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Coccinella septempunctata, left hind wing showing veins, resilin-containing areas and major folding and £exion lines. Digitized from a microscope slide. Terminology after Kukalova¨-Kukalova¨-Peck & Lawrence (1993). Rectangles a and b indicate joints shown in ¢gure 6. AA, anal anterior; CuA, cubitus anterior; af, anal fold; cf, claval fold; lf, longitudinal fold; mf, median £exion line; sf, sti¡ening fold, visible only in £ight and in £uorescent microscope; MP1 + 2, media posterior; RA, radius anterior; RP, radius posterior; ScA, subcosta anterior; tf1, ¢rst transverse fold; tf 2, second transverse fold.
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This account shows the distribution of elastic elements in hind wings in the scarabaeid Pachnoda marginata and coccinellid Coccinella septempunctata (both Coleoptera). Occurrence of resilin, a rubber-like protein, in some mobile joints together with data on wing unfolding and flight kinematics suggest that resilin in the beetle wing has multiple fu...
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... Particularly noteworthy is the large, untanned, resilin-dominated cuticular region of the aboral mandibular pad (amp) (figure 6x), which is consistent with the findings in other Odonata species [16,56,88]. This structure was supposed to provide flexibility and to allow elastic energy storage (energy generated by the 0md1 adductor muscle) and, thus, to prevent structural failure and support the masticating process [16,29,51,[89][90][91][92][93]. Furthermore, it was assumed to be used as a damper to secure the mandible against collisions with the head capsule due to large opening angles [56]. ...
The arthropod cuticle offers strength, protection, and lightweight. Due to its limit in expandability, arthropods have to moult periodically to grow. While moulting is beneficial in terms of parasite or toxin control, growth and adaptation to environmental conditions, it costs energy and leaves the soft animal's body vulnerable to injuries and desiccation directly after ecdysis. To investigate the temporal change in sclerotization and pigmentation during and after ecdysis, we combined macrophotography, confocal laser scanning microscopy, scanning electron microscopy and histological sectioning. We analysed the tarsal and mandibular cuticle of the blue emperor dragonfly to compare the progress of tanning for structures that are functionally involved during emergence (tarsus/tarsal claws) with structures whose functionality is required much later (mandibles). Our results show that: (i) the tanning of the tarsal and mandibular cuticle increases during emergence; (ii) the tarsal cuticle tans faster than the mandibular cuticle; (iii) the mandibles tan faster on the aboral than on the oral side; and (iv) both the exo- and the endocuticle are tanned. The change in the cuticle composition of the tarsal and mandibular cuticle reflects the demand for higher mechanical stability of these body parts when holding on to the substrate during emergence and during first walking or hunting attempts.
... Thus, phasic active spanwise-folding and deployment (PASFD) is found in beetle flight control. This newly discovered PASFD phenomenon further confirms and complements Hass's conjecture about the possible existence of phasic active claval-wing-folding in beetles (which was not experimentally confirmed due to the low frame rate of digital cameras at that time) [25]. When comparing the flapping motion with and without folding, we found that both the flapping frequencies were essentially stable and constant. ...
... Thus, phasic active spanwise-folding and deployment (PASFD) is found in beetle flight control. This newly discovered PASFD phenomenon further confirms and complements Hass's conjecture about the possible existence of phasic active clavalwing-folding in beetles (which was not experimentally confirmed due to the low frame rate of digital cameras at that time) [25]. ...
... Furthermore, we specially designed the cable-driven folding mechanism, using nitrile rubber as the elastic hinge. The elastic hinge mimics the function of resilin in biological wings, and is capable of storing elastic energy to provide the energy needed to unfold the wing [25]. Specifically, tightening of the nylon cord (from OB to OB′, see Figure 3c) results in elastic energy storage, and when the cord is loosened (from OB′ to OB) the elastic energy drives the deployment of the wing-tip. ...
Insects produce a variety of highly acrobatic maneuvers in flight owing to their ability to achieve various wing-stroke trajectories. Among them, beetles can quickly change their flight velocities and make agile turns. In this work, we report a newly discovered phasic wing-tip-folding phenomenon and its aerodynamic basis in beetles. The wings’ flapping trajectories and aerodynamic forces of the tethered flying beetles were recorded simultaneously via motion capture cameras and a force sensor, respectively. The results verified that phasic active spanwise-folding and deployment (PASFD) can exist during flapping flight. The folding of the wing-tips of beetles significantly decreased aerodynamic forces without any changes in flapping frequency. Specifically, compared with no-folding-and-deployment wings, the lift and forward thrust generated by bilateral-folding-and-deployment wings reduced by 52.2% and 63.0%, respectively. Moreover, unilateral-folding-and-deployment flapping flight was found, which produced a lateral force (8.65 mN). Therefore, a micro-flapping-wing mechanism with PASFD was then designed, fabricated, and tested in a motion capture and force measurement system to validate its phasic folding functions and aerodynamic performance under different operating frequencies. The results successfully demonstrated a significant decrease in flight forces. This work provides valuable insights for the development of flapping-wing micro-air-vehicles with high maneuverability.
... Among invertebrates, such mechanisms can be quite sophisticated, sometimes mediated by a latch (Ilton et al., 2018;Farley et al., 2019). Some arthropods have cuticle properties capable of storing elastic energy when deformed (Haas et al., 2000;B€ aumler and Büsse, 2019;Michels and Gorb, 2012). Many of these multifaceted power amplification systems, especially in small cryptic organisms, are not well described. ...
Springtails are notable for their jumping apparatus and latch-mediated spring mechanism. The challenge, in the light of the tiny size and rapid movement of these organisms, has been to understand the morphological intricacies of this spring system. This study takes an approach that integrates SEM, MicroCT, cLSM and high-speed video recordings to understand the composition and functionality of the jumping apparatus in Megalothorax minimus (Neelipleona), Dicyrtomina ornata and Dicyrtomina minuta (Symphypleona). We focus on reconstructing, describing, and understanding the functioning of structures such as basal plates, musculature and furca. The dimensions of the jumping apparatus in Dicyrtomina and Megalothorax differ significantly from those in elongated springtails. A hypothesis of functional coherence between taxa, based on muscle connections and basal plates, is postulated. High-speed video recordings provide information on: 1) furca release timing and function during jumping and self-righting; 2) performance properties of manubrium, dens and mucro in interaction with the ground and in take-off; 3) possible pre-release furca moves. The study underscores the need for further research employing a variety of visualization methods in order to explore additional aspects such as retinaculum unlatching and furca flexion/extension muscles.
... Resilin, a multipurpose biopolymer commonly found in insect wings [37][38][39][40] serves as an inspiration to expand this design space while enabling multistability. Resilin assigns mountain or valley folds to origamilike insect wings [41,42] and serves as an energy storage device, for example, in the (i) When β < 0, we have angular excess, and an e-Cone is observed. (ii) When β = 0, we are folding from a flat plane (traditional origami). ...
Non-Euclidean origami is a promising technique for designing multistable deployable structures folded from nonplanar developable surfaces. The impossibility of flat foldability inherent to non-Euclidean origami results in two disconnected solution branches each with the same angular deficiency but opposite handedness. We show that these regions can be connected via “crease stretching,” wherein the creases exhibit extensibility in addition to torsional stiffness. We further reveal that crease stretching acts as an energy storage method capable of passive deployment and control. Specifically, we show that in a Miura-Ori system with a single stretchable crease, this is achieved via two unique, easy to realize, equilibrium folding pathways for a certain wide set of parameters. In particular, we demonstrate that this connection mostly preserves the stable states of the non-Euclidean system, while resulting in a third stable state enabled only by the interaction of crease torsion and stretching. Finally, we show that this simplified model can be used as an efficient and robust tool for inverse design of multistable origami based on closed-form predictions that yield the system parameters required to attain multiple, desired stable shapes. This facilitates the implementation of multistable origami for applications in architecture materials, robotics, and deployable structures.
... This passive deformation mechanism needs to be clearly understood. Previous researches found that resilin-generally, the resilin-bearing vein jointswas found to contribute a lot to the chordwise flexibility of the wings of insects such as dragonflies, beetles, and bumblebees [6][7][8][9][10]. Although some of the conclusions are based on speculation, the functions of resilin on insect wings have attracted much attention. ...
The flexibility of insect wings should be considered in the design of bionic micro flapping-wing aircraft. The honeybee is an ideal biomimetic object because its wings are small and possess a concise vein pattern. In this paper, we focus on resilin, an important flexible factor in honeybees’ forewings. Both resilin joints and resilin stripes are considered in the finite element model, and their mechanical behaviors are studied comprehensively. Resilin was found to increase the static deflections in chordwise and spanwise directions by 1.4 times and 1.9 times, respectively. In modal analysis, natural frequencies of the first bending and first torsional modes were found to be decreased significantly—especially the latter, which was reduced from 500 Hz to 217 Hz—in terms of resilin joints and stripes, closely approaching flapping frequency. As a result, the rotational angle amplitude in dynamic responses is remarkable, with an amplification ratio of about six. It was also found that resilin joints and stripes together lead to well-cambered sections and improve the stress concentrations in dynamic deformation. As resilin is widespread in insect wings, the study could help our understanding of the flexible mechanism of wing structure and inspire the development of flexible airfoils.
... 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). The mechanisms often involve specific areas of elasticity within the wing cuticle, sometimes including lines or patches of the protein elastomer resilin (6,(30)(31)(32)(33)(34), but other examples depend on reversible thin-plate buckling, in which the shape and height as well as the elastic properties of the wing section are of major importance (1,8,15,19). The double-layer membrane structure described here can fit into the latter category. ...
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.
... Especially resilin, a viscoelastic, extracellular protein-matrix (Lerch et al., 2020;Weis-Fogh, 1960;Weis-Fogh, 1961), plays an essential role in a multitude of functions such as: (i) reducing wear and the risk of structural failure (e.g., Bäumler & Büsse, 2019;Haas et al., 2000aHaas et al., , 2000b, (ii) improving fatigue resistance (e.g., Haas et al., 2000aHaas et al., , 2000bRajabi et al., 2016a), (iii) allowing energy storage for high-speed movements (e.g., Burrows & Sutton, 2012;Büsse et al., 2021b;Gorb, 2004) or in insect attachment devices (e.g., Büscher et al., 2021;Büsse et al., 2019;Jandausch et al., 2018;Peisker et al., 2013;Petersen et al., 2018). The cuticular material composition of damselfly and dragonfly (Büsse et al., 2021a) larval mouthparts has recently been studied. ...
... Especially resilin, a viscoelastic, extracellular protein-matrix (Lerch et al., 2020;Weis-Fogh, 1960;Weis-Fogh, 1961), plays an essential role in a multitude of functions such as: (i) reducing wear and the risk of structural failure (e.g., Bäumler & Büsse, 2019;Haas et al., 2000aHaas et al., , 2000b, (ii) improving fatigue resistance (e.g., Haas et al., 2000aHaas et al., , 2000bRajabi et al., 2016a), (iii) allowing energy storage for high-speed movements (e.g., Burrows & Sutton, 2012;Büsse et al., 2021b;Gorb, 2004) or in insect attachment devices (e.g., Büscher et al., 2021;Büsse et al., 2019;Jandausch et al., 2018;Peisker et al., 2013;Petersen et al., 2018). The cuticular material composition of damselfly and dragonfly (Büsse et al., 2021a) larval mouthparts has recently been studied. ...
... F I G U R E 15 Anax imperator, visualization of the abduction movement of the labium (abdominal direction), with maximum mma (red), maximum paa (dark blue) and median paa (light blue); lateral view. congruent with observations made in different biomechanical systems of insects (Haas et al., 2000a;2000b;Michels, Vogt & Gorb, 2012;Rajabi et al., 2016aRajabi et al., , 2016b. Büsse and Gorb (2018) discussed the role of the mandibular pad in absorbing mechanical stress generated by the incisive teeth and the molar lobe, a function that could be attributed to the pad of A. imperator as well. ...
Insects evolved differently specialized mouthparts. We study the mouthparts of adult Anax imperator, one of the largest odonates found in Central Europe. Like all adult dragonflies, A. imperator possesses carnivorous‐type of biting‐chewing mouthparts. To gain insights into the feeding process, behavior and kinematics, living specimens were filmed during feeding using synchronized high‐speed videography. Additionally, the maximum angles of movement were measured using a measuring microscope and combined with data from micro‐computed tomography (µCT). The resulting visualizations of the 3D‐geometry of each mouthpart were used to study their anatomy and complement the existing descriptive knowledge of muscles in A. imperator to date. Furthermore, CLSM‐projections allow for estimation of differences in the material composition of the mouthparts' cuticle. By combining all methods, we analyze possible functions and underlying biomechanics of each mouthpart. We also analyzed the concerted movements of the mouthparts; unique behavior of the mouthparts during feeding is active participation by the labrum and distinct movement by the maxillary laciniae. We aim to elucidate the complex movements of the mouthparts and their functioning by combining detailed information on (1) in vivo movement behavior (supplemented with physiological angle approximations), (2) movement ability provided by morphology (morphological movement angles), (3) 3D‐anatomy, and (4) cuticle composition estimates. This article is protected by copyright. All rights reserved.
... The flexibility of both the stylus and basis probably serves as a shock absorber, when interacting with obstacles (see also 58,90 ), a mechanism also previously reported from other biological structures (e.g. [112][113][114]. ...
The radula, a chitinous membrane with embedded tooth rows, is the molluscan autapomorphy for feeding. The morphologies, arrangements and mechanical properties of teeth can vary between taxa, which is usually interpreted as adaptation to food. In previous studies, we proposed about trophic and other functional specialisations in taenioglossan radulae from species of African paludomid gastropods. These were based on the analysis of shape, material properties, force‑resistance, and the mechanical behaviour of teeth, when interacting with an obstacle. The latter was previously simulated for one species (Spekia zonata) by the finite‑element‑analysis (FEA) and, for more species, observed in experiments. In the here presented work we test the previous hypotheses by applying the FEA on 3D modelled radulae, with incorporated material properties, from three additional paludomid species. These species forage either on algae attached to rocks (Lavigeria grandis), covering sand (Cleopatra johnstoni), or attached to plant surface and covering sand (Bridouxia grandidieriana). Since the analysed radulae vary greatly in their general size (e.g. width) and size of teeth between species, we additionally aimed at relating the simulated stress and strain distributions with the tooth sizes by altering the force/volume. For this purpose, we also included S. zonata again in the present study. Our FEA results show that smaller radulae are more affected by stress and strain than larger ones, when each tooth is loaded with the same force. However, the results are not fully in congruence with results from the previous breaking stress experiments, indicating that besides the parameter size, more mechanisms leading to reduced stress/strain must be present in radulae.
... Beetles of the family Ptiliidae fold their wings by bending them along four oblique lines in the resilin-rich areas of the blade. Resilin gives elasticity to the wing membrane and, as can be expected, occurs in those wing areas which should bend during wing folding and quickly straighten during unfolding (Haas et al., 2000). First the beetle pulls its wings back and rotates them slightly around their axis while closing the elytra; then it bends the wings by moving its abdomen. ...
... Third, detailed investigation of the specialized morphology and material properties of the coupling mechanism may inspire the biomimetic design of insectinspired energy absorption devices. Last, in addition to the coupling, both elastin of the hind wings (Haas et al 2000) and the muscles at the roots of the elytra (Frantsevich 2010) may also influent the energy absorption, and how these functions while colliding on land can be extensively uncovered in our future project. ...
Some insects, such as bees, wasps, and bugs, have specialized coupling structures to synchronize the wing motions in flight. Some others, such as ladybirds, are equipped with coupling structures that work only at rest. By locking elytra into each other, such structures provide hindwings with a protective cover to prevent contamination. Here, we show that the coupling may play another significant role: contributing to energy absorption in falls, thereby protecting the abdomen against mechanical damage. In this combined experimental, numerical and theoretical study, we investigated free falls of ladybirds (Coccinella septempunctata), and discovered that upon collision to the ground, the coupling may fail and the elytra may unlock. This unlocking of the coupling increased the energy absorption by 33%, in comparison to when the elytra remain coupled. Using micro-CT scanning, we developed comparative models that enabled us to simulate impact scenarios numerically. Our results showed that unlocking of the coupling, here called elytra splitting, reduces both the peak impact force and rebound velocity. We fabricated the insect-inspired coupling mechanism using 3D printing and demonstrated its application as a damage preventing on system for quadcopters in accidental collisions.
