg mRNA expression in N. lugens adults compared with the control (without ABA spraying). Nlvg mRNA expression in N. lugens adults decreased significantly after third instar nymphs fed on ABA-treated (5, 20, and 40 mg/liter) TN1 for 1 and 2 d, and for IR42, after fed on ABA-treated (20 and 40 mg/liter) rice plants for 1 d and after fed on ABA-treated (5, 20, and 40 mg/liter) rice for 2 d decreased significantly. The callose contents showed no significant change for TN1, while for IR42, significantly increased in roots and sheathes after N. lugens infestation under ABA treatments (20 and 40 mg/liter) compared with the control. The decrease of Nlvg mRNA expression may be partially attributed to the increase of callose content of plants. The results provide a profile for concerning the effects of ABA-induced rice plants' defenses on phloem-feeding insects.
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DOI:10.1038/s41598-018-37424-7URLPMID:30692583 [本文引用: 1]
The tea plant (Camellia sinensis) suffers heavily from a harmful piercing pest, the tea green leafhopper (TLH) Empoasca (Matsumurasca) onukii Matsuda. In the present study, we studied the effect of an efficient elicitor of plant disease resistance, the beta-1,3-glucan laminarin, on the induced defense against TLH in tea plants. Defense responses elicited by laminarin in tea include the activation of mitogen-activated protein kinases and WRKY, the burst of H2O2, salicylic acid, and abscisic acid, and the accumulation of direct-defense chemicals (including chitinase, phenylalanine ammonia lyase, callose, polyphenol oxidase, and flavonol synthase), as well as the production of volatile compounds. The laminarin-treated tea plants reduced the performance of TLH and enhanced the attractiveness to the egg parasitoid wasp of TLH, Stethynium empoascae Subba Rao. In the field experiment, laminarin application effectively reduced the number of TLH by attracting parasitoids. These results suggest that laminarin can induce protection against TLH by regulating signaling pathways in tea plant. Our study also proposes an environment friendly strategy for the integrated management of an economically important piercing pest.
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DOI:10.1038/s41598-019-45201-3URLPMID:31222054 [本文引用: 1]
Almost all previous work on host-plant selection by insect herbivores has focused on adult behaviour; however, immature life stages can also play an active role in host discrimination. The important forest pest Lymantria dispar (gypsy moth) has three recognised subspecies: the European, Asian, and Japanese gypsy moth. Unlike the other two subspecies, the European subspecies is characterised by a loss of female flight ability, which might impose a selective pressure on larvae to actively engage in host-plant selection. We therefore explored the interactions of early-instar larvae from laboratory colonies of each subspecies with four potential hosts of differing quality: oak, beech, maple, and pine-measuring larval survival and performance, feeding preferences, responses to host-derived odour cues, and the propensity to disperse from hosts via ballooning. Compared to larvae from the Asian and Japanese subspecies, larvae from the (American-originated) European gypsy moth colony exhibited (i) significantly lower survival on the poorest quality host (pine), (ii) an ability to discriminate among hosts via olfactory cues; and (iii) higher propensity to disperse from sub-optimal hosts. These results are consistent with the hypothesis that larvae from flightless female European Gypsy moth subspecies play a more active role in host-plant selection.
DOI:10.1038/35069058URLPMID:11279494 [本文引用: 1]
Plants respond to insect herbivory by synthesizing and releasing complex blends of volatile compounds, which provide important host-location cues for insects that are natural enemies of herbivores. The effects of these volatile blends on herbivore behaviour have been investigated to only a limited extent, in part because of the assumption that herbivore-induced volatile emissions occur mainly during the light phase of the photoperiod. Because many moths-whose larvae are some of the most important insect herbivores-are nocturnal, herbivore-induced plant volatiles have not hitherto been considered to be temporally available as host-location cues for ovipositing females. Here we present chemical and behavioural assays showing that tobacco plants (Nicotiana tabacum) release herbivore-induced volatiles during both night and day. Moreover, several volatile compounds are released exclusively at night and are highly repellent to female moths (Heliothis virescens). The demonstration that tobacco plants release temporally different volatile blends and that lepidopteran herbivores use induced plant signals released during the dark phase to choose sites for oviposition adds a new dimension to our understanding of the role of chemical cues in mediating tritrophic interactions.
URLPMID:26377197 [本文引用: 1]
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DOI:10.1016/j.ibmb.2018.11.006URLPMID:30468768 [本文引用: 1]
Terpenoids emitted from herbivore-damaged plants were found to play an important role in regulating tritrophic interactions. How herbivores and their natural enemies perceive terpenoids has not been thoroughly elucidated to date. Using in vivo calcium imaging, we found in this study that farnesene activates one glomerulus in the antennal lobe of female Helicoverpa assulta. The response induced by a mixture of farnesene isomers is stronger than that elicited by E-beta-farnesene alone. In the Xenopus oocyte expression system, HassOR23/ORco is narrowly tuned to farnesene isomers and compounds with similar structures. Finally, the behavioral studies showed that the farnesene isomers have an inhibitory effect on oviposition of female H. assulta, but have an attractive effect on host searching of Campoletis chlorideae, the key endoparasitoid of H. assulta larvae. These results demonstrate that farnesene isomers are encoded by a labeled-line mode in the olfactory system of female H. assulta, suggesting that farnesene as a chemical signal from plants has important behavioral relevance and evolutionary implications in the tritrophic context.
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URLPMID:25378120 [本文引用: 1]
DOI:10.1007/s10886-007-9273-6URLPMID:17415625 [本文引用: 1]
Induced plant responses to attack by chewing insects have been intensively studied, but little is known about plant responses to nonchewing insects or to attack by multiple herbivores with different feeding habits. We examined volatile emissions by tobacco, Nicotiana tabacum, in response to feeding by the piercing-sucking insect western flower thrips (WFT), Frankliniella occidentalis, the chewing herbivore Heliothis virescens, and both herbivores simultaneously. In addition, we examined the effects of herbivore-induced plant defenses on host-plant selection by WFT. Plants responded to thrips feeding by consistently releasing five compounds. Simultaneous feeding by WFT and H. virescens elicited the same 11 compounds emitted in response to caterpillar feeding alone; however, two compounds, alpha-humulene and caryophyllene oxide, were produced in greater amounts in response to simultaneous herbivory. In choice tests, thrips consistently preferred uninduced plants over all other treatments and preferred plants damaged by caterpillars and those treated with caterpillar saliva over those treated with caterpillar regurgitant. The results are consistent with a previous finding that caterpillar regurgitant induces the release of significantly more volatile nicotine than plants damaged by caterpillars or plants treated with caterpillar saliva. A repellent effect of nicotine on WFT was confirmed by encircling unwounded plants with septa releasing volatile nicotine. Our results provide the first direct evidence that thrips feeding induces volatile responses and indicates that simultaneous herbivory by insects with different feeding habits can alter volatile emissions. In addition, the findings demonstrate that induced plant responses influence host-plant selection by WFT and suggest that the induction of volatile nicotine may play a role in this process.
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URLPMID:24248958 [本文引用: 1]
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DOI:10.1007/BF01021772URLPMID:24263497 [本文引用: 1]
A volatile kairomone emitted from lima bean plants (Phaseolus lunatus) infested with the spider miteTetranychus urticae, was collected on Tenax-TA and analyzed with GC-MS. Two components were identified as the methylene monoterpene (3E)-4,8-dimethyl-1,3,7-nonatriene and the methylene sesquiterpene (3E,7E)-4,8,12-dimethyl-1,3,7,11-tridecatetraene, respectively, after purification by preparative GC on a megabore column and recording of UV, IR, and [(1)H]NMR spectra. The response of two species of predatory mites towards the identified chemicals was tested in a Y-tube olfactometer. Four of the compounds tested, linalool (3,7-dimethyl-1,6-octadien-3-ol), (E)-beta-ocimene [(3E)-3,7-dimethyl-1,3,6-octatriene], (3E)-4,8-dimethyI-1,3,7-nonatriene, and methyl salicylate attracted females ofPhytoseiulus persimilis. Linalool and methyl salicylate attracted females ofAmblyseius potentillae. The response ofA. potentillae to these two kairomone components was affected by the rearing diet of the predators in the same way as was reported for the response to the natural kairomone blend: when reared on a carotenoid-deficient diet, the predators responded to the volatile kairomone ofT. urticae, but when reared on a carotenoid-containing diet they did not. The identified kairomone components are all known from the plant kingdom. They are not known to be produced by animals de novo. In addition to biological evidence, this chemical evidence suggests that the plant is involved in production of the kairomone. Based on the present study and literature data on the response ofT. urticae to infochemicals, it is concluded that the kairomone component linalool is also a component of a volatile spider-mite dispersing pheromone.
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DOI:10.1007/s10886-011-0047-9URLPMID:22170346 [本文引用: 1]
Herbivore-induced plant volatiles are important host finding cues for larval parasitoids, and similarly, insect oviposition might elicit the release of plant volatiles functioning as host finding cues for egg parasitoids. We hypothesized that egg parasitoids also might utilize HIPVs of emerging larvae to locate plants with host eggs. We, therefore, assessed the olfactory response of two egg parasitoids, a generalist, Trichogramma pretiosum (Tricogrammatidae), and a specialist, Telenomus remus (Scelionidae) to HIPVs. We used a Y-tube olfactometer to tests the wasps' responses to volatiles released by young maize plants that were treated with regurgitant from caterpillars of the moth Spodoptera frugiperda (Noctuidae) or were directly attacked by the caterpillars. The results show that the generalist egg parasitoid Tr. pretiosum is innately attracted by volatiles from freshly-damaged plants 0-1 and 2-3 h after regurgitant treatment. During this interval, the volatile blend consisted of green leaf volatiles (GLVs) and a blend of aromatic compounds, mono- and homoterpenes, respectively. Behavioral assays with synthetic GLVs confirmed their attractiveness to Tr. pretiosum. The generalist learned the more complex volatile blends released 6-7 h after induction, which consisted mainly of sesquiterpenes. The specialist T. remus on the other hand was attracted only to volatiles emitted from fresh and old damage after associating these volatiles with oviposition. Taken together, these results strengthen the emerging pattern that egg and larval parasitoids behave in a similar way in that generalists can respond innately to HIPVs, while specialists seems to rely more on associative learning.
URLPMID:30746603 [本文引用: 1]
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DOI:10.1111/pce.12959URLPMID:28370092 [本文引用: 1]
Volatile terpenoids play a key role in plant defence against herbivory by attracting parasitic wasps. We identified seven terpene synthase genes from lima bean, Phaseolus lunatus L. following treatment with either the elicitor alamethicin or spider mites, Tetranychus cinnabarinus. Four of the genes (Pltps2, Pltps3, Pltps4 and Pltps5) were up-regulated with their derived proteins phylogenetically clustered in the TPS-g subfamily and PlTPS3 positioned at the base of this cluster. Recombinant PlTPS3 was able to convert geranyl diphosphate and farnesyl diphosphate to linalool and (E)-nerolidol, the latter being precursor of the homoterpene (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT). Recombinant PlTPS4 showed a different substrate specificity and produced linalool and (E)-nerolidol, as well as (E,E)-geranyllinalool from geranylgeranyl diphosphate. Transgenic rice expressing Pltps3 emitted significantly more (S)-linalool and DMNT than wild-type plants, whereas transgenic rice expressing Pltps4 produced (S)-linalool, DMNT and (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene (TMTT). In laboratory bioassays, female Cotesia chilonis, the natural enemy of the striped rice stemborer, Chilo suppressalis, were significantly attracted to the transgenic plants and their volatiles. We further confirmed this with synthetic blends mimicking natural rice volatile composition. Our study demonstrates that the transformation of rice to produce volatile terpenoids has the potential to enhance plant indirect defence through natural enemy recruitment.
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DOI:10.1603/0046-225X-37.6.1410URLPMID:19161683 [本文引用: 2]
Seven synthetic herbivore-induced plant volatiles (HIPVs) and a mixture of nonanal + (Z)-3-hexen-1-ol were field tested for their ability to attract beneficial insects in an open cotton field. Eleven species of the main natural enemies of insect pests in cotton fields were studied. Significantly greater numbers of the ladybird beetle Coccinella septempunctata were trapped on (Z)-3-hexenyl acetate-baited cards than on others that were HIPV baited or the control cards. Erigonidium graminicolum was attracted to traps baited with nonanal, (Z)-3-hexenyl acetate, and methyl salicylate (MeSA). The predatory bug Deraeocoris punctulatus was only attracted to traps baited with octanal. The predatory bug Orius similis responded to traps baited with 3,7-dimethyl,1,3,6-octatriene, nonanal, (Z)-3-hexenyl acetate, nonanal + (Z)-3-hexen-1-ol, and MeSA. Dimethyl octatriene, nonanal + (Z)-3-hexen-1-ol, and octanal significantly attracted the syrphid fly Paragus quadrifasciatus. The ladybird beetle Propylaea japonica, the green lacewing Chrysopa sinica, the bigeyed bug Geocoris pallidipennis, the syrphid fly Epistrophe balteata, and the parasitic wasp Campoletis chlorideae did not respond to any of the HIPVs tested. These results are discussed with regard to the potential of HIPVs as tools for recruiting natural enemies into cotton fields.
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DOI:10.1007/s10265-018-1073-2URLPMID:30456735 [本文引用: 2]
The discovery of few isolated populations of Gymnospermium scipetarum (since now considered as an amphi-Adriatic endemic) in the S-Apennines prompted to investigate, also for conservation purposes, some aspects of its reproductive biology. We aim: (1) to determine if insects play an important role in pollination; (2) to describe the pollinator community; (3) to detect floral scent composition. Experiments of insect exclusion were carried out in the field using 24 flowering individuals: one raceme was capped whereas the nearest one was used as control to ascertain differences in seed set. Pollinator community was detected during the blooming phase of two consecutive flowering seasons by visual observation; insect identification was made at the highest possible taxonomic resolution with the help of digital photographs. In order to determine the chemical composition of the volatiles, we used SPME sampling of cultivated plants. Mann-Whitney U test reveals significant differences for treatment in mean seed set with very low values for capped flowers, thus clearly indicating as insects are crucial for successful pollination. During the 42 h of observations we detected 326 visitors belonging to only three guilds: 79% were Diptera, 20% Hymenoptera and 1% Coleoptera. We identified overall 36 floral organic compounds with only two compounds common to the other studied Berberidaceae. Ambrox was never identified before in the floral scents of any angiosperm. The presence in the scent of several aldehydes and one ketone (benzophenone) could be related to the detected dominance of muscoid flies as pollinators. Floral morphology and composition of the pollinators community indicate a generalist pollination behaviour probably related to its phenology and habitat preference. The possibility of being pollinated also by muscoid flies can be considered an advantage for the reproductive fitness of the species, since these Diptera are abundant in the mountain pastures surrounding the forest habitat of Gymnospermium.
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URLPMID:30191434 [本文引用: 3]
DOI:10.1111/j.1461-0248.2010.01451.xURLPMID:20337694 [本文引用: 1]
Plants have evolved a range of strategies to manipulate the behaviour of their insect partners. One powerful strategy is to produce signals that already have a role in the animals' own communication systems. To investigate to what extent the evolution of floral scents is correlated with chemical communication in insects, I analyse the occurrence, commonness, and evolutionary patterns of the 71 most common 'floral' volatile organic compounds (VOCs) in 96 plant families and 87 insect families. I found an overlap of 87% in VOCs produced by plants and insects. 'Floral' monoterpenes showed strong positive correlation in commonness between plants (both gymnosperms and angiosperms) and herbivores, whereas the commonness of 'floral' aromatics was positively correlated between angiosperms and both pollinators and herbivores. According to a multivariate regression analysis the commonness of 'floral' aromatics was best explained by their commonness in pollinators, whereas monoterpenes were best explained by herbivores. Among pollinator orders, aromatics were significantly more common in Lepidoptera than in Hymenoptera, whereas monoterpenes showed no difference among the two orders. Collectively, these patterns suggest that plants and insects converge in overall patterns of volatile production, both for attraction and defence. Monoterpenes seem to have evolved primarily for defence under selection by herbivores, whereas aromatics evolved signalling functions in angiosperms, primarily for pollinator attraction.
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DOI:10.1016/j.phytochem.2018.11.011URLPMID:30481663 [本文引用: 1]
Iris species can adopt different pollination strategies to attract their pollinators, generalized shelter-mimicking, specialized deceptive sexual-mimicking or food-rewarding. As attractive stimuli, Iris flowers may use their colours, large-size, symmetry, and volatile organic compounds (VOCs). However, relatively few studies investigated Iris floral olfactory cues in the context of plant-visitor/pollinator interactions. In the present study we combined the identification of the floral volatiles of the nectariferous I. planifolia with insects visiting its flowers to gather data on its biology. Floral volatiles were collected in the natural environment by dynamic headspace and analysed by gas chromatography-mass spectrometry (GC-MS). Insect visitors/pollinators were also recorded. The volatile bouquet was aromatic-dominated with 1,4 dimethoxybenzene as major compound. Among the insects visiting its flowers, bumble and honey bees were the most abundant followed by hover flies. Overall, our results suggest that I. planifolia advertises its food reward by an aromatic dominated volatile composition.
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DOI:10.1007/s10886-016-0688-9URLPMID:27074793 [本文引用: 1]
Inflorescences of Araceae pollinated by cyclocephaline scarab beetles are visited frequently by a wide array of other arthropods that exploit floral resources without taking part in pollination, including earwigs, flies, and true bugs. To date, nothing is known about the cues these insect visitors use to locate the inflorescences and whether or to what extent floral scents play a role. An aroid visited by large numbers of plant bugs (Miridae) in addition to cyclocephaline scarab beetle pollinators is the Neotropical species Dieffenbachia aurantiaca. We identified the plant bug species and investigated their behavior and arrival time on the inflorescences. To test the importance of olfactory cues in locating their host we conducted experiments with open and gauze-bagged inflorescences as well as natural scent samples of D. aurantiaca. Inflorescence scents were analyzed by gas chromatography linked to mass spectrometry (GC/MS), and the attractive potential of the main scent compound was determined by behavioral assays. Three species of Neella, the most common one being N. floridula, visited the inflorescences at nightfall, shortly after the beginning of scent emission, and showed feeding and copulation activity. Bagged inflorescences as well as natural scent samples attracted similar numbers of plant bugs as the non-bagged inflorescences, showing that olfactory cues are sufficient for them to locate their host. Cis-jasmone was the major component within the inflorescence scent bouquet. In two-choice field bioassays, this compound proved to be highly attractive to Neella, and thus obviously plays a key role in finding host plants.
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DOI:10.1126/science.1087835URLPMID:14564006 [本文引用: 1]
The
DOI:10.1007/s00114-005-0636-yURLPMID:15931514 [本文引用: 1]
A standing enigma in pollination ecology is the evolution of pollinator attraction without offering reward in about one third of all orchid species. Here I review concepts of pollination by deception, and in particular recent findings in the pollination syndromes of food deception and sexual deception in orchids. Deceptive orchids mimic floral signals of rewarding plants (food deception) or mating signals of receptive females (sexual deception) to attract pollen vectors. In some food deceptive orchids, similarities in the spectral reflectance visible to the pollinator in a model plant and its mimic, and increased reproductive success of the mimic in the presence of the model have been demonstrated. Other species do not mimic specific model plants but attract pollinators with general attractive floral signals. In sexually deceptive orchids, floral odor is the key trait for pollinator attraction, and behaviorally active compounds in the orchids are identical to the sex pheromone of the pollinator species. Deceptive orchids often show high variability in floral signals, which may be maintained by negative frequency-dependent selection, since pollinators can learn and subsequently avoid common deceptive morphs more quickly than rare ones. The evolution of obligate deception in orchids seems paradoxical in the light of the typically lower fruit set than in rewarding species. Pollination by deception, however, can reduce self-pollination and encourage pollen flow over longer distances, thus promoting outbreeding. Although some food deceptive orchids are isolated through postzygotic reproductive barriers, sexually deceptive orchids lack post-mating barriers and species isolation is achieved via specific pollinator attraction. Recent population genetic and phylogenetic investigations suggest gene-flow within subgeneric clades, but pollinator-mediated selection may maintain species-specific floral traits.
DOI:10.1105/tpc.010162URLPMID:11595805 [本文引用: 1]
Emission of methyl benzoate, one of the most abundant scent compounds of bee-pollinated snapdragon flowers, occurs in a rhythmic manner, with maximum emission during the day, and coincides with the foraging activity of bumblebees. Rhythmic emission of methyl benzoate displays a
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DOI:10.1016/j.cub.2017.03.017URLPMID:28434859 [本文引用: 1]
More than 87% of flowering plant species are animal-pollinated [1] and produce floral scents and other signals to attract pollinators. These floral cues may however also attract antagonistic visitors, including herbivores [2]. The dilemma is exacerbated when adult insects pollinate the same plant that their larvae consume. It remains largely unclear how plants maximize their fitness under these circumstances. Here we show that in the night-flowering wild tobacco Nicotiana attenuata, the emission of a sesquiterpene, (E)-alpha-bergamotene, in flowers increases adult Manduca sexta moth-mediated pollination success, while the same compound in leaves is known to mediate indirect defense against M. sexta larvae [3, 4]. Forward and reverse genetic analyses demonstrated that both herbivory-induced and floral (E)-alpha-bergamotene are regulated by the expression of a monoterpene-synthase-derived sesquiterpene synthase (NaTPS38). The expression pattern of NaTPS38 also accounts for variation in (E)-alpha-bergamotene emission among natural accessions. These results highlight that differential expression of a single gene that results in tissue-specific emission of one compound contributes to resolving the dilemma for plants when their pollinators are also herbivores. Furthermore, this study provides genetic evidence that pollinators and herbivores interactively shape the evolution of floral signals and plant defense.
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DOI:10.1017/S0007485316000833URLPMID:27628497 [本文引用: 1]
In tritrophic 'plants-herbivores-natural enemies' systems, there are relatively few reports concerning the role(s) of kairomones in pupal parasitism. Chouioia cunea Yang (Hymenoptera: Eulophidae), an endoparasitic chalcid wasp, parasitizes pupae of the fall webworm (Hyphantria cunea Drury). The role of host-related kairomones was investigated using electroantennogram (EAG) and behavioral techniques. Chemicals from some host stages (pupae) and host by-products (frass), induced arrestment behavior of female parasitoids, while chemicals from prepupae, were inactive. Gas chromatography-mass spectrometry analysis of volatiles collected from pupae, frass and prepupae using solid-phase microextration revealed seven compounds with carbon chain lengths ranging from C4 to C20. All of the chemicals elicited significant EAG responses in C. cunea. Y-tube olfactometer bioassays demonstrated a significant positive response of mated female C. cunea to 1-dodecene. These data provide a better understanding of the host location mechanisms of pupal parasitoid.
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DOI:10.1023/a:1013519003944URLPMID:11871395 [本文引用: 1]
The parasitoids Trichogramma chilonis (Hymenoptera: Trichogrammatidae) and Cotesia plutellae (Hymenoptera: Braconidae), and the predator Chrysoperla carnea (Neuroptera: Chrysopidae), are potential biological control agents for the diamondback moth, Plutella xylostella (Lepidoptera: Yponomeutidae). We present studies on the interactions between these bioagents and various host-associated volatiles using a Y olfactometer. T chilonis was attracted to a synthetic pheromone blend (Z11-16:Ald, Z11-16:Ac, and Z11-16:OH in a 1:1:0.01 ratio), to Z11-16:Ac alone, and to a 1:1 blend of Z11-16:Ac and Z11-16:Ald. C. plutellae responded to the blend and to Z11-16:Ac and Z11-16:Ald. Male and female C. carnea responded to the blend and to a 1:1 blend of the major components of the pheromone, although no response was elicited by single compounds. Among the four host larval frass volatiles tested (dipropyl disulfide, dimethyl disulfide, allyl isothiocyanate, and dimethyl trisulfide), only allyl isothiocyanate elicited significant responses in the parasitoids and predator, but C. plutellae and both sexes of C. carnea did respond to all four volatiles. Among the green leaf volatiles of cabbage (Brassica oleracea subsp. capitata), only Z3-6:Ac elicited significant responses from T. chilonis, C. plutellae, and C. carnea, but C. plutellae also responded to E2-6:Ald and Z3-6:OH. When these volatiles were blended with the pheromone, the responses were similar to those elicited by the pheromone alone, except for C. carnea males, which had an increased response. The effect of temperature on the response of the biological agents to a mixture of the pheromone blend and Z3-6:Ac was also studied. T. chilonis was attracted at temperatures of 25-35 degrees C, while C. plutellae and C. carnea responded optimally at 30-35 degrees C and 20-25 degrees C, respectively. These results indicate that the sex pheromone and larval frass volatiles from the diamondback moth, as well as volatile compounds from cabbage, may be used by these natural enemies to locate their diamondback moth host.
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DOI:10.1007/s00114-005-0624-2URLPMID:15812573 [本文引用: 1]
It is well documented that host-related odors enable many species of parasitoids and predatory insects to locate their prey and prey habitats. This study reports the first characterization of prey and prey host odor reception in two species of lacewings, Chrysoperla carnea (Say) and Chrysopa oculata L. 2-Phenylethanol, one of the volatiles emitted from their prey's host plants (alfalfa and corn) evoked a significant EAG response from antennae of C. carnea. Traps baited with this compound attracted high numbers of adult C. carnea, which were predominantly females. One of the sex pheromone components (1R,4aS,7S,7aR)-nepetalactol of an aphid species, Acyrthosiphon pisum (Harris) attracted only C. oculata adults. Single sensillum recordings showed that the olfactory neurons of C. carnea responded to both 2-phenylethanol and aphid sex pheromone components, but those of C. oculata only responded to the latter.
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DOI:10.1146/annurev-ento-120811-153635URLPMID:23020622 [本文引用: 1]
Our knowledge of the molecular basis of odorant reception in insects has grown exponentially over the past decade. Odorant receptors (ORs) from moths, fruit flies, mosquitoes, and the honey bees have been deorphanized, odorant-degrading enzymes (ODEs) have been isolated, and the functions of odorant-binding proteins (OBPs) have been unveiled. OBPs contribute to the sensitivity of the olfactory system by transporting odorants through the sensillar lymph, but there are competing hypotheses on how they act at the end of the journey. A few ODEs that have been demonstrated to degrade odorants rapidly may act in signal inactivation alone or in combination with other molecular traps. Although ORs in Drosophila melanogaster respond to multiple odorants and seem to work in combinatorial code involving both periphery and antennal lobes, reception of sex pheromones by moth ORs suggests that their labeled lines rely heavily on selectivity at the periphery.
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DOI:10.1016/j.cois.2014.10.006URLPMID:25584200 [本文引用: 1]
Olfactory signaling is a crucial component in the life history of insects. The development of precise and parallel mechanisms to analyze the tremendous amount of chemical information from the environment and other sources has been essential to their evolutionary success. Considerable progress has been made in the study of insect olfaction fueled by bioinformatics- based utilization of genomics along with rapid advances in functional analyses. Here we review recent progress in our rapidly emerging understanding of insect peripheral sensory reception and signal transduction. These studies reveal that the nearly unlimited chemical space insects encounter is covered by distinct chemosensory receptor repertoires that are generally derived by species-specific, rapid gene gain and loss, reflecting the evolutionary consequences of adaptation to meet their specific biological needs. While diverse molecular mechanisms have been put forth, often in the context of controversial models, the characterization of the ubiquitous, highly conserved and insect-specific Orco odorant receptor co-receptor has opened the door to the design and development of novel insect control methods to target agricultural pests, disease vectors and even nuisance insects.
DOI:10.1038/s41598-017-11646-7URLPMID:28931846 [本文引用: 1]
Perception of environmental and habitat cues is of significance for insect survival and reproduction. Odor detection in insects is mediated by a number of proteins in antennae such as odorant receptors (ORs), ionotropic receptors (IRs), odorant binding proteins (OBPs), chemosensory proteins (CSPs), sensory neuron membrane proteins (SNMPs) and odorant degrading enzymes. In this study, we sequenced and assembled the adult male and female antennal transcriptomes of a destructive agricultural pest, the diamondback moth Plutella xyllostella. In these transcriptomes, we identified transcripts belonging to 6 chemoreception gene families related to ordor detection, including 54 ORs, 16 IRs, 7 gustatory receptors (GRs), 15 CSPs, 24 OBPs and 2 SNMPs. Semi-quantitative reverse transcription PCR analysis of expression patterns indicated that some of these ORs and IRs have clear sex-biased and tissue-specific expression patterns. Our results lay the foundation for future characterization of the functions of these P. xyllostella chemosensory receptors at the molecular level and development of novel semiochemicals for integrated control of this agricultural pest.
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DOI:10.1016/j.bbrc.2018.12.007URLPMID:30545637 [本文引用: 1]
The peach fruit borers Carposina sasakii Matsumura (Lepidoptera: Carposinidae) are serious fruit borers in East Asia. Odorants have been reported to elicit their olfactory responses, including sex pheromone and host plant volatiles. However, little is known about their molecular aspects of olfactory perception to these odorants. Odorant binding proteins (OBPs) are believed to play an important role in insect olfaction, which recognize and bind odorants, and transport them to odorant receptors located on olfactory neurons. We identified 12 putative OBPs (CsasOBP1-12) expressed in intact heads (with antennae and mouthparts) of C. sasakii adult males and females by transcriptome analysis, including 4 putative pheromone binding proteins (PBPs) (CsasOBP1-2, 7, 11) and 4 putative general odorant binding proteins (GOBPs) (CsasOBP6, 9-10, 12). A phylogenetic tree was constructed to characterize these OBPs. An analysis using fluorescence quantitative PCR showed that CsasOBP3, 5, 7, 9-12 have the highest expression level in the intact heads, suggesting that among them there may be the OBPs playing key roles in C. sasakii olfaction. In the 7 OBPs, CsasOBP5, 7, 11-12 have higher level of expressions in intact heads of C. sasakii adult males than females, while CsasOBP3, 9-10 are expressed equally in the two sexes. Moreover, CsasOBP2 is expressed dominantly in the intact heads and wings, with an equal level between them. CsasOBP8 expression is highest in the wings, while CsasOBP1, 4, 6 are expressed dominantly in the abdomens. Our study is helpful for understanding C. sasakii OBPs' functions and C. sasakii olfaction from molecular view.
DOI:10.1016/j.ibmb.2019.01.005URLPMID:30654011 [本文引用: 1]
We examined the genome of the soybean aphid, Aphis glycines, and an updated genome assembly of the pea aphid, Acyrthosiphon pisum, for members of the three major families of chemoreceptors, the Odorant Receptors (ORs), Gustatory Receptors (GRs) and Ionotropic Receptors (IRs), as well as the Odorant Binding Proteins (OBPs). The soybean aphid has 47 ORs, 61GRs, 19 IRs, and 10 OBPs, compared with 87 ORs, 78 Grs, 19 IRs, and 18 OBPs in the pea aphid, with variable numbers of pseudogenes in the OR and GR families. Phylogenetic analysis reveals that while all of the IRs are simple orthologs between these two species, the OR, GR, and OBP families in the pea aphid have experienced major expansions of particular gene lineages and fewer losses of gene lineages. This imbalance in birth-and-death of chemosensory genes has led to the larger pea aphid gene repertoire, which might be related to the broader host range of pea aphids versus the specialization of soybean aphids on a single summer host plant. Examination of the expression levels of these chemosensory genes in parthenogenetic and sexual females and males of pea aphids revealed multiple genes that are differentially expressed in sexual females or males and might be involved in reproductive biology. Examination of the soybean aphid genes in parthenogenetic females under multiple stressors revealed multiple genes whose expression levels changed with heat or starvation stress, the latter potentially important in finding new food sources.
DOI:10.1002/arch.21542URLPMID:30820994 [本文引用: 1]
Our bioassays reviewed that antennae played crucial roles in the responses of maize weevil (Sitophilus zeamais) to food and sex volatiles. In order to identify the maize weevil odorant-binding protein (OBP) genes, we analyzed its antennal transcriptome. In total, 21,587,928 high-quality clean reads were obtained from RNA-seq, 52,206 unigenes were assembled, and 25,744 unigenes showed significant similarity ( E value < 10 (-5) ) to known proteins in the NCBI nonredundant protein database. From those unigenes, we identified 41 candidate OBP proteins, which could be categorized into dimeric OBPs subfamily, minus-C OBPs subfamily, and classical OBPs subfamily. Phylogenic analysis indicated that most maize weevil OBPs were closely related to their orthologues in other beetles of the Superfamily Curculionoidea. We further investigated the expression profiles of those candidate OBP genes by quantitative real-time polymerase chain reaction. Twenty-six of forty-one maize weevil OBP genes were highly expressed in the antennae or other parts of the head. The rest were expressed in the legs, wings, or other tested tissues. The antennal transcriptomic data and candidate OBP genes described here provide a basis for the functional studies of the maize weevil chemical perception, which are potential novel targets for pest control strategies.
DOI:10.1111/imb.12513URLPMID:29888835 [本文引用: 1]
Odorant binding proteins (OBPs) and chemosensory proteins (CSPs) play essential roles in insect chemosensory recognition. Here, we identified nine OBPs and nine CSPs from the Myzus persicae transcriptome and genome. Genomic structure analysis showed that the number and length of the introns are much higher, and this appears to be a unique feature of aphid OBP genes. Three M. persicae OBP genes (OBP3/7/8) as well as CSP1/4/6, CSP2/9 and CSP5/8 are tandem arrayed in the genome. Phylogenetic analyses of five different aphid species suggest that aphid OBPs and CSPs are conserved in single copy across all aphids (with occasional losses), indicating that each OBP and CSP class evolved from a single gene in the common ancestor of aphids without subsequent duplication. Motif pattern analysis revealed that aphid OBP and CSP motifs are highly conserved, and this could suggest the conserved functions of aphid OBPs and CSPs. Three OBPs (MperOBP6/7/10) are expressed antennae specifically, and five OBPs (MperOBP2/4/5/8/9) are expressed antennae enriched, consistent with their putative olfactory roles. M. persicae CSPs showed much broader expression profiles in nonsensory organs than OBPs. None of the nine MperCSPs were found to be antennae specific, but five of them (MperCSP1/2/4/5/6) showed higher expression levels in the legs than in other tissues. MperCSP10 mainly expressed in the antennae and legs. The broad and diverse expression patterns of M. persicae CSPs suggest their multifunctions in olfactory perception, development and other processes.
DOI:10.1111/1744-7917.12576URLPMID:29441682 [本文引用: 1]
Odorant-binding proteins (OBPs) and chemosensory proteins (CSPs) of insects are thought to play roles in olfactory recognition affecting host choice, copulation, reproduction and other behaviors. Previous descriptions of OBPs and CSPs in the whitefly Bemisia tabaci often provided no or incomplete genetic information. In this study, we present a genome-wide and transcriptome-wide investigation of the OBPs and CSPs in B. tabaci MEAM1 (Middle East-Asia Minor1 species). Eight OBP and 19 CSP genes were identified that covered all previous sequences. Phylogenetic analyses showed that the CSP genes had a lineage-specific expansion (BtabBCSP1, BtabBCSP3, BtabBCSP13, BtabBCSP17, BtabBCSP18 and BtabBCSP19). Expression profiling of OBPs and CSPs by transcriptome sequencing and quantitative real-time polymerase chain reaction (qPCR) revealed that expression patterns differed among developmental stages of B. tabaci MEAM1. Five OBP genes and 11 CSP genes significantly differed between males and females; four of the 19 CSP genes were highly expressed in adults, while two were highly expressed in nymphs. The expression profiles of the OBP and CSP genes in different tissues of B. tabaci MEAM1 adults were analyzed by qPCR. Four OBP genes found in B. tabaci MEAM1 were highly expressed in the head. Conversely, only two CSPs were enriched in the head, while the other six CSPs were specifically expressed in other tissues. Our results provide a foundation for future research on OBPs and CSPs in B. tabaci.
DOI:10.1007/s10886-007-9277-2URLPMID:17393279 [本文引用: 1]
Pheromone binding proteins (PBP) play an important role in insect pheromone communication. However, the PBP for the beet armyworm, Spodoptera exigua Hubner (Lepidoptera: Noctuidae), an important agricultural pest worldwide, remains unaddressed. We report the cloning of two PBP genes, SexigPBP1 and SexigPBP2, from the antennal cDNA of S. exigua by reverse transcriptase-polymerase chain reaction (RT-PCR) and rapid amplification of cDNA ends-PCR (RACE-PCR). The deduced PBP amino acid sequences are characteristic of the odorant binding protein (OBP) family, although the two PBPs are only 44% identical. From an analysis of the genomic DNA, two introns and a similar intron/extron structural pattern were identified in each of the two PBP genes. RT-PCR analysis revealed that the two PBP genes are only expressed in antennae. Real-time PCR further indicated that the expression of SexigPBP1 is much higher than that of SexigPBP2, regardless of sex. However, the female expression levels for SexigPBP1 and SexigPBP2 are about 39% and 73%, respectively, relative to male levels. Finally, phylogenetic analysis suggested that PBPs from the Noctuidae are divided into three distinct groups based on the primary sequences.
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DOI:10.1016/j.ibmb.2016.03.005URLPMID:27001069 [本文引用: 1]
Olfaction of Lepidopteran larvae has received little attention, compared to the damage to crops done by insects at this stage. We report that larvae of the diamondback moth Plutella xylostella are attracted to their natural sex pheromone and to their major component (Z)-11-hexadecenal, but only in a food context. For such task they use two general odorant-binding proteins (GOBPs), abundantly expressed in the three major sensilla basiconica of the larval antenna, as shown by whole-mount immunostaining and immunocytochemistry experiments. None of the three genes encoding pheromone-binding proteins (PBPs) are expressed at this stage. Both recombinant GOBPs bind (Z)-11-hexadecenal and the corresponding alcohol, but not the acetate. Binding experiments performed with five mutants of GOBP2, where aromatic residues in the binding pocket were replaced with leucine showed that only one or two amino acid substitutions can completely abolish binding to the pheromone shifting the affinity to plant-derived compounds. We hypothesise that detection of their species-specific pheromone may direct larvae to the sites of foraging chosen by their mother when laying eggs, to find better food, as well as to reduce competition with individuals of the same or other species sharing the same host plant. We also provide evidence that GOBP2 is a narrowly tuned binding protein, whose affinity can be easily switched from linear pheromones to branched plants terpenoids, representing a tool better suited for the simple olfactory system of larvae, as compared to the more sophisticated organ of adults.
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DOI:10.1016/j.pestbp.2019.04.011URLPMID:31378363 [本文引用: 1]
Sex pheromones are crucial for communication between females and males in moths, and pheromone receptors (PRs) play a key role in peripheral coding of sex pheromones. During the last decade, many PR candidates have been identified based on transcriptome sequencing and bioinformatic analysis, but their detailed functions remain mostly unknown. Here, focusing on four PR candidates of Athetis dissimilis (AdisOR1, AdisOR6, AdisOR11 and AdisOR14) identified in a previous study, we first cloned the full-length cDNAs and determined the tissue expression profiles by quantitative real-time PCR (qPCR). The results revealed that expression of three of these genes were male antennae-specific, while AdisOR11 was similar in expression between male and female antennae. Furthermore, the expression level of AdisOR1 was much higher than those of the other three genes. Then, functional analysis was conducted using Xenopus oocyte system. AdisOR1 responded strongly to the sex pheromone component Z9-14:OH and the potential pheromone component Z9,E12-14:OH, suggesting its important role in the sex pheromone perception; AdisOR14 showed specificity for Z9,E12-14:OH; while AdisOR6 and AdisOR11 did not respond to any of the pheromone components and analogs tested. Taken together, this study contributes to elucidate the molecular mechanism of sex pheromone reception and provides potential targets for development of OR based pest control techniques in A. dissimilis.
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DOI:10.1002/ps.5107URLPMID:29869368 [本文引用: 1]
BACKGROUND: The peach fruit moth, Carposina sasakii Matsumura (Lepidoptera: Carposinidae), is one of the most destructive pests of pome and stone fruits, while few studies of their molecular biology and physiology have been conducted. Research into CsasPBPs (Carposina sasakii pheromone binding proteins) and CsasGOBPs (Carposina sasakii general odorant binding proteins) may provide insights in to the mechanisms of olfaction in Carposina sasakii. RESULTS: In our study, results of real time quantitative polymerase chain reaction (qPCR) assays demonstrated that CsasPBP1-3 and CsasGOBP1-2 transcripts were abundantly expressed in the antennae of both sexes, suggesting they play a vital role in olfaction. In addition, to examine specific functional differences between pheromone binding proteins (PBPs) and general odorant binding proteins (GOBPs), fluorescence competitive binding assays were used to measured the binding affinities for the two sex pheromones and 18 apple plant volatiles. As a result, both PBPs and GOBPs showed stronger binding affinities to Z-7-eicosene-11-one than Z-7-nonadecene-11-one in two sex pheromones, whereas only PBP3 exhibited specific affinity towards both these two sex pheromone components, and PBP1 showed a high binding affinity to the sex pheromone components, and to other plant volatiles. In addition, GOBP1-2 displayed high binding affinity to general components of plant volatiles. CONCLUSION: Our study suggested CsasPBPs and CsasGOBPs play distinct physiological roles in the perception of sex pheromones and host plant volatiles. (c) 2018 Society of Chemical Industry.
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DOI:10.1016/j.gene.2016.06.052URLPMID:27374155 [本文引用: 1]
At the peripheral level of the insect olfaction, odorant binding proteins (OBPs) are thought to bind and transport exogenous hydrophobic volatiles to the odorant receptors (ORs) located on the dendrite membrane of the olfactory neurons. In this study, cDNA sequences of 29 OBP genes from Chilo suppressalis, a notorious rice pest, were identified, with 15 of them being newly reported. The tissue and temporal expression patterns of these CsupOBPs were determined by RT-PCR, revealing that CsupOBP8, 10 and 24 were specifically expressed in the heads at larval stage and in antennae of both sexes at adult stage. In addition, CsupOBP 6, 9 and 18 were expressed in much higher levels in heads than abdomen at the larval stage, while CsupPBP4 and CsupOBP1, 3, 8, 11 and 24 were specifically expressed in antennae of both sexes at the adult stage. Such expression profile strongly suggests olfactory roles of these genes. Furthermore, the function of CsupOBP8 was chosen to be investigated by the competitive fluorescence binding assay. Results showed that CsupOBP8 displayed high binding affinities (Ki=4.9-15.0muM) with plant volatile beta-ionone, nerolidol, farnesol and 2-hexanone, suggesting that CsupOBP8 plays olfactory roles through binding and transporting the plant volatiles. The study provides an important base for understanding the olfactory mechanisms in C. suppressalis.
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DOI:10.1371/journal.pone.0043034URLPMID:22952629 [本文引用: 1]
Insects are the most diverse group of animals on the planet, comprising over 90% of all metazoan life forms, and have adapted to a wide diversity of ecosystems in nearly all environments. They have evolved highly sensitive chemical senses that are central to their interaction with their environment and to communication between individuals. Understanding the molecular bases of insect olfaction is therefore of great importance from both a basic and applied perspective. Odorant binding proteins (OBPs) are some of most abundant proteins found in insect olfactory organs, where they are the first component of the olfactory transduction cascade, carrying odorant molecules to the olfactory receptors. We carried out a search for OBPs in the genome of the parasitoid wasp Nasonia vitripennis and identified 90 sequences encoding putative OBPs. This is the largest OBP family so far reported in insects. We report unique features of the N. vitripennis OBPs, including the presence and evolutionary origin of a new subfamily of double-domain OBPs (consisting of two concatenated OBP domains), the loss of conserved cysteine residues and the expression of pseudogenes. This study also demonstrates the extremely dynamic evolution of the insect OBP family: (i) the number of different OBPs can vary greatly between species; (ii) the sequences are highly diverse, sometimes as a result of positive selection pressure with even the canonical cysteines being lost; (iii) new lineage specific domain arrangements can arise, such as the double domain OBP subfamily of wasps and mosquitoes.
DOI:10.1016/j.bbrc.2015.10.008URLPMID:26454175
Asecodes hispinarum (Hymenoptera: Eulophidae) is an endoparasitoid and an efficient biological control agent which attacks larvae of Brontispa longissima, a serious insect pest of Palmae plants in China. Odorant binding proteins (OBPs) are believed to be important for transporting semiochemicals through the aqueous sensillar lymph to the olfactory receptor cells within the insect antennal sensilla. No previous study has reported on OBPs in A. hispinarum. In this study, we conducted the large-scale identification of OBP genes from the antennae of A. hispinarum by using transcriptome sequencing. Approximately 28.4 million total raw reads and about 27.3 million total clean reads were obtained, and then 46,363 unigenes were assembled. Of these unigenes, a total of 21,263 can be annotated in the NCBI non-redundant database. Among the annotated unigenes, 16,623 of them can be assigned to GO (Gene Ontology). Furthermore, we identified 8 putative OBP genes, and a phylogenetic tree analysis was performed to characterize the 8 OBP genes. In addition, the expression of the 8 OBP genes in different A. hispinarum body tissues was analyzed by real-time quantitative polymerase chain reaction (qRT-PCR). The results indicated that the 8 OBP genes were expressed accordingly to sexes and tissues, but all highly expressed in antennae. The finding of this study will lay the foundation for unraveling molecular mechanisms of A. hispinarum chemoperception.
DOI:10.1038/srep24078URLPMID:27090020
Insects are extremely successful animals whose odor perception is very prominent due to their sophisticated olfactory system. The main chemosensory organ, antennae play a critical role in detecting odor in ambient environment before initiating appropriate behavioral responses. The antennal chemosensory receptor genes families have been suggested to be involved in olfactory signal transduction pathway as a sensory neuron response. The Macrocentrus cingulum is deployed successfully as a biological control agent for corn pest insects from the Lepidopteran genus Ostrinia. In this research, we assembled antennal transcriptomes of M. cingulum by using next generation sequencing to identify the major chemosensory receptors gene families. In total, 112 olfactory receptors candidates (79 odorant receptors, 20 gustatory receptors, and 13 ionotropic receptors) have been identified from the male and female antennal transcriptome. The sequences of all of these transcripts were confirmed by RT-PCR, and direct DNA sequencing. Expression profiles of gustatory receptors in olfactory and non-olfactory tissues were measured by RT-qPCR. The sex-specific and sex-biased chemoreceptors expression patterns suggested that they may have important functions in sense detection which behaviorally relevant to odor molecules. This reported result provides a comprehensive resource of the foundation in semiochemicals driven behaviors at molecular level in polyembryonic endoparasitoid.
DOI:10.1016/j.gene.2018.07.065URLPMID:30107229 [本文引用: 1]
Chemoreception is critical for the survival of insects. Insects have a variety of behavioral responses, such as mating, host searching and ovipositing, in response to different odor signals detected in their living environment. Trichogramma chilonis, an egg parasitoid, acts as an efficient and effective biocontrol reagent for many agricultural and forestry insect pests in many parts of China. However, little is known about the molecular mechanism of the olfaction-evoked behavior in T. chilonis. In the present study, we conducted transcriptome profiling analysis of T. chilonis based on the Illumina high-throughput sequencing platform in order to explore differences of chemoreception between male and female T. chilonis. In this study, a total of 85 chemosensory genes were identified from transcriptomic data, including 45 odorant receptors (ORs), 22 odorant binding proteins (OBPs), 14 ionotropic receptors (IRs), 2 sensory neuron membrane proteins (SNMPs) and 2 chemosensory proteins (CSPs). From the analysis of the transcriptome, most of the candidate olfactory genes had similar expression levels in males and females, including a few OR and OBP genes (TchiOR38, TchiOR39, TchiOR40, TchiOR41, TchiOR42, TchiOR43, TchiOR44, TchiOR45, TchiOBP1, TchiOBP4, TchiOBP10, TchiOBP12, TchiOBP18 and TchiOBP19) which showed male-biased expression. Some annotated unigenes were chosen randomly to have qRT-PCR, which verified the correctness of analysis of transcriptome in T. chilonis. This is the first study to obtain and identify candidate genes related to chemoreception in T. chilonis. Our work lays a solid foundation for related future research on the chemosensory system of T. chilonis at the molecular level and helps advance the use of T. chilonis as biological control agents.
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DOI:10.1016/j.jinsphys.2018.10.004URLPMID:30336148 [本文引用: 1]
Chrysoperla sinica is an important natural predator of many notorious agricultural pests. Understanding its olfactory mechanism can help enhance the effectiveness of C. sinica in biological control. In the present study, we investigated the tissue expression patterns of 12 odorant-binding protein (OBP) genes from C. sinica (CsinOBPs). The results of quantitative real-time polymerase chain reaction (qPCR) showed that CsinOBP1, CsinOBP2, CsinOBP3, CsinOBP4, CsinOBP6, CsinOBP7, CsinOBP9, CsinOBP10, and CsinOBP12 were predominantly expressed in the antennae of both sexes, indicating their roles in olfaction. Additionally, the qPCR analysis revealed that the 12 CsinOBP genes had distinct expression patterns, while the motif-pattern investigation suggested that the OBPs had different ligands. The ligand-binding assay showed that CsinOBP1 and CsinOBP10 had broader binding spectra than did the other OBPs. Thus, CsinOBP1 was able to bind not only plant volatiles (such as farnesol, cis-3-hexenyl hexanoate, geranylacetone, beta-ionone, 2-tridecanone, and trans-nerolidol) but also the aphid alarm pheromone (E)-beta-farnesene. On the other hand, CsinOBP2 and CsinOBP6 exhibited relatively narrow binding spectra, only binding ethyl benzoate. The study also identified several compounds that can potentially be used to develop slow-release agents attracting C. sinica and to improve search strategies for insect pest control.
URLPMID:15882644 [本文引用: 1]
DOI:10.1534/genetics.107.078683URLPMID:18039874 [本文引用: 1]
The insect chemoreceptor superfamily comprises the olfactory receptor (Or) and gustatory receptor (Gr) multigene families. These families give insects the ability to smell and taste chemicals in the environment and are thus rich resources for linking molecular evolutionary and ecological processes. Although dramatic differences in family size among distant species and high divergence among paralogs have led to the belief that the two families evolve rapidly, a lack of evolutionary data over short time scales has frustrated efforts to identify the major forces shaping this evolution. Here, we investigate patterns of gene loss/gain, divergence, and polymorphism in the entire repertoire of approximately 130 chemoreceptor genes from five closely related species of Drosophila that share a common ancestor within the past 12 million years. We demonstrate that the overall evolution of the Or and Gr families is nonneutral. We also show that selection regimes differ both between the two families as wholes and within each family among groups of genes with varying functions, patterns of expression, and phylogenetic histories. Finally, we find that the independent evolution of host specialization in Drosophila sechellia and D. erecta is associated with a fivefold acceleration of gene loss and increased rates of amino acid evolution at receptors that remain intact. Gene loss appears to primarily affect Grs that respond to bitter compounds while elevated Ka/Ks is most pronounced in the subset of Ors that are expressed in larvae. Our results provide strong evidence that the observed phenomena result from the invasion of a novel ecological niche and present a unique synthesis of molecular evolutionary analyses with ecological data.
URLPMID:23614018 [本文引用: 1]
DOI:10.1016/j.cub.2017.04.035URLPMID:28528905 [本文引用: 1]
Many insect species use multi-component sex pheromones to discriminate among potential mating partners [1-5]. In moths, pheromone blends tend to be dominated by one or two major components, but behavioral responses are frequently optimized by the inclusion of less abundant minor components [6]. An increasing number of studies have shown that female insects use these chemicals to convey their mating availability to males, who can assess the maturity of females and thus decide when to mate [7, 8]. However, little is known about the biological mechanisms that enable males to assess female reproductive status. In this study, we found that females of Helicoverpa armigera avoid nonoptimal mating by inhibiting males with pheromone antagonist cis-11-Hexadecenol (Z11-16:OH). We also show that this antagonist-mediated optimization of mating time ensures maximum fecundity. To further investigate molecular aspects of this phenomenon, we used the CRISPR/Cas9 system to knock out odorant receptor 16 (OR16), the only pheromone receptor tuned to Z11-16:OH. In mutant males, electrophysiological and behavioral responses to Z11-16:OH were abolished. Inability to detect Z11-16:OH prompted the males to mate with immature females, which resulted in significantly reduced viability of eggs. In conclusion, our study demonstrates that the sensitivity of OR16 to Z11-16:OH regulates optimal mating time and thus ensures maximum fecundity. These results may suggest novel strategies to disrupt pest insect mating.
DOI:10.7554/eLife.31101URLPMID:29271742 [本文引用: 1]
Neurite initiation is the first step in neuronal development and occurs spontaneously in soft tissue environments. Although the mechanisms regulating the morphology of migratory cells on rigid substrates in cell culture are widely known, how soft environments modulate neurite initiation remains elusive. Using hydrogel cultures, pharmacologic inhibition, and genetic approaches, we reveal that paxillin-linked endocytosis and adhesion are components of a bistable switch controlling neurite initiation in a substrate modulus-dependent manner. On soft substrates, most paxillin binds to endocytic factors and facilitates vesicle invagination, elevating neuritogenic Rac1 activity and expression of genes encoding the endocytic machinery. By contrast, on rigid substrates, cells develop extensive adhesions, increase RhoA activity and sequester paxillin from the endocytic machinery, thereby delaying neurite initiation. Our results highlight paxillin as a core molecule in substrate modulus-controlled morphogenesis and define a mechanism whereby neuronal cells respond to environments exhibiting varying mechanical properties.
URLPMID:24632377 [本文引用: 1]
DOI:10.1038/srep18742URLPMID:26744070 [本文引用: 1]
Male moths efficiently recognize conspecific sex pheromones thanks to their highly accurate and specific olfactory system. The Heliothis/Helicoverpa species are regarded as good models for studying the perception of sex pheromones. In this study, we performed a series of experiments to investigate the peripheral mechanisms of pheromone coding in two-closely related species, Helicoverpa armigera and H. assulta. The morphology and distribution patterns of sensilla trichoidea are similar between the two species when observed at the scanning electron microscope, but their performances are different. In H. armigera, three functional types of sensilla trichoidea (A, B and C) were found to respond to different pheromone components, while in H. assulta only two types of such sensilla (A and C) could be detected. The response profiles of all types of sensilla trichoidea in the two species well matched the specificities of the pheromone receptors (PRs) expressed in the same sensilla, as measured in voltage-clamp experiments. The expressions of PRs in neighboring olfactory sensory neurons (OSNs) within the same trichoid sensillum were further confirmed by in situ hybridization. Our results show how the same pheromone components can code for different messages at the periphery of two Helicoverpa species.
DOI:10.1111/j.1365-2583.2010.01045.xURLPMID:20946532 [本文引用: 1]
Functional analyses of candidate Heliothis virescens pheromone odorant receptors (HvORs) were conducted using heterologous expression in Xenopus oocytes. HvOR6 was found to be highly tuned to Z9-14:Ald, while HvOR13, HvOR14 and HvOR16 showed specificity for Z11-16:Ald, Z11-16:OAc and Z11-16:OH, respectively. HvOR15, which had been considered a candidate receptor for Z9-14:Ald did not respond to any of the pheromone compounds tested, nor to 50 other general odorants. Thus, while HvOR15 is specifically expressed in H. virescens male antennae, its role in pheromone reception remains unknown. Based on our results and previous research we can now assign pheromone receptors in H. virescens males to each of the critical H. virescens agonistic pheromone compounds and two antagonistic compounds produced by heterospecific females.
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DOI:10.1016/j.ibmb.2013.05.009URLPMID:23751753 [本文引用: 1]
In moths, males can detect a distinct blend of several pheromone components by specialized olfactory receptor neurons (ORNs) on the antennae. Four candidate pheromone receptors (PR) with seven transmembrane domains were identified by homology cloning from the antennae of Spodoptera exigua (Sexi). Phylogenetic analyses reveal that all four odorant receptors (OR) belong to pheromone receptor subtypes. Expression patterns revealed that PRs were male-specific in the antenna except for SexiOR11, which was female antenna-biased. Functional analyses of these PRs were conducted using heterologous expression in Xenopus oocytes. SexiOR13 and SexiOR16 were all broadly activated by multiple pheromone components. SexiOR13 responded robustly to the critical pheromone component, Z9, E12-14:OAc and the minor pheromone component, Z9-14:OAc at a concentration of 10(-4) M. Dose-response studies indicate that SexiOR13 was approximately 4 times more sensitive to Z9,E12-14:OAc (EC50 = 3.158 x 10(-6) M) compared to Z9-14:OAc (EC50 = 1.203 x 10(-5) M). While, SexiOR16 responded robustly to the secondary pheromone component Z9-14:OH with high sensitivity (EC50 = 9.690 x 10(-7) M). However, similar tests of the five pheromones with SexiOR6 and SexiOR11 failed to elicit any response. These results provide basic knowledge to further advance research on the molecular mechanisms of pheromone reception.
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DOI:10.1111/imb.12033URLPMID:23679893 [本文引用: 1]
Olfaction plays an important role in insect behaviours. The odorant receptor (OR) repertoire, housed within the dendritic membrane of sensory neurons, is one of the primary determinants of odour recognition. ORs in moths could be classified into pheromone receptors (PRs) and non-pheromone receptors (non-PR ORs). Much research in the field of insect olfaction recently has been focused on PRs of the male moth, but few Lepidoptera studies have been done on the functional study of non-PR ORs. In the present study, we identified and characterized four non-PR ORs from Spodoptera litura (Lepidoptera: Noctuidae) antennae. The tissue expression pattern showed that the four ORs were mainly expressed in adult antennae and further in situ hybridization revealed SlituOR12 was expressed in both long and short sensilla trichodea and sensilla basiconica. A functional analysis of the four SlituORs was conducted in the heterologous expression system Xenopus oocytes. SlituOR12 was exclusively and sensitively tuned to cis-3-Hexenyl acetate and SlituOR19 slightly responded to 4'-Ethylacetophenone; however, SlituOR44 and SlituOR51 did not respond to any chemicals tested in this study. It is proposed that SlituOR12 might partially account for some key behaviours of the female, such as detection of host location and oviposition site.
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DOI:10.1371/journal.pone.0155029URLPMID:27163122 [本文引用: 1]
Odorant receptors (ORs) play an important role in insects to monitor and adapt to the external environment, such as host plant location, oviposition-site selection, mate recognition and natural enemy avoidance. In our study, we identified and characterized OR12 from three closely-related species, Helicoverpa armigera, Helicoverpa assulta, Heliothis virescens, sharing between 90 and 98% of their amino acids. The tissue expression pattern analysis in H. armigera showed that HarmOR12 was strongly expressed both in male and female antennae, but not in other tissues. Functional analysis performed in the heterologous Xenopus expression system showed that all three OR12 were tuned to six structurally related plant volatiles. Electroantennogram recordings from male and female antennae of H. armigera closely matched the data of in vitro functional studies. Our results revealed that OR12 has a conserved role in Heliothinae moths and might represent a suitable target for the control of these crop pests.
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DOI:10.7554/eLife.53706URLPMID:32436842 [本文引用: 1]
Antennae are often considered to be the nostrils of insects. Here, we sequenced the transcriptome of the pheromone gland-ovipositor complex of Helicoverpa assulta and discovered that an odorant receptor (OR) gene, HassOR31, had much higher expression in the ovipositor than in antennae or other tissues. To determine whether the ovipositor was involved in odorant detection, we co-expressed HassOR31 and its co-receptor, HassORco, in a Xenopus oocyte model system, and demonstrated that the OR was responsive to 12 plant odorants, especially Z-3-hexenyl butyrate. These odorants elicited electrophysiological responses of some sensilla in the ovipositor, and HassOR31 and HassORco were co-expressed within ovipositor sensilla. Two oviposition preference experiments showed that female moths lacking antennae still preferentially selected oviposition sites containing plant volatiles. We suggest that the expression of HassOR31 in the ovipositor of H. assulta helps females to determine precise egg-laying sites in host plants.
URLPMID:30058747 [本文引用: 1]
DOI:10.1016/j.ibmb.2017.09.001URLPMID:28912112 [本文引用: 1]
Anastatus japonicus is a parasitic wasp and natural enemy of the litchi pest Tessaratoma papillosa, and for decades in China, A. japonicus has been mass-reared inside the eggs of Antheraea pernyi to control T. papillosa. A series of experiments was performed to explore the olfactory mechanism underlying the oviposition behavior of A. japonicus. First, a transcriptomic analysis was performed on the antennae of A. japonicus, and the resulting assemblies led to the generation of 70,473 unigenes. Subsequently, 21,368 unigenes were matched to known proteins, 48 odorant receptors (ORs) (including Orco) and 13 antennal ionotropic receptors (IRs) (including the co-receptors IR8a and IR25a) were identified and predicted to form complete open reading frames (ORFs). The FPKM (fragments per Kb per million reads) values and RT-PCR results showed that AjapOrco, AjapOR10, AjapOR27, AjapOR33 and AjapOR35 were either highly abundant or expressed specifically in the olfactory organs. Furthermore, AjapOrco silencing resulted in a significant decrease in both the parasitism rate and the host-seeking time of A. japonicus, whereas dsRNA injection showed that IR8a and IR25a did not produce significant behavioral changes, suggesting that the oviposition behavior of A. japonicus is more reliant on OR-based pathways than IR-based pathways. Our previous GC-MS data derived twenty-nine compounds which were abundent from these host plants and host insects. We performed electrophysiological and oviposition assays on A. japonicus, and eight odorants were found to elicit a significant electroantennogram (EAG) response. Among these odorants, beta-Caryophyllene, Undecane, (E)-alpha-Farnesene (+)-Aromadendrene and Cis-3-Hexen-ol had strong attractant effects on oviposition, whereas 2-Ethyl-1-Hexan-ol, Ethyl Acetate and alpha-Caryophyllene had a strong repellant effects. Thus, these chemicals might influence oviposition guidance/repulsion behavior in A. japonicus. To further explore the target ORs that are tuned to the functional odorants, the nine candidate ORs described above were silenced by RNA interference, and the results showed that a large decrease in the EAG response of all the tested functional odorants in the AjapOrco-silencing group. In addition, the AjapOR35-silencing group showed a significant decrease in the EAG response to beta-Caryophyllene and (E)-alpha-Farnesene, indicating that AjapOR35 is tuned to these two oviposition attractants beta-Caryophyllene and (E)-alpha-Farnesene. Further binary-choice oviposition assays showed that the oviposition attractant effect of beta-Caryophyllene and (E)-alpha-Farnesene vanished after AjapOR35 was silenced, indicating that the emission of these attractants from host plants can guide A. japonicus to locate eggs for ovipositioning and indicated that AjapOR35 is correlated with the olfactory detection oviposition behavior of this species. This study provides a better understanding of the molecular basis and functional chemicals underlying the oviposition behavior of A. japonicus, and the results may help improve biocontrol approaches.
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URLPMID:20388299 [本文引用: 1]
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DOI:10.1007/s10886-005-6072-9URLPMID:16132208 [本文引用: 1]
Jasmonate signaling pathway plays an important role in induced plant defense against herbivores and pathogens, including the emission of volatiles that serve as attractants for natural enemies of herbivores. We studied the volatiles emitted from rice plants that were wounded and treated with jasmonic acid (JA) and their effects on the host-searching behavior of the rice brown planthopper, Nilaparvata lugens (Stal), and its mymarid egg parasitoid Anagrus nilaparvatae Pang et Wang. Female adults of N. lugens significantly preferred to settle on JA-treated rice plants immediately after release. The parasitoid A. nilaparvatae showed a similar preference and was more attracted to the volatiles emitted from JA-treated rice plants than to volatiles from control plants. This was also evident from greenhouse and field experiments in which parasitism of N. lugens eggs by A. nilaparvatae on plants that were surrounded by JA-treated plants was more than twofold higher than on control plants. Analyses of volatiles collected from rice plants showed that JA treatment dramatically increased the release of volatiles, which included aliphatic aldehydes and alcohols, monoterpenes, sesquiterpenes, methyl salicylate, n-heptadecane, and several as yet unidentified compounds. These results confirm an involvement of the JA pathway in induced defense in rice plants and demonstrate that the egg parasitoid A. nilaparvatae exploits plant-provided cues to locate hosts. We explain the use of induced plant volatiles by the egg parasitoid by a reliable association between planthopper feeding damage and egg presence.
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DOI:10.1016/s0958-1669(03)00025-9URLPMID:12732318 [本文引用: 1]
When attacked by herbivorous insects or mites, some plant species call on other arthropods for help. They emit mixtures of volatile compounds, dominated by terpenes, to attract carnivorous arthropods that prey on or parasitise herbivores and so reduce further damage. This fascinating defence strategy offers a new, environmentally friendly approach to crop protection. Using recent advances in the biochemistry and molecular genetics of terpene biosynthesis, it should now be possible to engineer crop plants that release terpenes for attracting herbivore enemies. By introducing or selectively altering the existing rate of terpene emission and composition, plant breeders could enable attacked plants to attract enemies and reduce additional herbivory, without compromising the effectiveness of other modes of defence.
DOI:10.1016/j.tplants.2009.12.002URLPMID:20047849 [本文引用: 1]
Attacks by herbivores elicit changes in the bouquet of volatiles released by plants. These herbivore-induced plant volatiles (HIPVs) have been interpreted as being indirect defenses. However, given that no studies have yet investigated whether HIPVs benefit the fitness of a plant, their defensive function remains to be established. Moreover, herbivores, pathogens, pollinators and competitors also respond to HIPVs and, in addition, neighbouring plants in native populations also emit volatiles that provide a background odour. These considerations enrich the evolutionary context of HIPVs and complicate predictions about their adaptive value. Molecular advances in our understanding of HIPV signaling and biosynthesis is enabling the creation of HIPV-'mute' and possibly HIPV-'deaf' plants. As we discuss here, such plants could be used for unbiased examination of the fitness value of HIPV emissions under natural conditions.
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DOI:10.1038/srep11183URLPMID:26108150 [本文引用: 1]
Insect pheromones offer potential for managing pests of crop plants. Volatility and instability are problems for deployment in agriculture but could be solved by expressing genes for the biosynthesis of pheromones in the crop plants. This has now been achieved by genetically engineering a hexaploid variety of wheat to release (E)-beta-farnesene (Ebetaf), the alarm pheromone for many pest aphids, using a synthetic gene based on a sequence from peppermint with a plastid targeting amino acid sequence, with or without a gene for biosynthesis of the precursor farnesyl diphosphate. Pure Ebetaf was produced in stably transformed wheat lines with no other detectable phenotype but requiring targeting of the gene produced to the plastid. In laboratory behavioural assays, three species of cereal aphids were repelled and foraging was increased for a parasitic natural enemy. Although these studies show considerable potential for aphid control, field trials employing the single and double constructs showed no reduction in aphids or increase in parasitism. Insect numbers were low and climatic conditions erratic suggesting the need for further trials or a closer imitation, in the plant, of alarm pheromone release.
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DOI:10.1023/a:1015253600083URLPMID:12049233 [本文引用: 1]
Herbivore-induced plant volatiles can function as indirect defense signals that attract natural enemies of herbivores. Several parasitoids are known to exploit these plant-provided cues to locate their hosts. One such parasitoid is the generalist Cotesia marginiventris, which is, among others, attracted to maize volatiles induced by caterpillar damage. Maize plants can be induced to produce the same blend of attractive volatiles by treating them with regurgitant of Spodoptera species. We collected and analyzed the regurgitant-induced emissions of two plant species (cowpea and maize) and of eight Mexican maize varieties and found significant differences among their volatile emissions, both in terms of total quantity and the quality of the blends. In a Y-tube olfactometer. the odors of the same artificially induced plant species and Mexican varieties were offered in dual choice experiments to naive mated females of C. marginiventris. Wasps preferred cowpea over maize odor and, in 3 of 12 combinations with the maize varieties, they showed a preference for the odors of one of the varieties, A comparison of the odor collection with results from the behavioral assays indicates that not only the quantity of the volatile emissions, but also the quality (composition) of the volatile blends is important for attraction of C. marginiventris. The results are discussed in the context of the possibility of breeding crop varieties that are particularly attractive to parasitoids.
DOI:10.1007/BF00979619URLPMID:24263303 [本文引用: 1]
Considerable basic information has been gathered on the interaction between the onion fly (Delia antiqua) and its host plant, the onion (Allium cepa). An attempt is underway to manipulate ovipositional behavior of this pest by treating onion seedlings with chemical deterrents while simultaneously providing deeply planted onion culls on which onion flies prefer to lay. This bipolar strategy of behavioral manipulation, termed
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DOI:10.1016/j.copbio.2013.12.006URLPMID:24445079 [本文引用: 1]
Farming systems for pest control, based on the stimulo-deterrent diversionary strategy or push-pull system, have become an important target for sustainable intensification of food production. A prominent example is push-pull developed in sub-Saharan Africa using a combination of companion plants delivering semiochemicals, as plant secondary metabolites, for smallholder farming cereal production, initially against lepidopterous stem borers. Opportunities are being developed for other regions and farming ecosystems. New semiochemical tools and delivery systems, including GM, are being incorporated to exploit further opportunities for mainstream arable farming systems. By delivering the push and pull effects as secondary metabolites, for example, (E)-4,8-dimethyl-1,3,7-nonatriene repelling pests and attracting beneficial insects, problems of high volatility and instability are overcome and compounds are produced when and where required.
