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

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 >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.To AH: Research grant from the Konrad-Adenauer-Foundation To DL: Research grant from the Cusanuswerk To WF: UK Biotechnology and Biological Sciences Research Council (BB/I008667/1)This is the final version of the article. It first appeared from RSC via http://dx.doi.org/10.1098/rsos.15012

Morphology of powerful suction organs from blepharicerid larvae living in raging torrents

AbstractBackgroundSuction organs provide powerful yet dynamic attachments for many aquatic animals, including octopus, squid, remora, and clingfish. While the functional morphology of suction organs from some cephalopods and fishes has been investigated in detail, there are only few studies on such attachment devices in insects. Here we characterise the morphology and ultrastructure of the suction attachment organs of net-winged midge larvae (genusLiponeura; Diptera: Blephariceridae) – aquatic insects that live on rocks in rapid alpine waterways where flow speeds can reach 3 m s− 1– using scanning electron microscopy, confocal laser scanning microscopy, and X-ray computed micro-tomography (micro-CT). Furthermore, we study the function of these organs in vivo using interference reflection microscopy.ResultsWe identified structural adaptations important for the function of the suction attachment organs inL. cinerascensandL. cordata. First, a dense array of spine-like microtrichia covering each suction disc comes into contact with the substrate upon attachment, analogous to hairy structures on suction organs from octopus, clingfish, and remora fish. These spine-like microtrichia may contribute to the seal and provide increased shear force resistance in high-drag environments. Second, specialised rim microtrichia at the suction disc periphery were found to form a continuous ring in close contact and may serve as a seal on a variety of surfaces. Third, a V-shaped cut on the suction disc (“V-notch“) is actively opened via two cuticular apodemes inserting on its flanks. The apodemes are attached to dedicated V-notch opening muscles, thereby providing a unique detachment mechanism. The complex cuticular design of the suction organs, along with specialised muscles that attach to them, allows blepharicerid larvae to generate powerful attachments which can withstand strong hydrodynamic forces and quickly detach for locomotion.ConclusionThe suction organs fromLiponeuraare underwater attachment devices specialised for resisting extremely fast flows. Structural adaptations from these suction organs could translate into future bioinspired attachment systems that perform well on a wide range of surfaces.</jats:sec

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.We are sincerely grateful to all our colleagues who readily shared published and unpublished data with us: Aaron M. Bauer, Jon Barnes, Niall Crawford, Thomas Endlein, Hanns Hagen Goetzke, Thomas E. Macrini, Anthony P. Russell & Joanna M. Smith. We also thank Casey Gilman, Dylan Briggs, Irina Showalter, Dan King and Mike Imburgia for their assistance with the collection of gecko toepad data. This study was supported by research grants from the UK Biotechnology and Biological Sciences Research Council (BB/I008667/1) to WF, the Human Frontier Science Programme (RGP0034/2012) to DI, AJC and WF, the Denman Baynes Senior Research Fellowship to DL and a Discovery Early Career Research Fellowship (DE120101503) to CJC.This is the author accepted manuscript. The final version is available from the National Academy of Sciences via http://dx.doi.org/ 10.1073/pnas.151945911

Sticking under wet conditions: the remarkable attachment abilities of the torrent frog, staurois guttatus

Tree frogs climb smooth surfaces utilising capillary forces arising from an air-fluid interface around their toe pads, whereas torrent frogs are able to climb in wet environments near waterfalls where the integrity of the meniscus is at risk. This study compares the adhesive capabilities of a torrent frog to a tree frog, investigating possible adaptations for adhesion under wet conditions. We challenged both frog species to cling to a platform which could be tilted from the horizontal to an upside-down orientation, testing the frogs on different levels of roughness and water flow. On dry, smooth surfaces, both frog species stayed attached to overhanging slopes equally well. In contrast, under both low and high flow rate conditions, the torrent frogs performed significantly better, even adhering under conditions where their toe pads were submerged in water, abolishing the meniscus that underlies capillarity. Using a transparent platform where areas of contact are illuminated, we measured the contact area of frogs during platform rotation under dry conditions. Both frog species not only used the contact area of their pads to adhere, but also large parts of their belly and thigh skin. In the tree frogs, the belly and thighs often detached on steeper slopes, whereas the torrent frogs increased the use of these areas as the slope angle increased. Probing small areas of the different skin parts with a force transducer revealed that forces declined significantly in wet conditions, with only minor differences between the frog species. The superior abilities of the torrent frogs were thus due to the large contact area they used on steep, overhanging surfaces. SEM images revealed slightly elongated cells in the periphery of the toe pads in the torrent frogs, with straightened channels in between them which could facilitate drainage of excess fluid underneath the pad

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

Pneumatosis intestinalis versus pseudo-pneumatosis: review of CT findings and differentiation

Pneumatosis intestinalis is defined as the presence of gas within the wall of the gastrointestinal tract. Originally described on plain abdominal radiographs, it is an imaging sign rather than a specific diagnosis and it is associated with both benign and life-threatening clinical conditions. The most common life-threatening cause of pneumatosis intestinalis is bowel ischaemia. Computed tomography (CT) is usually requested to detect underlying disease. The presence of pneumatosis intestinalis often leads physicians to make a diagnosis of serious disease. However, an erroneous diagnosis of pneumatosis intestinalis may be made (i.e. pseudo-pneumatosis) when intraluminal beads of gas are trapped within or between faeces and adjacent mucosal folds. The purpose of this pictorial essay is to review and describe the CT imaging findings of pneumatosis and pseudo-pneumatosis intestinalis and to discuss key discriminatory imaging features

PplD Is a De-N-Acetylase of the Cell Wall Linkage Unit of Streptococcal Rhamnopolysaccharides

The cell wall of the human bacterial pathogen Group A Streptococcus (GAS) consists of peptidoglycan decorated with the Lancefield group A carbohydrate (GAC). GAC is a promising target for the development of GAS vaccines. In this study, employing chemical, compositional, and NMR methods, we show that GAC is attached to peptidoglycan via glucosamine 1-phosphate. This structural feature makes the GAC-peptidoglycan linkage highly sensitive to cleavage by nitrous acid and resistant to mild acid conditions. Using this characteristic of the GAS cell wall, we identify PplD as a protein required for deacetylation of linkage N-acetylglucosamine (GlcNAc). X-ray structural analysis indicates that PplD performs catalysis via a modified acid/base mechanism. Genetic surveys in silico together with functional analysis indicate that PplD homologs deacetylate the polysaccharide linkage in many streptococcal species. We further demonstrate that introduction of positive charges to the cell wall by GlcNAc deacetylation protects GAS against host cationic antimicrobial proteins