Sridhar Ravi

Aerodynamics, sensing and control of insect-scale flapping-wing flight

There are nearly a million known species of flying insects and 13 000 species of flying warm-blooded vertebrates, including mammals, birds and bats. While in flight, their wings not only move forward relative to the air, they also flap up and down, plunge and sweep, so that both lift and thrust can be generated and balanced, accommodate uncertain surrounding environment, with superior flight stability and dynamics with highly varied speeds and missions. As the size of a flyer is reduced, the wing-to-body mass ratio tends to decrease as well. Furthermore, these flyers use integrated system consisting of wings to generate aerodynamic forces, muscles to move the wings, and sensing and control systems to guide and manoeuvre. In this article, recent advances in insect-scale flapping-wing aerodynamics, flexible wing structures, unsteady flight environment, sensing, stability and control are reviewed with perspective offered. In particular, the special features of the low Reynolds number flyers associated with small sizes, thin and light structures, slow flight with comparable wind gust speeds, bioinspired fabrication of wing structures, neuron-based sensing and adaptive control are highlighted.

Rolling with the flow: Bumblebees flying in unsteady wakes

SUMMARY Our understanding of how variable wind in natural environments affects flying insects is limited because most studies of insect flight are conducted in either smooth flow or still air conditions. Here, we investigate the effects of structured, unsteady flow (the von Karman vortex street behind a cylinder) on the flight performance of bumblebees (Bombus impatiens). Bumblebees are ‘all-weather’ foragers and thus frequently experience variable aerial conditions, ranging from fully mixed, turbulent flow to unsteady, structured vortices near objects such as branches and stems. We examined how bumblebee flight performance differs in unsteady versus smooth flow, as well as how the orientation of unsteady flow structures affects their flight performance, by filming bumblebees flying in a wind tunnel under various flow conditions. The three-dimensional flight trajectories and orientations of bumblebees were quantified in each of three flow conditions: (1) smooth flow, (2) the unsteady wake of a vertical cylinder (inducing strong lateral disturbances) and (3) the unsteady wake of a horizontal cylinder (inducing strong vertical disturbances). In both unsteady conditions, bumblebees attenuated the disturbances induced by the wind quite effectively, but still experienced significant translational and rotational fluctuations as compared with flight in smooth flow. Bees appeared to be most sensitive to disturbance along the lateral axis, displaying large lateral accelerations, translations and rolling motions in response to both unsteady flow conditions, regardless of orientation. Bees also displayed the greatest agility around the roll axis, initiating voluntary casting maneuvers and correcting for lateral disturbances mainly through roll in all flow conditions. Both unsteady flow conditions reduced the upstream flight speed of bees, suggesting an increased cost of flight in unsteady flow, with potential implications for foraging patterns and colony energetics in natural, variable wind environments.

Biomechanics and biomimetics in insect-inspired flight systems

Insect- and bird-size drones—micro air vehicles (MAV) that can perform autonomous flight in natural and man-made environments are now an active and well-integrated research area. MAVs normally operate at a low speed in a Reynolds number regime of 104–105 or lower, in which most flying animals of insects, birds and bats fly, and encounter unconventional challenges in generating sufficient aerodynamic forces to stay airborne and in controlling flight autonomy to achieve complex manoeuvres. Flying insects that power and control flight by flapping wings are capable of sophisticated aerodynamic force production and precise, agile manoeuvring, through an integrated system consisting of wings to generate aerodynamic force, muscles to move the wings and a control system to modulate power output from the muscles. In this article, we give a selective review on the state of the art of biomechanics in bioinspired flight systems in terms of flapping and flexible wing aerodynamics, flight dynamics and stability, passive and active mechanisms in stabilization and control, as well as flapping flight in unsteady environments. We further highlight recent advances in biomimetics of flapping-wing MAVs with a specific focus on insect-inspired wing design and fabrication, as well as sensing systems. This article is part of the themed issue ‘Moving in a moving medium: new perspectives on flight’.

Bumblebee flight performance in cluttered environments: effects of obstacle orientation, body size and acceleration.

ABSTRACT Locomotion through structurally complex environments is fundamental to the life history of most flying animals, and the costs associated with movement through clutter have important consequences for the ecology and evolution of volant taxa. However, few studies have directly investigated how flying animals navigate through cluttered environments, or examined which aspects of flight performance are most critical for this challenging task. Here, we examined how body size, acceleration and obstacle orientation affect the flight of bumblebees in an artificial, cluttered environment. Non-steady flight performance is often predicted to decrease with body size, as a result of a presumed reduction in acceleration capacity, but few empirical tests of this hypothesis have been performed in flying animals. We found that increased body size is associated with impaired flight performance (specifically transit time) in cluttered environments, but not with decreased peak accelerations. In addition, previous studies have shown that flying insects can produce higher accelerations along the lateral body axis, suggesting that if maneuvering is constrained by acceleration capacity, insects should perform better when maneuvering around objects laterally rather than vertically. Our data show that bumblebees do generate higher accelerations in the lateral direction, but we found no difference in their ability to pass through obstacle courses requiring lateral versus vertical maneuvering. In sum, our results suggest that acceleration capacity is not a primary determinant of flight performance in clutter, as is often assumed. Rather than being driven by the scaling of acceleration, we show that the reduced flight performance of larger bees in cluttered environments is driven by the allometry of both path sinuosity and mean flight speed. Specifically, differences in collision-avoidance behavior underlie much of the variation in flight performance across body size, with larger bees negotiating obstacles more cautiously. Thus, our results show that cluttered environments challenge the flight capacity of insects, but in surprising ways that emphasize the importance of behavioral and ecological context for understanding flight performance in complex environments. Highlighted Article: Large bumblebees exhibit impaired flight performance in cluttered environments compared with small bees; this is not due to the scaling of acceleration performance, however, and may be driven by the allometry of collision avoidance.

Hummingbird flight stability and control in freestream turbulent winds

ABSTRACT Airflow conditions close to the Earths surface are often complex, posing challenges to flight stability and control for volant taxa. Relatively little is known about how well flying animals can contend with complex, adverse air flows, or about the flight control mechanisms used by animals to mitigate wind disturbances. Several recent studies have examined flight in the unsteady von Kármán vortex streets that form behind cylinders, generating flow disturbances that are predictable in space and time; these structures are relatively rare in nature, because they occur only the immediate, downstream vicinity of an object. In contrast, freestream turbulence is characterized by rapid, unpredictable flow disturbances across a wide range of spatial and temporal scales, and is nearly ubiquitous in natural habitats. Hummingbirds are ideal organisms for studying the influence of freestream turbulence on flight, as they forage in a variety of aerial conditions and are powerful flyers. We filmed ruby-throated hummingbirds (Archilochus colubris) maintaining position at a feeder in laminar and strongly turbulent (intensity ∼15%) airflow environments within a wind tunnel and compared their mean kinematics of the head, body, tail and wing, as well as variability in these parameters. Hummingbirds exhibited remarkably stable head position and orientation in both smooth and turbulent flow while maintaining position at the feeder. However, the hummingbirds body was less stable in turbulent flow and appeared to be most sensitive to disturbances along the mediolateral axis, displaying large lateral accelerations, translations and rolling motions during flight. The hummingbirds mitigated these disturbances by increasing mean wing stroke amplitude and stroke plane angle, and by varying these parameters asymmetrically between the wings and from one stroke to the next. They also actively varied the orientation and fan angle of the tail, maintaining a larger mean fan angle when flying in turbulent flow; this may improve their passive stability, but probably incurs an energetic cost as a result of increased drag. Overall, we observed many of the same kinematic changes noted previously for hummingbirds flying in a von Kármán vortex street, but we also observed kinematic changes associated with high force production, similar to those seen during load-lifting or high-speed flight. These findings suggest that flight may be particularly costly in fully mixed, freestream turbulence, which is the flow condition that hummingbirds are likely to encounter most frequently in natural habitats. Summary: Hummingbirds are exceptional flyers that employ a combination of mean and transient flight control mechanisms to enable them to maintain flight in highly turbulent wind environments.

Nectar vs. pollen loading affects the tradeoff between flight stability and maneuverability in bumblebees

Significance Many insects carry substantial loads during flight. Although this behavior is often critical for survival and reproduction, few studies have examined its consequences for flight performance. Bumblebees regularly carry two types of loads in different locations: pollen on their legs or nectar in their abdomen. We found that when bees carry a pollen load rather than a nectar load, they are more stable but less maneuverable in flight, supporting the concept of a tradeoff between stability and maneuverability—a notion that is frequently suggested but has rarely been tested, particularly in flying insects. Our findings also suggest that wind conditions may influence what resources pollinators choose to collect, which would have important consequences for pollination dynamics and colony fitness. Bumblebee foragers spend a significant portion of their lives transporting nectar and pollen, often carrying loads equivalent to more than half their body mass. Whereas nectar is stored in the abdomen near the bee’s center of mass, pollen is carried on the hind legs, farther from the center of mass. We examine how load position changes the rotational moment of inertia in bumblebees and whether this affects their flight maneuverability and/or stability. We applied simulated pollen or nectar loads of equal mass to Bombus impatiens bumblebees and examined flight performance in a wind tunnel under three conditions: flight in unsteady flow, tracking an oscillating flower in smooth flow, and flower tracking in unsteady flow. Using an inertial model, we estimated that carrying a load on the legs rather than in the abdomen increases a bee’s moment of inertia about the roll and yaw axes but not the pitch axis. Consistent with these predictions, we found that bees carrying a load on their legs displayed slower rotations about their roll and yaw axes, regardless of whether these rotations were driven by external perturbations or self-initiated steering maneuvers. This allowed pollen-loaded bees to maintain a more stable body orientation and higher median flight speed in unsteady flow but reduced their performance when tracking a moving flower, supporting the concept of a tradeoff between stability and maneuverability. These results demonstrate that the types of resources collected by bees affect their flight performance and energetics and suggest that wind conditions may influence resource selection.

Bumblebees minimize control challenges by combining active and passive modes in unsteady winds

The natural wind environment that volant insects encounter is unsteady and highly complex, posing significant flight-control and stability challenges. It is critical to understand the strategies insects employ to safely navigate in natural environments. We combined experiments on free flying bumblebees with high-fidelity numerical simulations and lower-order modeling to identify the mechanics that mediate insect flight in unsteady winds. We trained bumblebees to fly upwind towards an artificial flower in a wind tunnel under steady wind and in a von Kármán street formed in the wake of a cylinder. Analysis revealed that at lower frequencies in both steady and unsteady winds the bees mediated lateral movement with body roll - typical casting motion. Numerical simulations of a bumblebee in similar conditions permitted the separation of the passive and active components of the flight trajectories. Consequently, we derived simple mathematical models that describe these two motion components. Comparison between the free-flying live and modeled bees revealed a novel mechanism that enables bees to passively ride out high-frequency perturbations while performing active maneuvers at lower frequencies. The capacity of maintaining stability by combining passive and active modes at different timescales provides a viable means for animals and machines to tackle the challenges posed by complex airflows.

Influence of large-scale freestream turbulence on the performance of a thin airfoil

Micro air vehicles are typically small in size and are remotely controlled or autonomous aircraft that fly slowly (50; 000 < Reynolds number < 200; 000) and very close to the Earths surface. Due to the combined influences of low Reynolds number effects and the high levels of freestream turbulence, present within the atmospheric boundary layer, micro air vehicles are very difficult to control/fly outdoors.

The gust-mitigating potential of flapping wings

Natures flapping-wing flyers are adept at negotiating highly turbulent flows across a wide range of scales. This is in part due to their ability to quickly detect and counterract disturbances to their flight path, but may also be assisted by an inherent aerodynamic property of flapping wings. In this study, we subject a mechanical flapping wing to replicated atmospheric turbulence across a range of flapping frequencies and turbulence intensities. By means of flow visualization and surface pressure measurements, we determine the salient effects of large-scale freestream turbulence on the flow field, and on the phase-average and fluctuating components of pressure and lift. It is shown that at lower flapping frequencies, turbulence dominates the instantaneous flow field, and the random fluctuating component of lift contributes significantly to the total lift. At higher flapping frequencies, kinematic forcing begins to dominate and the flow field becomes more consistent from cycle to cycle. Turbulence still modulates the flapping-induced flow field, as evidenced in particular by a variation in the timing and extent of leading edge vortex formation during the early downstroke. The random fluctuating component of lift contributes less to the total lift at these frequencies, providing evidence that flapping wings do indeed provide some inherent gust mitigation.

The answer is blowing in the wind: free flying honeybees can integrate visual and mechano-sensory inputs for making complex foraging decisions

ABSTRACT Bees navigate in complex environments using visual, olfactory and mechano-sensorial cues. In the lowest region of the atmosphere, the wind environment can be highly unsteady and bees employ fine motor-skills to enhance flight control. Recent work reveals sophisticated multi-modal processing of visual and olfactory channels by the bee brain to enhance foraging efficiency, but it currently remains unclear whether wind-induced mechano-sensory inputs are also integrated with visual information to facilitate decision making. Individual honeybees were trained in a linear flight arena with appetitive–aversive differential conditioning to use a context-setting cue of 3 m s−1 cross-wind direction to enable decisions about either a ‘blue’ or ‘yellow’ star stimulus being the correct alternative. Colour stimuli properties were mapped in bee-specific opponent-colour spaces to validate saliency, and to thus enable rapid reverse learning. Bees were able to integrate mechano-sensory and visual information to facilitate decisions that were significantly different to chance expectation after 35 learning trials. An independent group of bees were trained to find a single rewarding colour that was unrelated to the wind direction. In these trials, wind was not used as a context-setting cue and served only as a potential distracter in identifying the relevant rewarding visual stimuli. Comparison between respective groups shows that bees can learn to integrate visual and mechano-sensory information in a non-elemental fashion, revealing an unsuspected level of sensory processing in honeybees, and adding to the growing body of knowledge on the capacity of insect brains to use multi-modal sensory inputs in mediating foraging behaviour. Summary: In complex environments, free-flying honeybees are able to integrate both visual and wind-induced mechano-sensory cues to make decisions in a way that is suggestive of integrative processing by the brain.