雄性棉铃虫感受性信息素的分子和神经机制研究进展

Research progress in the molecular and neural mechanisms of sex pheromone reception in male Helicoverpa armigera(Lepidoptera:Noctuidae)

棉铃虫Helicoverpa armigera主要借助于性信息素通讯完成雌雄识别,实现交配和种群繁衍。关于棉铃虫感受性信息素机制的研究一直是我国化学生态学领域的热点和重心,研究结果有助于开发和改进棉铃虫防治的性引诱剂。本文将对棉铃虫雄虫感受雌虫释放的性信息素的机制进行综述,以期为深入研究棉铃虫及其他相关昆虫的性信息素感受的分子和神经机理提供参考。棉铃虫雌虫性信息素腺体合成和释放多种长链、饱和或非饱和的脂肪醛和醇等化合物,其中Z 11-16:Ald为主要性信息素成分,Z 9-16∶Ald和Z 9-14∶Ald为次要性信息素成分,不同组分按一定比例混合可明显增强对雄性棉铃虫的引诱效果,而化合物Z 11-16∶OH和高剂量的Z 9-14∶Ald对性信息素引诱活性具有明显的抑制效果。相应地,雄性棉铃虫触角上A,B和C 3种类型的毛形感器能够感受这些信息化合物。A类型毛形感器内表达受体OR13感受Z 11-16∶Ald,B类型毛形感器内表达OR14b感受Z 9-14∶Ald,C类型毛形感器内表达OR6和OR16感受Z 9-16∶Ald,Z 9-14∶Ald,Z 11-16:Ac和Z 11-16∶OH。受体的表达位置和功能与不同类型毛形感器的电生理反应特性相一致。钙离子成像证明在棉铃虫触角叶内的3个扩大型神经纤维球接受这些气味信息,其中神经纤维球云状体接受Z 11-16∶Ald,背中间后侧纤维球接受Z 9-16∶Ald,背中间前侧纤维球接受Z 9-14∶Ald,Z 11-16∶Ac和Z 11-16∶OH。这些研究成果在感器、受体和脑中枢水平上揭示了棉铃虫感受性信息素的机制,在这些研究基础上,我们认为需要深入开展以下方面的研究:(1)进一步鉴定相关性信息素受体的功能和定位;(2)深入研究脑内嗅觉高级中枢对性信息素信息的处理和整合神经机制;(3)明确棉铃虫性信息素感受受到寄主植物、光周期、温度、湿度等环境因素的影响及机制。

The cotton bollworm,Helicoverpa armigera,mainly relies on sex pheromone communication for mate orientation and reproduction.The mechanisms of pheromone reception in H.armigera are the hot topic in the field of chemical ecology in China.In this article,we reviewed the reception mechanism to female sex pheromone in male H.armigera,hoping to provide references for further investigation of the molecular and neural mechanisms of sex pheromone perception in H.armigera and other related insects.In the sex pheromone glands of female adult,long-chain,saturated or unsaturated fatty aldehydes,and alcohols were synthesized,among which Z 11-16∶Ald was used as the main sex pheromone component,while Z 9-16∶Ald and Z 9-14∶Ald as the secondary sex pheromone components.The blends of these components at certain ratios show enhanced attraction to male H.armigera.Z 11-16∶OH and a high-dose Z 9-14∶Ald inhibit the attractiveness of pheromone blends to male H.armigera.Correspondingly,three types of trichoid sensilla,types A,B,and C,on the antennae of male H.armigera could detect these compounds.The receptor OR13 responding to Z 11-16∶Ald is expressed in the type-A trichoid sensillum,OR14b responding to Z 9-14∶Ald is expressed in the type-B trichoid sensillum,and OR6 and OR16 responding to Z 9-16∶Ald,Z 9-14∶Ald,and Z 11-16∶OH are expressed in the type-C trichoid sensillum.The expression sites and functions of the receptors are consistent with the electrophysiological characteristics of different types of trichoid sensilla.In addition,the results of calcium imaging showed that three macroglomeruli tune to these odors,among which the glomerulus of cumulus tunes to Z 11-16∶Ald,the posterior dorsomedial glomerulus to Z 9-16∶Ald,and the anterior dorsomedial glomerulus to Z 9-14∶Ald,Z 11-16∶Ac and Z 11-16∶OH.These results revealed the mechanisms of pheromone reception in male H.armigera at the levels of sensillum,receptor and central nervous system.Based upon these studies,we think it is necessary to make further studies on the following aspects:(1)further identifying the function and location of the related sex pheromone receptors;(2)further studying the processing and integration of sex pheromone information in high brain center;(3)clarifying the influences and mechanisms of such environmental factors as host plant,photoperiod,temperature and humidity on the sex pheromone perception in H.armigera.