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Bioinspir. Biomim. 20 (2025) 016024 https://doi.org/10.1088/1748-3190/ad9535
RECEIVED
17 July 2024
REVISED
2 November 2024
ACC EPT ED FOR PUB LICATI ON
20 November 2024
PUBLISHED
23 December 2024
PAPER
Stability and agility trade-offs in spring-wing systems
James Lynch1, Ethan S Wold2, Jeff Gau3, Simon Sponberg2,4and Nick Gravish1,∗
1Department of Mechanical & Aerospace Engineering, University of California, San Diego, CA, United States of America
2School of Biological Sciences, Georgia Institute of Technology, Atlanta, GA, United States of America
3Interdisciplinary Bioengineering Graduate Program and George W. Woodruff School of Mechanical Engineering, Georgia Institute of
Technology, Atlanta, GA, United States of America
4School of Physics, Georgia Institute of Technology, Atlanta, GA, United States of America
∗Author to whom any correspondence should be addressed.
E-mail: ngravish@ucsd.edu
Keywords: insect flight, resonance, dynamic scaling, elasticity, robophysics
Abstract
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 with N) 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.
1. Introduction
Flapping flight is an extremely power-intensive mode
of locomotion, requiring both high frequency wing-
beats and large forces to produce lift and perform
agile maneuvers. Flying insects achieve efficient flight
through a combination of specialized flight muscles
[1] and elastic energy storage in the thorax [2–4]. The
insect flight system can thus be described as muscle
actuation of an elastic structure which oscillates wings
to generate aerodynamic forces. We call this combina-
tion of elastic, inertial, and aerodynamic mechanisms
a ‘spring-wing’ system [5]. While significant research
focus has been devoted to the aerodynamic force gen-
eration of flapping wings (see review in [6]), relat-
ively fewer studies have focused on understanding the
implications of elastic energy storage and return for
flight dynamics and control [3,4,7–10].
In the classic spring-mass-damper model, there
exists a particular actuation frequency which results
in the largest amplitude oscillation of a mass, the so-
called resonance frequency. In the performance con-
siderations for a ‘spring-wing’ system, there exist sev-
eral different resonant wingbeat frequencies at which
different forms of optimality (maximum amplitude,
lift, or efficiency, for example) are achieved [11].
Operating at a resonant frequency that maximizes
lift can enable significant performance advantage,
allowing insects to use smaller muscle force/power to
generate lift for flight. Indeed, roboticists designing
insect-scale flapping robots have found that incorpor-
ating elasticity and operating near resonance enables
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