寄生蜂调控寄主害虫免疫与发育机理的研究新进展

寄生蜂是重要且种类最为丰富的膜翅目昆虫类群之一,也是极具价值的害虫生物控制因子。寄生蜂携带有不同类型的活性因子,包括毒液、多分DNA病毒类病毒颗粒、卵巢蛋白、畸形细胞及幼虫分泌物等,用于调控寄主害虫的免疫反应、发育等重要生理过程,以确保成功寄生并确保其子代在寄主害虫体内(内寄生蜂)或体表(外寄生蜂)正常发育,最终可导致寄主害虫死亡,从而有效控制寄主害虫种群数量。目前已有诸多与寄生蜂调控寄主害虫内在机理相关的研究报道,该领域也已成为昆虫寄生学与生理学的研究热点之一。本文仅从寄生蜂寄生因子多样性、寄生蜂调控寄主害虫免疫及发育的机理等方面,对相关的最新研究进展作一概述。     

寄生蜂多分DNA病毒对寄主昆虫的免疫抑制作用

多分DNA病毒（PDV）是在膜翅目姬蜂科和茧蜂科寄生蜂体内的一类很独特的病毒,寄生蜂产卵时,PDV随同卵和萼液一起被注射入寄主体内,能干扰寄主的细胞免疫和体液免疫,该病毒直接侵染或间接作用于血细胞,主要是浆细胞和颗粒细胞,导致血细胞变圆或凋亡,PDV也能抑制血淋巴酚氧化酶活性,诱导抗菌因子的大量合成,最近有关研究主要集中在病毒基因的表达和伴随寄主血细胞功能失常的分子事件上,一些寄主蜂的PDV与其他因子,如卵巢蛋白,畸形细胞或蜂毒等协同发挥作用。     

中国寄生蜂研究及其在害虫生物防治中的应用

寄生蜂是一类重要的寄生性天敌昆虫,种类繁多、习性复杂,在害虫生物防治和综合治理中发挥着极其重要的作用。在产卵时,寄生蜂携带的毒液、多DNA病毒等寄生因子就会随之进入寄主体内,发挥调控寄主生长、发育、免疫、代谢、行为的作用,从而保障了寄生蜂后代的发育。本文主要针对我国寄生蜂的系统分类、资源普查、生物学、生态学、寄主调控、人工繁殖、释放应用、田间保护和助增等方面的基础研究和应用进行了概述和整理。     

昆虫寄主标记信息素

综述了昆虫寄主标记信息素的研究进展。昆虫寄主标记信息素是指由昆虫产生的用来标记寄主上有同种个体存在的化学物质。昆虫寄主标记信息素的主要生态学功能是调节昆虫的产卵行为,通过阻止自身或同种其它个体对已标记寄主的产卵选择,或减少产卵量来减少后代之间对寄主资源的竞争。寄主标记信息素也会给释放着带来不利的影响,如信息盗用和盗寄生现象等。昆虫寄主标记信息素也调节昆虫近缘种之间对共同寄主资源的竞争。近缘种昆虫对相互寄主标记信息素识别能力的差异反映了不同昆虫对同一寄主资源竞争能力的强弱。寄主标记信息素产生和贮存的部位一般与外分泌腺、消化系统或生殖系统相联系,杜氏腺、毒腺、前胸腺、腹腺、下唇腺、后产卵管、卵巢、中肠和后肠等是产生或贮存寄主标记信息素的常见部位。产生的寄主标记信息素一般在成虫产卵时由产卵器、口器或排泄口释放到寄主体内或体表。卵寄生蜂的寄主标记信息素一般标记在寄主的体表,雌成蜂用触角检测;其它寄生蜂的寄主标记信息素常产在寄主体内,用产卵器检测;植食性昆虫的寄主标记信息素只产在寄主表面,用触角或产卵器检测。昆虫的产卵器、口器、触角或跗节上着生有感受寄主标记信息素的化感器,可以检测到标记在寄主体内或体表的寄主标记信息素。昆虫寄主标记信息素的完全定性涉及活性化合物的分离、鉴定、合成以及行为测定等,已有几种昆虫的寄主标记信息素成分得到了分离鉴定。     

二种寄生蜂寄生对不同虫龄小菜蛾精巢的影响

为探明寄生蜂引起寄主寄生性去势的机制,本文选取携带不同多分DNA病毒(Polydnavirus,PDV)的2种内寄生蜂与共同寄主小菜蛾Plutella xylostella(L.)为寄生体系,研究不同虫龄小菜蛾被寄生后雄性生精细胞、精子束形态和精巢发育体积变化,系统比较寄生性去势程度和分别拥有2类PDV的寄生蜂在寄主精子发生和形成过程中的作用。结果表明:携带Bracovirus PDV的菜蛾盘绒茧蜂Cotesiave stalis(Haliday)或拥有Ichvovirus PDV的半闭弯尾姬蜂Diadegma semiclausum Hellén寄生对不同虫龄小菜蛾的精子发生和形成过程均产生明显的抑制作用,表现为不能产生精子束或精子束数量减少,但抑制程度以寄生低龄寄主时最明显。2种蜂寄生均能抑制小菜蛾精巢体积的增长,但对低龄寄主的抑制程度明显强于高龄寄主,寄生性去势程度取决于寄生时寄主虫龄。相比而言,寄生不同虫龄小菜蛾时,茧蜂引起小菜蛾寄生性去势的程度均强于姬蜂。     

寄生蜂携带的多DNA病毒的起源及其特性

病毒在通常意义上都是有害的寄生生物,但是有一类特殊的多DNA病毒(PDV)却成为在数量庞大的寄生蜂的生活史中不可或缺的共生病毒.PDV随着寄生蜂的卵一起被注入到寄主的体内,调控寄主的免疫和发育生理,从而确保幼蜂在寄主体内生存发育.PDV的基因组整合在寄生蜂的基因组内,因此PDV病毒粒子的形成并非在寄主受到侵染的组织内,而是位于寄生蜂卵巢的特定细胞(卵萼细胞)中,其编码的蛋白质几乎没有病毒结构性蛋白,主要的表达产物都用于调控寄主的生理活动.PDV基因编码序列比对分析结果显示,茧蜂病毒(BV)与致病性的裸病毒nudivirus和杆状病毒baculovirus存在一定的亲缘关系,然而这些同源基因已高度分化,具有不同的生物学特性,同时也不能确定这些基因是否还具有原始的功能.在寄生蜂基因组中尚有许多与裸病毒和杆状病毒类似的基因,其中一部分基因能够编码BV病毒粒子的结构蛋白,其表达模式亦与PDV病毒粒子的形成有关.由于PDV使用了与这2种病毒类似的传播途径,才使得寄生蜂的PDV基因转入寄主体内发挥作用,虽然这不能说明PDV属于这2种病毒,至少存在PDV从这2种病毒发展而来的可能性.     

寄生蜂畸形细胞特性及与蜂幼虫生长发育的关系

本文综述寄生蜂畸形细胞特性及其功能研究的最新进展。一些进化高等的膜翅目内寄生蜂,在胚胎发育过程中,包裹在胚胎周围的浆膜会随着蜂卵的孵化,以游离的细胞团或单个细胞的形式释放到寄主昆虫的血淋巴中。这些源自浆膜被称作“畸形细胞（Teratocyte）”的特殊类群,在伴随蜂幼虫的生长发育过程中,自身也经历一系列结构和功能的改变。它们主要通过分泌蛋白质或酶类,改变寄主的物质代谢途径等方式为发育中的蜂幼虫提供营养和能量。对寄生蜂重要的调控因子一畸形细胞的特性及与蜂幼虫相互关系的揭示,可以阐明寄生蜂一寄主昆虫协同进化的内在机理,也可为开发害虫生物防治新策略提供新的思路和途径。     

基于组学的寄生蜂与寄主免疫的相互作用

寄生蜂作为天敌昆虫,在害虫生物防治中起着极其重要的作用。现代分子生物学和第二代测序技术的迅速发展,为筛选寄生蜂与寄主免疫互作的关键分子,阐明寄生蜂逃避或抑制寄主免疫系统攻击的机理提供了新的机遇。昆虫的天然免疫可分为细胞免疫和体液免疫,寄生蜂在与寄主长期的协同进化过程中,依靠毒液(venom)、多分DNA病毒(PDV)、类病毒颗粒(VLP)、畸形细胞(teratocyte)等手段,一方面调控寄主血细胞的增殖和分化,抑制细胞包囊反应,从而破坏寄主的细胞免疫系统;另一方面,通过抑制酚氧化酶原激活干扰寄主的体液免疫,从而逃避黑化反应的攻击。寄生蜂在发育过程中也有相应的免疫防御机制。本文从细胞免疫、体液黑化反应以及免疫信号通路等方面,综述了近年来利用组学手段在寄生蜂调控寄主免疫方面取得的最新进展,以期为阐明天然免疫的分子机制、探索害虫防治新方法提供借鉴。     

基于IBM DB2的中国寄生性膜翅目昆虫资源数据库构建

以IBM DB2数据库作为开发平台,根据寄生性膜翅目昆虫数据的特点组建了分类地位表、寄主表、形态特征图和生物学特性表,将寄生性膜翅目的中文名、拉丁名、定名人、寄主、形态特征、图形、生物学特性等信息分别存储于这4个表中,构建出中国寄生性膜翅目昆虫资源数据库。用Visual Basic 6.0编制了友好的人机对话界面,即信息的录入和修改界面、信息查询界面和用户管理界面,使数据库的应用与维护更加便捷。     

多寄主型寄生性天敌昆虫的寄主适应性及其影响因素

寄生性天敌昆虫对不同寄主资源的适应能力是其存活和繁殖的必要条件,它们的寄主选择行为则是其重要的适应结果。多寄主型寄生性天敌昆虫虽然对某种特定寄主资源的利用效率可能不如单寄主型寄生蜂,但却有利于拓展更宽的寄主范围,因此对环境的适应能力更强,更容易在自然界维持其种群的生存。统计结果也表明多寄主型天敌的生物防治效果往往比专食性天敌更高。有时生物防治成功的关键可能并不在于所利用的天敌种类的不同,而在于天敌的不同生物型或地理宗。总结了多寄主型寄生性天敌昆虫的寄主适应能力及其影响因素。寄生性天敌昆虫不仅因地理隔离产生种群分化,也可能因寄生不同的寄主产生种群内的分化,从而更加适应寄主的生活特性和栖境条件。寄主种类、寄主发育阶段、寄主大小、寄主营养、寄主免疫反应、寄主逃避反应、其它天敌的竞争、寄主共生或共栖生物的存在、寄主植物、天敌自身的学习能力及其共生微生物等多种因素对寄生性天敌昆虫的寄主适应性可产生影响。展望了多寄主型寄生性天敌昆虫对新寄主资源的拓展利用能力和适应性在生产上的可能应用前景和途径,以期为明确天敌与寄主间的互作关系,人工驯化寄生性天敌昆虫增强对靶标害虫的控制作用,合理利用天敌提高生物防治效率提供新的思路和理论支持。     

Parasitic wasp assemblages associated with native and weedy plant species in an agricultural landscape

Abstract  The establishment and maintenance of suitable habitat on-farm or in the surrounding landscape can enhance the survival of beneficial parasitic Hymenoptera, thus improving the control of pest species. Both endemic and weedy non-crop plant species across a highly modified agricultural landscape supported species-rich and abundant parasitic wasp assemblages with diverse biology and host associations. It was also shown that isolated, recently planted, single-species stands of plants can rapidly accumulate diverse assemblages of parasitoids. Chalcidoidea was the most species-rich and abundant group, egg and larval parasitoids were the most speciose and abundant guilds, and parasitoids of herbivorous insects feeding on and inside plant tissue were the most species-rich and abundant functional groups. The hymenopteran assemblages associated with the majority of plant species were dominated by three parasitoid species: a Trichogrammatidae, a Scelionidae ( Telenomus sp.) and a Eulophidae ( Ceranisus sp.), all genera that contain many important biocontrol agents of pest Lepidoptera, Hemiptera and Thysanoptera. Results show that both native and weedy plant species may potentially provide an important reservoir of mobile parasitic wasps of benefit to crop protection.     

Biological control of tortricidae in tea fields in Japan using insect viruses and parasitoids

Tea is a perennial and evergreen plant. Cultivated tea trees provide a habitat for insect pests and their natural enemies. In Japan, granuloviruses (GVs) have successfully controlled two of the most important pests of tea, Adoxophyes honmai and Homona magnanima (Tortricidae: Lepidoptera). The GVs are produced in vivo and a single application sustains pesticidal efficacy throughout a year, which encompasses 4 to 5 discrete generations of both species. A. honmai and H. magnanima also have various natural enemies, especially hymenopteran parasitoids. Such resident natural enemies also play a role in reducing the pest density in virus-controlled fields, but the effect of virus infection on parasitoids sharing the same host larva has not been well studied. Survival of one of the major parasitoids ofA. honmai, Ascogaster reticulata (Braconidae: Hymenoptera), is reduced by virus infection of the host. Viruses, including GV and entomopoxvirus (EPV), and certain koinobiont endoparasitoids, including A. reticulata, are both known to regulate host endocrinology. However, the GV and EPV have distinct host regulation mechanisms, and consequently have different impacts on the survival of A. retuculata, when A. reticulata parasitizes a host that is infected with either GV or EPV. These additional effects on host regulation displayed by both viruses and parasitoids affect the outcome of virus-parasitoid interactions.     

寄生蜂寄主选择行为研究进展

寄生蜂是膜翅目昆虫中的重要类群,在自然界种类多,数量大。据文献报道全世界膜翅目昆虫大约有10万种”’,其中许多种类与农林害虫长期保持一定的制约关系,在害虫防治中占有重要地位。由于大量使用化学农药导致害虫的抗药性增加,利用天敌昆虫来防治害虫越来越显示出其重要性和优越性。但是,直到近20年来,寄生蜂的寄生行为才受到重视,并给予了广泛的、深入的研究”－‘’严格地讲,寄生蜂应该称为拟寄生蜂,它与通常所说的寄生昆虫像虱子、臭虫等有明显不同,主要区别在于：1．寄生仅仅发生在未成熟阶段,而成虫阶段营自由生活;2．寄…     

Behavioral mechanisms of parasitic wasps for searching concealed insect hosts

Wood borers are important forest insect pests and difficult to be controlled owing to their concealed behavior. However, parasitic wasps can effectively ascertain and parasitize wood borers as well as other concealed pests by using special searching, finding and attacking mechanisms, which have been developed during the course of long-term coevolution with their hosts. The present paper summarizes the behavioral mechanisms of parasitic wasps involved in searching and locating their concealed hosts. Parasitic wasps can accurately find the location of their hidden hosts and then parasitize them, usually by using olfactory semiochemicals from hosts (lavare and adults), host frass and symbiotic microorganisms in host galleries; visual signals from color contrast of plant surface; contact stimuli from characters of host physical defense; substrate vibrations produced by host feeding and movement; infrared radiation from host activities and metabolizability. Some parasitic wasps may integrate the information of several stimuli from different sources to enhance the reliability and accuracy of host locations. In addition, the potentials for utilizing the host location signals of parasitoids in biological control are discussed.     

寄生蜂寻找隐蔽性寄主害虫的行为机制

林木蛀干类害虫具有高度的隐蔽性,是林业上的一类重要害虫,也是目前世界上最难防治的害虫类群之一.寄生蜂在与寄主长期的协同进化过程中,形成了搜索、发现和攻击寄主害虫的独特机制,能够有效地找到并寄生它们.总结了寄生性天敌寻找、发现并成功定位隐蔽性寄主害虫的行为学机制.寄生蜂可以利用来自嗅觉的化学信息物质(如寄主、寄主粪便、虫道共生菌的挥发性气味)、寄主成虫的化学通讯物质、来自视觉的植物表面色差信息、来自触觉的寄主保护物性状特征、来自寄主取食和运动所产生的介质振动信号以及来自寄主活动和代谢的红外辐射等多种途径有效地发现隐蔽性害虫的位置,从而完成寄生行为.有些寄生蜂还能综合利用来源不同的多种信息,从而提高寄主定位的可靠性和准确性.本文还对寄生蜂寻找寄主的这些线索在生物防治上可能的利用途径和前景进行了讨论.这对促进我国在该领域的研究,充分利用天敌昆虫,提高生物防治效率具有参考价值.     

拟寄生蜂搜索产卵过程中对寄主的竞争

综述拟寄生蜂搜索产卵过程中对寄主竞争的最新研究进展.这类竞争具有四种方式,即标记寄主、杀卵和杀幼、守护寄主和捕食寄主.（1）标记寄主常涉及寄主标记信息素,这是由雌蜂在产卵前、产卵时或产卵后分泌的化学物质.寄主标记信息素常介导拟寄生蜂对已寄生和健康寄主的辨别、减少过寄生和多寄生、减轻种内和种间竞争压力.（2）雌蜂遇到已寄生寄主时,很多种类杀死前一雌蜂遗留的卵和幼虫,再产下自己的卵.雌蜂使用三种方法杀卵和杀幼,即产卵器穿刺、取食和使用有毒物质.通过杀卵和杀幼,产卵雌蜂清除了前一雌蜂遗留的后代,主动改善了寄主品质,从而有利于自身后代的生存.（3）守护寄主在肿腿蜂科、缘腹细蜂科、金小蜂科、缨小蜂科和茧蜂科中均有报道,守护者驱逐入侵者以保护后代及健康寄主.（4）捕食寄主不仅减少了健康寄主数量,且直接导致已寄生寄主中拟寄生蜂卵和幼虫的死亡.雌蜂一般在体内成熟卵量较少时捕食寄主.讨论了研究拟寄生蜂搜索产卵过程中竞争寄主的理论意义和实际应用价值.     

The parasitoid factor in the virulence and spread of lepidopteran baculoviruses

Insect parasitoids and baculoviruses play important roles in the natural and strategic biological control of insects. The two parasites are frequent competitors within common hosts and much research has focused on the negative impact that baculoviral host infections have on parasitoids. This review summarizes the impacts that parasitoids may have on the virulence and spread of lepidopteran baculoviruses. By changing host behavior and development, parasitoids have been shown to decrease baculovirus virulence and productivity within parasitized baculovirus-susceptible hosts; however, studies of the tools used by hymenopteran parasitoids to overcome their hosts' immune systems, suggest that parasitoids may, in some cases, facilitate baculoviral infections in less susceptible hosts. Laboratory and field research have demonstrated that parasitoids can mechanically transmit bacuioviruses between insects, and in this way, increase the efficacy of the viruses. Instances of new, more virulent isolates of baculoviruses have been recorded from specifically parasitoid-targeted hosts suggesting other possible benefits from the transmission or activation of baculoviruses by parasitoids.     

Histories of host shifts and cospeciation among free‐living parasitoids of Rhagoletis flies

Host shifts by specialist insects can lead to reproductive isolation between insect populations that use different hosts, promoting diversification. When both a phytophagous insect and its ancestrally associated parasitoid shift to the same novel host plant, they may cospeciate. However, because adult parasitoids are free living, they can also colonize novel host insects and diversify independent of their ancestral host insect. Although shifts of parasitoids to new insect hosts have been documented in ecological time, the long‐term importance of such shifts to parasitoid diversity has not been evaluated. We used a genus of flies with a history of speciation via host shifting (Rhagoletis [Diptera: Tephritidae]) and three associated hymenopteran parasitoid genera (Diachasma, Coptera and Utetes) to examine cophylogenetic relationships between parasitoids and their host insects. We inferred phylogenies of Rhagoletis, Diachasma, Coptera and Utetes and used distance‐based cophylogenetic methods (ParaFit and PACo) to assess congruence between fly and parasitoid trees. We used an event‐based method with a free‐living parasitoid cost model to reconstruct cophylogenetic histories of each parasitoid genus and Rhagoletis. We found that the current species diversity and host–parasitoid associations between the Rhagoletis flies and parasitoids are the primary result of ancient cospeciation events. Parasitoid shifts to ancestrally unrelated hosts primarily occur near the branch tips, suggesting that host shifts contribute to recent parasitoid species diversity but that these lineages may not persist over longer time periods. Our analyses also stress the importance of biologically informed cost models when investigating the coevolutionary histories of hosts and free‐living parasitoids.     

寄生蜂取食寄主特性及其在害虫生物防治中的作用

许多寄生性天敌昆虫的雌虫不仅寄生寄主, 而且还能取食寄主。在卵育型（synovigenic）寄生蜂类群中, 取食寄主行为是较为普遍的现象。本文综合近20年相关研究进展, 从寄生蜂类群、取食类型、生态学意义及影响因子等方面对寄生蜂的取食寄主行为进行了归纳总结。寄生蜂通过取食不仅可以杀死寄主, 直接起到控制害虫种群数量的作用, 还能通过取食策略为卵的成熟和再生提供营养来源, 对延长雌虫的寿命也有一定的帮助。对取食寄主行为的了解可为筛选优势寄生性天敌种类、评估寄生蜂在害虫生物防治中的作用提供重要信息。