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.

Thorax vibrations of a stingless bee ( Melipona seminigra). II. Dependence on sugar concentration.

Using a laser vibrometer we studied the influence of the food's sugar concentration on different parameters of the thorax vibrations produced by foragers of Melipona seminigra during trophallaxis in the nest. The concentrations tested (20-70% sugar w/w) were within the biologically relevant range. They substantially influenced different parameters of the thorax vibrations. An increase of energy gains at the food source due to an increased sugar concentration was followed by an increase of both the pulse duration and the duty cycle and by a decrease of the pause duration between two subsequent pulses. These findings further support the hypothesis that the temporal pattern of the thorax vibrations reflects the energy budget of a foraging trip rather than food source distance. Likewise, the steep increase of pulse duration variability with sugar concentration is hard to reconcile with the assumption that pulse duration conveys reliable information about food source distance when bees collect at high-quality food sources.

Thorax vibrations of a stingless bee ( Melipona seminigra). I. No influence of visual flow.

An important question in stingless bee communication is whether the thorax vibrations produced by foragers of the genus Melipona upon their return to the nest contain spatial information about food sources or not. As previously shown M. seminigra is able to use visual flow to estimate flight distances. The present study investigated whether foraging bees encode the visually measured distance in their thorax vibrations. Bees were trained to collect food in flight tunnels lined with a black-and-white pattern on their side walls and floor, which substantially influenced the image motion they experienced. When the bees had collected inside the tunnels the temporal pattern of their vibrations differed significantly from the pattern after collecting in a natural environment. These changes, however, were not associated with the visual flow experienced inside the tunnel. Bees collecting in tunnels offering little visual flow (stripes parallel to flight direction) modified their vibrations similarly to bees collecting in tunnels with high image motion (cross stripes). A higher energy expenditure due to drastically reduced flight velocities inside the tunnel is suggested to be responsible for changes in the thorax vibrations. The bees' vibrations would thus reflect the overall energetic budget of a foraging trip.

Arthropod touch reception: structure and mechanics of the basal part of a spider tactile hair.

Our previous studies on spider tactile hairs concentrated on the mechanical behavior of the hair shaft and the electrophysiological properties of the sensory cells. Here we focus on the structure and mechanical properties of the coupling of the hair shaft and the sensory terminals. 1. The functional "design" of the coupling provides for a combination of high sensitivity and protection against mechanical damage and overstimulation. The dendritic sheath is not directly coupled to the hair shaft. Rather, there is "terminal connecting material" between the dendrites and the hair shaft. 2. The hair shaft forms a first-order lever. Its acentric axis of rotation is located ca.3.5 microm from its inner end. Displacement of the hair tip is scaled down by a factor of ca.750:1, not even considering the outer hair shaft's bending. 3. At threshold the dendrite sheath displacement is ca. 0.05 microm by forces in the order of 0.4-4x10(-6) N. 4. The hair shaft bends within the socket even before contacting it. The elasticities representing its suspension and bending in the socket can be described quantitatively by measuring the hair's restoring moments (range: 10(-9) Nm) and bending at different degrees of deflection.

A stingless bee uses labial gland secretions for scent trail communication ( Trigona recursa Smith 1863).

The pheromones used by several species of stingless bees for scent trail communication are generally assumed to be produced by the mandibular glands. Here we present strong evidence that in Trigona recursa these pheromones originate from the labial glands, which are well developed in the heads of foragers. Analysis of the behavior involved in scent marking shows that a bee extends her proboscis and rubs it over the substrate. A single scent marking event lasts for 0.59+/-0.21 s while the bee runs a stretch of 1.04+/-0.37 cm on a leaf. According to choice experiments the bees are attracted by a feeder baited with labial gland extract (84.2+/-6% of the bees choose this feeder) but repelled from a feeder baited with mandibular gland extract (only 27.5+/-13.1% of the bees choose this feeder). They do not discriminate between two clean feeders (49.6+/-3% of the bees at a feeder). 87+/-5.1% of bees already feeding leave the feeder after the application of mandibular gland extract whereas only 6.2+/-4.9% and 2.6+/-4% do so when labial gland extract or pure solvent was applied.

A stingless bee (Melipona seminigra) uses optic flow to estimate flight distances.

Foragers of a stingless bee, Melipona seminigra, are able to use the optic flow experienced en route to estimate flight distance. After training the bees to collect food inside a flight tunnel with black-and-white stripes covering the side walls and the floor, their search behavior was observed in tunnels lacking a reward. Like honeybees, the bees accurately estimated the distance to the previously offered food source as seen from the sections of the tunnel where they turned around in search of the food. Changing the visual flow by decreasing the width of the flight tunnel resulted in the underestimation of the distance flown. The removal of image motion cues either in the ventral or lateral field of view reduced the bees' ability to gauge distances. When the feeder inside the tunnel was displaced together with the bees feeding on it while preventing the bee from seeing any image motion during the displacement the bees experienced different distances on their way to the food source and during their return to the nest. In the subsequent test the bees searched for the food predominantly at the distance associated with their return flight.

Arthropod touch reception: spider hair sensilla as rapid touch detectors.

Wandering spiders like Cupiennius salei are densely covered by tactile hairs. In darkness Cupiennius uses its front legs as tactile feelers. We selected easily identifiable hairs on the tarsus and metatarsus which are stimulated during this behavior to study tactile hair properties. Both the mechanical and electrophysiological hair properties are largely independent of the direction of hair displacement. Restoring torques measure 10(-9) to 10(-8) Nm. The torsional restoring constant S changes non-linearly with deflection angle. It is of the order of 10(-8) Nm/rad, which is about 10,000 times larger than for trichobothria. Angular thresholds for the generation of action potentials are ca.1 degrees. Electrophysiology reveals a slow and a fast sensory cell, differing in adaptation time. Both cells are movement detectors mainly responding to the dynamic phase (velocity) of a stimulus. When applying behaviorally relevant stimulus velocities (up to 11 cm s(-1)) threshold deflection for the elicitation of action potentials and maximum response frequency are reached as early as 1.2 ms after stimulus onset and followed by a rapid decline of impulse frequency. Obviously these hairs inform the spider on the mere presence of a stimulus but not on details of its time-course and spatial orientation.

Arthropod touch reception: stimulus transformation and finite element model of spider tactile hairs.

Striving towards an in depth understanding of stimulus transformation in arthropod tactile hairs, we studied the mechanical events associated with tactile stimulation. A finite element model was developed taking a tarsal tactile hair of the spider Cupiennius salei as an example. Considering hair diameter, wall thickness, and curvature, the hair is subdivided into six regions each with its specific mechanical properties. When the hair is touched from above with a flat surface oriented parallel to the tarsus the point of stimulus contact moves towards the hair base with increasing load and hair deflection. Thereby the effective lever arm is reduced protecting the hair against breaking near its base. At the same time the mechanical working range of the hair increases implying higher mechanical sensitivity for small deflections (about 5x10(-5) N/degrees) than for large deflections (about 1x10(-4) N/degrees). The major stresses within the hair shaft are axial stresses due to bending. The position of stress maxima moves along the shaft with the movement of the stimulus contact point. Remarkably, the amplitude of this maximum (about 1x10(5) N/m2) hardly changes with increasing loading force due to the way the hair shaft is deflected by the stimulus.

How to catch the wind: spider hairs specialized for sensing the movement of air.

Most arthropods are hairy creatures. Some of them have several hundreds of thousands of hairs on their exoskeleton which in the majority of cases serve mechanosensory functions. Filiform hairs or trichobothria, as they are called in the arachnids, respond to the slightest movement of the surrounding air. They have repeatedly been shown to be involved in the guidance of escape and prey capture behavior and are indeed among the most sensitive biosensors known to date. Accordingly, the mechanical interaction between the air and the hair which is deflected and thus adequately stimulated by viscous forces is very close and to a large extent follows principles known in fluid mechanics. Both the experimental and theoretical analysis of this interaction has reached considerable depth. Using spider trichobothria as the main example the present review article strives to explain in a simple way the main mechanical parameters to be considered and how hair morphology and mechanics bring about such remarkable sensitivity.

Early cortical activation indicates preparation for retrieval of memory for faces: an event-related potential study.

Event-related brain potentials (ERPs) were used to investigate the time course of memory processes following the presentation of faces. Following a phase in which subjects were asked to memorize faces presented on a computer screen (study phase) they had to distinguish the previously presented faces from others new to the experiment (test phase). We found that in a time period from 250 to 350 ms after onset of stimulus presentation ERPs show higher negativity for both repeated and novel faces in the test phase compared to the study phase. This situation dependent effect is most pronounced in occipito-temporal regions. We conclude that memory retrieval for faces is a sequential process. The early part of this process constitutes preparation for the retrieval of stored information, and a later part of the process comprises the discrimination between repeated and novel faces.

Fine structural correlates of sensitivity in the eyes of the ctenid spider, Cupiennius salei Keys.

We studied fine structural correlates of sensitivity in the principal and secondary eyes of the nocturnal hunting spider Cupiennius salei. In night-adapted eyes the four rhabdomeres of the principal eye photoreceptors are 58 microm long and occupy together 234 microm(2) in cross-section (average), whereas the two rhabdomeres of the secondary eye photoreceptors are about 49 microm long and measure 135-183 microm(2) in cross-section (average). The rhabdoms (photosensitive structures) consist of tightly packed microvilli (diameter 0.1 microm, maximum length 3.5 microm) and occupy up to 63% of the cross-sectional area of the retina. When calculating the amount of light the eyes of Cupiennius are able to capture according to their morphological characteristics, the values for sensitivity S(see Land, 1981, 1985) are between 78 and 109 microm(2). Cupiennius is more sensitive than any other hunting spider examined except Dinopis whose posterior median eyes are the most sensitive ones of all terrestrial arthropod eyes studied. In day-adapted eyes the rhabdomeral microvilli are almost completely degraded. The remaining microvillar surface amounts to only about one-tenth compared with the night-adapted state. Efferent synaptoid terminals have been found to contact the photoreceptors in all eyes of C. salei. The present fine structural data are compared to previous electrophysiological research and underline the significance of vision in Cupiennius.

Two visual systems in one brain: neuropils serving the secondary eyes of the spider Cupiennius salei.

Like other araneans, the wandering spider Cupiennius salei is equipped with one pair of principal eyes and three pairs of secondary eyes. Primary and secondary eyes serve two distinct sets of visual neuropils in the brain. This paper describes cellular organization in neuropils supplied by the secondary eyes, which individually send axons into three laminas resembling their namesakes serving insect superposition eyes. Secondary eye photoreceptors send axons to small-field projection neurons (L-cells) which extend from each lamina to supply three separate medullas. Each medulla is a vault of neuropil comprising only a few morphological types of neurons. These can be compared to a subset of retinotopic neurons in the medullas of calliphorid Diptera supplying giant motion-sensitive neurons in the lobula plate. In Cupiennius, neurons from secondary eye medullas converge at a single target neuropil called the "mushroom body." This region contains giant output neurons which, like their counterparts in the calliphorid lobula plate, lead to descending pathways that supply thoracic motor circuits. It is suggested that the cellular arrangements serving Cupiennius's secondary eyes are color independent pathways specialized for detecting horizontal motion. The present results do not support the classical view that the spider "mushroom body" is phylogenetically homologous or functionally analogous to its namesake in insects.

Two visual systems in one brain: neuropils serving the principal eyes of the spider Cupiennius salei.

Principal (anterior median) eyes of the wandering spider Cupiennius are served by three successive neuropils, the organization of which is distinct from those serving secondary eyes. Photoreceptors terminate in the first optic neuropil amongst second order neurons with overlapping dendritic fields. Second order neurons terminate at various depths in anterior median eye medulla where they are visited by bush-like dendritic trees of third order projection neurons. These supply tracts which extend into the "central body." This crescent-shaped neuropil lies midsagittally in the rear of the brain near its dorsal surface. It is organized into columns and it supplies both columnar and tangential efferents to other brain centers. The supply to and organization of the "central body" neuropil is reminiscent of retinotopic neuropils supplying the lobula of insects. Channels to the "central body" comprise one of two concurrent visual pathways, the other provided by the secondary eyes supplying the "mushroom body." We suggest that principal eye pathways may be involved in form and texture perception whereas secondary eyes detect motion, as is known for jumping spiders. Our data do not support Hanström's classical view that the "central body" is specifically associated with web-building, nor that it is homologous to its namesake in insect brains.

Fine structure of olfactory sensilla in myriapods and arachnids.

Structural features of various types of olfactory sensilla are reviewed. 1) Sensilla basiconica which differ in form and size are found on the antennae of centipedes and millipedes. Their walls show longitudinal slits or grooves that either open into the sensillum lumen or do not penetrate the cuticle. In other such sensilla the outer surface is pierced by pores and the inner surface grooved and pocketed. These sensilla are innervated by one to six sensory cells. Their unbranched outer dendritic segments extend to the tip of the sensillum. The sensory cells are surrounded by two or three sheath cells which terminate at the sensillum base or form a continuous tube around the entire length of the outer dendritic segments. 2) Temporal organs of centipedes are located between the insertion of the antenna and the ocelli. These sensilla consist of a shallow cuticular ring with a central sensory plate made up by a layer of unperforated cuticle or a capsule with a mushroom-shaped structure inside formed by fibrous-looking cuticle. A dozen sensory cells with unbranched outer dendritic segments innervate each sensillum. They extend toward the sensory cuticle and pass just below it. Numerous sheath cell processes run parallel to the outer dendritic segments up to the sensory cuticle. 3) Thread-like flagella of Pauropoda are found on the antennae. They possess a flexible unperforated cuticular wall. These sensilla contain nine sensory cells surrounded by several sheath cells which form a continuous cytoplasmic tube around the outer dendritic segments. 4) Single-walled sensilla with numerous plugged pores penetrating the cuticular wall occur on the tarsus of the first leg in ticks. Each sensillum is innervated by 4-15 sensory cells. Three sheath cells terminate in the base of the sensillum. 5) Double-walled sensilla with spoke canals are found on the first tarsus of ticks. Their shaft is longitudinally grooved. Pore canals lead inward from the bottom of the grooves and open into vase-shaped chambers. From its base these canals extend into the lumen of the sensillum which contains unbranched outer dendritic segments of 1-2 sensory cells. 6) Single-walled sensilla with pore openings occur on the distal tarsal segments of the first leg of whip spiders. These sensilla are innervated by 40-45 sensory cells. Their unbranched outer dendritic segments fill the shaft lumen and extend partly into the wall pores. Microvillus-shaped sheath cell processes line the inner surface of the cuticular wall.(ABSTRACT TRUNCATED AT 400 WORDS)

A versatile feedback controller for electro-mechanical stimulation devices.

Neurophysiological and behavioral work often requires that various laboratory stimulators be feedback-stabilized. We describe the design and performance of a versatile electronic controller that can be used to extend and flatten the frequency response of commercially available stimulating devices. The design includes flexible proportional-integral-derivative control action and active second-order, high-pass compensation. As an example application of this controller to 3 different electro-mechanical vibrator/transducer combinations demonstrates that the useful frequency response can be extended by more than a decade as compared with the uncontrolled device.

Vibratory communication through living plants by a tropical wandering spider.

Female Cupiennius salei pheromone on banana and Agave plants elicits patterned oscillations by the male. Resulting pulse trains of vibrations through the leaf average 76 hertz. The brief vibratory response by the otherwise immobile female hidden up to at least 1 meter away on another leaf guides the male across the plant to her location. Reciprocal signaling continues in the presence of random noise that masks the male's airborne sounds.

Dorsal giant fibre septum of earthworm. Fine structural details and further evidence for gap junctions.

The fine structure of the dorsal giant fibre septa of earthworm (Lumbricus terrestris and Eisenia foetida) was studied by thin sectioning. E-PTA, BIUL and ZIO impregnation were carried out to characterize the various membrane specializations found. A septum, which is formed by two axon membranes only ca. 7 nm apart, is a heterogeneous structure showing a number of different specializations (intermediate junctions, dense projection-like humps, septate regions, vesicles quite often associated with a widened intercellular gap and membrane thickening). Septal areas referred to as septal complexes are of particular interest. They comprise up to 20% of septal cross-sections and are characterized by membrane appositions (diameter ca. 25 nm) which bridge the septal gap, protrude 16-25 nm into the respective axoplasms, and form a more or less hexagonal array with a centre-to-centre spacing of ca. 33 nm (ca. 29 nm after E-EPTA treatment) in en face sections. We interpret the septal complexes as gap junctions, i.e. sites of electronic coupling, and emphasize their unusually large dimension. The vesicles found at the septum could not be stained by ZIO treatment.