Leif Ristroph

Discovering the flight autostabilizer of fruit flies by inducing aerial stumbles

Just as the Wright brothers implemented controls to achieve stable airplane flight, flying insects have evolved behavioral strategies that ensure recovery from flight disturbances. Pioneering studies performed on tethered and dissected insects demonstrate that the sensory, neurological, and musculoskeletal systems play important roles in flight control. Such studies, however, cannot produce an integrative model of insect flight stability because they do not incorporate the interaction of these systems with free-flight aerodynamics. We directly investigate control and stability through the application of torque impulses to freely flying fruit flies (Drosophila melanogaster) and measurement of their behavioral response. High-speed video and a new motion tracking method capture the aerial “stumble,” and we discover that flies respond to gentle disturbances by accurately returning to their original orientation. These insects take advantage of a stabilizing aerodynamic influence and active torque generation to recover their heading to within 2° in < 60 ms. To explain this recovery behavior, we form a feedback control model that includes the fly’s ability to sense body rotations, process this information, and actuate the wing motions that generate corrective aerodynamic torque. Thus, like early man-made aircraft and modern fighter jets, the fruit fly employs an automatic stabilization scheme that reacts to short time-scale disturbances.

Automated hull reconstruction motion tracking (HRMT) applied to sideways maneuvers of free-flying insects

SUMMARY Flying insects perform aerial maneuvers through slight manipulations of their wing motions. Because such manipulations in wing kinematics are subtle, a reliable method is needed to properly discern consistent kinematic strategies used by the insect from inconsistent variations and measurement error. Here, we introduce a novel automated method that accurately extracts full, 3D body and wing kinematics from high-resolution films of free-flying insects. This method combines visual hull reconstruction, principal components analysis, and geometric information about the insect to recover time series data of positions and orientations. The technique has small, well-characterized errors of under 3 pixels for positions and 5 deg. for orientations. To show its utility, we apply this motion tracking to the flight of fruit flies, Drosophila melanogaster. We find that fruit flies generate sideways forces during some maneuvers and that strong lateral acceleration is associated with differences between the left and right wing angles of attack. Remarkably, this asymmetry can be induced by simply altering the relative timing of flips between the right and left wings, and we observe that fruit flies employ timing differences as high as 10% of a wing beat period while accelerating sideways at 40% g.

Active and passive stabilization of body pitch in insect flight.

Flying insects have evolved sophisticated sensory–motor systems, and here we argue that such systems are used to keep upright against intrinsic flight instabilities. We describe a theory that predicts the instability growth rate in body pitch from flapping-wing aerodynamics and reveals two ways of achieving balanced flight: active control with sufficiently rapid reactions and passive stabilization with high body drag. By glueing magnets to fruit flies and perturbing their flight using magnetic impulses, we show that these insects employ active control that is indeed fast relative to the instability. Moreover, we find that fruit flies with their control sensors disabled can keep upright if high-drag fibres are also attached to their bodies, an observation consistent with our prediction for the passive stability condition. Finally, we extend this framework to unify the control strategies used by hovering animals and also furnish criteria for achieving pitch stability in flapping-wing robots.

Dynamic self-assembly of microscale rotors and swimmers

Biological systems often involve the self-assembly of basic components into complex and functioning structures. Artificial systems that mimic such processes can provide a well-controlled setting to explore the principles involved and also synthesize useful micromachines. Our experiments show that immotile, but active, components self-assemble into two types of structure that exhibit the fundamental forms of motility: translation and rotation. Specifically, micron-scale metallic rods are designed to induce extensile surface flows in the presence of a chemical fuel; these rods interact with each other and pair up to form either a swimmer or a rotor. Such pairs can transition reversibly between these two configurations, leading to kinetics reminiscent of bacterial run-and-tumble motion.

Stable hovering of a jellyfish-like flying machine

Ornithopters, or flapping-wing aircraft, offer an alternative to helicopters in achieving manoeuvrability at small scales, although stabilizing such aerial vehicles remains a key challenge. Here, we present a hovering machine that achieves self-righting flight using flapping wings alone, without relying on additional aerodynamic surfaces and without feedback control. We design, construct and test-fly a prototype that opens and closes four wings, resembling the motions of swimming jellyfish more so than any insect or bird. Measurements of lift show the benefits of wing flexing and the importance of selecting a wing size appropriate to the motor. Furthermore, we use high-speed video and motion tracking to show that the body orientation is stable during ascending, forward and hovering flight modes. Our experimental measurements are used to inform an aerodynamic model of stability that reveals the importance of centre-of-mass location and the coupling of body translation and rotation. These results show the promise of flapping-flight strategies beyond those that directly mimic the wing motions of flying animals.

Hydrodynamic schooling of flapping swimmers

Fish schools and bird flocks are fascinating examples of collective behaviours in which many individuals generate and interact with complex flows. Motivated by animal groups on the move, here we explore how the locomotion of many bodies emerges from their flow-mediated interactions. Through experiments and simulations of arrays of flapping wings that propel within a collective wake, we discover distinct modes characterized by the group swimming speed and the spatial phase shift between trajectories of neighbouring wings. For identical flapping motions, slow and fast modes coexist and correspond to constructive and destructive wing–wake interactions. Simulations show that swimming in a group can enhance speed and save power, and we capture the key phenomena in a mathematical model based on memory or the storage and recollection of information in the flow field. These results also show that fluid dynamic interactions alone are sufficient to generate coherent collective locomotion, and thus might suggest new ways to characterize the role of flows in animal groups.

Sculpting of an erodible body by flowing water

Erosion by flowing fluids carves striking landforms on Earth and also provides important clues to the past and present environments of other worlds. In these processes, solid boundaries both influence and are shaped by the surrounding fluid, but the emergence of morphology as a result of this interaction is not well understood. We study the coevolution of shape and flow in the context of erodible bodies molded from clay and immersed in a fast, unidirectional water flow. Although commonly viewed as a smoothing process, we find that erosion sculpts pointed and cornerlike features that persist as the solid shrinks. We explain these observations using flow visualization and a fluid mechanical model in which the surface shear stress dictates the rate of material removal. Experiments and simulations show that this interaction ultimately leads to self-similarly receding boundaries and a unique front surface characterized by nearly uniform shear stress. This tendency toward conformity of stress offers a principle for understanding erosion in more complex geometries and flows, such as those present in nature.

Self-similar evolution of a body eroding in a fluid flow

Erosion of solid material by flowing fluids plays an important role in shaping landforms, and in this natural context is often dictated by processes of high complexity. Here, we examine the coupled evolution of solid shape and fluid flow within the idealized setting of a cylindrical body held against a fast, unidirectional flow, and eroding under the action of fluid shear stress. Experiments and simulations both show self-similar evolution of the body, with an emerging quasi-triangular geometry that is an attractor of the shape dynamics. Our fluid erosion model, based on Prandtl boundary layer theory, yields a scaling law that accurately predicts the bodys vanishing rate. Further, a class of exact solutions provides a partial prediction for the bodys terminal form as one with a leading surface of uniform shear stress. Our simulations show this predicted geometry to emerge robustly from a range of different initial conditions, and allow us to explore its local stability. The sharp, faceted features of th...

Dynamics, Control, and Stabilization of Turning Flight in Fruit Flies

Complex behaviors of flying insects require interactions among sensory-neural systems, wing actuation biomechanics, and flapping-wing aerodynamics. Here, we review our recent progress in understanding these layers for maneuvering and stabilization flight of fruit flies. Our approach combines kinematic data from flying insects and aerodynamic simulations to distill reduced-order mathematical models of flight dynamics, wing actuation mechanisms, and control and stabilization strategies. Our central findings include: (1) During in-flight turns, fruit flies generate torque by subtly modulating wing angle of attack, in effect paddling to push off the air; (2) These motions are generated by biasing the orientation of a biomechanical brake that tends to resist rotation of the wing; (3) A simple and fast sensory-neural feedback scheme determines this wing actuation and thus the paddling motions needed for stabilization of flight heading against external disturbances. These studies illustrate a powerful approach for studying the integration of sensory-neural feedback, actuation, and aerodynamic strategies used by flying insects.

Fjords in viscous fingering: Selection of width and opening angle

Our experiments on viscous fingering of air into oil contained between closely spaced plates reveal two selection rules for the fjords of oil that separate fingers of air. (Fjords are the building blocks of solutions of the zero-surface-tension Laplacian growth equation.) Experiments in rectangular and circular geometries yield fjords with base widths lambda(c)/2, where lambda(c) is the most unstable wavelength from a linear stability analysis. Further, fjords open at an angle of 8.0 degrees +/- 1.0 degree. These selection rules hold for a wide range of pumping rates and fjord lengths, widths, and directions.