Taste adaptations associated with host specialization in the specialist Drosophila sechellia.

Chemosensory-driven host plant specialization is a major force mediating insect ecological adaptation and speciation. Drosophila sechellia, a species endemic to the Seychelles islands, feeds and oviposits on Morinda citrifolia almost exclusively. This fruit is harmless to D. sechellia but toxic to other Drosophilidae, including the closely related generalists D. simulans and D. melanogaster, because of its high content of fatty acids. While several olfactory adaptations mediating D. sechellia's preference for its host have been uncovered, the role of taste has been much less examined. We found that D. sechellia has reduced taste and feeding aversion to bitter compounds and host fatty acids that are aversive to D. melanogaster and D. simulans. The loss of aversion to canavanine, coumarin and fatty acids arose in the D. sechellia lineage, as its sister species D. simulans showed responses akin to those of D. melanogaster. Drosophila sechellia has increased taste and feeding responses towards M. citrifolia. These results are in line with D. sechellia's loss of genes that encode bitter gustatory receptors (GRs) in D. melanogaster. We found that two GR genes which are lost in D. sechellia, GR39a.a and GR28b.a, influence the reduction of aversive responses to some bitter compounds. Also, D. sechellia has increased appetite for a prominent host fatty acid compound that is toxic to its relatives. Our results support the hypothesis that changes in the taste system, specifically a reduction of sensitivity to bitter compounds that deter generalist ancestors, contribute to the specialization of D. sechellia for its host.

Phylogenetic diversity of two common Trypanosoma cruzi lineages in the Southwestern United States.

Trypanosoma cruzi is the causative agent of Chagas disease, a devastating parasitic disease endemic to Central and South America, Mexico, and the USA. We characterized the genetic diversity of Trypanosoma cruzi circulating in five triatomine species (Triatoma gerstaeckeri, T. lecticularia, T.indictiva, T. sanguisuga and T. recurva) collected in Texas and Southern Arizona using multilocus sequence typing (MLST) with four single-copy loci (cytochrome oxidase subunit II- NADH dehydrogensase subunit 1 region (COII-ND1), mismatch-repair class 2 (MSH2), dihydrofolate reductase-thymidylate synthase (DHFR-TS) and a nuclear gene with ID TcCLB.506529.310). All T. cruzi variants fall in two main genetic lineages: 75% of the samples corresponded to T. cruzi Discrete Typing Unit (DTU) I (TcI), and 25% to a North American specific lineage previously labelled TcIV-USA. Phylogenetic and sequence divergence analyses of our new data plus all previously published sequence data from those four loci collected in the USA, show that TcIV-USA is significantly different from any other previously defined T. cruzi DTUs. The significant level of genetic divergence between TcIV-USA and other T. cruzi DTUs should lead to an increased focus on understanding the epidemiological importance of this DTU, as well as its geographical range and pathogenicity in humans and domestic animals. Our findings further corroborate the fact that there is a high genetic diversity of the parasite in North America and emphasize the need for appropriate surveillance and vector control programs for Chagas disease in southern USA and Mexico.

Evolution of Olfactory Receptors Tuned to Mustard Oils in Herbivorous Drosophilidae.

The diversity of herbivorous insects is attributed to their propensity to specialize on toxic plants. In an evolutionary twist, toxins betray the identity of their bearers when herbivores coopt them as cues for host-plant finding, but the evolutionary mechanisms underlying this phenomenon are poorly understood. We focused on Scaptomyza flava, an herbivorous drosophilid specialized on isothiocyanate (ITC)-producing (Brassicales) plants, and identified Or67b paralogs that were triplicated as mustard-specific herbivory evolved. Using in vivo heterologous systems for the expression of olfactory receptors, we found that S. flava Or67bs, but not the homologs from microbe-feeding relatives, responded selectively to ITCs, each paralog detecting different ITC subsets. Consistent with this, S. flava was attracted to ITCs, as was Drosophila melanogaster expressing S. flava Or67b3 in the homologous Or67b olfactory circuit. ITCs were likely coopted as olfactory attractants through gene duplication and functional specialization (neofunctionalization and subfunctionalization) in S. flava, a recently derived herbivore.

Food-derived volatiles enhance consumption in Drosophila melanogaster.

Insects use multiple sensory modalities when searching for and accepting a food source, in particular odor and taste cues. Food-derived odorants are generally involved in mediating long- and short-range attraction. Taste cues, in contrast, act directly by contact with the food source, promoting the ingestion of nutritious food and the avoidance of toxic substances. It is possible, however, that insects integrate information from these sensory modalities during the process of feeding itself. Here, using a simple feeding assay, we investigated whether odors modulate food consumption in the fruit fly Drosophila melanogaster We found that the presence of both single food-derived odorants and complex odor mixtures enhanced consumption of an appetitive food. Feeding enhancement depended on the concentration and the chemical identity of the odorant. Volatile cues alone were sufficient to mediate this effect, as feeding was also increased when animals were prevented from contacting the odor source. Both males and females, including virgin females, increased ingestion in the presence of food-derived volatiles. Moreover, the presence of food-derived odorants significantly increased the consumption of food mixtures containing aversive bitter compounds, suggesting that flies integrate diverse olfactory and gustatory cues to guide feeding decisions, including situations in which animals are confronted with stimuli of opposite valence. Overall, these results show that food-derived olfactory cues directly modulate feeding in D. melanogaster, enhancing ingestion.

An inside look at the sensory biology of triatomines.

Although kissing bugs (Triatominae: Reduviidae) are perhaps best known as vectors of Chagas disease, they are important experimental models in studies of insect sensory physiology, pioneered by the seminal studies of Wigglesworth and Gillet more than eighty years ago. Since then, many investigations have revealed that the thermal, hygric, visual and olfactory senses play critical roles in the orientation of these blood-sucking insects towards hosts. Here we review the current knowledge about the role of these sensory systems, focussing on relevant stimuli, sensory structures, receptor physiology and the molecular players involved in the complex and cryptic behavioural repertoire of these nocturnal insects. Odours are particularly relevant, as they are involved in host search and are used for sexual, aggregation and alarm communication. Tastants are critical for a proper recognition of hosts, food and conspecifics. Heat and relative humidity mediate orientation towards hosts and are also important for the selection of resting places. Vision, which mediates negative phototaxis and flight dispersion, is also critical for modulating shelter use and mediating escape responses. The molecular bases underlying the detection of sensory stimuli started to be uncovered by means of functional genetics due to both the recent publication of the genome sequence of Rhodnius prolixus and the availability of modern genome editing techniques.

Neuroethology of Olfactory-Guided Behavior and Its Potential Application in the Control of Harmful Insects.

Harmful insects include pests of crops and storage goods, and vectors of human and animal diseases. Throughout their history, humans have been fighting them using diverse methods. The fairly recent development of synthetic chemical insecticides promised efficient crop and health protection at a relatively low cost. However, the negative effects of those insecticides on human health and the environment, as well as the development of insect resistance, have been fueling the search for alternative control tools. New and promising alternative methods to fight harmful insects include the manipulation of their behavior using synthetic versions of "semiochemicals", which are natural volatile and non-volatile substances involved in the intra- and/or inter-specific communication between organisms. Synthetic semiochemicals can be used as trap baits to monitor the presence of insects, so that insecticide spraying can be planned rationally (i.e., only when and where insects are actually present). Other methods that use semiochemicals include insect annihilation by mass trapping, attract-and- kill techniques, behavioral disruption, and the use of repellents. In the last decades many investigations focused on the neural bases of insect's responses to semiochemicals. Those studies help understand how the olfactory system detects and processes information about odors, which could lead to the design of efficient control tools, including odor baits, repellents or ways to confound insects. Here we review our current knowledge about the neural mechanisms controlling olfactory responses to semiochemicals in harmful insects. We also discuss how this neuroethology approach can be used to design or improve pest/vector management strategies.

The neural bases of host plant selection in a Neuroecology framework.

Understanding how animals make use of environmental information to guide behavior is a fundamental problem in the field of neuroscience. Similarly, the field of ecology seeks to understand the role of behavior in shaping interactions between organisms at various levels of organization, including population-, community- and even ecosystem-level scales. Together, the newly emerged field of "Neuroecology" seeks to unravel this fundamental question by studying both the function of neurons at many levels of the sensory pathway and the interactions between organisms and their natural environment. The interactions between herbivorous insects and their host plants are ideal examples of Neuroecology given the strong ecological and evolutionary forces and the underlying physiological and behavioral mechanisms that shaped these interactions. In this review we focus on an exemplary herbivorous insect within the Lepidoptera, the giant sphinx moth Manduca sexta, as much is known about the natural behaviors related to host plant selection and the involved neurons at several level of the sensory pathway. We also discuss how herbivore-induced plant odorants and secondary metabolites in floral nectar in turn can affect moth behavior, and the underlying neural mechanisms.

Hunger is the best spice: effects of starvation in the antennal responses of the blood-sucking bug Rhodnius prolixus.

Blood-sucking insects strongly rely on olfactory cues to find their vertebrate hosts. As in other insects with different lifestyles, it has been shown that endogenous and exogenous factors modulate olfactory responses. The triatomine bug Rhodnius prolixus is an important vector of Chagas disease and a classical model for studies of physiology and behavior. In this species, the behavioral response to host-derived odorants is modulated by both the time of the day and the starvation. Here I investigated the peripheral neural mechanisms underlying these modulatory effects. For this, I measured the electroantennogram (EAG) responses of insects towards different concentrations (from 0.5% to 75% vol/vol) of an attractive host-odorant, ammonia. I tested the responses of starved and fed animals during the middle of the day (when insects are inactive and aggregated in refuges) and at the beginning of the night (when insects become active and search for hosts). Regardless of the time of the day and the starvation status, EAG responses systematically increased with odorant concentration, thus accurately reflecting the response of olfactory receptor cells. Interestingly, the EAG responses of starved insects were larger than those of fed insects only during the night, with larger differences (6-7 times) observed at low-middle concentrations. This study is the first reporting modulation of sensory responses at the neural level in triatomines. This modulation, considering that triatomine hosts are mostly diurnal and are also potential predators, has an important adaptive value, ensuring that insects search for hosts only when they are hungry and at appropriate times.

Glomerular interactions in olfactory processing channels of the antennal lobes.

An open question in olfactory coding is the extent of interglomerular connectivity: do olfactory glomeruli and their neurons regulate the odorant responses of neurons innervating other glomeruli? In the olfactory system of the moth Manduca sexta, the response properties of different types of antennal olfactory receptor cells are known. Likewise, a subset of antennal lobe glomeruli has been functionally characterized and the olfactory tuning of their innervating neurons identified. This provides a unique opportunity to determine functional interactions between glomeruli of known input, specifically, (1) glomeruli processing plant odors and (2) glomeruli activated by antennal stimulation with pheromone components of conspecific females. Several studies describe reciprocal inhibitory effects between different types of pheromone-responsive projection neurons suggesting lateral inhibitory interactions between pheromone component-selective glomerular neural circuits. Furthermore, antennal lobe projection neurons that respond to host plant volatiles and innervate single, ordinary glomeruli are inhibited during antennal stimulation with the female's sex pheromone. The studies demonstrate the existence of lateral inhibitory effects in response to behaviorally significant odorant stimuli and irrespective of glomerular location in the antennal lobe. Inhibitory interactions are present within and between olfactory subsystems (pheromonal and non-pheromonal subsystems), potentially to enhance contrast and strengthen odorant discrimination.

Effects of starvation on the olfactory responses of the blood-sucking bug Rhodnius prolixus.

Blood-sucking insects use olfactory cues in a variety of behavioral contexts, including host-seeking and aggregation. In triatomines, which are obligated blood-feeders, it has been shown that the response to CO2, a host-associated olfactory cue used almost universally by blood-sucking insects, is modulated by hunger. Host-finding is a particularly dangerous task for these insects, as their hosts are also their potential predators. Here we investigated whether olfactory responses to host-derived volatiles other than CO2 (nonanal, α-pinene and (-)-limonene), attractive odorant mixtures (yeast volatiles), and aggregation pheromones (present in feces) are also modulated by starvation in the blood-sucking bug Rhodnius prolixus. For this, the responses of both non-starved and starved insects were individually tested at the beginning of the scotophase using a dual-choice "T-shaped" olfactometer, in which one of its arms presented odor-laden air and the other arm presented odorless air. We found that the response of non-starved insects toward host-odorants and odorant mixtures was odor-dependent: insects preferred the odor-laden arm of the maze when tested with α-pinene, the odorless arm of the maze when tested with (-)-limonene, and distributed at random when tested with yeast volatiles or nonanal. In contrast, starved insects significantly preferred the odor-laden arm of the maze when tested with host-odorants or yeast volatiles. When tested with aggregation be, while starved insects preferred the odorless arm of the maze; insects that were even more starved (8-9 weeks post-ecdysis) significantly preferred the odor-laden arm of the maze. We postulate that this odor- and starvation-dependent modulation of sensory responses has a high adaptive value, as it minimizes the costs and risks associated with the associated behaviors. The possible physiological mechanisms underlying these modulatory effects are discussed.

Species-specific effects of herbivory on the oviposition behavior of the moth Manduca sexta.

In Southwestern USA, the jimsonweed Datura wrightii and the nocturnal sphinx moth Manduca sexta form a pollinator-plant and herbivore-plant association. While certain plant volatile organic compounds (VOCs) attract moths for oviposition, it is likely that other host-derived olfactory cues, such as herbivore-induced VOCs, repel moths for oviposition. Here, we studied the oviposition preference of female M. sexta towards intact and damaged host plants of three species: D. wrightii, D. discolor (a less preferred feeding resource but also used by females for oviposition), and Solanum lycopersicum-tomato-(used by moths as an oviposition resource only). Damage was inflicted to the plants either by larval feeding or artificial damage. Mated females were exposed to an intact plant and a damaged plant and allowed to lay eggs for 10 min. Oviposition preferences of females were highly heterogeneous in all cases, but a larger proportion of moths laid significantly fewer eggs on feeding-damaged and artificially damaged plants of S. lycopersicum. Many females also avoided feeding-damaged D. discolor and D. wrightii plants induced by treatment with methyl jasmonate. Chemical analyses showed a significant increase in the total amount of VOCs released by vegetative tissues of feeding-damaged plants, as well as species-specific increases in emission of certain VOCs. In particular, feeding-damaged S. lycopersicum plants emitted (-)-linalool, an odorant that repels moths for oviposition. Finally, the emission of D. wrightii floral VOCs, which are important in mediating feeding by adult moths (and hence pollination), did not change in plants damaged by larval feeding. We propose that the observed differential effects of herbivory on oviposition choice are due to different characteristics (i.e., mutually beneficial or parasitic) of the insect-plant interaction.

The distribution and abundance of triatomine insects, potential vectors of Chagas Disease, in a metropolitan area in southern Arizona, United States.

Triatomine insects are a problem for human health in southwestern United States because of the moderate-to-severe allergic reactions their bites can cause and because they are potential vectors of Chagas Disease. Although both infected insects and wild mammalian reservoirs are plentiful in southern U.S., only seven cases of autochthonous transmission (plus 16 new presumed cases) of this disease have been reported to date. Therefore, the purpose of this study was to investigate triatomine distribution and abundance in a metropolitan area in southern Arizona. Species, life-stage, locality, and date of collection were recorded for 1,878 triatomine insects collected during 4 yr inside and around houses. For both sexes of the most abundant species, Triatoma rubida (Uhler) (>95% of triatomines collected), dispersal followed a typical year-to-year pattern: dispersal started at the beginning of May and peaked during the first-second week of June. T. rubida was found widely distributed in suburban areas. Triatomines of the two less abundant species, T. recurva (Stal) and T. protracta (Uhler), were collected in all suburban areas throughout the 4-yr survey. All of these population characteristics were observed both at a large (i.e., all collection sites pooled) and a small (i.e., single collection sites) scale. In total, approximately 55-60% of the triatomines were collected inside houses, and 30-35% of those were found in or near beds; thus, it is likely that they fed on humans. To our knowledge, this study is the first comprehensive multi-year analysis of triatomine distribution and abundance in the U.S., providing data that allow inferences about risks to human health.

An insight into the sialotranscriptome of Triatoma rubida (Hemiptera: Heteroptera).

The kissing bug Triatoma rubida (Uhler, 1894) is found in southwestern United States and parts of Mexico where it is found infected with Trypanosoma cruzi, invades human dwellings and causes allergies from their bites. Although the protein salivary composition of several triatomine species is known, not a single salivary protein sequence is known from T. rubida. Furthermore, the salivary diversity of related hematophagous arthropods is very large probably because of the immune pressure from their hosts. Here we report the sialotranscriptome analysis of T. rubida based on the assembly of 1,820 high-quality expressed sequence tags, 51% of which code for putative secreted peptides, including lipocalins, members of the antigen five family, apyrase, hemolysin, and trialysin families. Interestingly, T. rubida lipocalins are at best 40% identical in primary sequence to those of T. protracta, a kissing bug that overlaps its range with T. rubida, indicating the diversity of the salivary lipocalins among species of the same hematophagous genus. We additionally found several expressed sequence tags coding for proteins of clear Trypanosoma spp. origin. This work contributes to the future development of markers of human and pet exposure to T. rubida and to the possible development of desensitization therapies. Supp. Data 1 and 2 (online only) of the transcriptome and deducted protein sequences can be obtained from http://exon.niaid.nih.gov/transcriptome/Trubida/Triru-S1-web.xlsx and http://exon.niaid.nih.gov/transcriptome/Trubida/Triru-S2-web.xlsx.

The neurobiology of insect olfaction: sensory processing in a comparative context.

The simplicity and accessibility of the olfactory systems of insects underlie a body of research essential to understanding not only olfactory function but also general principles of sensory processing. As insect olfactory neurobiology takes advantage of a variety of species separated by millions of years of evolution, the field naturally has yielded some conflicting results. Far from impeding progress, the varieties of insect olfactory systems reflect the various natural histories, adaptations to specific environments, and the roles olfaction plays in the life of the species studied. We review current findings in insect olfactory neurobiology, with special attention to differences among species. We begin by describing the olfactory environments and olfactory-based behaviors of insects, as these form the context in which neurobiological findings are interpreted. Next, we review recent work describing changes in olfactory systems as adaptations to new environments or behaviors promoting speciation. We proceed to discuss variations on the basic anatomy of the antennal (olfactory) lobe of the brain and higher-order olfactory centers. Finally, we describe features of olfactory information processing including gain control, transformation between input and output by operations such as broadening and sharpening of tuning curves, the role of spiking synchrony in the antennal lobe, and the encoding of temporal features of encounters with an odor plume. In each section, we draw connections between particular features of the olfactory neurobiology of a species and the animal's life history. We propose that this perspective is beneficial for insect olfactory neurobiology in particular and sensory neurobiology in general.

Feeding and defecation behavior of Triatoma rubida (Uhler, 1894) (Hemiptera: Reduviidae) under laboratory conditions, and its potential role as a vector of Chagas disease in Arizona, USA.

Chagas disease is caused by the parasite Trypanosoma cruzi, which is transmitted to humans by blood-sucking triatomine insects. This disease is endemic throughout Mexico and Central and South America, but only a few autochthonous cases have been reported in the United States, despite the fact that infected insects readily invade houses and feed on humans. Competent vectors defecate during or shortly after feeding so that infective feces contact the host. We thus studied the feeding and defecation behaviors of the prevalent species in southern Arizona, Triatoma rubida. We found that whereas defecation during feeding was frequent in females (93%), it was very rare in immature stages (3%), and absent in males. Furthermore, more than half of the immature insects that exhibited multiple feeding bouts (62%) defecated during interruptions of feeding, i.e., while likely on or near the host. These results indicate that T. rubida potentially could transmit T. cruzi to humans.

Spiking patterns and their functional implications in the antennal lobe of the tobacco hornworm Manduca sexta.

Bursting as well as tonic firing patterns have been described in various sensory systems. In the olfactory system, spontaneous bursts have been observed in neurons distributed across several synaptic levels, from the periphery, to the olfactory bulb (OB) and to the olfactory cortex. Several in vitro studies indicate that spontaneous firing patterns may be viewed as "fingerprints" of different types of neurons that exhibit distinct functions in the OB. It is still not known, however, if and how neuronal burstiness is correlated with the coding of natural olfactory stimuli. We thus conducted an in vivo study to probe this question in the OB equivalent structure of insects, the antennal lobe (AL) of the tobacco hornworm Manduca sexta. We found that in the moth's AL, both projection (output) neurons (PNs) and local interneurons (LNs) are spontaneously active, but PNs tend to produce spike bursts while LNs fire more regularly. In addition, we found that the burstiness of PNs is correlated with the strength of their responses to odor stimulation--the more bursting the stronger their responses to odors. Moreover, the burstiness of PNs was also positively correlated with the spontaneous firing rate of these neurons, and pharmacological reduction of bursting resulted in a decrease of the neurons' responsiveness. These results suggest that neuronal burstiness reflects a physiological state of these neurons that is directly linked to their response characteristics.

Local interneuron diversity in the primary olfactory center of the moth Manduca sexta.

Local interneurons (LNs) play important roles in shaping and modulating the activity of output neurons in primary olfactory centers. Here, we studied the morphological characteristics, odor responses, and neurotransmitter content of LNs in the antennal lobe (AL, the insect primary olfactory center) of the moth Manduca sexta. We found that most LNs are broadly tuned, with all LNs responding to at least one odorant. 70% of the odorants evoked a response, and 22% of the neurons responded to all the odorants tested. Some LNs showed excitatory (35%) or inhibitory (33%) responses only, while 33% of the neurons showed both excitatory and inhibitory responses, depending on the odorant. LNs that only showed inhibitory responses were the most responsive, with 78% of the odorants evoking a response. Neurons were morphologically diverse, with most LNs innervating almost all glomeruli and others innervating restricted portions of the AL. 61 and 39% of LNs were identified as GABA-immunoreactive (GABA-ir) and non-GABA-ir, respectively. We found no correlations between odor responses and GABA-ir, neither between morphology and GABA-ir. These results show that, as observed in other insects, LNs are diverse, which likely determines the complexity of the inhibitory network that regulates AL output.

Histamine-immunoreactive local neurons in the antennal lobes of the hymenoptera.

Neural networks receive input that is transformed before being sent as output to higher centers of processing. These transformations are often mediated by local interneurons (LNs) that influence output based on activity across the network. In primary olfactory centers, the LNs that mediate these lateral interactions are extremely diverse. For instance, the antennal lobes (ALs) of bumblebees possess both gamma-aminobutyric acid (GABA)- and histamine-immunoreactive (HA-ir) LNs, and both are neurotransmitters associated with fast forms of inhibition. Although the GABAergic network of the AL has been extensively studied, we sought to examine the anatomical features of the HA-ir LNs in relation to the other cellular elements of the bumblebee AL. As a population, HA-ir LNs densely innervate the glomerular core and sparsely arborize in the outer glomerular rind, overlapping with the terminals of olfactory receptor neurons. Individual fills of HA-ir LNs revealed heavy arborization of the outer ring of a single "principal" glomerulus and sparse arborization in the core of other glomeruli. In contrast, projection neurons and GABA-immunoreactive LNs project throughout the glomerular volume. To provide insight into the selective pressures that resulted in the evolution of HA-ir LNs, we determined the phylogenetic distribution of HA-ir LNs in the AL. HA-ir LNs were present in all but the most basal hymenopteran examined, although there were significant morphological differences between major groups within the Hymenoptera. The ALs of other insect taxa examined lacked HA-ir LNs, suggesting that this population of LNs arose within the Hymenoptera and underwent extensive morphological modification.

Infection of kissing bugs with Trypanosoma cruzi, Tucson, Arizona, USA.

Triatomine insects (Hemiptera: Reduviidae), commonly known as kissing bugs, are a potential health problem in the southwestern United States as possible vectors of Trypanosoma cruzi, the causative agent of Chagas disease. Although this disease has been traditionally restricted to Latin America, a small number of vector-transmitted autochthonous US cases have been reported. Because triatomine bugs and infected mammalian reservoirs are plentiful in southern Arizona, we collected triatomines inside or around human houses in Tucson and analyzed the insects using molecular techniques to determine whether they were infected with T. cruzi. We found that 41.5% of collected bugs (n = 164) were infected with T. cruzi, and that 63% of the collection sites (n = 22) yielded >or=1 infected specimens. Although many factors may contribute to the lack of reported cases in Arizona, these results indicate that the risk for infection in this region may be higher than previously thought.

Antagonistic effects of floral scent in an insect-plant interaction.

In southwestern USA, the jimsonweed Datura wrightii and the nocturnal moth Manduca sexta form a pollinator-plant and herbivore-plant association. Because the floral scent is probably important in mediating this interaction, we investigated the floral volatiles that might attract M. sexta for feeding and oviposition. We found that flower volatiles increase oviposition and include small amounts of both enantiomers of linalool, a common component of the scent of hawkmoth-pollinated flowers. Because (+)-linalool is processed in a female-specific glomerulus in the primary olfactory centre of M. sexta, we hypothesized that the enantiomers of linalool differentially modulate feeding and oviposition. Using a synthetic mixture that mimics the D. wrightii floral scent, we found that the presence of linalool was not necessary to evoke feeding and that mixtures containing (+)- and/or (-)-linalool were equally effective in mediating this behaviour. By contrast, females oviposited more on plants emitting (+)-linalool (alone or in mixtures) over control plants, while plants emitting (-)-linalool (alone or in mixtures) were less preferred than control plants. Together with our previous investigations, these results show that linalool has differential effects in feeding and oviposition through two neural pathways: one that is sexually isomorphic and non-enantioselective, and another that is female-specific and enantioselective.