Guide Insect Symbiosis, Volume 3 (Contemporary Topics in Entomology)

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Due to the higher susceptibility of P. For each independent replicate for both survival and pathogen multiplication experiments , all the bacterial cultures were prepared separately. After the injections of L. In order to compare the L. Five microliters of hemolymph of each injected animals were sampled 24 h post-injection PI. Sampled hemolymph from S.


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All colonies grown on the plates were morphologically similar. However, to check whether the colonies on the plates were actually S. Sampled hemolymph from L. The plates were then incubated overnight and the CFUs were counted the next day. To check whether the colonies showing the proper morphology all belonged to L.

Asymbiotic dilatatus A along with symbiotic dilatatus A- w Con and dilatatus A- w Dil males Table 1 were injected either with the w VulC suspension obtained from crushed ovaries of infected females or with the control suspension obtained from crushed ovaries of uninfected females.

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The survival was recorded every 15 days starting at day 1 until day The latter can be sampled without killing the animals allowing us to use the same animal both to measure the quantity of Wolbachia and to inject pathogens. Total DNA was extracted using the protocol described by Kocher et al. The wsp copy number was estimated with the help of the standard curve plotted using a dilution of the wsp purified PCR product 2. The wsp copy number was then divided by the total DNA amount of the sample in ng in order to obtain normalized values for comparison between samples.

For each sample two independent technical replicates were made. A t -test with Bonferroni correction for multiple testing was used when the data followed a normal distribution and variances of the samples were homogenous. A Wilcoxon-rank test was performed when the data was not normally distributed.

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Bacterial injection i. As the experiments were performed on different independent groups of individuals, a block effect was included as a random effect in the models. The latter analysis showed a low variance between independent groups i. Each model was also fitted without control groups. This allowed us to compare the survival of the different asymbiotic and symbiotic lines when infected with the pathogenic bacteria.

Survival of the different lines was compared pairwise to assess the difference between their survival times using the log-rank test. Models were fitted for the days 60, 75, and PI in order to test differences in survival during the course of the infection of P. Survival was modeled using a mixed effects Cox proportional hazards model as described above. In both analyses, first a global model including all of the three lines i.

Then, further analysis was made to clarify the effect of the presence of each of the Wolbachia strains i. Additionally, the log-rank test was used to compare the survival times of each lineage pairwise.


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    • Females of symbiotic A. Wolbachia density in Armadillidium vulgare A and Porcellio dilatatus B.

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    NS, not significant. The symbiotic P. The density of Wolbachia in both lines was similar as already reported in Pigeault et al. The number of wsp copies in the w VulC bacterial suspensions injected for transinfection of P. For both isopod species, the global survival models including all treatment groups controls without injection of pathogens included , showed that L.

    The presence or absence of pathogenic bacteria in the injection explained an important part of the deviance between treatments. However, this was not the only significant explanatory factor. In the global model indeed, the symbiotic status i. TABLE 2. Survival analyses of the different A. Other explanatory factors were specific to each of the isopod species. Survival of the different A.

    Control groups were injected with liquid medium LB. Survival of the different P.

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    A Cox proportional hazards model was fitted using the survival data of symbiotic dilatatus A- w Con, dilatatus A- w Dil and asymbiotic dilatatus A P. Results for females A and for males B are presented separately. A reduced submodel was fitted after excluding the control groups i.

    The survival of P. Pairwise comparison of survival data with log-rank test showed that dilatatus A- w Dil animals infected with native Wolbachia survived better than both dilatatus A- w Con and asymbiotic dilatatus A ones. These results showed that only P. Twenty-four hours PI, hemolymph samples were taken from animals injected with S. In both A.

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    Moreover, in both A. CFUs of S. In addition, symbiotic WX w females survived significantly better than asymbiotic WXa when injected with L. Control groups were injected with liquid medium BHI. However, pairwise comparison of survival data showed a weak effect of Wolbachia status; there was actually no significant difference between the survival of asymbiotic and symbiotic females infected with L. A Cox proportional hazards model was fitted using the survival data of symbiotic dilatatus A- w Con, dilatatus A- w Dil and asymbiotic dilatatus A individuals after being injected with pathogenic bacteria L.

    Different letters indicate significant differences between the survival curves of different lineages based on log-rank test. Twenty-four hours PI, hemolymph was sampled from L. The mean of the pathogen load in the hemolymph was more than 60 times lower than the initially injected L. No significant difference between asymbiotic and symbiotic animals was detected in any of the isopod models for A. The negative effect of the w VulC infection on the survival of the animals was detected from day 90 PI Figure 7 ; Table 3. Influence of resident Wolbachia on the survival of P.

    Porcellio dilatatus males were injected with invasive w VulC Wolbachia strain and their survival was recorded every 15 days.


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    A Cox proportional hazards model was fitted. TABLE 3. Influence of resident Wolbachia on the survival and mobility of P. Leg tremors and seizures caused by the w VulC infection were observed during the course of experiment for all lines. The mobility of the animals injected with w VulC decreased compared to the control group, starting from the day 60 PI Table 3.

    This decrease continued until the last days of the infection monitoring in all lines. However, the decrease of mobility due to w VulC was lower for symbiotic dilatatus A- w Con individuals compared to the other symbiotic lineage dilatatus A- w Dil at both days 75 and PI Table 3 ; Figure 8. Influence of resident Wolbachia on the mobility of P.

    The mobility test was performed measuring the time the animals move in a Petri dish during a period of s. The potential mutualistic nature of Wolbachia as a protective symbiont was revealed few years ago by the interactions between Wolbachia and RNA viruses in Drosophila Hedges et al. Since then, the protection conferred by the presence of Wolbachia has been reported against various natural enemies.

    However, this has mainly been observed in artificially established host- Wolbachia associations either when a naturally naive host was transinfected with Wolbachia Kambris et al. The aim of our study was to find out whether a resident Wolbachia has a protective effect against intracellular bacterial pathogens in terrestrial isopods. This demonstrates that w VulC behaves as a protective symbiont for A. A prevailing effect of Wolbachia on A.

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    In 1-year-old A. Besides native w VulC Wolbachia presence leads to a down regulation of some immune genes in the whole body of A. The previously described immunodepressive effects of Wolbachia presence might be somehow compensated by the general up-regulation of immune genes specifically in the immune tissues hence causing the observed protective effect. However, the global picture of Wolbachia influence on A.

    Besides any immune system stimulation, such protection effect of the presence of Wolbachia could also be the result of negative interferences between resident symbiont and invasive bacteria in diverse possible ways. However, each resident Wolbachia increased survival against different invasive pathogenic bacteria: the native w Dil conferred better resistance to S. Previous assessments of the influence of both Wolbachia strains on some P. However, an alternative hypothesis not implying mediation via immunity may also explain such patterns of protection: w Dil could interact harder with Salmonella and counteracts its multiplication while w Con would interact harder with Listeria.

    For P.