Mechanics of trichobothria in orb-weaving spiders (Agelenidae,Araneae)

Summary When a fly is humming at a distance of about one centimetre from an orb-weaving spider (Agelenidae) the trichobothria on the spider's extremities are deflected by air streams and air vibrations. Frequency analysis of the hum of the two prey animals,Drosophila andMusca, shows that the effective sound velocities of the harmonics with frequencies inferior to some five hundred Hz exceeds that of higher frequencies by a factor of at least 5. Biologically relevant resonances would, therefore, have to be looked for in the range of a few hundred Hz. Frequency response diagrams show that single hairs have no resonance between a few Hz and approximately 2 kHz. The maximal relative amplitude of hairs of different lengths shifts from the longer to the shorter hairs with increasing frequency. As this is only a minor effect, however, it appears that there is no frequency discrimination by the mechanical apparatus. Constant air streams with a velocity of 40 mm/s cause hair deflection of about 10 degrees (the hair's bend is neglibible). Similarly, near-field particle velocity of sound fields up to a few hundred Hz is well transmitted. The mechanical directional sensitivity does not depend on the azimuthal angle of deflection. Thus, information about direction and velocity of stationary and near-field air movements is transmitted without deformation by the mechanical apparatus. This is well matched with the fact that the hair is multiply innervated.Supported by grants of the Deutsche Forschungsmeinschaft in the field of the Schwerpunktprogramm Rezeptorphysiologie     

Direction sensitivity of the filiform hair population of the cricket cereal system

Each cercus on the adult cricket Acheta domesticus bears 1000–2000 filiform hair mechanoreceptors. In order to determine the extent of identifiability of individual hair receptors, the structural characteristics of ten putative identified hairs were measured in 21–25 different animals. For these ten hairs, the sets of preferred directions and circumferential locations had standard deviations of 6.8° and 5.9°, respectively. There was no significant inter-animal covariance between a hair's preferred direction and its circumferential location. The preferred directions of 246 different identified hairs were then measured from 16 animals in order to characterize the distribution of preferred directions of hairs on a single typical cercus. These data were transformed from the frame of reference of the cercus into that of the cricket, generating an estimate of the representation of air-current stimulus direction provided by the entire ensemble of filiform hairs on both cerci. The distribution of hair directional sensitivities was continuous but extremely non-uniform, and more complex than previous studies had suggested.Abbreviations MT medial-transverse - LT lateral-transverse - AL anterior-longitudinal - PL posterior-longitudinal - MAO medial-anterior-oblique - MPO medial-posterior-oblique - LAO lateral-anterior-oblique - LPO lateral-posterior-oblique - T transverse - L longitudinal - O oblique - CNS central nervous system     

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.     

The tibial thread-hairs ofAcheta domesticus L. (Saltatoria,Gryllidae)

Summary The ultrastructure of the thread-like hairs (sensilla) on the tibia of the front leg ofAcheta domesticus (Gryllidae) Saltatoria was examined by serial sectioning. The presence of a tubular body indicates that these sensilla are mechanosensitive; electrophysiological measurements also confirmed this. The opposing forces on the articulating apparatus of single hairs and the sensitivity of the single receptor cell were measured after deflection of the hair in different directions. The articulating apparatus is characterized by three cuticular elements: a joint membrane, suspension fibers, and a socket septum. These elements form the basis for a structural bilateral symmetry along whose plane of symmetry the direction line of both the minimum receptor sensitivity and the minimum opposing forces lie. The tubular body embedded in the tip of the socket septum is attached to the base of the hair shaft. The hair provides the leverage for displacing the tubular body and the socket septum limits the extent to which it may be laterally displaced.These investigations have been supported by the Deutsche Forschungsgemeinschaft     

The structure of a hair mechanoreceptor in the antennule of crayfish (Crustacea)

Summary In this study we examine the fine structure of mechanosensory hairs in the antennule of crayfish. The sensory hair is a stiff shaft with feather-like filaments. The hair's base is a large expansion of membrane which allows the hair shaft to deflect. The sensory transducing elements are located far from the hair, but are coupled mechanically with the hair shaft by a fine extracellular chorda. The sensory element is a type of scolopidium which consists of a scolopale cell and three sensory cells with a 9 + 0 type ciliary process.This type of scolopidium is characteristic of the chordotonal organ that has no cuticular structure on the surface of the exoskeleton. In this crustacean hair receptor, the deflection of the cuticular hair is transmitted through the chorda to the scolopidium which is a tension-sensitive transducer. The present study reveals that the mechanosensory hair of decapod crustaceans is a chordotonal organ accompanied by a cuticular hair structure. We also discuss comparative aspects of cuticular and subcuticular chordotonal organs in arthropods.     

Central projections of labellar taste hairs in the blowfly,Phormia regina Meigen

Summary We have traced the central projections of the receptor neurons associated with each of the eleven largest taste hairs on the labellum of the blowfly, Phormia regina (Meigen), by staining them with cobaltous lysine. The eleven hairs fall into three groups which reflect their peripheral locations and their branching patterns in the subesophageal ganglion. Group 1, consisting of the anterior hairs (numbers 1 and 2) and Group 3, consisting of the posterior hairs (numbers 9–11) project bilaterally, while Group 2, consisting of the middle hairs (numbers 3–8) projects primarily ipsilaterally. The central projections of the hairs within a single group are similar. Each hair houses four chemoreceptors, which have differing chemical sensitivities and behavioral roles, and one mechanoreceptor. In some cases, there were indications that the different cells within a single hair have different central branching patterns. For some hairs, however, it was clear that a single central branching region and pattern was shared by more than one receptor cell. We failed to find either a continuous somatotopic representation of a hair's position on the periphery, or an anatomical segregation of receptors coding for different modalities. Behavioral experiments indicate that the fly is informed both of the identity of the hair stimulated and of the chemical nature of the stimulus. Our results suggest that this information is not represented on a gross anatomical level.     

Morphology and physiology of a hair plate sensory organ located on the antenna of the rock lobster Palinurus vulgaris

At the level of the J1 joint of each antenna of the rock lobster Palinurus vulgaris a hair plate sensory organ (hp) similar to those described in insects has been observed. The hp is located on the internal side of the S2 segment of the antenna, close to the soft articulating membrane of the J1 joint. It is formed by a triangular surface of cuticle about 3mm2 in area, covered with numerous hairs of different lengths (Figs. 1 and 2). Details of the hp were studied by scanning electron microscopy (Fig. 2). Physiological stimulation of the hp hairs occurs during medial movement of the J1 joint. Under this condition the soft articulating membrane rolls over the hairs and bends them progressively back onto the cuticle. Flexion of all the hairs corresponds to a medial movement of the J1 through 40 degrees. During this type of movement, the number of successively flexed hairs increases linearly (Fig. 3). Electrophysiological recordings of the hp sensory nerve correlated with selective mechanical stimulation of individual hairs demonstrated that each hair is innervated by a single sensory fiber. This sensory neurone responds phasically when the hair is flexed back onto the cuticle (as during an S2 medial movement) and when it returns to its resting position (as during an S2 lateral movement). Most of the sensory neurones are sensitive to the movement velocity of the hairs (Figs. 4 and 5). When the hair is maintained flexed its sensory neurone discharges tonically (Fig. 4). Electrical stimulation of the hp sensory nerve induced reflex actions in the external and internal rotator muscles of the segment S1. These effects were found to selectively activate the tonic motor command of these muscles (Fig. 6).     

Directional selectivities of near-field filiform hair mechanoreceptors on the crayfish tailfan (Crustacea: Decapoda)

Directional selectivities of mechanoreceptors that innervate filiform hairs on the crayfish tailfan were investigated with unidirectional, sinusoidal, water-motion stimuli. These recordings provide the first representative sample from filiform hair sensilla on the entire tailfan. The filiform hair receptors exhibit unimodal directional selectivity patterns that were well fitted by a cardioid function with a half-width of 122°. The preferred directions correspond to the major axis of hair motion, and are perpendicular to the orientation of lateral branches on the main hair shaft. Pooled plots of preferred directions demonstrate quadrimodal patterns on the telson and endopods which are associated with hair location, and a bimodal pattern on the exopods. For each appendage, the combination of the overall pattern of preferred directions with “coarse coding” of direction by individual receptors provides sensitivity to a full 0–360° range of water motion and the potential to discriminate the direction of water motion throughout this range. The results demonstrate several similarities to the wind-sensitive cercal receptor system in orthopteroid insects, and suggest that crustacean filiform hair receptors provide a sufficient sensory basis for behavioral orientation to water currents and shorelines. Accepted: 5 January 1998     

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.     

Long,smooth hair sensilla on the spider leg coxa: Sensory physiology,central projection pattern,and proprioceptive function (Arachnida,Araneida)

Summary Rows of long, smooth hair sensilla situated on both sides of the leg coxae were examined in the spider Cupiennius salei (Ctenidae). The hair shafts point into the space between adjacent legs and are deflected when the hairs of one coxa touch the cuticle of the neighboring coxa. 1. Unlike the serrated hair shafts of the ubiquitous tactile and chemosensitive setae of spiders, these hairs are entirely smooth. At their base they are articulated in a socket with an asymmetrical groove that determines the direction of hair deflection. Hair shafts are up to 1000 m long. The exact grouping of smooth hairs in rows is typical of the coxae for each pair of legs. 2. Unlike the other, multiply innervated cuticular sensilla of spiders, smooth hairs are supplied by only a single mechanosensitive neuron. This is confirmed by electrophysiological recordings from single hairs. Threshold deflection to elicit a spike response lies near 1°. The response to maintained, step-like stimuli declines rapidly. 3. All central endings of these hair receptors in the fused segmental ganglia are confined to dorsal neuropil of the ipsilateral neuromere. The specific arborization pattern resembles an elongated, three-pronged fork with a long central prong. Topography, natural stimulus situation, and the phasic response characteristic of smooth hairs suggest that spiders use these sensilla to monitor the relative distance between leg coxae during locomotion.     

Reaction of single cercal mechanoreceptors of the cricket, Gryllus bimaculatus, to mechanical stimulation

Naked tungsten microelectrodes were introduced through the chitinous wall close to the cell body of the receptor cells of trichoid hair sensillae, and responses to airpuffs, and rectangular, and trapezoidal displacements of the hair were recorded. Receptors of dorsal zone are activated during lateral deflection, those of ventral zone--during medial deflection and receptors of medial and lateral zones--during deflection to the cercal base. Sensitivity of receptors to the air-puffs is a function of hair length, the largest hairs being most sensitive. During trapezoidal displacement of the hair with different velocities of the slope, discharge frequency of the dynamic response is a function of velocity and angle in the range of angles up to 3-5 degrees (fig. 1). Discharge frequency of the stationary phase (corresponding to the plateau of the stimulus) is mainly a function of velocity in the range up to 6 degrees (fig. 2). The presence of sensillae with different hair length, and hence sensitivity, and definite directionality of receptors in different hair length, and hence sensitivity, and definite directionality of receptors in different zones may provide a basis for amplitude, velocity and direction discrimination of air-puffs or low-frequency mechanical stimulation by the cercal system of crickets.     

A virtual hair cell, I: addition of gating spring theory into a 3-D bundle mechanical model

We have developed a virtual hair cell that simulates hair cell mechanoelectrical transduction in the turtle utricle. This study combines a full three-dimensional hair bundle mechanical model with a gating spring theory. Previous mathematical models represent the hair bundle with a single degree of freedom system which, we have argued, cannot fully explain hair bundle mechanics. In our computer model, the tip link tension and fast adaptation modulator kinetics determine the opening and closing of each channel independently. We observed the response of individual transduction channels with our presented model. The simulated results showed three features of hair cells in vitro. First, a transient rebound of the bundle tip appeared when fast adaptation dominated the dynamics. Second, the dynamic stiffness of the bundle was minimized when the response-displacement (I-X) curve was steepest. Third, the hair cell showed "polarity", i.e., activation decreased from a peak to zero as the forcing direction rotated from the excitatory to the inhibitory direction.     

北京幽灵蛛体表微感受器的类型、结构和分布

北京幽灵蛛(Pholcus beijingensis)体表的微感受器包括毛状感受器(触毛、听毛、味觉毛和刺)、裂缝状感受器(单个裂缝器、竖琴器)和跗节器等.扫描电镜观察显示,北京幽灵蛛体表的毛状感受器数量最多,分布最广;其次是裂缝感受器;此外,每个跗节末端具有一个跗节器.除触毛在整个身体表面均有分布外,其他毛状感受器(...     

Mechanosensory afferents innervating the swimmerets of the lobster

Feathered hair sensilla fringe both rami of the lobster (Homarus americanus) swimmeret. The sensory response to hair displacement was characterized by recording afferent impulses extracellularly from the swimmeret sensory nerve while deflecting sensilla with a rigidly-coupled probe or controlled water movements. Two populations of hairs were observed: "distal" hairs localized to the distal 1/3 of each ramus and "proximal" hairs near its base. Distal hairs are not innervated by a mechanosensory neuron but instead act as levers producing strain within adjacent cuticle capable of activating a nearby hypodermal mechanoreceptor. Hair deflections of 25 degrees or more are required to evoke an afferent response and this response is dependent on hair deflection direction. The frequency and duration of the afferent discharge evoked are determined by the velocity of hair displacement. Each proximal hair is innervated by a single mechanosensory neuron responding phasically to hair deflections as small as 0.2 degrees in amplitude. Deflection at frequencies up to 5 Hz elicits a single action potential for each hair movement; at higher frequencies many deflections fail to evoke an afferent response. These sensilla, which are mechanically coupled, may be activated by the turbulent flow of water produced by the swimmerets during their characteristic beating movements.     

Coxal hair plates in spiders: physiology,fine structure,and specific central projections

Coxal hair-plate sensilla in the spider Cupiennius salei are described with respect to their innervation, central projection pattern, electrical response to mechanical stimulation, and putative behavioral function.

Mobilities of the cercal wind-receptor hairs of the cricket, Gryllus bimaculatus

The deflection sensitivities of cercal filiform hairs of the cricket, Gryllus bimaculatus, were determined by direct measurement. The tangential velocity of deflecting hair shafts in response to stimulus air motion was measured in situ by a laser-Doppler velocimeter with surface scattering of the shaft. The velocity of the stimulus air motion in a small wind tunnel was calibrated by the same velocimeter with smoke from a joss-stick. The mobility of the hair was obtained from former measurements with reference to the latter calibration of the single apparatus. A Gaussian white noise signal was employed as a stimulus waveform, and the stimulus-response transfer function was calculated through a cross-correlation method, which provides greater precision and wider frequency for a longer period of measurement. The mobility of hair was expressed in deflection amplitudes and phase shifts in reference to the velocity sinusoid of a stimulus at various frequencies. The measurements established the following conclusions. The wind receptor hairs comprise an array of mechanical band-pass filters whose best frequencies are inversely proportional to the length. The motion dynamics of the wind-receptor hairs have strong damping. Accepted: 24 February 1998     

Differential effects of nitric oxide on the responsiveness of tactile hairs

The responses of tactile hairs located on legs of the desert locust Schistocerca gregaria (Forskål) are modulated by nitric oxide (NO). There are two types of tactile hair on the tibia of the hind leg of the locust which differ in their thresholds for mechanical stimulation, their location on the leg and in the effect of NO on their responses to deflection. The spike response rates of mechanosensory neurons of low-threshold hairs decreased when exposed to elevated NO levels caused by perfusion of the leg with saline containing the NO donor PAPANONOate. In contrast, in high-threshold hairs, which show low responsiveness under control conditions, an increase in spike rates was observed during PAPANONOate application. These opposing effects of NO reduce the differences in the spike responses of the two types of tactile hairs to mechanical stimulation and are likely to have an impact on behaviours elicited by mechanical stimulation of the legs.     

蜚蠊尾须单个细毛型感受器对机械刺激的反应

用钨丝微电极插入蜚蠊尾须细毛感受器的园盘作胞外记录.研究了感受器自发脉冲的统计特性.利用电磁驱动装置控制毛杆的偏转,研究了感受器电位和脉冲对正弦波、方波和阶梯波机械刺激的反应特性,并测定了六列细毛的兴奋方向.     

Viscosity-mediated motion coupling between pairs of trichobothria on the leg of the spider <Emphasis Type="Italic">Cupiennius salei</Emphasis>

Arachnids and insects use long, thin hairs as motion sensors to detect signals contained in the movement of the surrounding air. These hairs often form groups with a small spacing of tens to hundreds of micrometers between them. For air oscillation frequencies of biological interest, the potential exists for viscosity-mediated coupling among hairs in a group affecting their response characteristics. Even a small diameter hair can, in principle, affect the flow field around it and the dynamics of the hairs in its neighborhood. The viscosity-mediated coupling between a pair of hairs is investigated here both experimentally and theoretically. The conditions for the existence of the coupling effect, and its magnitude as a function of relevant parameters, are determined. In the range of biologically relevant frequencies (30–300 Hz), viscous coupling between pairs of hairs is only very small in the case of the spider Cupiennius salei. Theoretical analysis points to the relatively large spacing between hairs (20 to 50 hair diameters) and the tuning of the hairs to the above-mentioned frequencies to explain the practical absence of coupling.     

Nonlinear mechanical responses of mouse cochlear hair bundles.

The stiffness of sensory hair bundles of both inner (IHC) and outer (OHC) hair cells was measured with calibrated silica fibres in mouse cochlear cultures to test the hypothesis that the mechanical properties of the hair bundle reflect processes underlying mechanotransduction. For OHCs, the displacement of the hair bundle relaxed with time constants of 6 ms for displacements which open transducer channels and 4 ms for displacements which close the channels. The corresponding values of the time constants for IHCs were 10 ms and 8 ms, respectively. A displacement-dependent change in the stiffness of the hair bundle was not observed when the bundle was displaced orthogonally to the direction of excitation. The stiffness of the hair bundle as a function of nanometre displacements from the resting position was remarkably nonlinear. The stiffness declined to a minimum from the resting stiffness by about 12% for OHCs and 20% for IHCs when the hair bundle was displaced by about 20 nm in the excitatory direction, and it increased by a similar amount when the bundle was displaced by 20 nm in the inhibitory direction. The displacement at which the stiffness reached a minimum was within the most sensitive region of the hair-cell transducer function (receptor potential as a function of hair-bundle displacement), and the displacement at which the stiffness reached a maximum was at the point of saturation of the transducer function in the inhibitory direction. The nonlinear displacement-dependent compliance change is reversibly abolished, and the time constant of relaxation of the bundle for excitatory displacements is reversibly reduced, when mechanotransduction is blocked by the addition of either neomycin sulphate or cobalt chloride to the solution bathing the hair cells. The displacement-dependent compliance change was not apparently reduced when the receptor potential was attenuated through the substitution of sodium in the bathing solution with a less permeant cation, tetraethylammonium. These findings suggest that the nonlinear mechanical properties of the hair bundle are associated with aspects of the hair-cell mechanotransducer process. The mechanical properties of the hair bundle are discussed in relation to the 'gating-spring' hypothesis of hair-cell transduction.