Female fruit flies (Drosophila melanogaster) oviposit at communal sites where the larvae may cooperate or compete for resources depending on group size. This offers a model system to determine how females assess quantitative social information. We show that the concentration of pheromones found on a substrate increases linearly with the number of adult flies that have visited that site. Females prefer oviposition sites with pheromone concentrations corresponding to an intermediate number of previous visitors, whereas sites with low or high concentrations are unattractive. This dose-dependent decision is based on a blend of 11-cis-Vaccenyl Acetate (cVA) indicating the number of previous visitors and heptanal (a novel pheromone deriving from the oxidation of 7-Tricosene), which acts as a dose-independent co-factor. This response is mediated by detection of cVA by odorant receptor neurons Or67d and Or65a, and at least five different odorant receptor neurons for heptanal. Our results identify a mechanism allowing individuals to transform a linear increase of pheromones into a non-linear behavioral response.

Social interactions are generally regulated by pheromones that convey information about the identity, physiological state, and location of an individual. The fruit fly, Drosophila melanogaster , offers a powerful model system to study the mechanisms through which pheromones modulate social interactions. Most of the fruit fly’s social behavior is demonstrably modulated by pheromones, and many of the chemical compounds composing its pheromonal profile have been characterized. This chapter describes several behavioral bioassays that can be used to determine the function of contact and short-range volatile pheromones in D. melanogaster’s social behavior. The chapter first provides instructions on how to rear flies for pheromonal experimentation and how to generate flies that cannot produce cuticular hydrocarbons. Afterward, protocols on how to determine the function of pheromones in courtship behavior and mate choice are provided, followed by protocols to determine whether pheromones function as volatile or contact cues during oviposition site selection. Finally, the last section of the chapter gives general advice on how to work with pheromones in the laboratory.

Adaptation to different environments can result in reproductive isolation between populations and the formation of new species. Food resources are among the most important environmental factors shaping local adaptation. The chemosensory system, the most ubiquitous sensory channel in the animal kingdom, not only detects food resources and their chemical composition, but also mediates sexual communication and reproductive isolation in many taxa. Chemosensory divergence may thus play a crucial role in resource-mediated adaptation and speciation. Understanding how the chemosensory system can facilitate resource-mediated ecological speciation requires integrating mechanistic studies of the chemosensory system with ecological studies, to link the genetics and physiology of chemosensory properties to divergent adaptation. In this review, we use examples of insect research to present seven key questions that can be used to understand how the chemosensory system can facilitate resource-mediated ecological speciation in consumer populations.

Insects searching for resources are exposed to a complexity of mixed odours, often involving both attractant and repellent substances. Understanding how insects respond to this complexity of cues is crucial for understanding consumer-resource interactions, but also to develop novel tools to control harmful pests. To advance our understanding of insect responses to combinations of attractive and repellent odours, we formulated three qualitative hypotheses; the response-ratio hypothesis, the repellent-threshold hypothesis and the odour-modulation hypothesis. The hypotheses were tested by exposing Drosophila melanogaster in a wind tunnel to combinations of vinegar as attractant and four known repellents; benzaldehyde, 1-octen-3-ol, geosmin and phenol. The responses to benzaldehyde, 1-octen-3-ol and geosmin provided support for the response-ratio hypothesis, which assumes that the behavioural response depends on the ratio between attractants and repellents. The response to phenol, rather supported the repellent-threshold hypothesis, where aversion only occurs above a threshold concentration of the repellent due to overshadowing of the attractant. We hypothesize that the different responses may be connected to the localization of receptors, as receptors detecting phenol are located on the maxillary palps whereas receptors detecting the other odorants are located on the antennae.

Background: Wetlands are habitats where variation in soil moisture content and associated environmental conditions can strongly affect the survival of herbivorous insects by changing host plant quality and natural enemy densities. In this study, we combined natural enemy exclusion experiments with random survival forest analyses to study the importance of local variation in host plant quality and predation by natural enemies on the egg and larval survival of the leaf beetle Galerucella sagittariae along a soil moisture gradient.

Results: Our results showed that the exclusion of natural enemies substantially increased the survival probability of G. sagittariae eggs and larvae. Interestingly, the egg survival probability decreased with soil moisture content, while the larval survival probability instead increased with soil moisture content. For both the egg and larval survival, we found that host plant height, the number of eggs or larvae, and vegetation height explained more of the variation than the soil moisture gradient by itself. Moreover, host plant quality related variables, such as leaf nitrogen, carbon and phosphorus content did not influence the survival of G. sagittariae eggs and larvae.

Conclusion: Our results suggest that the soil moisture content is not an overarching factor that determines the interplay between factors related to host plant quality and factors relating to natural enemies on the survival of G. sagittariae in different microhabitats. Moreover, the natural enemy exclusion experiments and the random survival forest analysis suggest that natural enemies have a stronger indirect impact on the survival of G. sagittariae offspring than host plant quality.

The Queensland fruit fly, Bactrocera tryoni, is considered one of the worst horticultural pests in Australia attacking a large variety of fruit crops. To defeat pest insects, olfactory attractants have been developed and widely used in lure and kill strategies. Male B. tryoni are strongly attracted to the compound raspberry ketone and its synthetic analog, cuelure. Despite the strong behavioral response, a recent study failed to show any activation of antennal receptors to cuelure. Therefore, we hypothesized that cuelure may be detected by an accessory olfactory organ, the maxillary palp. Combining behavioral and physiological experiments we clearly demonstrate that male flies, but not female flies, primarily use the maxillary palps and not the antennae to detect and respond to cuelure. Furthermore, regardless of satiety status, male flies always preferred cuelure over a sugar rich source, unless the maxillary palps were excised.

Parasitoid fitness is influenced by the ability to overcome host defense strategies and by the ability of parasitoid females to select high-quality host individuals. When females are unable to differentiate among hosts, their fitness will decrease with an increasing abundance of resistant hosts. To understand the effect of mixed host populations on female fitness, it is therefore necessary to investigate the ability of female parasitoids to select among hosts. Here, we used behavioral assays, headspace volatile collection, and electrophysiology to study the ability of Asecodes parviclava to use olfactory cues to select between a susceptible host (Galerucella calmariensis) and a resistant host (Galerucella pusilla) from a distance. Our studies show that parasitoid females have the capacity to distinguish the two hosts and that the selection behavior is acquired through experiences during earlier life stages. Further, we identified two volatiles (-terpinolene and [E]--ocimene) which amounts differ between the two plant-herbivore systems and that caused behavioral and electrophysiological responses. The consequence of this selection behavior is that females have the capacity to avoid laying eggs in G.pusilla, where the egg mortality is higher due to much stronger immune responses toward A.parviclava than in larvae of G.calmariensis.

Many insect species have limited sensory abilities and may not be able to perceive the quality of different resource types while approaching patchily distributed resources. These restrictions may lead to differences in selection rates between separate patches and between different resource types within a patch, which may have consequences for associational effects between resources. In this study, we used an oviposition assay containing different frequencies of apple and banana substrates divided over two patches to compare resource selection rates of wild-type Drosophila melanogaster at the between- and within-patch scales. Next, we compared the wild-type behavior with that of the olfactory-deficient strain Orco(2) and the gustatory-deficient strain Poxn(Delta M22-B5) and found comparable responses to patch heterogeneity and similarly strong selection rates for apple at both scales for the wild-type and olfactory-deficient flies. Their oviposition behavior translated into associational susceptibility for apple and associational resistance for banana. The gustatory-deficient flies, on the other hand, no longer had a strong selection rate for apple, strongly differed in between- and within-patch selection rates from the wild-type flies, and caused no associational effects between the resources. Our study suggests that differences in sensory capabilities can affect resource selection at different search behavior scales in different ways and in turn underlie associational effects between resources at different spatial scales.

Associational effects occur when the attack rate on a resource depends on neighbouring resources in the environment. These effects are predicted to result from mismatches experienced by the consumer organism in resource selection along hierarchical search levels. As resource selection depends on sensory information used during search behaviour, we expected that different physiological states of an insect might modulate the outcome of associational effects due to differences in resource selection. We used Drosophila melanogaster, as a model organism for olfactory-guided behaviour in insects, to study the effects of mating induced behavioural changes on associational effects between two alternative resources. We found that mating has no effect on the ability of D. melanogaster to locate resource patches, but rather affects the perception of the resources within the patch. Consequently, we only found associational effects in the experiments with unmated females and not in the experiments with mated females. Our results suggest that the lack of associational effects for mated females resulted from changes in the use of short-range olfactory cues, leading to random selection among the resources. In conclusion, our results suggest that the physiological state of an insect modulates associational effects by affecting resource selection rates within the patch.

Many insects face the challenge to select oviposition sites in heterogeneous environments where biotic and abiotic factors can change over time. One way to deal with this complexity is to use sensory experiences made during developmental stages to locate similar habitats or hosts in which larval development can be maximized. While various studies have investigated oviposition preference and larval performance relationships in insects, they have largely overlooked that sensory experiences made during the larval stage can affect such relationships. We addressed this issue by determining the role of natal experience on oviposition preference and larval performance relationships in a tritrophic system consisting of Galerucella sagittariae, feeding on the two host plants Potentilla palustris and Lysimachia thyrsiflora, and its larval parasitoid Asecodes lucens. We firstly determined whether differences in host-derived olfactory information could lead to divergent host selection, and secondly, whether host preference could result in higher larval performance based on the natal origin of the insects. Our results showed that the natal origin and the quality of the current host are both important aspects in oviposition preference and larval performance relationships. While we found a positive relationship between preference and performance of natal Lysimachia beetles, natal Potentilla larvae showed no such relationship and developed better on L. thyrsiflora. Additionally, the host selection by the parasitoid was mainly affected by the natal origin, while its performance was higher on Lysimachia larvae. With this study we showed that the relationship between oviposition preference and larval performance depends on the interplay between the natal origin of the female and the quality of the current host. However, without incorporating the full tritrophic context of these interactions, their implication in insect fitness and potential adaptation cannot be fully understood.