Natural interspecific and intraspecific horizontal transfer of parthenogenesis-inducing Wolbachia in Trichogramma wasps

The intracellular bacterium Wolbachia is one of the most common symbionts in arthropods and, because of its manipulative effects on host reproduction, is assumed to be an important factor in several evolutionary processes. These bacteria are mainly vertically transmitted from mother to daughter through the egg cytoplasm, and horizontal transmission is generally assumed to be rare. Here, we show natural inter- and intraspecific horizontal transfer of parthenogenesis-inducing Wolbachia between parasitoid wasps of the genus Trichogramma. Horizontal transfer was observed when infected and uninfected larvae shared the same host egg. This is the first report, to our knowledge, on interspecific horizontal transfer of Wolbachia between closely related sympatric species. Some originally uninfected immature wasps acquired Wolbachia while inside the host egg, but not all of these newly infected females exhibited the parthenogenesis phenotype. In general, intraspecific horizontal transfer was more successful than interspecific transfer. Wolbachia underwent vertical transmission in the new species but the infection tended to be lost within several generations. Our results have important implications for understanding the evolution of Wolbachia-host associations.     

卷蛾赤眼蜂体内共生Wolbachia对寄主产雌孤雌生殖行为的影响

在30℃的条件下,连续饲养卷蛾赤眼蜂 Trichogramma cacoeciae Marchall 10代,观察每代蜂的生物学特性,并进行Wolbachia的wsp基因的分子检测,以研究卷蛾赤眼蜂产雌孤雌生殖行为与其体内共生菌Wolbachia的关系.结果表明,高温下卷蛾赤眼蜂体内的Wolbachia被抑制,4代以后wsp基因的PCR检测结果变为阴性;同时赤眼蜂的产雌孤雌生殖行为的改变程度也随着世代的增加呈递增趋势.这显示卷蛾赤眼蜂的产雌孤雌生殖行为与其体内的共生菌Wolbachia具有一定程度的相关性.     

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。     

An ancient horizontal gene transfer between mosquito and the endosymbiotic bacterium Wolbachia pipientis

The extent and biological relevance of horizontal gene transfer (HGT) in eukaryotic evolution remain highly controversial. Recent studies have demonstrated frequent and large-scale HGT from endosymbiotic bacteria to their hosts, but the great majority of these transferred genes rapidly become nonfunctional in the recipient genome. Here, we investigate an ancient HGT between a host metazoan and an endosymbiotic bacterium, Wolbachia pipientis. The transferred gene has so far been found only in mosquitoes and Wolbachia. In mosquitoes, it is a member of a gene family encoding candidate receptors required for malaria sporozoite invasion of the mosquito salivary gland. The gene copy in Wolbachia has substantially diverged in sequence from the mosquito homolog, is evolving under purifying selection, and is expressed, suggesting that this gene is also functional in the bacterial genome. Several lines of evidence indicate that the gene may have been transferred from eukaryotic host to bacterial endosymbiont. Regardless of the direction of transfer, however, these results demonstrate that interdomain HGT may give rise to functional, persistent, and possibly evolutionarily significant new genes.     

Independent origins and horizontal transfer of bacterial symbionts of aphids

Many insect groups have obligate associations with primary endosymbionts: mutualistic bacteria that are maternally transmitted and derived from an ancient infection. Often, the same insects are hosts to 'secondary' bacterial symbionts which are maternally transmitted but relatively labile within host lineages. To explore the dynamics of secondary symbiont associations in aphids, we characterized bacteria infecting 15 species of macrosiphine aphids using DNA sequencing, diagnostic polymerase chain reaction (PCR), diagnostic restriction digests, phylogenetic analyses, and electron microscopy to examine aphids from nature and from laboratory colonies. Three types of bacteria besides Buchnera were found repeatedly; all three fall within the Enterobacteriaceae. The R-type has a 16S rDNA less than 0.1% different from that of the secondary symbiont previously reported from Acyrthosiphon pisum and is related to Serratia species. The T-type includes a symbiont previously reported from a whitefly; the U-type comprises a new cluster near the T-type. The T-type was found in every one of 40 Uroleucon ambrosiae clones collected throughout the United States. In contrast, A. pisum individuals were infected by any combination of the three symbiont types. Secondary symbionts were maternally transmitted for 11 months within laboratory-reared A. pisum clones and were present in sexually produced eggs. PCR screens for a bacteriophage, APSE-1, indicated its presence in both A. pisum and U. ambrosiae containing secondary symbionts. Electron microscopy of R-type and T-type bacteria in A. pisum and in U. ambrosiae revealed rod-shaped organisms that attain extremely high densities within a few bacteriocytes.     

Nocturnal parasitism of moth eggs by Trichogramma wasps

Parasitoid wasps of the genus Trichogramma are used worldwide as biological control agents against lepidopteran pests. Trichogramma wasps develop inside eggs of a wide range of host species, most of them moths. They are generally considered as diurnal insects. Here, we investigated whether Trichogramma wasps can also successfully parasitise host eggs at night under controlled laboratory conditions. Eggs of the moth Ephestia kuehniella were offered under dark conditions (scotophase) to females of Trichogramma brassicae and Trichogramma evanescens either from 9:00 PM to 9:00 AM or from 11:00 AM to 5:00 PM at four different temperatures (5°C, 10°C, 15°C and 20°C). Both species are known to parasitise E. kuehniella eggs in the photophase during daytime. The results show that T. brassicae did not parasitise eggs in the scotophase at night and only very few in the artificially induced scotophase during daytime from 10°C to 20°C. In contrast, T. evanescens parasitised more eggs in the dark both at night and artificially induced scotophase during daytime. Parasitism in the scotophase already started at 5°C, with more eggs being parasitised and more offspring being produced at higher temperatures. T. evanescens displayed higher parasitism activity in the induced scotophase during daytime than in the scotophase at night. The present study suggests that Trichogramma are capable of successfully parasitising host eggs at night, even at low temperatures, but that nocturnal activity with respect to parasitism varies between wasp species.     

Side-stepping secondary symbionts: widespread horizontal transfer across and beyond the Aphidoidea

To elucidate the co-evolutionary relationships between phloem-feeding insects and their secondary, or facultative, bacterial symbionts, we explore the distributions of three such microbes--provisionally named the R-type (or PASS, or S-sym), T-type (or PABS), and U-type--across a number of aphid and psyllid hosts through the use of diagnostic molecular screening techniques and DNA sequencing. Although typically maternally transmitted, phylogenetic and pairwise divergence analyses reveal that these bacteria have been independently acquired by a variety of unrelated insect hosts, indicating that horizontal transfer has helped to shape their distributions. Based on the high genetic similarity between symbionts in different hosts, we argue that transfer events have occurred recently on an evolutionary timescale. In several instances, however, closely related symbionts associate with related hosts, suggesting that horizontal transfer between distant relatives may be rarer than transmission between close relatives. Our findings on the prevalence of these symbionts within many aphid taxa, along with published observations concerning their effects on host fitness, imply a significant role of facultative symbiosis in aphid ecology and evolution.     

Dynamics of Wolbachia populations in transfected lines of Trichogramma

Fluorescence in situ hybridization was tested to specifically detect symbionts of the genus Wolbachia in Trichogramma and to allow for semiquantitative estimations of symbiont abundance. Extraction solutions used for horizontal transfers of symbionts contain a high abundance of Wolbachia, but Wolbachia have a low and decreasing abundance in microinjected lines (transfected lines). Moreover, eggs of microinjected lines were shown to be polymorphic for the infection. In naturally infected lines, Wolbachia are localized at the posterior pole of the eggs; they are scattered during the early stages of larval development and then concentrated in the ovaries at the end of the female pupal development. Scattering and concentration are probably not active but rather the result of replications or morphogenesis. Conversely, Wolbachia are not concentrated at the posterior pole of eggs in microinjected lines. Comparison of the within-family and between-family variances of the symbiont abundance in a microinjected line did not lead us to conclude that this character shows a genetic variability.     

Effects of endosymbiotic Wolbachia on the diapause in Trichogramma hosts and effects of the diapause on Wolbachia

Abstract Used as an agent in biological control, Trichogramma which has been infected by endosymbiotic bacteria belonging to the genus Wolbachia and inducing the thelytokous mode of reproduction, increases rearing productivity in commercial insectaries. These insectaries can now make use of Trichogramma diapause to spread out and optimise their production. We therefore studied the effect of Wolbachia on the diapause in Trichogramma evanescens. The symbionts show no beneficial or harmful effect on the proportion of emergence in diapausing individuals or on their fecundity. Therefore, infected strains can easily be used in crop protection. On the other hand, low temperatures exert a slightly negative effect on Wolbachia, as detected from the proportion of females in the offspring of symbiotic diapausing females.     

Efficacy of Trichogramma wasps for controlling tomato leaf miner Tuta absoluta

Tomato is considered as one of the most essential vegetables after potato in Egypt. Tomato is infested by many insects; one of these serious insects is tomato leaf miner Tuta absoluta, which causes yield losses up to 100%. T absoluta is attacking leaves, flowers, stems and fruits. Trichogramma species is playing an important role in reducing this insect infestation. Efficiency of Trichogramma evanescens alone and in combination with insecticides were assessed against T. absoluta in tomato fields. Three release rates 500, 1000 and 1500 parasitized egg/card as well as 2 insecticides (Dimilin and Corgen) were applied. Number of normal and parasitized eggs, live and dead larvae and tunnel numbers were recorded. The results revealed that T. absoluta eggs were higher in control plots than those counted in Trichogramma plots. Among Trichogramma release rates, higher release rates, obtained higher parasitism rates of Tuta eggs (reached to 50%). Companying Trichogramma with IGR Dimilin had a little side effect on parasitism rate which it was smaller compared to plots of Trichogramma alone. Dimilin (IGR) caused reducing in Tuta larval population, which it differed significantly compared to control plots. Another IGR, Coragen resulted increasing the mortality of Tuta larvae than other treatments. In Trichogramma release rates, higher release rates, lower number of tunnels were found. In plots of companying Trichogramma with Coragen, the lowest tunnel numbers were recorded. Field release of T. evancences would be a promise biocontrol agent to suppress T. absoluta. These findings may encourage using T. evancences as a favorable strategy for integrated management of T. absoluta to produce safe tomato.     

短期高温对感染Wolbachia的松毛虫赤眼蜂发育和繁殖的影响

感染了沃尔巴克氏体Wolbachia的松毛虫赤眼蜂Trichogramma dendrolimi Matsumura营孤雌产雌生殖方式,但高温可能影响这一特性。为了模拟该赤眼蜂在田间遇到短期高温后的受影响情况,研究了柞蚕卵繁殖的Wolbachia感染了Wolbachia的松毛虫赤眼蜂孤雌产雌品系,于蛹中期和蛹后期经历6h32,35,38℃高温单次冲击处理,对当代羽化出蜂率、单卵出蜂数、单卵雌蜂率、寿命4个指标和处理子代羽化出蜂率、有效繁殖个体、单卵出蜂数、单卵雌蜂率4个指标的影响。结果表明,短期高温刺激会对孤雌产雌的松毛虫赤眼蜂产生不利影响,尤其是38℃高温不利影响明显,主要表现在羽化出蜂率和单卵出蜂数两指标明显降低;高温冲击对子代蜂各指标影响不明显。2代赤眼蜂均未有雄峰出现,表明短期高温冲击不能对Wolbachia调控其宿主生殖方式的作用产生影响。     

Wolbachia在螟黄赤眼蜂种群内的分布及其遗传稳定性

本文通过对Wolbachia的wsp及16S rDNA基因的PCR扩增及克隆测序,明确了Wolbachia在华南、华北不同作物田间螟黄赤眼蜂Trichogramma chilonis种群内的分布情况.结果表明,在所有检测的螟黄赤眼蜂个体内都存在着Kue和Pip两组Wolbachia,双重感染率达到了100%.在不同温度下用米蛾Corcyra cephalonica Stainton卵饲养多代后,螟黄赤眼蜂体内的Wolbachia类型及感染率未发生改变,表明Wolbachia能够在螟黄赤眼蜂体内稳定遗传,不受环境温度的直接影响.本研究中螟黄赤眼蜂为两性生殖或孤雌产雄生殖,Wolbachia未能引起赤眼蜂的产雌孤雌生殖.首次报道了Wolbachia在我国螟黄赤眼蜂野生种群内的分布情况及其遗传稳定性.     

Insect symbionts and applications: The paradigm of cytoplasmic incompatibility-inducing Wolbachia

Wolbachia is a group of Gram‐negative, obligatory intracellular and maternally transmitted alpha‐Proteobacteria. They have been reported to establish symbiotic relationships with a great variety of species of the most diverse animal class, the insects, as well as with several other arthropods and with filarial nematodes. The reproductive alterations Wolbachia causes in its hosts account for its widespread distribution. These alterations include parthenogenesis, feminization, male killing, and cytoplasmic incompatibility (CI). CI is the most frequent and best studied effect Wolbachia has on its hosts. CI is a form of male sterility, ultimately resulting in embryo lethality in diplodiploid host species. As a consequence of CI, Wolbachia infections spread and lead to the replacement of uninfected populations. CI was used nearly four decades ago to control important disease vectors with very encouraging results, and a number of more recent studies have confirmed the effectiveness of CI as a pest population suppression tool as well as a driving mechanism. Furthermore, recent advancements in the field encourage the development of Wolbachia‐based methods for the biological control of insect pests and disease vectors of agricultural, environmental and medical importance.     

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...     

A case for ancestral transfer of symbionts between cockroaches and termites

Living species of the cockroach family Cryptocercidae have intestinal symbionts that are congeneric with some of the gut protozoa found in Isoptera. Presence of such closely related symbionts in cryptocercids and in termites has been frequently interpreted as a uniquely derived homologous character shared between the two xylophagous groups. This may not be the most parsimonious interpretation. Cryptocercus nymphs placed into Zootermopsis (dampwood termite) colonies were killed and eaten by the termites. Termites placed into a Cryptocercus nest box were also fully consumed. Modern Cryptocercus punctulatus and Zootermopsis are often found in the same decaying logs in the Pacific Northwest of the U.S.A., and it is likely that their ancestors also cohabited in at least a portion of their ranges. By occasionally killing and consuming an intruder from the other group, gut protozoa could have been acquired and exchanged between termites and Cryptocercus or their ancestors, under natural conditions and before the life histories of the protozoa became specialized within the host orders. Implications for assessing the phylogeny of the two dictyopteroid groups are also discussed.     

卷蛾赤眼蜂体内共生菌Wolbachia对寄主产雌孤雌生殖行为的影响

在30℃的条件下,连续饲养卷蛾赤眼蜂Trichogramma cacoeciaeMarchall10代,观察每代蜂的生物学特性,并进行Wolbachia的wsp基因的分子检测,以研究卷蛾赤眼蜂产雌孤雌生殖行为与其体内共生菌Wolbachia的关系。结果表明,高温下卷蛾赤眼蜂体内的Wolbachia被抑制,4代以后wsp基因的PCR检测结果变为阴性;同时赤眼蜂的产雌孤雌生殖行为的改变程度也随着世代的增加呈递增趋势。这显示卷蛾赤眼蜂的产雌孤雌生殖行为与其体内的共生菌Wolbachia具有一定程度的相关性。     

Wolbachia infection in six species of gall wasps and their parasitoids

Wolbachia are endosymbiotic bacteria that are widely present in nematodes and arthropods and sometimes have a significant impact on the evolution, ecology, and biology of their hosts. The co-occurrence of Wolbachia within both Cynipid gall wasps and their parasitoids has rarely been studied. In this study, we report the occurrence of six species of gall wasps and 10 species of their parasitoids in central China. Wolbachia detection using the wsp gene showed that Wolbachia infected two species of gall wasps as well as their parasitoids, indicating that horizontal transmission of Wolbachia occurs between gall wasps and their parasitoids. Given that parasitoids will kill their hosts, Wolbachia may be horizontally transferred from gall wasps to their parasitoids. Using multilocus sequence typing (MLST) analysis, five new strains of Wolbachia were identified, all of which belonged to supergroup A. The strains of Wolbachia that infected gall wasps were not the same as those that infected their parasitoids. This result indicated that Wolbachia may evolve independently in parasitoids after they have been transferred from the host gall wasps.