Visual gaze control during peering flight manoeuvres in honeybees

As animals travel through the environment, powerful reflexes help stabilize their gaze by actively maintaining head and eyes in a level orientation. Gaze stabilization reduces motion blur and prevents image rotations. It also assists in depth perception based on translational optic flow. Here we describe side-to-side flight manoeuvres in honeybees and investigate how the bees’ gaze is stabilized against rotations during these movements. We used high-speed video equipment to record flight paths and head movements in honeybees visiting a feeder. We show that during their approach, bees generate lateral movements with a median amplitude of about 20 mm. These movements occur with a frequency of up to 7 Hz and are generated by periodic roll movements of the thorax with amplitudes of up to ±60°. During such thorax roll oscillations, the head is held close to horizontal, thereby minimizing rotational optic flow. By having bees fly through an oscillating, patterned drum, we show that head stabilization is based mainly on visual motion cues. Bees exposed to a continuously rotating drum, however, hold their head fixed at an oblique angle. This result shows that although gaze stabilization is driven by visual motion cues, it is limited by other mechanisms, such as the dorsal light response or gravity reception.     

Optomotor regulation of ground velocity in moths during flight to sex pheromone at different heights

ABSTRACT. Males of two species of moths ( Grapholitha molesta (Busck) and Heliothis virescens (F.)) were flown in a sustained-flight tunnel in horizontal pheromone plumes. The up-tunnel velocity of the moths increased with increasing height of flight and for G.molesta was independent of tunnel wind velocities. Use of moving ground patterns verified that the height of flight above the ground was the factor related to the changes in up-tunnel velocity. Even though up-tunnel velocity increased with increased flight height, angular velocity of image motion did not. Males appeared to use visual cues from the ground pattern and from other sources to determine their up-tunnel velocities. The relationship of preferred retinal velocities to optomotor anemotaxis is discussed.     

Active knee joint velocity replication measures are stable and accurate in healthy individuals

The purpose of this study was to determine the stability and accuracy of active knee joint velocity replication methods in healthy subjects. We used a repeated measures design with 14 healthy volunteers. Measures of velocity replication were performed in two ranges of knee joint flexion (0°–30° and 60°–90°), across four testing velocities (5, 10, 15, and 30°/s) in two movement directions (flexion and extension). Statistical analysis included intraclass correlation coefficients (ICCs; 2, k) and associated standard error of the measures calculated between day 1 and 2. We performed z-tests between all possible combinations of ICC pairs using Fisher’s Z transformations to determine if any significant differences existed between observed ICCs. We also calculated correlation ratios (η²) to explain the source of variability in the calculated ICCs. To assess measurement accuracy, we calculated constant error and absolute error between criterion and replication velocities. Results on ICCs and standard error of the measurements (SEMs) ranged from r = ?0.44 ± 7.00 to 0.88 ± 0.72°/s. Calculated z-tests indicated six paired ICCs were significantly different (?p < 0.1). In all six pairs, the faster test velocity had a lower ICC magnitude. The η² calculations demonstrated that inconsistent performance between day 1 and 2 caused the low ICC magnitudes observed with faster testing velocities. Significantly more absolute error occurred at 30 and 15°/s compared with 5°/s. Significantly less constant error was observed for 30°/s compared with 15°/s. A significant direction by range of motion interaction indicated less constant error for flexion movements in the 60°–90° range of motion (ROM) as compared with extension movements in either ROM. Healthy individuals could actively replicate slower criterion velocities in the mid and end ranges of knee joint motion in both movement directions with an acceptable amount of consistency and accuracy. The data support the use of velocity replication in future investigations on proprioceptive function.     

Pesticide exposure affects flight dynamics and reduces flight endurance in bumblebees

The emergence of agricultural land use change creates a number of challenges that insect pollinators, such as eusocial bees, must overcome. Resultant fragmentation and loss of suitable foraging habitats, combined with pesticide exposure, may increase demands on foraging, specifically the ability to collect or reach sufficient resources under such stress. Understanding effects that pesticides have on flight performance is therefore vital if we are to assess colony success in these changing landscapes. Neonicotinoids are one of the most widely used classes of pesticide across the globe, and exposure to bees has been associated with reduced foraging efficiency and homing ability. One explanation for these effects could be that elements of flight are being affected, but apart from a couple of studies on the honeybee (Apis mellifera), this has scarcely been tested. Here, we used flight mills to investigate how exposure to a field realistic (10 ppb) acute dose of imidacloprid affected flight performance of a wild insect pollinator—the bumblebee, Bombus terrestris audax. Intriguingly, observations showed exposed workers flew at a significantly higher velocity over the first ¾ km of flight. This apparent hyperactivity, however, may have a cost because exposed workers showed reduced flight distance and duration to around a third of what control workers were capable of achieving. Given that bumblebees are central place foragers, impairment to flight endurance could translate to a decline in potential forage area, decreasing the abundance, diversity, and nutritional quality of available food, while potentially diminishing pollination service capabilities.     

Vision is Important for Plant Location by the Phytophagous Aquatic Specialist Euhrychiopsis lecontei Dietz (Coleoptera: Curculionidae)

The aquatic milfoil weevil Euhrychiopsis lecontei Dietz (Coleoptera: Curculionidae) is a specialist on Myriophyllum spp. and is used as a biological control agent for Eurasian watermilfoil (Myriophyllum spicatum L.), an invasive aquatic macrophyte. We show evidence that visual cues are important for plant detection by these weevils. Weevils had difficulty locating plants in dark conditions and were highly attracted to plant stems in the light, even when the plant sample was sealed in a vial. However, weevils were equally attracted to both M. spicatum and another aquatic macrophyte, coontail (Ceratophyllum demersum L.) in vials. Turbidity (0–100 NTU) did not significantly influence visual plant detection by the weevils. This work fills a void in the literature regarding visual plant location by aquatic specialists and may help lead to a better understanding of when and where these weevils will find, accept, and damage their target host-plants.     

Stabilizing responses to sideslip disturbances in Drosophila melanogaster are modulated by the density of moving elements on the ground

Stabilizing responses to sideslip disturbances are a critical part of the flight control system in flies. While strongly mediated by mechanoreception, much of the final response results from the wide-field motion detection system associated with vision. In order to be effective, these responses must match the disturbance they are aimed to correct. To do this, flies must estimate the velocity of the disturbance, although it is not known how they accomplish this task when presented with natural images or dot fields. The recent finding, that motion parallax in dot fields can modulate stabilizing responses only if perceived below the fly, raises the question of whether other image statistics are also processed differently between eye regions. One such parameter is the density of elements moving in translational optic flow. Depending on the habitat, there might be strong differences in the density of elements providing information about self-motion above and below the fly, which in turn could act as selective pressures tuning the visual system to process this parameter on a regional basis. By presenting laterally moving dot fields of different densities we found that, in Drosophila melanogaster, the amplitude of the stabilizing response is significantly affected by the number of elements in the field of view. Flies countersteer strongly within a relatively low and narrow range of element densities. But this effect is exclusive to the ventral region of the eye, and dorsal stimuli elicit an unaltered and stereotypical response regardless of the density of elements in the flow. This highlights local specialization of the eye and suggests the lower region may play a more critical role in translational flight stabilization.     

Nocturnal insects use optic flow for flight control

To avoid collisions when navigating through cluttered environments, flying insects must control their flight so that their sensory systems have time to detect obstacles and avoid them. To do this, day-active insects rely primarily on the pattern of apparent motion generated on the retina during flight (optic flow). However, many flying insects are active at night, when obtaining reliable visual information for flight control presents much more of a challenge. To assess whether nocturnal flying insects also rely on optic flow cues to control flight in dim light, we recorded flights of the nocturnal neotropical sweat bee, Megalopta genalis, flying along an experimental tunnel when: (i) the visual texture on each wall generated strong horizontal (front-to-back) optic flow cues, (ii) the texture on only one wall generated these cues, and (iii) horizontal optic flow cues were removed from both walls. We find that Megalopta increase their groundspeed when horizontal motion cues in the tunnel are reduced (conditions (ii) and (iii)). However, differences in the amount of horizontal optic flow on each wall of the tunnel (condition (ii)) do not affect the centred position of the bee within the flight tunnel. To better understand the behavioural response of Megalopta, we repeated the experiments on day-active bumble-bees (Bombus terrestris). Overall, our findings demonstrate that despite the limitations imposed by dim light, Megalopta-like their day-active relatives-rely heavily on vision to control flight, but that they use visual cues in a different manner from diurnal insects.     

To keep on track during flight,fruitflies discount the skyward view

When small flying insects go off their intended course, they use the resulting pattern of motion on their eye, or optic flow, to guide corrective steering. A change in heading generates a unique, rotational motion pattern and a change in position generates a translational motion pattern, and each produces corrective responses in the wingbeats. Any image in the flow field can signal rotation, but owing to parallax, only the images of nearby objects can signal translation. Insects that fly near the ground might therefore respond more strongly to translational optic flow that occurs beneath them, as the nearby ground will produce strong optic flow. In these experiments, rigidly tethered fruitflies steered in response to computer-generated flow fields. When correcting for unintended rotations, flies weight the motion in their upper and lower visual fields equally. However, when correcting for unintended translations, flies weight the motion in the lower visual fields more strongly. These results are consistent with the interpretation that fruitflies stabilize by attending to visual areas likely to contain the strongest signals during natural flight conditions.     

Visual ground pattern modulates flight speed of male Oriental fruit moth Grapholita molesta

Insects flying in a horizontal pheromone plume must attend to visual cues to ensure that they make upwind progress. Moreover, it is suggested that flying insects will also modulate their flight speed to maintain a constant retinal angular velocity of terrestrial contrast elements. Evidence from flies and honeybees supports such a hypothesis, although tests with male moths and beetles flying in pheromone plumes are not conclusive. These insects typically fly faster at higher elevations above a high‐contrast ground pattern, as predicted by the hypothesis, although the increase in speed is not sufficient to demonstrate quantitatively that they maintain constant visual angular velocity of the ground pattern. To test this hypothesis more rigorously, the flight speed of male oriental fruit moths (OFM) Grapholita molesta Busck (Lepidoptera: Tortricidae) flying in a sex pheromone plume in a laboratory wind tunnel is measured at various heights (5–40 cm) above patterns of different spatial wavelength (1.8–90°) in the direction of flight. The OFM modulate their flight speed three‐fold over different patterns. They fly fastest when there is no pattern in the tunnel or the contrast elements are too narrow to resolve. When the spatial wavelength of the pattern is sufficiently wide to resolve, moths fly at a speed that tends to maintain a visual contrast frequency of 3.5 ± 3.2 Hz rather than a constant angular velocity, which varies from 57 to 611° s^?1. In addition, for the first time, it is also demonstrated that the width of a contrast pattern perpendicular to the flight direction modulates flight speed.     

Visual and olfactory disruption of orientation by the western pine beetle to attractant-baited traps

Olfactory deterrents have been proposed as tree protectants against attack by bark beetles, but their development has been hindered by a lack of knowledge of host selection behavior. Among the primary tree-killing (aggressive) Dendroctonus, vision appears to be an integral part of the host selection process. We evaluated the importance of vision in host finding by D. brevicomis LeConte, and our ability to affect it by modifying the visual stimulus provided by attractant-baited multiple-funnel traps. White-painted traps caught 42% fewer D. brevicomis than black traps in California, USA (P < 0.05). Visual treatments were less effective (P < 0.0001) than olfactory disruptants (verbenone with ipsdienol), which reduced catch by about 78%. When combined, olfactory and visual disruptants resulted in 89% fewer D. brevicomis being caught, but this combination was not more effective than olfactory disruptants alone (P > 0.05). Our results demonstrate that the visual component of D. brevicomis host finding behavior can be manipulated, but that D. brevicomis may be more affected by olfactory than visual disruptants. In contrast, visual disruption is more pronounced in the southern pine beetle, Dendroctonus frontalis Zimmermann, suggesting that non-insecticidal tree protection strategies for these related species should differ.     

The dynamics of height stabilization in Drosophila

ABSTRACT. Frequency analysis of the height stabilizing system in Drosophila hydei Sturtevant was performed by moving a striped pattern sinusoidally up and down around free-flying flies. The results show that height is controlled by the flies' vertical acceleration which is set linearly by the velocity of image movement past their eyes in the vertical direction, with a lag of 0.05 s at 25C. No evidence was found of any reaction to the position of the striped pattern relative to the eyes. The gain of the vertical acceleration-controlling reaction decreased with increasing frequency of visual stimulation. The dynamics of this decrease showed that it was not brought about by an integration step in the movement detector, but was due to the flies reaching the limit to the rate at which they could change their acceleration. The reaction time of the flies decreased with increasing temperature from 0.1 s at 15C to 0.04 s at 30C; the gain of the reaction increasing with temperature. When the vertical acceleration and the vertical velocity were determined every 0.02 s, the acceleration rather than the velocity was proportional to the flies' lift, and they took at least 0.3 s to reach their terminal velocity. This means that the acceleration-controlling response found here is equivalent to the lift-controlling responses of tethered flies.     

Visual system of the stalk-eyed fly,Cyrtodiopsis quinqueguttata (Diopsidae,Diptera): an anatomical investigation of unusual eyes

Diopsid flies have eye stalks up to a centimeter in length, displacing the retina laterally from the rest of the head. This bizarre condition, called hypercephaly, is rare, but has evolved independently among several insect orders and is most common in flies (Diptera). Earlier studies of geometrical optics and behavior have led to various hypotheses about possible adaptive advantages of eye stalks, such as enhanced stereoscopic vision while other hypothesis suggest that eye stalks are an outcome of sexual selection. Here, we focus on how these curious distortions of head/eye morphology are accompanied by changes in the neural organization of the visual system of Cyrtodiopsis quinqueguttata. Histological examinations reveal that the optic lobes, lamina (La), medulla (Me), lobula (Lo), and lobula plate (LP) are contained entirely within the fly's eye bulbs, which are located at the distal ends of the eye stalks. We report that the organization of the peripheral visual system (La and Me) is similar to that of other Diptera (e.g., Musca and Drosophila), but deeper visual areas (Lo and LP) have been more strongly modified. For example, in both the lobula and lobula plate, fewer but larger giant collector neurons are found. The most pronounced difference is the reduction in the number of wide-field vertical cells of the lobula plate, where there are only four relatively large fibers, as opposed to 11 in Musca. The “fewer but larger” neural organization may enhance the conduction velocities of these cells, but may result in a loss of spatial resolution. At the base of the eye bulb, axon bundles collect and form a long optic nerve that extends the length of the eye stalk. We suggest that this organization of the diopsid visual system provides evidence for the costs of possessing long eye stalks. © 1998 John Wiley & Sons, Inc. J Neurobiol 37: 449–468, 1998     

Three‐dimensional flight tracking shows how a visual target alters tsetse fly responses to human breath in a wind tunnel

Tsetse flies Glossina spp. (Diptera; Glossinidae) are blood‐feeding vectors of disease that are attracted to vertebrate hosts by odours and visual cues. Studies on how tsetse flies approach visual devices are of fundamental interest because they can help in the development of more efficient control tools. The responses of a forest tsetse fly species Glossina brevipalpis (Newstead) to human breath are tested in a wind tunnel in the presence or absence of a blue sphere as a visual target. The flight responses are video recorded with two motion‐sensitive cameras and characterized in three dimensions. Although flies make meandering upwind flights predominantly in the horizontal plane in the plume of breath alone, upwind flights are highly directed at the visual target presented in the plume of breath. Flies responding to the visual target fly from take‐off within stricter flight limits at lower ground speeds and with a significantly lower variance in flight trajectories in the horizontal plane. Once at the target, flies fly in loops principally in the horizontal plane within 40 cm of the blue sphere before descending in spirals beneath it. Successful field traps designed for G. brevipalpis take into account the strong horizontal component in local search behaviour by this species at objects. The results suggest that trapping devices should also take into account the propensity of G. brevipalpis to descend to the lower parts of visual targets.     

Adipokinetic hormone functions that are not associated with insect flight

Abstract This review deals with some lesser known functions of adipokinetic hormones (AKHs), specifically those that are not associated directly with flight activity. The data summarized and discussed relate to AKHs in insects that have lost the ability to fly and use exclusively and/or mostly walking for their locomotion; and to activation of pathways that do not lead directly to production and subsequent rapid consumption of energy, but help the insect to combat stress situations. Emphasis is placed on AKH‐stimulated walking activities in Pyrrhocoris apterus, Gryllus bimaculatus, Periplaneta americana and Drosophila mellanogaster; diel fluctuations in AKH activities; the actions of AKH in alternative stress situations in which infection, toxins and other kinds of stressors interact; and the role of AKHs in anabolic processes and egg production. Possible mechanisms of action are proposed when justified by available knowledge.     

The spatiotemporal dynamics of Tribolium castaneum (Herbst): adult flight and gene flow

Tribolium castaneum (Herbst) has been used as a model organism to develop and test important ecological and evolutionary concepts and is also a major pest of grain and grain products globally. This beetle species is assumed to be a good colonizer of grain storages through anthropogenic movement of grain, and active dispersal by flight is considered unlikely. Studies using T. castaneum have therefore used confined or walking insects. We combine an ecological study of dispersal with an analysis of gene flow using microsatellites to investigate the spatiotemporal dynamics and adult flight of T. castaneum in an ecological landscape in eastern Australia. Flying beetles were caught in traps at grain storages and in fields at least 1 km from the nearest stored grain at regular intervals for an entire year. Significantly more beetles were trapped at storages than in fields, and almost no beetles were caught in native vegetation reserves many kilometres from the nearest stored grain. Genetic differentiation between beetles caught at storages and in fields was low, indicating that although T. castaneum is predominantly aggregated around grain storages, active dispersal takes place to the extent that significant gene flow occurs between them, mitigating founder effects and genetic drift. By combining ecological and molecular techniques, we reveal much higher levels of active dispersal through adult flight in T. castaneum than previously thought. We conclude that the implications of adult flight to previous and future studies on this model organism warrant consideration.     

Winging it: moth flight behavior and responses of olfactory neurons are shaped by pheromone plume dynamics

Terrestrial odor plumes have a physical structure that results from turbulence in the fluid environment. The rapidity of insect flight maneuvers within a plume indicates that their responses are dictated by fleeting (<1 s) rather than longer (>1 s) exposures to odor imposed by physical variables that distribute odor molecules in time and space. Even though encounters with pheromone filaments are brief, male moths responding to female-produced pheromones are remarkably able to extract information relating to the biological properties of these olfactory signals. These properties include the types of molecule present and their relative abundances. Thus, peripheral and central olfactory neurons are capable of representing these biological properties of a pheromone plume within the context of a temporally irregular and unpredictable signal. The mechanisms underlying olfactory processing of these signals with respect to their biological and physical properties are discussed in the context of a behavioral framework.     

Alterations in flightin phosphorylation inDrosophila flight muscles are associated with myofibrillar defects engendered by actin and myosin heavy-chain mutant alleles

Flightin is a 20-kD myofibrillar protein found in the stretch-activated flight muscles ofDrosophila melanogaster. Nine of the eleven isoelectric variants of flightin are generatedin vivo by multiple phosphorylations. The accumulation of these isoelectric variants is affected differently by mutations that eliminate thick filaments or thin filaments. Mutations in the myosin heavy-chain gene that prevent thick filament assembly block accumulation of all flightin variants except N1, the unphosphorylated precursor, which is present at much reduced levels. Mutations in the flight muscle-specific actin gene that block actin synthesis and prevent thin filament assembly disrupt the temporal regulation of flightin phosphorylation, resulting in premature phosphorylation and premature accumulation of flightin phosphovariants. Cellular fractionation of fibers that are devoid of thin filaments show that flightin remains associated with the thick filamentrich cytomatrix. These results suggest that flightin is a structural component of the thick filaments whose regulated phosphorylation is dependent upon the presence of thin filaments.This work was supported by National Science Foundation Grant IBN-9253045.     

Spatial heterogeneity in water velocity drives leaf litter dynamics in streams

1. Stream hydro-morphology refers to the heterogeneous distribution of hydrologic conditions that occur above a complex benthic surface such as a streambed.
2. We hypothesised that hydro-morphological conditions will influence the retention, re-distribution, and microbial-driven decomposition of leaf litter inputs in stream ecosystems because each process varies with overlying water velocity.
3. We tested this hypothesis using: (1) the spatial distribution of water velocity within a stream reach as a surrogate of stream hydro-morphology; (2) leaf tracer (i.e. Ginkgo biloba L.) additions with serial recovery to examine the relationship between benthic retention and overlying velocity; and (3) measurements of leaf litter decomposition (i.e. Alnus glutinosa [L.] Gaertn.) under different water velocity conditions.
4. Results demonstrate that water velocity exerts a significant influence on the retention and re-distribution of leaf litter inputs within the reach. The observed range of water velocity (from c. 0 to 0.92 cm/s) also strongly influences the range of leaf litter decomposition rates (0.0076–0.0222/day).
5. Our findings illustrate that water velocity influences leaf litter dynamics in streams by controlling leaf litter transport, retention and re-distribution as well as how leaves decompose within recipient stream reaches. Ultimately, the results show that the efficiency of leaf litter inputs in supporting stream ecosystem function is dependent on the hydro-morphological characteristics of the receiving stream ecosystems.

     

Root growth dynamics of Nicotiana attenuata seedlings are affected by simulated herbivore attack

Many studies demonstrate resource-based trade-offs between growth and defence on a large timescale. Yet, the short-term dynamics of this growth reaction are still completely unclear, making it difficult to explain growth-defence trade-offs mechanistically. In this study, image-based non-destructive methods were used to quantify root growth reactions happening within hours following simulated herbivore attack. The induction of wound reactions in Nicotiana attenuata in the seedling stage led to transiently decreased root growth rates. Application of the oral secretion of the specialist herbivore Manduca sexta to the leaves led to a transient decrease in root growth that was more pronounced than if a mere mechanical wounding was imposed. Root growth reduction was more pronounced than leaf growth reduction. When fatty acid-amino acid conjugates (FACs) were applied to wounds, root growth reduction occurred in the same intensity as when oral secretion was applied. Timing of the transient growth reduction coincided with endogenous bursts of jasmonate (JA) and ethylene emissions reported in literature. Simulation of a wound response by applying methyl jasmonate (MeJA) led to more prolonged negative effects on root growth. Increased nicotine concentrations, trichome lengths and densities were observed within 72 h in seedlings that were treated with MeJA or that were mechanically wounded. Overall, these reactions indicate that even in a very early developmental stage, the diversion of plant metabolism from primary (growth-sustaining) to secondary (defence-related) metabolism can cause profound alterations of plant growth performance.