Behavioral Choice Tests Research Articles

Suitability and Profitability of a Cereal Aphid for the Parasitoid Aphidius platensis in the Context of Conservation Biological Control of Myzus persicae in Orchards.

The use of cover crops can promote the abundance and early arrival of populations of natural enemies. Cereal cover crops between orchards rows could encourage the early arrival of the parasitoid Aphidius platensis, as they offer alternative winter hosts (e.g., Rhopalosiphum padi), enhancing the control of Myzus persicae in spring. However, the preference for and suitability of the alternative host must be addressed beforehand. To evaluate the potential of this strategy, we assessed host preference using behavioural choice tests, as well as no-choice tests measuring fitness traits, when developing on both host species. One source field for each aphid population from the above hosts was chosen. There was a clear choice for R. padi compared to M persicae, independently of the source, probably due to more defensive behaviours of M. persicae (i.e., kicks and escapes). Nevertheless, both aphid species were suitable for parasitoids’ development. The female progeny developed on R. padi were larger in size, irrespective of their origin. According to our results, in peach orchards with cereals sown between peach trees during the autumn, where we expect when R. padi populations will no longer be available during spring, A. platensis should be able to switch to M. persicae.

Novel GPR120 agonist TUG891 modulates fat taste perception and preference and activates tongue-brain-gut axis in mice

GPR120 is implicated as a lipid receptor in the oro-sensory detection of dietary fatty acids. However, the effects of GPR120 activation on dietary fat intake or obesity are not clearly understood. We investigated to determine whether the binding of TUG891, a novel GPR120 agonist, to lingual GPR120 modulates fat preference in mice. We explored the effects of TUG891 on obesity-related hormones and conducted behavioral choice tests on mice to better understand the physiologic relevance of the action of TUG891. In cultured mouse and human taste bud cells (TBCs), TUG891 induced a rapid increase in Ca2+ by acting on GPR120. A long-chain dietary fatty acid, linoleic acid (LA), also recruited Ca2+ via GPR120 in human and mouse TBCs. Both TUG891 and LA induced ERK1/2 phosphorylation and enhanced in vitro release of glucagon-like peptide-1 from cultured human and mouse TBCs. In situ application of TUG891 onto the tongue of anesthetized mice triggered the secretion of pancreatobiliary juice, probably via the tongue-brain-gut axis. Furthermore, lingual application of TUG891 altered circulating concentrations of cholecystokinin and adipokines, associated with decreased circulating LDL, in conscious mice. In behavioral tests, mice exhibited a spontaneous preference for solutions containing either TUG891 or LA instead of a control. However, addition of TUG891 to a solution containing LA significantly curtailed fatty acid preference. Our study demonstrates that TUG891 binds to lingual GPR120 receptors, activates the tongue-brain-gut axis, and modulates fat preference. These findings may support the development of new fat taste analogs that can change the approach to obesity prevention and treatment.

How do familiarity and relatedness influence mate choice in Armadillidium vulgare?

Mate choice is an important process in sexual selection and usually prevents inbreeding depression in populations. In the terrestrial isopod Armadillidium vulgare, the close physical proximity between individuals may increase the risk of reproducing with siblings. Moreover, individuals of this species can be infected with the feminizing bacteria of Wolbachia, which influence male mate choice. However, little is known about the kinship or familiarity assessment of the selected partner that occurs when a male can choose between females with or without Wolbachia. To investigate the potential mechanisms leading to mate choice and the potential impact of the parasite, we performed behavioral choice tests on males where they could choose between sibling vs. nonsibling females, familiar vs. unfamiliar females, and sibling familiar vs. unfamiliar nonsibling females. To investigate the costs of inbreeding, we compared the reproductive success of both sibling and nonsibling mates. Our results revealed that male copulation attempts were higher for familiar females and for nonsibling females when both females were Wolbachia-infected, but the duration was longer when both females were Wolbachia-free. When males mated with a sibling female, their fecundity was severely decreased, consistent with inbreeding depression. Overall, we observed copulations with all types of females and demonstrated discrimination capacities and potential preferences. We highlight the complexity of the tradeoff between kinship, familiarity and parasite transmission assessment for mate choice.

Blue petrels recognize the odor of their egg.

Most studies on avian olfactory communication have focused on mate choice, and the importance of olfaction in subsequent nesting stages has been poorly explored. In particular, the role of olfactory cues in egg recognition has received little attention, despite eggs potentially being spread with parental odorous secretions known to elicit individual discrimination. Here, we used behavioral choice tests to determine whether female blue petrels (Halobaena caerulea) can discriminate the odor of their own egg from the odor of a conspecific egg. Females preferentially approached the odor of their own egg, suggesting that blue petrels can recognize their own egg using odor cues. This finding raises the question of the adaptive value of this mechanism, and may inspire further research on odor-based egg discrimination in species suffering brood parasitism.

Inhibition of biochemical terpene pathways in <i>Achillea millefolium</i> flowers differently affects the behavior of bumblebees (<i>Bombus terrestris</i>) and flies (<i>Lucilia sericata</i>)

Floral scents serve multiple functions in the interactions with organisms. Flowers of Achillea millefolium (Asteraceae) emit scent bouquets dominated by terpenoids. These flowers are mainly visited by flies and beetles, whereas bumblebees, common visitors at other Asteraceae, are absent from A. millefolium flowers. In order to test how a reduced mono- and sesquiterpenoid emission affect insect behaviour we inhibited the biochemical pathways towards the production of terpenoids of A. millefolium plants and conducted behavioural choice tests. The inhibition resulted in reduced emission rates of most mono- and sesquiterpenes and thus altered the olfactory phenotype of the flowers. In a flight cage, flies usually chose flowers with a natural scent bouquet, bumblebees clearly preferred flowers treated with inhibitors. These findings confirm that floral scents play a pronounced role in foraging decisions of flower visiting insects and support the notion that responses towards scent are animal species-specific emphasising the role of scents as floral filters.

The differential response of workers and queens of the ant Lasius niger to an environment marked by workers: Ants dislike the unknown

It is well known that ants can use cuticular hydrocarbons (CHCs) as a specific recognition cue. Most previous studies addressed the perception of CHCs occurring on the cuticle. However, the presence of CHCs in the environment (e.g., on the substrate) and the role of these compounds as a signal cue are less clear.In this study of the ant L. niger, CHCs were extracted from corpses of workers, from the legs of workers and from filter paper marked by workers. Behavioural choice tests were then performed to study the reactions of workers and queens to CHC passively deposited on the substrate by walking workers.A GC–MS analysis of the CHCs revealed a complex mixture of 28 peaks composed of linear alkanes, alkenes and mono-, di- or trimethylalkane. This mixture was consistently present in the three sources tested (corpses of workers, the tarsus of workers and paper marked by walking workers). Nevertheless, discriminant analyses distinguished clearly among the three types of profiles.The results of our behavioural experiments showed that the workers were able to distinguish between marked and unmarked papers but that they exhibited no preferences for different types of marked papers (papers marked by workers from their own colony or papers marked by workers from a different colony).In queens, the perception of marked papers depended on age or physiological state. Just-mated queens did not discriminate between unmarked papers and papers marked by an alien colony. In contrast, the egg-laying queens 3days after mating significantly favoured the marked papers. We discuss the ecological relevance of the differences in decision-making between castes.

Male and female Trybliographa rapae (Hymenoptera: Figitidae) behavioural responses to food plant, infested host plant and combined volatiles

Many parasitoids use volatiles produced by plants as important cues during their food and host search process. We investigated the attraction of the parasitic wasp Trybliographa rapae Westwood (Hymenoptera: Figitidae) to volatiles emitted from plants infested by the cabbage root fly Delia radicum L. (Diptera: Anthomyiidae), as well as to volatiles from a nectar food plant. Behavioural choice tests showed that male parasitoids were not attracted to any volatiles from plants infested by D. radicum or from nectar plants, while females showed clear attraction to both volatile sources. Young females were more attracted to combined volatiles of host and food plants over those from only the host plant, whereas older females showed no differences in attraction to the two odour sources. This suggests that intercropping attractive flowers with host plants could potentially be used to recruit newly emerged parasitoids from surrounding fields while older parasitoids invest more energy in host location than in additional food search. Volatiles from a whole infested plant were chosen over those emitted from separated above- and below-ground parts from infested plants. It is important to consider the availability of both energy and host resources for parasitoids when designing an eco-compatible management of a vegetable crop system.

The Fecal Odor of Sick Hedgehogs (Erinaceus europaeus) Mediates Olfactory Attraction of the Tick Ixodes hexagonus

Parasite loads of animals vary among individuals, but the underlying mechanisms have not been fully identified. Here, we investigated whether health status of hedgehogs (Erinaceus europaeus) is correlated with tick burden, and whether chemical cues linked to the health status of the host mediate attraction of the tick Ixodes hexagonus. An ecological survey conducted over 10 years, involving 226 wild hedgehogs, revealed a strong association between health status and tick burden of hedgehogs, with healthy animals being less likely to carry ticks than unhealthy ones. Behavioral choice tests demonstrated that ticks display a preference for the fecal odor from sick hedgehogs compared with healthy ones. Chemical analysis of fecal odors using gas chromatography-mass spectrometry showed differences in the odor profile between sick and healthy hedgehogs. Sick animals tended to exhibit raised levels of the volatile aromatic heterocyclic compound indole in their feces. Ticks were attracted to indole when given the choice between indole and a solvent control. However, fecal matter from healthy hosts, with the addition of indole, was not attractive to ticks, suggesting that indole interacts with other, undetected compounds in mediating attraction. This study implies that it is the attraction to fecal odor that causes higher tick burdens in sick hedgehogs. Ticks might benefit from this preference by avoiding possible repulsion mechanisms of healthy hosts. We suggest that ticks potentially choose their host based on odor linked to the host's health status.

Host Plant Phenology Affects Performance of an Invasive Weevil,Phyllobius oblongus(Coleoptera: Curculionidae), in a Northern Hardwood Forest

We investigated how host plant phenology and plant species affected longevity, reproduction, and feeding behavior of an invasive weevil. Phyllobius oblongus L. (Coleoptera: Curculionidae) is common in northern hardwood forests of the Great Lakes Region. Adults emerge in spring, feed on foliage of woody understory plants, and oviposit in the soil. Preliminary data indicate that adults often feed on sugar maple, Acer saccharum Marshall, foliage early in the season, then feed on other species such as raspberry, Rubus spp. Whether this behavior reflects temporal changes in the quality of A. saccharum tissue or merely subsequent availability of later-season plants is unknown. We tested adult P. oblongus in laboratory assays using young (newly flushed) sugar maple foliage, old (2-3 wk postflush) sugar maple foliage, and raspberry foliage. Raspberry has indeterminate growth, thus always has young foliage available for herbivores. Survival, oviposition, and leaf consumption were recorded. In performance assays under no-choice conditions, mated pairs were provided one type of host foliage for the duration of their lives. In behavioral choice tests, all three host plants were provided simultaneously and leaf area consumption was compared. Adults survived longer on and consumed greater amounts of young maple and raspberry foliage than old maple foliage. P. oblongus preferred young maple foliage to old maple foliage early in the season, however, later in the growing season weevils showed less pronounced feeding preferences. These results suggest how leaf phenology, plant species composition, and feeding plasticity in host utilization may interact to affect P. oblongus population dynamics.

Too close for comfort: effect of trap spacing distance and pattern on statistical inference of behavioral choice tests in the field

AbstractBehavioral choice tests provide a powerful and commonly used technique for evaluating the biological activity of chemical signals. Despite the widespread application of this approach, relatively few studies have evaluated a key assumption, that is, relative independence among treatments. Previous work has demonstrated that both the number of choices and their physical arrangement can affect the results of choice tests with leaf‐feeding insects in laboratory assays. Here, we consider another spatial component, the distance between treatments, in a field assay, using a bark beetle as our model. We used three geometries of trap arrangements, two spacing levels, and both ‘low activity’ lures and a ‘high activity’ lure in our behavioral assays. We found that proximity to an attractive treatment generated unexpectedly high trap catches at relatively non‐attractive treatments, even in the presence of a uniform treatment effect and relatively constant insect population size. Increases in traps baited with ‘low activity’ lures proximate to a highly attractive treatment ranged from 4 to 7× the catch observed in configurations with traps spaced wider apart. Moreover, even in the absence of ‘high activity’ lures, a lure catching less than one insect per day on average could obscure the effect of a control trap at proximate spacing. In our example, a spacing distance of 15 m appears to provide independence among traps used to sample the bark beetle Ips pini (Say) (Coleoptera: Curculionidae: Scolytinae), but 3 m does not. Our broader intent is to provide a useful approach to the design and evaluation of behavioral choice experiments in the field.

Intraspecific interactions and their effect on habitat utilisation by the praying mantid Ciulfina biseriata (Mantodea: Liturgusidae)

Tree trunks are spatially separated habitat patches for the tree-running mantid Ciulfina biseriata (Mantodea: Liturgusidae). This paper investigates how intraspecific interactions may affect spacing patterns in C. biseriata. Many mantid species inhabit tree trunk surfaces, but little is known about how they interact with this type of habitat. Behavioural observations and choice tests investigated the effect of a conspecific on habitat utilisation. Field surveys were used to investigate patterns of movement in the field. The decision to inhabit a tree trunk may be affected by the presence of a conspecific. Whereas adult males seek out females when selecting habitat, females will actively avoid males. Habitat choice decisions are primarily based on aspects of their mating system rather than on competition for space or food resources. The behavioural interactions of C. biseriata are described, including novel observations of same-sex mating behaviour in adult males.

Nutritional Manipulation of Adult Female <I>Orius pumilio</I> (Hemiptera: Anthocoridae) Enhances Initial Predatory Performance

Commercial shipments of Orius spp. (Hemiptera: Anthocoridae) commonly include water and protein, the latter typically supplied by eggs from a moth such as Ephestia kuehniella (Zeller). To determine whether modified dietary conditions might improve predation, newly eclosed adult female Orius pumilio (Champion) were fed on E. kuehniella eggs plus encapsulated water, encapsulated 5% sucrose solution only, or encapsulated water only for periods of 24, 48, or 72 h. Feeding activity was assessed by measuring the area of a crop in digital images of dissected digestive tracts. Adult females fed continuously on eggs had larger crops than did females fed on encapsulated sucrose solution. When females were prefed encapsulated water or sucrose and then fed 3 h on eggs, their crops became highly engorged and were larger than those in females fed continuously on eggs for the same periods. In behavioral choice tests, adult females prefed on encapsulated water or 5% sucrose solution spent a larger portion of time in contact with eggs, presumably feeding, whereas females prefed on eggs showed no preference between eggs or encapsulated water. After overnight shipping, females prefed on water or sucrose and held for 48 h total consumed 3.6- and 4.3-fold, respectively, more western flower thrips, Frankliniella occidentalis (Pergande), in 3 h than those prefed on eggs. Survival rates when prefed on sugar or water were comparable with prefeeding on eggs. Thus, inundative releases of Orius can be enhanced by starvation, because females initially feed much more voraciously after shipment with no apparent reduction in fitness.

Dose-Dependent Pheromone Responses of Mountain Pine Beetle in Stands of Lodgepole Pine

We conducted seven behavioral choice tests with Lindgren multiple-funnel traps in stands of mature lodgepole pine in British Columbia, from 1988 to 1994, to determine the dose-dependent responses of the mountain pine beetle, Dendroctonus ponderosae Hopkins, to its pheromones. A multifunctional dose-dependent response was exhibited by D. ponderosae to the pheromones cis- and trans-verbenol in areas with low population numbers. In an area with a high population level of D. ponderosae, the response was directly proportional to release rates. No dose-dependent response was exhibited by D. ponderosae to exo-brevicomin at low release rates. At rates of release >0.5 mg/d, exo-brevicomin interrupted the attraction of D. ponderosae in a dose-dependent fashion. The bark beetle predators, Enoclerus sphegeus (F.) and Thanasimus undatulus (Say), showed dose-dependent responses to only a few pheromones, with trap catches directly proportional to release rates. The multi-functional response of D. ponderosae to verbenols is consistent with an optimal attack density hypothesis.

Dose-Dependent Pheromone Responses ofIps pini,Orthotomicus latidens(Coleoptera: Scolytidae), and Associates in Stands of Lodgepole Pine

We conducted four behavioral choice tests in stands of mature lodgepole pine in British Columbia, from 1988 to 1991, to determine the dose-dependent responses of Orthotomicus latidens (LeConte) and Ips pini (Say) to their respective pheromones. Dose-dependent responses were exhibited by I. pini to (±)-ipsdienol and lanierone, with trap catches directly proportional to pheromone release rates. We found differences in sex ratio of I. pini in their responses to ipsdienol but not lanierone. There was no dose-dependent response exhibited by O. latidens to its pheromone, (±)-ipsenol, with or without the presence of the host kairomone, (−)-&bgr;-phellandrene. The bark beetle predator, Enoclerus lecontei (Wolcott), showed a dose-dependent response to lanierone. Thanasimus undatulus (Say) was attracted to ipsenol in a dose-dependent fashion only when traps were also baited with &bgr;-phellandrene. Neither T. undatulus nor E. sphegeus (F.) exhibited dose-dependent responses to ipsdienol.

Could Species Segregation be a Consequence of Aggregation Processes? Example ofPeriplaneta americana(L.) andP. fuliginosa(Serville)

AbstractOur aim was to show how interindividual interactions, based on specific recognition signals that induce aggregation, can lead to the emergence of a segregation pattern between species. Groups including two cockroach species,Periplaneta americana(L.) (P.a.) andPeriplaneta fuliginosa(Serville) (Dictyoptera: Blattidae) (P.f.) were tested. Behavioural choice tests between two resting sites demonstrated aggregation behaviour in the two species. Aggregation in both species is based on chemical cues.Periplaneta fuliginosahas a higher aggregation level thanP.a.and responds only to its conspecific odour.Periplaneta americanais attracted by the odour of both species presented alone. In the presence of the odours of both species,P.f.selects its own odour, butP.a.neither selects its own odour, nor aggregates under these conditions. This could mean that the perceived chemical signals impair the aggregation process. In mixed groups including larvae of these two species, segregation between species occurs. We assumed that the first species that happened to form an aggregate on one of the resting sites induced the other species to occupy the remaining site. The emergence of a segregation pattern between the two species could be a consequence of the aggregation processes.

How many choices can your test animal compare effectively? Evaluating a critical assumption of behavioral preference tests

Behavioral choice tests comprise one of the most commonly used experimental designs in ecology. However a critical assumption of these assays, that the outcome is independent of the number of choices, has not been tested explicitly. We developed a methodology for testing this assumption, and discuss how it can be incorporated into experimental design. The model with which we performed this test consisted of an insect herbivore, the gypsy moth Lymantria dispar L., feeding on a clonal host plant, Populus. We established a dose-response feeding gradient by amending leaves of a single age class with defined concentrations of a diterpene, isopimaric acid, that exhibits feeding deterrent properties. We selected various concentrations that elicited different levels of feeding for subsequent tests in which we modified the number of choices. A sample size of 30 assay units per test generated statistically significant separations in two-way choice tests, yielded statistically significant but somewhat inconsistent results when four concentrations were offered, and failed to provide complete separation when five concentrations were offered. Other factors associated with the number of choices that affected results included specific combinations of doses, physical arrangement of choices, and total consumption per assay unit. We used our results to develop procedures for estimating the sample sizes needed to compare a specified number of choices. We based these methods on power considerations, the requirements for data transformation and inclusion of covariates. We develop a general approach for estimating the number of replicates needed to support a particular number of choices for a test organism, and discuss factors to be considered when relating this approach to various types of behavioral choice assays.

Taste detection of phytoecdysteroids in larvae of Bombyx mori, Spodoptera littoralis and Ostrinia nubilalis

A number of plants produce significant amounts of phytoecdysteroids that can disrupt the hormonal levels of insects feeding upon them. Insects equipped with taste receptors sensitive to phytoecdysteroids are able to avoid such plants. How common is this strategy? By recording from the lateral and medial sensilla styloconica in two polyphagous species ( Ostrinia nubilalis and Spodoptera littoralis) and in a monophagous species ( Bombyx mori), we tested whether the receptors could detect three commonly occurring phytoecdysteroids 20-hydroxyecdysone (20E), ponasterone A (ponA) and ecdysone (E). In B. mori, 20E and ponA elicited dose-dependent responses with a threshold of 1 μM only in the medial sensilla. In O. nubilalis, 20E, E and ponA elicited responses at threshold of 1 μM in both sensilla. In S. littoralis, 20E elicited responses with a threshold of 10 μM in both sensilla. By means of behavioural choice tests, we show that 20E is an effective feeding deterrent for O. nubilalis and S. littoralis first instar larvae. This suggests that the perception of phytoecdysteroids is more common among phytophagous lepidoptera than previously thought, although their toxicity or antifeedancy varies between species.

Effects of elicitation treatment and genotypic variation on induced resistance in Populus: impacts on gypsy moth (Lepidoptera: Lymantriidae) development and feeding behavior.

We examined the effects of various wounding treatments and genotypic variation on induced resistance in Populus (Salicales: Salicaceae) against herbivory by the gypsy moth, Lymantria dispar L. (Lepidoptera: Lymantriidae). Second-instar larvae grew and consumed less on leaves from induced than non-induced trees. Likewise, larvae preferred leaf disks from non-induced trees. Among induction treatments, gypsy moth feeding had the strongest and most consistent effect in behavioral choice tests. Mechanical wounding of leaves and mechanical wounding plus application of gypsy moth regurgitant had intermediate effects, while application of jasmonic acid had the weakest overall effect. Under no-choice conditions, there were no consistent trends across clones in the ability of various treatments to elicit plant responses affecting the herbivore. Levels of constitutive and inducible resistance to herbivory varied significantly among 12 Populus clones. Larvae grew up to 30-fold more, and consumed up to 250-fold more on the most suitable than the least suitable clone. Prior feeding by gypsy moths reduced larval feeding up to 71.4% on the most highly inducible clone, but it had little or no effect for the least inducible clones. There was no evidence for a relationship between levels of inducible and constitutive resistance, or between inducible resistance and phylogenetic relatedness among clones. We discuss implications for the ecology and evolution of plant-insect interactions and the management of insect pests.

African Plants as Antifeedants Against Stored-Product Insect Pests

The antifeeding activity of 39 plant extracts against selected species of stored-product insect pests was evaluated. The extracts were prepared from a collection of traditional medicine plants representing 9 African species collected in Zaire and belonging to the Meliaceae, Acanthaceae, Simarroubaceae and Fabaceae familes. Adults and larvae of Tribbolium confusum Duv., adults of Sitophilus granarius L., and larvae of Trogoderma granarium Ev. were used in the feeding-deterrent tests using standard behavioral choice and no-choice tests of indicated food. The most effective plant antifeedants were extracts from Entandrophragma spp., Khaya spp. (Meliaceae) and Quassia africana (Simaroubaceae).

Responses of Gypsy Moth (Lepidoptera: Lymantriidae) and Forest Tent Caterpillar (Lepidoptera: Lasiocampidae) to Transgenic Poplar, Populus spp., Containing a Bacillus thuringiensisd-Endotoxin Gene

Transgenic Populus plants (clone NC5339) expressing a Bacillus thuringiensisd-endotoxin gene significantly reduced feeding and weight gain by gypsy moth, Lymantria dispar (L.), and the forest tent caterpillar, Malacosoma disstria Hubner. Forest tent caterpillar larvae also experienced significant mortality following exposure to the transgenic d-endotoxin. Gypsy moths were better able to recover after transfer to artificial diet than were forest tent caterpillars. Transgenic leaves deterred feeding by both species in behavioral choice tests. Four additional transformants contained, but did not express the d-endotoxin gene. Results are discussed in terms of efficacy, plant deployment strategies, ecosystem stability, and tree improvement.