Host genotype–endosymbiont associations and their relationship with aphid parasitism at the field level

1. The relationship between endosymbionts and insects represent complex eco‐evolutionary interactions. Vertically transmitted endosymbionts can be a source of evolutionary novelty by conferring ecologically important traits to their insect hosts, such as protection against natural enemies. Host–endosymbiont associations could constitute an adaptive complex (holobiont) on which selective pressures present in the environment can act, being transferred to the next generation. 2. Although several laboratory‐based studies have confirmed host genotype × symbiont interactions, few studies have been directed at those associations in the natural populations and their ability to protect themselves from parasitism pressure at the field level. 3. A field‐based approach to study the aphid genotype–endosymbiont associations and its relationship with the total parasitism in the grain aphid Sitobion avenae was conducted. From the field study, experiments were carried out to study the defensive effect of the two most common facultative endosymbionts (Regiella insecticola and Hamiltonella defensa) present in S. avenae against one of the most important parasitoid species, Aphidius ervi. 4. Evidence is presented here of a high specificity of the aphid clone–endosymbiont associations in the field; however, the field and experimental results here do not support a relationship between the aphid clone–endosymbiont associations and a proxy of total parasitism in S. avenae. These findings highlight the importance of particular host clone–endosymbiont couplings as a key factor in gaining an understanding of the coevolutionary dynamics of endosymbionts in nature and their effect on the invasive potential of pest insects.     

Species and endosymbiont diversity of Bemisia tabaci (Homoptera: Aleyrodidae) on vegetable crops in Senegal

Bemisia tabaci‐transmitted geminiviruses are one of the major threats on cassava and vegetable crops in Africa. However, to date, few studies are available on the diversity of B. tabaci and their associated endosymbionts in Africa. More than 28 species have been described in the complex of B. tabaci cryptic species; among them, 2 are invasive pests worldwide: MED and MEAM1. In order to assess the species diversity of B. tabaci in vegetable crops in Senegal, several samplings in different localities, hosts and seasons were collected and analyzed with nuclear (microsatellite) and mitochondrial (COI) markers. The bacterial endosymbiont community was also studied for each sample. Two species were detected: MED Q1 and MEAM1 B. Patterns of MED Q1 (dominance on most of the samples and sites, highest nuclear and mitochondrial diversity and broader secondary endosymbiont community: Hamiltonella, Cardinium, Wolbachia and Rickettsia), point toward a predominant resident begomovirus vector group for MED Q1 on market gardening crops. Furthermore, the lower prevalence of the second species MEAM1 B, its lower nuclear and mitochondrial diversity and a narrower secondary endosymbiont community (Hamiltonella/Rickettsia), indicate that this genetic group is exotic and results from a recent invasion in this area.     

Wolbachia distribution in selected beetle taxa characterized by PCR screens and MLST data

Wolbachia (Alphaproteobacteria) is an inherited endosymbiont of arthropods and filarial nematodes and was reported to be widespread across insect taxa. While Wolbachia's effects on host biology are not understood from most of these hosts, known Wolbachia‐induced phenotypes cover a spectrum from obligate beneficial mutualism to reproductive manipulations and pathogenicity. Interestingly, data on Wolbachia within the most species‐rich order of arthropods, the Coleoptera (beetles), are scarce. Therefore, we screened 128 species from seven beetle families (Buprestidae, Hydraenidae, Dytiscidae, Hydrophilidae, Gyrinidae, Haliplidae, and Noteridae) for the presence of Wolbachia. Our data show that, contrary to previous estimations, Wolbachia frequencies in beetles (31% overall) are comparable to the ones in other insects. In addition, we used Wolbachia MLST data and host phylogeny to explore the evolutionary history of Wolbachia strains from Hydraenidae, an aquatic lineage of beetles. Our data suggest that Wolbachia from Hydraenidae might be largely host genus specific and that Wolbachia strain phylogeny is not independent to that of its hosts. As this contrasts with most terrestrial Wolbachia–arthropod systems, one potential conclusion is that aquatic lifestyle of hosts may result in Wolbachia distribution patterns distinct from those of terrestrial hosts. Our data thus provide both insights into Wolbachia distribution among beetles in general and a first glimpse of Wolbachia distribution patterns among aquatic host lineages.     

Sex ratio, mating behaviour and Wolbachia infections in a sheetweb spider

The solitary sheetweb spider Pityohyphantes phrygianus has a female-biased primary sex ratio. Earlier studies have indicated that female post-copulatory position may influence the degree of bias, but the adaptive significance of the departure from a Fisherian 1 : 1 sex ratio is not known. In this study we show that there is a significant relationship between abdominal position and offspring sex ratio in undisturbed females. Moreover, female mating behaviour (and thus sex ratio) is correlated with female size, but this relationship may shift from negative to positive depending on the presence of the maternally acquired endosymbiotic bacterium Wolbachia in both sexes. A survey of wild-caught individuals shows that the incidence of Wolbachia infection in natural populations is high. This indicates that Wolbachia may be a significant factor influencing female post-copulatory behaviour and sex ratio in natural populations. We discuss various adaptive reasons for sex-ratio bias and behavioural plasticity in females.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 181–186.     

Trichogramma parasitoids for control of Lepidopteran borers in Taiwan: species,life‐history traits and Wolbachia infections

Trichogramma, polyphagous endoparasitoids of lepidopteran eggs, are used against a variety of crop pests throughout the world including those of sugar cane and corn in south‐eastern Asia. Their ability to be easily and economically reared on factitious hosts and their wide host range have contributed to their widespread use in pest control. The overall aim of this study was to select strains for eventual release in crop areas for control of lepidopteran borer pests of sugar cane and corn. To this end, we identified common Trichogramma species emerging from corn borer egg masses throughout south‐western Taiwan, compared their life‐history characteristics, assessed their thermal limits and identified the Wolbachia infection status of collected Trichogramma parasitoids. Trichogramma ostriniae was the most commonly collected species on corn, with occasional detection of T. chilonis and an unidentified species designated as T. sp. y. Although the sex ratio varied widely between sites, Wolbachia infection was detected only at a single site in one species (T. ostriniae). Wolbachia‐infected T. ostriniae were tolerant to high temperature stress. Trichogramma chilonis had lowest fecundity of the three species tested, and a Wolbachia‐infected T. ostriniae strain had lower fecundity than an uninfected strain. Given the limited availability of distribution and historical data for Trichogramma species in Taiwan, the current study provides a baseline for future work and also highlights the importance of accurately identifying species when establishing colonies of natural enemies for biocontrol.     

The value of an egg: resource reallocation in ladybirds (Coleoptera: Coccinellidae) infected with male-killing bacteria

Male-killing bacteria are thought to persist in host populations by vertical transmission and conferring direct and/or indirect fitness benefits to their hosts. Here, we test the role of indirect fitness benefits accrued from resource reallocation in species that engage in sibling egg cannibalism. We found that a single-egg meal significantly increased larval survival in 12 ladybird species, but the value of an egg (to survival) differed substantially between species. Next, we tested the impact of three male-killing bacteria on larval survival in one ladybird species, Adalia bipunctata. Spiroplasma reduced larval survival, whereas Wolbachia and Rickettsia had no effect. However, Spiroplasma-infected larvae showed the greatest response to a single-egg meal. The indirect fitness benefit obtained from a single egg is thus so large that even male-killing bacteria with direct fitness costs can persist in host populations. This study supports the hypothesis that fitness compensation via resource reallocation can explain male-killing bacteria persistence.     

Extremely low levels of genetic polymorphism in endosymbionts (Buchnera) of aphids (Pemphigus)

Molecular evolutionary studies have suggested that vertically transmitted endosymbionts are subject to accumulation of deleterious mutations through genetic drift. Predictions of this hypothesis for patterns of intraspecific polymorphism were borne out in the single relevant study available, on the symbiont Buchnera aphidicola of Uroleucon ambrosiae. In order to examine the generality of this result, we surveyed DNA sequence variation in Buchnera of the distantly related aphid, Pemphigus obesinymphae. In contrast to Uroleucon species, Pemphigus species have complex life cycles with few dispersal stages. Despite these differences, P. obesinymphae showed patterns of variation at two Buchnera loci and one mitochondrial locus that were remarkably similar to those reported previously for Buchnera of U. ambrosiae. In the western US, Buchnera was nearly monomorphic, and in the eastern US, synonymous divergence ranged from 0.08 to 0.16%. Most polymorphisms involved rare alleles, consistent with a recent range of ancestral polymorphism, probably due to demographic fluctuations in aphid populations. These results support the generality of small effective population size in Buchnera and their aphid hosts.     

Quantitative steps in symbiogenesis and the evolution of homeostasis

The merging of two independent populations of heterotrophs and autotrophs into a single population of mixotrophs has occurred frequently in evolutionary history. It is an example of a wide class of related phenomena, known as symbiogenesis. The physiological basis is almost always (reciprocal) syntrophy, where each species uses the products of the other species. Symbiogenesis can repeat itself after specialization on particular assimilatory substrates. We discuss quantitative aspects and delineate eight steps from two free-living interacting populations to a single fully integrated endosymbiotic one. The whole process of gradual interlocking of the two populations could be mimicked by incremental changes of particular parameter values. The role of products gradually changes from an ecological to a physiological one. We found conditions where the free-living, epibiotic and endobiotic populations of symbionts can co-exist, as well as conditions where the endobiotic symbionts outcompete other symbionts. Our population dynamical analyses give new insights into the evolution of cellular homeostasis. We show how structural biomass with a constant chemical composition can evolve in a chemically varying environment if the parameters for the formation of products satisfy simple constraints. No additional regulation mechanisms are required for homeostasis within the context of the dynamic energy budget (DEB) theory for the uptake and use of substrates by organisms. The DEB model appears to be dosed under endosymbiosis. This means that when each free-living partner follows DEB rules for substrate uptake and use, and they become engaged in an endosymbiotic relationship, a gradual transition to a single fully integrated system is possible that again follows DEB rules for substrate uptake and use.