Predator efficacy and attraction to herbivore-induced volatiles determine insect pest selection of inferior host plant.

Unlike mammals, most invertebrates provide no direct parental care for their progeny, which makes a well-selected oviposition site crucial. However, little is known about the female evaluation of opportunities and threats during host selection. Leveraging the wide range of host plants used by the polyphagous pest, Spodoptera littoralis, we investigate oviposition choice between two plants of different nutritional quality. Females prefer to lay their eggs on the host plant, which has inferior larval development and more natural enemies but provides lower predation rates. On the superior host plant, a major predator shows more successful search behavior and is more attracted to herbivore-induced volatiles. Our findings show that predator efficacy and odor-guided attraction, rather than predator abundance, determine enemy free space. We postulate that predators' behaviors contribute to the weak correlation between preference and performance during host plant selection in S. littoralis and in polyphagous insects in general.

Specific response to herbivore-induced de novo synthesized plant volatiles provides reliable information for host plant selection in a moth.

Animals depend on reliable sensory information for accurate behavioural decisions. For herbivorous insects it is crucial to find host plants for feeding and reproduction, and these insects must be able to differentiate suitable from unsuitable plants. Volatiles are important cues for insect herbivores to assess host plant quality. It has previously been shown that female moths of the Egyptian cotton leafworm, Spodoptera littoralis (Lepidoptera: Noctuidae), avoid oviposition on damaged cotton Gossypium hirsutum, which may mediated by herbivore-induced plant volatiles (HIPVs). Among the HIPVs, some volatiles are released following any type of damage while others are synthesized de novo and released by the plants only in response to herbivore damage. In behavioural experiments we here show that oviposition by S. littoralis on undamaged cotton plants was reduced by adding volatiles collected from plants with ongoing herbivory. Gas chromatography-electroantennographic detection (GC-EAD) recordings revealed that antennae of mated S. littoralis females responded to 18 compounds from a collection of headspace volatiles of damaged cotton plants. Among these compounds, a blend of the seven de novo synthesized volatile compounds was found to reduce oviposition in S. littoralis on undamaged plants under both laboratory and ambient (field) conditions in Egypt. Volatile compounds that are not produced de novo by the plants did not affect oviposition. Our results show that ovipositing females respond specifically to the de novo synthesized volatiles released from plants under herbivore attack. We suggest that these volatiles provide reliable cues for ovipositing females to detect plants that could provide reduced quality food for their offspring and an increased risk of competition and predation.

Modulation of the temporal pattern of calling behavior of female Spodoptera littoralis by exposure to sex pheromone.

We have examined the timing of calling behavior in the female Egyptian cotton leafworm, Spodoptera littoralis and its modification by exposure to sex pheromone. The calling rhythm of the female moth was found to be circadian, persistent for at least 4 days once it has been entrained, and could be phase shifted by altering the light:dark regime. We also found that female exposure to pheromone affected the rate and duration of calling. A brief exposure to pheromone gland extract increased the proportion of females calling in a constant dim light and this effect persisted for at least 2 days. In response to pheromone exposure, significantly more females also called late into scotophase when most unexposed control females had ceased calling. The adaptive significance of responding to conspecific sex pheromone is discussed.

Attraction modulated by spacing of pheromone components and anti-attractants in a bark beetle and a moth.

Orientation for insects in olfactory landscapes with high semiochemical diversity may be a challenging task. The partitioning of odor plumes into filaments that are interspersed with pockets of 'clean air' may help filament discrimination and upwind flight to attractive sources in the face of inhibitory signals. We studied the effect of distance between odor sources on trap catches of the beetle, Ips typographus, and the moth, Spodoptera littoralis. Insects were tested both to spatially separated pheromone components [cis-verbenol and 2-methyl-3-buten-2-ol for Ips; (Z,E)-9,11-tetradecadienyl acetate and (Z,E)-9,12-tetradecadienyl acetate for Spodoptera], and to separated pheromone and anti-attractant sources [non-host volatile (NHV) blend for Ips; (Z)-9-tetradecenyl acetate for Spodoptera]. Trap catch data were complemented with simulations of plume structure and plume overlap from two separated sources using a photo ionization detector and soap bubble generators. Trap catches of the beetle and the moth were both affected when odor sources in the respective traps were increasingly separated. However, this effect on trap catch occurred at smaller (roughly by an order of magnitude) odor source separation distances for the moth than for the beetle. This may reflect differences between the respective olfactory systems and central processing. For both species, the changes in trap catches in response to separation of pheromone components occurred at similar spacing distances as for separation of pheromone and anti-attractant sources. Overlap between two simulated plumes depended on distance between the two sources. In addition, the number of detected filaments and their concentration decreased with downwind distance. This implies that the response to separated odor sources in the two species might take place under different olfactory conditions. Deploying multiple sources of anti-attractant around a pheromone trap indicated long-distance (meter scale) effects of NHV on the beetle and a potential use for NHV in forest protection.

Glomerular representation of plant volatiles and sex pheromone components in the antennal lobe of the female Spodoptera littoralis.

We studied the projection patterns of antennal lobe (AL) interneurones sensitive to plant volatiles and female-produced sex pheromone components in the female moth, Spodoptera littoralis. Ten compounds (eight plant-derived compounds and two sex pheromone components) were singly applied to the antenna and, using intracellular recording and staining techniques, the physiological and morphological characteristics of responding neurones were investigated. In addition, ALs stained with a synapsin antibody were optically sectioned using confocal microscopy, and a three-dimensional map of glomeruli in the anterior aspect of the AL was reconstructed. We used the map as a reference for identification of glomeruli innervated by projection neurones (PNs) that respond to plant volatiles and/or pheromone components. Nineteen PNs, responding to one to seven compounds of the ten tested stimuli, were stained with neurobiotin. These neurones each arborised in a single glomerulus in the frontal side of the AL. PNs responding to the same compound arborised in different glomeruli and PNs arborising in the same glomerulus responded to different compounds. Accordingly, glomeruli harbouring the dendritic arborisations of PNs responding to each of the tested compounds constituted a unique array of glomeruli that were not necessarily adjacent. It was thus clear that, at the output level, a single plant volatile or a sex pheromone component was not represented within a single glomerulus in the AL. We expect complex patterns of glomeruli to be involved in the coding of plant-derived compounds, as well as sex pheromone components, in female S. littoralis.