Biomechanical adaptations enable phoretic mite species to occupy distinct spatial niches on host burying beetles.

Niche theory predicts that ecologically similar species coexist by minimizing interspecific competition through niche partitioning. Therefore, understanding the mechanisms of niche partitioning is essential for predicting interactions and coexistence between competing organisms. Here, we study two phoretic mite species, Poecilochirus carabi and Macrocheles nataliae that coexist on the same host burying beetle Nicrophorus vespilloides and use it to 'hitchhike' between reproductive sites. Field observations revealed clear spatial partitioning between species in distinct host body parts. Poecilochirus carabi preferred the ventral side of the thorax, whereas M. nataliae were exclusively found ventrally at the hairy base of the abdomen. Experimental manipulations of mite density showed that each species preferred these body parts, largely regardless of the density of the other mite species on the host beetle. Force measurements indicated that this spatial distribution is mediated by biomechanical adaptations, because each mite species required more force to be removed from their preferred location on the beetle. While P. carabi attached with large adhesive pads to the smooth thorax cuticle, M. nataliae gripped abdominal setae with their chelicerae. Our results show that specialist biomechanical adaptations for attachment can mediate spatial niche partitioning among species sharing the same host.

Bumblebees negotiate a trade-off between nectar quality and floral biomechanics.

How and why pollinators choose which flowers to visit are fundamental, multifaceted questions in pollination biology, yet most studies of floral traits measure simple relative preferences. Here, we used vertically and horizontally oriented slippery-surfaced artificial flowers to test whether bumblebees could make a trade-off between floral handling difficulty and nectar sucrose concentration. We quantified foraging energetics, thereby resolving the rationale behind the bees' foraging decisions. The bees chose flowers with either a high handling cost or low sucrose concentration, depending on which was the energetically favorable option. Their behavior agreed with the critical currency being the rate of energy return (net energy collected per unit time), not energetic efficiency (net energy collected per unit energy spent). This suggests that bumblebees prioritize immediate carbohydrate flow to the nest rather than energy gain over the working lifespan of each bee. Trade-off paradigms like these are a powerful approach for quantifying pollinator trait preferences.

Extreme suction attachment performance from specialised insects living in mountain streams (Diptera: Blephariceridae).

Suction is widely used by animals for strong controllable underwater adhesion but is less well understood than adhesion of terrestrial climbing animals. Here we investigate the attachment of aquatic insect larvae (Blephariceridae), which cling to rocks in torrential streams using the only known muscle-actuated suction organs in insects. We measured their attachment forces on well-defined rough substrates and found that their adhesion was less reduced by micro-roughness than that of terrestrial climbing insects. In vivo visualisation of the suction organs in contact with microstructured substrates revealed that they can mould around large asperities to form a seal. We have shown that the ventral surface of the suction disc is covered by dense arrays of microtrichia, which are stiff spine-like cuticular structures that only make tip contact. Our results demonstrate the impressive performance and versatility of blepharicerid suction organs and highlight their potential as a study system to explore biological suction mechanisms.

Suction cups are widely used to attach objects to surfaces in bathrooms and kitchens. They work well on tiles and other smooth surfaces, but do not stick well to rougher materials like brick or wood because they are unable to form an air-tight seal. Researchers have been searching for ways to improve these cups by studying how octopuses, remora fish and other sea animals use muscle-powered suction organs to stick to wet and rough surfaces. However, the experiments needed to understand the detailed mechanics of suction organs are difficult to perform on living specimens of these animals. The aquatic larvae of a family of insects known as the net-winged midges also have suction organs that are powered by muscles. These insects survive in fast flowing mountain streams where they use their suction organs to stick to rocks underwater. However, it remained unclear how these suction organs work. Here, Kang et al. found that net-winged midge larvae attach extremely well to a variety of surfaces. The larvae were able to withstand forces over one thousand times their body weight when attached to smooth surfaces. Even on rough materials, where human-made suction cups attach poorly, the larvae were able to withstand forces up to 240-times their body weight. Further experiments using several microscopy approaches revealed that the suction organs of the larvae are covered in multiple spine-like structures called microtrichia that interlock with bumps and dips on a surface to help the organ remain in place. Similar structures have previously been found on the suction organs of remora fish, but are not as tightly packed together. These findings demonstrate that net-winged midge larvae may be useful model systems to study how natural suction organs operate. Furthermore, they provide a new source of inspiration for scientists and engineers to design and manufacture suction cups capable of attaching to a wider variety of surfaces.

How a sticky fluid facilitates prey retention in a carnivorous pitcher plant (Nepenthes rafflesiana).

Nepenthes pitcher plants grow in nutrient-poor soils and produce large pitfall traps to obtain additional nutrients from animal prey. Previous research has shown that the digestive secretion in N. rafflesiana is a sticky viscoelastic fluid that retains insects much more effectively than water, even after significant dilution. Although the retention of prey is known to depend on the fluid's physical properties, the details of how the fluid interacts with insect cuticle and how its sticky nature affects struggling insects are unclear. In this study, we investigated the mechanisms behind the efficient prey retention in N. rafflesiana pitcher fluid. By measuring the attractive forces on insect body parts moved in and out of test fluids, we show that it costs insects more energy to free themselves from pitcher fluid than from water. Moreover, both the maximum force and the energy required for retraction increased after the first contact with the pitcher fluid. We found that insects sink more easily into pitcher fluid than water and, accordingly, the surface tension of N. rafflesiana pitcher fluid was lower than that of water (60.2 vs. 72.3 mN/m). By analysing the pitcher fluid's wetting behaviour, we demonstrate that it strongly resists dewetting from all surfaces tested, leaving behind residual films and filaments that can facilitate re-wetting. This inhibition of dewetting may be a further consequence of the fluid's viscoelastic nature and likely represents a key mechanism underlying prey retention in Nepenthes pitcher plants. STATEMENT OF SIGNIFICANCE: Carnivorous Nepenthes pitcher plants secrete sticky viscoelastic fluids that prevent insects from escaping after falling into the pitcher. What physical mechanisms are responsible for the fluid's retentive function? First, insects sink and drown more readily in N. rafflesiana pitcher fluid due to its reduced surface tension. Second, once within the fluid, our force measurements show that it costs more energy to separate insects from pitcher fluid than from water. Third, the fluid strongly resists dewetting, making it harder for insects to extract themselves and covering their cuticle with residues that facilitate re-wetting. Such striking inhibition of dewetting may represent a previously unrecognised mechanism of prey retention by Nepenthes. Pitcher fluid fulfils a well-defined biological function and may serve as a model for studying the mechanics of complex fluids.

Role of legs and foot adhesion in salticid spiders jumping from smooth surfaces.

Many spiders and insects can perform rapid jumps from smooth plant surfaces. Here, we investigate how jumping spiders (Pseudeuophrys lanigera and Sitticus pubescens) avoid slipping when accelerating. Both species differed in the relative contribution of leg pairs to the jump. P. lanigera accelerated mainly with their long third legs, whereas their short fourth legs detached earlier. In contrast, S. pubescens accelerated mainly with their long fourth legs, and their short third legs detached earlier. Because of the different orientation (fourth-leg tip pointing backward, third-leg tip pointing forward), the fourth-leg tarsus pushed, whereas the third-leg tarsus pulled. High-speed video recordings showed that pushing and pulling was achieved by different attachment structures. In P. lanigera, third-leg feet made surface contact with setae on their distal or lateral claw tuft, whereas fourth-leg feet engaged the proximal claw tuft, and the distal tuft was raised off the ground. S. pubescens showed the same division of labour between proximal and distal claw tuft for pushing and pulling, but the claw tuft contact lasted longer and was more visible in the fourth than in the third legs. Experimental ablation of claw tufts caused accelerating spiders to slip, confirming that adhesion is essential for jumps from smooth substrates.

Disentangling the role of surface topography and intrinsic wettability in the prey capture mechanism of Nepenthes pitcher plants.

Nepenthes pitcher plants capture prey with leaves specialised as pitfall traps. Insects are trapped when they 'aquaplane' on the pitcher rim (peristome), a surface structured with macroscopic and microscopic radial ridges. What is the functional significance of this hierarchical surface topography? Here, we use insect pad friction measurements, photolithography, wetting experiments and physical modelling to demonstrate that the ridges enhance the trap's efficacy by satisfying two functional demands on prey capture: Macroscopic ridges restrict lateral but enhance radial spreading of water, thereby creating continuous slippery tracks which facilitate prey capture when little water is present. Microscopic ridges, in turn, ensure that the water film between insect pad and peristome remains stable, causing insects to aquaplane. In combination, the hierarchical ridge structure hence renders the peristome wettable, and water films continuous, so avoiding the need for a strongly hydrophilic surface chemistry, which would compromise resistance to desiccation and attract detrimental contamination.

The mechanics of nectar offloading in the bumblebee Bombus terrestris and implications for optimal concentrations during nectar foraging.

Nectar is a common reward provided by plants for pollinators. More concentrated nectar is more rewarding, but also more viscous, and hence more time-consuming to drink. Consequently, theory predicts an optimum concentration for maximizing energy uptake rate, dependent on the mechanics of feeding. For social pollinators such as bumblebees, another important but little-studied aspect of foraging is nectar offloading upon return to the nest. Studying the bumblebee Bombus terrestris, we found that the relationship between viscosity (µ) and volumetric transfer rates (Q) of sucrose solutions differed between drinking and offloading. For drinking, Q ∝ µ-0.180, in good agreement with previous work. Although offloading was quicker than drinking, offloading rate decreased faster with viscosity, with Q ∝ µ-0.502, consistent with constraints imposed by fluid flow through a tube. The difference in mechanics between drinking and offloading nectar leads to a conflict in the optimum concentration for maximizing energy transfer rates. Building a model of foraging energetics, we show that including offloading lowers the maximum rate of energy return to the nest and reduces the concentration which maximizes this rate by around 3%. Using our model, we show that published values of preferred nectar sugar concentrations suggest that bumblebees maximize the overall energy return rather than the instantaneous energy uptake during drinking.

Shear-sensitive adhesion enables size-independent adhesive performance in stick insects.

The ability to climb with adhesive pads conveys significant advantages and is widespread in the animal kingdom. The physics of adhesion predict that attachment is more challenging for large animals, whereas detachment is harder for small animals, due to the difference in surface-to-volume ratios. Here, we use stick insects to show that this problem is solved at both ends of the scale by linking adhesion to the applied shear force. Adhesive forces of individual insect pads, measured with perpendicular pull-offs, increased approximately in proportion to a linear pad dimension across instars. In sharp contrast, whole-body force measurements suggested area scaling of adhesion. This discrepancy is explained by the presence of shear forces during whole-body measurements, as confirmed in experiments with pads sheared prior to detachment. When we applied shear forces proportional to either pad area or body weight, pad adhesion also scaled approximately with area or mass, respectively, providing a mechanism that can compensate for the size-related loss of adhesive performance predicted by isometry. We demonstrate that the adhesion-enhancing effect of shear forces is linked to pad sliding, which increased the maximum adhesive force per area sustainable by the pads. As shear forces in natural conditions are expected to scale with mass, sliding is more frequent and extensive in large animals, thus ensuring that large animals can attach safely, while small animals can still detach their pads effortlessly. Our results therefore help to explain how nature's climbers maintain a dynamic attachment performance across seven orders of magnitude in body weight.

Dynamic biological adhesion: mechanisms for controlling attachment during locomotion.

The rapid control of surface attachment is a key feature of natural adhesive systems used for locomotion, and a property highly desirable for man-made adhesives. Here, we describe the challenges of adhesion control and the timescales involved across diverse biological attachment systems and different adhesive mechanisms. The most widespread control principle for dynamic surface attachment in climbing animals is that adhesion is 'shear-sensitive' (directional): pulling adhesive pads towards the body results in strong attachment, whereas pushing them away from it leads to easy detachment, providing a rapid mechanical 'switch'. Shear-sensitivity is based on changes of contact area and adhesive strength, which in turn arise from non-adhesive default positions, the mechanics of peeling, pad sliding, and the targeted storage and controlled release of elastic strain energy. The control of adhesion via shear forces is deeply integrated with the climbing animals' anatomy and locomotion, and involves both active neuromuscular control, and rapid passive responses of sophisticated mechanical systems. The resulting dynamic adhesive systems are robust, reliable, versatile and nevertheless remarkably simple. This article is part of the theme issue 'Transdisciplinary approaches to the study of adhesion and adhesives in biological systems'.

Strong Wet and Dry Adhesion by Cupped Microstructures.

Recent advances in bio-inspired microfibrillar adhesives have resulted in technologies that allow reliable attachment to a variety of surfaces. Because capillary and van der Waals forces are considerably weakened underwater, fibrillar adhesives are however far less effective in wet environments. Although various strategies have been proposed to achieve strong reversible underwater adhesion, strong adhesives that work both in air and underwater without additional surface treatments have yet to be developed. In this study, we report a novel design-cupped microstructures (CM)-that generates strong controllable adhesion in air and underwater. We measured the adhesive performance of cupped polyurethane microstructures with three different cup angles (15, 30, and 45°) and the same cup diameter of 100 µm in dry and wet conditions in comparison to standard mushroom-shaped microstructures (MSMs) of the same dimensions. In air, 15°CM performed comparably to the flat MSM of the same size with an adhesion strength (force per real contact area) of up to 1.3 MPa, but underwater, 15°CM achieved 20 times stronger adhesion than MSM (∼1 MPa versus ∼0.05 MPa). Furthermore, the cupped microstructures exhibit self-sealing properties, whereby stronger pulls lead to longer stable attachment and much higher adhesion through the formation of a better seal.

Froghoppers jump from smooth plant surfaces by piercing them with sharp spines.

Attachment mechanisms used by climbing animals facilitate their interactions with complex 3D environments and have inspired novel types of synthetic adhesives. Here we investigate one of the most dynamic forms of attachment, used by jumping insects living on plants. Froghopper insects can perform explosive jumps with some of the highest accelerations known among animals. As many plant surfaces are smooth, we studied whether Philaenus spumarius froghoppers are able to take off from such substrates. When attempting to jump from smooth glass, the insects' hind legs slipped, resulting in weak, uncontrolled jumps with a rapid forward spin. By contrast, on smooth ivy leaves and smooth epoxy surfaces, Philaenus froghoppers performed strong jumps without any slipping. We discovered that the insects produced traction during the acceleration phase by piercing these substrates with sharp spines of their tibia and tarsus. High-speed microscopy recordings of hind legs during the acceleration phase of jumps revealed that the spine tips indented and plastically deformed the substrate. On ivy leaves, the spines of jumping froghoppers perforated the cuticle and epidermal cell walls, and wounds could be visualized after the jumps by methylene blue staining and scanning electron microscopy. Improving attachment performance by indenting or piercing plant surfaces with sharp spines may represent a widespread but previously unrecognized strategy utilized by plant-living insects. This attachment mechanism may also provide inspiration for the design of robotic grippers.

Scaling of claw sharpness: mechanical constraints reduce attachment performance in larger insects.

Claws are the most widespread attachment devices in animals, but comparatively little is known about the mechanics of claw attachment. A key morphological parameter in determining attachment ability is claw sharpness; however, there is a conflict between sharpness and fracture resistance. Sharper claws can interlock on more surfaces but are more likely to break. Body size interacts with this conflict such that larger animals should have much blunter claws and consequently poorer attachment ability than smaller animals. This expected size-induced reduction in attachment performance has not previously been investigated, and it is unclear how animals deal with this effect, and whether it indeed exists. We explored the scaling of claw sharpness with body size using four insect species (Nauphoeta cinerea, Gromphadorhina portentosa, Atta cephalotes and Carausius morosus) each covering a large size range. The scaling of claw sharpness varied significantly between species, suggesting that they face different pressures regarding claw function. Attachment forces were measured for A. cephalotes and G. portentosa (which had different scaling of claw sharpness) on several rough surfaces using a centrifuge setup. As expected, attachment performance was poorer in larger animals. Firstly, larger animals were more likely to slip, although this effect depended on the scaling of claw sharpness. Secondly, when they gripped, they attached with smaller forces relative to their weight. This size-induced reduction in attachment performance has significant implications for the attachment ability of larger animals on rough surfaces.

Coping with the climate: cuticular hydrocarbon acclimation of ants under constant and fluctuating conditions.

Terrestrial arthropods achieve waterproofing by a layer of cuticular hydrocarbons (CHCs). At the same time, CHCs also serve as communication signals. To maintain waterproofing under different climate conditions, insects adjust the chemical composition of their CHC layer, but this may affect the communication via CHCs. The detailed acclimatory changes of CHCs and how these influence their physical properties are still unknown. Here, we studied acclimation in two closely related ant species with distinct CHC profiles, Myrmica rubra and Myrmica ruginodis, in response to constant or fluctuating temperature and humidity regimes. We measured how acclimation affected CHC composition and viscosity, and the ants' drought survival. In both species, CHC composition showed strong, predictable responses to temperature regimes. Warm-acclimated individuals had higher proportions of linear alkanes, and less methyl-branched or unsaturated CHCs. These changes coincided with higher solid content and viscosity of CHCs in warm-acclimated ants. Temperature fluctuation caused effects similar to those observed under constant-cool conditions in Mrubra, but led to entirely different profiles in Mruginodis, suggesting that fluctuating and constant conditions pose very different challenges. Acclimation to dry conditions led to higher absolute amounts of CHCs, which increased the ants' drought survival, whereas temperature acclimation did not. Hence, the temperature-induced CHC changes cannot be explained by the need for waterproofing alone. Although these changes could be non-adaptive, we propose that they serve to maintain a constant CHC viscosity, which may be essential for communication and other functions.

Jumping without slipping: leafhoppers (Hemiptera: Cicadellidae) possess special tarsal structures for jumping from smooth surfaces.

Many hemipteran bugs can jump explosively from plant substrates, which can be very smooth. We therefore analysed the jumping performance of froghoppers (Philaenus spumarius, Aphrophoridae) and leafhoppers (Aphrodes bicinctus/makarovi, Cicadellidae) taking off from smooth (glass) and rough (sandpaper, 30 µm asperity size) surfaces. On glass, the propulsive hind legs of Philaenus froghoppers slipped, resulting in uncontrolled jumps with a fast forward spin, a steeper angle and only a quarter of the velocity compared with jumps from rough surfaces. By contrast, Aphrodes leafhoppers took off without their propulsive hind legs slipping, and reached low take-off angles and high velocities on both substrates. This difference in jumping ability from smooth surfaces can be explained not only by the lower acceleration of the long-legged leafhoppers, but also by the presence of 2-9 soft pad-like structures (platellae) on their hind tarsi, which are absent in froghoppers. High-speed videos of jumping showed that platellae contact the surface briefly (approx. 3 ms) during the acceleration phase. Friction force measurements on individual hind tarsi on glass revealed that at low sliding speeds, both pushing and pulling forces were small, and insufficient to explain the recorded jumps. Only when the tarsi were pushed with higher velocities did the contact area of the platellae increase markedly, and high friction forces were produced, consistent with the observed jumps. Our findings show that leafhoppers have special adhesive footpads for jumping from smooth surfaces, which achieve firm grip and rapid control of attachment/detachment by combining anisotropic friction with velocity dependence.

Biomechanics of shear-sensitive adhesion in climbing animals: peeling, pre-tension and sliding-induced changes in interface strength.

Many arthropods and small vertebrates use adhesive pads for climbing. These biological adhesives have to meet conflicting demands: attachment must be strong and reliable, yet detachment should be fast and effortless. Climbing animals can rapidly and reversibly control their pads' adhesive strength by shear forces, but the mechanisms underlying this coupling have remained unclear. Here, we show that adhesive forces of stick insect pads closely followed the predictions from tape peeling models when shear forces were small, but strongly exceeded them when shear forces were large, resulting in an approximately linear increase of adhesion with friction. Adhesion sharply increased at peel angles less than ca 30°, allowing a rapid switch between attachment and detachment. The departure from classic peeling theory coincided with the appearance of pad sliding, which dramatically increased the peel force via a combination of two mechanisms. First, partial sliding pre-stretched the pads, so that they were effectively stiffer upon detachment and peeled increasingly like inextensible tape. Second, pad sliding reduces the thickness of the fluid layer in the contact zone, thereby increasing the stress levels required for peeling. In combination, these effects can explain the coupling between adhesion and friction that is fundamental to adhesion control across all climbing animals. Our results highlight that control of adhesion is not solely achieved by direction-dependence and morphological anisotropy, suggesting promising new routes for the development of controllable bio-inspired adhesives.

Elasto-capillarity in insect fibrillar adhesion.

The manipulation of microscopic objects is challenging because of high adhesion forces, which render macroscopic gripping strategies unsuitable. Adhesive footpads of climbing insects could reveal principles relevant for micro-grippers, as they are able to attach and detach rapidly during locomotion. However, the underlying mechanisms are still not fully understood. In this work, we characterize the geometry and contact formation of the adhesive setae of dock beetles (Gastrophysa viridula) by interference reflection microscopy. We compare our experimental results to the model of an elastic beam loaded with capillary forces. Fitting the model to experimental data yielded not only estimates for seta adhesion and compliance in agreement with previous direct measurements, but also previously unknown parameters such as the volume of the fluid meniscus and the bending stiffness of the tip. In addition to confirming the primary role of surface tension for insect adhesion, our investigation reveals marked differences in geometry and compliance between the three main kinds of seta tips in leaf beetles.

Extreme positive allometry of animal adhesive pads and the size limits of adhesion-based climbing.

Organismal functions are size-dependent whenever body surfaces supply body volumes. Larger organisms can develop strongly folded internal surfaces for enhanced diffusion, but in many cases areas cannot be folded so that their enlargement is constrained by anatomy, presenting a problem for larger animals. Here, we study the allometry of adhesive pad area in 225 climbing animal species, covering more than seven orders of magnitude in weight. Across all taxa, adhesive pad area showed extreme positive allometry and scaled with weight, implying a 200-fold increase of relative pad area from mites to geckos. However, allometric scaling coefficients for pad area systematically decreased with taxonomic level and were close to isometry when evolutionary history was accounted for, indicating that the substantial anatomical changes required to achieve this increase in relative pad area are limited by phylogenetic constraints. Using a comparative phylogenetic approach, we found that the departure from isometry is almost exclusively caused by large differences in size-corrected pad area between arthropods and vertebrates. To mitigate the expected decrease of weight-specific adhesion within closely related taxa where pad area scaled close to isometry, data for several taxa suggest that the pads' adhesive strength increased for larger animals. The combination of adjustments in relative pad area for distantly related taxa and changes in adhesive strength for closely related groups helps explain how climbing with adhesive pads has evolved in animals varying over seven orders of magnitude in body weight. Our results illustrate the size limits of adhesion-based climbing, with profound implications for large-scale bio-inspired adhesives.

On Heels and Toes: How Ants Climb with Adhesive Pads and Tarsal Friction Hair Arrays.

Ants are able to climb effortlessly on vertical and inverted smooth surfaces. When climbing, their feet touch the substrate not only with their pretarsal adhesive pads but also with dense arrays of fine hairs on the ventral side of the 3rd and 4th tarsal segments. To understand what role these different attachment structures play during locomotion, we analysed leg kinematics and recorded single-leg ground reaction forces in Weaver ants (Oecophylla smaragdina) climbing vertically on a smooth glass substrate. We found that the ants engaged different attachment structures depending on whether their feet were above or below their Centre of Mass (CoM). Legs above the CoM pulled and engaged the arolia ('toes'), whereas legs below the CoM pushed with the 3rd and 4th tarsomeres ('heels') in surface contact. Legs above the CoM carried a significantly larger proportion of the body weight than legs below the CoM. Force measurements on individual ant tarsi showed that friction increased with normal load as a result of the bending and increasing side contact of the tarsal hairs. On a rough sandpaper substrate, the tarsal hairs generated higher friction forces in the pushing than in the pulling direction, whereas the reverse effect was found on the smooth substrate. When the tarsal hairs were pushed, buckling was observed for forces exceeding the shear forces found in climbing ants. Adhesion forces were small but not negligible, and higher on the smooth substrate. Our results indicate that the dense tarsal hair arrays produce friction forces when pressed against the substrate, and help the ants to push outwards during horizontal and vertical walking.

Functional morphology and efficiency of the antenna cleaner in Camponotus rufifemur ants.

Contamination of body surfaces can negatively affect many physiological functions. Insects have evolved different adaptations for removing contamination, including surfaces that allow passive self-cleaning and structures for active cleaning. Here, we study the function of the antenna cleaner in Camponotus rufifemur ants, a clamp-like structure consisting of a notch on the basitarsus facing a spur on the tibia, both bearing cuticular 'combs' and 'brushes'. The ants clamp one antenna tightly between notch and spur, pull it through, and subsequently clean the antenna cleaner itself with the mouthparts. We simulated cleaning strokes by moving notch or spur over antennae contaminated with fluorescent particles. The notch removed particles more efficiently than the spur, but both components eliminated more than 60% of the particles with the first stroke. Ablation of bristles, brush and comb strongly reduced the efficiency, indicating that they are essential for cleaning. To study how comb and brush remove particles of different sizes, we contaminated antennae of living ants, and anaesthetized them immediately after they had performed the first cleaning stroke. Different-sized beads were trapped in distinct zones of the notch, consistent with the gap widths between cuticular outgrowths. This suggests that the antenna cleaner operates like a series of sieves that remove the largest objects first, followed by smaller ones, down to the smallest particles that get caught by adhesion.

Rate-dependence of 'wet' biological adhesives and the function of the pad secretion in insects.

Many insects use soft adhesive footpads for climbing. The surface contact of these organs is mediated by small volumes of a liquid secretion, which forms thin films in the contact zone. Here, we investigate the role of viscous dissipation by this secretion and the 'bulk' pad cuticle by quantifying the rate-dependence of the adhesive force of individual pads. Adhesion increased with retraction speed, but this effect was independent of the amount of pad secretion present in the contact zone, suggesting that the secretion's viscosity did not play a significant role. Instead, the rate-dependence can be explained by relating the strain energy release rate to the speed of crack propagation, using an established empirical power law. The 'wet' pads' behaviour was akin to that of 'dry' elastomers, with an equilibrium energy release rate close to that of dry van-der-Waals contacts. We suggest that the secretion mainly serves as a 'release layer', minimising viscous dissipation and thereby reducing the time- and 'loading-history'-dependence of the adhesive pads. In contrast to many commercial adhesives which derive much of their strength from viscous dissipation, we show that the major modulator of adhesive strength in 'wet' biological adhesive pads is friction, exhibiting a much larger effect than retraction speed. A comparison between 'wet' and 'dry' biological adhesives, using both results from this study and the literature, revealed a striking lack of differences in attachment performance under varying experimental conditions. Together, these results suggest that 'wet' and 'dry' biological adhesives may be more similar than previously thought.