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寄生蜂与非适应性寄主之间的发育和免疫互作研究
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摘要
昆虫拥有一套虽然原始却复杂有效的免疫系统,所以当寄生蜂寄生寄主昆虫的时候,会面临非常严峻的考验。为此,寄生蜂进化产生了许多寄生因子来抑制寄主的免疫系统并操控寄主的生长发育,这些因子包括多分DNA病毒(Polydnavirus,PDV)、毒液(Venom)以及寄生蜂卵孵化时释放到寄主血腔中的畸形细胞(Teratocyte)等,它们能够为寄生蜂后代创造一种有利的生长环境。本论文以家蚕Bombyx mori幼虫作为毁侧沟茧蜂Microplitis demolitor和菜蛾盘绒茧蜂Cotesia vestalis的非适应性寄主,通过研究它们之间的发育与免疫互作关系,完善寄生蜂利用PDV调控寄主的分子机理,为PDV这一类特殊病毒与寄主之间的协同进化研究提供新的实验证据和理论依据,同时还可以加深对天敌昆虫成功控制寄主和种间竞争取胜根本原因的认识。本研究得到的结果和结论如下:
     (1)在毁侧沟茧蜂一家蚕这个寄生系统中,随着家蚕幼虫龄期的增加,它对寄生的抵抗能力也不断增加。低龄家蚕(一龄和二龄初)被寄生后大量死亡,而高龄家蚕(三龄末和四龄)几乎不受寄生的影响。解剖死亡的或将要死亡的被寄生家蚕时,发现它们体内的寄生蜂蜂卵全部孵化,而且一龄幼蜂都没有被包囊,同时在血淋巴中还发现了大量的成熟的畸形细胞,说明这些家蚕的免疫系统丧失了对寄生蜂的包囊作用。解剖不受寄生影响的家蚕时,发现寄生蜂很快就被血细胞包囊并杀死。部分被寄生的低龄家蚕还出现了龄期增加的现象,即经历了六龄才结茧化蛹。
     (2)毁侧沟茧蜂寄生家蚕幼虫后,绝大部分蜂卵的胚胎都能发育,但发育速度慢于在适应性寄主大豆夜蛾Pseudoplusia includens体内的胚胎。孵化为一龄幼蜂后,寄生蜂随即停止发育,直至被寄主血细胞包囊或与寄主一起死亡。这些结果说明家蚕体内的生理生化和营养条件可以支持毁侧沟茧蜂蜂卵的发育,但不支持毁侧沟茧蜂幼蜂的发育。
     (3)单独注射毁侧沟茧蜂多分DNA病毒MdBV(M.demolitor bracovirus)到二龄初未寄生的家蚕体内不能造成家蚕的死亡,但向二龄初被寄生的家蚕体内注射MdBV会使家蚕的死亡率从60%左右增加到100%,这说明MdBV不是被寄生家蚕死亡的直接原因,但其剂量增加可以提高被寄生家蚕的死亡率。MdBV病毒基因组DNA在家蚕体内是可以长期存在的,也可以进行转录、表达,但是病毒蛋白表达的量受到了抑制。病毒蛋白在有些血细胞中表达量很高,这些血细胞没有延展,成圆形,不能对大肠杆菌进行吞噬,说明MdBV病毒蛋白可以作用于家蚕血细胞;但病毒蛋白在有些血细胞中表达量非常低,这些血细胞有延展,可以对细菌进行吞噬,这说明在这些血细胞内MdBV病毒蛋白表达量的减少削弱了病毒对血细胞的免疫抑制作用。另外,MdBV主要感染家蚕幼虫的颗粒血细胞。
     (4)从菜蛾盘绒茧蜂多分DNA病毒CvBV(C.vestalis bracovirus)cDNA文库中筛选了两个新基因,分别命名为CvBV1和CvBV2,它们分别位于CvBV基因组片段CvBV-S5(14.5 kb)和CvBV-S51(17.5 kb)上。经软件预测它们都编码信号肽序列,可能是分泌蛋白。在同一基因组的其他片段上也发现了与它们同源的基因,说明有基因重复的现象出现。RT-PCR结果显示在寄生后0.5小时就能在寄主小菜蛾幼虫体内检测到它们的转录,并一直持续到寄生后6天。真核表达结果显示在细胞培养基中可以检测到这两个基因编码的蛋白,证实它们是分泌到胞外的病毒蛋白。
     (5)用菜蛾盘绒茧蜂寄生二龄初家蚕幼虫后,发现它们的生长发育与对照家蚕无明显差别,没有死亡现象发生,说明该蜂的寄生对家蚕的生长发育没有明显影响。在不同时间点解剖被寄生家蚕,收集寄生蜂的蜂卵,发现它们都没有孵化,胚胎发育只进行到胚带形成这个阶段,然后胚胎死亡,蜂卵被家蚕血细胞包囊。通过RT-PCR的方法在家蚕体内检测CvBV基因CvBV1、CvBV2、EP1、EP2、ORF805和Lectin的表达情况,发现CvBV2、ORF805和EP2基因都没有表达,CvBV1和Lectin基因只在寄生后1天有表达,只有EP1基因在整个过程都有表达。这说明家蚕在转录水平就对CvBV病毒基因进行了抑制。噬菌实验证明病毒对家蚕血细胞没有作用。
After parasitoids lay eggs into their hosts,their progenies will face the challenges by the efficient innate immune system of the hosts.Parasitoids have evolved an amazing array of mechanisms to manipulate the development and immune response of their hosts.Polydnavirus(PDV),venom,ovarian proteins and teratocytes are the universal regulatory factors employed by many endoparasitoids to create an environment that is favorable for the development of the parasitoid larvae.Here we used the silkworm Bombyx mori as a nonpermissive host of Microplitis demolitor and Cotesia vestalis to study the developmental and immunological interactions between parasitoids and their nonpermissive host,especially the mechanism of the effects of PDV on their host.The results and conclusions are summarized as follows:
     (1) In M.demolitor-B.mori system,B.mori larvae became increasingly resistant to the parasitism as they aged.Parasitism almost had no effect on old third or fourth instar larvae,while most parasitized first and early second instar larvae died after they molted to the next instar.Free first instar parasitoid larvae and teratocytes were found in the dying or dead B.mori larvae,which indicated that the host lost the ability of encapsulation.In non-affected B.mori larvae,the parasitoid larvae or eggs were encapsulated and killed by the hemocytes of the hosts.In addition,parasitism could induce supernumerary instars in some young B.mori larvae.
     (2) Most M.demolitor eggs could develop and hatch in B.mori larvae,although they developed more slowly than the eggs developed in the permissive host Pseudoplusia includens.After egg hatched,the first instar parasitoid larvae ceased development until they were encapsulated by the hosts or died because of the death of the hosts. These results indicated that the physiological,biochemistrical and nutrient conditions in B.mori larvae could support the development of M.demolitor eggs but not the M. demolitor larvae.
     (3) The injection of MdBV(M.demolitor bracovirus) could not cause the non-parasitized early second instar B.mori larvae to die,but could increase the death rate of the parasitized second instar B.mori larvae from 60%to 100%.MdBV DNA could persist in B.mori larvae and its genes could be expressed,but the level of expression was suppressed.The expression level of MdBV genes was very high in some hemocytes whose abilities of spreading and phagocytosis were affected;while the expression level was relatively low in other hemocytes that could still spread and phagocytize.These results indicated that MdBV could suppress the cellular immune response but this effect was diminished by the low expression level of MdBV genes.
     (4) Two new genes,CvBV1 and CVBV2,were isolated in CvBV(C.vestalis bracovirus) cDNA library.CvBV1 was located on segment CvBV-S5 with a size of 790 bp,while CvBV2 was located on segment CvBV-S51 with a size of 459 bp.Both these two genes were predicted to have an N-terminal signal peptide.Gene duplication occurred in both genes,tandem gene duplication for CvBV1 and segmental duplication for CVBV2.Gene transcripts of the two genes were detected in hosts as early as 0.5 h post-parasitization and continued to be detected for six days. These two genes were expressed in High Five cells and the recombinant proteins could be detected in the medium,which confirmed that these two proteins were secretive.
     (5) Parasitism by C.vestalis could neither affect the development of early second instar B.mori larvae obviously nor cause the larvae to die.No C.vestalis egg hatched and embryogenesis proceeded up to germband extension but thereafter development ceased.The embryos died and were encapsulated by the hemocytes of B.mori larvae. The expression of CvBV gene CvBVl,CvBV2,EP1,EP2,ORF805 and Lectin was examined by RT-PCR.,of whcih CvBV2,ORF805 and EP2 could not be detected, CvBV1 and Lectin could only be detected on 1 day post-parasitization,and only EP1 gene could be detected at all the examined time points.CvBV had no effect on the ability of phagocytosis.These results indicate that the expression of CvBV genes in B. mori larvae was inhibited and the virus had no effect on the cellular immune response of B.mori larvae.
引文
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