摘要
本文通过构建蝗虫触角转录组文库,利用生物信息学方法首次鉴定出沙漠蝗的离子型受体(IRs),并对其序列特征、时空表达特征、细胞定位等特征进行了研究；首次鉴定出了飞蝗的求偶相关的信息素及相应的气味分子结合蛋白(LmigOBP4),并且综合利用分子生物学、行为学、电生理学等方法,证明了LmigOBP4参与了飞蝗成虫对飞蝗求偶信息素的感受过程。主要结果如下：
I.沙漠蝗(Schistocerca gregaria)离子型受体IR8a和IR25a的克隆、时空表达和细胞定位
通过对沙漠蝗触角转录组文库进行生物信息学分析,组装得到两个可能的离子受体IR8a和IR25a基因的部分片段。首次在沙漠蝗触角中克隆到SgreIR8a ORF的全长序列,和SgreIR25a的部分序列,序列比对显示它们与其他昆虫的IR8a和IR25a有很高的序列同源性。其次,RT-PCR的结果表明,SgreIR8a和SgreIR25a在蝗蝻期和成虫期的触角中均有表达。再次,通过原位杂交实验,证明SgreIR8a和SgreIR25a表达在腔锥形感器下的神经元细胞中。最后,双荧光原位杂交实验表明,SgreIR8a和SgreIR25a在腔锥形感器下的部分细胞中存在共表达现象。另外还发现,SgreIR25a表达在部分刺形感器下的神经元细胞中。但是两个IRs都不表达在锥形感器和毛形感器下的细胞里。这一点,通过与SgreORco探针的双荧光原位杂交得到进一步证明,即表达SgreIR8a和SgreIR25a的细胞,都不表达SgreORco。综上,证明沙漠蝗的触角采用了两类不同的嗅觉感受神经元来感受气味：位于腔锥形感器中表达IR的神经元以及位于锥形感器和毛形感器中表达OR的神经元,为揭示蝗虫的嗅觉机制奠定了基础。
Ⅱ. LmigOBP4参与了飞蝗(Locusta migratoria)对信息素的感受过程
前人的研究表明KT(一种化合物,暂时保密)是雄成虫体表的特异挥发物,能引起毛形感器神经元的兴奋或抑制性反应。电生理实验证明其能够引起雌雄成虫的触角电生理反应。行为学实验表明,KT对5龄幼虫没有任何的作用,而在一定剂量下,对雌成虫有吸引作用,对雄成虫有排斥作用,且这种作用模式与一组雄虫(30头)吸引单头雌虫是类似的,推测其可能为求偶相关的信息素。在飞蝗触角cDNA文库中筛选到一个高丰度表达的气味分子结合蛋白(OBP),命名为LmigOBPA,对该蛋白进行原核表达和纯化。利用重组蛋白进行荧光竞争结合实验,结果表明LmigOBPA与KT有很强的结合能力。Western blotting结果表明该蛋白只表达在嗅觉器官---触角上；原位杂交和免疫电镜的结果表明LmigOBP4在三种类型的嗅觉感器(锥形、毛形、腔锥形感器)的淋巴液中都有表达。初步表明LmigOBP4可能参与了KT的识别。通过RNAi的方法显著地降低OBP4的表达量后,发现显著降低了雌雄虫触角对KT的电生理反应。进一步地,行为学测定结果表明,干扰OBP4表达的飞蝗对KT的行为反应发生了显著的改变：OBP4的缺失逆转了KT对雌成虫的吸引反应和对雄成虫的排斥反应,转变为,KT吸引雄成虫而排斥雌成虫,当KT剂量升高时,突变型蝗虫的行为反应并没有恢复到与野生型一致。综上结果表明,LmigOBP4参与了东亚飞蝗对信息素KT的感受过程,且对雌雄虫做出正确的求偶反应起到了关键性作用。
本文以沙漠蝗为材料,首次对其离子型受体的表达特征进行的深入的研究,这是继果蝇之后,第一次在昆虫中进行如此细致的研究,为完善昆虫的嗅觉机制,以及未来蝗虫的嗅觉机制研究奠定了基础。本文首次对飞蝗的求偶信息素及相应的结合蛋白进行了鉴定,初步明确了其生理功能,为从飞蝗求偶分子机制方向设计控制蝗虫物种特异的分子靶标奠定了基础,这对制定防治蝗虫的绿色防控体系提供了依据。
Two novel ionotropic receptors, IR8a and IR25a were firstly identified in grasshopper, Schistocerca gregaria. Further research was conducted in spatiotemporal expression pattern, cell localization and so on. A putative courtship related pheromone was defined and a possible pheromone binding protein-OBP4may be involved in the process of sensing this pheromone, details are as follows,
I. Identification and Characterization of IR8a and IR25a in locust, Schistocerca gregaria
In this study, we have identified the genes of S. gregaria which encode homologues of co-receptors for the variant ionotropic receptors, the subtypes IR8a and IR25a. It was found that both subtypes, SgreIR8a and SgreIR25a, was expressed in the antennae of all five nymphal stages and adults. Attempts to assign the relevant cell types by means of in situ hybridization revealed that SgreIR8a and SgreIR25a are expressed in cells of sensilla coeloconica. Double fluorescence in situ hybridization experiments disclosed that the two IR-subtypes are co-expressed in some cells of the coeloconic sensilla. Expression of SgreIR25a was also found in some of the sensilla chaetica, however, neither SgreIR25a nor SgreIRSa was found to be expressed in basiconic or trichoid sensilla. This observation was substantiated by the results of double FISH experiments demonstrating that cells expressing ISgreIR8a or SgreIB25a do not express SgreORco. These results support the notion that the antenna of the desert locust employs two different populations of OSNs to sense odors:cells which express IRs in sensilla coeloconica and cells which express ORs in sensilla basiconica and sensilla trichodea.
II. A dual-role male pheromone sensed by odorant-binding protein in locust
Here for the first time, we identified KT as one courtship related pheromoene, a component of male adult body volatiles of locust, Locusta migratoria. KT has been reported by Cui et al., that can excite or inhibit neurons in a portion of sensilla trichodea. This compound elicited high electrophysiological responses of male and female antenna. Behavioural assays showed that KT evoked sexually dimorphic behaviours at certain dosage, attracting females and repelling males but5instar nymphs did not show any response to KT. Together these results suggested that this compound is likely to be one courtship related pheromone of L. migratoria. Consequently, this pheromone was demonstrated to have binding affinity to a novel odorant binding protein which was identified from locust, named as LmigOBP4, highly expressed in olfactory sensilla observed in in situ hybridyzation and immunocytochemistry localization experiments. Furthermore, the defect of LmigOBP4by RNAi technique diminished significantly electroantennogram responses of both male and female locusts to KT. Consistently, behavioural tendencies in the selection of KT at certain doses by both sexes were reversed with that of the comparison of wild types. Additionally, behavioural impairment of KT by RNA interference of Lmigobp4could not be restored and compensated by increasing doses. Therefore, we have characterized a dual-role locust male-specific pheromone, KT which can elicit courtship behaviour, attracting females and repelling males, and the pheromone is sensed by a novel odorant binding protein, LmigOBP4. Our findings support that odorant binding protein plays key roles in the process of pheromone detection, and locust has highly efficient courtship communication system which is more similar to fly, other than moth. This provides the evidence to understand the molecular evolution of animal courtship, as well as to develop new methods for controling locust plagues based on functions of molecules in locust courtship behavior.
引文
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