Visual control of flight speed in Drosophila melanogaster

Flight control in insects depends on self-induced image motion (optic flow), which the visual system must process to generate appropriate corrective steering maneuvers. Classic experiments in tethered insects applied rigorous system identification techniques for the analysis of turning reactions in the presence of rotating pattern stimuli delivered in open-loop. However, the functional relevance of these measurements for visual free-flight control remains equivocal due to the largely unknown effects of the highly constrained experimental conditions. To perform a systems analysis of the visual flight speed response under free-flight conditions, we implemented a `one-parameter open-loop' paradigm using `TrackFly' in a wind tunnel equipped with real-time tracking and virtual reality display technology. Upwind flying flies were stimulated with sine gratings of varying temporal and spatial frequencies, and the resulting speed responses were measured from the resulting flight speed reactions. To control flight speed, the visual system of the fruit fly extracts linear pattern velocity robustly over a broad range of spatio–temporal frequencies. The speed signal is used for a proportional control of flight speed within locomotor limits. The extraction of pattern velocity over a broad spatio–temporal frequency range may require more sophisticated motion processing mechanisms than those identified in flies so far. In Drosophila, the neuromotor pathways underlying flight speed control may be suitably explored by applying advanced genetic techniques, for which our data can serve as a baseline. Finally, the high-level control principles identified in the fly can be meaningfully transferred into a robotic context, such as for the robust and efficient control of autonomous flying micro air vehicles

The production of elevated flight force compromises manoeuvrability in the fruit fly Drosophila melanogaster

In this study, we have investigated how enhanced total flight force production compromises steering performance in tethered flying fruit flies, Drosophila melanogaster. The animals were flown in a closed-loop virtual-reality flight arena in which they modulated total flight force production in response to vertically oscillating visual patterns. By simultaneously measuring stroke amplitude and stroke frequency, we recorded the ability of each fly to modulate its wing kinematics at different levels of aerodynamic force production. At a flight force that exactly compensates body weight, the temporal deviations with which fruit flies vary their stroke amplitude and frequency are approximately 27° and 4.8 Hz of their mean value, respectively. This variance in wing kinematics decreases with increasing flight force production, and at maximum force production fruit flies are restricted to a unique combination of stroke amplitude, stroke frequency and mean force coefficient. This collapse in the kinematic envelope during peak force production could greatly attenuate the manoeuvrability and stability of animals in free flight

Spatial organization of visuomotor reflexes in Drosophila

In most animals, the visual system plays a central role in locomotor guidance. Here, we examined the functional organization of visuomotor reflexes in the fruit fly, Drosophila, using an electronic flight simulator. Flies exhibit powerful avoidance responses to visual expansion centered laterally. The amplitude of these expansion responses is three times larger than those generated by image rotation. Avoidance of a laterally positioned focus of expansion emerges from an inversion of the optomotor response when motion is restricted to the rear visual hemisphere. Furthermore, motion restricted to rear quarter-fields elicits turning responses that are independent of the direction of image motion about the animal's yaw axis. The spatial heterogeneity of visuomotor responses explains a seemingly peculiar behavior in which flies robustly fixate the contracting pole of a translating flow field

Flight Mechanics and Control of Escape Manoeuvres in Hummingbirds. I. Flight Kinematics

Hummingbirds are nature’s masters of aerobatic manoeuvres. Previous research shows that hummingbirds and insects converged evolutionarily upon similar aerodynamic mechanisms and kinematics in hovering. Herein, we use three-dimensional kinematic data to begin to test for similar convergence of kinematics used for escape flight and to explore the effects of body size upon manoeuvring. We studied four hummingbird species in North America including two large species (magnificent hummingbird, Eugenes fulgens, 7.8 g, and blue-throated hummingbird, Lampornis clemenciae, 8.0 g) and two smaller species (broad-billed hummingbird, Cynanthus latirostris, 3.4 g, and black-chinned hummingbirds Archilochus alexandri, 3.1 g). Starting from a steady hover, hummingbirds consistently manoeuvred away from perceived threats using a drastic escape response that featured body pitch and roll rotations coupled with a large linear acceleration. Hummingbirds changed their flapping frequency and wing trajectory in all three degrees of freedom on a stroke-by-stroke basis, likely causing rapid and significant alteration of the magnitude and direction of aerodynamic forces. Thus it appears that the flight control of hummingbirds does not obey the ‘helicopter model’ that is valid for similar escape manoeuvres in fruit flies. Except for broad-billed hummingbirds, the hummingbirds had faster reaction times than those reported for visual feedback control in insects. The two larger hummingbird species performed pitch rotations and global-yaw turns with considerably larger magnitude than the smaller species, but roll rates and cumulative roll angles were similar among the four species

Moth-inspired navigation algorithm in a turbulent odor plume from a pulsating source

Some female moths attract male moths by emitting series of pulses of pheromone filaments propagating downwind. The turbulent nature of the wind creates a complex flow environment, and causes the filaments to propagate in the form of patches with varying concentration distributions. Inspired by moth navigation capabilities, we propose a navigation strategy that enables a flier to locate a pulsating odor source in a windy environment using a single threshold-based detection sensor. The strategy is constructed based on the physical properties of the turbulent flow carrying discrete puffs of odor and does not involve learning, memory, complex decision making or statistical methods. We suggest that in turbulent plumes from a pulsating point source, an instantaneously measurable quantity referred as a "puff crossing time", improves the success rate as compared to the navigation strategy based on "internal counter" that does not use this information. Using computer simulations of fliers navigating in turbulent plumes of the pulsating point source for varying flow parameters: turbulent intensities, plume meandering and wind gusts, we obtained trajectories qualitatively resembling male moths flights towards the pheromone sources. We quantified the probability of a successful navigation as well as the flight parameters such as the time spent searching and the total flight time, with respect to different turbulent intensities, meandering or gusts. The concepts learned using this model may help to design odor-based navigation of miniature airborne autonomous vehicles

Living IoT: A Flying Wireless Platform on Live Insects

Sensor networks with devices capable of moving could enable applications ranging from precision irrigation to environmental sensing. Using mechanical drones to move sensors, however, severely limits operation time since flight time is limited by the energy density of current battery technology. We explore an alternative, biology-based solution: integrate sensing, computing and communication functionalities onto live flying insects to create a mobile IoT platform. Such an approach takes advantage of these tiny, highly efficient biological insects which are ubiquitous in many outdoor ecosystems, to essentially provide mobility for free. Doing so however requires addressing key technical challenges of power, size, weight and self-localization in order for the insects to perform location-dependent sensing operations as they carry our IoT payload through the environment. We develop and deploy our platform on bumblebees which includes backscatter communication, low-power self-localization hardware, sensors, and a power source. We show that our platform is capable of sensing, backscattering data at 1 kbps when the insects are back at the hive, and localizing itself up to distances of 80 m from the access points, all within a total weight budget of 102 mg.Comment: Co-primary authors: Vikram Iyer, Rajalakshmi Nandakumar, Anran Wang, In Proceedings of Mobicom. ACM, New York, NY, USA, 15 pages, 201

The wake dynamics and flight forces of the fruit fly Drosophila melanogaster

We have used flow visualizations and instantaneous force measurements of tethered fruit flies (Drosophila melanogaster) to study the dynamics of force generation during flight. During each complete stroke cycle, the flies generate one single vortex loop consisting of vorticity shed during the downstroke and ventral flip. This gross pattern of wake structure in Drosophila is similar to those described for hovering birds and some other insects. The wake structure differed from those previously described, however, in that the vortex filaments shed during ventral stroke reversal did not fuse to complete a circular ring, but rather attached temporarily to the body to complete an inverted heart-shaped vortex loop. The attached ventral filaments of the loop subsequently slide along the length of the body and eventually fuse at the tip of the abdomen. We found no evidence for the shedding of wing-tip vorticity during the upstroke, and argue that this is due to an extreme form of the Wagner effect acting at that time. The flow visualizations predicted that maximum flight forces would be generated during the downstroke and ventral reversal, with little or no force generated during the upstroke. The instantaneous force measurements using laser-interferometry verified the periodic nature of force generation. Within each stroke cycle, there was one plateau of high force generation followed by a period of low force, which roughly correlated with the upstroke and downstroke periods. However, the fluctuations in force lagged behind their expected occurrence within the wing-stroke cycle by approximately 1 ms or one-fifth of the complete stroke cycle. This temporal discrepancy exceeds the range of expected inaccuracies and artifacts in the measurements, and we tentatively discuss the potential retarding effects within the underlying fluid mechanics

Enhanced flight performance by genetic manipulation of wing shape in Drosophila

Insect wing shapes are remarkably diverse and the combination of shape and kinematics determines both aerial capabilities and power requirements. However, the contribution of any specific morphological feature to performance is not known. Using targeted RNA interference to modify wing shape far beyond the natural variation found within the population of a single species, we show a direct effect on flight performance that can be explained by physical modelling of the novel wing geometry. Our data show that altering the expression of a single gene can significantly enhance aerial agility and that the Drosophila wing shape is not, therefore, optimized for certain flight performance characteristics that are known to be important. Our technique points in a new direction for experiments on the evolution of performance specialities in animals

Flight Mechanics and Control of Escape Manoeuvres in Hummingbirds. II. Aerodynamic Force Production, Flight Control and Performance Limitations

The superior manoeuvrability of hummingbirds emerges from complex interactions of specialized neural and physiological processes with the unique flight dynamics of flapping wings. Escape manoeuvring is an ecologically relevant, natural behaviour of hummingbirds, from which we can gain understanding into the functional limits of vertebrate locomotor capacity. Here, we extend our kinematic analysis of escape manoeuvres from a companion paper to assess two potential limiting factors of the manoeuvring performance of hummingbirds: (1) muscle mechanical power output and (2) delays in the neural sensing and control system. We focused on the magnificent hummingbird (Eugenes fulgens, 7.8 g) and the black-chinned hummingbird (Archilochus alexandri, 3.1 g), which represent large and small species, respectively. We first estimated the aerodynamic forces, moments and the mechanical power of escape manoeuvres using measured wing kinematics. Comparing active-manoeuvring and passive-damping aerodynamic moments, we found that pitch dynamics were lightly damped and dominated by the effect of inertia, while roll dynamics were highly damped. To achieve observed closed-loop performance, pitch manoeuvres required faster sensorimotor transduction, as hummingbirds can only tolerate half the delay allowed in roll manoeuvres. Accordingly, our results suggested that pitch control may require a more sophisticated control strategy, such as those based on prediction. For the magnificent hummingbird, we estimated that escape manoeuvres required muscle mass-specific power 4.5 times that during hovering. Therefore, in addition to the limitation imposed by sensorimotor delays, muscle power could also limit the performance of escape manoeuvres

Locomotion response of airborne, ambulatory and aquatic insects to thermal stimulation using piezoceramic microheaters

This paper reports the locomotion response of airborne, ambulatory and aquatic insects to thermal stimulation. A finite element model has been developed to predict the variation of insect–stimulator interface temperature with input power. Piezothermal stimulators have been fabricated from lead zirconate titanate (PZT) using a batch mode micro ultrasonic machining process. Typical sizes range from 200 µm to 3.2 mm. For PZT stimulators, the temperature and thermal efficiency reach the maximum value around the resonance frequency which is typically in the range of 650 kHz to 47 MHz. Experiments have been conducted on green June beetles (GJBs), Madagascar hissing roaches and green diving beetles (GDBs) in order to show the versatility of the proposed technique. The stimulators have been implanted near the antennae of the GJBs and on either side of the thorax of the Madagascar hissing roaches and GDBs, respectively. In all cases, the insects move away from the direction of the actuated stimulator. The left and right turns are statistically similar. Thermal stimulation achieves an overall success rate of 78.7%, 92.8% and 61.6% in GJBs, roaches and GDBs, respectively. On average, thermal stimulation results in an angle turn of about 13.7°–16.2° on GJBs, 30°–45° on the roaches and 30°–50° on GDBs. The corresponding average input power is 360, 330 and 100 mW for GJBs, roach and GDBs, respectively. Scaling limits of the PZT stimulators for operating these stimulators are also discussed.Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/90797/1/0960-1317_21_12_125002.pd