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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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Flexural rigidity of hawkmoth antennae depends on the bending direction
To probe its environment, the flying insect controllably flexes, twists, and maneuvers its antennae by coupling mechanical deformations with the sensory output. We question how the materials properties of insect antennae could influence their performance. A comparative study was conducted on four hawkmoth species: Manduca sexta, Ceratomia catalpae, Manduca quinquemaculata, and Xylophanes tersa. The morphology of the antennae of three hawkmoths that hover while feeding and one putatively non-nectar-feeding hawkmoth (Ceratomia catalpa) do not fundamentally differ, and all the antennae are comb-like (i.e., pectinate), markedly in males but weakly in females. Applying different weights to the free end of extracted cantilevered antennae, we discovered anisotropy in flexural rigidity when the antenna is forced to bend dorsally versus ventrally. The flexural rigidity of male antennae was less than that of females. Compared with the hawkmoths that hover while feeding, Ceratomia catalpae has almost two orders of magnitude lower flexural rigidity. Tensile tests showed that the stiffness of male and female antennae is almost the same. Therefore, the differences in flexural rigidity are explained by the distinct shapes of the antennal pectination. Like bristles in a comb, the pectinations provide extra rigidity to the antenna. We discuss the biological implications of these discoveries in relation to the flight habits of hawkmoths. Flexural anisotropy of antennae is expected in other groups of insects, but the targeted outcome may differ. Our work offers promising new applications of shaped fibers as mechanical sensors.
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- Award ID(s):
- 2042937
- PAR ID:
- 10595942
- Editor(s):
- Wagner, William R
- Publisher / Repository:
- Elsevier
- Date Published:
- Journal Name:
- Acta Biomaterialia
- Edition / Version:
- 1
- Volume:
- 184
- Issue:
- C
- ISSN:
- 1742-7061
- Page Range / eLocation ID:
- 273 to 285
- Subject(s) / Keyword(s):
- Insect antennae, Bending, Hawkmoths, Flexural rigidity, Lepidoptera
- Format(s):
- Medium: X Size: 2MB Other: pdf
- Size(s):
- 2MB
- Sponsoring Org:
- National Science Foundation
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- Free Publicly Accessible Full Text
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Accepted Manuscript1.0
- Journal Article:
- https://doi.org/10.1016/j.actbio.2024.06.036
