Figure - uploaded by Hamed Rajabi
Content may be subject to copyright.
Graphical Abstract
Source publication
Insects thrived soon after they acquired the ability to fly. Beyond the reach of the non-flying competitors, flying insects colonized a wide variety of habitats. Although flight is an efficient way to disperse and escape predators, it is energetically costly. Hence, various strategies are served to enhance flight efficiency as much as possible. A s...
Citations
... When we approach nature from an engineering perspective and focus on the mechanical design of natural systems, we uncover collective mechanical behaviours and emergent mechanical properties resulting from their specialized design. For example, observations of snake locomotion systems [9][10][11], fish body armours [12][13][14], gecko adhesive pads [15][16][17][18], insect flight systems [19][20][21][22][23][24], beetle fighting mechanisms [25,26] and many more indicate that natural mechanical systems consist of complicated material composition, nano-and micro-architecture and structural elements. This complicatedness can arise from two underlying design principles: (1) a network of simple design elements that collectively form a complex system and/or (2) a collection of complicated subsystems that together constitute the entire system. ...
Presenting a novel framework for sustainable and regenerative design and development is a fundamental future need. Here we argue that a new framework, referred to as complexity biomechanics, which can be used for holistic analysis and understanding of natural mechanical systems, is key to fulfilling this need. We also present a roadmap for the design and development of intelligent and complex engineering materials, mechanisms, structures, systems, and processes capable of automatic adaptation and self-organization in response to ever-changing environments. We apply complexity biomechanics to elucidate how the different structural components of a complex biological system as dragonfly wings, from ultrastructure of the cuticle, the constituting bio-composite material of the wing, to higher structural levels, collaboratively contribute to the functionality of the entire wing system. This framework not only proposes a paradigm shift in understanding and drawing inspiration from natural systems but also holds potential applications in various domains, including materials science and engineering, biomechanics, biomimetics, bionics, and engineering biology.
... [1] Understanding the complex morphology of insect wings is not merely an academic pursuit in multiple scientific domains, such as biomechanics, animal behavior, physiology, and ecology, [2][3][4][5][6][7] but an inspiration for innovative technologies in the field of engineering. These include the development of robust load-bearing structures, [8][9][10] bioinspired joints, [11][12][13][14] insect-inspired hinges, [15] bioinspired wing, [16,17] insectinspired composites, [18,19] bioinspired attachment strategies, [20][21][22] and bioinspired grippers. [23] As our understanding of the wing structures deepens, manual analysis methods face challenges in accurately capturing the complex geometric features to characterize wing morphology [24,25] comprehensively. ...
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.
... [196] Their flexible, resilin-bearing bases and their rather stiff, hooked tips enable the hamuli to maintain a tight but flexible forewing-hindwing connection at alternating wing movements and forewing-hindwing tilt angles (allowing large angle variations of 135-235°between the wings), which is crucial for the wings' aerodynamic efficiency (especially by building a zone of ventral flexion (claval flexion line and connection line between forewing and hindwing) and thereby allowing strong camber reversal during the upstroke). [122,[196][197][198] In addition, the movability of the hamuli allows certain lateral sliding along the ridge and was suggested to be advantageous for decoupling and re-coupling. [196] In this context, it is interesting that bees are able to re-couple their wings even in flight when they were uncoupled because of, e.g., contact to the vegetation (www.ibycter.wordpress.com/tag/hamuli/ ...
... Furthermore, the resilindominated membrane was suggested to increase the durability and reduce the risk of failure of the hamuli structures throughout a bee's lifetime. [196,198] The number of distal hamuli varies widely within the Hymenoptera (ranging from two to over sixty), and sometimes differences in the number of distal hamuli occur even within one species and sometimes between the two hindwings of one individual. [193,199] Schwarz proposed that the number of hamuli can be related to body weight and (foraging) flight range, which is in agreement with the observation that the foraging flight range correlates with the number of hamuli in the alfalfa leafcutting bee Megachile femorata (Megachilidae). ...
Compared to wingless insects, pterygote insects profit from numerous wing‐related benefits including a wider distribution range, the exploitation of various food resources and the escape from water‐ or land‐confined predators. In order to maintain the wings´ functionality, the wing design and resistance to material fatigue are of key importance. This is even more essential for survival when considering that wings are used for millions of wing beat cycles but cannot be repaired and do not contain inner muscles so that their aerodynamic performance is mainly based on passive, structure‐based wing deformations. One of the components serving this purpose is the endowment of certain wing components with the elastomeric protein resilin building stable and complex material composites with the tanned cuticle. Resilin endows the respective structures with, e.g., higher flexibility and compliance and enables elastic energy storage. In this study, the occurrence of resilin in the insect flight system is reviewed based on previous studies of several insect orders including Odonata, Orthoptera, Hymenoptera, Coleoptera, Dermaptera, and Diptera, and the function of resilin is discussed with reference to the respective structures.
... One-way hinges and asymmetric bending and twisting are widespread in insect wings (1,2,12,19,(24)(25)(26)(27). They operate in flight, allowing the wings to deform automatically and asymmetrically between the upstroke and the downstroke and also in controlling the precise, complex patterns of folding and unfolding in the hind wings of beetles (Coleoptera) and of earwigs (Dermaptera) (17,18,28,29). ...
Insect wings are deformable airfoils, in which deformations are mostly achieved by complicated interactions between their structural components. Due to the complexity of the wing design and technical challenges associated with testing the delicate wings, we know little about the properties of their components and how they determine wing response to flight forces. Here, we report an unusual structure from the hind-wing membrane of the beetle Pachnoda marginata. The structure, a transverse section of the claval flexion line, consists of two distinguishable layers: a bell-shaped upper layer and a straight lower layer. Our computational simulations showed that this is an effective one-way hinge, which is stiff in tension and upward bending but flexible in compression and downward bending. By systematically varying its design parameters in a computational model, we showed that the properties of the double-layer membrane hinge can be tuned over a wide range. This enabled us to develop a broad design space, which we later used for model selection. We used selected models in three distinct applications, which proved that the double-layer hinge represents a simple yet effective design strategy for controlling the mechanical response of structures using a single material and with no extra mass. The insect-inspired, one-way hinge is particularly useful for developing structures with asymmetric behavior, exhibiting different responses to the same load in two opposite directions. This multidisciplinary study not only advances our understanding of the biomechanics of complicated insect wings but also informs the design of easily tunable engineering hinges.
... In recent years, studies on the biomechanics and functionality of the fore-and hindwing coupling have been mainly focused on three types of wing-coupling mechanisms, including hook-furrow coupling (caddisflies, bark lice, and bees), [1][2][3][4][5][6][7] clamp-furrow coupling (bugs) [8][9][10][11][12][13] and amplexiform coupling (butterflies). 14 The above literature demonstrated the comparative morphology and biomechanical properties (including sliding-friction reduction and anti-structural-damage strategies) of wing-couplings among two or more insect groups. ...
... These literature data aimed to elucidate the working principles that the insects applied in their evolution to keep a firm foreand hindwing coupling and overcome millions of wingflapping cycles in flight without damage. 15,16 Since 2019, Prof. Gorb and his collaborators have consistently studied the wing-coupling mechanisms of insects, especially the hook-furrow coupling of honeybees [5][6][7]17 (this study). In detail, the bee wing-coupling is generally composed of a set of hamuli (hooks) at the leading edge of the hindwing and a posterior recurved margin (PRM) at the trailing edge of the forewing (Fig. 1). ...
... In detail, the bee wing-coupling is generally composed of a set of hamuli (hooks) at the leading edge of the hindwing and a posterior recurved margin (PRM) at the trailing edge of the forewing (Fig. 1). Until now, the structure, material composition, biomechanics, aerodynamics and even bioinspired application of the bee wing-coupling, to some extent, have been systematically investigated, [5][6][7]17 devoted to revealing the likely significance of cuticle sclerotization and hamulus morphology on the performance of the coupling. However, one issue which needed to be improved on the mechanical properties of the coupling was only focusing on the mechanical performance of hamuli without considering the PRM deformation as well as the likelihood of its structural failure. ...
Fore- and hindwings of honeybees are coupled and synchronized to flap by means of a forewing posterior recurved margin (PRM) and hindwing hamuli which constitute a hook-furrow coupling. Morphological analysis shows that the PRM is composed of a thickened and sclerotized membrane with the Archimedean spiral configuration and hamuli are a set of tiny, sclerotized hooks with flexible bases. By developing a theoretical PRM model, the influence of cuticle sclerotization and membrane-thickening on a deforming pattern and maximal coupling force was comparatively simulated, indicating that the real PRM is capable of bearing the highest coupling force and the membrane thickening makes more contribution than cuticle sclerotization on augmenting the maximal coupling force that the PRM can resist. In addition, four combined strategies, i.e. the hook shape, Archimedean spiral, rich resilin concentration, and cuticle sclerotization in different parts of the whole system were proposed, and deemed to endow the honeybee wing-coupling with remarkable stability and durability to eliminate a potential structural failure of the coupling over millions of wing flapping cycles across the honeybee lifespan. This study assists us in the comprehensive understanding of the functionality of the hook-furrow wing-coupling and shows us new avenues for biomimetics of mobile coupling mechanisms in modern engineering.
... Both mosquito antennae and honey bee tongues are long-lasting functional organs with high work intensity, subjected to continuous mechanical stress. For these animal appendages, the addition of soft material to a hard structure can effectively protect them from wear and tear [35] . The antennae of a mosquito are covered in functional fibrillae that served to enhance auditory sense [36] . ...
The honey bee, Apis mellifera ligustica, uses the specialized tongue structured by ∼120 segmental units, coated by bushy hairs, to dip varying concentration nectar flexibly at small scales. While dipping, the segmental units elongate by 20%, coordinated with rhythmical erection of hairs, the pattern of which is demonstrated to be capable of both increasing nectar intake rate and saving energy. The compliance in the segmental units allows extension of the tongue, which however, challenges the structural stability while traveling through the viscous fluid. In this combined experimental and theoretical investigation, we apply scanning electron microscopy (SEM), confocal laser scanning microscopy (CLSM), micro-computed tomography scanning (micro-CT), atomic force microscopy (AFM), and mechanical models to reveal the structural and material specialisations in a bee tongue for meeting the functionally contradictive demands. We find that each segmental unit is a complex structure, which is composed of an intersegmental membrane (ISM) and a ring-like hair base (RHB), with spatially distributed discrete changes in material properties. The combination of these two components makes the tongue multifunctional, in which the ISMs characterized by resilin-rich material make the segmental units compliant, while the RHBs with rigid sclerotized material provide stable supporting for hairs. Our study may enlighten deployable mechanisms with correlative functional components, especially the microscopic mechanisms applied in viscous fluid tranport.
Statement of significance
The honey bee tongue is a versatile tool that extends to probe into varying-shaped corollas, retracting with 3,000 glossal hairs staying erected to load nectar. The combined requirement of both deformability and structural stability imposes opposing demands on structural stiffness. Here we show that glossal hairs are supported by rigid continuum ring-like hair bases, embedded in the elastic resilient intersegmental membrane, making the whole tongue both flexible and rigid at the same time. Our findings extend our understanding of relationship between morphology, material composition and biomechanics of dynamic biological surfaces, which may inspire design paradigms of multifunctional deployable mechanisms coordinating deformability and structural stability.
... For this purpose, we estimated the elastic modulus of 600 uniformly distributed points on the rostrum and then measured their mean value. The Poisson's ratio of the model was set as 0.3, which is the same as that of many previously investigated biological materials [20] and has frequently been used for models of insect cuticle [21][22][23]. ...
Elongated rostra (snouts) are remarkable features of many female weevils. The female of Curculio glandium uses the snout to excavate channels in acorns to oviposit. Considering the slenderness of the rostrum, the excavation of channels in solid substrates without buckling is a challenging task from both engineering and biological points of view. Here we aimed to examine the roles of the material properties and morphology of the rostrum in its buckling resistance. We employed microscopy techniques, non-destructive material characterisation and finite element (FE) modelling to shed more light on the excavation mechanics of the rostrum. We found that sexual dimorphisms are present not only in the length but also in the material, particularly the elastic modulus, and morphological features, particularly the curvature and thickness of the cuticular layers. Our FE modelling showed that those factors play essential roles to maximise the buckling resistance and minimise the bending resistance of the female rostrum. Considering that during excavation, the rostrum needs to be straightened without buckling, the functionality of the rostrum is likely to be a compromise between the flexibility and stiffness.
... Furthermore, the interlocking structures were supposed to not contain any structures with large proportions of resilin (Ma et al., 2019). The visualization of different exoskeleton autofluorescences was also performed in the third study that suggests that some exoskeleton material located at the bases of the hamuli might contain large resilin proportions, although this was not clearly visualized (Toofani et al., 2020). It is not only obvious that the results of these three studies are contradictory but also noticeable that they are in part not consistent with the commonly known nature of such structures that is described below. ...
... All results described so far are in accordance with the results obtained from earlier analyses of the autofluorescence composition of the wing-interlocking structures of honeybees (Ma et al., 2019;Toofani et al., 2020). In clear contrast to the results of Ma et al. (2019), indicating that the interlocking structures do not contain any structure with a large proportion of resilin, the present study confirms the assumption of Toofani et al. (2020) with respect to the presence of structures with large resilin proportions. ...
... All results described so far are in accordance with the results obtained from earlier analyses of the autofluorescence composition of the wing-interlocking structures of honeybees (Ma et al., 2019;Toofani et al., 2020). In clear contrast to the results of Ma et al. (2019), indicating that the interlocking structures do not contain any structure with a large proportion of resilin, the present study confirms the assumption of Toofani et al. (2020) with respect to the presence of structures with large resilin proportions. It clearly reveals that each hamulus base is surrounded by and embedded in material that features large proportions of resilin and is located in strongly sclerotised socket-like structures on the radius (e.g., Figs. ...
Hymenoptera are characterised by the presence of one forewing pair and one hindwing pair. The two wings of each body side are coupled to each other during flight making the morphologically four-winged insects functionally two-winged. This coupling is formed by a row of hook-like structures, called hamuli, that are located at the leading edge of the hindwing and interlock with a thickened and recurved margin present at the trailing edge of the forewing. In this study, autofluorescence analyses performed with confocal laser scanning microscopy revealed differences in the exoskeleton material composition of the interlocking structures. While the wing veins and the recurved margin are strongly sclerotised and chitinous, the wing membranes mainly contain the elastomeric protein resilin. The hamuli are composed of sclerotised chitinous material, and each hamulus base is surrounded by and embedded in material that features large proportions of resilin and is located in strongly sclerotised socket-like wing vein structures. This exoskeleton organisation likely allows movements of the hamuli and, in combination with the exoskeleton material gradients visualized in the other interlocking structures, is assumed to guarantee an effective wing coupling and to simultaneously decrease the risk of wear and damage under mechanical loads occurring in flight, coupling and decoupling situations.
... This setae is located well before the beginning of the fringe, and may function as part of a wing coupling apparatus during flight for better efficiency, or at rest. A variety of wing coupling mechanisms were reported in four-winged insects (Paraneoptera, Lepidoptera, Trichoptera and some Hymenoptera) (Stocks, 2008), and the structure and function of these mechanisms were subject to many studies (New, 1974;Ellington, 1980;D'Urso and Ippolito, 1994;D'Urso, 2002;Stocks, 2010aStocks, , 2010bOgawa and Yoshizawa, 2018;Toofani et al., 2020). The potential wing coupling apparatus observed in P. spinosum gen. ...
Four new protopsyllidioid species, Paraprotopsyllidium spinosum gen. et sp. nov., Angustipsyllidium minutum gen. et sp. nov., Burmapsyllidium setosum gen. et sp. nov., and Maliawa akrawna gen et sp. nov. are described from the mid-Cretaceous Burmese amber, and assigned to a new family, Paraprotopsyllidiidae fam. nov., that we establish. These taxa are characterized by their narrowed fore and hind wings, bearing a fringe of long setae; the possible functions of these bristles are explored in this paper. The population dynamics, the potential feeding diet and the biogeographical distribution of this family are briefly discussed.
... FE software packages were developed to simplify often complicated simulation processes. They are especially very common in engineering applications [3][4][5] and are becoming increasingly popular in the investigation of the mechanical behavior of biological structures, such as complex human and animal body parts [6][7][8][9][10][11][12][13]. ...
Simple Summary
Manual modeling of complicated insect wings presents considerable practical challenges. To overcome these challenges, therefore, we developed WingMesh. This is an application for simple yet precise automatic modeling of insect wings. Using a series of examples, we showed the performance of our application in practice. We expect WingMesh to be particularly useful in comparative studies, especially where the modeling of a large number of insect wings is required within a short time.
Abstract
The finite element (FE) method is one of the most widely used numerical techniques for the simulation of the mechanical behavior of engineering and biological objects. Although very efficient, the use of the FE method relies on the development of accurate models of the objects under consideration. The development of detailed FE models of often complex-shaped objects, however, can be a time-consuming and error-prone procedure in practice. Hence, many researchers aim to reach a compromise between the simplicity and accuracy of their developed models. In this study, we adapted Distmesh2D, a popular meshing tool, to develop a powerful application for the modeling of geometrically complex objects, such as insect wings. The use of the burning algorithm (BA) in digital image processing (DIP) enabled our method to automatically detect an arbitrary domain and its subdomains in a given image. This algorithm, in combination with the mesh generator Distmesh2D, was used to develop detailed FE models of both planar and out-of-plane (i.e., three-dimensionally corrugated) domains containing discontinuities and consisting of numerous subdomains. To easily implement the method, we developed an application using the Matlab App Designer. This application, called WingMesh, was particularly designed and applied for rapid numerical modeling of complicated insect wings but is also applicable for modeling purposes in the earth, engineering, mathematical, and physical sciences.
