Ultrahydrophobicity indicates a non-adhesive default state in gecko setae

Geckos may represent the world’s most demanding adhesives application. The adhesive setae on the toes of climbing geckos must adhere strongly yet avoid fouling or attachment at inappropriate times. We tested the hypothesis that gecko setae are non-adhesive in their unloaded default state by comparing the water droplet contact angle (θ) of isolated setal arrays to the smooth surface of eye spectacle scales of tokay geckos (Gekko gecko). At equilibrium, θ was 98.3 ± 3.4° in spectacle scales of live geckos and 93.3 ± 3.5° in isolated spectacles. Isolated setal arrays were ultrahydrophobic, with θ of 160.6 ± 1.3° (means ± SD). The difference in θ of setal arrays and smooth spectacles indicates a very low contact fraction. Using Cassie’s law of surface wettability, we infer that less than 6.6% of the surface of unloaded setae is solid and at least 93.4% is air space. We calculated that the contact fraction must increase from 6.6% in the unloaded state to 46% in the loaded state to account for previously measured values of adhesion. Thus gecko setae may be non-sticky by default because only a very small contact fraction is possible without mechanically deforming the setal array.     

Friction and adhesion in the tarsal and metatarsal scopulae of spiders

Friction and adhesion forces of the ventral surface of tarsi and metatarsi were measured in the bird spider Aphonopelma seemanni (Theraphosidae) and the hunting spider Cupiennius salei (Ctenidae). Adhesion measurements revealed no detectable attractive forces when the ventral surfaces of the leg segments were loaded and unloaded against the flat smooth glass surface. Strong friction anisotropy was observed: friction was considerably higher during sliding in the distal direction. Such anisotropy is explained by an anisotropic arrangement of microtrichia on setae: only the setal surface facing in the distal direction of the leg is covered by the microtrichia with spatula-like tips. When the leg is pushed, the spatula-shaped tips of microtrichia contact the substrate, whereas, when the leg is pulled over a surface, setae bend in the opposite direction and contact the substrate with their spatulae-lacking sides. In an additional series of experiments, it was shown that desiccation has an effect on the friction force. Presumably, drying of the legs results in reduction of the flexibility of the setae, microtrichia, spatulae, and underlying cuticle; this diminishes the ability to establish proper contact with the substrate and thus reduces the contact forces.     

Setal entanglement: an undescribed method of stridulation by a neotropical tarantula (Araneae: Theraphosidae)

The theraphosid spider Theraphosa leblondi (Latreille) produces a sibilant, hissing sound during defensive displays. This sound is produced using a previously undescribed method of stridulation: setal entanglement. The opposing surfaces of the femora of the pedipalps, first, and second pair of legs are clothed in unique setae which have their distal portion bearing hooks and a shaft clothed in filaments. Experimental ablations described here showed these plumose setae to be the site of sound production. Based on examination of the fine structure of these setae, we propose that the hissing sound is produced by the entanglement and pulling apart of the hooks at the ends of the setae on one leg surface and the long, plumose filaments on the median part of the setae on the opposing leg surface. Evidence presented suggests that this is a case of an acoustic aposematic display directed at vertebrate predators.     

Tarsal attachment devices of the southern green stink bug Nezara viridula (Heteroptera: Pentatomidae)

Based on analyses with cryo‐scanning and transmission electron microscopy, the present study reports on the morphology and ultrastructure of the attachment structures of the green stinkbug Nezara viridula L. (Heteroptera: Pentatomidae), a cosmopolitan pest of different crops in most areas of the world. In addition, the presence and distribution of large proportions of the elastic protein resilin in these structures was revealed by confocal laser scanning microscopy. The attachment structures of each leg comprise two sclerotised claws, a pair of smooth flexible pulvilli and a hairy adhesive pad located at the ventral side of the basitarsus. No sexual dimorphism is evident. Contact areas of resting individuals on a smooth surface show that N. viridula creates contact to the substrate with the ventral surface of (a) the distal portions of the pulvilli, (b) the setae of the hairy adhesive pad, (c) the two paraempodia representing mechanosensory setae, and (d) the tips of the claws. Each pulvillus is a sac‐like structure formed by complex cuticular layers that vary in their structure and resilin content. The dorsal side consists of sclerotised chitinous material, while the ventral cuticle consists mainly of resilin and shows a very thin epicuticle and a thick exocuticle. The setae of the hairy adhesive pad are pointed and socketed. They exhibit a pronounced longitudinal gradient in the material composition, with large proportions of resilin being present in the setal tips. In most of these setae, especially in those of the distal‐most part of the pad, also a transverse gradient in the material composition is visible.     

Contact behaviour of setal tips in the hairy attachment system of the fly Calliphora vicina (Diptera, Calliphoridae): a cryo-SEM approach

The fly Calliphora vicina (Diptera, Calliphoridae) bears attachment pads (pulvilli) covered with setae on their ventral sides. These structures enable attachment to smooth vertical surfaces and ceilings. The contact between the terminal setal tips (spatulae) and various substrates was visualised using various experimental techniques combined with conventional scanning electron microscopy (SEM) and cryo-SEM. The results show that the setal endplates are highly flexible structures that form contact with the surface by bending their tips in the distal direction. With conventional SEM, a comparison of partly attached endplates with unattached endplates demonstrated the presence of a distinct marginal bulge. As observed with cryo-SEM, the bulge continuously disappeared as a larger area of the endplate came into contact. Two explanations of this result are suggested. First, the volume between the bulge, the mid-part of the endplate and the substrate may be filled with a fluid secretion that is released into the contact area in the endplate region. Second, the flexible central part of the endplate may jump into contact with the substrate during contact formation.     

Flagellar setae of the second antennae in decapod shrimps: sexual dimorphism and possible role in detection of contact sex pheromones

Summary

The antenna 2 (antennal) flagella of decapod shrimps are chemotactile, and their setae are proposed as sensilla involved in recognition of females by males via a contact sex pheromone on the surface of the female. Male recognition of females receptive to mating occurs in many caridean species upon contact of male antennal flagella with the surface of a newly molted parturial female. The hypothesis of sexual dimorphism in the number and kind of setae on the antennal flagella of four caridean and one penaeid shrimp species was tested with setal counts and observations on setal morphology. Unique male antennal setae (“male-specific sensilla”) were not identified in any of the species investigated. However, the abundance of antennal setae was significantly greater in males than in breeding females in the palaemonid carideans Palemonetes pugio and Macrobrachium ohione. In the hippolytid caridean Thor manningi and alpheid caridean Alpheus normanni, no sexual dimorphism in setal abundance was demonstrated. In the penaeoid Rimapenaeus similis, males had a higher abundance of antennal setae than the larger breeding females but so did juvenile females, similar in size to males. The sexual dimorphism in antennal sensilla in the palaemonid species and its absence in A. normanni might be related to their different mating systems, but no such association is suggested for T. manningi and R. similis. Setal morphology suggestive of chemoreceptive function (a terminal pore) was observed in all species.     

Morphology and distribution of setae on the antennules of the Caribbean spiny lobster Panulirus argus reveal new types of bimodal chemo-mechanosensilla

This study describes the morphology and distribution of setae on the lateral and medial flagella of the antennules of the spiny lobster Panulirus argus in an effort to identify antennular chemoreceptors in addition to the well-studied aesthetasc chemosensilla. Setae were examined using light and electron microscopy, and their distribution on flagellar annuli was analyzed. We identified ten setal types based on external morphology: hooded, plumose, short setuled, long simple, medium simple, short simple, aesthetasc, guard, companion, and asymmetric setae, with the last four types being unique to the "tuft" located on the distal half of the lateral flagellum. The three setal types whose ultrastructure was examined--hooded, long simple, and medium simple setae--had characteristics of bimodal (chemo-mechanoreceptive) sensilla. The antennules have four distinct annular types based on their setal complement, as shown by cluster analysis. This basic distribution of non-tuft setal types is similar for both lateral and medial flagella. Annuli in the tuft region have tuft setal types superimposed on a basic organization of non-tuft setal types. These results show that the antennules possess a diverse set of setae, that these setae have a highly ordered arrangement on the antennules, that at least four (and probably many more) of these setal types are chemosensilla, and suggest that most antennular chemosensilla are bimodally sensitive.     

Prey finding behavior and mechanosensilla of larval Toxorhynchites brevipalpis Theobald (Diptera : Culicidae)

Behavioral experiments demonstrated that starved 3rd-instar Toxorhynchites brevipalpis (Diptera : Culicidae) will attack a glass probe in response to vibrations alone. Frequencies in the range of 80–200 Hz elicited 85% of the attacks. The observation that most attacks occurred while the larva was drifting forward towards the probe, substantiates that the predatory behavior of these organisms is basically opportunistic.The main setae on the thorax and abdomen arise from setal support plates and are of 4 general morphological types: dendritic, smooth, spinulate spiniform, and unbranched aciculate setae. The numbers and lengths of the setae on 3rd- and 4th-instar larvae are given. Each seta is innervated by a bipolar neuron with a tubular body in the dendrite, a characteristic of mechanosensilla. By virtue of their abundance and location, it seems probable that one or more types of these setae sense the water vibrations capable of eliciting an attack response.The tubular bodies of the 4 types of setae are somewhat unusual in that they lack electron-dense material, except near sites of attachment of the microtubules to the dendritic membrane, yet possess a large number of complexly arranged microtubules. Among the setal types, variations were observed in the prominence of microtubular attachment sites, proportion of tubules associated with the sites, and orientation of microtubules. In spinulate spiniform setae, the arrangement of microtubules varies within the same tubular body; peripherally most of the microtubules are in one direction with little convergence and have only one distinct attachment site, whereas medially they converge considerably resulting in 2 groups, each associated with its own well-defined site.The attachment sites, which presumably through linkage with the dendritic membrane provide a means of initiating depolarization, are associated with the distal ends of almost all of the microtubules. This suggests that in these tubular bodies depolarization may be initiated through mechanical force acting along the length of the microtubules, that is, stretching and/or compression.     

Biomechanism of adhesion in gecko setae

The study of the adhesion of millions of setae on the toes of geckos has been advanced in recent years with the emergence of new technology and measurement methods. The theory of the mechanism of adhesion by van der Waals forces is now accepted and broadly understood. However, this paper presents limitations of this theory and gives a new hypothesis of the biomechanism of gecko adhesion. The findings are obtained through measurements of the magnitude of the adhesion of setae under three different conditions, to show the close relationship between adhesion and status of the setae. They are reinforced by demonstrating two setal structures, follicle cells and hair, the former making the setae capable of producing bioelectrical charges, which play an important role in attachment and detachment processes. It is shown that the abundant muscular tissues at the base of the setae cells, which are controlled by peripheral nerves, are instrumental in producing the foot movement involved in attachment and detachment. Our study will further uncover the adhesion mechanism of geckos, and provide new ideas for designing and fabricating synthetic setae.     

Structure of dactyl sensilla in the kelp crab,Pugettia producta

In the kelp crab, Pugettia producta, flat plate setae cover all but the ventral surfaces of the walking leg dactyls. Dendrites enter the setal shaft located inside the plate superstructure, and extend to a region of the setal tip that contains a system of minute pores resembling the pore systems found in chemosensory sensilla of insects. Presumably, much of the chemosensitivity of the dactyls in the kelp crab is mediated by the plate setae. In the interior of the dactyl, supporting cells and the neurons innervating plate setae, other types of setae, and other presumptive sensilla form scolopidia. Large scolopidia, containing as many as 12 dendrites, appear to innervate some of the plate setae and also large ventral rodlike setae that might be chemosensory. Two of the dendrites of large scolopidia usually have more densely packed microtubules, longer ciliary axonemes, slightly larger rootlets, and dark A fibers with arms, characteristics indicative of mechanosensory function. Some dactyl setae, therefore, could be both mechanosensory and chemosensory. Small scolopidia containing two or three dendrites that exhibit mechanosensory characteristics appear to innervate small, rodlike setae, which presumably are strictly mechanosensory. The two types of structures located on the epicuticular cap, elliptical structures resembling campaniform sensilla and small cones in pits resembling CAP organs, appear to be dually innervated and presumably are mechanosensory, although other functions are possible. The internal positions of the scolopidia, together with the support afforded by an extracellular dendritic sheath, by the scolopale, and by desmosomelike and septate junctions, may serve to protect internal portions of setal dendrites, some of which appear to remain functional in nonmolting adults that have abraded setae.     

Subdigital and subcaudal microornamentation in chamaeleonidae—A comparative study

Locomotion on horizontal and vertical substrates requires effective attachment systems. In three clades of arboreal and rupicolous Iguanidae, Gekkota and Scincidae adhesive systems consisting of microscopic hair‐like structures (setae) have been evolved independently. Also the substrate contacting sites on toes and tails of chameleons (Chamaeleonidae) are covered with setae. In the present comparative scanning electron microscopy study, we show that representatives from the chamaeleonid genera Calumma, Chamaeleo, Furcifer, and Trioceros feature highly developed setae that are species‐specific and similar on their feet and tail. These 10 μm long, unbranched setae rather resemble those in anoline and scincid lizards than the larger and branched setae of certain gecko species. In contrast to the thin triangular tips of other lizards, all examined species of the genera Furcifer and Calumma and one of the five examined species of the genus Trioceros have spatulate tips. All other examined species of genera Trioceros and Chamaeleo bear setae with narrowed, fibrous tips. Unlike the setae of other lizards, chamaeleonid setal tips do not show any orientation along the axis of the toes, but they are flexible to bend in any direction. With these differences, the chameleon's unique microstructures on the zygodactylous feet and prehensile tail rather increase friction for arboreal locomotion than being a shear‐induced adhesive system as setal pads of other lizards. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

Integrative functional morphology of the gekkotan adhesive system (reptilia: gekkota)

Climbing assisted by adhesive subdigital pads in gekkotan lizardshas been the subject of intrigue and study for centuries. Manyhypotheses have been advanced to explain the mechanism of adhesion,and recently this phenomenon has been investigated at the levelof individual setae. The ability to isolate, manipulate andrecord adhesive forces from individual setae has provided newinsights, not only into the mechanism of attachment, but alsointo the physical orientation of these structures necessaryto establish attachment, maximize adhesive force, and effectsubsequent release. This, in turn, has enabled a reassessmentof the overall morphology and mode of operation of the adhesivesystem. Digital hyperextension has often been noted as a behavioralcharacteristic associated with the deployment of the gekkotanadhesive system—this is now understandable in the contextof setal attachment and release kinematics, and in the contextof the evolution of this pattern of digital movement from theprimitive pattern of saurian digital kinematics. The perpendicularand parallel preloads associated with setal attachment are nowreconcilable with other morphological aspects of the gekkotanadhesive system—the lateral digital tendon complex andthe vascular sinus network, respectively. Future investigationsof the integrated adhesive system will help to further elucidatethe interdependence of its structural and functional components.     

The setae of Austropotamobius pallipes (Crustacea: Astacidae)

The author presents here the results of an introductory study into the setal armature of the British species of crayfish, Austropotamobius pallipes (Lereboullet, 1858). Using an ordinary optical microscope, it is demonstrated that a considerable variety of setae exist on the body and appendages of A. pallipes. These setae are described, and their distribution indicated. The setae fall into two main groups:
(1) Setae with relatively thick walls and inconspicuous ampulla, and having no basal septum.
(2) Those setae with relatively thin walls, a well-developed ampulla and basal septum. Further subdivisions of these two groups are made on the bases of outgrowths and outlines of the setal wall. Finally, the significance of the distribution and the variation, is discussed in relation to crayfish behaviour and function, linking the observed morphological facts with the results of some current electro-physiological experiments performed on related decapods.     

Ultrastructure and development of forked and capillary setae in the polychaetes Orbinia bioreti and Orbinia latreillii (Annelida: Orbiniidae)

Abstract. Recent investigations into chaetogenesis of certain types of annelid setae provide important results for unravelling the phylogenetic relationships within several taxa of poly-chaetous annelids. This paper presents data on ultrastructure and development of 2 types of orbiniid setae. The analysis of the crenulate capillaries in Orbinia latreillii reveals a formation process which clearly differs from the development of Equisetum -like setae of lingulid bra-chiopods. For the investigation of forked setae, which up to now have been neglected in the discussion on the phylogenetic significance of annelid setae, notopodial setal sacs of O. latreillii and O. bioreti were studied by light- and electron microscopy. In the setal sacs, stages of forked setae are restricted to a dorsocaudal pouch, which represents the site of setal formation. The 2 diverging, stout tines of the fork bear spines on their inner margins, each of which is preformed by a single microvillus. After retraction of the microvilli, a characteristic pattern of the setal canals inside of the spines remains. The present study belongs to a series of comparative studies into chaetogenesis of forked setae. These special setae are also found in other orbiniid taxa as well as some paraonids, scalibregmatids, and nephtyids. Ultrastructural investigations into the development of these forked setae might suggest homology.     

Surface structures of the antennular flagella of the hermit crab Pagurus alaskensis (Benedict): A light and scanning electron microscopy study

The surface structures of the antennular flagella of Pagurus alaskensis are described in detail. Attention is directed towards the surface morphology of two types of possible sensilla: (1) exoskeletal pores (1.0–3.0 μm in diameter); (2) setae of various kinds. In addition, small (0.1–0.2 μm) pits occur in the exoskeleton which are not considered to be sensory in function. The exoskeletal pores are found at fairly specific locations on both the inner and outer flagella, particularly on the short segments of the outer flagella. Neither the inner nor the outer flagella are bilaterally symmetrical with respect to their setal armature. On the outer flagellum six groups of setae may be distinguished: lateralmesial; dorsal; ventral; accessory; aesthetasc; setae of the distal segment. On the inner flagellum setae of the mesial and lateral rows form distinctive groups. The morphology, orientation and locations of all the flagellar setae are defined and where possible the numbers of the various morphological types within the specific setal groups are given. It is noteworthy that many setal types have obvious apical pores and yet no pores could be found in the chemoreceptive aesthetasc setae. The functions of the various setae are discussed in relation to their topographical position and to existing electrophysiological and behavioral data. Some suggestions are made about future experiments to demonstrate the central connections of specific sensilla or groups of sensilla and to show their significance in the whole animal.     

Setal morphology of grooming appendages in the spider crab,Libinia dubia

In crustaceans, grooming behaviors decrease fouling by removing debris from the exoskeleton and body structures; these grooming behaviors improve respiration, sensory reception, movement, and reproduction. Setal morphologies of the following grooming appendages in the decapod crustacean spider crab Libinia dubia are examined including the first pereiopod (cheliped), first, second, and third maxillipeds (mouthparts), and first, second, and third epipods (internal extensions of the maxillipeds). The objective of this study was to describe setal morphologies of these grooming appendages and to elucidate possible functions and efficiencies of setal structures. Spider crabs are hypothesized to have elaborate setal morphologies, mainly for cleaning specialized decorating setae as well as for cleaning inside the gill chamber, which has a higher likelihood of becoming fouled compared to other decapods such as shrimps. Fourteen setal types are documented and included several varieties of serrate and pappose setae as well as simple setae, cuspidate setae, papposerrate setae, and canoe setae. Maxillipodal epipods in the gill chamber are free of fouling, suggesting the setation on the third maxilliped protopod has an efficient functional morphology in removing debris before water enters the gill chamber. Serrate setae may function for detangling and separating structures whereas pappose setae may function for fine detailed grooming. The cheliped is the only grooming appendage that can reach decorating setae and it contains only pappose setae; thus decorating setae is not likely groomed in a manner that would greatly decrease fouling. J. Morphol. 277:1045–1061, 2016. © 2016 Wiley Periodicals, Inc.     

Structure, mechanism and mechanical properties of pupal attachment in Greta oto (Lepidoptera: Nymphalidae: Ithomiinae)

The structure and mechanism of pupal attachment are described for the nymphalid Greta oto using electron microscopy, and high‐speed and time‐lapse photography. The cremaster is composed of a 3‐D array of hooked setae that engage with silk fibers spun into layers in a pad on the lower leaf surface. Each seta comprises a shaft terminating in a strongly curved hook, tipped with two lateral barbs. These hook into the silk pad, which is densely laid and built‐up in the central portion, flattening out peripherally. Time‐lapse photography showed that silk pad construction by fifth instar larvae is completed in four distinct spinning movements, producing a random fiber arrangement. It is proposed that such a fiber arrangement provides isotropic strength, giving greater flexibility to the attachment. The cremaster is attached to the silk pad by a series of lateral movements of the pupa's posterior abdomen. This movement, together with the shape of the setal hooks, is thought to be integral to the attachment process. Tensile loading tests showed that attachment failure is due to the breakage of the silk pad, which undergoes gradual destruction before releasing the cremaster. The attachment was found to have high tensile strength and fracture toughness, both of which suggest that it has evolved for the dual purpose of preventing the pupa being pulled from the leaf by a predator and preventing the attachment being weakened by wind, which causes the pupa to swing.