Beetle Exoskeleton Research Articles

Numerical investigation on energy absorption characteristics of impact-resistant lightweight structure of bio-mimetic micro aerial vehicle

With the increasing development of micro aerial vehicle, the impact resistance of the structure has gradually become a research hotspot. The beetle's exoskeleton not only provides important protection for its body and flexible wings but also ensures its good flight performance. Therefore, the beetle's exoskeleton has become an important research content in the field of bio-mimetic micro aerial vehicles. In this study, taking the internal structure of the beetle's exoskeleton as the research object, a kind of bio-mimetic lightweight thin-walled impact-resistant structure that can be used in the micro aerial vehicle is proposed. The energy absorption characteristics of the bio-mimetic structure under axial impact loading are calculated and analyzed by the numerical simulation method, and the important parameters such as impact angle, protective layer thickness, and wall thickness are optimized and discussed. The main results include that the filling method of the hollow column has a certain correlation with the energy absorption characteristics of the structure under different impact angles, and among the different cross-sectional shapes, the hollow column with a circular cross-sectional configuration has the best energy absorption ability. In addition, as the internal space of the structure increase exponentially, the decrease of specific energy absorption value of the bio-mimetic structure is relatively small. Finally, the wall thickness with good energy absorption characteristics is about 1.5 mm, while the preferred span length of the structure is between 90 and 120 mm.

Anti-impact performance of bionic tortoiseshell-like composites

Biological stiff-soft phase composite structures, such as tortoise shells and conch shells, have excellent anti-impact performance to withstand various types of high-stress events encountered in nature. A remarkable 3D interdigitating zigzag architecture in the tortoise shell, named occlusal suture structure, makes the tortoise shell simultaneously possess high mechanical strength and toughness, which is also an ideal stiff-soft phase composite prototype to mimic. In this study, four types of bionic anti-impact composite plates that feature the hard-soft phase architecture of tortoiseshell (Bio-T), mollusk shell (Bio-M), beetle exoskeleton (Bio-B) and nacre (Bio-N) and a homogeneous structure plate (HSP) are designed, to systematically examine and compare their roles in impact energy dissipation. Through an integrated approach combining additive manufacturing and drop tower testing, the peak force, residual velocity, and energy absorption of these bionic composite plates are studied and compared. Experimental results indicate that the Bio-T composite plate has the best impact resistance, compared with the other three bionic composites. Furthermore, the influence of the ply angle and the dimension of suture structure on the impact resistance of the Bio-T composite plate is identified. The [0°/30°/0°] arrangement is able to resist higher loads before failure. The 3D Bio-T composite plate provides a significant enhancement of anti-impact performance than the 2D Bio-T composite plate. Finally, the crack propagation mode in the suture structure of the Bio-T composite plates is examined, enhancing our understanding of the underlying mechanisms during impact. Such findings may prove useful for the design of future protective apparatus for soldiers and aircrafts, with improved anti-impact performance.

Temporal changes in the physical and mechanical properties of beetle elytra during maturation

Changes in physical properties of Tenebrio molitor and Tribolium castaneum elytra (hardened forewings) were studied to understand how the development of microstructure and chemical interactions determine cuticle mechanical properties. Analysis of these properties supports a model in which cuticular material is continuously secreted from epidermal cells to produce an extracellular matrix so that the outermost layers mature first. It is hypothesized that enzymatic crosslinking and pigmentation reactions along with dehydration help to stabilize the protein-chitin network within the initial layers of cuticle shortly after eclosion. Mature layers are proposed to bear most of the mechanical loads. The frequency dependence of the storage modulus and the tan δ values decreased during the beginning of maturation, reaching constant values after 48 h post-eclosion. A decrease of tan δ indicates an increase in crosslinking of the material. The water content declined from 75% to 31%, with a significant portion lost from within the open spaces between the dorsal and ventral cuticular layers. Dehydration had a less significant influence than protein crosslinking on the mechanical properties of the elytron during maturation. When Tribolium cuticular protein TcCP30 expression was decreased by RNAi, the tan δ and frequency dependence of E’ of the elytron did not change during maturation. This indicates that TcCP30 plays a role in the crosslinking process of the beetle's exoskeleton. This study was inspired by previous work on biomimetic multicomponent materials and helps inform future work on creating robust lightweight materials derived from natural sources. Statement of significanceExamination of changes in the physical properties of the elytra (hardened forewings) of two beetle species advanced understanding of how the molecular interactions influence the mechanical properties of the elytra. Physical characterization, including dynamic mechanical analysis, determined that the outer portion of the elytra matured first, while epidermal cells continued to secrete reactive components until the entire structure reached maturation. RNA interference was used to identify the role of a key protein in the elytra. Suppression of its expression reduced the formation of crosslinked polymeric components in the elytra. Identifying the molecular interactions in the matrix of proteins and polysaccharides in the elytra together with their hierarchical architecture provides important design concepts in the development of biomimetic materials.

Performance of 3D-Printed Bionic Conch-Like Composite Plate under Low-Velocity Impact.

Biological armors can provide an effective protection against predators. In this study, inspired by conch shell, beetle exoskeleton, and nacre, three different types of bionic composites plates were fabricated: Bio-S, Bio-B, and Bio-N, as well as an equivalent monolithic plate formed from the same stiff material designed and manufactured by additive manufacturing, respectively. Low velocity impact tests using drop tower were conducted to study their impact resistance. Experimental findings indicated that the Bio-S composite had superior impact resistance compared with the other bionic composites and the monolithic plate. Furthermore, the influence of the ply angle on the impact resistance of the Bio-S composite plate was investigated. The (0°/30°/0°/30°) arrangement was able to provide the highest impact resistance. Finally, the crack propagation mode in Bio-S composites plates was analyzed, enhancing our understanding of the underlying mechanisms during impact. Such findings may lead to the development of superior lightweight protective structures with improved anti-impact performance.

Microstructural design for mechanical–optical multifunctionality in the exoskeleton of the flower beetle Torynorrhina flammea

Biological systems have a remarkable capability of synthesizing multifunctional materials that are adapted for specific physiological and ecological needs. When exploring structure-function relationships related to multifunctionality in nature, it can be a challenging task to address performance synergies, trade-offs, and the relative importance of different functions in biological materials, which, in turn, can hinder our ability to successfully develop their synthetic bioinspired counterparts. Here, we investigate such relationships between the mechanical and optical properties in a multifunctional biological material found in the highly protective yet conspicuously colored exoskeleton of the flower beetle, Torynorrhina flammea Combining experimental, computational, and theoretical approaches, we demonstrate that a micropillar-reinforced photonic multilayer in the beetle's exoskeleton simultaneously enhances mechanical robustness and optical appearance, giving rise to optical damage tolerance. Compared with plain multilayer structures, stiffer vertical micropillars increase stiffness and elastic recovery, restrain the formation of shear bands, and enhance delamination resistance. The micropillars also scatter the reflected light at larger polar angles, enhancing the first optical diffraction order, which makes the reflected color visible from a wider range of viewing angles. The synergistic effect of the improved angular reflectivity and damage localization capability contributes to the optical damage tolerance. Our systematic structural analysis of T. flammea's different color polymorphs and parametric optical and mechanical modeling further suggest that the beetle's microarchitecture is optimized toward maximizing the first-order optical diffraction rather than its mechanical stiffness. These findings shed light on material-level design strategies utilized in biological systems for achieving multifunctionality and could thus inform bioinspired material innovations.

Earwig preying on ambrosia beetle: Evaluating predatory process and prey preference

AbstractEarwigs (Dermaptera), such as Forficula auricularia L., are important euryphagous predators for a wide variety of prey and can markedly influence the populations of orchard pests. Most previous studies on earwig feeding behaviour have not used adult beetles of the prey species; few researchers have focused on prey preference in earwigs. Some fragments of beetle exoskeleton and an earwig adult, Anisolabella marginalis (Dohrn), were found in the same cage, where adults of ambrosia beetle, Euwallacea interjectus (Blandford), were emerging from the logs of a fig tree infected with Ceratocystis canker (fig wilt disease). Thus, A. marginalis was suspected of being a predator of E. interjectus. To shed light on this issue, in the laboratory, we set up a test arena and observed and recorded behavioural interactions between A. marginalis and E. interjectus. E. interjectus was collected from the logs of fig trees and reared on an artificial diet, along with six different ambrosia beetle species, which were collected from a trap (baited with ethanol) and a fallen maple tree. A series of laboratory experiments demonstrated that A. marginalis is actually a predator of E. interjectus and other species of ambrosia beetle, indicating its a potential for use in effective pest control in the field. The predators frequently consume and tend to select their prey depending on prey size, rather than sex and beetle species. Furthermore, earwigs have alternative predatory strategies for dealing with seven different species, although they use their forceps to cut the body of most tested beetles.

Fracture modes and hybrid toughening mechanisms in oscillated/twisted plywood structure

Twisted or oscillated plywood structure can be often found in biological composites such as claws of lobsters, bone of mammals, dactyl club of mantis shrimps, and exoskeleton of beetles, which exhibits a combination of high stiffness, high fracture toughness and low density. However, there lacks a quantitative understanding of the relationship between the fracture toughness of the composite and its internal geometry. In this article, we propose that a combination of crack tilting and crack bridging determines the effective fracture toughness of the fiber-reinforced composite with the plywood structure. During the fracturing process, a crack plane initially propagates in the matrix-fiber interface following the twisted fiber alignment. Such crack tilting mechanism can significantly enlarge the area of cracking surface and thus enhance the effective fracture toughness of the composite. With the propagation of the tilted crack plane, the local energy release rate becomes too small to maintain the growth of the tilted crack plane, leading the crack to grow into the matrix, crossing the fibers. Because of the high strength of the fiber, a few fibers can maintain unbroken behind the crack tip, corresponding to crack bridging mechanism. Based on our quantitative analysis, it is found that the effective fracture toughness of the composite can be maximized for a certain pitch angle of the oscillated/twisted plywood structure, which agrees well with experiments. Statements of SignificanceFiber-reinforced composites can be widely found in nature and engineering applications. Recently, it has been discovered that many fiber-reinforced composites in biology have twisted or oscillated plywood structure with high fracture toughness, high mechanical stiffness and low density. Detailed experiments have indicated that an optimal pitch angle may exist for the plywood structure to maximize the fracture toughness of the composites. However, there lacks a quantitative model of revealing such pitch angle-dependent fracture toughness. In this work, we propose a hybrid toughening mechanism and predict the optimal pitch angle in twisted/oscillated plywood structure for maximizing the fracture toughness. Our predictions agree reasonably well with experimental results. As such, the theory may help the design of better fiber-reinforced composites.

Solid-State NMR Study of the Cicada Wing.

Wings of flying insects are part of the cuticle which forms the exoskeleton. The primary molecular components of cuticle are protein, chitin, and lipid. How these components interact with one another to form the exoskeleton is not completely understood. The difficulty in characterizing the cuticle arises because it is insoluble and noncrystalline. These properties severely limit the experimental tools that can be used for molecular characterization. Solid-state nuclear magnetic resonance experiments have been used in the past to characterize the exoskeleton of beetles and have found that chitin and protein make comparable contributions to the molecular matrix. However, little work has been done to characterize the components of the wing, which includes vein and membrane. In this work, solid-state NMR was used to characterize the wing of the 17-year cycle cicada (Magicicada cassini) that appeared in northern West Virginia during the summer of 2016. The NMR results show noticeable differences between the molecular components of the vein and membrane.

Exoskeleton may influence the internal body temperatures of Neotropical dung beetles (Col. Scarabaeinae).

The insect exoskeleton is a multifunctional coat with a continuum of mechanical and structural properties constituting the barrier between electromagnetic waves and the internal body parts. This paper examines the ability of beetle exoskeleton to regulate internal body temperature considering its thermal permeability or isolation to simulated solar irradiance and infrared radiation. Seven Neotropical species of dung beetles (Coleoptera, Scarabaeinae) differing in colour, surface sculptures, size, sexual dimorphism, period of activity, guild category and altitudinal distribution were studied. Specimens were repeatedly subjected to heating trials under simulated solar irradiance and infrared radiation using a halogen neodymium bulb light with a balanced daylight spectrum and a ceramic infrared heat emitter. The volume of exoskeleton and its weight per volume unit were significantly more important for the heating rate at the beginning of the heating process than for the asymptotic maximum temperature reached at the end of the trials: larger beetles with relatively thicker exoskeletons heated more slowly. The source of radiation greatly influences the asymptotic temperature reached, but has a negligible effect in determining the rate of heat gain by beetles: they reached higher temperatures under artificial sunlight than under infrared radiation. Interspecific differences were negligible in the heating rate but had a large magnitude effect on the asymptotic temperature, only detectable under simulated sun irradiance. The fact that sun irradiance is differentially absorbed dorsally and transformed into heat among species opens the possibility that differences in dorsal exoskeleton would facilitate the heat gain under restrictive environmental temperatures below the preferred ones. The findings provided by this study support the important role played by the exoskeleton in the heating process of beetles, a cuticle able to act passively in the thermal control of body temperature without implying energetic costs and metabolic changes.

Design principles of the beetle's exoskeleton

Design principles of the beetle's exoskeleton

Beetle Exoskeleton May Facilitate Body Heat Acting Differentially across the Electromagnetic Spectrum.

Exoskeletons of beetles and their associated morphological characteristics can serve many different functions, including thermoregulation. We study the thermal role of the exoskeleton in 13 Geotrupidae dung beetle species using heating experiments under controlled conditions. The main purpose was to measure the influence of heating sources (solar radiance vs. infrared), animal position (dorsal exposure vs. ventral exposure), species identity, and phylogenetic relationships on internal asymptotic temperatures and heating rates. The thermal response was significantly influenced by phylogenetic relatedness, although it was not affected by the apterous condition. The asymptotic internal temperature of specimens was not affected by the thoracic volume but was significantly higher under simulated sunlight conditions than under infrared radiation and when exposed dorsally as opposed to ventrally. There was thus a significant interaction between heating source and body position. Heating rate was negatively and significantly influenced by thoracic volume, and, although insignificantly slower under simulated sunlight, it was significantly affected by body position, being faster under dorsal exposure. The results constitute the first evidence supporting the hypothesis that the beetle exoskeleton acts differentially across the electromagnetic spectrum determining internal body temperatures. This interesting finding suggests the existence of a kind of passive physiology imposed by the exoskeleton and body size, where interspecific relationships play a minor role.

Tunable nano-wrinkling of chiral surfaces: Structure and diffraction optics.

Periodic surface nano-wrinkling is found throughout biological liquid crystalline materials, such as collagen films, spider silk gland ducts, exoskeleton of beetles, and flower petals. These surface ultrastructures are responsible for structural colors observed in some beetles and plants that can dynamically respond to external conditions, such as humidity and temperature. In this paper, the formation of the surface undulations is investigated through the interaction of anisotropic interfacial tension, swelling through hydration, and capillarity at free surfaces. Focusing on the cellulosic cholesteric liquid crystal (CCLC) material model, the generalized shape equation for anisotropic interfaces using the Cahn-Hoffman capillarity vector and the Rapini-Papoular anchoring energy are applied to analyze periodic nano-wrinkling in plant-based plywood free surfaces with water-induced cholesteric pitch gradients. Scaling is used to derive the explicit relations between the undulations' amplitude expressed as a function of the anchoring strength and the spatially varying pitch. The optical responses of the periodic nano-structured surfaces are studied through finite difference time domain simulations indicating that CCLC surfaces with spatially varying pitch reflect light in a wavelength higher than that of a CCLC's surface with constant pitch. This structural color change is controlled by the pitch gradient through hydration. All these findings provide a foundation to understand structural color phenomena in nature and for the design of optical sensor devices.

Feeding Effectiveness of Megaphobema mesomelas (Araneae, Theraphosidae) on Two Prey Types

Prey selection is essential for individual fitness; therefore, it would be expected that a predator would select prey of a higher rank (energy/time) when exposed to prey of differing quality. In this paper, we compare the feeding effectiveness (biomass consumed/time) of Megaphobema mesomelas (O. P.-Cambridge, 1892) in captivity, and the preference between two prey types: beetles and crickets. Spiders are more effective when feeding on crickets. The heavy exoskeleton of beetles increases prey-handling time in order to access a relatively smaller amount of edible tissue. Effectiveness also increases with spider and prey size (mass), with larger spiders feeding more effectively on larger prey. Spiders show a strong preference for feeding upon crickets over beetles when both prey types are offered at the same time.

Cuticle structure of the scarab beetle Cetonia aurata analyzed by regression analysis of Mueller-matrix ellipsometric data

Since one hundred years it is known that some scarab beetles reflect elliptically and near-circular polarized light as demonstrated by Michelson for the beetle Chrysina resplendens. The handedness of the polarization is in a majority of cases left-handed but also right-handed polarization has been found. In addition, brilliant colors with metallic shine are observed. The polarization and color effects are generated in the beetle exoskeleton, the so-called cuticle. The objective of this work is to demonstrate that structural parameters and materials optical functions of these photonic structures can be extracted by advanced modeling of spectral multi-angle Mueller-matrix data recorded from beetle cuticles. A dual-rotating compensator ellipsometer is used to record normalized Mueller-matrix data in the spectral range 400 - 800 nm at angles of incidence in the range 25-75°. Analysis of data measured on the scarab beetle Cetonia aurata are presented in detail. The model used in the analysis mimics a chiral nanostructure and is based on a twisted layered structure. Given the complexity of the nanostructure, an excellent fit between experimental and model data is achieved. The obtained model parameters are the spectral variation of the refractive indices of the cuticle layers and structural parameters of the chiral structure.

Reply to Roberts et al.: Reflectivity and pointillist structural color on land and in water

Roberts et al. (1) point out the dominance of water creatures when it comes to multilayer interference in nature, both qualitatively and in terms of quantity. This is perhaps unsurprising when we consider that 70% of the earth is covered by water and that marine life has had a considerably longer time frame than terrestrial plants in which to develop and refine optical innovations. In our comparison of the reflectivity of the Pollia fruit to that of structurally colored animals, we focused only on terrestrial species. Although these are most conspicuous to us, Roberts et al. are correct in pointing out the dominance of fish and squid. An accurate revision of our text (2) would state: “The bright blue coloration of this fruit is more intense than that of many previously described biological materials” and “this is the highest reported reflectivity of any land-based biological organism including beetle exoskeleton, bird feathers, and the famously intense blue of Morpho butterfly scales.”

Exoskeletal Microstructure of the Bark Beetle Ips acuminatus (Coleoptera: Curculionidae: Scolytinae)

Bark beetles are minute beetles found in forest and can make galleries directly into the heartwood of trees. As a part of research for controlling this pest insect, here we demonstrate the fine structural aspects of their external bodies of the bark beetle Ips acuminatus using the field emission scanning electron microscopy (FESEM). This beetle has well developed sensory system respond to both of visual and chemical stimuli. The mouthparts which consisted of a labrum, a pair of mandibles, a pair of maxillae and the labium are very effective devices for penetration and boring holes. The thorax contains a particularly hard exoskeleton and hard elytra including the powerful muscles that operate both the wings and legs. The beetle's exoskeleton is made up of numerous plates called sclerites, separated by thin sutures. This structure can provide strong armor of this beetle while maintaining flexibility.

External exoskeletal cavities in Coleoptera and their possible mycangial functions

AbstractThis paper reviews the occurrence of external exoskeletal cavities in beetles and provides critical reassessment of their possible mycangial function. In most reported cases, the decision to attribute mycangial function to exoskeletal cavities was based solely on the combination of two factors: (i) observation of these cavities on beetle's body; and (ii) knowledge that this particular beetle species uses fungi as a food source. Such reasoning resulted in the assumption, occasionally premature and not supported by other evidence, that exoskeletal pits in the following families may function as mycangia: Rhysodidae, Ptiliidae, Staphylinidae, Hybosoridae, Scarabaeidae, Derodontidae, Ptinidae, Jacobsoniidae, Boganiidae, Cryptophagidae, Endomychidae, Erotylidae, Latridiidae, Nitidulidae, Phloeostichidae, Silvanidae, Sphindidae, Pyrochroidae, Anthribidae, Attelabidae and Curculionidae. We conclude that only two beetle families include species with adequately documented cases of external exoskeletal mycangia: (i) Curculionidae (some Scolytinae and Platypodinae); and (ii) the structurally complex mycangia of Attelabidae (Euops females). One or more species of Sphindidae, Erotylidae, Silvanidae, and Latridiidae have likely functional mycangia. Exoskeletal pits with uncertain function are additionally reported from the following families: Cupedidae, Ommatidae, Lepiceridae, Carabidae, Histeridae, Hydraenidae, Leiodidae, Elmidae, Artematopodidae, Throscidae, Elateridae, Rhinorhipidae, Biphyllidae, Cerylonidae, Cyclaxyridae, Monotomidae, Mycetophagidae and Zopheridae. We also discuss the transport of fungal spores and conidia by waxy exudates or debris build up on beetle exoskeleton, as well as their passive attachment to the body.

Stable hydrogen-isotope ratios in beetle chitin: preliminary European data and re-interpretation of North American data

Beetle exoskeletons contain chitin, a poly amino-sugar that is biosynthesized incorporating hydrogen isotopes from diet and water. As the stable isotope ratios D/H (or 2H/ 1H, expressed as δ D values) of precipitation and diet are jointly influenced by climate, the biochemically recorded hydrogen-isotope ratio in fossil beetle exoskeleton has the potential to be used for paleoclimatic reconstruction. New δ D data from modern beetles are presented as a preliminary database for Europe, with a re-evaluation of earlier North American data. We present correlated matrices of δ D values in modern beetle chitin and modern precipitation to demonstrate the concept. We review the pertinent literature to highlight the history, utility, and likely future research directions for the use of chitin's stable isotopes in entomological paleoclimatology.

Competitive Interactions among Symbiotic Fungi of the Southern Pine Beetle

The southern pine beetle, a damaging pest of conifers, is intimately linked to three symbiotic fungi. Two fungi, Ceratocystiopsis ranaculosus and Entomocorticium sp. A, are transported within specialized structures (mycangia) in the beetle exoskeleton and are mutualists of the beetle. A third fungus, Ophiostoma minus, is transported externally on the beetle exoskeleton (phoretically) and is an antagonist of the beetle. This study examined competitive interactions among these three fungi. The results of de Wit replacement series and primary and secondary resource capture assays with these fungi provide strong evidence for differential competition between the phoretic and mycangial fungi. O. minus was the most able to capture both uncolonized and colonized resources. Entomocorticium sp. A and C. ranaculosus, although equal to one another in competitive abilities, differed in their ability to compete with O. minus. Entomocorticium sp. A was able to maintain space free of O. minus to a much greater degree than was C. ranaculosus. The outcome of such competitive interactions may have significant impacts on the biology of this ecologically and economically important beetle.