Biomedical subjects

Stanislav Gorb

Publications and source records attributed to Stanislav Gorb.

10 recordsLinked to original sources

PubMed2006

Material structure, stiffness, and adhesion: why attachment pads of the grasshopper (Tettigonia viridissima) adhere more strongly than those of the locust (Locusta migratoria) (Insecta: Orthoptera).

The morphology, ultrastructure, effective elastic modulus, and adhesive properties of two different smooth-type attachment pads were studied in two orthopteran species. Tettigonia viridissima (Ensifera) and Locusta migratoria (Caelifera) have a similar structural organization of their attachment pads. They both possess a flexible exocuticle, where the cuticular fibrils are fused into relatively large rods oriented at an angle to the surface. The compliant material of the pad contributes to the contact formation with the substrate. However, the pad material structure was found to be different in these two species. L. migratoria pads bear a thick sub-superficial layer, as well as a higher density of rods. The indentation experiments showed a higher effective elastic modulus and a lower work of adhesion for L. migratoria pads. When the indentations were made at different depths, a higher effective elastic modulus was revealed at lower indentation depths in both species. This effect is explained by the higher stiffness of the superficial pad layer. The obtained results demonstrate a clear correlation between density of the fibres, thickness of the superficial layer, compliance of the pad, and its adhesive properties. Such material structures and properties may be dependent on the preferred environment of each species.

Adhesiveness↗

PubMed2006

Courtship dances in the flies of the genus Lispe (Diptera: Muscidae): from the fly's viewpoint.

Two predatory fly species, Lispe consanguinea Loew, 1858 and L. tentaculata DeGeer, 1776, inhabit the supralittoral zone at the shore of a fresh-water reservoir. Both species look alike and possess similar "badges," reflective concave silvery scales on the face. Flies occupy different lek habitats. Males of the first species patrol the bare wet sand on the beach just above the surf. Males of the second species reside on the more textured heaps of algae and stones. Courtship and aggressive behaviour of males was video-recorded and analysed frame by frame. Visual stimuli provided by the conspecific partner were computed in the body-fixed space of a fly observer. Males of L. consanguinea perform long pedestrian dances of pendulating circular arcs (frequency 2 s(-1), median radius 2.5 cm, linear velocity 0.130 m/s). Right and left side runs are equally probable. Circular runs are interrupted by standby intervals of average duration 0.35 s. The female views the male as a target covering 2 by 2 ommatidia, moving abruptly with the angular velocity over 200 degrees/s in a horizontal direction down the path of about 50 degrees till the next standpoint. Dancing is evenly distributed around the female. On the contrary, the male fixates the image of the female within the narrow front sector (median +/-10 degrees); the target in his view has 6-7 times less angular velocity and angular span of oscillations, and its image in profile overlays 6-8 by 2 ommatidia. If the female walks, the male combines tracking with voluntary circular dances. Rival males circle about one another at a distance shorter than 15 mm, but not in close contact. Males of L. tentaculata are capable of similar circular courting dances, but do so rarely. Usually they try to mount any partner immediately. In the latter species, male combat consists of fierce wrestling. Flies of both species often walk sideward and observe the partner not in front but at the side.

Aggression↗

PubMed2005

Adhesion design maps for bio-inspired attachment systems.

Fibrous surface structures can improve the adhesion of objects to other surfaces. Animals, such as flies and geckos, take advantage of this principle by developing "hairy" contact structures which ensure controlled and repeatable adhesion and detachment. Mathematical models for fiber adhesion predict pronounced dependencies of contact performance on the geometry and the elastic properties of the fibers. In this paper the limits of such contacts imposed by fiber strength, fiber condensation, compliance, and ideal contact strength are modeled for spherical contact tips. Based on this, we introduce the concept of "adhesion design maps" which visualize the predicted mechanical behavior. The maps are useful for understanding biological systems and for guiding experimentation to achieve optimum artificial contacts.

Adhesiveness↗

PubMed2005

Adhesion of echinoderm tube feet to rough surfaces.

Echinoderms attach strongly and temporarily to the substratum by means of specialized organs, the podia or tube feet. The latter consist of a basal extensible cylinder, the stem, which bears an apical flattened disc. The disc repeatedly attaches to and detaches from the substratum through adhesive and de-adhesive secretions. In their activities, echinoderms have to cope with substrata of varying degrees of roughness as well as with changing hydrodynamic conditions, and therefore their tube feet must adapt their attachment strength to these environmental constraints. This study is the first attempt to evaluate the influence of substratum roughness on the temporary adhesion of echinoderm tube feet and to investigate the material properties of their contact surface. It was demonstrated that tube foot discs are very soft (E-modulus of 6.0 and 8.1 kPa for sea stars and sea urchins, respectively), have viscoelastic properties and adapt their surface to the substratum profile. They also show increased adhesion on a rough substratum in comparison to its smooth counterpart, which is due mostly to an increase in the geometrical area of contact between the disc and the surface. Tenacity (force per unit area) increases with roughness [e.g. 0.18 and 0.34 MPa on smooth polymethyl-methacrylate (PMMA), 0.21 and 0.47 MPa on rough PMMA for sea stars and sea urchins, respectively] if only the projected surface area of the adhesive footprint is considered. However, if this tenacity is corrected to take into account the actual substratum 3-D profile, surface roughness no longer influences significantly the corrected adhesion strength (e.g. 0.18 and 0.34 MPa on smooth PMMA, 0.19 and 0.42 MPa on rough PMMA for sea stars and sea urchins, respectively). It can be hypothesized that, under slow self-imposed forces, disc material behaves viscously to adapt to substratum roughness while the adhesive fills out only very small surface irregularities (in the nanometer range). It is deposited as a thin film ideal for generation of strong adhesion. Under short pulses of wave-generated forces, attached discs probably behave elastically, distributing the stress along the entire contact area, in order to avoid crack generation and thus precluding disc peeling and tube foot detachment.

Adhesiveness↗

PubMed2004

Adhesion forces measured at the level of a terminal plate of the fly's seta.

The attachment pads of fly legs are covered with setae, each ending in small terminal plates coated with secretory fluid. A cluster of these terminal plates contacting a substrate surface generates strong attractive forces that hold the insect on smooth surfaces. Previous research assumed that cohesive forces and molecular adhesion were involved in the fly attachment mechanism. The main elements that contribute to the overall attachment force, however, remained unknown. Multiple local force-volume measurements were performed on individual terminal plates by using atomic force microscopy. It was shown that the geometry of a single terminal plate had a higher border and considerably lower centre. Local adhesion was approximately twice as strong in the centre of the plate as on its border. Adhesion of fly footprints on a glass surface, recorded within 20 min after preparation, was similar to adhesion in the centre of a single attachment pad. Adhesion strongly decreased with decreasing volume of footprint fluid, indicating that the layer of pad secretion covering the terminal plates is crucial for the generation of a strong attractive force. Our data provide the first direct evidence that, in addition to Van der Waals and Coulomb forces, attractive capillary forces, mediated by pad secretion, are a critical factor in the fly's attachment mechanism.

Adhesiveness↗

PubMed2004

Structure and properties of the glandular surface in the digestive zone of the pitcher in the carnivorous plant Nepenthes ventrata and its role in insect trapping and retention.

Carnivorous plants of the genus Nepenthes grow in nutrient-poor habitats and have evolved specialised trapping organs, known as pitchers. These are composed of different surface zones serving the functions of attraction, capture and digestion of insects, which represent a main source of nitrogen. To investigate the role of the glandular digestive zone in the trapping mechanism of the pitcher, structural, mechanical and physico-chemical studies were applied to N. ventrata and combined with insect behavioural experiments. It was found that the glandular surface is microscopically rough since it is regularly structured with multicellular glands situated in epidermal depressions. The presence of downward-directed 'hoods' over the upper part of glands and sloped depressions in the proximal direction of the pitcher causes a marked anisotropy of the surface. The glandular zone surface is composed of relatively stiff material (Young's modulus, 637.19+/-213.44 kPa). It is not homogeneous, in terms of adhesive properties, and contains numerous areas without adhesion as well as adhesive areas differing greatly in tenacity values (range, 1.39-28.24 kPa). The surface is readily wettable with water (contact angle, 31.9-36.0 degrees C) and has a high surface free energy (56.84-61.93 mN m(-1)) with a relatively high polar component (33.09-52.70 mN m(-1)). To examine the effect of the glandular secretion on attachment systems of insects having hairy and smooth adhesive pads, forces generated on different surfaces by Calliphora vicina flies and Pyrrhocoris apterus bugs, respectively, were measured. Flies attached equally well to both fresh and air-dried glandular surfaces whereas bugs generated a significantly lower force on the fresh glandular surface compared with the air-dried one. It is assumed that the contribution of the glandular surface to insect retention, due to its effect on insect attachment, differs depending on insect weight and the type of insect attachment system. Surface anisotropy does not facilitate effective claw interlocking so that insects possessing only claws are probably not able to cling to the glandular surface. However, stiffness of the pitcher wall material in the digestive zone can provide claw clinging via punching of the pitcher wall by claws. Small insects lacking pads may use adhesive areas on the plant surface to attach themselves, but such solitary points with very strong adhesion possibly impede their overall locomotion and chance of escape. Pad-bearing insects are presumably able to attach to smooth parts of the glandular surface located between glands. High free surface energy of the plant substrate may promote adhesion. Gland secretion may decrease attachment ability in insects with smooth adhesive pads but not influence attachment of insects with hairy attachment systems.

Adhesiveness↗

PubMed2003

From micro to nano contacts in biological attachment devices.

Animals with widely varying body weight, such as flies, spiders, and geckos, can adhere to and move along vertical walls and even ceilings. This ability is caused by very efficient attachment mechanisms in which patterned surface structures interact with the profile of the substrate. An extensive microscopic study has shown a strong inverse scaling effect in these attachment devices. Whereas microm dimensions of the terminal elements of the setae are sufficient for flies and beetles, geckos must resort to sub-microm devices to ensure adhesion. This general trend is quantitatively explained by applying the principles of contact mechanics, according to which splitting up the contact into finer subcontacts increases adhesion. This principle is widely spread in design of natural adhesive systems and may also be transferred into practical applications.

Animals↗

PubMed2003

Tarsal movements in flies during leg attachment and detachment on a smooth substrate.

In order to understand the attachment mechanism of flies, it is important to clarify the question of how the adhesive pad (pulvillus) builds and breaks the contact with the substrate. By using normal and high-speed video recordings, the present study revealed that pulvilli are positioned on the surface in a particular way. The pulvilli are apparently loaded or pressed upon the substrate after leg contact, as evidenced by splaying of the claws. Detachment of pulvilli from the substrate may be achieved in four different modes depending on the leg (fore-, mid- or hindleg): pulling, shifting, twisting, and lifting. Lifting is the only detachment mode depending on the claws' action. Kinematics of the tarsal chain is studied in leg preparations, in which the tendon of the claw flexor muscle was pulled by tweezers and video recorded. The morphological background of tarsal movements during attachment and detachment is studied by scanning electron microscopy, fluorescent microscopy, and bright field light microscopy followed by serial semithin sectioning of pretarsal structures. Several resilin-bearing springs are involved in the recoil of the tarsal segments to their initial position, when the tendon is released after pull.

Animals↗

PubMed2002

Contact behaviour of tenent setae in attachment pads of the blowfly Calliphora vicina (Diptera, Calliphoridae).

To enable strong attachment forces between pad and substrata, a high proximity between contacting surfaces is required. One of the mechanisms, which can provide an intimate contact of solids, is a high flexibility of both materials. It has been previously presumed that setae of hairy attachment pads of insects are composed of flexible cuticle, and are able to replicate the surface profile. The aim of this work was to visualise the contact behaviour of the setae by freezing-substitution technique to understand setal mechanics while adhering to a smooth surface. This approach revealed considerable differences in the area of the setal tips between contacting and non-contacting pulvilli. Based on the assumption that setae behave like a spring pushed by the tip, a spring constant of 1.31 N m(-1) was calculated from direct measurements of single setae by atomic force microscopy. In order to explain the relationship between the behaviour of the attachment setae at a microscale and leg movements, high-speed video recordings were made of walking flies. This data show that some proximal movement of the leg is present during contact formation with the substrate.

Animal Structures↗

PubMed2002

Arcus as a tensegrity structure in the arolium of wasps (Hymenoptera: Vespidae).

The unfolding of the hymenopteran attachment pad (arolium) may be achieved in two ways, hydraulic and mechanical. The first was confirmed in experiments with pressure applied to more proximal leg parts and on immersion in hypotonic solutions. Presumably, this way of unfolding does not play an important role for a living hornet. Mechanical unfolding was studied experimentally with the aid of a micromanipulator pulling the tendon of the musculus retractor unguis. Ablation experiments on different parts of the arolium indicated that the arcus is the most crucial element for mechanical unfolding. The shape of the arcus in closed and open conditions was measured using a 3D measurement microscope and reconstructed by means of 3D computer graphics. The arcus coils up upon being freed from the arolium tissues, and coils up even more after immersion into a 10% aqueous solution of NaOH. Geometrical models of the arcus are proposed, from which the rotational moment of elasticity is derived. Conformations and deformations of the arcus are quantified in order to explain its role in the folding and unfolding processes of the arolium. The diversity of approaches supports the idea that the arcus is a prestressed (tensegrity) structure providing immediate, soft, and graded transmission of forces during folding and unfolding action of the arolium.

Journal Article↗