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Flying insects are thought to achieve energy-efficient flapping flight by storing and releasing elastic energy in their muscles, tendons, and thorax. However, ‘spring-wing’ flight systems consisting of elastic elements coupled to nonlinear, unsteady aerodynamic forces present possible challenges to generating stable and responsive wing motion. The energetic efficiency from resonance in insect flight is tied to the Weis-Fogh number (N), which is the ratio of peak inertial force to aerodynamic force. In this paper, we present experiments and modeling to study how resonance efficiency (which increases withN) influences the control responsiveness and perturbation resistance of flapping wingbeats. In our first experiments, we provide a step change in the input forcing amplitude to a series-elastic spring-wing system and observe the response time of the wing amplitude increase. In our second experiments we provide an external fluid flow directed at the flapping wing and study the perturbed steady-state wing motion. We evaluate both experiments across Weis-Fogh numbers from 1 < N < 10. The results indicate that spring-wing systems designed for maximum energetic efficiency also experience trade-offs in agility and stability as the Weis-Fogh number increases. Our results demonstrate that energetic efficiency and wing maneuverability are in conflict in resonant spring-wing systems, suggesting that mechanical resonance presents tradeoffs in insect flight control and stability.
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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.
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- Award ID(s):
- 2100858
- PAR ID:
- 10342645
- Date Published:
- Journal Name:
- Biology Letters
- Volume:
- 18
- Issue:
- 5
- ISSN:
- 1744-957X
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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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
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Abstract Powering small-scale flapping flight is challenging, yet insects sustain exceptionally fast wingbeats with ease. Since insects act as tiny biomechanical resonators, tuning their wingbeat frequency to the resonant frequency of their springy thorax and wings could make them more efficient fliers. But operating at resonance poses control problems and potentially constrains wingbeat frequencies within and across species. Resonance may be particularly limiting for the many orders of insects that power flight with specialized muscles that activate in response to mechanical stretch. Here, we test whether insects operate at their resonant frequency. First, we extensively characterize bumblebees and find that they surprisingly flap well above their resonant frequency via interactions between stretch-activation and mechanical resonance. Modeling and robophysical experiments then show that resonance is actually a lower bound for rapid wingbeats in most insects because muscles only pull, not push. Supra-resonance emerges as a general principle of high-frequency flight across five orders of insects from moths to flies.more » « less
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In the last decade, roboticists have had significant success building centimeter-scale flapping wing micro aerial vehicles (FWMAVs) inspired by the flight of insects. Evidence suggests that insects store and release energy in the thoracic exoskeleton to improve energy efficiency by flapping at resonance. Insect-inspired micro flying robots have also leveraged resonance to improve efficiency, but they have discovered that operating at the resonant frequency leads to issues with flight control. This research seeks to investigate the roles that elasticity, aerodynamics, and muscle dynamics play in the emergent dynamics of flapping flight by studying elastic flapping spring-wing systems using dynamically-scaled robophysical models of springwings. Studying the dynamics of a robot with comparable features enables the validation of models from biology that are otherwise difficult to test in living insects, the generation of new hypotheses, and the development of novel FWMAV designs. In Chapter 1, the spring-wing system is characterized via a nonlinear spring-mass-damper model. A robophysical model validates that such systems gain energetic benefits from operating at resonance, but reveals that the benefit scales with an underappreciated dimensionless ratio of inertial to aerodynamic forces, the Weis-Fogh number. We show through dimensional analysis that any real system, living or robotic, must balance the mechanical advantage gained from operating at resonance with diminishing returns in efficiency. Chapter 2 further explores the impact of the Weis-Fogh number on flapping dynamics, showing that responsiveness to control inputs is reduced and resistance to environmental perturbations is increased as the dimensionless ratio increases. Together with calculations of Weis-Fogh number in insects, these studies illustrate tradeoffs that drive evolution of resonant flight in nature and guide development of future FWMAVs with elastic energy exchange. In the second half of the thesis, muscle dynamics are introduced in the form of a simplified model of self-excited asynchronous insect muscle. In Chapter 3, a form of velocity feedback, adapted from experiments on insect flight muscle, is developed and integrated with the springwing model, producing a system that generates steady flapping via limit-cycle oscillations despite the absence of periodic control inputs. The model is explored analytically, in simulation, and via implementation on the robotic spring-wing. Novel dynamic characteristics that enable adaptation to damage and passive response to wing collisions are described. Chapter 4 leverages the asynchronous feedback model as part of an interdisciplinary study of the evolution of asynchronous muscle. Phylogenetic analysis, direct measurement of insect muscle dynamics, and experiments on the robophysical system show that evolutionary transitions between periodicallyforced and self-excited insect muscle were likely made possible by a ”bridge” in the dynamic parameter space that could be traversed under specific conditions. The asynchronous spring-wing model provides new insight into the flight and evolution of some of the most agile insects in nature, and presents a novel adaptive control scheme for future FWMAVs.more » « less
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In most instances, flapping wing robots have emulated the “synchronous” actuation of insects in which the wingbeat timing is generated from a time-dependent, rhythmic signal. The internal dynamics of asynchronous insect flight muscle enable high-frequency, adaptive wingbeats with minimal direct neural control. In this paper, we investigate how the delayed stretch-activation (dSA) response of asynchronous insect flight muscle can be transformed into a feedback control law for flapping wing robots that results in stable limit cycle wingbeats. We first demonstrate - in theory and simulation - the mechanism by which asynchronous wingbeats self-excite. Then, we implement the feedback law on a dynamically-scaled robophysical model as well as on an insect-scale robotic flapping wing. Experiments on large- and small-scale robots demonstrate good agreement with the theory results and highlight how dSA parameters govern wingbeat amplitude and frequency. Lastly, we demonstrate that asynchronous actuation has several advantages over synchronous actuation schemes, including the ability to rapidly adapt or halt wingbeats in response to external loads or collisions through low-level feedback control.more » « less
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Accepted Manuscript1.0
- Journal Article:
- https://doi.org/10.1098/rsbl.2022.0063
