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Spider webs that serve as snares are one of the most fascinating and abundant type of animal architectures. In many cases they include an adhesive coating of silk lines-so-called viscid silk-for prey capture. The evolutionary switch from silk secretions forming solid fibres to soft aqueous adhesives remains an open question in the understanding of spider silk evolution. Here we functionally and chemically characterized the secretions of two types of silk glands and their behavioural use in the cellar spider, Pholcus phalangioides. Both being derived from the same ancestral gland type that produces fibres with a solidifying glue coat, the two types produce respectively a quickly solidifying glue applied in thread anchorages and prey wraps, or a permanently tacky glue deployed in snares. We found that the latter is characterized by a high concentration of organic salts and reduced spidroin content, showing up a possible pathway for the evolution of viscid properties by hygroscopic-salt-mediated hydration of solidifying adhesives. Understanding the underlying molecular basis for such radical switches in material properties not only helps to better understand the evolutionary origins and versatility of ecologically impactful spider web architectures, but also informs the bioengineering of spider silk-based products with tailored properties.

Hair-like attachment structures are frequently used by animals to create stable contact with rough surfaces. Previous studies focused primarily on axisymmetric biomimetic models of artificial spatulas, such as those with a mushroom-shaped and cylinder-shaped geometry, in order to simulate the so-called gecko effect. Here, two geometric prototypes of artificial adhesive structures with non-axisymmetric properties were designed. The investigation of the prototype's interactions with rough surfaces was carried out using the finite element software ABAQUS. Under increasing vertical displacement, the effect of asperity size on the contact pressure evolution of the spatula was investigated. It has been demonstrated that the contact behaviour is greatly affected by the flexibility of the spatula, which is caused by its variable thickness. The thinner spatula shows a higher nominal contact area and attaches more strongly to various rough surfaces. Although a thicker spatula is more susceptible to the 'leverage' phenomenon, which occurs when excessively applied displacements prematurely reduce the nominal contact area, it obtains the ability to regulate attachment during unidirectional loading. Two non-axisymmetric prototypes provide different design concepts for the artificial adhesives. It is hoped that this study will provide fresh viewpoints and innovations that contribute to the development of biologically inspired adhesives., Competing Interests: We declare we have no competing interests., (© 2024 The Author(s).)

Cimicidae are well-known for traumatic insemination, and males pierce females with their parameres and transfer sperm through them. The shape of parameres is relatively stable in the family, but in some genera, the paramere is elongated, appearing less resistant against lateral deflection. To understand the mechanical limitations of the paramere, we studied its penetration mechanics of the common bed bug, Cimex lectularius . We examined the post-abdominal morphology, paramere geometry and material properties and conducted breaking stress experiments on the paramere under wet and dry conditions. Mechanical property gradients are present with the paramere tip as the stiffest region and the base as the most flexible one. These mechanical properties relate to the presence of Ca, Zn and Si. The basal wing-shaped structure is flexible, enabling it to interlock with the anal region during mating. The paramere is slightly twisted; the tip region is circular in cross-section, and the geometry of the rest is rather complex. In the mechanical tests, wet parameres mainly buckled, while dried parameres broke off. The level of structural failures depended on directions from which the compression forces were applied. Structural, material and mechanical strengthening mechanisms preventing the paramere from mechanical failure are discussed.

Radular teeth have to cope with wear, when interacting with ingesta. In some molluscan taxa, wear-coping mechanisms, related to the incorporation of high contents of iron or silica, have been previously determined. For most species, particularly for those which possess radulae without such incorporations, wear-coping mechanisms are understudied. In the present study, we documented and characterized the wear on radular teeth in the model species Loligo vulgaris (Cephalopoda). By applying a range of methods, the elementary composition and mechanical properties of the teeth were described, to gain insight into mechanisms for coping with abrasion. It was found that the tooth regions that are prone to wear are harder and stiffer. Additionally, the surfaces interacting with the ingesta possessed a thin coating with high contents of silicon, probably reducing abrasion. The here presented data may serve as an example of systematic study of radular wear, in order to understand the relationship between the structure of radular teeth and their properties., Competing Interests: We declare we have no competing interests., (© 2024 The Authors.)

Leafcutter ant colonies are divided into castes with the individuals performing different tasks, based mostly on size. With the mandibles, the small minims care for the brood or the fungus, whereas the larger minors and mediae cut and transport plant material, with the ant size positively related to the material size. The mechanical properties and composition of the mandible cuticle have been previously tested in the soldiers as the largest caste, revealing that the cutting edges contained high contents of the cross-linking transition metal zinc (Zn). With regard to the smaller castes, no data are present. To study how the mandible size and function relates to its mechanical properties, we here tested the mandibles of minims, minors and mediae by nanoindentation. We found that the hardness (H) and Young's modulus (E) values increased with increasing ant size and that the mandible cutting edges in each caste have the highest H- and E-values. To gain insight into the origins of these properties, we characterized the elemental composition by energy-dispersive X-ray analysis, revealing that minors and mediae possessed higher content of Zn in the cutting edges in contrast to the minims containing significantly less Zn. This shows, that Zn content relates to higher mechanical property values. Additionally, it shows that all of these parameters can differ within a single species., Competing Interests: We declare we have no competing interests., (© 2024 The Authors.)

Mandible morphology has an essential role in biting performance, but the mandible cuticle can have regional differences in its mechanical properties. The effects of such a heterogeneous distribution of cuticle material properties in the mandible responses to biting loading are still poorly explored in chewing insects. Here, we tested the mechanical properties of mandibles of the ant species Formica cunicularia by nanoindentation and investigated the effects of the cuticular variation in Young's modulus (E) under bite loading with finite-element analysis (FEA). The masticatory margin of the mandible, which interacts with the food, was the hardest and stiffest region. To unravel the origins of the mechanical property gradients, we characterized the elemental composition by energy-dispersive X-ray spectroscopy. The masticatory margin possessed high proportions of Cu and Zn. When incorporated into the FEA, variation in E effectively changed mandible stress patterns, leading to a relatively higher concentration of stresses in the stiffer mandibular regions and leaving the softer mandible blade with relatively lower stress. Our results demonstrated the relevance of cuticle E heterogeneity in mandibles under bite loading, suggesting that the accumulation of transition metals such as Cu and Zn has a relevant correlation with the mechanical characteristics in F . cunicularia mandibles., Competing Interests: We declare we have no competing interests., (© 2024 The Authors.)

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., Competing Interests: We declare we have no competing interests., (© 2024 The Author(s).)

Enhanced attachment ability is common in plants on islands to avoid potential fatal passive dispersal. However, whether island insects also have increased attachment ability remains unclear. Here we measured the attachment of a flightless weevil, Pachyrhynchus sarcitis kotoensis , from tropical islands, and compared it with documented arthropods from the mainland. We examined the morphology and material gradient of its attachment devices to identify the specific adaptive modifications for attachment. We find that the weevil has much stronger attachment force and higher safety factor than previously studied arthropods, regardless of body size and substrate roughness. This probably results from the specific flexible bases of the adhesive setae on the third footpad of the legs. This softer material on the setal base has not been reported hitherto and we suggest that it acts as a flexible hinge to form intimate contact to substrate more effectively. By contrast, no morphological difference in tarsomeres and setae between the weevil and other beetles is observed. Our results show the remarkably strong attachment of an island insect and highlights the potential adaptive benefits of strong attachment in windy island environment. The unique soft bases of the adhesive hairs may inspire the development of strong biomimetic adhesives.

Green dock beetles Gastrophysa viridula exhibit sexual dimorphism in tarsal attachment setae: females have only pointed, lanceolate and spatula-like setae, while males additionally possess discoidal ones. The sexual dimorphism is probably attributed to the necessity of male discoidal setae to adhere to the smooth back of the female during copulation. We aimed to understand its possible mechanism of attachment with G. viridula . Pull-off forces of both females and males were measured on (i) alive females, (ii) dead and dried females, and (iii) resin replicas of fresh females. The attachment ability tended to increase on dead and replicated female surfaces in both sexes, which indicates that the epicuticular grease layer on the integument of alive intact beetles decreases the attachment. This tendency was prominent in females. The present study clearly showed that in G. viridula discoidal setae enable the males to adhere stronger to female surfaces. The divergent performance found between the sexes differing in their setal composition is probably caused by the stiffness difference between the setae types and by the specific shape of the setal tips. A peculiar reproductive biology in G. viridula is probably attributed to this remarkable divergence of labour in their attachment pads between the sexes.

Molluscs forage with their radula, a chitinous membrane with teeth. Adaptations to hard or abrasive ingesta were well studied in Polyplacophora and Patellogastropoda, but for other taxa there are large gaps in knowledge. Here, we investigated the nudibranch gastropods Felimare picta and Doris pseudoargus , both of which feed on Porifera. Tooth morphologies were documented by scanning electron microscopy, and mechanical properties were tested by nanoindentation. We found that these parameters are rather similar in both species, indicating that teeth are similar in their function. To study the composition, teeth were visualized using confocal laser scanning microscopy (CLSM), to determine the degree of tanning, and analysed with energy-dispersive X-ray spectroscopy, to test the elemental composition. The emitted autofluorescence signal and the inorganic content differed between the species. This was especially prominent when studying the inner and outer tooth surfaces (leading and trailing edges). In F. picta , we detected high proportions of Si, whereas teeth of D. pseudoargus contained high amounts of Ca, which influenced the autofluorescence signal in CLSM. Employing nanoindentation, we determined high Young's modulus and hardness values for the leading edges of teeth, which relate to the Si and Ca content. This highlights that teeth with a similar morphology and mechanical properties can be mechanically enhanced via different chemical pathways in Nudibranchia.

The radula is the structure used for food processing in Mollusca. It can consist of a membrane with stiffer teeth, which is, together with alary processus, muscles and odontophoral cartilages, part of the buccal mass. In malacology, it is common practice to infer potential tooth functions from morphology. Thus, past approaches to explain functional principles are mainly hypothesis driven. Therefore, there is an urgent need for a workflow testing hypotheses on the function of teeth and buccal mass components and interaction of structures, which can contribute to understanding the structure as a whole. Here, in a non-conventional approach, we introduce a physical and dynamic radular model, based on morphological data of Spekia zonata (Gastropoda, Paludomidae). Structures were documented, computer-modelled, three-dimensional-printed and assembled to gather a simplistic but realistic physical and dynamic radular model. Such a bioinspired design enabled studying of radular kinematics and interaction of parts when underlain supporting structures were manipulated in a similar manner as could result from muscle contractions. The presented work is a first step to provide a constructional manual, paving the way for even more realistic physical radular models, which could be used for understanding radular functional morphology and for the development of novel gripping devices.

Sandy pitfall traps of antlions are elaborate constructions to capture prey. Antlions exploit the interactions between the particles in their habitat and build a stable trap. This trap is close to the unstable state; prey items will slide towards the centre-where the antlion ambushes-when entering the trap. This is efficient but requires permanent maintenance. According to the present knowledge, antlions throw sand, mainly to cause sandslides towards the centre of the pit. We hypothesized that: (i) sand-throwing causes sandslides towards the centre of the pit and (ii) sand-throwing constantly maintains the pitfall trap and thus keeps its efficiency high. Using laboratory experiments, as well as finite-element analysis, we tested these hypotheses. We show, experimentally and numerically, that sand that accumulates at the centre of the pit will be removed continuously by sand-throwing, this maintenance is leading to slope condition close to an unstable state. This keeps the slope angle steep and the efficiency of the trap constant. Furthermore, the resulting sandslides can relocate the trapped prey towards the centre of the pit. This study adds further insights from specific mechanical properties of a granular medium into the behavioural context of hunting antlion larvae.

Successful insect pollination can be achieved by a sequence of numerous attachment and detachment events at various biological surfaces. However, the quantitative measurements of pollen adhesion on biological surfaces have been poorly studied so far. We performed atomic force microscopy adhesion measurements of pollen on two most important floral parts for Asteraceae in a course of pollination: the stigma and style of Hypochaeris radicata plant. The results indicated distinct adhesive properties of them-the pollen adhesion on stigmatic surfaces drastically increased over prolonged contact time, while the pollen adhesion increase on stylar surfaces was rather restrained. Based on the observation with cryo-scanning electron microscopy, we explained the experimental results by the presence of morphological features in form of flexible stigmatic papillae that may play a crucial role in enhancing both capillary attraction and van der Waals forces. The distinct adhesive properties seemingly originate from the unique adhesive tasks that each of the floral parts requires to achieve successful pollination. The insights into the adhesive interaction between pollen and the floral parts, obtained in the present study, may lead to better understanding of pollination mechanisms, which are strongly related to our food production. Additionally, the novel pollen adhesive mechanisms learned from the stigma of the studied Asteraceae plant can inspire biomimetic designs of spontaneous gripping systems.

The antennae of mosquitoes are model systems for acoustic sensation, in that they obey general principles for sound detection, using both active feedback mechanisms and passive structural adaptations. However, the biomechanical aspect of the antennal structure is much less understood than the mechano-electrical transduction. Using confocal laser scanning microscopy, we measured the fluorescent properties of the antennae of two species of mosquito- Toxorhynchites brevipalpis and Anopheles arabiensis-and, noting that fluorescence is correlated with material stiffness, we found that the structure of the antenna is not a simple beam of homogeneous material, but is in fact a rather more complex structure with spatially distributed discrete changes in material properties. These present as bands or rings of different material in each subunit of the antenna, which repeat along its length. While these structures may simply be required for structural robustness of the antennae, we found that in FEM simulation, these banded structures can strongly affect the resonant frequencies of cantilever-beam systems, and therefore taken together our results suggest that modulating the material properties along the length of the antenna could constitute an additional mechanism for resonant tuning in these species.

When sitting and walking, the feet of wandering spiders reversibly attach to many surfaces without the use of gluey secretions. Responsible for the spiders' dry adhesion are the hairy attachment pads that are built of specially shaped cuticular hairs (setae) equipped with approximately 1 µm wide and 20 nm thick plate-like contact elements (spatulae) facing the substrate. Using synchrotron-based scanning nanofocus X-ray diffraction methods, combining wide-angle X-ray diffraction/scattering and small-angle X-ray scattering, allowed substantial quantitative information to be gained about the structure and materials of these fibrous adhesive structures with 200 nm resolution. The fibre diffraction patterns showed the crystalline chitin chains oriented along the long axis of the attachment setae and increased intensity of the chitin signal dorsally within the seta shaft. The small-angle scattering signals clearly indicated an angular shift by approximately 80° of the microtrich structures that branch off the bulk hair shaft and end as the adhesive contact elements in the tip region of the seta. The results reveal the specific structural arrangement and distribution of the chitin fibres within the attachment hair's cuticle preventing material failure by tensile reinforcement and proper distribution of stresses that arise upon attachment and detachment.

According to literature data, porous substrates can cause a reduction of insect attachment ability. We carried out traction experiments with adult ladybird beetles Harmonia axyridis on the smooth solid glass sample and rough porous Al 2 O 3 membrane to prove the primary effect of absorption of the insect pad secretion by porous media, rather than surface roughness, on the attachment force on the porous sample. With each insect individual, a set of five experiments was conducted: (1) on glass; (2) on the porous membrane; (3-5) on glass immediately after the test on the porous surface, then after 30 min and 1 h of recovery time. On the porous substrate, the forces, being similar in females and males, were greatly reduced compared to those measured on glass. A significant difference between the force values obtained in the first (before the test on the porous sample) and second (immediately after the experiment on the porous sample) tests on glass was observed. After 30 min recovery time, beetles completely regained their attachment ability. Females produced significantly lower forces than males in all experiments on glass: the differences are probably caused by the sexual dimorphism in the microstructure of their adhesive pads. The obtained results are of fundamental importance for further application in biomimetics of novel insect-repelling surfaces and in plant protection by using porous materials.

Springtails (Collembola) are known to exhibit complex hierarchical nanostructures of their exoskeleton surface that repels water and other fluids with remarkable efficiency. These nanostructures were previously widely studied due to their structure, chemistry and fluid-repelling properties. These ultrastructural and chemical studies revealed the involvement of different components in different parts of the nanopattern, but the overall process of self-assembly into the complex rather regular structures observed remains unclear. Here, we model this process from a theoretical point of view partially using solutions related to the so-called Tammes problem. By using densities of three different reacting substances, we obtained a typical morphology that is highly similar to the ones observed on the cuticle of some springtail species. These results are important not only for our understanding of the formation of hierarchical nanoscale structures in nature, but also for the fabrication of novel surface coatings., (© 2018 The Author(s).)

Nature has evolved structures with high load-carrying capacity and long-term durability. The principles underlying the functionality of such structures, if studied systematically, can inspire the design of more efficient engineering systems. An important step in this process is to characterize the material properties of the structure under investigation. However, direct mechanical measurements on small complex-shaped biological samples involve numerous technical challenges. To overcome these challenges, we developed a method for estimation of the elastic modulus of insect cuticle, the second most abundant biological composite in nature, through simple light microscopy. In brief, we established a quantitative link between the autofluorescence of different constituent materials of insect cuticle, and the resulting mechanical properties. This approach was verified using data on cuticular structures of three different insect species. The method presented in this study allows three-dimensional visualisation of the elastic modulus, which is impossible with any other available technique. This is especially important for precise finite-element modelling of cuticle, which is known to have spatially graded properties. Considering the simplicity, ease of implementation and high-resolution of the results, our method is a crucial step towards a better understanding of material-function relationships in insect cuticle, and can potentially be adapted for other graded biological materials., (© 2018 The Author(s).)

Surfaces with re-entrant topographies can repel liquids even of extremely low surface tension, almost independently of the material's inherent wettability. We show that this topography-based wetting resistance can also be applied to underwater applications, reducing the permanent adhesion of marine hardfouling organisms. Having combined a biofouling assay in the marine environment with microscopic analyses, we demonstrate how a synergistic effect of a soft silicone-based material with a re-entrant mushroom-shaped surface topography strongly increases the fouling release ability of such coatings compared with a smooth control made from the same material. Our coating inhibited the complete wetting of the solidifying glues produced by marine organisms, which resulted in a decreased contact area and, consequently, low adhesion. Our work suggests that topography-induced wetting resistance of surfaces may become a viable approach in preventing permanent adhesion of marine hardfouling organisms., (© 2018 The Author(s).)

Active tactile exploration behaviour is constrained to a large extent by the morphological and biomechanical properties of the animal's somatosensory system. In the model organism Carausius morosus , the main tactile sensory organs are long, thin, seemingly delicate, but very robust antennae. Previous studies have shown that these antennae are compliant under contact, yet stiff enough to maintain a straight shape during active exploration. Overcritical damping of the flagellum, on the other hand, allows for a rapid return to the straight shape after release of contact. Which roles do the morphological and biomechanical adaptations of the flagellum play in determining these special mechanical properties? To investigate this question, we used a combination of biomechanical experiments and numerical modelling. A set of four finite-element (FE) model variants was derived to investigate the effect of the distinct geometrical and material properties of the flagellum on its static (bending) and dynamic (damping) characteristics. The results of our numerical simulations show that the tapered shape of the flagellum had the strongest influence on its static biomechanical behaviour. The annulated structure and thickness gradient affected the deformability of the flagellum to a lesser degree. The inner endocuticle layer of the flagellum was confirmed to be essential for explaining the strongly damped return behaviour of the antenna. By highlighting the significance of two out of the four main structural features of the insect flagellum, our study provides a basis for mechanical design of biomimetic touch sensors tuned to become maximally flexible while quickly resuming a straight shape after contact., (© 2018 The Author(s).)

In its evolution, the diverse group of stick and leaf insects (Phasmatodea) has undergone a rapid radiation. These insects evolved specialized structures to adhere to different surfaces typical for their specific ecological environments. The cuticle of their tarsal attachment pads (euplantulae) is known to possess a high diversity of attachment microstructures (AMS) which are suggested to reflect ecological specializations of different groups within phasmids. However, the origin of these microstructures and their developmental background remain largely unknown. Here, based on the detailed scanning electron microscopy study of pad surfaces, we present a theoretical approach to mathematically model an outstanding diversity of phasmid AMS using the reaction-diffusion model by Alan Turing. In general, this model explains pattern formation in nature. For the first time, we were able to identify eight principal patterns and simulate the transitions among these. In addition, intermediate transitional patterns were predicted by the model. The ease of transformation suggests a high adaptability of the microstructures that might explain the rapid evolution of pad characters. We additionally discuss the functional morphology of the different microstructures and their assumed advantages in the context of the ecological background of species., (© 2018 The Author(s).)

Friction anisotropy is an important property of many surfaces that usually facilitate the generation of motion in a preferred direction. Such surfaces are very common in biological systems and have been the templates for various bio-inspired materials with similar tribological properties. So far friction anisotropy is considered to be the result of an asymmetric arrangement of surface nano- and microstructures. However, here we show by using bio-inspired sawtooth-structured surfaces that the anisotropic friction properties are not only controlled by an asymmetric surface topography, but also by the ratio of the sample-substrate stiffness, the aspect ratio of surface structures, and by the substrate roughness. Systematically modifying these parameters, we were able to demonstrate a broad range of friction anisotropies, and for specific sample-substrate combinations even an inversion of the anisotropy. This result highlights the complex interrelation between the different material and topographical parameters on friction properties and sheds new light on the conventional design paradigm of tribological systems. Finally, this result is also of great importance for understanding functional principles of biological materials and surfaces, as such inversion of friction anisotropy may correlate with gait pattern and walking behaviour in climbing animals, which in turn may be used in robotic applications., (© 2018 The Author(s).)

Insect cuticle is a biological composite with a high degree of complexity in terms of both architecture and material composition. Given the complex morphology of many insect body parts, finite-element (FE) models play an important role in the analysis and interpretation of biomechanical measurements, taken by either macroscopic or nanoscopic techniques. Many previous studies show that the interpretation of nanoindentation measurements of this layered composite material is very challenging. To develop accurate FE models, it is of particular interest to understand more about the variations in the stiffness through the thickness of the cuticle. Considering the difficulties of making direct measurements, in this study, we use the FE method to analyse previously published data and address this issue numerically. For this purpose, sets of continuous or discontinuous stiffness profiles through the thickness of the cuticle were mathematically described. The obtained profiles were assigned to models developed based on the cuticle of three insect species with different geometries and layer configurations. The models were then used to simulate the mechanical behaviour of insect cuticles subjected to nanoindentation experiments. Our results show that FE models with discontinuous exponential stiffness gradients along their thickness were able to predict the stress and deformation states in insect cuticle very well. Our results further suggest that, for more accurate measurements and interpretation of nanoindentation test data, the ratio of the indentation depth to cuticle thickness should be limited to 7% rather than the traditional '10% rule'. The results of this study thus might be useful to provide a deeper insight into the biomechanical consequences of the distinct material distribution in insect cuticle and also to form a basis for more realistic modelling of this complex natural composite., (© 2017 The Author(s).)

Silks play an important role in the life of various arthropods. A highly neglected prerequisite to make versatile use of silks is sufficient attachment to substrates. Although there have been some studies on the structure and mechanics of silk anchorages of spiders, for insects only anecdotal reports on attachment-associated spinning behaviour exist. Here, we experimentally studied the silk attachment of the pupae and last instar caterpillars of the tea bagworm Eumeta minuscula (Butler 1881) (Lepidoptera, Psychidae) to the leaves of its host plant Ilex chinensis We found that the bagworms spin attachment discs, which share some structural features with those of spiders, like a plaque consisting of numerous overlaid, looped glue-coated silk fibres and the medially attaching suspension thread. Although the glue, which coats the fibres, cannot spread and adhere very well to the leaf surface, high pull-off forces were measured, yielding a mean safety factor (force divided by the animal weight) of 385.6. Presumably, the bagworms achieve this by removal of the leaf epidermis prior to silk attachment, which exposes the underlying tissue that represents a much better bonding site. This ensures a reliable attachment during the immobile, vulnerable pupal stage. This is the first study on the biomechanics and structure of silk attachments to substrates in insects., (© 2017 The Author(s).)

During slithering locomotion the ventral scales at a snake's belly are in direct mechanical interaction with the environment, while the dorsal scales provide optical camouflage and thermoregulation. Recent work has demonstrated that compared to dorsal scales, ventral scales provide improved lubrication and wear protection. While biomechanic adaption of snake motion is of growing interest in the fields of material science and robotics, the mechanism for how ventral scales influence the friction between the snake and substrate, at the molecular level, is unknown. In this study, we characterize the outermost surface of snake scales using sum frequency generation (SFG) spectra and near-edge X-ray absorption fine structure (NEXAFS) images collected from recently shed California kingsnake (Lampropeltis californiae) epidermis. SFG's nonlinear optical selection rules provide information about the outermost surface of materials; NEXAFS takes advantage of the shallow escape depth of the electrons to probe the molecular structure of surfaces. Our analysis of the data revealed the existence of a previously unknown lipid coating on both the ventral and dorsal scales. Additionally, the molecular structure of this lipid coating closely aligns to the biological function: lipids on ventral scales form a highly ordered layer which provides both lubrication and wear protection at the snake's ventral surface., (© 2015 The Author(s).)

Previous research demonstrated that arrays of vertically aligned carbon nanotubes (VACNTs) exhibit strong frictional properties. Experiments indicated a strong decrease of the friction coefficient from the first to the second sliding cycle in repetitive measurements on the same VACNT spot, but stable values in consecutive cycles. VACNTs form clusters under shear applied during friction tests, and self-organization stabilizes the mechanical properties of the arrays. With increasing load in the range between 300 µN and 4 mN applied normally to the array surface during friction tests the size of the clusters increases, while the coefficient of friction decreases. To better understand the experimentally obtained results, we formulated and numerically studied a minimalistic model, which reproduces the main features of the system with a minimum of adjustable parameters. We calculate the van der Waals forces between the spherical friction probe and bunches of the arrays using the well-known Morse potential function to predict the number of clusters, their size, instantaneous and mean friction forces and the behaviour of the VACNTs during consecutive sliding cycles and at different normal loads. The data obtained by the model calculations coincide very well with the experimental data and can help in adapting VACNT arrays for biomimetic applications.

Intergenomic evolutionary conflicts increase biological diversity. In sexual conflict, female defence against males is generally assumed to be resistance, which, however, often leads to trait exaggeration but not diversification. Here, we address whether tolerance, a female defence mechanism known from interspecific conflicts, exists in sexual conflict. We examined the traumatic insemination of female bed bugs via cuticle penetration by males, a textbook example of sexual conflict. Confocal laser scanning microscopy revealed large proportions of the soft and elastic protein resilin in the cuticle of the spermalege, the female defence organ. Reduced tissue damage and haemolymph loss were identified as adaptive female benefits from resilin. These did not arise from resistance because microindentation showed that the penetration force necessary to breach the cuticle was significantly lower at the resilin-rich spermalege than at other cuticle sites. Furthermore, a male survival analysis indicated that the spermalege did not impose antagonistic selection on males. Our findings suggest that the specific spermalege material composition evolved to tolerate the traumatic cuticle penetration. They demonstrate the importance of tolerance in sexual conflict and genitalia evolution, extend fundamental coevolution and speciation models and contribute to explaining the evolution of complexity. We propose that tolerance can drive trait diversity., (© 2015 The Author(s) Published by the Royal Society. All rights reserved.)

While adult green dock leaf beetles Gastrophysa viridula use tarsal adhesive setae to attach to and walk on smooth vertical surfaces and ceilings, larvae apply different devices for similar purposes: pretarsal adhesive pads on thoracic legs and a retractable pygopod at the 10th abdominal segment. Both are soft smooth structures and capable of wet adhesion. We studied attachment ability of different larval instars, considering the relationship between body weight and real contact area between attachment devices and the substrate. Larval gait patterns were analysed using high-speed video recordings. Instead of the tripod gait of adults, larvae walked by swinging contralateral legs simultaneously while adhering by the pygopod. Attachment ability of larval instars was measured by centrifugation on a spinning drum, revealing that attachment force decreases relative to weight. Contributions of different attachment devices to total attachment ability were investigated by selective disabling of organs by covering them with melted wax. Despite their smaller overall contact area, tarsal pads contributed to a larger extent to total attachment ability, probably because of their distributed spacing. Furthermore, we observed different behaviour in adults and larvae when centrifuged: while adults gradually slipped outward on the centrifuge drum surface, larvae stayed at the initial position until sudden detachment.

The stalked barnacle Dosima fascicularis secretes foam-like cement, the amount of which usually exceeds that produced by other barnacles. When Dosima settles on small objects, this adhesive is additionally used as a float which gives buoyancy to the animal. The dual use of the cement by D. fascicularis requires mechanical properties different from those of other barnacle species. In the float, two regions with different morphological structure and mechanical properties can be distinguished. The outer compact zone with small gas-filled bubbles (cells) is harder than the interior one and forms a protective rind presumably against mechanical damage. The inner region with large, gas-filled cells is soft. This study demonstrates that D. fascicularis cement is soft and visco-elastic. We show that the values of the elastic modulus, hardness and tensile stress are considerably lower than in the rigid cement of other barnacles.

The morphology of the toe epithelium of the rock frog, Staurois parvus (Family Ranidae), was investigated using a variety of microscopical techniques. The toe pad epithelium is stratified (four to five cell layers), the apical parts of the cells of the outermost layer being separated by fluid-filled channels. The surface of these cells is covered by a dense array of nanopillars, which also cover the surface of subarticular tubercles and unspecialized ventral epithelium of the toes, but not the dorsal epithelium. The apical portions of the outer two layers contain fibrils that originate from the nanopillars and are oriented approximately normal to the surface. This structure is similar to the pad structure of tree frogs of the families Hylidae and Rhacophoridae, indicating evolutionary convergence and a common evolutionary design for reversible attachment in climbing frogs. The main adaptation to the torrent habitat seems to be the straightness of the channels crossing the toe pad, which will assist in drainage of excess water. The presence of nanopillar arrays on all ventral surfaces of the toes resembles that on clingfish suckers and may be a specific adaptation for underwater adhesion and friction. The relevance of these findings to the development of new biomimetically inspired reversible adhesives is discussed.

One of the important problems appearing in experimental realizations of artificial adhesives inspired by gecko foot hair is so-called clusterization. If an artificially produced structure is flexible enough to allow efficient contact with natural rough surfaces, after a few attachment-detachment cycles, the fibres of the structure tend to adhere one to another and form clusters. Normally, such clusters are much larger than original fibres and, because they are less flexible, form much worse adhesive contacts especially with the rough surfaces. Main problem here is that the forces responsible for the clusterization are the same intermolecular forces which attract fibres to fractal surface of the substrate. However, arrays of real gecko setae are much less susceptible to this problem. One of the possible reasons for this is that ends of the seta have more sophisticated non-uniformly distributed three-dimensional structure than that of existing artificial systems. In this paper, we simulated three-dimensional spatial geometry of non-uniformly distributed branches of nanofibres of the setal tip numerically, studied its attachment-detachment dynamics and discussed its advantages versus uniformly distributed geometry.

Dosima fascicularis is the only barnacle which can drift autonomously at the water surface with a foam-like cement float. The cement secreted by the animal contains numerous gas-filled cells of different size. When several individuals share one float, their size and not their number is crucial for the production of both volume and mass of the float. The gas content within the cells of the foam gives positive static buoyancy to the whole float. The volume of the float, the gas volume and the positive static buoyancy are positively correlated. The density of the cement float without gas is greater than that of seawater. This study shows that the secreted cement consists of more than 90% water and the gas volume is on average 18.5%. Our experiments demonstrate that the intact foam-like cement float is sealed to the surrounding water.

Microstructures responsible for temporary arresting of contacting surfaces are widely distributed on surfaces in different organisms. Recent morphological studies show that these structures have different density of outgrowths and not ideal distribution pattern on both complementary parts of the contact. One can suggest that this difference is optimized by natural selection to get stronger mechanical arrest within the system. In this paper, we simulate such a system numerically, both in the frames of continuous contact and discrete dynamical models to prove this hypothesis and elucidate other aspects of optimization of such mechanical adhesive systems., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

Wet adhesive systems of insects strongly rely for their function on the formation of capillary bridges with the substrate. Studies on the chemical composition and evaporation dynamics of tarsal secretions strongly suggest a difference in chemistry of secretion in beetles and flies, both possessing hairy attachment devices. This difference is assumed to influence the viscosity of the secretion. Here, we applied a microrheological technique, based on the immersion of nanometric beads in the collected tarsal footprints, to estimate secretion viscosity in a beetle (Coccinella septempunctata) and a fly (Calliphora vicina). Both species studied possess distinct differences in viscosity, the median of which was calculated as 21.8 and 10.9 mPa s, respectively. We further present an approximate theoretical model to calculate the contact formation time of spatula-like terminal contact elements using the viscosity data of the covering fluid. The estimated contact formation time is proportional to the tarsal secretion viscosity and to the square of the contact radius of the contact element., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

Araneomorph spiders have evolved different silks with dissimilar material properties, serving different purposes. The two-compound pyriform secretion is used to glue silk threads to substrates or to other threads. It is applied in distinct patterns, called attachment discs. Although ubiquitously found in spider silk applications and hypothesized to be strong and versatile at low material consumption, the performance of attachment discs on different substrates remains unknown. Here, we analyse the detachment forces and fracture mechanics of the attachment discs spun by five different species on three different substrates, by pulling on the upstream part of the attached thread. Results show that although the adhesion of the pyriform glue is heavily affected by the substrate, even on Teflon it is frequently strong enough to hold the spider's weight. As plant surfaces are often difficult to wet, they are hypothesized to be the major driving force for evolution of the pyriform secretion., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

The bird feather vane can be separated into two parts by pulling the barbs apart. The original state can be re-established easily by lightly stroking through the feather. Hooklets responsible for holding vane barbs together are not damaged by multiple zipping and unzipping cycles. Because numerous microhooks keep the integrity of the feather, their properties are of great interest for understanding mechanics of the entire feather structure. This study was undertaken to estimate the separation force of single hooklets and their arrays using force measurement of an unzipping feather vane. The hooklets usually separate in some number synchronously (20 on average) with the highest observed separation force of 1.74 mN (average force 0.27 mN), whereas the single hooklet separation force was 14 μN. A simple numerical model was suggested for a better understanding of zipping and unzipping behaviour in feathers. The model demonstrates features similar to those observed in experiments.

In this study, we investigate the morphology and mechanical features of Octopus vulgaris suckers, which may serve as a model for the creation of a new generation of attachment devices. Octopus suckers attach to a wide range of substrates in wet conditions, including rough surfaces. This amazing feature is made possible by the sucker's tissues, which are pliable to the substrate profile. Previous studies have described a peculiar internal structure that plays a fundamental role in the attachment and detachment processes of the sucker. In this work, we present a mechanical characterization of the tissues involved in the attachment process, which was performed using microindentation tests. We evaluated the elasticity modulus and viscoelastic parameters of the natural tissues (E ∼ 10 kPa) and measured the mechanical properties of some artificial materials that have previously been used in soft robotics. Such a comparison of biological prototypes and artificial material that mimics octopus-sucker tissue is crucial for the design of innovative artificial suction cups for use in wet environments. We conclude that the properties of the common elastomers that are generally used in soft robotics are quite dissimilar to the properties of biological suckers.

Leafhoppers (Insecta, Hemiptera, Cicadellidae) actively coat their integuments with buckyball-shaped submicron proteinaceous secretory particles, called brochosomes. Here, we demonstrate that brochosomal coats, recently shown to be superhydrophobic, act as non-stick coatings and protect leafhoppers from contamination with their own sticky exudates--filtered plant sap. We exposed 137 wings of Alnetoidia alneti (Dahlbom), from half of which brochosomes were removed, to the rain of exudates under a colony of live A. alneti. One hundred and fifty-two droplets became stuck to the bared wings and only three to the intact wings. Inspection of the wings with a scanning electron microscope confirmed that the droplets that had hit the intact wings had rolled or bounced off the brochosomal coats. This is the first experimental study that tested a biological function of the brochosomal coats of leafhopper integuments. We argue that the production of brochosomes in leafhoppers and production of epidermal wax blooms in other sap-sucking hemipterans are alternative solutions, both serving to protect these insects from entrapment by their exudates.

During sexual conflict, males and females are expected to evolve traits and behaviours with a sexually antagonistic function. Recently, sexually antagonistic coevolution was proposed to occur between male and female diving beetles (Dytiscidae). Male diving beetles possess numerous suction cups on their forelegs whereas females commonly have rough structures on their elytra. These rough structures have been suggested to obstruct adhesion from male suction cups during mating attempts. However, some diving beetle species are dimorphic, where one female morph has a rough elytra and the other has a smooth elytra. Here, we used biomechanics to study the adhesive performance of male suction cups on the female morphs in two diving beetle species: Dytiscus lapponicus and Graphoderus zonatus. We compared adhesion on the rough and the smooth female morphs to infer the function of the rough elytral modifications. We found that the adhesive force on the rough structures was much lower than on other surfaces. These findings support the suggestion of sexual conflict in diving beetles and a sexually antagonistic function of the rough female structures. In addition, males differed in their adhesive capacity on different female surfaces, indicating a male trade-off between adhering to smooth and rough female morphs.

Hook-like surface structures, observed in some plant species, play an important role in the process of plant growth and seed dispersal. In this study, we developed an elastic model and further used it to investigate the mechanical behaviour of fruit hooks in four plant species, previously measured in an experimental study. Based on Euler-Bernoulli beam theory, the force-displacement relationship is derived, and its Young's modulus is obtained. The result agrees well with the experimental data. The model aids in understanding the mechanics of hooks, and could be used in the development of new bioinspired Velcro-like materials.

Reliable attachment ability of insect adhesive pads is proposed to be due to pad secretion. It has been shown that surface roughness strongly reduces adhesion forces of insect pads. This effect has been explained by decreased contact area and rapid fluid absorption from the pad surface by rough surfaces. However, it remains unclear how the fluid flows on rough substrates having different roughness parameters and surface energy. In this paper, we numerically studied the fluid flow on rough substrates during contact formation. The results demonstrate that an increase in the density of the substrate structures leads to an increase in fluid loss from the pad: substrates with a fine roughness absorb pad fluid faster. Decreased affinity of the solid substrate to the fluid has a more remarkable effect on the fluid loss and leads to a decrease in the fluid loss. With an increase in the aspect ratio of the substrate irregularities (porosity), the fluid loss is decreased. The numerical results obtained agree well with previous observations on insects and experimental results on nanoporous substrata. The significance of the obtained results for understanding biological wet adhesives is discussed.

On the basis of structural and experimental data, it was previously demonstrated that the snake integument consists of a hard, robust, inflexible outer surface (Oberhäutchen and β-layer) and softer, flexible inner layers (α-layers). It is not clear whether this phenomenon is a general adaptation of snakes to limbless locomotion or only to specific conditions, such as habitat and locomotion. The aim of the present study was to compare the structure and material properties of the outer scale layers (OSLs) and inner scale layers (ISLs) of the exuvium epidermis in four snake species specialized to live in different habitats: Lampropeltis getula californiae (terrestrial), Epicrates cenchria cenchria (generalist), Morelia viridis (arboreal) and Gongylophis colubrinus (sand-burrowing). Scanning electron microscopy (SEM) of skin cross sections revealed a strong variation in the epidermis structure between species. The nanoindentation experiments clearly demonstrated a gradient of material properties along the epidermis in the integument of all the species studied. The presence of such a gradient is a possible adaptation to locomotion and wear minimization on natural substrates. In general, the difference in both the effective elastic modulus and hardness of the OSL and ISL between species was not large compared with the difference in epidermis thickness and architecture.

Scanning white light interferometry and micro-force measurements were applied to analyse stimulus transformation in strain sensors in the spider exoskeleton. Two compound or 'lyriform' organs consisting of arrays of closely neighbouring, roughly parallel sensory slits of different lengths were examined. Forces applied to the exoskeleton entail strains in the cuticle, which compress and thereby stimulate the individual slits of the lyriform organs. (i) For the proprioreceptive lyriform organ HS-8 close to the distal joint of the tibia, the compression of the slits at the sensory threshold was as small as 1.4 nm and hardly more than 30 nm, depending on the slit in the array. The corresponding stimulus forces were as small as 0.01 mN. The linearity of the loading curve seems reasonable considering the sensor's relatively narrow biological intensity range of operation. The slits' mechanical sensitivity (slit compression/force) ranged from 106 down to 13 nm mN(-1), and gradually decreased with decreasing slit length. (ii) Remarkably, in the vibration-sensitive lyriform organ HS-10 on the metatarsus, the loading curve was exponential. The organ is thus adapted to the detection of a wide range of vibration amplitudes, as they are found under natural conditions. The mechanical sensitivities of the two slits examined in this organ in detail differed roughly threefold (522 and 195 nm mN(-1)) in the biologically most relevant range, again reflecting stimulus range fractionation among the slits composing the array.

To shed light on the role of suction in adhesion of microstructure with mushroom-shaped terminal elements, we compared pull-off forces measured at different retraction velocities on structured and smooth surfaces under different pressure conditions. The results obtained allow us to suggest that suction may contribute up to 10 per cent of the pull-off force measured on the structured surfaces at high velocities. We therefore conclude that the attachment ability of this biomimetic adhesive must not be solely based on van der Waals forces. Our experiments also suggest a change in visco-elastic properties of the structured surfaces compared with the bulk material. Based on the results obtained, it is assumed that this adhesive may be suitable in dynamic pick-and-drop processes even under vacuum conditions at which sufficiently high adhesive capability is maintained.

Traction experiments with adult seven-spotted ladybird beetles Coccinella septempunctata (L.) were carried out to study the influence of surface structure on insect attachment. Force measurements were performed with tethered walking insects, both males and females, on five different substrates: (i) smooth glass plate, (ii) smooth solid Al(2)O(3) (sapphire) disc, and (iii-v) porous Al(2)O(3) discs (anodisc membranes) with the same pore diameter but different porosity. The traction force of beetles ranged from 0.16 to 16.59 mN in males and from 0.32 to 8.99 mN in females. In both sexes, the highest force values were obtained on smooth solid surfaces, where males showed higher forces than females. On all three porous substrates, forces were significantly reduced in both males and females, and the only difference within these surfaces was obtained between membranes with the highest and lowest porosity. Males produced essentially lower forces than females on porous samples. The reduction in insect attachment on anodisc membranes may be explained by (i) possible absorption of the secretion fluid from insect adhesive pads by porous media and/or (ii) the effect of surface roughness. Differences in attachment between males and females were probably caused by the sexual dimorphism in the terminal structure of adhesive setae.