Sex pheromones play a pivotal role in the communication of many sexually reproducing organisms. Accordingly, speciation is often accompanied by pheromone diversification enabling proper mate finding and recognition. Current theory implies that chemical signals are under stabilizing selection by the receivers who thereby maintain the integrity of the signals. How the tremendous diversity of sex pheromones seen today evolved is poorly understood. Here we unravel the genetics of a newly evolved pheromone phenotype in wasps and present results from behavioural experiments indicating how the evolution of a new pheromone component occurred in an established sender-receiver system. We show that male Nasonia vitripennis evolved an additional pheromone compound differing only in its stereochemistry from a pre-existing one. Comparative behavioural studies show that conspecific females responded neutrally to the new pheromone phenotype when it evolved. Genetic mapping and gene knockdown show that a cluster of three closely linked genes accounts for the ability to produce this new pheromone phenotype. Our data suggest that new pheromone compounds can persist in a sender's population, without being selected against by the receiver and without the receiver having a pre-existing preference for the new pheromone phenotype, by initially remaining unperceived. Our results thus contribute valuable new insights into the evolutionary mechanisms underlying the diversification of sex pheromones. Furthermore, they indicate that the genetic basis of new pheromone compounds can be simple, allowing them to persist long enough in a population for receivers to evolve chemosensory adaptations for their exploitation.

Interspecific mating can cause severe fitness costs due to the fact that hybrids are often non-viable or less fit. Thus, theory predicts the selection of traits that lessen reproductive interactions between closely related sympatric species. Males of the parasitic wasp Nasonia vitripennis differ from all other Nasonia species by an additional sex pheromone component, but the ecological selective forces underlying this pheromone diversification are unknown. Here we present data from lab experiments suggesting that costly interspecific sexual interactions with the sympatric species N. giraulti might have been responsible for the pheromone evolution and some courtship-related behavioural adaptations in N. vitripennis. Most N. giraulti females are inseminated already within the host, but N. giraulti males still invest in costly sex pheromones after emergence. Furthermore, they do not discriminate between N. vitripennis females and conspecifics during courtship. Therefore, N. vitripennis females, most of which emerge as virgins, face the risk of mating with N. giraulti resulting in costly all-male broods due to Wolbachia-induced cytoplasmic incompatibility. As a counter adaptation, young N. vitripennis females discriminate against N. giraulti males using the more complex conspecific sex pheromone and reject most of them during courtship. With increasing age, however, N. vitripennis females become less choosy, but often compensate mating errors by re-mating with a conspecific. By doing so, they can principally avoid suboptimal offspring sex ratios, but a microcosm experiment suggests that under more natural conditions N. vitripennis females cannot completely avoid fitness costs due to heterospecific mating. Our study provides support for the hypothesis that communication interference of closely related sympatric species using similar sexual signals can generate selective pressures that lead to their divergence.

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BackgroundGraphistrength© C100 multiwalled carbon nanotubes (MWCNT) provide superior electrical and mechanical properties for various applications. The evaluation of the intrinsic hazard properties of Graphistrength© C100 is an essential step for safe use. A general feature of multiwalled carbon nanotubes after inhalation or intratracheal exposures is the induction of an inflammatory reaction in the lungs sometimes associated with local genotoxic effects.MethodsAfter investigating different parameters for the aerosol generation and performing a 5-day inhalation range finding study, male and female Wistar rats were exposed nose-only for 90 days to target concentrations of 0.05, 0.25 and 5.0 mg/m3 air of Graphistrength© C100 and sacrificed 24 h and 90 days after the last exposure. Broncho-alveolar lavage fluid (BALF) was also collected and analyzed for inflammatory parameters. Twenty-four hours post-exposure, chromosomal aberrations in the bone marrow cells were evaluated by the micronucleus test and DNA damages in the lung, kidney and liver cells by both the standard and the human 8-oxoguanine DNA N-glycosylase 1 (hOGG1)-modified comet assay. All studies were performed according to the OECD test guidelines.ResultsAn inflammatory lung reaction and the release of inflammatory factors in the BALF were observed in all rats exposed to 5.0 mg/m3, associated with changes in the differential white blood cells counts. The slight changes in BALF parameters at 0.25 mg/m3 recovered and signs of lung clearance of the MWCNT were observed. No pathological changes were observed on the pleura. Neither increase in the number of micronucleated polychromatic erythrocytes nor increase in percent DNA damage were observed at any concentration.ConclusionsLung inflammation characteristic of an overload with insoluble particles was observed after a 90-day exposure to 5.0 mg/m3 of Graphistrength© C100. Clear signs of clearance and recovery were observed at 0.25 mg/m3. No genotoxicity was detected locally in lung and distally in bone marrow, liver and kidney. Therefore, Graphistrength© C100 appears of low concern in term of local and systemic genotoxicity and a No-Observed Adverse Effect Concentration (NOAEC) of 0.25 mg/m3 (0.28 mg/m3 as actual concentration) was established for the repeated-dose toxicity.Electronic supplementary materialThe online version of this article (doi:10.1186/s12989-015-0096-2) contains supplementary material, which is available to authorized users.

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