skip to main content

Explore Research Products in the NSF-PAR

Title: The hawkmoth wingbeat is not at resonance
Flying insects have elastic materials within their exoskeletons that could reduce the energetic cost of flight if their wingbeat frequency is matched to a mechanical resonance frequency. Flapping at resonance may be essential across flying insects because of the power demands of small-scale flapping flight. However, building up large-amplitude resonant wingbeats over many wingstrokes may be detrimental for control if the total mechanical energy in the spring-wing system exceeds the per-cycle work capacity of the flight musculature. While the mechanics of the insect flight apparatus can behave as a resonant system, the question of whether insects flap their wings at their resonant frequency remains unanswered. Using previous measurements of body stiffness in the hawkmoth, Manduca sexta , we develop a mechanical model of spring-wing resonance with aerodynamic damping and characterize the hawkmoth's resonant frequency. We find that the hawkmoth's wingbeat frequency is approximately 80% above resonance and remains so when accounting for uncertainty in model parameters. In this regime, hawkmoths may still benefit from elastic energy exchange while enabling control of aerodynamic forces via frequency modulation. We conclude that, while insects use resonant mechanics, tuning wingbeats to a simple resonance peak is not a necessary feature for all centimetre-scale flapping flyers.  more » « less
Award ID(s):
2100858
NSF-PAR ID:
10342645
Author(s) / Creator(s):
; ; ; ;
Date Published:
Journal Name:
Biology Letters
Volume:
18
Issue:
5
ISSN:
1744-957X
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. ; ; ; ; ; ( , Proceedings of the Royal Society B: Biological Sciences)
    null (Ed.)
    Centimetre-scale fliers must contend with the high power requirements of flapping flight. Insects have elastic elements in their thoraxes which may reduce the inertial costs of their flapping wings. Matching wingbeat frequency to a mechanical resonance can be energetically favourable, but also poses control challenges. Many insects use frequency modulation on long timescales, but wingstroke-to-wingstroke modulation of wingbeat frequencies in a resonant spring-wing system is potentially costly because muscles must work against the elastic flight system. Nonetheless, rapid frequency and amplitude modulation may be a useful control modality. The hawkmoth Manduca sexta has an elastic thorax capable of storing and returning significant energy. However, its nervous system also has the potential to modulate the driving frequency of flapping because its flight muscles are synchronous. We tested whether hovering hawkmoths rapidly alter frequency during perturbations with vortex rings. We observed both frequency modulation (32% around mean) and amplitude modulation (37%) occurring over several wingstrokes. Instantaneous phase analysis of wing kinematics revealed that more than 85% of perturbation responses required active changes in neurogenic driving frequency. Unlike their robotic counterparts that abdicate frequency modulation for energy efficiency, synchronous insects use wingstroke-to-wingstroke frequency modulation despite the power demands required for deviating from resonance. 
    more » « less
  2. ; ; ; ( , Journal of The Royal Society Interface)
    null (Ed.)
    Flapping-wing insects, birds and robots are thought to offset the high power cost of oscillatory wing motion by using elastic elements for energy storage and return. Insects possess highly resilient elastic regions in their flight anatomy that may enable high dynamic efficiency. However, recent experiments highlight losses due to damping in the insect thorax that could reduce the benefit of those elastic elements. We performed experiments on, and simulations of, a dynamically scaled robophysical flapping model with an elastic element and biologically relevant structural damping to elucidate the roles of body mechanics, aerodynamics and actuation in spring-wing energetics. We measured oscillatory flapping-wing dynamics and energetics subject to a range of actuation parameters, system inertia and spring elasticity. To generalize these results, we derive the non-dimensional spring-wing equation of motion and present variables that describe the resonance properties of flapping systems: N , a measure of the relative influence of inertia and aerodynamics, and K ^ , the reduced stiffness. We show that internal damping scales with N , revealing that dynamic efficiency monotonically decreases with increasing N . Based on these results, we introduce a general framework for understanding the roles of internal damping, aerodynamic and inertial forces, and elastic structures within all spring-wing systems. 
    more » « less
  3. ; ; ; ( , Journal of The Royal Society Interface)
    Muscles act through elastic and dissipative elements to mediate movement, which can introduce dissipation and filtering which are important for energetics and control. The high power requirements of flapping flight can be reduced by an insect's exoskeleton, which acts as a spring with frequency-independent material properties under purely sinusoidal deformation. However, this purely sinusoidal dynamic regime does not encompass the asymmetric wing strokes of many insects or non-periodic deformations induced by external perturbations. As such, it remains unknown whether a frequency-independent model applies broadly and what implications it has for control. We used a vibration testing system to measure the mechanical properties of isolated Manduca sexta thoraces under symmetric, asymmetric and band-limited white noise deformations. The asymmetric and white noise conditions represent two types of generalized, multi-frequency deformations that may be encountered during steady-state and perturbed flight. Power savings and dissipation were indistinguishable between symmetric and asymmetric conditions, demonstrating that no additional energy is required to deform the thorax non-sinusoidally. Under white noise conditions, stiffness and damping were invariant with frequency, suggesting that the thorax has no frequency-dependent filtering properties. A simple flat frequency response function fits our measured frequency response. This work demonstrates the potential of materials with frequency-independent damping to simplify motor control by eliminating any velocity-dependent filtering that viscoelastic elements usually impose between muscle and wing. 
    more » « less
  4. ; ; ( , Biomimetics)
    Small-scale flapping-wing micro air vehicles (FWMAVs) are an emerging robotic technology with many applications in areas including infrastructure monitoring and remote sensing. However, challenges such as inefficient energetics and decreased payload capacity preclude the useful implementation of FWMAVs. Insects serve as inspiration to FWMAV design owing to their energy efficiency, maneuverability, and capacity to hover. Still, the biomechanics of insects remain challenging to model, thereby limiting the translational design insights we can gather from their flight. In particular, it is not well-understood how wing flexibility impacts the energy requirements of flapping flight. In this work, we developed a simple model of an insect drive train consisting of a compliant thorax coupled to a flexible wing flapping with single-degree-of-freedom rotation in a fluid environment. We applied this model to quantify the energy required to actuate a flapping wing system with parameters based off a hawkmoth Manduca sexta. Despite its simplifications, the model predicts thorax displacement, wingtip deflection and peak aerodynamic force in proximity to what has been measured experimentally in flying moths. We found a flapping system with flexible wings requires 20% less energy than a flapping system with rigid wings while maintaining similar aerodynamic performance. Passive wing deformation increases the effective angle of rotation of the flexible wing, thereby reducing the maximum rotation angle at the base of the wing. We investigated the sensitivity of these results to parameter deviations and found that the energetic savings conferred by the flexible wing are robust over a wide range of parameters. 
    more » « less
  5. ; ( , Physics of Fluids)

    Odor-guided navigation is an indispensable aspect of flying insects' behavior, facilitating crucial activities such as foraging and mating. The interaction between aerodynamics and olfaction plays a pivotal role in the odor-guided flight behaviors of insects, yet the interplay of these two functions remains incompletely understood. In this study, we developed a fully coupled three-way numerical solver, which solves the three-dimensional Navier–Stokes equations coupled with equations of motion for the passive flapping wings, and the odorant advection–diffusion equation. This numerical solver is applied to investigate the unsteady flow field and the odorant transport phenomena of a fruit fly model in odor-guided upwind surge flight over a broad spectrum of reduced frequencies (0.325–1.3) and Reynolds numbers (90–360). Our results uncover a complex dependency between flight velocity and odor plume perception, modulated by the reduced frequency of flapping flight. At low reduced frequencies, the flapping wings disrupt the odor plume, creating a saddle point of air flow near the insect's thorax. Conversely, at high reduced frequencies, the wing-induced flow generates a stagnation point, in addition to the saddle point, that alters the aerodynamic environment around the insect's antennae, thereby reducing odor sensitivity but increasing the sampling range. Moreover, an increase in Reynolds number was found to significantly enhance odor sensitivity due to the synergistic effects of greater odor diffusivity and stronger wing-induced flow. These insights hold considerable implications for the design of bio-inspired, odor-guided micro air vehicles in applications like surveillance and detection.

     
    more » « less
  • Citation Formats
  • MLA
  • APA
  • Chicago
  • BibTeX
  • Save / Share this Record
  • Send to Email