A computational fluid dynamics model of viscous coupling of hairs

Arrays of arthropod filiform hairs form highly sensitive mechanoreceptor systems capable of detecting minute air disturbances, and it is unclear to what extent individual hairs interact with one another within sensor arrays. We present a computational fluid dynamics model for one or more hairs, coupled to a rigid-body dynamics model, for simulating both biological (e.g., a cricket cercal hair) and artificial MEMS-based systems. The model is used to investigate hair–hair interaction between pairs of hairs and quantify the extent of so-called viscous coupling. The results show that the extent to which hairs are coupled depends on the mounting properties of the hairs and the frequency at which they are driven. In particular, it is shown that for equal length hairs, viscous coupling is suppressed when they are driven near the natural frequency of the undamped system and the damping coefficient at the base is small. Further, for certain configurations, the motion of a hair can be enhanced by the presence of nearby hairs. The usefulness of the model in designing artificial systems is discussed.     

Modeling arthropod filiform hair motion using the penalty immersed boundary method

Crickets are able to sense their surrounding environment through about 2000 filiform hairs located on a pair of abdominal cerci. The mechanism by which the cricket is able to sense a wide range of input signals using these filiform hairs of different length and orientation is of great interest. Most of the previous filiform hair models have focused on a single, rigid hair in an idealized air field. Here, we present a model of the cercus and filiform hairs that are mechanically coupled to the surrounding air, and the model equations are based on the penalty immersed boundary method. The key difference between the penalty immersed boundary method and the traditional immersed boundary method is the addition of forces to account for density differences between the immersed solid (the filiform hairs) and the surrounding fluid (air). The model is validated by comparing the model predictions to experimental results, and then the model is used to examine the interactions between multiple hairs. With multiple hairs, there is little interaction when the hairs are separated by more than 1mm, and, as they move closer, they interact through viscous coupling, which reduces the deflection of the hairs due to the air movement. We also examine the computational scalability of the algorithm and show that the computational costs grow linearly with the number of hairs being modeled.     

Quantitative characterization of the filiform mechanosensory hair array on the cricket cercus

Background

Crickets and other orthopteran insects sense air currents with a pair of abdominal appendages resembling antennae, called cerci. Each cercus in the common house cricket Acheta domesticus is approximately 1 cm long, and is covered with 500 to 750 filiform mechanosensory hairs. The distribution of the hairs on the cerci, as well as the global patterns of their movement vectors, have been characterized semi-quantitatively in studies over the last 40 years, and have been shown to be very stereotypical across different animals in this species. Although the cercal sensory system has been the focus of many studies in the areas of neuroethology, development, biomechanics, sensory function and neural coding, there has not yet been a quantitative study of the functional morphology of the receptor array of this important model system.

Methodology/Principal Findings

We present a quantitative characterization of the structural characteristics and functional morphology of the cercal filiform hair array. We demonstrate that the excitatory direction along each hair''s movement plane can be identified by features of its socket that are visible at the light-microscopic level, and that the length of the hair associated with each socket can also be estimated accurately from a structural parameter of the socket. We characterize the length and directionality of all hairs on the basal half of a sample of three cerci, and present statistical analyses of the distributions.

Conclusions/Significance

The inter-animal variation of several global organizational features is low, consistent with constraints imposed by functional effectiveness and/or developmental processes. Contrary to previous reports, however, we show that the filiform hairs are not re-identifiable in the strict sense.     

Structure of the afferent terminals in terminal ganglion of a cricket and persistent homology

We use topological data analysis to investigate the three dimensional spatial structure of the locus of afferent neuron terminals in crickets Acheta domesticus. Each afferent neuron innervates a filiform hair positioned on a cercus: a protruding appendage at the rear of the animal. The hairs transduce air motion to the neuron signal that is used by a cricket to respond to the environment. We stratify the hairs (and the corresponding afferent terminals) into classes depending on hair length, along with position. Our analysis uncovers significant structure in the relative position of these terminal classes and suggests the functional relevance of this structure. Our method is very robust to the presence of significant experimental and developmental noise. It can be used to analyze a wide range of other point cloud data sets.     

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     

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     

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.     

Agonistic signals received by an arthropod filiform hair allude to the prevalence of near-field sound communication

Arthropod filiform hairs respond to air particle movements and are among the most sensitive animal sensory organs. In many species, they are tuned to detect predators or prey and trigger escape or prey capture behaviours. Here we show for the first time that these hairs also receive intraspecific near-field sound signals in an arachnid. During agonistic encounters, whip spiders (Arachnida, Amblypygi) perform antenniform leg vibration (ALV) displays that have significantly longer duration in contest winners than losers. During an ALV display: (i) the vibrating antenniform leg of the displaying whip spider is positioned close to the trichobothria (filiform hairs) on its opponent's walking legs, (ii) the vibrating antenniform leg can excite these trichobothria via air movements and without direct contact, (iii) the antenniform leg of the displaying whip spider vibrates at a frequency that causes particularly strong, sustained excitation and little adaptation in the trichobothria, and (iv) the duration of an ALV display can be extracted from the response of a trichobothrium. Since filiform hairs are widespread among arthropods, communication via such hairs could be extremely prevalent.     

A Model of Filiform Hair Distribution on the Cricket Cercus

Crickets and other orthopteran insects sense air currents with a pair of abdominal appendages resembling antennae, called cerci. Each cercus in the common house cricket Acheta domesticus is covered with between 500 to 750 filiform mechanosensory hairs. The distribution of the hairs on the cerci, as well as the global patterns of their movement axes, are very stereotypical across different animals in this species, and the development of this system has been studied extensively. Although hypotheses regarding the mechanisms underlying pattern development of the hair array have been proposed in previous studies, no quantitative modeling studies have been published that test these hypotheses. We demonstrate that several aspects of the global pattern of mechanosensory hairs can be predicted with considerable accuracy using a simple model based on two independent morphogen systems. One system constrains inter-hair spacing, and the second system determines the directional movement axes of the hairs.     

Relative contributions of organ shape and receptor arrangement to the design of cricket’s cercal system

Understanding the relative contributions of the shape of a sensory organ and the arrangement of receptors to the overall performance of the organ has long been a challenge for sensory biologists. We tackled this issue using the wind-sensing system of crickets, the cerci, two conical abdominal appendages covered with arrays of filiform hairs. Scanning electron microscopy coupled with 3D reconstruction methods were used for mapping of all cercal filiform hairs. The hairs are arranged according to their diameter in a way that avoids collisions with neighbours during hair deflection: long hairs are regularly spaced, whereas short hairs are both randomly and densely distributed. Particle image velocimetry showed that the variation in diameter of the cercus along its length modifies the pattern of fluid velocities. Hairs are subject to higher air flow amplitudes at the base than at the apex of the cercus. The relative importance of interactions between receptors and the air flow around the organ may explain the performance of the cricket's cercal system: it is characterised by a high density of statistically non-interacting short hairs located at the base of the cercus where sensitivity to air currents is the highest.     

Central projections of campaniform sensilla on the cerci of crickets and cockroaches

Summary Campaniform sensilla associated with filiform hairs comprise an important receptor type of the multimodal sensory system of the cerci of crickets and cockroaches. Their axon projections were investigated using iontophoretic cobalt injection into single sensilla.In crickets (Gryllus bimaculatus, Acheta domestica), six different types of cereal campaniform sensilla projections can be distinguished on the basis of their axonal arborizations and terminations. Typically, a proportion of cereal campaniform sensilla, associated with long filiform hairs, give rise to axons that ascend as through fibres from the terminal ganglion to reach the sixth abdominal ganglion. Cereal campaniform sensilla associated with clavate hairs have projections restricted to the terminal ganglion alone.Whereas in crickets axons of cercal campaniform sensilla invade only certain segmental neuropils in the terminal ganglion, in cockroaches (Periplaneta americana) axons from cercal campaniform sensilla branch in every segmental neuropil. A proportion of cereal campaniform sensilla in this species also gives rise to through fibres to the fifth abdominal ganglion.We discuss morphological and functional interpretations of differences between crickets and cockroaches and consider the significance of this type of receptor in the context of previous studies of the cercal system.     

中国黄耆属糙叶组叶表皮毛状体微形态学特征

采用体视显微镜和扫描电子显微镜,对中国产黄耆属糙叶组34种植物的叶表皮毛状体的形态特征及其分布进行了观察,以探讨中国黄耆属糙叶组物种表皮毛微观形态的变异式样及其分类学意义,为糙叶组物种的分类鉴定提供依据。结果表明：除乳白花黄耆（A. galactites）仅在叶下表面被毛外,其余种类上下表皮均有毛,且毛的分布、形态、长度和臂比在不同种类间存在差异。叶表皮毛为伏贴或开展的中空管状毛,伏贴毛着生部位基部膨大,整体形态从中部向两端由粗逐渐变细,开展的毛大部分为细长的丝状或线状,光滑或具条纹状纹饰,上面分布有乳突。研究认为,叶表皮表面伏贴、短而平直、其上光滑具密集乳突、臂比值小、整体宽扁的毛是祖先特征;而叶表皮表面向上开展、长而密集、外形呈丝状或线形、其上具条纹状纹饰、乳突稀疏、臂比值大的毛是进化特征。     

Cercal sensory system and giant interneurons in Gryllodes sigillatus

The external morphologies of two cricket species, Gryllodes sigillatus and Gryllus bimaculatus, were investigated. Despite its small body length, G. sigillatus possessed longer cerci and longer cercal filiform hairs than G. bimaculatus. The estimated number of filiform hairs on a cercus was also larger in G. sigillatus than in G. bimaculatus. Wind-sensitive interneurons receiving sensory inputs from cercal filiform hairs and running in the ventral nerve cord (VNC) were investigated in G. sigillatus both morphologically and physiologically. By intracellular staining, these interneurons were proved to be morphologically homologous with previously identified giant interneurons (GIs 8-1, 9-1, 9-2, 9-3, 10-2, and 10-3) in G. bimaculatus and Acheta domesticus. In G. sigillatus, the intensity-response relationship (I-R curve) for each GI was investigated using a unidirectional air current stimulus. The stimulus was applied from 12 different directions, and an I-R curve was obtained for each stimulus direction. Each GI showed a characteristic I-R curve depending on stimulus direction. The directionality curve expressed in terms of threshold velocity showed that each GI had a distinctive directional characteristic. The functional properties of GIs in G. sigillatus, such as I-R curve, threshold velocity, and directional characteristics, were compared with those of homologous GIs in G. bimaculatus in Discussion.     

A scanning electron microscopic study of the dorsal surface of the human tongue

To study the dorsal surface of the human tongue using a scanning electron microscopy (SEM), tissue specimens were taken from the anterior part of the tongues of 15 individuals aged from 21- to 28-years-old. The formalin-fixed samples were processed routinely for SEM. With SEM the surface of the normal tongue mucosa was shown to be rather evenly covered by filiform papillae, with some fungiform papillae scattered among them. Filiform papillae consisted of two parts: the body and hairs. The mucosal surface of the body was smooth; the squamous epithelial cells were polygonal, and their boundaries were prominent. On the surface of the superficial epithelial cells were parallel or branching microplicae. Each filiform papilla had 6-10 hairs, which were scaled and covered by an extensive plaque of microorganism. The upper surface of the fungiform papillae was smooth; only a few desquamating cells were seen. The superficial cells had a pitted appearance and cell boundaries overlapped. Taste pores, up to 3 pores in a single papilla, were found on the upper surface. Desquamation was more pronounced on the base of the fungiform papillae than on the upper surface. In almost all fungiform papillae some hairs protruded from the base. Parallel microplicae were found on the surface of the superficial cells of the base. The structure and function of the human tongue, as well as the microplicae of its superficial cells, are compared to those of various species of animals.     

Ectopic sensory neurons in mutant cockroaches compete with normal cells for central targets.

The cercus of the first instar cockroach, Periplaneta americana, bears two filiform hairs, lateral (L) and medial (M), each of which is innervated by a single sensory neuron. These project into the terminal ganglion of the CNS where they make synaptic connections with a number of ascending interneurons. We have discovered mutant animals that have more hairs on the cercus; the most typical phenotype, called "Space Invader" (SI), has an extra filiform hair in a proximo-lateral position on one of the cerci. The afferent neuron of this supernumerary hair (SIN) "invades the space" occupied by L in the CNS and makes similar synaptic connections to giant interneurons (GIs). SIN and L compete for these synaptic targets: the size of the L EPSP in a target interneuron GI3 is significantly reduced in the presence of SIN. Morphometric analysis of the L afferent in the presence or absence of SIN shows no anatomical concomitant of competition. Ablation of L afferent allows SIN to increase the size of its synaptic input to GI3. Less frequently in the mutant population, we find animals with a supernumerary medical (SuM) sensillum. Its afferent projects to the same neuropilar region as the M afferent, makes the same set of synaptic connections to GIs, and competes with M for these synaptic targets. The study of these competitive interactions between identified afferents and identified target interneurons reveals some of the dynamic processes that go on in normal development to shape the nervous system.     

A neuromorphic hair sensor model of wind-mediated escape in the cricket

Crickets are able to extract directional information about a wind stimulus through the filiform hairs located on their cerci. This paper describes the design and testing of a neuromorphic sensor that aims to achieve a close correlation with both the physical and functional properties of these hairs. An integrate and fire neural network is used to process the sensory information in real time. The resulting system is shown to be capable of extracting directional information from a wind stimulus and producing an appropriate motor control pattern.     

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     

Evidence for air movement signals in the agonistic behaviour of a nocturnal arachnid (order Amblypygi)

Many arthropods possess filiform hair sensilla (termed trichobothria in arachnids), which are extremely sensitive detectors of medium particle displacement. Electrophysiological evidence in some taxa suggests that these sensilla can detect air particle displacements resulting from intraspecific communication signals. However, it has not yet been shown for any species that the air particle displacements detected by the filiform hairs are themselves perceived as a 'signal' (i.e. that individuals make behavioural decisions based upon the responses of these organs to the displays of conspecifics). We investigate the agonistic behaviour of the whip spider Phrynus marginemaculatus and the role of its trichobothria in receiving agonistic signals. Whip spiders have extremely elongated 'antenniform' first legs, which they vibrate close to their opponents during agonistic interactions, inducing air movements that excite their opponents' trichobothria. We find that ablation of the trichobothria causes significant increases in: (I) contest duration, and (II) the probability of contest escalation past aggressive displays to physical fighting. Therefore, in the absence of air movement-sensitive sensilla, contest assessment is impaired. This suggests that whip spiders exploit true air movement signals during agonistic interactions, and that these are received by the trichobothria. Furthermore, these results indicate that, in whip spiders, such signals help mitigate the cost of agonistic interaction.     

Sensitivity of an insect mechanoreceptor during moulting

ABSTRACT. The fine structure and the behavioural threshold for vibration sensitivity of the eight thoracic filiform hairs of Barathra brassicae caterpillars were investigated through an intermoult/moult cycle. Associated with each filiform hair is one bipolar sensory cell and three enveloping cells. The outer dendritic segment terminates in an ecdysial canal in the hair base and a tubular body lies at its distal end. Shortly before apolysis the dendrite elongates. By this means the connection between the sensory cell and the old cuticular apparatus is maintained while the epithelium and the old thoracic cuticle are separating. The new cuticular apparatus of the filiform hair is formed in the second half of the larval stage by the three enveloping cells. A second tubular body in the elongated outer dendritic segment is formed at the base of the replacement hair 10 h before next ecdysis, so that the new hair functions as soon as ecdysis is completed, the old cuticular apparatus with the old tubular bodies being torn away with the exuvia during ecdysis. Sensitivity to a 300 Hz tone was tested in the standing wave of a Kundt's tube. Throughout most of the larval instar the threshold was 2.0 ± 0.3 μm particle displacement amplitude until 1–2h before ecdysis when it rose to 6.8 ± 1.3 μm and at 10–30 min before the beginning of ecdysis no reaction to sound could be detected. Once the old cuticle was shed maximum sensitivity returned as soon as the replacement hairs were erect. The sensilla are therefore physiologically functional at all developmental stages except for 30–60 min during actual ecdysis.