Michael H. Dickinson

Spanwise flow and the attachment of the leading-edge vortex on insect wings

The flow structure that is largely responsible for the good performance of insect wings has recently been identified as a leading-edge vortex. But because such vortices become detached from a wing in two-dimensional flow, an unknown mechanism must keep them attached to (three-dimensional) flapping wings. The current explanation, analogous to a mechanism operating on delta-wing aircraft, is that spanwise flow through a spiral vortex drains energy from the vortex core. We have tested this hypothesis by systematically mapping the flow generated by a dynamically scaled model insect while simultaneously measuring the resulting aerodynamic forces. Here we report that, at the Reynolds numbers matching the flows relevant for most insects, flapping wings do not generate a spiral vortex akin to that produced by delta-wing aircraft. We also find that limiting spanwise flow with fences and edge baffles does not cause detachment of the leading-edge vortex. The data support an alternative hypothesis—that downward flow induced by tip vortices limits the growth of the leading-edge vortex.

A Long-Term Depression of AMPA Currents in Cultured Cerebellar Purkinje Neurons

Cerebellar long-term depression (LTD) is a model of synaptic plasticity in which conjunctive stimulation of parallel fiber and climbing fiber inputs to a Purkinje neuron induces a persistent depression of the parallel fiber-Purkinje neuron synapse. We report that an analogous phenomenon may be elicited in the cultured mouse Purkinje neuron when iontophoretic glutamate application and depolarization of the Purkinje neurons are substituted for parallel fiber and climbing fiber stimulation, respectively. The induction of LTD in these cerebellar cultures requires activation of both ionotropic (AMPA) and metabotropic quisqualate receptors, together with depolarization in the presence of external Ca2+. This postsynaptic alteration is manifest as a depression of glutamate or AMPA currents, but not aspartate or NMDA currents. These results strengthen the contention that the expression of cerebellar LTD is at least in part postsynaptic and provide evidence that activation of both ionotropic and metabotropic quisqualate receptors are necessary for LTD induction.

High-throughput Ethomics in Large Groups of Drosophila

We present a camera-based method for automatically quantifying the individual and social behaviors of fruit flies, Drosophila melanogaster, interacting in a planar arena. Our system includes machine-vision algorithms that accurately track many individuals without swapping identities and classification algorithms that detect behaviors. The data may be represented as an ethogram that plots the time course of behaviors exhibited by each fly or as a vector that concisely captures the statistical properties of all behaviors displayed in a given period. We found that behavioral differences between individuals were consistent over time and were sufficient to accurately predict gender and genotype. In addition, we found that the relative positions of flies during social interactions vary according to gender, genotype and social environment. We expect that our software, which permits high-throughput screening, will complement existing molecular methods available in Drosophila, facilitating new investigations into the genetic and cellular basis of behavior.

Unsteady forces and flows in low Reynolds number hovering flight:two-dimensional computations vs robotic wing experiments

SUMMARY We compare computational, experimental and quasi-steady forces in a generic hovering wing undergoing sinusoidal motion along a horizontal stroke plane. In particular, we investigate unsteady effects and compare two-dimensional (2D) computations and three-dimensional (3D) experiments in several qualitatively different kinematic patterns. In all cases, the computed drag compares well with the experiments. The computed lift agrees in the cases in which the sinusoidal changes in angle of attack are symmetrical or advanced with respect to stroke positions, but lags behind the measured 3D lift in the delayed case. In the range of amplitudes studied here, 3–5 chords, the force coefficients have a weak dependence on stroke amplitude. As expected, the forces are sensitive to the phase between stroke angle and angle of attack, a result that can be explained by the orientation of the wing at reversal. This dependence on amplitude and phase suggests a simple maneuver strategy that could be used by a flapping wing device. In all cases the unsteady forces quickly reach an almost periodic state with continuous flapping. The fluid forces are dominated by the pressure contribution. The force component directly proportional to the linear acceleration is smaller by a factor proportional to the ratio of wing thickness and stroke amplitude; its net contribution is zero in hovering. The ratio of wing inertia and fluid force is proportional to the product of the ratio of wing and fluid density and the ratio of wing thickness and stroke amplitude; it is negligible in the robotic wing experiment, but need not be in insect flight. To identify unsteady effects associated with wing acceleration, and coupling between rotation and translation, as well as wake capture, we examine the difference between the unsteady forces and the estimates based on translational velocities, and compare them against the estimate of the coupling between rotation and translation, which have simple analytic forms for sinusoidal motions. The agreement and disagreement between the computed forces and experiments offer further insight into when the 3D effects are important. A main difference between a 3D revolving wing and a 2D translating wing is the absence of vortex shedding by a revolving wing over a distance much longer than the typical stroke length of insects. No doubt such a difference in shedding dynamics is responsible in part for the differences in steady state force coefficients measured in 2D and 3D. On the other hand, it is unclear whether such differences would have a significant effect on transient force coefficients before the onset of shedding. While the 2D steady state force coefficients underpredict 3D forces, the transient 2D forces measured prior to shedding are much closer to the 3D forces. In the cases studied here, the chord is moving between 3 to 5 chords, typical of hovering insect stroke length, and the flow does not appear to separate during each stroke in the cases of advanced and symmetrical rotation. In these cases, the wing reverses before the leading edge vortex would have time to separate even in 2D. This suggests that the time scale for flow separation in these strokes is dictated by the flapping frequency, which is dimensionally independent. In such cases, the 2D unsteady forces turn out to be good approximations of 3D experiments.

Force production and flow structure of the leading edge vortex on flapping wings at high and low Reynolds numbers

SUMMARY The elevated aerodynamic performance of insects has been attributed in part to the generation and maintenance of a stable region of vorticity known as the leading edge vortex (LEV). One explanation for the stability of the LEV is that spiraling axial flow within the vortex core drains energy into the tip vortex, forming a leading-edge spiral vortex analogous to the flow structure generated by delta wing aircraft. However, whereas spiral flow is a conspicuous feature of flapping wings at Reynolds numbers (Re) of 5000, similar experiments at Re=100 failed to identify a comparable structure. We used a dynamically scaled robot to investigate both the forces and the flows created by a wing undergoing identical motion at Re of∼ 120 and ∼1400. In both cases, motion at constant angular velocity and fixed angle of attack generated a stable LEV with no evidence of shedding. At Re=1400, flow visualization indicated an intense narrow region of spanwise flow within the core of the LEV, a feature conspicuously absent at Re=120. The results suggest that the transport of vorticity from the leading edge to the wake that permits prolonged vortex attachment takes different forms at different Re.

The influence of wing-wake interactions on the production of aerodynamic forces in flapping flight.

SUMMARY We used two-dimensional digital particle image velocimetry (DPIV) to visualize flow patterns around the flapping wing of a dynamically scaled robot for a series of reciprocating strokes starting from rest. The base of the wing was equipped with strain gauges so that the pattern of fluid motion could be directly compared with the time history of force production. The results show that the development and shedding of vortices throughout each stroke are highly stereotyped and influence force generation in subsequent strokes. When a wing starts from rest, it generates a transient force as the leading edge vortex (LEV) grows. This early peak, previously attributed to added-mass acceleration, is not amenable to quasi-steady models but corresponds well to calculations based on the time derivative of the first moment of vorticity within a sectional slice of fluid. Forces decay to a stable level as the LEV reaches a constant size and remains attached throughout most of the stroke. The LEV grows as the wing supinates prior to stroke reversal, accompanied by an increase in total force. At stroke reversal, both the LEV and a rotational starting vortex (RSV) are shed into the wake, forming a counter-rotating pair that directs a jet of fluid towards the underside of the wing at the start of the next stroke. We isolated the aerodynamic influence of the wake by subtracting forces and flow fields generated in the first stroke, when the wake is just developing, from those produced during the fourth stroke, when the pattern of both the forces and wake dynamics has reached a limit cycle. This technique identified two effects of the wake on force production by the wing: an early augmentation followed by a small attenuation. The later decrease in force is consistent with the influence of a decreased aerodynamic angle of attack on translational forces caused by downwash within the wake and is well explained by a quasi-steady model. The early effect of the wake is not well approximated by a quasi-steady model, even when the magnitude and orientation of the instantaneous velocity field are taken into account. Thus, the wake capture force represents a truly unsteady phenomenon dependent on temporal changes in the distribution and magnitude of vorticity during stroke reversal.

Electrical activity in cerebellar cultures determines Purkinje cell dendritic growth patterns

In primary dissociated cultures of mouse cerebellum a number of Purkinje cell-specific marker proteins and characteristic ionic currents appear at the appropriate developmental time. During the first week after plating, Purkinje cell dendrites elongate, but as electrical activity emerges the dendrites stop growing and branch. If endogenous electrical activity is inhibited by chronic tetrodotoxin or high magnesium treatment, dendrites continue to elongate, as if they were still immature. At the time that branching begins, intracellular calcium levels become sensitive to tetrodotoxin, suggesting that this cation may be involved in dendrite growth. Even apparently mature Purkinje cells alter their dendritic growth in response to changes in activity, suggesting long-term plasticity.

Active flight increases the gain of visual motion processing in Drosophila

We developed a technique for performing whole-cell patch-clamp recordings from genetically identified neurons in behaving Drosophila. We focused on the properties of visual interneurons during tethered flight, but this technique generalizes to different cell types and behaviors. We found that the peak-to-peak responses of a class of visual motion–processing interneurons, the vertical-system visual neurons (VS cells), doubled when flies were flying compared with when they were at rest. Thus, the gain of the VS cells is not fixed, but is instead behaviorally flexible and changes with locomotor state. Using voltage clamp, we found that the passive membrane resistance of VS cells was reduced during flight, suggesting that the elevated gain was a result of increased synaptic drive from upstream motion-sensitive inputs. The ability to perform patch-clamp recordings in behaving Drosophila promises to help unify the understanding of behavior at the gene, cell and circuit levels.

Wing transmission for a micromechanical flying insect

Flapping wings provide unmatched manoeuvrability for flying microrobots. Recent advances in modelling insect aerodynamics show that adequate wing rotation at the end of the stroke is essential for generating adequate flight forces. We developed a thorax structure using four bar frames combined with an extensible fan-fold wing to provide adequate wing stroke and rotation. Flow measurements on a scale model of the beating wing show promising aerodynamics. Calculations using a simple resonant mechanical circuit model show that piezoelectric actuators can generate sufficient power, force and stroke to drive the wings at 150 Hz.

Serotonin integrates the feeding behavior of the medicinal leech

Summary1.We have investigated functional roles of serotonin (5-hydroxytryptamine, 5-HT) in the feeding behavior of the medicinal leech,Hirudo medicinalis. Bathing intact leeches in serotonin (3×10−5 mol/l, 20 min) significantly alters three components of their feeding behavior. Leeches exposed to serotonin initiate swimming toward a vibrating point with a shorter latency, bite a warm surface more frequently, and ingest more blood than control animals.2.Thermal stimulation of the prostomium evokes pharyngeal peristalsis, jaw movements and an obvious salivation by semidissected preparations. The application of serotonin (10−8 to 10−6 mol/l) directly activates and increases the rates of pharyngeal peristalsis and salivary secretion in these preparations. Serotonin also produces jaw movements and a constriction of the crop when centrally applied.3.Vibrational stimuli, which evoke the swimming phase of feeding behavior, synaptically excite Retzius cells within segmental ganglia. Thermal stimuli, which evoke the biting phase, synaptically excite 5-HT-containing neurons within the subesophageal ganglia. Intracellular stimulation of identified 5-HT-containing neurons produces pharyngeal peristalsis reliably and enhances salivation.4.Pharmacological lesions of particular 5-HT neurons abolishes feeding behavior. However, a brief exposure to exogenous 5-HT restores biting behavior. Well-fed leeches do not normally bite, but a brief bath in 5-HT will evoke biting behavior by them as well.5.Serotonin is localized to a limited population of identifiable neurons in the C.N.S., and plays a pivotal and mandatory role in organizing the feeding behavior of the medicinal leech.