Insect Wings Research Articles

Flow sensing on dragonfly wings.

One feature of animal wings is their embedded mechanosensory system that can support flight control. Insect wings are particularly interesting as they are highly deformable yet the actuation is limited to the wing base. It is established that strain sensors on insect wings can directly mediate reflexive control; however, little is known about airflow sensing by insect wings. What information can flow sensors capture and how can flow sensing benefit flight control? Here, we use the dragonfly (Sympetrum striolatum) as a model to explore the function of wing sensory bristles in the context of flight control. Combining our detailed anatomical reconstructions of both the sensor microstructures and wing architecture, we used computational fluid dynamics simulations to ask the following questions. (1) Are there strategic locations on wings that sample flow for estimating aerodynamically relevant parameters such as the local effective angle of attack? (2) Is the sensory bristle distribution on dragonfly wings optimal for flow sensing? (3) What is the aerodynamic effect of microstructures found near the sensory bristles on dragonfly wings? We discuss the benefits of flow sensing for flexible wings and how the evolved sensor placement affects information encoding.

Quasi-steady aerodynamic modeling and dynamic stability of mosquito-inspired flapping wing pico aerial vehicle.

Recent exploration in insect-inspired robotics has generated considerable interest. Among insects navigating at low Reynolds numbers, mosquitoes exhibit distinct flight characteristics, including higher wingbeat frequencies, reduced stroke amplitudes, and slender wings. This leads to unique aerodynamic traits such as trailing edge vortices via wake capture, diminished reliance on leading vortices, and rotational drag. This paper shows the energetic analysis of a mosquito-inspired flapping-wing Pico aerial vehicle during hovering, contributing insights to its future design and fabrication. The investigation relies on kinematic and quasi-steady aerodynamic modeling of a symmetric flapping-wing model with a wingspan of approximately 26 mm, considering translational, rotational, and wake capture force components. The control strategy adapts existing bird flapping wing approaches to accommodate insect wing kinematics and aerodynamic features. Flight controller design is grounded in understanding the impact of kinematics on wing forces. Additionally, a thorough analysis of the dynamic stability of the mosquito-inspired PAV model is conducted, revealing favorable controller response and maneuverability at a small scale. The modified model, incorporating rigid body dynamics and non-averaged aerodynamics, exhibits weak stability without a controller or sufficient power density. However, the controller effectively stabilizes the PAV model, addressing attitude and maneuverability. These preliminary findings offer valuable insights for the mechanical design, aerodynamics, and fabrication of RoboMos, an insect-inspired flapping wing pico aerial vehicle developed at UPM Malaysia.

The AAA-ATPase Ter94 regulates wing size in Drosophila by suppressing the Hippo pathway

Insect wing development is a fascinating and intricate process that involves the regulation of wing size through cell proliferation and apoptosis. In this study, we find that Ter94, an AAA-ATPase, is essential for proper wing size dependently on its ATPase activity. Loss of Ter94 enables the suppression of Hippo target genes. When Ter94 is depleted, it results in reduced wing size and increased apoptosis, which can be rescued by inhibiting the Hippo pathway. Biochemical experiments reveal that Ter94 reciprocally binds to Mer, a critical upstream component of the Hippo pathway, and disrupts its interaction with Ex and Kib. This disruption prevents the formation of the Ex-Mer-Kib complex, ultimately leading to the inactivation of the Hippo pathway and promoting proper wing development. Finally, we show that hVCP, the human homolog of Ter94, is able to substitute for Ter94 in modulating Drosophila wing size, underscoring their functional conservation. In conclusion, Ter94 plays a positive role in regulating wing size by interfering with the Ex-Mer-Kib complex, which results in the suppression of the Hippo pathway.

Machine learning reveals the control mechanics of an insect wing hinge.

Insects constitute the most species-rich radiation of metazoa, a success that is due to the evolution of active flight. Unlike pterosaurs, birds and bats, the wings of insects did not evolve from legs1, but are novel structures that are attached to the body via a biomechanically complex hinge that transforms tiny, high-frequency oscillations of specialized power muscles into the sweeping back-and-forth motion of the wings2. The hinge consists of a system of tiny, hardened structures called sclerites that are interconnected to one another via flexible joints and regulated by the activity of specialized control muscles. Here we imaged the activity of these muscles in a fly using a genetically encoded calcium indicator, while simultaneously tracking the three-dimensional motion of the wings with high-speed cameras. Using machine learning, we created a convolutional neural network3 that accurately predicts wing motion from the activity of the steering muscles, and an encoder-decoder4 that predicts the role of the individual sclerites on wing motion. By replaying patterns of wing motion on a dynamically scaled robotic fly, we quantified the effects of steering muscle activity on aerodynamic forces. A physics-based simulation incorporating our hinge model generates flight manoeuvres that are remarkably similar to those of free-flying flies. This integrative, multi-disciplinary approach reveals the mechanical control logic of the insect wing hinge, arguably among the most sophisticated and evolutionarily important skeletal structures in the natural world.

Allometric Scaling Reveals Evolutionary Constraint on Odonata Wing Cellularity via Critical Crack Length.

Scaling in insect wings is a complex phenomenon that seems pivotal in maintaining wing functionality. In this study, the relationship between wing size and the size, location, and shape of wing cells in dragonflies and damselflies (Odonata) is investigated, aiming to address the question of how these factors are interconnected. To this end, WingGram, the recently developed computer-vision-based software, is used to extract the geometric features of wing cells of 389 dragonflies and damselfly wings from 197 species and 16 families. It has been found that the cell length of the wings does not depend on the wing size. Despite the wide variation in wing length (8.42 to 56.5 mm) and cell length (0.1 to 8.5 mm), over 80% of the cells hada length ranging from 0.5 to 1.5 mm, which waspreviously identified as the critical crack length of the membrane of locust wings. An isometric scaling of cells is also observed with maximum size in each wing, which increased as the size increased. Smaller cells tended to be more circular than larger cells. The results have implications for bio-mimetics, inspiring new materials and designs for artificial wings with potential applications in aerospace engineering androbotics.

Experimental Investigation on Aerodynamic Performance of Inclined Hovering with Asymmetric Wing Rotation.

This study presents a model experiment method that can accurately reproduce the flapping motion of insect wings and measure related unsteady aerodynamic data in real time. This method is applied to investigate the aerodynamic characteristics of inclined hovering, which distinguishes it from normal hovering by having asymmetric wing rotation during the two half strokes. In the study of the aerodynamic influence of the downstroke rotational angle, it is found that the rotational angle affects lift generation by changing the angle between the wing surface and the horizontal plane in the mid-downstroke. When the wing is almost parallel to the horizontal plane in the mid-downstroke, the vortex structure can maintain structural integrity and a large magnitude, which is conducive to the generation of high lift. In the study of the aerodynamic effect of the upstroke rotational angle, the windward conversion mechanism is proposed to explain the influence of the upstroke rotational angle on the direction and magnitude of thrust. Obtaining the rotational angle that is most conducive to maintaining the flight state of hovering in the present study can provide guidance for the structural design and kinematic control of micro aerial vehicles.

Bioinspired compliant joints for passive-automatic adaptability

Modern engineering requires structural components that adapt to continuously changing conditions. Adaptability leads to multi-functionality, enhanced durability, and functional efficiency of mechanical systems. That is why adaptive structures have received increasing attention, especially in robotics, aerospace, automotive, and biomedical engineering. Current adaptive structures require significant active actuations and controls, factors that add to their complexity, weight, and production costs. In contrast, most biological adaptive systems respond to external stimuli in a passive-automatic way. Here, inspired by a striking biological example, i.e. insect wings and specifically their vein-joints, we develop structures in which adaptability is determined by the presence of a network of bioinspired compliant joints. By systematically changing the key geometric parameters of the bioinspired joints, we show that they can be designed in a variety of patterns to passively control the adaptability of the structures under loading. We further test the performance of the bioinspired joints in practice. The simplicity, scalability, and adjustability are the features that facilitate the widespread community uptake of our designs.

WingSegment: A Computer Vision‐Based Hybrid Approach for Insect Wing Image Segmentation and 3D Printing

This article introduces WingSegment, a MATLAB app‐designed tool employing a hybrid approach of computer vision and graph theory for precise insect wing image segmentation. WingSegment detects cells, junctions, Pterostigma, and venation patterns, measuring geometric features and generating Voronoi patterns. The tool utilizes region‐growing, thinning, and Dijkstra's algorithms for boundary detection, junction identification, and vein path extraction. It provides histograms and box plots of geometric features, facilitating comprehensive wing analysis. WingSegment's efficiency is validated through comparisons with established tools and manual measurements, demonstrating accurate results. The tool further enables exporting detected boundaries as FreeCAD macro files for 3D modeling and printing, supporting finite element analysis. Beyond advancing insect wing morphology understanding, WingSegment holds broader implications for diverse planar structures, including leaves and geocells. This tool not only enhances automated geometric analysis and 3D model generation in insect wing studies but also contributes to the broader advancement of analysis, 3D printing, and modeling technologies across various planar structures.

Multidimensional vibrational circular dichroism for insect wings: Comparison of species.

This study reports the microscopic measurements of vibrational circular dichroism (VCD) on four different insect wings using a quantum cascade laser VCD system equipped with microscopic scanning capabilities (named multi-dimensional VCD [MultiD-VCD]). Wing samples, including (i) beetle, Anomala albopilosa (female), (ii) European hornet, Verspa crabro flavofasciata Cameron, 1903 (female), (iii) tiny dragonfly, Nannophya pygmae Rambur, 1842 (male), and (iv) dragonfly, Symetrum gracile Oguma, 1915 (male), were used in this study. Two-dimensional patterns of VCD signals (~10 mm × 10mm) were obtained at a spatial resolution of 100 μm. Measurements covered the absorption peaks assigned to amides I and II in the range of 1500-1740 cm-1 . The measurements were based on the enhancement of VCD signals for the stereoregular linkage of peptide groups. The patterns were remarkably dependent on the species. In samples (i) and (ii), the wings comprised segregated domains of protein aggregates of different secondary structures. The size of each microdomain was approximately 100 μm. In contrast, no clear VCD spectra were detected in samples (iii) and (iv). One possible reason was that the chain of stereoregular polypeptides was too short to achieve VCD enhancement in samples (iii) and (iv). Notably, the unique features were only observed in the VCD spectra because the IR spectra were nearly the same among the species. The VCD results hinted at the connection of protein microscopic structures with the wing flapping mechanisms of each species.

Novel Computational Design of Polymer Micromachined Insect-Mimetic Wings for Flapping-Wing Nano Air Vehicles.

The flapping wings of insects undergo large deformations caused by aerodynamic forces, resulting in cambering. Insect-mimetic micro wings for flapping-wing nano air vehicles mimic these characteristic deformations. In this study, a 2.5-dimensional insect-mimetic micro wing model for flapping-wing nano air vehicles is proposed to realize this type of wing. The proposed model includes a wing membrane, a leading edge, a center vein, and a root vein, all of which are modeled as shell elements. The proposed wing is a 2.5-dimensional structure and can thus be fabricated using polymer micromachining. We conducted a design window search to demonstrate the capabilities of the wing. The design windows, which are areas of desirable design solutions in the design parameter space, are iteratively searched using nonlinear finite-element analysis under quasi-steady aerodynamic modeling. Here, thickness is selected as a design parameter. The properties of real insects, polymer materials, and fabrication conditions are used to determine the other parameters. A fabricable design solution that generates sufficient camber is found from the design windows.

Osiris17 is essential for stable integrin localization and function during insect wing epithelia remodeling

The dynamic adhesion between cells and their extracellular matrix is essential for the development and function of organs. During insect wing development, two epithelial sheets contact each other at their basal sites through the interaction of βPS integrins with the extracellular matrix. We report that Osiris17 contributes to the maintenance of βPS integrins localization and function in developing wing of Drosophila and locust. In flies with reduced Osiris17 expression the epithelia sheets fail to maintain the integrity of basal cytoplasmic junctional bridges and basal adhesion. In contrast to the continuous basal integrin localization in control wings, this localization is disrupted during late stages of wing development in Osiris17 depleted flies. In addition, the subcellular localization revealed that Osiris17 co-localizes with the endosomal markers Rab5 and Rab11. This observation suggests an involvement of Osiris17 in endosomal recycling of integrins. Indeed, Osiris17 depletion reduced the numbers of Rab5 and Rab11 positive endosomes. Moreover, overexpression of Osiris17 increased co-localization of Rab5 and βPS integrins and partially rescued the detachment phenotype in flies with reduced βPS integrins. Taken together, our data suggest that Osiris17 is an endosome related protein that contributes to epithelial remodeling and morphogenesis by assisting basal integrins localization in insects.

On the effects of rotor induced vibrational stability on helicopter flight dynamics

This article investigates vibrational stabilization effects in rotorcraft flight dynamics. This study is motivated by recent results in flapping-wing flight, which showed that the time-varying aerodynamic and inertial loads due to the insect wing periodic motion induce a vibrational stabilization mechanism in hover. The dynamics of flapping-wing flyers and rotary-wing vehicles are both described by time-periodic systems so vibrational stabilization mechanisms can also have an effect on stability characteristics of rotary-wing vehicles. The article extends the use of the harmonic decomposition method to vibrational stability analysis of rotorcraft. Two cases are considered: vibrational stability due blade imbalance at hover, and vibrational stability due to number-of-blades-per-rotor-revolution (Nb\\documentclass[12pt]{minimal} \\usepackage{amsmath} \\usepackage{wasysym} \\usepackage{amsfonts} \\usepackage{amssymb} \\usepackage{amsbsy} \\usepackage{mathrsfs} \\usepackage{upgreek} \\setlength{\\oddsidemargin}{-69pt} \\begin{document}$$N_b$$\\end{document}/rev) in high-speed forward flight. Results show that while vibrations induced by rotor blade imbalance do not stabilize the hovering dynamics of a helicopter, these vibrations can still have a significant effect on the hovering dynamics. Rotor blade imbalance results in a symmetric effect on the roll and pitch axes, in that it tends to decrease the frequency of the subsidence modes of the hovering cubic, while the unstable oscillatory modes tend to increase in frequency and decrease in damping (destabilizing effect). On the other hand, the yaw/heave dynamics are relatively unaffected compared to the lateral and longitudinal axes. Moreover, Nb\\documentclass[12pt]{minimal} \\usepackage{amsmath} \\usepackage{wasysym} \\usepackage{amsfonts} \\usepackage{amssymb} \\usepackage{amsbsy} \\usepackage{mathrsfs} \\usepackage{upgreek} \\setlength{\\oddsidemargin}{-69pt} \\begin{document}$$N_b$$\\end{document}/rev rotor loads in forward flight are shown to reduce the damping of the coupled roll/pitch oscillation mode.

I-GLAD: a new strategy for fabricating antibacterial surfaces

The paper uses inverted glancing angle deposition (I-GLAD) for creating antibacterial surfaces. Antibacterial surfaces are found in nature, such as on insect wings, eyes, and plant leaves. Since the bactericidal mechanism is purely physical for these surfaces, the antimicrobial resistance of bacteria to traditional chemical antibiotics can be overcome. The technical problem is how to mimic, synthesize, and scale up the naturally occurring antibacterial surfaces for practical applications, given the fact that most of those surfaces are composed of three-dimensional hierarchical micro-nano structures. This paper proposes to use I-GLAD as a novel bottom-up nanofabrication technique to scale up bio-inspired nano-structured antibacterial surfaces. Our innovative I-GLAD nanofabrication technique includes traditional GLAD deposition processes alongside the crucial inverting process. Following fabrication, we explore the antibacterial efficacy of I-GLAD surfaces using two types of bacteria: Escherichia coli (E. coli), a gram-negative bacterium, and Staphylococcus aureus (S. aureus), a gram-positive bacterium. Scanning electron microscopy (SEM) shows the small tips and flexible D/P (feature size over period) ratio of I-GLAD nanoneedles, which is required to achieve the desired bactericidal mechanism. Antibacterial properties of the I-GLAD samples are validated by achieving flat growth curves of E. coli and S. aureus, and direct observation under SEM. The paper bridges the knowledge gaps of seeding techniques for GLAD, and the control/optimization of the I-GLAD process to tune the morphologies of the nano-protrusions. I-GLAD surfaces are effective against both gram-negative and gram-positive bacteria, and they have tremendous potentials in hospital settings and daily surfaces.

Wing expansion functional analysis of ion transport peptide gene in Sogatella furcifera (Horváth) (Hemiptera: Delphacidae)

Ion transport peptide (ITP), a superfamily of arthropod neuropeptides, serves a crucial role in regulating various physiological processes such as diuresis, ecdysis behavior, and wing expansion. However, the molecular characteristics, expression profile, and role of ITP in Sogatella furcifera are poorly understood. To elucidate the characteristics and biological function of ITP in S. furcifera, we employed reverse transcription-polymerase chain reaction (RT-PCR) and RNA interference (RNAi) methods. The identified SfITP gene encodes 117 amino acids. The expression of SfITP gradually increased followed the formation of 3-day-old of 5th instar nymph, peaking initially at 40 min after eclosion, and reaching another peak 24 h after eclosion, with particularly high expression levels in thorax and wing tissues. Notably, SfITP RNAi in 3rd instar nymphs of S. furcifera significantly inhibited the transcript levels of SfITP, resulting in 55% mortality and 78% wing deformity. These findings suggests that SfITP is involved in the regulation of wing expansion in S. furcifera, providing insights into the regulation of insect wing expansion and contributing to the molecular understanding of this process.

Intraspecific Variation in the Placement of Campaniform Sensilla on the Wings of the Hawkmoth Manduca Sexta.

Flight control requires active sensory feedback, and insects have many sensors that help them estimate their current locomotor state, including campaniform sensilla (CS), which are mechanoreceptors that sense strain resulting from deformation of the cuticle. CS on the wing detect bending and torsional forces encountered during flight, providing input to the flight feedback control system. During flight, wings experience complex spatio-temporal strain patterns. Because CS detect only local strain, their placement on the wing is presumably critical for determining the overall representation of wing deformation; however, how these sensilla are distributed across wings is largely unknown. Here, we test the hypothesis that CS are found in stereotyped locations across individuals of Manduca sexta, a hawkmoth. We found that although CS are consistently found on the same veins or in the same regions of the wings, their total number and distribution can vary extensively. This suggests that there is some robustness to variation in sensory feedback in the insect flight control system. The regions where CS are consistently found provide clues to their functional roles, although some patterns might be reflective of developmental processes. Collectively, our results on intraspecific variation in CS placement on insect wings will help reshape our thinking on the utility of mechanosensory feedback for insect flight control and guide further experimental and comparative studies.

Design and Integration of a Flapping Wing Apparatus

Abstract In this paper, we presented the design, integration, and experimental verification of a flapping wing apparatus. The purpose for this apparatus is to provide a framework to study the applicability of various types of sensors on flapping wings in the presence of forward speed. This work is inspired by the discoveries of mechanosensory hairs on insect wings that perform like strain-gauge sensors. To design the apparatus, we started by kinematic analysis of a crank-slider mechanism to actuate the wings. After that, we constructed the equations of motion of the entire system to find the proper gear ratio, motor properties, and other geometric dimensions. For the aerodynamic modeling, we used a quasi-steady formulation and presented a closed-form solution for the aerodynamic torque. Then, we explained the integration process and manufacturing of the main parts and presented two prototypes for the apparatus. At the end, we showed the final constructed versions of the apparatus and presented the experimental response and compared them with the simulation.

Bacterial elimination via cell membrane penetration by violet phosphorene peripheral sub-nanoneedles combined with oxidative stress.

The effectiveness of an antibacterial agent is strongly influenced by its antibacterial mechanism, which, in turn, depends on the agent's topological structure. In the natural world, the nanoprotrusions on the surface of insect wings give them excellent antimicrobial properties through physical penetration while being compatible with host cells. Inspired by the novel nanostructure of insect wings, violet phosphorus (VP), a new member of the phosphorus family, has antibacterial potential due to the sub-nanoneedle on its edge. Here, we demonstrate that VP and its exfoliated product, violet phosphorene nanosheets (VPNSs), have superior antibacterial capability against pathogens via cell membrane penetration induced by peripheral sub-nanoneedles combined with oxidative stress. The results show that VPNSs can inactivate more than 99.9% of two common pathogens (Escherichia coli and Staphylococcus aureus) and more than 99.9% of two antibiotic-resistant bacteria (Escherichia coli pUC19 and methicillin-resistant Staphylococcus aureus), while showing almost no toxicity toward normal cells at a high concentration of 2.0 mg mL-1. Moreover, VPNSs can achieve effective treatment of induced skin wound infections and bacterial keratitis (BK) by Staphylococcus aureus and methicillin-resistant Staphylococcus aureus, respectively, showing promising potential for ocular and skin wound infection theragnostics.

Multifunctionally disordered TiO2 nanoneedles prevent periprosthetic infection and enhance osteointegration by killing bacteria and modulating the osteoimmune microenvironment.

Rationale: Total hip arthroplasty (THA) and total knee arthroplasty (TKA) are effective interventions for end-stage osteoarthritis; however, periprosthetic infection is a devastating complication of arthroplasty. To safely prevent periprosthetic infection and enhance osteointegration, the surface modification strategy was utilized to kill bacteria, modulate the osteoimmune microenvironment, and improve new bone formation. Methods: We used the hydrothermal method to fabricate a bionic insect wing with the disordered titanium dioxide nanoneedle (TNN) coating. The mussel-inspired poly-dopamine (PDA) and antibacterial silver nanoparticles (AgNPs) were coated on TNN, named AgNPs-PDA@TNN, to improve the biocompatibility and long-lasting bactericidal capacity. The physicochemical properties of the engineered specimen were evaluated with SEM, AFM, XPS spectrum, and water contact assay. The biocompatibility, bactericidal ability, and the effects on macrophages and osteogenic differentiation were assessed with RT-qPCR, Western blotting, live/dead staining, immunofluorescent staining, etc. Results: The AgNPs-PDA@TNN were biocompatible with macrophages and exhibited immunomodulatory ability to promote M2 macrophage polarization. In addition, AgNPs-PDA@TNN ameliorated the cytotoxicity caused by AgNPs, promoted cell spreading, and increased osteogenesis and matrix deposition of BMSCs. Furthermore, AgNPs-PDA@TNN exhibited bactericidal ability against E. coli and S. aureus by the bionic nanostructure and coated AgNPs. Various imaging analyses indicated the enhanced bactericidal ability and improved new bone formation by AgNPs-PDA@TNN in vivo. H&E, Gram, and Masson staining, verified the improved bone formation, less inflammation, infection, and fibrosis encapsulation. The immunofluorescence staining confirmed the immunomodulatory ability of AgNPs-PDA@TNN in vivo. Conclusion: The bionic insect wing AgNPs-PDA@TNN coating exhibited bactericidal property, immunomodulatory ability, and enhanced osteointegration. Thus, this multidimensional bionic implant surface holds promise as a novel strategy to prevent periprosthetic infection.

A hydrogen isoscape for tracing the migration of herbivorous lepidopterans across the Afro-Palearctic range.

Many insect species undertake multigenerational migrations in the Afro-tropical and Palearctic ranges, and understanding their migratory connectivity remains challenging due to their small size, short life span and large population sizes. Hydrogen isotopes (δ2 H) can be used to reconstruct the movement of dispersing or migrating insects, but applying δ2 H for provenance requires a robust isotope baseline map (i.e. isoscape) for the Afro-Palearctic. We analyzed the δ2 H in the wings (δ2 Hwing ) of 142 resident butterflies from 56 sites across the Afro-Palearctic. The δ2 Hwing values were compared to the predicted local growing-season precipitation δ2 H values (δ2 HGSP ) using a linear regression model to develop an insect wing δ2 H isoscape. We used multivariate linear mixed models and high-resolution and time-specific remote sensing climate and environmental data to explore the controls of the residual δ2 Hwing variability. A strong linear relationship was found between δ2 Hwing and δ2 HGSP values (r2 = 0.53). The resulting isoscape showed strong patterns across the Palearctic but limited variation and high uncertainty for the Afro-tropics. Positive residuals of this relationship were correlated with dry conditions for the month preceding sampling whereas negative residuals were correlated with more wet days for the month preceding sampling. High intra-site δ2 Hwing variance was associated with lower relative humidity for the month preceding sampling and higher elevation. The δ2 Hwing isoscape is applicable for tracing herbivorous lepidopteran insects that migrate across the Afro-Palearctic range but has limited geolocation potential in the Afro-tropics. The spatial analysis of uncertainty using high-resolution climatic data demonstrated that many African regions with highly variable evaporation rates and relative humidity have δ2 Hwing values that are less related to δ2 HGSP values. Increasing geolocation precision will require new modeling approaches using more time-specific environmental data and/or independent geolocation tools.

Thrust force is tuned by the rigidity distribution in insect-inspired flapping wings

We study the aerodynamics of a flapping flexible wing with a two-vein pattern that mimics the elastic response of insect wings in a simplified manner. The experiments reveal a non-monotonic variation of the thrust force produced by the wings when the angle between the two veins is varied. An optimal configuration is consistently reached when the two veins are spaced at an angle of about 20 degrees. This value is in the range of what has been measured in the literature for several insect species. The deformation of the wings is monitored during the experiment using video recordings, which allows to pinpoint the physical mechanism behind the non-monotonic behaviour of the force curve and the optimal distribution of the vein network in terms of propulsive force.