Coriolis and centrifugal forces drive haltere deformations and influence spike timing.

The halteres of flies are mechanosensory organs that serve a crucial role in the control of agile flight, providing sensory input for rapid course corrections to perturbations. Derived from hind wings, halteres are actively flapped and are thus subject to a variety of inertial forces as the fly undergoes complex flight trajectories. Previous analyses of halteres modelled them as a point mass, showing that Coriolis forces lead to subtle deflections orthogonal to the plane of flapping. By design, these models could not consider the effects of force gradients associated with a mass distribution, nor could they reveal three-dimensional spatio-temporal patterns of strain that result from those forces. In addition, diversity in the geometry of halteres, such as shape and asymmetries, could not be simply modelled with a point mass on a massless rod. To study the effects of mass distributions and asymmetries, we examine the haltere subject to both flapping and body rotations using three-dimensional finite-element simulations. We focus on a set of simplified geometries, in which we vary the stalk and bulb shape. We find that haltere mass distribution gives rise to two unreported deformation modes: (i) halteres twist with a magnitude that strongly depends on stalk and bulb geometry and (ii) halteres with an asymmetric mass distribution experience out-of-plane bending due to centrifugal forces, independent of body rotation. Since local strains at the base of the haltere drive deformations of mechanosensory neurons, we combined measured neural encoding mechanisms with our structural analyses to predict the spatial and temporal patterns of neural activity. This activity depends on both the flapping and rotation dynamics, and we show how the timing of neural activity is a viable mechanism for rotation-rate encoding. Our results provide new insights in haltere dynamics and show the viability for timing-based encoding of fly body rotations by halteres.

High-resolution, high-throughput imaging with a multibeam scanning electron microscope.

Electron-electron interactions and detector bandwidth limit the maximal imaging speed of single-beam scanning electron microscopes. We use multiple electron beams in a single column and detect secondary electrons in parallel to increase the imaging speed by close to two orders of magnitude and demonstrate imaging for a variety of samples ranging from biological brain tissue to semiconductor wafers.

A new twist on gyroscopic sensing: body rotations lead to torsion in flapping, flexing insect wings.

Insects perform fast rotational manoeuvres during flight. While two insect orders use flapping halteres (specialized organs evolved from wings) to detect body dynamics, it is unknown how other insects detect rotational motions. Like halteres, insect wings experience gyroscopic forces when they are flapped and rotated and recent evidence suggests that wings might indeed mediate reflexes to body rotations. But, can gyroscopic forces be detected using only changes in the structural dynamics of a flapping, flexing insect wing? We built computational and robotic models to rotate a flapping wing about an axis orthogonal to flapping. We recorded high-speed video of the model wing, which had a flexural stiffness similar to the wing of the Manduca sexta hawkmoth, while flapping it at the wingbeat frequency of Manduca (25 Hz). We compared the three-dimensional structural dynamics of the wing with and without a 3 Hz, 10° rotation about the yaw axis. Our computational model revealed that body rotation induces a new dynamic mode: torsion. We verified our result by measuring wing tip displacement, shear strain and normal strain of the robotic wing. The strains we observed could stimulate an insect's mechanoreceptors and trigger reflexive responses to body rotations.

Fluid-structure interaction in compliant insect wings.

Insect wings deform significantly during flight. As a result, wings act as aeroelastic structures wherein both the driving motion of the structure and the aerodynamic loading of the surrounding fluid potentially interact to modify wing shape. We explore two key issues associated with the design of compliant wings: over a range of driving frequencies and phases of pitch-heave actuation, how does wing stiffness influence (1) the lift and thrust generated and (2) the relative importance of fluid loading on the shape of the wing? In order to examine a wide range of parameters relevant to insect flight, we develop a computationally efficient, two-dimensional model that couples point vortex methods for fluid force computations with structural finite element methods to model the fluid-structure interaction of a wing in air. We vary the actuation frequency, phase of actuation, and flexural stiffness over a range that encompasses values measured for a number of insect taxa (10-90 Hz; 0-π rad; 10(-7)-10(-5) N m(2)). We show that the coefficients of lift and thrust are maximized at the first and second structural resonant frequencies of the system. We also show that even in regions of structural resonance, fluid loading never contributes more than 20% to the development of flight forces.