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            Free, publicly-accessible full text available July 1, 2026
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            Free, publicly-accessible full text available January 2, 2026
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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 Shrimps locomote through water using five pairs of appendages known as pleopods, which beat in a coordinated metachronal motion. Each pleopods consists of two membranous rami, a medial endopod and lateral exopod whose edges are lined with fine hair-like setae. Because of their close spacing and density, the setae act as an impermeable membrane. During swimming, each pleopod executes a power stroke, propelling water backward, followed by a recovery stroke to reset its position. During the power stroke, the exopods, endopods, and setae spread out, forming a propulsor with a larger area. In contrast, the rami close and overlap during the recovery stroke, reducing the effective area. In this study, we simulate natural shrimp swimming based on high-speed recordings under Reynolds number (Re) of 1980, using an in-house computational fluid dynamics (CFD) solver. We compared a model based on a natural swimming shrimp with a model with pleopod area fixed at the maximum area. Our results reveal that the model incorporating spread-out motion achieves a notable reduction of 49.84% in cycle-averaged hydrodynamic power while sacrificing only 23% of cycle-averaged thrust when compared to the fixed-pleopod area model. Furthermore, the effect of spread-out motion decreases the cost of transportation by 41.72% through reducing body drag by 12%. Additionally, our analysis observed the presence of a high-speed zone behind the second pleopod during stroke motion, particularly near the tangent plane of the lowest tip trajectory, and a low-speed zone in front of that pleopods.more » « lessFree, publicly-accessible full text available November 17, 2025
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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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            Abstract Ctenophores swim using flexible rows of appendages called ctenes that form the metachronal paddling. To generate propulsion, each appendage operates a power stroke that strokes backward, followed by a recovery stroke that allows the appendage to readjust its position. Notably, strokes of most metachronal swimmers are asymmetric, with faster power strokes while slower recovery strokes. Previously, the material properties are assumed as isotropic. So, the faster power stoke will lead to more pronounce deformation and the slower recovery stroke will lead to less deformation. However, this contradicts with the observations that power-stroking ctenes have the least deformation and recover deforms more, indicating an anisotropic material behavior. Such anisotropic material is hard to be manufactured, but the anisotropic behavior may be achieved by making the initial structural shape curved. The pre-curved ctene, that bending towards downstream, will be straighten in power stoke while easy to bend during recovery stroke. Our study aims to demonstrate the feasibility of using pre-curved shapes to achieve anisotropic material properties during metachronal swimming. Treating it as fluid-structure interaction (FSI) problem, we integrate our in-house computational fluid dynamics (CFD) solver with a finite element method (FEM) solver, utilizing strong coupling methods for convergence. By comparing the performance of pre-curved ctenes with straight ones, which represent isotropic material properties, we found that the curved ctenes exhibited 26.05% to 65.69% higher cycle-averaged thrust compared to the straight one as stiffness is lower. However, as stiffness increased, the pre-curved ctenes produced 3.92% to 30.58% less thrust than the straight ones. Similar trends were observed in propulsive efficiency, with the pre-curved ctenes demonstrating 46.97% better efficiency at the lowest stiffness but dropping to 34.02% less efficient as stiffness rise. Thus, while the pre-curved initial shape led to better performance at lower stiffness, exceeding a certain stiffness threshold resulted in worse performance compared to straight ctenes. The thrust enhancement from pre-curve shape is due to the drag reduction during recovery stroke, where the curved shape mitigate part of force to point more downward.more » « lessFree, publicly-accessible full text available November 17, 2025
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            Abstract Ctenophores employ flexible rows of appendages called ctenes that form the metachronal beating pattern. A complete cycle of such paddling consists of a power stroke that strokes backward to produce propulsion and a recovery stroke that allows the appendage to recover its initial position. Effective locomotion in these creatures relies on maximizing propulsion during the power stroke while minimizing drag in the recovery stroke. Unlike rigid oars, the ctenes are flexible during both the power stroke and the recovery stroke, and notably, their strokes are asymmetric, with faster movement during the power stroke. As previous research assumed uniform material properties. This assumption will eventually make the ctene deform more intensively in the power stroke than the recovery stroke due to the asymmetrical hydrodynamic forces. However, observations contradict these assumptions. One explanation posits that ctenes stiffen during the power stroke, enhancing their propulsive force, and become more flexible in the recovery stroke, reducing drag by minimizing the water-countering area. This study focusses on the influence of asymmetric stiffness on their propulsion mechanism. Inspired by nature, we conducted three-dimensional fluid-structure interaction (FSI) using an in-house immersed-boundary-method-based flow solver integrated with a nonlinear finite-element solid-mechanics solver. This integrated solver uses a two-way coupling that ensures a higher accuracy regarding the complexity due to the involvement of the multiple ctenes in a ctene row. The preliminary results show that the anisotropic stiffness of the ctene have better accuracy of deformation as compared to the deformation recorded by the high-speed camera. The asymmetric properties of the ctene material allow both the spatial and temporal asymmetry of the ctene beating pattern. Our investigation suggests that while symmetrical beating can only generate negative net thrust, a slightly asymmetrical beating can make the thrust positive. We find that power stroke period that cost 30% whole period can generates the highest thrust. As multiple ctenes involves, the interaction among ctenes can amplified the effects of the asymmetrical beating, so that the thrust generation is enhanced by 9 to 13 times because of it.more » « less
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            Abstract Metachronous rowing is a swimming mechanism widely adopted by small marine invertebrate like comb jellies, in which rows of appendages perform propulsive strokes sequentially in a coordinated manner with a fixed phase difference. To simulate metachronous rowing at intermediate Reynolds number, in this paper, a row of flexible cilia models was placed inside the flow field, with their roots stroke at a sinusoidal function of time and a fixed phase difference. A fully coupled two-way numerical solver was developed, which solves the Navier-Stokes equations for the fluid field coupled with the differential equation for the flexible cilia model. This numerical solver is applied to investigate how the row of cilia models are deformed by the hydrodynamic forces (pressure and shear) and momentum and thus impact hydrodynamic performance. Results show that the passive deformation of cilia potentially improve the hydrodynamic performance compared to the rigid cilia. With the metachronous rowing mechanism, the cilia generate the thrust to move forward. The approach used in this study presents a general way to explore the fluid dynamics of complex fluid-structure interaction problems.more » « less
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