Biomedical subjects

S A Combes

Publications and source records attributed to S A Combes.

4 recordsLinked to original sources

PubMed2003

Into thin air: Contributions of aerodynamic and inertial-elastic forces to wing bending in the hawkmoth Manduca sexta.

During flapping flight, insect wings must withstand not only fluid-dynamic forces, but also inertial-elastic forces generated by the rapid acceleration and deceleration of their own mass. Estimates of overall aerodynamic and inertial forces vary widely, and the relative importance of these forces in determining passive wing deformations remains unknown. If aeroelastic interactions between a wing and the fluid-dynamic forces it generates are minor compared to the effects of wing inertia, models of insect flight that account for passive wing flexibility would be far simpler to develop. We used an experimental approach to examine the contributions of aerodynamic and inertial-elastic forces to wing bending in the hawkmoth Manduca sexta. We attached fresh Manduca wings to a motor and flapped them at a realistic wing-beat frequency and stroke amplitude. We compared wing bending in normal air versus helium (approx. 15% air density), in which the contribution of fluid-dynamic forces to wing deformations is significantly reduced. This 85% reduction in air density produced only slight changes in the pattern of Manduca wing deformations, suggesting that fluid-dynamic forces have a minimal effect on wing bending. We used a simplified finite element model of a wing to show that the differences observed between wings flapped in air versus helium are most likely due to fluid damping, rather than to aerodynamic forces. This suggests that damped finite element models of insect wings (with no fluid-dynamic forces included) may be able to predict overall patterns of wing deformation prior to calculations of aerodynamic force production, facilitating integrative models of insect flight.

Animals↗

PubMed2003

Flexural stiffness in insect wings. I. Scaling and the influence of wing venation.

During flight, many insect wings undergo dramatic deformations that are controlled largely by the architecture of the wing. The pattern of supporting veins in wings varies widely among insect orders and families, but the functional significance of phylogenetic trends in wing venation remains unknown, and measurements of the mechanical properties of wings are rare. In this study, we address the relationship between venation pattern and wing flexibility by measuring the flexural stiffness of wings (in both the spanwise and chordwise directions) and quantifying wing venation in 16 insect species from six orders. These measurements show that spanwise flexural stiffness scales strongly with the cube of wing span, whereas chordwise flexural stiffness scales with the square of chord length. Wing size accounts for over 95% of the variability in measured flexural stiffness; the residuals of this relationship are small and uncorrelated with standardized independent contrasts of wing venation characters. In all species tested, spanwise flexural stiffness is 1-2 orders of magnitude larger than chordwise flexural stiffness. A finite element model of an insect wing demonstrates that leading edge veins are crucial in generating this spanwise-chordwise anisotropy.

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PubMed2003

Flexural stiffness in insect wings. II. Spatial distribution and dynamic wing bending.

The dynamic, three-dimensional shape of flapping insect wings may influence many aspects of flight performance. Insect wing deformations during flight are largely passive, and are controlled primarily by the architecture and material properties of the wing. Although many details of wing structure are well understood, the distribution of flexural stiffness in insect wings and its effects on wing bending are unknown. In this study, we developed a method of estimating spatial variation in flexural stiffness in both the spanwise and chordwise direction of insect wings. We measured displacement along the wing in response to a point force, and modeled flexural stiffness variation as a simple mathematical function capable of approximating this measured displacement. We used this method to estimate flexural stiffness variation in the hawkmoth Manduca sexta, and the dragonfly Aeshna multicolor. In both species, flexural stiffness declines sharply from the wing base to the tip, and from the leading edge to the trailing edge; this variation can be approximated by an exponential decline. The wings of M. sexta also display dorsal/ventral asymmetry in flexural stiffness and significant differences between males and females. Finite element models based on M. sexta forewings demonstrate that the measured spatial variation in flexural stiffness preserves rigidity in proximal regions of the wing, while transferring bending to the edges, where aerodynamic force production is most sensitive to subtle changes in shape.

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PubMed2001

Shape, flapping and flexion: wing and fin design for forward flight.

Both kinematics and morphology are critical determinants of performance in flapping flight. However, the functional consequences of changes in these traits are not yet well understood. Traditional aerodynamic studies of planform wing shape have suggested that high-aspect-ratio wings generate more force per area and perform more efficiently than low-aspect-ratio wings, but these analyses may neglect critical components of flapping flight such as unsteady fluid dynamics and wing or fin flexion. In this paper, we use an unsteady potential flow analysis that incorporates wing flexion to test predictions of optimal wing shape under varying degrees of unsteady motion and wing flexion. We focus on forward flapping flight and examine the effects of wing/fin morphology and movements on thrust generation and efficiency. We test the model by comparing our predictions with kinematic data derived from the aquatic flight of the ratfish Hydrolagus colliei. Our analyses show that aspect ratio and the proportion of area in the outer one-fifth of the wing can characterize wing shape in terms of aero- or hydrodynamic performance. By comparing the performance of wings that vary in these two parameters, we find that traditional predictions of optimal wing shape are valid only under limited circumstances (when flapping frequency is low, wings are stiff or wings are tapered at the tips). This indicates a complex relationship between locomotor traits and performance and helps explain the diversity of wing kinematics and morphologies observed in nature.

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