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            Abstract One of the most ancient and evolutionarily conserved behaviors in the animal kingdom involves utilizing wind-borne odor plumes to track essential elements such as food, mates, and predators. Insects, particularly flies, demonstrate a remarkable proficiency in this behavior, efficiently processing complex odor information encompassing concentrations, direction, and speed through their olfactory system, thereby facilitating effective odor-guided navigation. Recent years have witnessed substantial research explaining the impact of wing flexibility and kinematics on the aerodynamics and flow field physics governing the flight of insects. However, the relationship between the flow field and olfactory functions remains largely unexplored, presenting an attractive frontier with numerous intriguing questions. One such question pertains to whether flies intentionally manipulate the flow field around their antennae using their wing structure and kinematics to augment their olfactory capabilities. To address this question, we first reconstructed the wing kinematics based on high-speed video recordings of wing surface deformation. Subsequently, we simulated the unsteady flow field and odorant transport during the forward flight of blue bottle flies (Calliphora vomitoria) by solving the Navier–Stokes equations and odorant advection–diffusion equations using an in-house computational fluid dynamics solver. Our simulation results demonstrated that flexible wings generated greater cycle-averaged aerodynamic forces compared to purely rigid flapping wings, underscoring the aerodynamic advantages of wing flexibility. Additionally, flexible wings produced 25% greater odor intensity, enhancing the insect’s ability to detect and interpret olfactory cues. This study not only advances our understanding of the intricate interplay between wing motion, aerodynamics, and olfactory capabilities in flying insects but also raises intriguing questions about the intentional modulation of flow fields for sensory purposes in other behaviors.more » « less
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            Abstract As insects fly, their wings generate complex wake structures that play a crucial role in their aerodynamic force production. This study focuses on utilizing reduced order modeling techniques to gain valuable insights into the fluid dynamic principles underlying insect flight. Specifically, we used an immersed-boundary-method-based computational fluid dynamics (CFD) solver to simulate a hovering hawkmoth’s wake, and then identified the most energetic modes of the wake using proper orthogonal decomposition (POD). Furthermore, we employed a sparse identification of nonlinear dynamics (SINDy) approach to find a simple reduced order model that relates the most energetic POD modes. Through this process, we formulated multiple different models incorporating varying numbers of POD modes. To compare the accuracy of these models, we utilized a force survey method to estimate the aerodynamic forces produced by the hawkmoth’s wings. This force survey method uses an impulse-based approach to calculate the aerodynamic lift and drag based solely on the velocity and vorticity information provided by the model. By comparing the estimated aerodynamic force with the actual force production calculated by the CFD solver, we were able to find the simplest model that still provides an accurate representation of the complex wake produced by the hovering hawkmoth wings. We also evaluated the stability and accuracy of this model as the number of flapping cycles increases with time. The reduced order modeling of insect flight has important implications for the design and control of bio-inspired micro-aerial vehicles. In addition, it holds the potential to reduce the computational cost associated with high-fidelity CFD simulations of complex flows.more » « less
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            Free, publicly-accessible full text available August 1, 2026
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            Insects rely on their olfactory systems to detect odors and locate odor sources through highly efficient flapping-wing mechanisms. While previous studies on bio-inspired unsteady flows have primarily examined the aerodynamic functions of flapping wings, they have largely overlooked the effects of wing-induced unsteady flows on airborne odor stimuli. This study aims to explore how flapping kinematics influence odorant transport. Computational fluid dynamics simulations were employed to investigate unsteady flow fields and odorant transport by solving the Navier–Stokes and odor advection–diffusion equations. Both two-dimensional (2D) and three-dimensional (3D) simulations were conducted to visualize the flow fields and odor concentration distributions generated by pitching–plunging airfoils. Our findings reveal that higher Strouhal numbers, characterized by increased flapping frequency, produce stronger flow jets that enhance odor advection and dissipation downstream, while reducing odor concentration on the airfoil surface. In 2D simulations, symmetry breaking at high Strouhal numbers causes oblique advection of vortices and odor plumes. In contrast, 3D simulations exhibit bifurcated horseshoe-like vortex rings and corresponding odor plume bifurcations. These findings highlight the intricate coupling between unsteady aerodynamics and odor transport, offering valuable insights for bio-inspired designs and advanced olfactory navigation systems.more » « lessFree, publicly-accessible full text available March 1, 2026
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            Flying insects are equipped with complex olfactory systems, which they utilize to seek food, identify mates, and evade predators. It is suspected that insects flap their wings to draw odor plumes toward their antennae, a behavior akin to mammals' sniffing, aimed at enhancing olfactory sensitivity. However, insects' wing kinematics change drastically as their flight speed increases, and it is unknown how these changes affect the insect's odorant perception. Addressing this gap in knowledge is crucial to a full understanding of the interplay between insects' aerodynamic performance and sensory perception. To this end, we simulated odor-tracking hawkmoth flight at 2 and 4 m/s using an in-house computational fluid dynamics solver. This solver incorporated both the Navier–Stokes equations that govern the flow, as well as the advection-diffusion equation that governs the odor transport process. Findings indicate that hawkmoths enhance odor intensity along their antennae using their wings, with peak odor intensity being 39% higher during 2 m/s flight compared to 4 m/s flight. This demonstrates there is a trade-off between rapid transport and olfaction, which is attributable to differences in wing kinematics between low- and high-speed flights. Despite literature suggesting hawkmoths are limited to steady forward flights at speeds below 5 m/s—about half of what is theoretically predicted based on body mass—this study reveals that slower flight speeds improve their olfactory capabilities during navigation. Our findings offer insights into the evolution of flight and sensory capabilities in hawkmoths, as well as provide inspiration for the development of bio-inspired odor-guided navigation technologies.more » « lessFree, publicly-accessible full text available February 1, 2026
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            Flying insects exhibit remarkable capabilities in coordinating their olfactory sensory system and flapping wings during odour plume-tracking flights. While observations have indicated that their flapping wing motion can ‘sniff’ up the incoming plumes for better odour sampling range, how flapping motion impacts the odour concentration field around the antennae is unknown. Here, we reconstruct the body and wing kinematics of a forwards-flying butterfly based on high-speed images. Using an in-house computational fluid dynamics solver, we simulate the unsteady flow field and odourant transport process by solving the Navier–Stokes and odourant advection-diffusion equations. Our results show that, during flapping flight, the interaction between wing leading-edge vortices and antenna vortices strengthens the circulation of antenna vortices by over two-fold compared with cases without flapping motion, leading to a significant increase in odour intensity fluctuation along the antennae. Specifically, the interaction between the wings and antennae amplifies odour intensity fluctuations on the antennae by up to 8.4 fold. This enhancement is critical in preventing odour fatigue during odour-tracking flights. Further analysis reveals that this interaction is influenced by the inter-antennal angle. Adjusting this angle allows insects to balance between resistance to odour fatigue and the breadth of odour sampling. Narrower inter-antennal angles enhance fatigue resistance, while wider angles extend the sampling range but reduce resistance. Additionally, our findings suggest that while the flexibility of the wings and the thorax's pitching motion in butterflies do influence odour fluctuation, their impact is relatively secondary to that of the wing–antenna interaction.more » « lessFree, publicly-accessible full text available November 10, 2025
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            In this paper, we examine the coupling between odor dynamics and vortex dynamics around undulating bodies, with a focus on bio-inspired propulsion mechanisms. Utilizing computational fluid dynamics simulations with an in-house immersed boundary method solver, we investigate how different waveform patterns, specifically carangiform and anguilliform, influence the dispersion of chemical cues in both water and air environments. Our findings reveal that vortex dynamics significantly impact the overall trajectory of odor spots, although the alignment between odor spots and coherent flow structures is not always precise. We also evaluate the relative contributions of diffusion and convection in odor transport, showing that convection dominates in water, driven by higher Schmidt numbers, while diffusion plays a more prominent role alongside convection in air. Additionally, the anguilliform waveform generally produces stronger and farther-reaching chemical cues compared to carangiform swimmers. The critical roles of Strouhal number and Reynolds number in determining the efficiency of odor dispersion are also explained, offering insights that could enhance the design of more efficient, adaptive, and intelligent autonomous underwater vehicles by integrating sensory and hydrodynamic principles inspired by fish locomotion.more » « lessFree, publicly-accessible full text available November 1, 2025
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