Editorial: Bioadhesion

Materials and systems preventing the separation of two surfaces may be defined as adhesives. There are a variety of natural adhesive devices based on entirely mechanical principles, while others additionally rely on the chemistry of polymers and colloids. Adhesive organs are functional systems, the purpose of which is either temporary or permanent attachment of an organism to the substrate surface, to another organism, or temporary interconnection of body parts within an organism. Their design varies enormously and is subject to different functional loads. There is no doubt that many functional solutions have evolved independently in different lineages of organisms. Many species of animals and plants are supplied with diverse adhesive surfaces, the morphology of which depends on the species biology, and the particular function in which the adhesive device is involved. There are numerous publications on cell adhesion phenomena, but much fewer references are devoted to the non-specific adhesion of living organisms. Because of the structural, mechanical, and chemical complexity of biological surfaces related to adhesion, exact working mechanisms have been clarified only for some systems. In this Research Topic (RT) “Bioadhesion”, we aimed at collecting articles dealing with biological surfaces and systems specialized for adhesion enhancement. These contributions discuss adhesive function of biological surfaces and their relationship with the structure, contact mechanics and chemistry of surfaces. Because of the diversity of functions in adhesion-related biological surfaces, biology could provide interesting inspirations for a broad range of topics in physics, chemistry, and engineering. In the following, we briefly introduce the 13 articles comprising this RT. They can be broadly divided into four categories: 1) bacterial adhesion, 2) plant adhesion, 3) animal adhesion, and 4) biomimetic adhesion

Mandibular gnathobases of marine planktonic copepods – feeding tools with complex micro- and nanoscale composite architectures

Copepods are dominant members of the marine zooplankton. Their diets often comprise large proportions of diatom taxa whose silicified frustules are mechanically stable and offer protection against grazers. Despite of this protection, many copepod species are able to efficiently break even the most stable frustule types. This ability requires specific feeding tools with mechanically adapted architectures, compositions and properties. When ingesting food, the copepods use the gnathobases of their mandibles to grab and, if necessary, crush and mince the food items. The morphology of these gnathobases is related to the diets of the copepods. Gnathobases of copepod species that mainly feed on phytoplankton feature compact and stable tooth-like structures, so-called teeth. In several copepod species these gnathobase teeth have been found to contain silica. Recent studies revealed that the siliceous teeth are complex microscale composites with silica-containing cap-like structures located on chitinous exoskeleton sockets that are connected with rubber-like bearings formed by structures with high proportions of the soft and elastic protein resilin. In addition, the silica-containing cap-like structures exhibit a nanoscale composite architecture. They contain some amorphous silica and large proportions of the crystalline silica type α-cristobalite and are pervaded by a fine chitinous fibre network that very likely serves as a scaffold during the silicification process. All these intricate composite structures are assumed to be the result of a coevolution between the copepod gnathobases and diatom frustules in an evolutionary arms race. The composites very likely increase both the performance of the siliceous teeth and their resistance to mechanical damage, and it is conceivable that their development has favoured the copepods’ dominance of the marine zooplankton observed today

Role of Fruit Epicuticular Waxes in Preventing Bactrocera oleae (Diptera: Tephritidae) Attachment in Different Cultivars of Olea europaea

The olive fruit fly Bactrocera oleae (Diptera: Tephritidae) is the major pest of cultivated olives (Olea europaea L.), and a serious threat in all of the Mediterranean Region. In the present investigation, we demonstrated with traction force experiments that B. oleae female adhesion is reduced by epicuticular waxes (EWs) fruit surface, and that the olive fruit fly shows a different ability to attach to the ripe olive surface of different cultivars of O. europaea (Arbequina, Carolea, Dolce Agogia, Frantoio, Kalamata, Leccino, Manzanilla, Picholine, Nostrale di Rigali, Pendolino and San Felice) in terms of friction force and adhesion, in relation with different mean values of olive surface wettability. Cryo-scanning morphological investigation revealed that the EW present on the olive surface of the different analyzed cultivars are represented by irregular platelets varying in the orientation, thus contributing to affect the surface microroughness and wettability in the different cultivars, and consequently the olive fruit fly attachment. Further investigations to elucidate the role of EW in olive varietal resistance to the olive fruit fly in relation to the olive developmental stage and environmental conditions could be relevant to develop control methods alternative to the use of harmful pesticides

Jumping mechanism in the marsh beetles (Coleoptera: Scirtidae)

The jumping mechanism with supporting morphology and kinematics is described in the marsh beetle Scirtes hemisphaericus (Coleoptera: Scirtidae). In marsh beetles, the jump is performed by the hind legs by the rapid extension of the hind tibia. The kinematic parameters of the jump are: 139-1536 m s-2 (acceleration), 0.4-1.9 m s-1 (velocity), 2.7-8.4 ms (time to take-off), 0.2-5.4 × 10-6 J (kinetic energy) and 14-156 (g-force). The power output of a jumping leg during the jumping movement is 3.5 × 103 to 9.6 × 103 W kg-1. A resilin-bearing elastic extensor ligament is considered to be the structure that accumulates the elastic strain energy. The functional model of the jumping involving an active latching mechanism is proposed. The latching mechanism is represented by the conical projection of the tibial flexor sclerite inserted into the corresponding socket of the tibial base. Unlocking is triggered by the contraction of flexor muscle pulling the tibial flexor sclerite backwards which in turn comes out of the socket. According to the kinematic parameters, the time of full extension of the hind tibia, and the value of the jumping leg power output, this jumping mechanism is supposed to be latch-mediated spring actuation using the contribution of elastically stored strain energy

The frictional and adhesive properties of the shells of terrestrial hairy snails

Hair‐like structures of the shell in some terrestrial snails have been suggested to represent an adaptive advantage. One of the recently proposed explanations was the hypothesis that the possession of hairs facilitates the snails' adherence to the leaves of the plants during foraging, when humidity levels are high. In order to obtain some reliable data to test this contact mechanics hypothesis, the present paper deals with the frictional and adhesive properties of the smooth and hairy shells of two species of terrestrial mollusks, Trochulus hispidus and T. villosulus. Additionally, actively generated traction force of the snails was measured on different substrates. The conducted experiments revealed that in wet conditions the hairiness indeed increases friction and that adhesion through the water film is high enough to retain the weight of the snail. However, possession of long hairs decreases the adhesion, in comparison with the animals with short hairs. Our data suggest that hair‐like structures require more energy expenditure during locomotion and only provide the advantage of higher adhesion on wet substrates. Obtained maximal traction forces were about 30 times higher than friction forces of the shell surface dragged over the substrate. Neither wetting condition of the substrate nor surface roughness had considerable effect on the traction forces of snails. Therefore, the function of the hair‐like structures seems not to be related with snail locomotion and adhesion

Control Strategies of Gecko’s Toe in Response to Reduced Gravity

Shear-induced adhesion is one of the key properties for the gecko moving safely and quickly in a three-dimensional environment. The control strategies of such locomotion strongly relying on adhesion are still not well understood. In this study, we measured foot alignment and three-dimensional reaction forces of the single toes of the Tokay gecko running on the ground freely (gravity condition) and running in a situation where the gravity force was counterbalanced (reduced gravity condition). The forelimb rotated from the outward position to the front-facing position and the hindlimb rotated from the outward position to the rear-facing position, when running with balanced force, which indicated that the adhesive system was employed behaviorally through the modulation of the foot alignment. The toe was compressed and pulled in the gravity condition, but it was tensed and pulled in the reduced gravity condition. There was an approximately linear relationship between peak normal forces and the corresponding shear forces in both the reduced gravity condition (FN = −0.40FS − 0.008) and the gravity condition (FN = 2.70FS − 0.12). The footpad was compressed and pushed in the gravity condition, whereas it was tensed and pulled in the reduced gravity condition. There was an approximately linear relationship between peak normal forces and the corresponding shear forces in both the reduced gravity condition (FN = −0.39FS − 0.001) and in the gravity condition (FN = −2.80FS − 0.08). The shear-induced adhesion of the gecko footpad is controlled by the coupling of the normal force and shear forces: that is why in this system adhesion was shear-sensitive and friction was load-sensitive. Our measurements of single toe reaction forces also show that geckos control their footpad attachment using ‘toe rolling-in and gripping’ motion in both gravity and reduced gravity conditions

Subdivision of the neotropical Prisopodinae Brunner von Wattenwyl, 1893 based on features of tarsal attachment pads (Insecta, Phasmatodea)

The euplantulae of species from all five genera of the Prisopodinae Brunner von Wattenwyl, 1893 were examined using scanning electron microscopy with the aim to reveal the significance of attachment pads regarding their phylogenetic relationships. The split into the conventional two sister groups is supported by the two-lobed structure of the euplantulae with a smooth surface in the Prisopodini and a nubby surface microstructure in the Paraprisopodini. The two lineages are well distinguishable by this feature, as well as by the shape of the euplantulae themselves. The functional importance of the attachment pad surface features is discussed

Attachment devices and the tarsal gland of the bug Coreus marginatus (Hemiptera: Coreidae)

AbstractThe present ultrastructural investigation using scanning and transmission electron microscopy as well as light and fluorescence microscopy describes in detail the attachment devices and tarsal gland of the bug Coreus marginatus (L.) (Hemiptera: Coreidae). In particular, the fine structure of pulvilli reveals a ventral surface rich with pore channels, consistent with fluid emission, and a folded dorsal surface, which could be useful to enhance the pulvillus contact area during attachment to the substrate. The detailed description of the tarsal gland cells, whose structure is coherent with an active secretory function, allows us to consider the tarsal gland as the plausible candidate for the adhesive fluid production. Scolopidia strictly adhering to the gland cells are also described. On the basis of the fine structure of the tarsal gland, we hypothesise a fluid emission mechanism based on changes of the hydraulic pressure inside the gland, due to the unguitractor tendon movements. This mechanism could provide the fluid release based on compression of the pad and capillary suction, as demonstrated in other insects. The data here reported can contribute to understanding of insect adhesive fluid production, emission and control of its transport

Mechanical interaction of the egg parasitoid Anastatus bifasciatus (Hymenoptera: Eupelmidae) with artificial substrates and its host egg

Egg parasitoids play an important role in biological control of pest species attacking and killing their hosts at an early stage of their development. During the antagonistic coevolution with their hosts, egg parasitoids have developed a great ability to locate their host using chemical cues. A considerable amount of literature is available on this topic, while nothing is known about a possible adaptation of egg parasitoids to topography and mechanical properties of egg surface features and its shape when attaching to the host egg for oviposition. In the present investigation, the attachment ability of adults of both sexes of the egg parasitoid Anastatus bifasciatus (Hymenoptera: Eupelmidae) to artificial (polishing paper, flat glass, glass beads as dummies of the host egg) and natural surfaces (eggs of Halyomorpha halys and Nezara viridula, both Heteroptera: Pentatomidae), with different roughness and wettability, was measured using centrifugal force tester and traction force experiments. The parasitoid attachment devices and the egg surfaces were examined under cryo scanning electron microscope, wettability and roughness of natural and artificial substrates were characterised. We detected differences in the attachment devices and attachment ability of the two sexes. The collected data revealed a special ability of the female to attach to the eggs of the host species, thus suggesting an adaptation of the A. bifasciatus female to the surface features of the eggs during oviposition

Elasticity of the hair cover in air-retaining Salvinia surfaces

Immersed in water superhydrophobic surfaces (e.g., lotus) maintain thin temporary air films. In certain aquatic plants and animals, these films are thicker and more persistent. Floating ferns of the genus Salvinia show elaborated hierarchical superhydrophobic surface structures: a hairy cover of complex trichomes. In the case of S. molesta, they are eggbeater shaped and topped by hydrophilic tips, which pin the air–water interface and prevent rupture of contact. It has been proposed that these trichomes can oscillate with the air–water interface, when turbulences occur and thereby stabilize the air film. The deformability of such arrays of trichomes requires a certain elasticity of the structures. In this study, we determined the stiffness of the trichome coverage of S. molesta and three other Salvinia species. Our results confirm the elasticity of the trichome coverage in all investigated Salvinia species. We did not reveal a clear relationship between the time of air retention and stiffness of the trichome coverage, which means that the air retention function is additionally dependent on different parameters, e.g., the trichome shape and surface free energy. These data are not only interesting for Salvinia biology, but also important for the development of biomimetic air-retaining surfaces