We found that amounts of MSP1 and MSP3 components quantified in the air using headspace sampling increased with the number of matings Tables 1 , 2 ; Supplementary Figure 1. We could not compare the role of MSP2 as the latter was not detected in headspace extracts. Hence, mating success is associated to larger amounts of MSP components as quantified in headspace extracts, which possibly increase in relation to increased courtship activity.
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Table 1. Table 2. Summary of models testing for the effects of sex ratio and mating number on MSP1, and MSP3, where male and female age were used as a random factors. We hypothesized that if MSP emission is actively controlled by males, MSP headspace amounts should correlate with courtship activity, though not with general mobility i. We contrasted two types of male behaviors that were recorded during mating experiments: male sexual activity as represented by male courtship fluttering and thrusting and male general movements walking, flying as an internal control.
Males thus appear to actively emit MSPs during courtship activity, while simple wing movements produced during flight have an opposite effect on MSP emission. No MSP2 was found to be emitted. Figure 4.
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Table 3. Summary of models testing for the effect of courtship behaviors and general movements on MSP1 and MSP3 headspace amounts, where male and female age were used as random factors. Our results show that solvent extraction, one of the most common methods used to quantify olfactory signals, does not reflect MSP quantities as they are available in the air for female perception.
We also showed that male mating success correlated to increasing amounts of MSP1 and MSP3 when collected using headspace sampling, but not using solvent extracts, and males mated despite the absence of detectable levels of MSP2 in headspace extracts. Absence of a correlation between male solvent extracts and mating success is unlikely due to methodological differences, because male wings were extracted directly after behavioral experiments had ended, as in previous studies on B.
Finally, MSP amounts emitted in the air were found to increase with courtship activity i. Males could thus actively control the emission of MSP when courting females. How do the discrepancies between headspace and solvent extracts affect our understanding of sexual selection acting on olfactory communication in B.
The most striking difference between MSP solvent and headspace extracts was the absence of MSP2 in the latter, whereas on average 1. Absence of MSP2 in headspace samples could be due either to technical limitations or to a behavioral decision by males, which were indeed found to control MSP1 and MSP3 emission. Technical limitations are unlikely for several reasons. MSP2 should thus have been found in headspace samples if it had been emitted by males.
We further used males of different ages and densities of up to ten males. Absence of MSP2 in headspace samples thus suggests that MSP2 emission is actively controlled by males and that the experimental environment used did not elicit active emission of MSP2. A plausible explanation for the lack of MSP2 production by males may be the limited volume of the entrainment chamber 1. The role of limited cage size and repeated male-female interactions has recently been shown to artificially strengthen the relative importance of male-male competition over expression of female mate preference for determining mating success in this butterfly Holveck et al.
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Females were not able to escape this unnaturally small arena, providing males with an overall high chance of mating Holveck et al. Such a small-sized and crowded environment may lead males to not invest in the emission of the MSP2 component Nieberding and Holveck, , and refs therein. Further experiments using a range of cage volumes, including much larger cages allowing flight and escape to take place, coupled to headspace extractions, will be needed to tease these two explanations apart.
As MSP1 and MSP3 amounts quantified from headspace, but not solvent, extracts correlated with mating success, our results suggest that headspace sampling of MSP components is a better proxy of what females perceive to assess male quality than solvent extracts, when both methods of quantification are compared with a robust statistical approach.
Original Research ARTICLE
Compared to experiments using headspace extracts, a much larger sample size thus appears needed to spot differences in mating success due to variation in MSP levels when solvent extracts are used e. We can reasonably conclude that headspace extracts are more reliable estimates of olfactory signals as they are perceived, and under sexual selection by females, compared to solvent extracts. Our results highlight that our understanding of how sexual selection shapes olfactory sexual communication in this model butterfly may be biased by our sampling methodology.
Relative proportions of MSP components are known to be of great importance for species recognition in sexual interactions of many Lepidoptera Groot et al. In addition, sexually selected traits involved in assessing male quality are usually under strong directional selection for increasing amounts e. Hence, we may have been biased in previous studies with B.
While MSP3 amount did increase in some but not all; van Bergen et al. We may thus underestimate the role of chemical communication in mating success and sexual selection, and particularly of MSP2 amounts in B. Although pheromones that are present in the air may not equate to what is perceived by females because little is known about MSP olfactory receptor proteins on the Bicyclus antennae and Bicyclus neurobiology , this study reveals potential limitations of using solvent extracts of olfactory tissues and organs as proxies for olfactory communication as it evolves under sexual selection.
BV analyzed the data, wrote and edited the manuscript. ID produced technical illustrations and edited the manuscript. GL discussed and edited the manuscript. CN conceived and designed the research, analyzed the data, wrote and edited the manuscript. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. We would like to thank two reviewers for their helpful suggestions on an earlier draft of this paper.
We are further grateful to Christophe Pels for assistance with the butterfly rearing. We thank Dr. Erika A. Wallin and Prof. This paper was posted on bioRxiv prior to publication Visser et al. Supplementary Figure 1. Andersson, J. Male sex pheromone release and female mate choice in a butterfly. Bacquet, P.
Selection on male sex pheromone composition contributes to butterfly reproductive isolation. B Biol. Birch, M. Scents and eversible scent structures of male moths. Boughman, J. Divergent sexual selection enhaces reproductive isolation in sticklebacks. Nature , — Brakefield, P. The African butterfly Bicyclus anynana : a model for evolutionary genetics and evolutionary developmental biology. Cold Spring Harb. Byrne, K. Parapak-Q collection of airborne organic compounds serving as models for insect pheromones.
Google Scholar.
Clearwater, J. Chemistry and function of a pheromone produced by the male of the southern armyworm Pseudaletia separata. Insect Physiol. Costanzo, K. The use of chemical and visual cues in female choice in the butterfly Bicyclus anynana. Cross, J. Parapak-Q collection of phermone components and isolation of Z - and E methylhexadecenal, sex pheromone components, from females of four species of Trogoderma Coleoptera: Dermestidae.
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