Thorax vibrations of a stingless bee (<Emphasis Type="Italic">Melipona seminigra</Emphasis>). I. No influence of visual flow

An important question in stingless bee communication is whether the thorax vibrations produced by foragers of the genus Melipona upon their return to the nest contain spatial information about food sources or not. As previously shown M. seminigra is able to use visual flow to estimate flight distances. The present study investigated whether foraging bees encode the visually measured distance in their thorax vibrations. Bees were trained to collect food in flight tunnels lined with a black-and-white pattern on their side walls and floor, which substantially influenced the image motion they experienced. When the bees had collected inside the tunnels the temporal pattern of their vibrations differed significantly from the pattern after collecting in a natural environment. These changes, however, were not associated with the visual flow experienced inside the tunnel. Bees collecting in tunnels offering little visual flow (stripes parallel to flight direction) modified their vibrations similarly to bees collecting in tunnels with high image motion (cross stripes). A higher energy expenditure due to drastically reduced flight velocities inside the tunnel is suggested to be responsible for changes in the thorax vibrations. The bees' vibrations would thus reflect the overall energetic budget of a foraging trip.     

Bee vision of pattern and 3D. The Bidder Lecture 1994

Insect vision is nothing if not active. The regular head movements, called saccades, enable the fly Drosophila to keep a straight path in flight despite inequalities in the thrust of the wings. Using their own motion, bees in flight measure the ranges of nearby objects. A long history of research shows that bees discriminate visually in ways that depend on their activity or task, so we must distinguish between vision during flying, fixating or hovering and landing. Bees return again and again for a reward of sugar solution and use their eyes to find their way. In an apparatus that makes them discriminate between two simulataneously visible but regularly interchanged targets, seen at a distance of 27 cm, bees are able to distinguish a remarkable number of simple patterns, but they fail in certain critical cases. The results can be explained with the hypothesis that bees have several broadly tuned overlapping filters with large fields that respond to the predominant orientation in a region of the image, and others for radial and circular patterns. Together with colour, these filters are independent of range. Bees prefer to use landmarks where they can, then global pattern at the largest scale, and lastly the detail around the goal. The way that discrimination of one visual feature is independent of other variables can be explained by models analogous to the colour triangle in colour discrimination.     

Influence of flight time and flight environment on distance communication by dancing honey bees

Forager honey bees communicate the distance of food sources to nest mates through waggle dances, but how do bees measure these distances? Recent work suggests that bees measure distance flown in terms of the extent of image motion (integrated optic flow) that is experienced during flight. However, it is known that optic flow also regulates the speed of flight. Therefore, the duration of the flight to a destination is likely to co-vary with the optic flow that is experienced en route. This makes it difficult to tease apart the potential roles of flight duration and optic flow as cues in estimating distance flown. Here we examine whether flight duration alone can serve as an indicator of distance. We trained bees to visit feeders at two sites located in optically different environments, but positioned such that the flight durations to the two sites were similar. The distance estimates for the two sites, as reported in the waggle dance, were compared. We found that dances differed significantly between the two sites, even though flight times were similar. Flight time perse was a poor predictor of waggle dance behaviour. We conclude that foraging bees do not simply signal flight time as their measure of distance in the waggle dance; the environment through which they fly plays a dominant role. Received 11 April 2005; revised 16 May 2005; accepted 3 June 2005.     

The control of optic flow during learning flights

Hymenopteran insects perform systematic learning flights on departure from their nest, during which they acquire a visual representation of the nest environment. They back away from and pivot around the nest in a series of arcs while turning to view it in their fronto-lateral visual field. During the initial stages of the flights, turning rate and arc velocity relative to the nest are roughly constant at 100–200° s⁻¹ and are independent of distance, since the insects increase their flight speed as they back away from the pivoting centre. In this paper I analyse how solitary wasps control their flight by having them perform learning flights inside a rotating striped drum. The wasps' turning velocity is under visual control. When the insects fly inside a drum that rotates around the nest as a centre, their average turning rate is faster than normal when they fly an arc into the direction of drum rotation and slower when they fly in the opposite direction. The average slip speed they experience lies within 100–200° s⁻¹. The wasps also adjust their flight speed depending on the rotation of the drum. They modulate their distance from the pivoting centre accordingly and presumably also their height above ground, so that maximal ground slip is on average 200°␣s⁻¹. The insects move along arcs by short pulses of translation, followed by rapid body turns to correct for the change in retinal position of the nest entrance. Saccadic body turns follow pulses of translation with a delay of 80–120 ms. The optomotor response is active during these turns. The control of pivoting flight most likely involves three position servos, to control the retinal position of both the azimuth and the altitude of nest and the direction of flight relative to it, and two velocity servos, one constituting the optomotor reflex and the other one serving to clamp ground slip at about 200° s⁻¹. The control of ground slip is the prime source of the dynamic constancy of learning flights, which may help wasps to scale the pivoting parallax field they produce during these flights. Constant pivoting rate may in addition be important for the acquisition of a regular sequence of snapshots and in scanning for compass cues. Accepted : 31 July 1996     

A stingless bee (Melipona seminigra) uses optic flow to estimate flight distances

Foragers of a stingless bee, Melipona seminigra, are able to use the optic flow experienced en route to estimate flight distance. After training the bees to collect food inside a flight tunnel with black-and-white stripes covering the side walls and the floor, their search behavior was observed in tunnels lacking a reward. Like honeybees, the bees accurately estimated the distance to the previously offered food source as seen from the sections of the tunnel where they turned around in search of the food. Changing the visual flow by decreasing the width of the flight tunnel resulted in the underestimation of the distance flown. The removal of image motion cues either in the ventral or lateral field of view reduced the bees' ability to gauge distances. When the feeder inside the tunnel was displaced together with the bees feeding on it while preventing the bee from seeing any image motion during the displacement the bees experienced different distances on their way to the food source and during their return to the nest. In the subsequent test the bees searched for the food predominantly at the distance associated with their return flight.     

The effect of optic flow cues on honeybee flight control in wind

To minimize the risk of colliding with the ground or other obstacles, flying animals need to control both their ground speed and ground height. This task is particularly challenging in wind, where head winds require an animal to increase its airspeed to maintain a constant ground speed and tail winds may generate negative airspeeds, rendering flight more difficult to control. In this study, we investigate how head and tail winds affect flight control in the honeybee Apis mellifera, which is known to rely on the pattern of visual motion generated across the eye—known as optic flow—to maintain constant ground speeds and heights. We find that, when provided with both longitudinal and transverse optic flow cues (in or perpendicular to the direction of flight, respectively), honeybees maintain a constant ground speed but fly lower in head winds and higher in tail winds, a response that is also observed when longitudinal optic flow cues are minimized. When the transverse component of optic flow is minimized, or when all optic flow cues are minimized, the effect of wind on ground height is abolished. We propose that the regular sidewards oscillations that the bees make as they fly may be used to extract information about the distance to the ground, independently of the longitudinal optic flow that they use for ground speed control. This computationally simple strategy could have potential uses in the development of lightweight and robust systems for guiding autonomous flying vehicles in natural environments.     

Effects of flight behaviour on body temperature and kinematics during inter-male mate competition in the solitary desert bee Centris pallida

Abstract. Body temperatures and kinematics are measured for male Centris pallida bees engaged in a variety of flight behaviours (hovering, patrolling, pursuit) at a nest aggregation site in the Sonoran Desert. The aim of the study is to test for evidence of thermoregulatory variation in convective heat loss and metabolic heat production and to assess the mechanisms of acceleration and forward flight in field conditions. Patrolling males have slightly (1–3 °C) cooler body temperatures than hoverers, despite similar wingbeat frequencies and larger body masses, suggesting that convective heat loss is likely to be greater during patrolling flight than during hovering. Comparisons of thorax and head temperature as a function of air temperature (T_a) indicate that C. pallida males are thermoregulating the head by increasing heat transfer from the thorax to the head at cool T_a. During patrolling flight and hovering, wingbeat frequency significantly decreases as T_a increases, indicating that variation in metabolic heat production contributes to thermal stability during these behaviours, as has been previously demonstrated for this species during flight in a metabolic chamber. However, wingbeat frequency during brief (1–2 s) pursuits is significantly higher than during other flight behaviours and independent of T_a. Unlike most other hovering insects, C. pallida males hover with extremely inclined stroke plane angles and nearly horizontal body angles, suggesting that its ability to vary flight speed depends on changes in wingbeat frequency and other kinematic mechanisms that are not yet described.     

Maximal horizontal flight performance of hummingbirds: effects of body mass and molt

Hovering and fast forward flapping represent two strenuous types of flight that differ in aerodynamic power requirement. Maximal capabilities of ruby-throated hummingbirds (Archilochus colubris) in hovering and forward flight were compared under varying body mass and wing area. The capability to hover in low-density gas mixtures was adversely affected by body mass, whereas the capability to fly in a wind tunnel did not show any adverse mass effect. Molting birds that lost primary flight feathers and reduced wing area also displayed mass loss and loss of aerodynamic power and flight speed. This suggests that maximal flight speed is insensitive to short-term perturbations of body mass but that molting is stressful and reduces the birds' speed and capacity for chase and escape. Hummingbirds' flight behavior in confined space was also investigated. Birds reduced their speeds flying through a narrow tube to approximately one-fifth of that in the wind tunnel and did not display differences under varying body mass and wing area. Hence, performance in the flight tube was submaximal and did not correlate with performance variation in the wind tunnel. For ruby-throated hummingbirds, both maximal mass-specific aerodynamic power derived from hovering performance in low-density air media and maximal flight velocity measured in the wind tunnel were invariant with body mass.     

On the mechanism of speed and altitude control in Drosophila melanogaster

ABSTRACT The total power output of tethered flying Drosophila melanogaster in still air depends on translational velocity components of image flow on the eye, whereas the orientation of the average flight force in the midsagittal plane of the fly is widely independent of visual input (Götz, 1968). The fly does not seem to control the vertical and the horizontal force component independently. Freely flying flies nevertheless generate different ratios between lift and thrust, simply by changing the inclination of their body. By the combined adjustment of the body angle and the total power output a fly appears to be able to stabilize height and speed (David, 1985). Here a possible mechanism is proposed by which the appropriate torque about the transverse body axis could be generated. Translational pattern motion influences the posture of the abdomen and the plane of wing oscillation. Thus the position of the centre of gravity relative to the flight force vector is changed. When abdomen and stroke plane deviate from an equilibrium state, a lever is generated by which the force vector will rotate the fly about its transverse axis.     

A Mathematical Model for Flight Guidance in Honeybee Swarms

When a colony of honeybees relocates to a new nest site, less than 5?% of the bees (the scout bees) know the location of the new nest. Nevertheless, the small minority of informed bees manages to provide guidance to the rest and the entire swarm is able to fly to the new nest intact. The streaker bee hypothesis, one of the several theories proposed to explain the guidance mechanism in bee swarms, seems to be supported by recent experimental observations. The theory suggests that the informed bees make high-speed flights through the swarm in the direction of the new nest, hence conspicuously pointing to the desired direction of travel. This work presents a mathematical model of flight guidance in bee swarms based on the streaker bee hypothesis. Numerical experiments, parameter studies, and comparison with experimental data are presented.     

Orientation flights of solitary wasps (Cerceris; Sphecidae; Hymenoptera)

Bees and wasps are known to use a visual representation of the nest environment to guide the final approach to their nest. It is also known that they acquire this representation during an orientation flight performed on departure.A detailed film analysis shows that orientation flights in solitary wasps of the genus Cerceris consist of a systematic behavioural sequence: after lift-off from the nest entrance, wasps fly in ever increasing arcs around the nest. They fly along these arcs obliquely to their long axis and turn so that the nest entrance is held in the left or right visual field at retinal positions between 30° and 70° from the midline. Horizontal distance from the nest and height above ground increase throughout an orientation flight so that the nest is kept at retinal elevations between 45° and 60° below the horizon. The wasps' rate of turning is constant at between 100°/s and 200°/s independent of their distance from the nest and their ground velocity increases with distance. The consequence of this is that throughout the flight wasps circle at a constant angular velocity around the nest.Orientation flights are strongly influenced by landmark lay-out. Wasps adjust their flight-path and their orientation in a way that allows them to fixate the nest entrance and to hold the closest landmark in their frontal visual field.The orientation flight generates a specific topography of motion parallax across the visual field. This could be used by wasps to acquire a series of snapshots that all contain the nest position, to acquire snapshots of close landmarks only (distance filtering), to exclude shadow contours from their visual representation (figure-ground discrimination) or to gain information on the distance of landmarks relative to the nest.     

Optic flow cues guide flight in birds

Although considerable effort has been devoted to investigating how birds migrate over large distances, surprisingly little is known about how they tackle so successfully the moment-to-moment challenges of rapid flight through cluttered environments [1]. It has been suggested that birds detect and avoid obstacles [2] and control landing maneuvers [3-5] by using cues derived from the image motion that is generated in the eyes during flight. Here we investigate the ability of budgerigars to fly through narrow passages in a collision-free manner, by filming their trajectories during flight in a corridor where the walls are decorated with various visual patterns. The results demonstrate, unequivocally and for the first time, that birds negotiate narrow gaps safely by balancing the speeds of image motion that are experienced by the two eyes and that the speed of flight is regulated by monitoring the speed of image motion that is experienced by the two eyes. These findings have close parallels with those previously reported for flying insects [6-13], suggesting that some principles of visual guidance may be shared by all diurnal, flying animals.     

Ambush Predation of Stingless Bees (<Emphasis Type="Italic">Tetragonisca angustula</Emphasis>) by the Solitary-Foraging Ant <Emphasis Type="Italic">Ectatomma tuberculatum</Emphasis>

Social insect colonies are high-value foraging targets for insectivores, prompting the evolution of complex colony defensive adaptations as well as specialized foraging tactics in social insect predators. Predatory ants that forage on other social insects employ a diverse range of behaviors targeted at specific prey species. Here, we describe a solitary foraging strategy of the ant Ectatomma tuberculatum, on nest guards of the stingless bee Tetragonisca angustula. We observed multiple instances of E. tuberculatum ambushing and successfully capturing the hovering and standing guards of T. angustula near nest entrances. The unique hovering behavior of the guard caste of this bee species, an adaptation to frequent cleptoparasitism by other stingless bees, may make these guards particularly vulnerable to ground-based, ambush attacks by E. tuberculatum. Likewise, the behavior of the foraging ants appears to adaptively exploit the defensive formations and activity patterns of these bees. These observations suggest an adaptive and targeted predatory strategy aimed at gathering external guard bees as prey from these heavily fortified nests.     

凹唇壁蜂成蜂体重与取食对其飞行能力的影响

凹唇壁蜂Osmia excavata Alfken被广泛应用于我国北方果树的传粉, 而其飞行能力是影响其传粉效率的重要因素。本研究通过飞行磨吊飞试验, 评估了凹唇壁蜂雌蜂和雄蜂飞行能力的差异以及取食对其飞行能力的影响。结果表明, 凹唇壁蜂雌蜂体重（116.30 mg）显著大于雄蜂（59.80 mg） (P<0.001), 雌蜂的最大飞行速度（3.44 km/h）显著大于雄蜂（2.36 km/h）, 雄蜂的飞行距离和最大飞行速度与其飞行前体重成显著的正相关性, 雌蜂的飞行时间与其飞行前体重成显著正相关性, 而雌蜂的平均飞行速度与其体重成显著负相关性; 雌蜂的日平均飞行距离为0.23 km, 根据雌蜂以巢为中心, 采集花粉繁殖后代的生物学习性, 蜂巢之间的放置距离应少于100 m。取食蜂蜜后, 雌雄壁蜂的飞行距离、 飞行时间、 最大飞行速度均有提高的趋势, 建议在田间应用时, 可在蜂巢附近放置蜂蜜或种植其他蜜源植物给初羽化的凹唇壁蜂提供食物补充能量。本研究明确了雌、 雄壁蜂的飞行能力和出茧后补充食物对于壁蜂飞行的促进作用, 为有效地利用凹唇壁蜂进行传粉提供了理论依据。     

Visuomotor Response to Object Expansion in Free-Flying Bumble Bees

The flight control systems of flying insects enable many kinds of sophisticated maneuvers, including avoidance of midair collisions. Visuomotor response to an approaching object, received as image expansion on insects’ retina, is a complex event in a dynamic environment where both animals and objects are moving. There are intensive free flight studies on the landing response in which insects receive image expansion by their own movement. However, few studies have been conducted regarding how freely flying insects respond to approaching objects. Here, using common laboratory insects for behavioral research, the bumblebee Bombus ignitus, we examined their visual response to an approaching object in the free-flying condition. While the insect was slowly flying in a free-flight arena, an expanding stripe was projected laterally from one side of the arena with a high-speed digital mirror device projector. Rather than turning away reported before, the bumble bees performed complex flight maneuvers. We synchronized flight trajectories, orientations and wing stroke frequencies with projection parameters of temporal resolution in 0.5 ms, and analyzed the instantaneous relationship between visual input and behavioral output. In their complex behavioral responses, we identified the following two visuomotor behaviors: increasing stroke frequency when the bumble bees confront the stripe expansion, and turning towards (not away) the stripe expansion when it is located laterally to the bee. Our results suggested that the response to object expansion is not a simple and reflexive escape but includes object fixation, presumably for subsequent behavioral choice.     

Ground-nesting bees determine the location of their nest relative to a landmark by other than angular size cues

Bees and wasps acquire a visual representation of their nest's environment and use it to locate their nest when they return from foraging trips. This representation contains among other features cues to the distance of near-by landmarks. We worked with two species of ground-nesting bees, Lasioglossum malachurum (Hymenoptera: Halictidae), Dasypoda hirtipes (Hymenoptera: Melittidae) and asked which cues to landmark distance they use during homing. Bees learned to associate a single cylindrical landmark with their nest's location. We subsequently tested returning bees with landmarks of different sizes and thus introduced large discrepancies between the angular size of the landmark as seen from the nest during training and its distance from the nest. The bees' search behaviour and their choice of dummy nest entrances show that both species of ground-nesting bees consistently search for their nest at the learned distance from landmarks. The influence of the apparent size of landmarks on the bees' search and choice behaviour is comparatively weak. We suggest that the bees exploit cues derived from the apparent speed of the landmark's image at their retina for distance evaluation.     

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 scanning behaviour in honeybees

Freely flying bees were rewarded with sugar solution on a variety of black-and-white shapes as well as on coloured gratings in various training situations. In subsequent dual-choice tests, the bees' discrimination between the various shapes was measured. In addition, the bees were video-filmed while flying in front of the shapes. The scanning patterns thus obtained were then quantified in order to characterize scanning behaviour and its relationship to the geometrical parameters of the scanned shapes, investigate whether scanning plays a role in pattern discrimination and examine the influence of training on the characteristics of scanning. The scanning patterns clearly mirror the contours of the scanned shape in all cases, i.e. the bees fly along the contours contained in the shape. This behaviour does not depend on whether the scanned shape is one that was previously rewarded, or one that is completely novel to the bees. Comparison of the results of quantifying the scanning patterns with the results of dual-choice tests reveals that scanning behaviour is independent of discrimination performance. On the average, horizontal scanning directions occur more often than vertical directions. Variations of the training situation produce measurable differences in scanning behavior. However, except in the case of vertical scanning on a vertical grating, these differences are quite small, indicating that following contours is a largely stereotyped behaviour. Horizontal gratings are very well discriminated from vertical ones even if they offer contrast to only one receptor type, i.e. blue or green, demonstrating that the direction of contours is visible to the pattern recognition system even under these conditions. However, vertical and horizontal coloured gratings offering only blue-contrast do not elicit contour-following, whereas gratings offering only green-contrast do. Thus, the bees' scanning behaviour is colour-blind and most probably governed by the green receptors. We suggest that contour-following is the by-product of a behavioural mode which serves to prevent retinal image movement during flight in front of a contoured visual pattern.     

The ocelli control the flight course in honeybees

Abstract Fully-sighted honeybees and bees with all ocelli occluded were trained to fly through an arena to arrive at a feeding place. After training, the bees were exposed to side-light flashes during their feeding flights. The flight paths were recorded on video and analysed frame by frame at 40 ms intervals with reference to the main parameters, the coordinates of the thorax and the yaw angle of the bee. Course angles, translational course velocities and accelerations were calculated, and the responses to side light flashes evaluated with respect to 'on' and 'off.
Immediately after light on, fully-sighted bees respond slightly positively by yawing and flying toward the side light. Bees in which all ocelli are occluded are greatly disturbed and respond with negative yawing and flight path directions.
The ocelli apparently help to control phototactic alertness in the bee. They determine whether phototactic orienting or pattern-induced orienting behaviour is more important in a particular state of motivation. They help to minimize the level of disturbance in flight course control, obviously by activating a neuronal circuit with comparator attributes. It is assumed that this kind of compensation or suppression of phototactically guided reflexes occurs only for a few 100 ms. Consequently, the biological significance of light flashes shorter than 400 ms is very slight.
Fully-sighted bees decelerate strongly when a side light is switched on. Bees in which the ocelli are occluded behave less cautiously: they generally fly faster and need more reaction time. Thus, the ocelli help the bee to react photokinetically to photic stimuli in a much shorter time than do the compound eyes alone.