摘要
棉铃虫是我国棉花生产上的重大致灾性害虫。研究表明，棉铃虫能够感受空气中的挥发性化合物，并将此作为交配、觅食和寻找生殖场所的信息，这一特性可以用来监测和诱杀棉铃虫。近年来，对棉铃虫的行为化学模式进行了广泛的研究，但是对棉铃虫嗅觉识别的分子机制国内外几乎未见报道。对这方面的研究不但可以阐明棉铃虫行为反应的本质原因，为研制高效的引诱剂或驱避剂提供理论依据，而且对于研究脊椎动物的嗅觉识别机制具有重要的启示作用。本论文利用基因克隆、表达的方法获得了纯化的棉铃虫气味结合蛋白，在此基础上制备了棉铃虫气味结合蛋白高度特异性的抗体，对两种气味结合蛋白在棉铃虫触角中的分布进行了免疫定位研究；同时利用差异显示PCR结合RACE技术克隆了一个与嗅觉有关的触角特异表达基因。主要结果如下：
(1)扫描和透射电镜观察结果表明：棉铃虫触角呈线状，由柄节、梗节和鞭节组成。在棉铃虫雌雄蛾触角上观察到了以下5类感器：即毛形感器、锥形感器、刺形感器、耳形感器、腔锥感器。棉铃虫触角上的嗅觉感器基本上由表皮、感觉神经元、感受器淋巴液以及鞘细胞组成。棉铃虫触角上的两类嗅觉感器，即毛形感器和锥形感器的内部形态结构具有明显的区别。
(2)利用PCR结合RACE技术克隆了3个棉铃虫OBP基因(PBP-Harm、GOBP1-Harm、GOBP2-Harm，GenBank中序列号分别为AJ278992、AY049739和AJ278991)和一个甜菜夜蛾GOBP2基因(GOBP2-Sexi，GenBank中序列号为AJ294809)。
(3)Northern杂交结果表明：PBP-Harm和GOBP2-Harm都在棉铃虫触角中特异性表达。PBP-Harm在雄蛾触角中表达量明显比雌蛾中高，在触角发育的过程中，PBP-Harm大约在成虫羽化前四天开始表达，在成虫羽化前两天达到峰值，这一高水平表达一直延续到成虫羽化。GOBP2-Harm在雌雄蛾触角中表达量相同，在触角发育的过程中，GOBP2-Harm大约在成虫羽化前三天开始表达，在成虫羽化前一天达到峰值，这一高水平表达一直延续到成虫羽化。Southern杂交表明：PBP-Harm和GOBP2-Harm都非性别连锁，这两个基因都是以单拷贝的形式存在。
(4)构建了棉铃虫PBP-Harm和GOBP2-Harm的原核表达载体pGEX／PBP-Harm和
PGEX／GOBPZ－Harm，并在大肠杆菌中成功进行了表达，经亲合层析和凝胶过滤获得
了高纯度的目的蛋白，免疫兔子制备了高度专一性的抗体，然后对PBPHarm和
GOBPZ－Harm在棉铃虫触角感器中的分布进行了免疫定位。结果表明：PBP和 GOBPZ
存在于触角毛形感器或锥形感器的淋巴液和感受器着生腔中，不存在于树突或感受器
表皮壁中。在雄蛾触角中，P＊P主要存在于毛形感器中，不存在于锥形感器中：而
GOBPZ主要存在于锥形感器和一些中等大小的毛形感器中。然而，在雌蛾触角中，
PBP存在于少部分的锥形感器和中等大小的毛形感器中，长形毛形感器中不存在
PBP；而GOBPZ不但存在于大部分的锥形感器和部分中等大小的毛形感器中，而且
还存在于部分的长形毛形感器中。
＊厉用差异显示PCR的方法获得了3条棉铃虫触角专一性CDNA片段。其中克隆
A4与昆虫谷肤甘肽七－转移酶基因具有明显的同源性，表明克隆A4属于GST超
基因家族。利用5’－RACE克隆A41的5’端片段，拼接以后获得了该基因的序列，命
名为GSThaoto在GenBank中序列号为AY058242。Northern杂交表明，GST山aolf
在棉铃虫触角中专一性表达，并且在雄蛾触角中表达量明显比雌蛾触角中的表达量
高。生物测定结果也表明雄蛾触角中的GST酶活性明显高于雌蛾触角。
Helicoverpa armigera (Htlbner) is an eruptive agricultural pest in China. It can perceive certain chemical volatiles in the environment as the chemical information sources for its behavioral reactions such as to locate their mates, food sources and oviposition and hibernation sites, and to avoid dangerous situations or unsuitable habitats and hosts, and so on, and these volatile chemicals can be used to montior and trap-kill pest. In recent years, the behavioral-chemical models of H.armigera have been widely researched, however, there is nearly no report so far concerned with the molecular mechanism of the odor recognition in H.armigera. So to explore the molecular mechanism of insect olfaction is not only helpful for
opening out the intrinsical law of insect behavior and providing theoretical evidences in developing effective attractant or deterrent, but also offer an ideal model system for studying the molecular details of olfaction for invertebrate or vertebrate. In this paper, recombinant OBPs of H.armigera were obtained by gene expression. Based on which high specific antibody was prepared following the distribution of PBP and GOBP2 in olfactory sensilla was studied by immuno-electron microscopy. At the same time, an antennal-specific gene in Helicoverpa armigera was cloned by dd-PCR and RACE. The primary results are as follows:
(1) Antennae of H.armigera are in linear shape and made up of scapus, pedicle and flagella. Both the antennae of male and female moth contain five kinds of antennal sensilla, namely, sensillum trichodeum, sensillum basiconicum, sensillum chaeticum, ear-shaped sensillum and sensillum coeloconicum. Olfaction Sensillum in antennae of H.armigera consists of cuticular-wall, dendrites, sensillum lymph and sheath cells. Sensillum trichodeum and sensillum basiconicum is the main chemical odor receptors on antennae of H.armigera and there are significant differences between the internal morpha structures of them.
(2) Three OBP genes of H.armigera (the accession numbers ofPBP-Harm, GOBPl-Harm and GOBP2-Harm are AJ278992, AY049739 and AJ278991 in GenBank, respectively)and a GOBP2 gene of Spodoptera exigua (the accession number of GOBP2-Sexi is AJ294809 in GenBank) were cloned by PCR and RACE techniques.
(3) Northern blot showed that PBP-Harm and GOBP2-Harm are specifically expressed in the antenna of H.armigera. However, PBP-Harm is more abundantly expressed in male antenna than that in female antenna. During the antennal development, PBP-Harm is firstly expressed about 4d prior to adult eclosion and rises to a plateau 2d prior to adult eclosion.
Contrast with PBP-Harm, expression level of GOBP2-Harm in male antenna is the same as the in female antenna. GOBP2-Harm is firstly expressed about 3d prior to adult eclosion and rises to a plateau Id prior to adult eclosion. Southern blot analysis showed that neither PBP-Harm nor GOBP2-Harm is sex-linked and two genes exist as a single copy in genome.
(4) Bacterical expression vectors of PBP-Harm and GOBP2-Harm genes were constructed and successfully expressed in E.coli. The recombant proteins were purified by affinity chromatography and gel filtration. Polyclonal antibodies against PBP-Harm and GOBP2-Harm were prepared to mark the distribution of the protein in olfactory sensilla by immuno-electron microscopy. In labeled sensilla, very strong labeling was always observed in the sensillum lymph, in the hair lumen as well as in the sensillum lymph cavity below the hair base. In the male, PBP-Harm is mainly expressed in sensilla trichodea and not in sensilla basiconica. GOBP2Harm is mainly expressed in sensilla basiconica and in some medium-sized sensilla trichodea. While in the female, PBP-Harm is expressed in a few sensilla basiconica and
medium-sized sensilla trichodea. GOBP2Harm is expressed in partial sensilla basiconica, medium-sized sensilla trichodea and long sensilla trichodea.
(5) Three antennal-specialized cDNA fragment from the antennae of H.armigera were obtained by differential display polymerase
引文
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