Effect of Mechanical Stimulus Spread across Glabrous Skin of Raccoon and Squirrel Monkey Hand on Tactile Primary Afferent Fiber Discharge

The role of spread of skin deformation in activating cutaneous mechanoreceptors at a distance from their threshold receptive fields (RFs) was examined in glabrous skin of the North American raccoon and the squirrel monkey. One feedback-controlled mechanical stimulus probe was used to indent the skin to a controlled depth at a constant velocity, at varying distances from a second probe, which was used to monitor vertical displacement depth and velocity at this distant site. In many instances, the monitor probe was positioned over the RF of a cutaneous mechanoreceptor, and single-unit action potentials were simultaneously recorded from individual fibers of the median or ulnar nerve.

With distance from the site of stimulation, there was a systematic, monotonic decline in indentation depth and velocity; velocity fell off with distance more rapidly than depth. The degree of diminution with distance varied with the size, shape, and curvature of the digital or palm pad stimulated. Spread of indentation was more restricted on digital than on palm pads, and was more restricted across monkey skin than across raccoon skin. Spread was less with higher-velocity than with lower-velocity indentations, but was seemingly unaffected by indentation depth.

As expected from the findings noted above, the number of spikes discharged by slowly adapting mechanoreceptive afferent fibers declined more rapidly with distance between stimulus site and RF for digital than for palmar RFs, in squirrel monkey than in raccoon skin, and with higher-velocity than with lower-velocity stimuli. Furthermore, the number of spikes occurring during either ramp or early static indentation phases of stimulation dropped to zero more rapidly with distance than did either vertical indentation depth or velocity. Decreases with distance in both indentation depth and velocity acted to restrict the size of suprathreshold RFs. For most units, horizontal components of mechanical stimulation subtracted from the effects of vertical components.

It is suggested, on the basis of this and other studies, that many neural and perceptual phenomena usually attributed to central mechanisms of afferent inhibition may be attributable, at least in part, to mechanical properties of the skin. In addition, the present data suggest that regional variations in the two-point limen may be associated with variations in spread of mechanical deformation. The conclusion that glabrous skin and subjacent soft tissues act as a low-pass filter system provides a mechanical basis for the relative efficacy of high-frequency vibratory stimuli in tactile pattern perception. Finally, the view is presented that the skin and subjacent tissues should be considered, along with cutaneous mechanoreceptors, as forming a tactile receptor organ system.     

Neurophysiology of Ganglia of Auerbach's Plexus

The enteric plexuses of the automatic nervous system may beconsidered, on the basis of both function and morphology, tobe a simple integrative nervous system of vertebrate animals.Microelectrcde studies of single unit activity within entericganglia reveal four distinct types of ganglion cells distinguishedon the basis of pattern of spike discharge. These are (i) burst-typeunits which spontaneously discharge bursts of spikes at periodicintervals; (ii) fast- and slowly-adapting mechanoreceptors;(iii) tonic-type units which respond to mechanical stimulationwith prolonged, all-or-nothing trains of spikes; (iv) single-spikeunits which spontaneously discharge single action potentialsat variable intervals. The enteric plexuses are adapted forcontrol of the intestinal musculature which behaves as an electricalsyncytium activated by myogenic pacemaker potentials. The mechanismof neural control is integration of continuous neurogenic inhibitionof the inherently excitable musculature.     

Discharge patterns of chicken cochlear ganglion neurons following kanamycin-induced hair cell loss and regeneration

Hair cells in the basal, high frequency region (>1100 Hz) of the chicken cochlea were destroyed with kanamycin (400 mg/kg/d × 10 d) and allowed to regenerate. Afterwards, single unit recordings were made from cochlear ganglion neurons at various times post-treatment. During the first few weeks post-treatment, only neurons with low characteristic frequencies (<1100 Hz) responded to sound. Despite the fact that the low frequency region of the cochlea was not destroyed, neurons with low characteristic frequencies had elevated thresholds, abnormally broad U-shaped or W-shaped tuning curves and low spontaneous discharge rates. At 2 days post-treatment, the spontaneous discharge rates of some acoustically unresponsive units fluctuated in a rhythmical manner. As recovery time increased, thresholds decreased, tuning curves narrowed and developed a symmetrical V-shape, spontaneous rate increased and neurons with higher characteristic frequencies began to respond to sound. In addition, the proportion of interspike interval histograms with regularly spaced peaks increased. These improvements progressed along a low-to-high characteristic frequency gradient. By 10–20 weeks post-treatment, the thresholds and tuning curves of neurons with characteristic frequencies below 2000 Hz were within normal limits; however, the spontaneous discharge rates of the neurons were still significantly lower than those from normal animals.Abbreviations KM kanamycin - BrdU bromodeoxyuridine - CF characteristic frequency - CAP compound action potential - ISI interspike interval     

Processing of mechanosensory information from gustatory receptors on a hind leg of the locust

Gustatory receptors (basiconic sensilla) on the legs of the desert locust, Schistocerca gregaria, are innervated by chemosensory afferents and by a mechanosensory afferent. We show, for the first time, that these mechanosensory afferents form an elaborate detector system with the following properties: 1) they have low threshold displacement angles that decrease with increasing stimulus frequency in the range 0.05–1 Hz, 2) they respond phasically to deflections of the receptor shaft and adapt rapidly to repetitive stimulation, 3) they encode the velocity of the stimulus in their spike frequency and have velocity thresholds lower than 1°/s, and 4) they are directionally sensitive, so that stimuli moving proximally towards the coxa elicit the greatest response.The mechanosensory afferents, but not the chemosensory afferents, make apparently monosynaptic connections with spiking local interneurones in a population with somata at the ventral midline of the metathoracic ganglion. They evoke excitatory synaptic potentials that can sum to produce spikes in the spiking local interneurones. Stimulation of the single mechanosensory afferent of a gustatory receptor can also give rise to long lasting depolarizations, or to bursts of excitatory postsynaptic potentials in the interneurones that can persist for several seconds after the afferent spikes. These interneurones are part of the local circuitry involved in the production of local movements of a leg. The mechanosensory afferents from gustatory receptors must, therefore, be considered as part of the complex array of exteroceptors that provide mechanosensory information to these local circuits for use in adjusting, or controlling locomotion.     

Tactile receptor discharge and mechanical properties of glabrous skin

Current knowledge of the functional properties of mammalian cutaneous mechanoreceptors is reviewed with special reference to receptors associated with the glabrous skin of the raccoon and squirrel monkey hand. Four physiologically defined mechanoreceptor types are recognized: Pacinian afferents, rapidly adapting (RA), and slowly adapting type I (SAI), and slowly adapting type II (SAII). The SAI category is divided into moderately slowly adapting and very slowly adapting (VSA) types in terms of the duration of their response to a prolonged mechanical displacement of skin. Although both RA and SA units are capable of signaling displacement ramp velocity, the pattern of discharge during ramp stimulation may vary widely among units. SAI units also code the depth of skin displacement, but there is no best-fitting function describing the relationship. Static discharge is also markedly influenced by prior ramp velocity. Both raccoon and squirrel monkey VSA units show wide variation in the regularity of their discharge during static displacement. The rate of adaptation of SAI units is less when constant force stimuli are applied to the skin than when constant displacement stimuli are applied. This is partly attributable to mechanical properties of the skin. When either constant force or constant displacement stimuli are spaced too closely in time, there is a progressive (trial-to-trial) decrement in response rate, accounted for in part by failure of the skin to recover to its initial resting level.     

Branchial mechanoreceptor activity during spontaneous ventilation in channel catfish

Extracellular afferent neural activity was recorded in vivo from cranial nerve IX (glossopharyngeal) from mechanoreceptors in the first gill arch of anesthetized, spontaneously breathing channel catfish (Ictalurus punctatus). Single unit and paucifiber recordings show that both phasic and tonic receptors were active during normal ventilation. Phasic receptors were characterized as having a burst of activity during some phase of the ventilatory cycle. Most of these occurred during peak adduction or peak abduction. Phasic receptors were not active during spontaneous apnic periods. Tonic receptors were always active, even during apneas, firing frequency was modulated by breathing movements with peak activity occurring during adduction. Flow-sensitive mechanoreceptors were identified in anesthetized, paralyzed catfish. These receptors decreased activity when the ventilatory water flow was stopped. Hypercapnia (5% CO(2) in air) stimulated ventilatory rate and amplitude but had no effect on mechanoreceptor activity. The discharge characteristics of branchial mechanoreceptors indicate that they could be involved in the timing and coordination of ventilatory movements and maintenance of the 'gill curtain' to minimize ventilatory dead space. Unlike ventilatory mechanoreceptors in the air breathing organs of gar and lungs of lungfish and tetrapods, branchial mechanoreceptors were insensitive to hypercapnia.     

Electrophysiological identification of cold receptors on the antennae of the ground beetle Pterostichus aethiops

Abstract.  In single-sensillum extracellular electrophysiological recordings, terminal campaniform sensilla at the tip of antennae of the ground beetle Pterostichus aethiops (Pz., 1797) show action potentials of three sensory cells, A-, B- and C-cells, distinguished by differences in their spike amplitudes. Only the A-cell, with the largest spike amplitude, is highly sensitive to temperature fluctuations, showing remarkable changes in its firing rate induced by changes in temperature of 0.1 °C. The firing rate of A-cells at 23 °C varies from 15–52 Hz among different beetles. Mean impulse frequency of A-cells is found to be a function of steady temperature, the firing rate decreasing with temperature increase. A-cells respond to a rapid temperature drop with a strong phasic-tonic reaction; larger decreases in temperature evoke higher peak frequency values. Maximum peak frequencies, varying from 380–630 Hz in different beetles, are induced by temperature decreases of 3–10 °C, whereas temperature rise strongly inhibits impulse activity of the A-cell. The first manifestation of rapid warming in the nerve impulse sequence is a very long interspike period, followed by diminished activity. Both the length of the long interspike period and the rate of following impulse activity are functions of temperature change; hence, A-cells respond to temperature changes as typical cold receptors, similar to coeloconic and short hair-like sensilla in other insects.     

Neurophysiology of the vibration sense in locusts and bushcrickets: Response characteristics of single receptor units

The response characteristics of the vibration receptors in the legs of the migratory locust, Locusta migratoria, and the tettigoniid Decticus verrucivorus were investigated electro-physiologically by single cell recordings. The legs were stimulated by sinusoidal vibrations. There are four types of vibration receptor in each leg of Locusta and Decticus, which can be classified physiologically. One type—most probably campaniform sensilla—shows a phase-locked response to vibrations from 30 to 200 Hz, its threshold reflecting the displacement. A second type shows similar responses in the same frequency range, but its reactions depend on the stimulus acceleration. The receptor cells of the subgenual organ are very sensitive to vibration from 30 to at least 5000 Hz, and their responses depend on acceleration. There are two types of subgenual receptors, one of which shows a clear maximum of sensitivity between 200 and 1000 Hz, with a threshold below 0.01 m/sec^?2 acceleration. Subgenual receptors with different thresholds and different characteristic frequencies occur in each leg. The receptors of each leg pair have quite similar mean sensitivities and characteristic frequencies. However, in the front legs of tettigoniids the more sensitive subgenual receptors and an additional receptor type also respond to low-frequency airborne sound up to 10 kHz.     

Effects of temperature on a moth auditory receptor

Measurements of the thoracic temperature and recordings of the spike activity of the most sensitive auditory receptor (A1 cell) were made in Empyreuma pugione (Arctiidae, Ctenuchinae). The temperature range tested (19–36 °C) is relevant for the behavior and ecology of this species. Experiments were performed during the hours of maximal flying activity in the wild: sunrise and sunset. The thoracic temperature during rest reflects that of the surrounding air; there is an increase of 3–4 °C immediately after ceasing free flying in the laboratory. The spike activity of the tympanic organ was recorded with a stainless-steelhook electrode placed beneath the tympanic nerve in the mesothorax. The A1 cell activity was studied without acoustic stimulation (spontaneous) and in response to 35-kHz acoustic pulses of 20, 40, or 100 ms duration. At all of these durations A1 cell response to saturating stimulus was analysed, while with 40-ms pulses different stimulus intensities were used (20–90 dB SPL in 10-dB steps). The number of action potentials per pulse, mean spike rate, maximal instantaneous discharge, and latency period depend strongly on air temperature, while the variation coefficients of the interspike intervals during the responses were not temperature dependent and vary non-monotonically with stimulus intensity. During responses to a saturating stimulus, the stimulus duration does not affect the activation energy, calculated from an Arrhenius plot, of different physiological features. Adaptation, studied in the responses to 100-ms pulses, is also temperature dependent. This phenomenon has two components, each of which shows different activation energies, suggesting a different membrane origin. High stimulus intensity (90 dB SPL) significantly affects the activation energy of the action potentials and mean spike rate, while the activation energy, of the maximal instantaneous discharge and latency period do not show this strong dependency. The spontaneous A1 cell spike rate varies with temperature, as does the value of the mode of the relative frequency distribution of the interspike interval. The activation energy of the spike rates measured at A1 cell responses to saturating stimuli is in good agreement with that described in amphibian innerear hair cells. It is suggested that this moth auditory receptor cell also has mechanosensitive protein channels.Abbreviations AP/p action potentials per pulse - AP/s action potentials per second - CI confidence interval - E _a activation energy - ISI interspike interval - SD standard deviation - VC variation coefficient     

Local anesthetic action of phentolamine on insect mechanoreceptors

1. The effect of phentolamine on the response properties of insect mechanoreceptors and on the conduction in their axons was examined using electrophysiological techniques. 2. Phentolamine blocked conduction of action potentials along axons, an effect which exhibited 3 characteristics typical of local anesthetics: the effect was frequency-dependent, reversible and varied for nerves with different diameters. 3. The concentration of phentolamine required to block axonal conduction (1-2 x 10(-3) M) was significantly higher than that required to abolish the response of receptors to mechanical stimulation (3-5 x 10(-4) M). 4. All mechanoreceptors that were examined in Locusta migratoria and Periplaneta americana were inactivated by phentolamine (Table 1). The type I receptors (chordotonal, campaniform and hair sensilla) were inactivated within 5-15 min following phentolamine application. The only type II receptor examined (forewing stretch-receptor) underwent a phase of repetitive discharge before being inactivated. 5. Tolazoline and metoclopramide inactivated, like phentolamine, mechanoreceptors at lower concentrations than necessary to block axonal conduction. However, yohimbine and chlorpromazine inactivated mechanoreceptors and blocked axonal conduction at similar concentrations. 6. These findings suggest that phentolamine affects sense-organ specific ionic processes that are more sensitive to the drug than the ionic processes along the axons.     

Demonstration of mucous mechanoreceptors in the lower esophageal sphincter. Comparison with muscle mechanoreceptors

In anaesthetized cats, vagal unitary discharges originating from the Lower Oesophageal Sphincter (L.O.S.) were recorded in nodose ganglia by means of glass microelectrodes. Numerous mechanoreceptors located both in mucosa and muscular layers were found in L.O.S. The mucus mechanoreceptors (high threshold receptors) were activated by strong compressions and distensions, by rapid passage of liquid through the oesophagus and by striking the mucosa. The muscular mechanoreceptors (low threshold receptors) responded to contraction and distension of L.O.S. Both receptors were connected to nonmyelinated fibres (conduction velocity: 0.9-1.4 m/sec).     

Some Models of Neuronal Variability

The pattern of nerve action potentials produced by unit permeability changes (quantal inputs) occurring at random is considered analytically and by computer simulation methods. The important parameters of a quantal input are size and duration. Varying both the mean and the probability density function of these parameters has calculable effects on the distribution of interspike intervals. Particular attention is paid to the relation between the mean rate of excitatory inputs and the mean frequency of nerve action potentials (input-output curve) and the relation between the coefficient of variation for the interval distribution and the mean interval (variability curve). In the absence of action potentials one can determine the parameters of the voltage distribution including the autocorrelation function and the power spectrum. These parameters can sometimes be used to approximate the variability of interspike intervals as a function of the threshold voltage. Different neuronal models are considered including one containing the Hodgkin-Huxley membrane equations. The negative feedback inherent in the Hodgkin-Huxley equations tends to produce a small negative serial correlation between successive intervals. The results are discussed in relation to the interpretation of experimental results.     

Neuromodulation of central inhibition on peripheral mechanosensory afferents

100 g reserpine, suspended in sunflower oil, was injected into the crop of six leeches (Hirudo medicinalis). Five leeches were injected with oil alone. The efficacy of delivering reserpine in this manner was assessed by measuring swimming bout duration. Four days after injection, swimming bouts were significantly longer in reserpine-treated leeches than in oil-treated or normal leeches. The amplitude of excitatory potentials elicited in velocity sensitive skin mechanoreceptors (T cells) by action potentials in mechanosensory cells which respond to sustained deformation of the skin (P cells) was no different in reserpine-treated leeches than in untreated leeches. The amplitude of inhibitory potentials in these T cells, produced by interneurons excited by P cells, was significantly larger in reserpine-treated leeches. Depletion of monoamines enhances inhibition in T cells. This is consistent with the established roles played by monoamines in promoting leech swimming and feeding behaviour.     

The sensory basis for the trap-jaw mechanism in the ant Odontomachus bauri

Ants of the ponerine genus Odontomachus have evolved a mechanism that allows them to instantaneously close their long mandibles to catch prey or defend themselves. This trap-jaw action is triggered by contact of trigger hairs with a potential prey item. Two of these long mechanosensory hair sensilla reside proximally on each mandible and are supplied by giant sensory cells.Extracellular recordings demonstrate that the sensory cells respond to tactile stimulation. Their phasic responses encode amplitude and velocity of hair-deflection away from the midline, but not hair position. The discharge of action potentials follows stimulus frequencies of more than 300 Hz. During sinusoidal stimulation, the cells adapt very little, sustain discharge rates of more than 200 Hz for more than 20 s, and reach peak spike rates of about 450 Hz.The afferent axons of these sensory cells give rise to huge axon terminals within the suboesophageal ganglion. One of the afferents has a prominent contralateral branch, the other is confined to ipsilateral neuropil. Anatomical data indicate that the 4 afferents may be coupled and may serve as the substrate for a very fast reflex.Abbreviations HRP horseradish peroxidase - LGS lateral giant sensillum - MGS median giant sensillum - SEM scanning electron microscopy - SOG suboesophageal ganglion     

Modulatory effects of oligodendrocytes on the conduction velocity of action potentials along axons in the alveus of the rat hippocampal CA1 region

Like neurons and astrocytes, oligodendrocytes have a variety of neurotransmitter receptors and ion channels. However, except for facilitating the rapid conduction of action potentials by forming myelin and buffering extracellular K(+), little is known about the direct involvement of oligodendrocytes in neuronal activities. To investigate their physiological roles, we focused on oligodendrocytes in the alveus of the rat hippocampal CA1 region. These cells were found to respond to exogenously applied glutamate by depolarization through N-methyl-D-aspartate (NMDA) receptors and non-NMDA receptors. Electrical stimulation of the border between the alveus and stratum oriens evoked inward currents through several routes involving glutamate receptors and inward rectifier K(+) channels. Moreover, electrical stimulation resembling in vivo activity evoked long-lasting depolarization. To examine the modulatory effects of oligodendrocytes on neuronal activities, we performed dual, whole-cell recording on CA1 pyramidal neurons and oligodendrocytes. Direct depolarization of oligodendrocytes shortened the latencies of action potentials evoked by antidromic stimulation. These results indicate that oligodendrocytes increase the conduction velocity of action potentials by a mechanism additional to saltatory conduction, and that they have active roles in information processing in the brain.     

Spontaneous activity of passive muscle spindles of the cat triceps surae muscle

The hypothesis that one possible cause of the spontaneous discharge of muscle spindles in the totally relaxed and de-efferented triceps surae muscle, with its tendon divided, is the mechanical factor due to extrafusal-intrafusal interaction was tested in experiments on anesthetized cats. Responses of spontaneously active units were similar with respect to many indices to responses of silent receptors capable of generating a long discharge in response to an increase in static length of the muscle. Isotonic contraction of the relaxed muscle during direct or indirect stimulation was accompanied by a pause in spontaneous activity. The prolonged increase in discharge frequency sometimes arising as a result of pressure on the muscle in the region of the receptor also was abolished by weak isotonic contraction of the muscle. If a long posttetanic sensory discharge was induced in a spontaneously active receptor by intensive tetanization of the nerve to the muscle, and it was then abolished by short stretching of the muscle, the initial spontaneous discharge frequency was restored. The dynamic thresholds of spontaneously active receptors were lower than for silent receptors. In some spontaneously active receptors an initial slowing of the discharge was observed during linear or stepwise stretching of the muscle. It is suggested that the ability of sensory endings to generate a long spontaneous discharge is due to initial stretching of the spindle inside the relaxed muscle.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 10, No. 3, pp. 287–294, May–June, 1978.     

Mechanoreceptors innervating soft cuticle in the abdomen of the hermit crab, <Emphasis Type="Italic">Pagurus pollicarus</Emphasis>

Mechanoreceptors in the soft cuticle of the 4th abdominal segment of the hermit crab, Pagurus pollicarus, that are associated with reflex activation of abdominal postural motoneuron, were studied to determine whether their properties are consistent with a feedback control of abdominal stiffness. Three classes of receptors were identified: (1) setal dome receptors, (2) hypodermal receptors, and (3) funnel-canal receptors. The hypodermal receptors, which have the largest extracellular action potentials, were selected for further study. Their axons innervate the entire ipsilateral half of a segment; receptive fields of receptors with different amplitudes show extensive overlap. They are phasic and show significant adaptation; at higher frequencies they signal displacement rather than velocity. Although they are activated by changing muscle tension, their threshold for cuticular displacement is much lower than for forces generated by postural muscles. These features suggest that they are primarily involved in signaling cuticular displacement and shearing forces as they contact the columella of the shell in which the hermit crab lives.     

Perception and the Characteristics of the Response of Honey Bees,Paper Wasps,and Red Wood Ants to a Low-Frequency Electric Field

Bees, wasps, and ants have no specialized receptors for the perception of an electric field. An appropriate response to naturally occurring electric fields in bees and ants is associated with atmospheric exposure, amplified by the approach of the front of a thunderstorm. The primary transducers of mechanoreceptors that respond to displacement are related to the perception of low-frequency electric fields of high intensity by insects. The non-specific mechanism of perception of electric fields is based on irritation by induced currents that flow in the locations of their contact with each other and/or conductive surfaces. The frequency dependence of the electric field sensitivity is determined mainly by the magnitude of the current induced by it and the intensity of its contact action. The magnitude of the current induced in the outer part of the insect body is non-linearly related to the frequency of the electric field. The region with the highest sensitivity to electric fields is close to 500 Hz, which is consistent with the maximum magnitude of the induced current. At the same time, the threshold of the sensitivity to an electric field in wasps is approximately 0.04 kV/m, while in bees it is 0.45 kV/m. Ants react to the action of an electric field of 7–10 kV/m by slowing their movement. Magnetic fields and ionization, which accompany the generation of an electric field whose intensity reaches 15–20 kV/m, do not stimulate changes in the behavior of insects.

     

Adaptive cardiovascular modifications in the western chipmunks genusEutamias

Summary Although cutaneous type I and type II mechanoreceptors in the cat respond at progressively higher frequencies to increasingly rapid skin indentations of suprathreshold intensity, their thresholds are not lowered when these more rapid stimuli are applied. Since these receptors do not selectively detect rapid stimuli of small amplitude, even though they respond much more vigorously to a suprathreshold stimulus that is rapid, different parameters of the stimulus are signalled depending on whether it is near threshold or clearly suprathreshold.This work was supported by grant GB42643 from the National Science Foundation and by grants NS08769, NS07938 and NS05244 from the U.S. Public Health ServiceThe authors thank John Fisher, Gary Frederickson, Tom O'Leary, Robert Perry, and Jane Burgess for their valuable help.     

Cutaneous sensory afferents recorded from the nervus intramandibularis ofGallus gallus vardomesticus

Summary The responses of single sensory afferent nerve fibres were recorded from small nerve bundles of the intramandibular nerve of the chicken following thermal and mechanical stimulation of the beak. Thermoreceptors, nociceptors and mechanoreceptors were identified and their responses characterized.Of the thermoreceptors identified 11 units were classified as cold receptors, which responded to cooling the receptive field by increasing the discharge rate and had conduction velocities in the range 0.83 to 4.4 m/s. Only one warm unit was identified.Two classes of nociceptors were identified: mechano-thermal (polymodal) nociceptors and high threshold mechanical nociceptors. The discharge characteristics and stimulus-response curves of both types were described. While the mechanothermal nociceptors were exclusively C-fibres (c.v. 0.4 to 1.86 m/s), the high threshold mechanoreceptors contained both C and A delta fibres (c.v. 1 to 5.5 m/s). Thermal response thresholds for the mechano-thermal units ranged from 41 to 50 °C with mechanical thresholds of 2 to over 50 g. Mechanical thresholds for the high threshold units ranged from 5 to over 50 g.The mechanoreceptors were either slowly or rapidly adapting. The pattern of response together with stimulus-response curves were presented for the slowly adapting units. Conduction velocities of the slowly adapting units varied from 0.7 to 20 m/s and mechanical threshold from 0.1 to 2 g. On the basis of their response to a vibrating, and a ramp-and-hold mechanical stimulus, the rapidly adapting units were divided into Herbst and Grandry units with only the Herbst units responding accurately to the vibrating stimulus. Both units had fibres conducting in the 50 m/s range with thresholds in the 0.1 to 10 g range.The results are discussed in relation to the receptors found in other avian species and mammalian peripheral sensory afferents.Abbreviations c.v. conduction velocity - RA rapidly adapting (receptors) - SA slowly adapting (receptors)