共生菌Wolbachia引起宿主细胞质不亲和的研究进展

Wolbachia 是一类广泛存在于节肢动物以及线虫体内细胞质中呈母系遗传的共生细菌,能够在宿主中产生细胞质不亲和、孤雌生殖、雌性化及杀雄等多种生殖调控作用,其中细胞质不亲和是指被 Wolbachia 感染的雄性个体与未感染的雌性个体（单向不亲和）,或者感染不同株系 Wolbachia 的雌性个体（双向不亲和）交配后不能或很少产生后代,或者后代偏雄性的现象。细胞质不亲和作用使感染的雌性个体在种群中具有很大的生殖优势,凭借这种生殖优势,Wolbachia 能够迅速在宿主种群中扩张。细胞质不亲和的机理探索主要集中在细胞学水平上,其中广为接受的精子“修饰”和“拯救”理论认为,精巢中的 Wolbachia 能够修饰宿主的精细胞,使其不能和卵细胞正常融合,但是当母本感染相同的 Wolbachia 时,就能够将“修饰”过的精子细胞“拯救”过来,使其恢复与卵细胞的正常融合。而分子机理上的探索也开始在转录组、基因组和miRNA水平上对部分昆虫展开了研究。影响细胞质不亲和的因素有很多,包括宿主遗传背景、 Wolbachia 株系、Wolbachia 基因型、共生菌密度（浓度、滴度）、雄虫年龄、环境因素以及共生菌在宿主生殖组织的分布等。近年来,人类也应用细胞质不亲和控制害虫（主要是蚊虫）和人类疾病,取得了较好的进展。     

节肢动物内共生菌Wolbachia的研究进展

沃尔巴克氏体Wolbachia为母系传播的胞内共生菌,可通过对宿主产生多种调控方式扩大其自身在宿主种群的传播。据推测,有40%~60%的节肢动物都感染有Wolbachia,并可根据不同株系间的系统发育关系将其分为多个超群。为了有助于深入研究Wolbachia对其宿主的调控方式及其调控机制及提出更为有效的害虫生物防治策略,本文综述了节肢动物内共生菌Wolbachia的研究现状。1924年Wolbachia被报道首次发现于尖音库蚊Culex pipiens的生殖组织中,1971年确认其与宿主的胞质不亲和现象有关。Wolbachia可以通过胞质不亲和、杀雄、雌性化、孤雌生殖等作用方式调控宿主的生殖。除生殖调控之外,Wolbachia对宿主的调控方式还包括调控宿主新陈代谢、抵制病原菌、影响宿主生殖力等。Wolbachia调控的胞质不亲和现象可用“修饰-营救”(modification-rescue)模型解释,且已有与Wolbachia诱导宿主胞质不亲和相关的功能基因被报道。wMel株系是首个公布全基因组序列的Wolbachia株系,随后又有数十种不同株系的Wolbachia基因组陆续被破译。wMel株系Wolbachia可起到抑制登革热病毒传播的作用;同时,Wolbachia和昆虫不育技术的结合对白纹伊蚊Aedes albopictus野外种群起到良好的控制效果。鉴于目前节肢动物内共生菌Wolbachia的研究现状,我们认为未来应开展以下研究：(1)Wolbachia基因组及生殖调控作用关键功能基因的研究;(2)Wolbachia与宿主间互作机制的研究;(3)Wolbachia在生物防治方面的应用。     

节肢动物内共生菌Wolbachia的研究进展

沃尔巴克氏体Wolbachia为母系传播的胞内共生菌,可通过对宿主产生多种调控方式扩大其自身在宿主种群的传播。据推测,有40%～60%的节肢动物都感染有Wolbachia,并可根据不同株系间的系统发育关系将其分为多个超群。为了有助于深入研究Wolbachia对其宿主的调控方式及其调控机制及提出更为有效的害虫生物防治策略,本文综述了节肢动物内共生菌Wolbachia的研究现状。1924年Wolbachia被报道首次发现于尖音库蚊Culex pipiens的生殖组织中,1971年确认其与宿主的胞质不亲和现象有关。Wolbachia可以通过胞质不亲和、杀雄、雌性化、孤雌生殖等作用方式调控宿主的生殖。除生殖调控之外,Wolbachia对宿主的调控方式还包括调控宿主新陈代谢、抵制病原菌、影响宿主生殖力等。Wolbachia调控的胞质不亲和现象可用"修饰-营救"(modification-rescue)模型解释,且已有与Wolbachia诱导宿主胞质不亲和相关的功能基因被报道。wMel株系是首个公布全基因组序列的Wolbachia株系,随后又有数十种不同株系的Wolbachia基因组陆续被破译。wMel株系Wolbachia可起到抑制登革热病毒传播的作用;同时,Wolbachia和昆虫不育技术的结合对白纹伊蚊Aedes albopictus野外种群起到良好的控制效果。鉴于目前节肢动物内共生菌Wolbachia的研究现状,我们认为未来应开展以下研究:(1)Wolbachia基因组及生殖调控作用关键功能基因的研究;(2)Wolbachia与宿主间互作机制的研究;(3)Wolbachia在生物防治方面的应用。     

Wolbachia与昆虫精卵细胞质不亲和

Wolbachia是广泛分布在昆虫体内的一类共生菌,能通过多种机制调节宿主的生殖方式,包括诱导宿主精卵细胞质不亲和(CI)、孤雌生殖、雌性化、杀雄等,其中细胞质不亲和为最普遍的表型,即感染Wolbachia的雄性和未感染或感染不同品系Wolbachia的雌性宿主交配后,受精卵不能正常发育,在胚胎期死亡。多数CI胚胎在第1次分裂时,来自父本的染色质浓缩缺陷,导致父本遗传物质无法正常分配到子细胞中,因而引起胚胎死亡。守门员模型认为,产生CI可能需要有两种因子,其中之一使得精子发生修饰改变,导致受精后雄性原核发育滞后。第2种因子可能与Wolbachia的原噬菌体有关,在胚胎发育后期导致胚胎死亡。近期的研究已发现,在Wolbachia感染的宿主中,一些与生殖细胞发生和繁殖相关基因的表达发生了显著改变,Wolbachia可能因此对宿主的生殖产生重大影响,进而导致CI的产生。本文主要综述了CI的细胞学表型、解释CI的模型及其分子机理,向读者展示一个小小的细菌是如何通过精妙的策略影响昆虫宿主的繁殖,从而实现其自身的生存和传播的。     

果蝇细胞内共生菌Wolbachia研究进展

Wolbachia是一类存在于包括绝大多数昆虫在内的节肢动物以及线虫体内的细胞内共生菌,通过母系传递给子代。该菌能以多种方式调控宿主的生殖活动,包括诱导细胞质不亲和、孤雌生殖、杀雄、雌性化和增加雌性生殖力。本文对Wolbachia的基础生物学及其分布、以及该菌对宿主果蝇的影响等进行了评述,并对与果蝇相关的Wolbachia的研究现状及趋势进行了讨论。     

Wolbachia引起黑腹果蝇产生“修饰-挽救”因子的初步探究

沃尔巴克氏体(Wolbachia)是一种广泛分布于昆虫体内的共生细菌,对宿主的生殖具有多种调控作用,其中,最常见的调控方式称为细胞质不亲和(Cytoplasmic Incompatibility,CI),即感染了Wolbachia的雄性宿主与未感染或感染不同品系Wolbachia的雌性宿主交配后,胚胎在发育早期死亡。关于CI产生的原因,目前人们认为是一种"精子修饰-卵子挽救"机制,本研究初步探索了黑腹果蝇中参与"修饰-挽救"机制的重要因子。本研究采用定量RT-PCR方法,检测了3种基因:yem-α(yemanuclein)、uif(uninflatable)、Prosα4T1(proteasomeα4 subunit,Testis-specific 1)在感染和未感染果蝇中的表达差异。结果发现,Wolbachia感染能够显著上调雌蝇中yem-α和uif的表达,并且显著下调雄蝇中yem-α的表达。尽管Prosα4T1在雄性果蝇体内特异表达,但是Wolbachia感染对Prosα4T1表达无明显影响。因此,yem-α可能是Wolbachia产生"修饰-挽救"机制的重要因子之一,Wolbachia可能通过调控果蝇体内yem-α的表达,影响卵子发生和精子发生进程,导致CI的产生。     

Wolbachia在熊蜂中的双重感染

Wolbachia是一类广泛存在于节肢动物体内细胞质遗传的细菌,它可以通过诱导产雌孤雌生殖、引起细胞质不亲和、遗传雄性的雌性化、雄性致死和增强生殖力等作用方式引起其寄主生殖行为的改变.本文以16S rDNA为标记检测了3种熊蜂不同组织(头,胸,足,卵巢或雄外生殖器)的Wolbachia感染.其中明亮熊蜂Bombus l...     

寄生蜂体内共生菌Wolbachia研究进展

Wolbachia是广泛分布在节肢动物生殖组织中的细胞质遗传细菌,在寄生蜂中可以通过诱导胞质不亲和、孤雌生殖、雄性雌性化和杀雄等多种方式调节寄主的生殖.本文在综述了寄生蜂体内共生菌Wolbachia的类群和分化、传播方式及其对寄主生物学特性的影响、温度通过对Wolbachia的作用而影响寄生蜂的表现等基础上,讨论了如何...     

Wolbachia属共生菌及其对节肢动物宿主适合度的影响

Wolbachia是广泛分布于节肢动物体生殖组织内呈母质遗传的一类共生细菌。近30多年来,大量的研究主要集中于Wolbachia对宿主生殖方式的调控方面;近年来的研究发现,Wolbachia对节肢动物宿主的适合度具有不同程度的影响。现对Wolbachia的宿主分布、存在部位及其对节肢动物宿主种群适合度的影响等方面进行了综述,探讨了Wolbachia在该领域的研究意义和潜在的应用价值。     

昆虫共生细菌Wolbachia的研究进展

Wolbachia是广泛分布在节肢动物生殖组织的一类细胞质遗传的细菌,它们能通过多种机制调节寄生的生殖,包括诱导细胞质不亲和,孤雌生殖和遗传上的雄性雌性化,目前对这些微生物的研究已取了可喜的进展,通过主要综述Wolbachia的分布,功能,系统发育,多样性及时空动态,也讨论了将来的研究方向     

Wolbachia infections are distributed throughout insect somatic and germ line tissues

Wolbachia are intracellular microorganisms that form maternally-inherited infections within numerous arthropod species. These bacteria have drawn much attention, due in part to the reproductive alterations that they induce in their hosts including cytoplasmic incompatibility (CI), feminization and parthenogenesis. Although Wolbachia's presence within insect reproductive tissues has been well described, relatively few studies have examined the extent to which Wolbachia infects other tissues. We have examined Wolbachia tissue tropism in a number of representative insect hosts by western blot, dot blot hybridization and diagnostic PCR. Results from these studies indicate that Wolbachia are much more widely distributed in host tissues than previously appreciated. Furthermore, the distribution of Wolbachia in somatic tissues varied between different Wolbachia/host associations. Some associations showed Wolbachia disseminated throughout most tissues while others appeared to be much more restricted, being predominantly limited to the reproductive tissues. We discuss the relevance of these infection patterns to the evolution of Wolbachia/host symbioses and to potential applied uses of Wolbachia.     

Wolbachia属共生细菌及其对节肢动物生殖活动的调控作用

Wolbachia属是广泛分布于节肢动物生殖组织内的一类共生细菌。这些共生菌通过卵的细胞质传播并参与多种调控其宿主生殖活动的机制,包括:诱导生殖不亲和、诱导孤雌生殖、雌性化、雄性致死和调节繁殖力。Wolbachia被认为与性别决定、共生关系和物种形成等重要生物学问题密切相关,是探索这些研究领域的新线索。而且Wolbachia可作为特定的载体对其宿主种群进行遗传调控,如增强寄生蜂在害虫生物防治中的作用,控制线虫引起的疾病传播。该文综述了Wolbachia的形态学及存在部位、基因组结构、系统发育、种的命名、水平传递和Wolbachia对其宿主生殖活动的调控作用,并分析了Wolbachia研究的科学意义和发展趋势,以期引起我国生物学家对Wolbachia研究的注意和快速切入。     

Modes of acquisition of Wolbachia: horizontal transfer, hybrid introgression, and codivergence in the Nasonia species complex

Wolbachia are maternally inherited bacteria that infect a large number of insects and are responsible for different reproductive alterations of their hosts. One of the key features of Wolbachia biology is its ability to move within and between host species, which contributes to the impressive diversity and range of infected hosts. Using multiple Wolbachia genes, including five developed for Multi-Locus Sequence Typing (MLST), the diversity and modes of movement of Wolbachia within the wasp genus Nasonia were investigated. Eleven different Wolbachia were found in the four species of Nasonia , including five newly identified infections. Five infections were acquired by horizontal transmission from other insect taxa, three have been acquired by hybridization between two Nasonia species, which resulted in a mitochondrial- Wolbachia sweep from one species to the other, and at least three have codiverged during speciation of their hosts. The results show that a variety of transfer mechanisms of Wolbachia are possible even within a single host genus. Codivergence of Wolbachia and their hosts is uncommon and provides a rare opportunity to investigate long-term Wolbachia evolution within a host lineage. Using synonymous divergence among codiverging infections and host nuclear genes, we estimate Wolbachia mutation rates to be approximately one-third that of the nuclear genome.     

Wolbachia affects oviposition and mating behaviour of its spider mite host

Wolbachia bacteria are transmitted from mother to offspring via the cytoplasm of the egg. When mated to males infected with Wolbachia bacteria, uninfected females produce unviable offspring, a phenomenon called cytoplasmic incompatibility (CI). Current theory predicts that ‘sterilization’ of uninfected females by infected males confers a fitness advantage to Wolbachia in infected females. When the infection is above a threshold frequency in a panmictic population, CI reduces the fitness of uninfected females below that of infected females and, consequently, the proportion of infected hosts increases. CI is a mechanism that benefits the bacteria but, apparently, not the host. The host could benefit from avoiding incompatible mates. Parasite load and disease resistance are known to be involved in mate choice. Can Wolbachia also be implicated in reproductive behaviour? We used the two‐spotted spider mite – Wolbachia symbiosis to address this question. Our results suggest that uninfected females preferably mate to uninfected males while infected females aggregate their offspring, thereby promoting sib mating. Our data agrees with other results that hosts of Wolbachia do not necessarily behave as innocent bystanders – host mechanisms that avoid CI can evolve.     

Wolbachia and reproductive conflict in Exorista sorbillans

Many arthropods harbour endosymbiotic bacteria of the genus Wolbachia. These endosymbionts are transmitted vertically from one generation to the next and are obligatory in several Dipterans that have been studied to date. These bacteria induce an array of reproductive isolation mechanisms that are implicated in pest management to evolutionary biology of respective hosts. The uzifly, Exorista sorbillans, a tachinid endoparasitoid of the silkworm, Bombyx mori L. (Lepidoptera: Bombycidae), causes enormous losses to the silk industry; now it is known that it harbours Wolbachia endobacteria. The elimination of Wolbachia by antibiotics interrupts embryogenesis and causes various reproductive conflicts such as (1) a reduction of fecundity of uninfected female, (2) cytoplasmic incompatibility in the uninfected females crossed with infected males, (3) genomic incompatibility in crosses between males and females from uninfected population, and (4) sex-ratio distortion in uninfected females irrespective of the presence of Wolbachia in males. These results suggest that the relationship of Wolbachia with its uzifly host is one of mutual symbiosis as it controls the reproductive physiology of its host.     

The effect of Wolbachia-induced cytoplasmic incompatibility on host population size in natural and manipulated systems

Obligate, intracellular bacteria of the genus Wolbachia often behave as reproductive parasites by manipulating host reproduction to enhance their vertical transmission. One of these reproductive manipulations, cytoplasmic incompatibility, causes a reduction in egg-hatch rate in crosses between individuals with differing infections. Applied strategies based upon cytoplasmic incompatibility have been proposed for both the suppression and replacement of host populations. As Wolbachia infections occur within a broad range of invertebrates, these strategies are potentially applicable to a variety of medically and economically important insects. Here, we examine the interaction between Wolbachia infection frequency and host population size. We use a model to describe natural invasions of Wolbachia infections, artificial releases of infected hosts and releases of sterile males, as part of a traditional sterile insect technique programme. Model simulations demonstrate the importance of understanding the reproductive rate and intraspecific competition type of the targeted population, showing that releases of sterile or incompatible individuals may cause an undesired increase in the adult number. In addition, the model suggests a novel applied strategy that employs Wolbachia infections to suppress host populations. Releases of Wolbachia-infected hosts can be used to sustain artificially an unstable coexistence of multiple incompatible infections within a host population, allowing the host population size to be reduced, maintained at low levels, or eliminated.     

Evidence for widespread Wolbachia infection in isopod crustaceans: molecular identification and host feminization.

Wolbachia are maternally inherited, intracellular, alpha proteobacteria that infect a wide range of arthropods. They cause three kinds of reproductive alterations in their hosts: cytoplasmic incompatibility, parthenogenesis and feminization. There have been many studies of the distribution of Wolbachia in arthropods, but very few crustacean species are known to be infected. We investigated the prevalence of Wolbachia in 85 species from five crustacean orders. Twenty-two isopod species were found to carry these bacteria. The bacteria were found mainly in terrestrial species, suggesting that Wolbachia came from a continental environment. The evolutionary relationships between these Wolbachia strains were determined by sequencing bacterial genes and by interspecific transfers. All the bacteria associated with isopods belonged to the Wolbachia B group, based on 16S rDNA sequence data. All the terrestrial isopod symbionts in this group except one formed an independent clade. The results of interspecific transfers show evidence of specialization of Wolbachia symbionts to their isopod hosts. They also suggest that host species plays a more important role than bacterial phylogeny in determining the phenotype induced by Wolbachia infection.     

Wolbachia对赤眼蜂的性别调控机制及生理影响

沃尔巴克氏体(Wolbachia)是广泛分布于节肢动物生殖组织内的一类共生细菌。它们通过共生诱导宿主赤眼蜂的产雌孤雌生殖,在生物防治上具有重要意义。文章概述了Wolbachia在赤眼蜂中的分布状况,对赤眼蜂的性别调控机制,在赤眼蜂种群中的传播及限制因素,以及对赤眼蜂适合度的影响。     

Recombination in Wolbachia

Wolbachia are widely distributed intracellular bacteria that cause a number of reproductive alterations in their eukaryotic hosts. Such alterations include the induction of parthenogenesis, feminization, cytoplasmic incompatibility, and male killing [1-11]. These important bacteria may play a role in rapid speciation in insects [12-14], and there is growing interest in their potential uses as tools for biological control and genetic manipulation of pests and disease vectors [15-16]. Here, we show recombination in the Wolbachia outer surface protein gene (wsp) between strains of Wolbachia. In addition, we find a possible ecological context for this recombination. Evidence indicates either genetic exchange between Wolbachia in a parasitoid wasp and in the fly that it parasitizes or horizontal transfer of Wolbachia between the parasitoid and the fly, followed by a recombination event. Results have important implications for the evolution of these bacteria and the potential use of Wolbachia in biological control.