Hair Follicle Afferents Research Articles

Physiological properties of primary sensory neurons appropriately and inappropriately innervating skin in the adult rat

1. We have studied the physiology of sensory neurons innervating skin of the rat hindlimb, in three groups of animals: 1) normal animals; 2) animals in which the sural nerve (Sn) had regenerated to its original cutaneous target; and 3) animals in which the gastrocnemius muscle nerve (Gn) had previously been cut and cross anastomosed with the distal stump of the cut Sn so that its axons regenerated to a foreign target, skin. 2. Single-unit recordings were made from 222 afferents in normal, intact animals. They had conduction velocities of 0.5-53.1 m/s. The conduction velocity distribution had distinct peaks at approximately 37.5, 2.5, and 1.25 m/s, presumably corresponding to A alpha beta-, A delta-, and C-fiber populations. Eighty-two percent of the characterized myelinated fibers had low-threshold mechanosensitive receptive fields, whereas 16% were high threshold, and only 2% appeared to have no receptive field. The very large majority of low-threshold mechanosensitive receptive fields (87%) were rapidly adapting hair follicle afferents. 3. In animals with regenerated Sn, 308 afferents were recorded with conduction velocities of 0.4-58.8 m/s. However, the mean conduction velocity was lower than in control animals (P less than 0.05), and only one peak, at 27.5 m/s, was apparent for myelinated fibers. Eighty-six percent of myelinated fibers were low-threshold mechanosensitive afferents, 8.5% were high-threshold mechanoreceptors (HTMRs), and 5.5% appeared to have no receptive fields. Fewer low-threshold mechanoreceptors (LTMRs; compared with controls) were activated by hair movement (63 vs. 87%). Most of the remainder appeared to be field receptors (which were therefore more commonly observed here than in normal animals). 4. In animals in which the Gn had regenerated to skin, 430 afferents were recorded. These had conduction velocities ranging from 0.6 to 71.4 m/s, and again only one peak was apparent in the myelinated conduction velocity histogram, at approximately 17.5 m/s. Of the myelinated fibers, 79% had low-threshold mechanosensitive receptive fields in skin and 10% high-threshold mechanosensitive receptive fields. The remaining 11% apparently had no receptive field (cf. 5.5% in regenerated Sn). In contrast to normal or regrown sural afferents, only 58% of low-threshold gastrocnemius afferents in skin were rapidly adapting. Of the 42% slowly adapting afferents, many surprisingly responded to hair movement. Thus some gastrocnemius afferents seemed to have retained the adaptation properties characteristic of muscle afferents. Also surprisingly, given that the Gn contains fewer fibers than the Sn, receptive-field areas were not significantly different from regrown or normal sural fibers.(ABSTRACT TRUNCATED AT 400 WORDS)

Collateral sprouting of the central terminals of cutaneous primary afferent neurons in the rat spinal cord: Pattern, morphology, and influence of targets

The capacity of the central terminals of primary afferents to sprout into denervated areas of neonatal spinal cord and the morphology of any novel terminals has been investigated. In rats which had undergone sciatic nerve section on the day of birth, 12 of 18 physiologically characterized intact saphenous hair follicle afferents (HFAs) were labelled intra-axonally with horseradish peroxidase (HRP) were shown to sprout up to 2,000 microns into the deafferented sciatic terminal field. The morphology of these sprouts depended on which area of the sciatic nerve territory was invaded by the afferent sprouts. Six HFAs sprouted into areas normally innervated by glabrous skin afferents and the morphology of the collateral sprouts in this region resembled that of rapidly adapting (RA) afferents. The other six saphenous HFAs had sprouted into sciatic "hairy" skin areas and the morphology of these sprouts, although abnormal, was flame shaped. In rats whose sural, saphenous, and superficial peroneal nerves were cut at birth, 4 of 7 single HRP labelled RA afferents had central terminals that had sprouted into regions of cord normally devoted to "hairy" input. These showed clear signs of HFA morphology despite their peripheral receptive fields remaining in the glabrous skin. The results show collateral sprouting of single cutaneous sensory afferent axons into adjacent inappropriate central target regions following neonatal deafferentation. Such plasticity may provide some compensation following neonatal injury. The morphology of the sprouted terminals is appropriate to the new target area rather than to its functional class and is also independent of the peripheral receptive field location providing an example of central rather than peripheral control over afferent growth patterns.

Neonatal capsaicin treatment induces invasion of the substantia gelatinosa by the terminal arborizations of hair follicle afferents in the rat dorsal horn

Capsaicin, administered on the day of birth, was found to alter laminar distribution, but not the receptive field properties or the morphology of the collateral arborizations of hair follicle afferents (HFAs) intra-axonally injected with horseradish peroxidase (HRP). Of the 65 HFA terminal arbors in capsaicin treated rats, 46 (71%) were found to enter the substantia gelatinosa (in control rats, 44/165, 27%). All of the collaterals projected to somatotopically normal areas of cord. Dorsal horn shrinkage (21%), as estimated by planimetric measurements of Nissl and acetylcholinesterase-stained material, was only a partial explanation of this result. This idea was supported by the statistically significant increase (27%, P less than 0.05) in the absolute dorsoventral length of collaterals. The results show that the destruction of unmyelinated fibres during the early postnatal period by capsaicin induces HFA invasion into the area that C fibres normally occupy. This invasion suggests that the laminar termination sites for different primary afferent fibres are not altogether specified and that intact neonatal primary afferents have the capacity to sprout into denervated regions of spinal cord.

Correlation of monosynaptic field potentials evoked by single action potentials in single primary afferent axons and their bouton distributions in the dorsal horn

The relationship between structure and function of the projections of single identified primary cutaneous axons was investigated by recording cord dorsum potentials at 4 sites in response to electrical stimulation of the single axon and visualizing the boutons of the axon stained by intracellular injection of horseradish peroxidase. The rostrocaudal extent of boutons differed from fiber to fiber ranging from 4.14-11.50 mm; their location in the dorsal horn also varied in agreement with the known somatotopy of the presynaptic neuropil and dorsal horn neurons. Rostrocaudal distributions of cord dorsum potentials and boutons of individual fibers revealed good agreement. Cord dorsum potential amplitude and length of the spinal projection were positively correlated with number of boutons, but no correlation with bouton density was found. The spinal projection of afferents innervating slowly adapting type 1 mechanoreceptors exhibited a greater rostrocaudal extent (mean: 8.48 mm) than those innervating rapidly adapting mechanoreceptors (i.e., hair follicle and field receptors: mean: 5.87 mm). Although the mean total number of boutons was greater for axons with slowly adapting receptors (7,250/fiber) than for axons of rapidly adapting receptors (4,677/fiber), no differences in the longitudinal density of boutons (boutons/mm) were observed. Likewise, summed amplitudes of cord dorsum potentials at the 4 recording electrodes were larger for SA1 afferents than for those of field and hair follicle afferents. A major role for the number of boutons in determining these differences is supported by the finding that the calculated average contribution per bouton to cord dorsum potentials (expressed as an amplitude coefficient a) was similar for slowly and rapidly adapting afferents. No evidence was found for regions in which boutons did not contribute to the cord dorsum potential.

Structure-function relationships in identified afferent neurones

The review deals with structure-function relationships in primary afferent and spinal cord neurones that were intracellularly injected with a marker substance (mostly HRP) after physiological identification. At the level of dorsal root ganglion (DRG) cells, there is a significant correlation between soma size and conduction velocity (or diameter) of the afferent fibre for most subpopulations of DRG cells, but the scatter of data is considerable, so that the size of a DRG cell soma cannot be predicted from the diameter of its axon or vice versa. The spinal terminations of primary afferent fibres are the best example of a relationship between structure and function, since most of the afferent units possess characteristic patterns of spinal arborization, e.g. the "flame-shaped arbors" of hair follicle afferents in lamina III of the dorsal horn, or the projection of nociceptive afferents onto lamina I. The morphological features of spinal cord neurones can be used only to a limited extent for functional identification. Thus, many SCT neurones can be recognized by their triangular dendritic tree and STT cells in lamina VII/VIII by their dendritic projection into the white matter. It is still not possible, however, to distinguish a nociceptive STT cell from a low-threshold mechanoreceptive one on the basis of morphological criteria.

Morphology and somatotopic organization of the central terminals of hindlimb hair follicle afferents in the rat lumbar spinal cord

The morphology of the central collateral arborizations of 24 A-beta hair follicle afferents (HFAs) innervating different regions of the skin of the hindlimb were studied by the intra-axonal injection of horseradish peroxidase (HRP) in adult rats. A total of 236 collaterals were recovered. These fell into three classes--complex, simple, and blind-ending--based on numbers of boutons and terminal branch patterns. The morphology of the HFA central arbors innervating the lateral and medial leg and dorsum of the foot was flame-shaped. Afferents with receptive fields on the glabrous-hairy skin border consistently had extra terminal branches running ventromedially into laminae IV/V. Differences in the width of terminal arbors were found. HFA terminals innervating the lateral leg formed narrower sheets than those innervating the dorsum of the foot and toes. The somatotopic organization of the collaterals and terminal arborizations of individual afferents were analyzed both by considering all the collaterals along an axon's rostrocaudal extent and by only examining arbors with boutons (the complex and simple arbors). Thirty-seven percent of blind-ending and 18% of simple collaterals were found to overlap in the rostrocaudal direction with the complex arborizations of afferents whose receptive fields were in a different cutaneous nerve territory. There was no overlap between complex arborizations of afferents from different nerve territories. However, the complex arbors of afferents with receptive fields within a particular nerve territory showed considerable terminal overlap even if they had nonadjacent peripheral receptive fields. The topographical organization of the central terminals of HFAs, forms a coarse somatotopic map of overlapping terminals whereby a particular region of dorsal horn has a maximal, but not exclusive, input from a particular area of skin.

Facilitation of individual gamma-motoneurones by the discharge of single slowly adapting type 1 mechanoreceptors in cats.

1. Cross-correlation of the discharges of individual neurones has been used to investigate the influence of identified cutaneous afferents on gamma-motoneurones below the level of complete spinal section in decerebrated cats. Discharges of single, sural nerve afferents from the heel were recorded in the dorsal root ganglia. Discharges of gamma-motoneurones were recorded from cut filaments of the muscle nerve to gastrocnemius medialis of the same leg. gamma-Motoneurones had a background discharge in the absence of intentional stimulation. 2. Correlograms involving slowly adapting afferents were formed during steady application of a probe to the receptive field for repeated periods of 10 s. Afferent synchronization was minimized by rejecting any period of probe movement. Correlograms involving rapidly adapting afferents required continuous movement of the probe to sustain afferent discharge. 3. Statistically significant primary peaks in correlations were observed for twenty-one pairings of gamma-motoneurones with seventeen out of thirty-nine slowly adapting, type 1 (SA1) mechanoreceptors. Primary peaks had widths at half-maximum in the range of 2-7 ms. No such short duration peaks were seen for fourteen pairings of gamma-motoneurones with eleven slowly adapting type 2 (SA2) receptors or for thirty-five pairings with twenty-nine hair follicle (HF) afferents. Broad correlations with peaks extending over tens of milliseconds were seen for HF afferents and could be generated in correlograms for slowly adapting afferents by moving the probe. 4. The short duration peaks were delayed with respect to the SA1 afferent discharges. Subtraction of peripheral conduction times gave central delays for the increased probability of gamma-motoneurone firing ranging from 2.7 to 6.5 ms (mean = 4.0 ms). These values were not significantly different from the central delays of gamma-motoneurone excitation in response to electrical stimulation of the sural nerve at a strength 1.2 times threshold. 5. The increase in probability of gamma-motoneurone discharge given a single SA1 afferent discharge ranged from 0.005 to 0.156 with a mean value of 0.052. The rise time of the peak ranged from 1 to 4 ms with a mean value of 1.9 ms (n = 9). 6. The properties of the correlogram peaks were not related to the axon conduction velocity of either the SA1 afferent or gamma efferent neurones. 7. The SA1 afferents that facilitated gamma-motoneurone discharge had axon conduction velocities in the range 29-81 m/s and could not be distinguished from SA1 afferents lacking correlations.(ABSTRACT TRUNCATED AT 400 WORDS)

Gamma-aminobutyric acid and afferent inhibition in the cat and rat ventrobasal thalamus

Extracellular single neuron recordings were made in the ventrobasal thalami of anaesthetized rats and cats. Physiological stimulation of vibrissa and hair follicle afferents was performed with an air jet (10–20 ms duration) directed at a single vibrissa or small area of hairy skin. Paired conditioning and test air jets delivered to the excitatory portion of receptive fields revealed inhibition of the response of ventrobasal thalamic neurons to test stimuli following the excitatory response to the conditioning stimulus. Such inhibitions could last up to 500 ms. An increase in neuronal excitability was sometimes observed following this inhibitory period. In addition, it was possible to produce inhibition without an excitatory response using conditioning stimuli delivered adjacent to the excitatory receptive field. Iontophoretic application of bicuculline methochloride, with currents that were adequate to antagonize iontophoretically applied GABA, was found to reduce the inhibition of test responses evoked by conditioning stimuli in almost all of the neurons studied. In most cases, no excitatory responses to conditioning stimuli directed outside the original excitatory receptive field were revealed by application of the GABA antagonist. In rats, bicuculline also led to a decrease in the post-inhibitory excitation, whereas in cats the converse appeared to be the case. These results suggest that GABAergic transmission may underlie inhibitory responses of cat and rat ventrobasal thalamus neurons to physiological stimulation of somatosensory afferents. Furthermore, removal of such inhibition does not appear to reveal excitatory inputs from outside of the original excitatory receptive field.

Functional specialization of central projections from identified primary afferent fibers.

1. Somata of primary afferent fibers were impaled in the L7 or S1 dorsal root ganglion in cats anesthetized with alpha-chloralose. Individual cells (n = 182) were characterized according to receptive field (RF) and by the peripheral mechanoreceptor they innervated. They were then stimulated intracellularly while recording the evoked cord dorsum potentials (CDPs) simultaneously at four sites. CDPs were recorded in response to single fiber stimulation while varying both the frequency and the numbers of action potentials (APs) evoked per trial. Stimulus parameters included: 1) single APs evoked at both high (18 Hz) and low (0.67 Hz) frequencies, 2) pairs of APs (50-ms ISI) delivered at 0.67 Hz, and 3) trains of four APs (20-ms ISI) also delivered at 0.67 Hz. The properties of the CDPs and their relationship to receptor type innervated by the fiber were determined. 2. CDPs evoked by 18-Hz stimulation consisted of an axon fiber spike followed at a short latency [600 +/- 9.9 (SEM) microseconds] by the onset of a monophasic negative wave. A-beta-afferent fibers innervating slowly adapting type 1 and 2 receptors (SA1 and SA2) evoked the largest amplitude CDPs followed by field and hair follicle afferents while A-delta-fibers rarely produced measurable CDPs at this frequency of stimulation. 3. The magnitude of the CDP varied at the four recording sites with clear evidence in individual experiments that the projections of individual fibers are somatotopically organized. 4. CDPs evoked by 0.67-Hz stimulation had the same onset latency but were larger in amplitude and had longer time-to-peak and durations than those elicited by high frequency stimulation. Under these conditions A-beta-fibers innervating hair follicles produced the largest CDPs followed by field receptors and SA1s and SA2s. A-beta- and A-delta-fibers innervating high threshold mechanoreceptors (HTMRs) and A-delta-fibers innervating down hair follicles (D-hairs) produced CDPs of amplitude and duration similar to those evoked by slower A-beta-fibers. 5. The response to a test stimulus delivered 50 ms after a conditioning stimulus to the same single fiber was always depressed in the case of A-beta-fibers supplying rapidly adapting receptors. The conditioning stimulus exerted relatively little effect on the response to the test stimulus for A-beta- fibers innervating slowly adapting receptors. The test response to stimulation of A-beta- and A-delta- fibers innervating HTMRs was consistently facilitated while stimulation of A-delta- D-hairs evoked either marked facilitation or slight depression.(ABSTRACT TRUNCATED AT 400 WORDS)

Structure and development of central arborizations of hair follicle primary afferent fibers.

The present study describes the structure and development of the flame-shaped central arborizations of hair follicle (HF) afferents in the lumbosacral spinal cord of the rat. Tissue was processed according to the rapid Golgi method at successive stages of development from embryonic day 17 through postnatal day 30. Collaterals of most HF afferents were found to enter the dorsal horn via a characteristic U-shaped pathway which often parallels the vascular pattern. The HF collaterals can first be identified at embryonic day 19 and by postnatal day 5 they have established the dorso-ventral and rostro-caudal limits of their field of arborization. Dorsally the arbors extend no further than the inner zone of lamina II (IIi) at any stage of development. Short aberrant branches were observed on some HF collaterals during the prenatal period but none of these developed synaptic terminals or contributed branches to other arbors. Each HF collateral formed a single well defined flame-shaped arbor with a distribution and branching pattern which could be distinguished from that of other afferents throughout the postnatal developmental period. Two types of HF collaterals were observed. Structure and distribution patterns suggest that type I collaterals are derived from G and T HF afferents while collaterals in the type II category are probably derived from both G and T as well as D (A-delta) HF afferents. Type I collaterals divide into well defined medial and lateral collateral branches which arborize mainly in lamina III with a few branches to lamina IV. Type II collaterals are characterized by a delicate arbor which is more vertically oriented than type I arbors.(ABSTRACT TRUNCATED AT 250 WORDS)

Central terminations of cutaneous mechanoreceptive afferents in the rat lumbar spinal cord.

The morphology in the dorsal horn of the lumbar spinal cord of the collateral branches and terminal arborizations of three different types of low-threshold mechanoreceptor innervating the skin of the rat hindlimb has been studied by the intracellular injection of horseradish peroxidase into physiologically characterized afferent fibres. The central terminals of five rapidly adapting glabrous skin mechanoreceptors (RA), six hair follicle afferents (HFA), and four slowly adapting type I afferent fibres (SA I; two from glabrous and two from hairy skin) were recovered for detailed analysis. The number of collaterals per axon varied from eight to 12, and the length of axon stained from 3.5 to 4.7 mm. In each afferent, while the majority (52-67%) of the terminal arborizations displayed extensive branching with large numbers of en passant and terminal synaptic boutons, the arborizations at the caudal and rostral extremes of the terminal field tended to be much simpler and less profuse, with few and in some cases no boutons. This did not appear to be the consequence of inadequate filling. The general pattern of the terminal arborizations was one of mediolaterally compressed, rostrocaudally oriented sheets. In the case of the RA afferents there was no overlap between the adjacent terminal arborizations; a few of the arborizations of the SA I afferents overlapped; most, but not all, of the HFA terminal arborizations overlapped. The terminal arborizations of the HFAs had a distinctive morphology identical to the flame-shaped arbors described in earlier Golgi studies and included synaptic boutons extending from inner lamina II to lamina IV. The morphologies of the RA and SA terminal arborizations were similar to each other, but the former tended to be concentrated in lamina IV with branches in III and V, whereas the latter tended to V with few branches more superficial than lamina IV. There were also differences in the intercollateral spacing between these two types of afferent fibre. While there are similarities between the morphology of the central terminals of cutaneous low-threshold mechanoreceptors in the rat and those previously described in the cat (for example, the longitudinally continuous arrangement of the mediolaterally restricted flame-shaped HFA arborizations and the discontinuous RA arborizations arising from a dorsally located axon), there are also some major differences: the large number of HFA arbors extending to lamina IIi and to lamina IV rather than being restricted to lamina III, the deeper location of the RA arbors (in laminae IV and V rather than lamina III),(ABSTRACT TRUNCATED AT 400 WORDS)

Distribution of activity in the spinal terminations of single hair follicle afferent fibers to somatotopically identified regions of the cat spinal cord

Single-unit spike-triggered averaging has been used to study the distribution of activity in the central processes of 11 single hair follicle afferent (HFA) fibers in relation to the somatotopic organization of dorsal horn neurons (DHNs). Central responses could be recorded from all but one HFA, and the waveforms of these responses were similar to the biphasic, monophasic positive and compound terminal potentials (TPs), the triphasic, positive-negative-positive axonal (preterminal) potentials (APs), and the focal synaptic potentials (FSPs) described by other workers in different preparations. No central responses could be recorded from one HFA even though this axon was shown to be intact throughout the experiment and noise levels in the averaged records were below 3.1 microV. The spatial organization of TPs and APs mirrored the anatomical organization determined using intraaxonal staining only in that the region containing these potentials was a longitudinally orientated narrow (mean width = 405 microns) strip of dorsal horn. Within this strip large TPs, APs, and FSPs were usually found only in those regions in which the receptive field (RF) of the HFA was relatively central to the RFs of DHNs. The region in which the RFs of DHNs encompassed the RF of the HFA, the somatotopically appropriate region, was also organized into a longitudinally orientated strip of dorsal horn of approximately the same width as the strip containing TPs and APs. In any single experiment the strips formed by the somatotopically appropriate region and the TP-AP region occupied the same mediolateral position, but in contrast, the rostral and/or caudal boundaries of these strips often occurred at different levels along the dorsal horn. In some cases the TP-AP strip extended rostrally and/or caudally beyond the somatotopically appropriate region or ended at the same rostrocaudal level. In other cases the somatotopically appropriate region extended rostrally and/or caudally beyond the TP-AP strip. These results are discussed in relation to the rostrocaudal spread of the dendritic trees and the RF organization of DHNs. The results show clearly that in intact cats, anesthetized with alpha-chloralose, some HFAs give rise to collaterals in somatotopically inappropriate regions of the dorsal horn and that at least some parts of these collaterals are invaded by incoming action potentials. The question of whether some HFAs give rise to collaterals that are either infrequently invaded or not invaded at all is discussed.

Receptive field size of hair follicle afferents in rats treated neonatally with capsaicin

Receptive field parameters of G- and D-hair units were investigated in adult rats given capsaicin as neonates. An air jet stimulator mounted on a two-way micromanipulator was used for accurate mapping of receptive fields. Diameters of G- and D-hair unit receptive fields were similar in both control and treated groups averaging 2.14 and 2.44 mm respectively with a 1.5 × threshold stimulus. It is concluded that the reported expansion of rat spinal cord cutaneous receptive fields following capsaicin treatment does not have any significant peripheral component.

Morphology and synaptic relationships of physiologically identified low-threshold dorsal root axons stained with intra-axonal horseradish peroxidase in the cat and monkey.

The arborizations and synaptic relationships of intra-axonally stained horseradish peroxidase- (HRP) labeled primary afferent fibers to the dorsal horn of the cat and monkey spinal cord have been studied by light and electron microscopic methods. The light microscopic arborizations of the afferent fiber types (hair follicle afferents, pacinian corpuscle afferents, type I and type II slowly adapting afferents) are similar to those described by Brown and his colleagues (1) in the cat. The synaptic profiles formed by labeled afferents contain rounded synaptic vesicles. In serial thin sections, it was found that single dorsal root axons may make hundreds or thousands of synapses with neuronal structures of the dorsal horn. The vast majority of synaptic contacts are on the dendritic trees of dorsal horn neurons. The synapses made by these low-threshold afferent axons are almost all in the deeper laminae (III-VI) of the dorsal horn. The hair follicle afferent axons and the pacinian corpuscle afferents have numerous vesicle-containing structures that synapse on them to form either axoaxonal synapses or dendroaxonal synapses. The slowly adapting afferent axons are less often found to be postsynaptic to axons or dendrites. It is concluded that different physiological classes of primary afferent axons have different morphological characteristics, both at the light and electron microscopic level.

Behavior of cutaneous mechanoreceptors recorded in mandibular division of Gasserian ganglion of the rabbit during movements of lower jaw.

1. The activity of cutaneous mechanoreceptors was recorded extracellularly in the mandibular division of the Gasserian ganglion of hemidecerebrate anesthetized rabbits. Fifty-four hair follicle afferents and 80 skin mechanoreceptor afferents were functionally identified. Their receptive-field characteristics were described and their activity patterns were recorded while the jaw was displaced by hand in the vertical, horizontal, and anterior-posterior directions, and during masticatory movements. 2. All hair follicle afferents were classified as rapidly adapting, whereas skin afferents could be divided into two categories: rapidly adapting and slowly adapting. Rapidly and slowly adapting receptors were found in all regions of the mandibular skin. Only one hair follicle afferent and four skin afferents fired spontaneously with the jaw at rest. 3. Fifty-eight percent (29/50) of the hair follicle afferents were activated by imposed displacement of the jaw and all of these tested also discharged during chewing. They were sensitive to movement in all three axes and their firing frequency was linearly related to the velocity, regardless of direction. 4. Only 10% (8/80) of the skin afferents were activated by imposed displacement of the jaw if their receptive fields were not directly contacted. Firing frequency was not well related to either the amplitude or velocity of movement. During chewing, the discharge frequency was variable but, in general, firing was restricted to the jaw-closing phase. Most skin afferents that were active during jaw movement had receptive fields close to the corner of the mouth. 5. The results are discussed within the context of a possible involvement of cutaneous afferents in kinesthesia and in the control of jaw movements. Hair afferents could make specific contributions to the appreciation of movement because their discharge pattern was clearly related to one of its parameters. The lack of activity of most skin afferents, as well as the absence of a strong correlation between the firing frequency of those that were active and the movement parameters, suggest that they do not make specific contributions to the awareness of movement. 6. Hair and skin afferents may also make different contributions to the control of ongoing movements. It is suggested that skin afferent activity during jaw closure may elicit a reflex reduction in the velocity or duration of this phase. Some evidence from other studies suggests that hair afferent activity may indirectly influence movement via effects on elevator fusimotor neurons.

Characteristics of primate spinothalamic tract neurons receiving viscerosomatic convergent inputs in T3-T5 segments.

Characteristics of primate spinothalamic tract neurons receiving viscerosomatic convergent inputs in T3-T5 segments.