Dynamic visual cues induce jaw opening and closing by tiger beetles during pursuit of prey

In dynamic locomotory contexts, visual cues often trigger adaptive behaviour by the viewer, yet studies investigating how animals determine impending collisions typically employ either stationary viewers or objects. Here, we describe a dynamic situation of visually guided prey pursuit in which both impending prey contact and escape elicit observable adaptive behaviours in the pursuer, a predatory beetle. We investigated which visual cues may independently control opening and closing of the beetle''s jaws during chases of prey dummies. Jaw opening and closing typically occur when prey is within the 60° binocular field, but not at specific distances, angular sizes or time-to-collision. We show that a sign change in the expansion rate of the target image precedes jaw opening (16 ms) and closing (35 ms), signalling to the beetle that it is gaining on the target or that the target is getting away. We discuss the ‘sloppiness'' of such variation in the lag of the behavioural response, especially jaw closing, as an adaptation to uncertainty about target position due to degradation of the target image by motion blur from the fast-running beetle.     

Static antennae act as locomotory guides that compensate for visual motion blur in a diurnal,keen-eyed predator

High visual acuity allows parallel processing of distant environmental features, but only when photons are abundant enough. Diurnal tiger beetles (Carabidae: Cicindelinae) have acute vision for insects and visually pursue prey in open, flat habitats. Their fast running speed causes motion blur that degrades visual contrast, forces stop-and-go pursuit and potentially impairs obstacle detection. We demonstrate here that vision is insufficient for obstacle detection during running, and show instead that antennal touch is both necessary and sufficient for obstacle detection. While running, tiger beetle vision appears to be photon-limited in a way reminiscent of animals in low-light habitats. Such animals often acquire wide-field spatial information through mechanosensation mediated by longer, more mobile appendages. We show that a nocturnal tiger beetle species waves its antennae in elliptical patterns typical of poorly sighted insects. While antennae of diurnal species are also used for mechanosensation, they are rigidly held forward with the tips close to the substrate. This enables timely detection of path obstructions followed by an increase in body pitch to avoid collision. Our results demonstrate adaptive mechanosensory augmentation of blurred visual information during fast locomotion, and suggest that future studies may reveal non-visual sensory compensation in other fast-moving animals.     

Do earthworms help to sustain the slug predator Pterostichus melanarius (Coleoptera: Carabidae) within crops? Investigations using monoclonal antibodies

Earthworms provide a major potential source of alternative food for polyphagous predators, such as carabid beetles, that are natural enemies of slugs, aphids and other agricultural pests. Non-pest prey may foster larger numbers of natural enemies, which then help to control pests, or alternatively may help to divert the predators away from pest control. An earthworm-specific monoclonal antibody was developed to study carabid-earthworm interactions in the field and assess the role of earthworms as alternative prey. The antibody could identify as little at 7 ng of earthworm protein in an ELISA, and could detect earthworm remains in the foregut of the carabid beetle Pterostichus melanarius for 64 h after consumption. Thirty-six per cent of field-collected beetles contained earthworm remains. Quantities of earthworm proteins in the beetle foreguts were negatively related to total foregut biomass, suggesting that earthworm consumption increased as total prey availability declined. There was also a negative relationship between foregut biomass and beetle numbers, but both quantities and concentrations of earthworm proteins in beetle foreguts were positively related to beetle numbers. This suggests that as beetle activity-density increased, total prey availability declined, or, as prey availability declined, beetles spent more time searching. In these circumstances, beetles fed to a greater extent on earthworms, an acceptable but nonpreferred food item. Earthworms may, therefore, provide an ideal alternative prey for P. melanarius, helping to sustain it when pest numbers are low but allowing it to perform a 'lying-in-wait' strategy, ready to switch back to feeding on pests when they become available.     

Sensory Basis of Food Detection in Wild <Emphasis Type="BoldItalic">Microcebus murinus</Emphasis>

Very little is known about how nocturnal primates find their food. Here we studied the sensory basis of food perception in wild-caught gray mouse lemurs (Microcebus murinus) in Madagascar. Mouse lemurs feed primarily on fruit and arthropods. We established a set of behavioral experiments to assess food detection in wild-born, field-experienced mouse lemurs in short-term captivity. Specifically, we investigated whether they use visual, auditory, and motion cues to find and to localize prey arthropods and further whether olfactory cues are sufficient for finding fruit. Visual cues from motionless arthropod dummies were not sufficient to allow reliable detection of prey in choice experiments, nor did they trigger prey capture behavior when presented on the feeding platform. In contrast, visual motion cues from moving prey dummies attracted their attention. Behavioral observations and experiments with live and recorded insect rustling sounds indicated that the lemurs make use of prey-generated acoustic cues for foraging. Both visual motion cues and acoustic prey stimuli on their own were sufficient to trigger approach and capture behavior in the mouse lemurs. For the detection of fruit, choice experiments showed that olfactory information was sufficient for mouse lemurs to find a piece of banana. Our study provides the first experimental data on the sensory ecology of food detection in mouse lemurs. Further research is necessary to address the role of sensory ecology for food selection and possibly for niche differentiation between sympatric Microcebus species.     

Role of visual and olfactory cues from agricultural hedgerows in the orientation behavior of multicolored Asian lady beetle (Coleoptera: Coccinellidae)

Harmonia axyridis Pallas is an introduced lady beetle common in eastern North American agroecosystems. Two-choice behavioral bioassays were performed to determine whether visual and olfactory stimuli from prey and host habitats could elicit taxis in wild-collected H. axyridis adults and whether beetles exhibit a preference among stimuli. Soybean aphid (Aphis glycines Matsumura) spends much of the year in agricultural hedgerows residing on buckthorn (Rhamnus cathartica L), and H. axyridis is frequently observed feeding on aphids in this habitat. Olfactory bioassays were performed in a Y-tube olfactometer and tested the response of beetles to the odor of buckthorn leaves, apple leaves (Malus domestica Borkh.), and buckthorn leaves both naturally and artificially infested with A. glycines. No differences were observed between the numbers of beetles moving toward the odor of buckthorn artificially infested with A. glycines and uninfested buckthorn, but more beetles preferred naturally infested buckthorn over uninfested buckthorn. Visual bioassays were performed in an acrylic tube arena,and tested beetle response to silhouettes and to apple and buckhorn leaves. Beetles were significantly more likely to choose silhouettes over blank space in visual trials. Significantly more beetles moved toward buckthorn leaves than blank space, but beetles did not discern between apple and buckthorn until olfactory cues were also included. This study lays the foundation for future work examining the response of H. axyridis to visual and olfactory cues in Ontario agroecosystems, which could help enhance effectiveness of H. axyridis as a biological control and mitigate its impacts as a pest species.     

Chasing a dummy target: smooth pursuit and velocity control in male blowflies

Male blowflies chase and catch other flies in fast acrobatic flights. To unravel the underlying control system, we presented a black moving sphere instead of a real fly as a pursuit target. By varying the size and speed of the target, we were able to systematically analyse the decisive visual determinants that guide chasing behaviour. Flies pursue targets of a wide range of sizes and velocities. The percentage of pursuits resulting in target capture decreases with increasing target size and speed. Chasing male flies adjust their forward velocity depending on the retinal size of the target, indicating that retinal size is a relevant input variable of the control system. The chasing fly focuses the target with great accuracy in the frontal part of its visual field by means of a smooth pursuit control system using the retinal position of the target to determine the flight direction. We conclude that for a comprehensive understanding of chasing control different time lags in the control systems of angular and forward velocity together with the impact of inertia on fly movements need to be taken into account.     

The pattern and range of movement of a checkered beetle predator relative to its bark beetle prey

Theoretical studies of predator‐prey population dynamics have increasingly centered on the role of space and the movement of organisms. Yet, empirical studies have been slow to follow suit. Herein, we quantified the long‐range movement of a checkered beetle, Thanasimus dubius, which is an important predator of a pernicious forest pest, the southern pine beetle, Dendroctonus frontalis. Adult checkered beetles were marked and released at five sites and subsequently recaptured at traps baited with pine and pine beetle semiochemicals and located at distances up to 2 km away from the release point. While the pattern of recaptures‐with‐distance at each site provided a modest fit to a simple random‐diffusion model, there was a consistent discrepancy between observed and expected recaptures: a higher than expected proportion of beetles were recaptured at the more distant traps. To account for this deviation, we developed a model of diffusion that allowed for simple heterogeneity in the population of marked beetles; i.e., a slow and fast moving form of the checkered beetle. This model provided a significantly better fit to the data and formed the basis for our estimates of intra‐forest movement. We estimated that on average, one half of the checkered beetles dispersed at least 1.25 km, one third dispersed>2 km, and 5% dispersed>5 km. The source of the heterogeneous dispersal rates were partially due to differences in beetle size: smaller beetles (for both males and females) were more likely to be recaptured away from the release site than larger beetles. The southern pine beetle (prey for the checkered beetle) exhibited no significant heterogeneity in dispersal ability and provided a very good fit to the simple diffusion model. The only difference in dispersal between these two species was that checkered beetles were undergoing greater long‐distance dispersal than the pine beetles (the radius containing 95% of the dispersing individuals was 5.1 km for the checkered beetle and 2.3 km for the pine beetle). Data on the movement of these two species is used to evaluate a general model of spatial pattern formation in a homogeneous environment, and the potential of the checkered beetle as a biological control agent for the southern pine beetle.     

Turbulence, trees and semiochemicals: wind-tunnel orientation of the predator, Rhizophagus grandis, to its barkbeetle prey, Dendroctonus micans

Abstract. Insects apparently responding to the visual stimulus offered by a tree or other object may also be responding to the variety of physical effects caused by the obstruction, including turbulence, a reduction in wind velocity and eddies. The relative importance of prey odour associated with the bark beetle Dendroctonus micans (Kugelann) (Scolytidae), prey-host-tree odour, silhouette, and physical barrier to the wind, in the orientation and landing behaviour of a predatory beetle, Rhizophagus grandis Gyllenhall (Rhizophagidae), were investigated. R. grandis responded very positively to the frass of its prey when presented on a 'tree' but not to the same quantity of frass presented alone. Frass on black plastic uPVC pipes was significantly more attractive than frass on real host tree logs, suggesting that host-tree volatiles do not enhance the attractiveness of prey frass.
The beetles responded to 'transparent' Mylar three-dimensional cylinders in the same way as they did to the black plastic pipes. Beetles were also tested with frass and two-dimensional stimuli offering a visual stimulus or physical barrier alone or in combination. Frass with a two-dimensional visual stimulus was no better than frass alone: few insects landed. Significantly more insects were attracted to the same two-dimensional visual stimulus with a hidden upwind barrier producing turbulence, but a similar number were attracted to the frass in front of a hidden barrier alone. As many plants are similar in size to the logs used in these experiments, comparable effects might influence the behaviour of other phytophagous insects, their predators and parasitoids.     

Emergent effects of ground beetles size diversity on the strength of prey suppression

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Host finding byCryptolaemus montrouzieri (Col., Coccinellidae) a predator of mealybusgs (Hom., Pseudococcidae)

AdultC. montrouzieri were able to detect their prey by visual and chemical stimuli. Presence of mealybug caused a significant increase in the number of turns and a sharp decrease in flight attempts suggesting detection by olfaction. In the light, adult predators found polystyrene dummies significantly faster than under dark (red light) conditions, but light had no effect on the time taken for adults to pass across an invisible test point. Fourth instar larvae could only perceive the prey by physical contact.      

Diurnal Tiger Beetles (Coleoptera: Cicindelidae) Capture Prey Without Sight

Although much research has examined the process of prey capture by tiger beetles, an underlying assumption in this work is that tiger beetles are principally, or even exclusively, visual predators. Because this assumption is untested, we performed a series of experiments on four diurnally active tiger beetles in the Genus Cicindela. Individual beetles were placed in chambers in complete darkness and allowed to forage on apterous Drosophila for 6 h. Contrary to expectations, adults of all tested species captured more than 90% of prey items. These results show that other modalities can be used by tiger beetles during prey capture. Beyond potentially providing an explanation for observed night activity in tiger beetle species, the significance of these findings lies in the need to test underlying assumptions of even well-studied organisms.     

Predation by Pterostichus melanarius (Illiger) (Coleoptera: Carabidae) on immature Rhagoletis mendax Curran (Diptera: Tephritidae) in semi-field and field conditions

Practices that enhance abundance and diversity of generalist predators are often employed with the objective of improving biological control of insect pests. Ground beetles and other predators can prey on blueberry maggot, an important pest of blueberries, when mature larvae pupate in the ground. We conducted mesocosm and field experiments to determine if Pterostichus melanarius, a common predatory ground beetle, lowers maggot numbers in compost mulch or when predator and alternative prey abundances are manipulated. At background (field) densities of alternative prey, increasing densities of P. melanarius did not significantly reduce pest numbers in mesocosms containing compost or soil. When alternative prey were removed from compost, beetles reduced pest numbers by up to 35%. In field experiments, maggot numbers were higher when beetles and other predators were excluded from soil plots, but beetle exclusion had no effect in compost plots where both predator and alternative prey numbers were high. Our results indicate that there can be some reduction of blueberry maggot by P. melanarius and other potential predators when there are few alternative prey. However, despite attracting large numbers of predators compost mulch did not lead to a significant reduction in blueberry maggot; in fact, the high abundance of alternative food associated with compost appeared to interfere with beetle predation on blueberry maggot.     

Impacts on the predatory carabid beetle Ctenognathus novaezelandiae of pure and mixed diets of natural field-collected prey and Spodoptera litura fed control or transgenic avidin tobacco

The potential compatibility with biological control of transgenic insecticidal plants expressing the biotin-binding protein avidin was investigated in tri-trophic experiments with the predatory carabid beetle, Ctenognathus novaezelandiae. Beetles were provided with pure and mixed diets of 33%, 67% or 100% of Spodoptera litura larvae, fed either avidin-expressing or isogenic control tobacco, and invertebrates field-collected from the forest floor. Beetles given only tobacco-fed S. litura, whether avidin was present or not, had lower fecundity, egg fertility, body mass and male survival than beetles that received some field food. Fewer of the avidin tobacco-fed prey were consumed than the control tobacco-fed, whatever mixture or proportion offered, probably as a result of the reduced quality of biotin-deprived prey. Beetles consuming 100% avidin tobacco-fed prey had lower fecundity than those given 100% control tobacco-fed prey, although predation on eggs as well as reduced prey quality could have contributed to this result. Despite the nutritionally limiting nature of an exclusive diet of tobacco-fed prey, there was no effect of avidin on fecundity in beetles consuming 67% or 33% avidin prey, or any effect on female or male mass, survival or egg fertility, even in the 100% avidin prey treatment. Fecundity in beetles fed 33% field food with 67% tobacco-fed prey was lower than in those fed 67% or 100% field food. However, there was no added impact of avidin on fecundity, mass or survival, or egg fertility of the 33% field food diet, suggesting that under field conditions, where a mixture of prey is available, negative impacts of avidin-fed prey are unlikely.     

Features effective in course control during object fixation by walking Colorado beetles

A test paradigm is designed to determine which features of viewed objects (a disk 10° in diameter) are responsible for preferential fixation by Colorado beetles (Leptinotarsa decemlineata). The beetle walks on a locomotion compensator that permits free choice of direction in a visual surround containing three different objects. The beetles use both colour and contrast information in these choice experiments. The preferred colours are yellow and orange; their attractiveness is considerably enhanced when they appear in a pattern with highly contrasting features such as stripes. In its optimal form, the object amounts to a dummy beetle, with a pattern that resembles the markings on the elytra of this species. Since the beetles appear to employ a contact pheromone for sex recognition, the function of orientation to these patterns is unclear. However, the patterns might accelerate the identification of conspecifics or be effective at a distance. The reaction to the dummies is telotactic — that is, the beetles walk towards the dummy. One other object, a 10°-wide black stripe, elicits menotactic orientation, in which the beetle maintains a stable course at an angle to the object.     

Adults of the cerambycid beetle Megacyllene caryae use both olfactory and visual information to locate mates

Many species of beetles in the family Cerambycidae use volatile pheromones to facilitate the location of mates. Visual cues may also influence the location of mates, as the adults of many species of cerambycids are often brightly patterned and diurnal. Theory predicts that combining signals or cues of different modalities (e.g., chemical, visual) to transmit information will increase the likelihood of an organism responding to this information, compared to when the signal or cue is presented alone. Here, we test the hypothesis that attraction of adults of the cerambycid beetle Megacyllene caryae (Gahan) (Coleoptera: Cerambycidae, Clytini) to their pheromones will be increased when visual cues are present. Consistent with that hypothesis, the number of beetles caught by traps baited with pheromones was increased 3.4‐fold when a dead adult beetle of that species was attached to the trap, relative to those with just pheromone alone. Capture of M. caryae in our study was also influenced by the position of traps within forest stands, with traps at 100 m within stands catching 2.5× as many beetles as traps at the forest edge. These findings suggest that vision and visual cues play an important role in the location of mates by cerambycid beetles and warrant further research. Also, the inclusion of visual cues on traps may enhance the efficacy of trapping cerambycid beetles, such as the detection of species that are non‐native and potentially invasive, or when monitoring species that are native and of conservation concern.     

Orientation behavior of Propylaea japonica toward visual and olfactory cues from its prey–host plant combination

The lady beetle Propylaea japonica (Thunberg) (Coleoptera: Coccinellidae) is an important predator of aphids in agroecosystems. The inundative release of coccinellid beetles can be an effective biological control strategy. An understanding of how biological control agents perceive and use stimuli from host plants is the key to successfully implement commercially produced predators. Here, we studied the relative role of visual and volatile cues. Dual‐choice assays using foraging‐naïve and foraging‐experienced P. japonica adults were conducted using cotton plants [Gossypium hirsutum L. (Malvaceae)] with or without infestation by the cotton aphid, Aphis gossypii (Glover) (Hemiptera: Aphididae). Overall, experienced beetles were more attracted than naïve beetles toward cues associated with aphid‐infested plants. Experienced beetles were also more responsive to olfactory cues compared with naïve beetles. Both foraging‐naïve and ‐experienced lady beetles integrate olfactory and visual cues from plants infested with aphids, with an apparently greater reliance on olfactory cues. The results suggest that foraging experience may increase prey location in P. japonica.     

Prey perception of predation risk: volatile chemical cues mediate non-consumptive effects of a predator on a herbivorous insect

Predators can affect prey in two ways—by reducing their density (consumptive effects) or by changing their behavior, physiology or other phenotypic traits (non-consumptive effects). Understanding the cues and sensory modalities prey use to detect predators is critical for predicting the strength of non-consumptive effects and the outcome of predator–prey encounters. While predator-associated cues have been well studied in aquatic systems, less is known about how terrestrial prey, particularly insect larvae, detect their predators. We evaluated how Colorado potato beetle, Leptinotarsa decemlineata, larvae perceive predation risk by isolating cues from its stink bug predator, the spined soldier bug, Podisus maculiventris. When exposed to male “risk” predators that were surgically manipulated so they could hunt but not kill, beetles reduced feeding 29 % compared to controls. Exposure to risk females caused an intermediate response. Beetles ate 24 % less on leaves pre-exposed to predators compared to leaves never exposed to predators, indicating that tactile and visual cues are not required for the prey’s response. Volatile odor cues from predators reduced beetle feeding by 10 % overall, although male predators caused a stronger reduction than females. Finally, visual cues from the predator had a weak effect on beetle feeding. Because multiple cues appear to be involved in prey perception of risk, and because male and female predators have differential effects, beetle larvae likely experience tremendous variation in the information about risk from their local environment.     

Motion Control in Saccade and Smooth Pursuit for Bionic Eye Based on Three-dimensional Coordinates

Saccade and smooth pursuit are two important functions of human eye.In order to enable bionic eye to imitate the two functions,a control method that implements saccade and smooth pursuit based on the three-dimensional coordinates of target is proposed.An optimal observation position is defined for bionic eye based on three-dimensional coordinates.A kind of motion planning method with high accuracy is developed.The motion parameters of stepper motor consisting of angle acceleration and turning time are computed according to the position deviation,the target's angular velocity and the stepper motor's current angular velocity in motion planning.The motors are controlled with the motion parameters moving to given position with desired angular velocity in schedule time.The experimental results show that the bionic eye can move to optimal observation positions in 0.6 s from initial location and the accuracy of 3D coordinates is improved.In addition,the bionic eye can track a target within the error of less than 20 pixels based on three-dimensional coordinates.It is verified that saccade and smooth pursuit of bionic eye based on three-dimensional coordinates are feasible.     

Prey-mediated effects of transgenic canola on a beneficial, non-target, carabid beetle

Transgenic plants producing insecticidal proteins from Bacillus thuringiensis (Bt) can control some major insect pests and reduce reliance on sprayed insecticides. However, large scale adoption of this technology has raised concerns about potential negative effects, including evolution of pest resistance to Bt toxins, transgene flow from Bt crops to other plants, and harm to non-target beneficial organisms. Furthermore, concern has also been expressed over the effects this technology may have on biodiversity in general. Ecologically relevant risk assessment is therefore required (Risk = Hazard × Exposure). Transgenic plants that produce Bt toxins to kill insect pests could harm beneficial predators. This might occur directly by transmission of toxin via prey, or indirectly by toxin-induced reduction in prey quality (Hazard). To test these hypotheses, we determined the effects of Bt-producing canola on a predatory ground beetle (Pterostichus madidus) fed larvae of diamondback moth (Plutella xylostella) that were either susceptible or resistant to the Bt toxin. Survival, weight gain, and adult reproductive fitness did not differ between beetles fed prey reared on Bt-producing plants and those fed prey from control plants. Furthermore, while Bt-resistant prey was shown to deliver high levels of toxin to the beetle when they were consumed, no significant impact upon the beetle was observed. Subsequent investigation showed that in choice tests (Exposure), starved and partially satiated female beetles avoided Bt-fed susceptible prey, but not Bt-fed resistant prey. However, in the rare cases when starved females initially selected Bt-fed susceptible prey, they rapidly rejected them after beginning to feed. This prey type was shown to provide sufficient nutrition to support reproduction in the bioassay suggesting that Bt-fed susceptible prey is acceptable in the absence of alternative prey, however adults possess a discrimination ability based on prey quality. These results suggest that the direct effects of Bt-producing canola on predator life history was minimal, and that predators’ behavioural preferences may mitigate negative indirect effects of reduced quality of prey caused by consumption of Bt-producing plants. The results presented here therefore suggest that cultivation of Bt canola may lead to conservation of non-target predatory and scavenging organisms beneficial in pest control, such as carabids, and may therefore provide more sustainable agricultural systems than current practices. In addition, minimal impacts on beneficial carabids in agro-ecosystems suggest that Bt canola crops are likely to be compatible with integrated pest management (IPM) systems.     

A model of visually-guided smooth pursuit eye movements based on behavioral observations

We report a model that reproduces many of the behavioral properties of smooth pursuit eye movements. The model is a negative-feedback system that uses three parallel visual motion pathways to drive pursuit. The three visual pathways process image motion, defined as target motion with respect to the moving eye, and provide signals related to image velocity, image acceleration, and a transient that occurs at the onset of target motion. The three visual motion signals are summed and integrated to produce the eye velocity output of the model. The model reproduces the average eye velocity evoked by steps of target velocity in monkeys and humans and accounts for the variation among individual responses and subjects. When its motor pathways are expanded to include positive feedback of eye velocity and a switch, the model reproduces the exponential decay in eye velocity observed when a moving target stops. Manipulation of this expanded model can mimic the effects of stimulation and lesions in the arcuate pursuit area, the middle temporal visual area (MT), and the medial superior temporal visual area (MST).