Studying the deformation of arachnid slit sensilla by a fracture mechanical approach

Slit sensilla are sensory organs which measure strains in the exoskeleton of arachnids. They occur as isolated slits, in loose groups and in close parallel arrangements known as lyriform organs or compound slit sensilla. The deformations of the slits' faces induced by far-field strains acting on groups of slits are studied using Kachanov's analytical approximations for the opening displacements of cracks, a method developed within the framework of fracture mechanics. The accuracy of the approach is assessed by comparisons with results obtained by finite element analysis. The limits of its applicability to slit sensilla are found to be reached when the lateral spacing between interacting slits is less than half their length, i.e., the method is suitable for studying single slits and loose groups but not lyriform organs. The influence of a number of geometrical parameters of slit sensilla on the deformation patterns of the faces of parallel slits in generic arrangements is studied, viz., spacing between slits, longitudinal shifts between slits, and slit length. The results are presented as opening distances along the length of the cracks and in terms of normalized diagrams that relate the opening distances at mid-length of the slits to the geometrical parameters. In addition, Kachanov's method is used to find a set of slit lengths that give rise to prescribed opening distances.     

Proprioceptors and sensory nerves in the legs of a spider,Cupiennius salei (Arachnida,Araneida)

Summary Retrograde CoS-impregnation was used to trace and map the course of sensory nerves and the distribution and innervation of the various proprioceptor types in all leg segments of Cupiennius salei, a Ctenid spider.1. Sensory nerve branches. In both the tibia and femur, axons of all proprioceptor types ascend in just two lateral nerves which do not merge with the main leg nerve until they reach the next proximal joint region. In the short segments — coxa, trochanter, patella, and tarsus — axons of the internal joint receptors often run separately from those of the other sensilla. Axons of the large lyriform slit sense organ at the dorsal metatarsus and of the trichobothria join with only a few hair axons and form their own nerve branches (Figs. 1, 2, 3).2. Proprioceptors. Each of the seven leg joints is supplied with at least one set of the well-known internal joint receptors, slit sensilla (single slits and lyriform organs), and long cuticular hairs. In addition, we found previously unnoticed hair plates on both sides of the coxa, near the prosoma/coxa joint; they are deflected by the articular membrane during joint movements (Fig. 4).3. Sensory cells and innervation. CoS-impregnation shows that each slit of the slit sense organs — be it a single slit or several slits in a lyriform organ — is innervated by two bipolar sensory cells (Fig. 6). We also confirm previous reports of multiple innervation in the internal joint receptors and in the long joint hairs and cuticular spines.Most of the ascending nerve branches run just beneath the cuticle for at least a short distance (Fig. 5); hence they are convenient sites for electrophysiological recordings of sensory activity even in freely walking spiders.     

Ein atlas der spaltsinnesorgane von Cupiennius salei keys. Chelicerata (Araneae)

To facilitate further physiological investigation, a survey was undertaken of all the slit sense organs to be found on the body of the spider Cupiennius salei. We counted and mapped more than 3 000 sensory slits in the cuticle about half of which are combined to small groups of up to 29 slits forming compound or lyriform organs.     

Slit sense organs and kinesthetic orientation

Summary In the hunting spider Cupiennius salei Keys, kinesthetic orientation towards a catching site from which it has previously been chased away is observed. This ability strongly depends on the lyriform slit sense organs found on femur and tibia of the walking legs. The animals miss the original catching site, if these organs are destroyed on all legs. The mean angular deviation of the starting angles of the returns increases significantly as compared with intact spiders (P<0.005). Also, the directions of the mean vectors of the starting angles change (P<0.05).Supported by a grant of the Deutsche Forschungsgemeinschaft.     

Central nervous projections of mechanoreceptors in the spider Cupiennius salei Keys.

Summary The basic organization of sensory projections in the suboesophageal central nervous system of a spider (Cupiennius salei Keys.) was analyzed with anterograde cobalt fills and a modified Golgi rapid method. The projections of three lyriform slit sense organs and of tactile hairs located proximally on the legs are described and related to central nerve tracts. There are five main longitudinal sensory tracts in the central region of the suboesophageal nervous mass arranged one above the other. Whereas the three dorsal ones contain fibers from the lyriform organs, the two ventral ones contain axons from the hair receptors. Axons from all three lyriform organs have typical shapes and widely arborizing ipsilateral intersegmental branches and a few contralateral ones. The terminal branches of the afferent projections from identical lyriform organs on each leg form characteristic longitudinal pathways, typical of each organ: U-shaped, O-shaped, or two parallel bundles. The terminations of the hair sensilla are ipsilateral and intersegmental. Two large bilaterally arranged longitudinal sensory association tracts receive inputs from all legs including the dense arborizations from tactile hairs, lyriform organs, and other sense organs. These tracts may serve as important integrating neuropils of the suboesophageal central nervous system.     

Detection of vibrations in sand by tarsal sense organs of the nocturnal scorpion,Paruroctonus mesaensis

Summary The scorpionParuroctonus mesaensis locates prey by orienting to substrate vibrations produced by movements of the prey in sand. At the end of each walking leg of this scorpion there are two sense organs, the basitarsal compound slit sensillum and tarsal sensory hairs (Figs. 1, 3) that are excited by substrate vibrations conducted through sand. The slit sensilla appear to be most sensitive to surface (Rayleigh) waves while the tarsal sensory hairs respond best to compressional waves (Fig. 7). Both mechanoreceptors were activated by nearby disturbances of the substrate (Fig. 6) but only the slit sensilla responded to insects moving more than 15 cm away. Both receptors are highly sensitive to small amplitude (less than 10 Å) mechanical stimuli applied to the tarsus (Fig. 5).Behavioral studies of scorpions with ablated sense organs (Fig. 2) indicate that the basitarsal compound slit sensilla are necessary for determining vibration source direction.Abbreviation BCSS basitarsal compound slit sensillum (a) Supported by PHS Environmental Science and Regents Intern Fellowships (PB), and by intramural research funds from the University of California (RDF)     

Finite element modeling of arachnid slit sensilla: II. Actual lyriform organs and the face deformations of the individual slits

Arachnid slit sensilla respond to minute strains in the exoskeleton. After having applied Finite Element (FE) analysis to simplified arrays of five straight slits (Hößl et al. J Comp Physiol A 193:445–459, 2007) we now present a computational study of the effects of more subtle natural variations in geometry, number and arrangement of slits on the slit face deformations. Our simulations show that even minor variations in these parameters can substantially influence a slit’s directional response. Using white-light interferometric measurements of the surface deformations of a lyriform organ, it is shown that planar FE models are capable of predicting the principal characteristics of the mechanical responses. The magnitudes of the measured and calculated slit face deformations are in good agreement. At threshold, they measure between 1.7 and 43 nm. In a lyriform organ and a closely positioned loose group of slits, the detectable range of loads increases to approximately 3.5 times the range of the lyriform organ alone. Stress concentration factors (up to ca. 29) found in the vicinity of the slits were evaluated from the models. They are mitigated due to local thickening of the exocuticle and the arrangement of the chitinous microfibers that prevents the formation of cracks under physiological loading conditions.     

Proprioceptor distribution and control of a muscle reflex in the tibia of spider legs

In spiders, retrograde cobalt staining was used to clarify the distribution and detailed innervation of the three types of proprioceptors in the tibio-metatarsal leg joint: internal joint receptors, lyriform slit sense organs, and cuticular spines and hairs. The axons of all these receptors run in just two lateral, ascending nerves, which had previously been associated only with the internal receptors. Each nerve contains several hundred axons ranging in diameter from 0.1 micron to ca. 10 micron. Each slit of the four tibial lyriform organs is innervated by two bipolar sensory neurons. The lateral nerves are entirely sensory and run just beneath the cuticle, a convenient site for electrophysiological recording. We demonstrate simultaneous nerve and muscle recordings from intact spiders; these, in combination with selective sensory ablations, show that a resistance reflex in the flexor metatarsi muscles is elicited by internal joint-receptor units.     

Finite element modeling of arachnid slit sensilla—I. The mechanical significance of different slit arrays

Arachnid strain sensitive slit sensilla are elongated openings in the cuticle with aspect ratios (slit length l / slit width b) of up to 100. Planar Finite Element (FE) models are used to calculate the relative slit face displacements, D _c, at the centers of single slits and of arrangements of mechanically interacting slits under uni-axial compressive far-field loads. Our main objective is to quantitatively study the role of the following geometrical parameters in stimulus transformation: aspect ratio, slit shape, geometry of the slits‘ centerlines, load direction, lateral distance S, longitudinal shift λ, and difference in slit length Δl between neighboring slits. Slit face displacements are primarily sensitive to slit length and load direction but little affected by aspect ratios between 20 and 100. In stacks of five parallel slits at lateral distances typical of lyriform organs (S = 0.03 l) the longitudinal shift λ substantially influences slit compression. A change of λ from 0 to 0.85 l causes changes of up to 420% in D _c. Even minor morphological variations in the arrangements can substantially influence the stimulus transformation. The site of transduction in real slit sensilla does not always coincide with the position of maximum slit compression predicted by simplified models. An erratum to this article can be found at     

Central nervous projection patterns of trichobothria and other cuticular sensilla in the wandering spider Cupiennius salei (Arachnida,Araneae)

Summary Central projections of mechano-and chemoreceptors on the legs and pedipalps of the wandering spider Cupiennius salei were traced by anterograde cobalt fills. The primary afferent fibres from trichobothria, tactile hairs, lyriform organs and contact chemoreceptive hairs enter the leg ganglia and pedipalpal ganglia ventrally. On their way through these ganglia there is very little arborization. The main areas of arborization are in the sensory longitudinal tracts in the suboesophageal nervous mass. The central projections of all mechano-and chemoreceptors examined show somatotopic organization. Sensilla located proximally on the legs are represented in dorsally located sensory longitudinal tracts, whereas those located on distal leg segments enter more ventral tracts. The afferent fibres of receptors of identifical modality on a specific segment of all legs and of the pedipalps overlap in the same tracts. No indication for a tonotopic arrangement of the trichobothrial afferences was found, which might have been associated with the mechanical frequency tuning of the trichobothria known from other experiments. The convergence of the projections of different types of receptors in the sensory longitudinal tracts is considered to be an anatomical basis for their functional interaction in behaviour. Both the convergence of the projections of receptors from the same segment of different legs and the somatotopy are connectivity patterns possibly associated with the orientation of the spiders towards mechanical or chemical cues.     

The sensory equipment of a spider – A morphological survey of different types of sensillum in both sexes of Argiope bruennichi (Araneae,Araneidae)

Spiders show a wide range of sensory capabilities as evidenced by behavioural observations. Accordingly, spiders possess diverse sensory structures like mechano-, hygro-, thermo- or chemoreceptive sensilla. As to chemoreceptive structures, only trichoid tip-pore sensilla were found so far that were tested for gustation. That spiders are also able to receive airborne signals is corroborated by numerous behavioural experiments but the responsible structures have not been determined yet. Here, we provide sensilla distribution maps of pedipalps and walking legs of both sexes of the wasp spider Argiope bruennichi whose biology and mating system is well explored. By means of scanning electron microscopy, we scrutinized whether there is in fact only one type of trichoid pore sensillum and if so, if there are deviations in the outer structure of the tip-pore sensilla depending on their position on the body. We also describe the external structure and distribution of slit sense organs, trichobothria and tarsal organs. Our study shows that all four sensillum types occur on pedipalps and walking legs of both sexes. As to chemosensory organs, only tip-pore sensilla were found, suggesting that this sensillum type is used for both gustation and olfaction. The highest numbers of tip-pore sensilla were observed on metatarsi and tarsi of the first two walking legs. Mechanosensitive slit sense organs occur as single slit sensilla in rows along all podomers or as lyriform organs next to the joints. The mechanosensitive trichobothria occur on the basal part of tibiae and metatarsi. Tarsal organs occur on the dorsal side of all tarsi and the male cymbium. The distribution maps of the sensilla are the starting point for further exploration of internal, morphological differences of the sensilla from different regions on the body. Cryptic anatomical differences might be linked to functional differences that can be explored in combination with electrophysiological analyses. Consequently, the maps will help to elucidate the sensory world of spiders.     

Lyriform slit sense organs on the pedipalps and spinnerets of spiders

Lyriform slits sense organs (LSSO) are a precise assembly of stress detecting cuticular slit sensilla found on the appendages of arachnids. While these structures on the legs of the wandering spiderCupennius salei are well studied in terms of morphology, function and contribution to behaviour, their distribution on pedipalps and spinnerets of spiders is not well explored. A study was therefore carried out to observe the distribution of LSSO on pedipalps and spinnerets of some spider species. Haplogyne spiders belonging to familyPholcidae have a simple complement of LSSOs represented by one or two LSSOs on their femur. The entelegyne spiders possess a complex assembly of LSSOs on the distal segments of their pedipalps. Various types of LSSOs are found on the pedipalps indicating a capacity for analysis of complex cuticular stress. It is suggested that the complexity of LSSOs on pedipalps of entelegyne spiders relates to courtship and spermatophore transfer and may help in reproductive isolation. Lack of LSSOs on the distal segments of pedipalps leads us to infer that unlike legs, pedipalps are less likely to receive vibratory input through their distal segments. Spinnerets have a relatively simple complement of LSSOs. One LSSO is found only on anterior spinnerets and it is a common feature observed among spiders, irrespective of the variations in web building behaviour. The orb-weaving araneidArgiope pulchella, however, has two LSSOs on the anterior spinneret. As non-web builders and orb weavers do not differ markedly in terms of LSSOs on the spinnerets and LSSOs are simple in nature (type A), it is likely that spinning and weaving are not largely regulated by sensory input from LSSOs on the spinnerets.     

From stress and strain to spikes: mechanotransduction in spider slit sensilla

This review focuses on the structure and function of a single mechanoreceptor organ in the cuticle of spiders. Knowledge emerging from the study of this organ promises to yield general principles that can be applied to mechanosensation in a wide range of animal systems. The lyriform slit sense organ on the antero-lateral leg patella of the spider Cupiennius salei is unusual in possessing large sensory neurons, whose cell bodies are close to the sites of sensory transduction, and accessible to intracellular recording during mechanotransduction. This situation, combined with recent technical developments, has made it possible to observe and experiment with all the major stages of mechanosensation. Important findings include the approximate size, number and ionic selectivity of the ion channels responsible for mechanotransduction, the types of voltage-activated ion channels responsible for action potential encoding, and the mechanisms controlling the dynamic properties of transduction and encoding. Most recently, a complex efferent system for peripheral modulation of mechanosensation has been discovered and partially characterized. Much remains to be learned about mechanosensation, but the lyriform slit sense organ system continues to offer important opportunities to advance our understanding of this crucial sense.     

Species richness and biogeography of non-acarine arachnids in Namibia

A total of 821 species, 296 genera, and 69 families of non-acarine arachnids (Araneae, Solifugae, Scorpiones, Pseudoscorpiones, Opiliones and Amblypygi) are presently known from Namibia. Patterns of spider, solifuge and scorpion species richness and endemism are summarized relative to the 14 major Namibian types of vegetation. Spiders are most speciose in the higher rainfall areas, solifuges in the dry areas, and scorpions in rocky areas. Namibia probably has the world's greatest diversity of solifuges. Many arachnids endemic to Namibia are associated with the Namib Desert sand dunes.     

Histologic Structure of the Harvestman Leiobunum japonicum (Arachnida: Opiliones: Sclerosomatidae)

Opiliones are an order of arachnids commonly known as harvestmen. Although they belong to the class of arachnids, harvestmen are not spiders. Harvestmen are well known for their exceptionally long legs compared to body size, and the made up of two main parts, cephalothorax and abdomen. Although over 6,400 species of harvestmen have been discovered worldwide, the research on these arachnids is still an unexplored field comparing to spiders or insects. The Harvestmen is the Arachnids disregarded to Korean researchers. Therefore, in Korea, it doesn't become nearly the investigation of the harvestmen. Histological data about the harvestmen tries to be provided through the research of the internal structure of the harvestmen. The study of structures of the opiliones was made with the histological image and it focused mainly on the eye, leg, and genital organs.     

Model studies on the mechanical significance of grouping in compound spider slit sensilla (Chelicerata,Araneida)

Summary The mechanical implications of various types of slit arrangements found among the strain-sensitive slit sensilla in the arachnid exoskeleton (Fig. 3) were studied by measuring the deformation of model slits, cut into plastic discs, under static load applied in the plane of the disc and from varying directions (Figs. 1, 2).1. Close parallel, lyriform arrangements. Compression of slits (adequate stimulus) reaches much higher values than dilatation. It is highest with the load direction at right angle to the slit axes. Also, in the majority of slits the range of load angles resulting in compression is considerably larger than that leading to dilatation. Length distribution and lateral shift of slits in the models have a pronounced effect on slit deformability (Figs. 4-5): (a) In the oblique bar arrangement with slits of equal length and regular lateral shift (Fig. 4A) deformation of all slits is very similar at all load directions. In all slits compression results from a range of load angles larger than 120°. (b) In arrangements with a regular increase in slit length and a triangular outline shape deformability differs greatly among the slits at all load angles (Fig. 4B). (c) The slit configuration with a heartshaped outline (Fig. 4C) is peculiar for the large spread of load angles at which the compression of the different slits is highest. — These properties recommend different arrangements for the solution of different strain measuring problems, with for instance, the particular need of a wide angular working range (arrangement a), of a large spectrum of absolute sensitivities (b), or of the analysis of load direction (c).2. Angle and distance between slits. Due to the mechanical directionality inherent in an elongated slit the divergence of slit axes within a group of slits is likely to indicate the importance of the analysis of strain direction (Fig. 6). The mechanical interaction between closely neighbouring slits decreases with their distance from each other. In a parallel arrangement of equally long slits it disappears if the distance is about 1.5 times the slit length (Fig. 7).3. Aiming towards a mechanical model which would explain the complex deformation found in a lyriform organ, we consider the outline of the organ as a hole traversed by beams of material. Slit deformation can be calculated from the elastic lines of the beams which separate the slits and information drawn from photoelastic experiments (Figs. 8-11).     

Intracellular recording from a spider vibration receptor

The present study introduces a new preparation of a spider vibration receptor that allows intracellular recording of responses to natural mechanical or electrical stimulation of the associated mechanoreceptor cells. The spider vibration receptor is a lyriform slit sense organ made up of 21 cuticular slits located on the distal end of the metatarsus of each walking leg. The organ is stimulated when the tarsus receives substrate vibrations, which it transmits to the organ’s cuticular structures, reducing the displacement to about one tenth due to geometrical reasons. Current clamp recording was used to record action potentials generated by electrical or mechanical stimuli. Square pulse stimulation identified two groups of sensory cells, the first being single-spike cells which generated only one or two action potentials and the second being multi-spike cells which produced bursts of action potentials. When the more natural mechanical sinusoidal stimulation was applied, differences in adaptation rate between the two cell types remained. In agreement with prior extracellular recordings, both cell types showed a decrease in the threshold tarsus deflection with increasing stimulus frequency. Off-responses to mechanical stimuli have also been seen in the metatarsal organ for the first time.     

Spider mechanoreceptors

Spiders have highly developed mechanosensory systems, some of which provide access to forms of stimulation alien to our own sensations. Studies of hair-shaped air movement detectors (trichobothria) and tactile sensors have uncovered an outstanding refinement of the processes of stimulus uptake and stimulus transformation, which reflect details of both stimulus physics and behavioral significance. They also emphasize the potential contained in the seemingly simple Bauplan of arthropod cuticular hairs. Embedded into the spider exoskeleton are several thousands of strain detectors (slit sensilla) measuring compressive exoskeletal strains induced by various forms of loads and forces. A compound slit sensillum (lyriform organ) on the leg has become an important model system for studies of mechanoreceptor primary processes at the cellular and membrane level.     

A Spider's Vibration Receptor: Its Anatomy and Physiology

The common house spider, Achaearanea tepidariorum, has a sensitivevibration receptor on each of its eight legs. This receptoris the tarso-metatarsal lyriform organ, and while it may notbe the only vibration receptor the spider has, it appears tobe the most sensitive. The 10 receptor units of which the senseorgan is composed are all sharply tuned to specific frequenciesbetween 60 and 1400 cps, but the sensitivity of each receptorunit depends upon the tension of the slits of the lyriform organ.Physiological experiments have shown that the tuning of thereceptors occurs only in response to air-borne sound; no discriminationis made of frequencies of web-borne vibration. One can speculatethat the high sensitivity to air-borne sound is a consequenceof the relatively poor transmission of vibration through Achaearanea'sweb.     

How to stay on mummy's back: Morphological and functional changes of the pretarsus in arachnid postembryonic stages

A specific type of maternal care occurs in several groups of Arachnida: mothers carry their offspring on their back (pulli-carrying behaviour). In scorpions, whip scorpions and whip spiders it is the prenymphal stage that settles on the mother. The prenymph is not yet fully developed for a free life and very limited in its mobility, but its feet are equipped with special adhesive organs (arolia) that become lost at the nymphal stage. Here we study the morphology, ultrastructure and mechanical function of the arolia. In scorpions (Scorpiones) the contact area between arolia and substrate and thus adhesion of the pad is controlled by the antagonistic work of hydrostatic pressure and muscular retraction. Arolia of whip scorpions (Thelyphonida) do not require muscular action for strong attachment. Arrays of long, branching fibres in the mesocuticle lead to high compliancy of the pad. In whip spiders (Amblypygi) the prenymphal pretarsus is already equipped with sclerites and claws. Its arolium is retained in nymphs and adults in some taxa, but acquires a more complex structure. These results contribute to our knowledge on the postembryonic development of arachnids and to the understanding of attachment pad evolution among arthropods. Some of the described developmental, structural, and mechanical phenomena are not known from other animals and might be of potential interest for further biomimetic developments.