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PubMed1990

Physiological characteristics of responses of wide dynamic range spinal neurones to cutaneously applied vibration in the cat.

Extracellular single-unit recordings were made from wide dynamic range neurones in the lumbar dorsal horn of anaesthetized or decerebrated cats. Vibration applied to the skin at a frequency of 80 Hz could evoke 3 distinct types of response--excitation, depression or a biphasic response consisting of excitation followed by depression. By applying vibration at different sites, a given neurone was found to show more than one type of response. Parametric studies of the depressant and biphasic responses were made because previous studies indicated that adenosine mediates the depression in these types of response. Thus, amplitude- and frequency-response relationships were determined at individual stimulation sites: amplitude was varied from 0.001 to 1.0 mm (frequency, 80 Hz) and the frequencies studied were 10, 20, 40, 80, 120 and 240 Hz (amplitude, 0.15 mm). Vibration at amplitudes greater than 0.15 mm caused a decrease in the rate of discharge during the period of stimulation, the magnitude of this decrease varying directly with amplitude; at amplitudes of 0.15 mm and less vibration had no statistically significant effect. With regard to the frequency-response relationship, a decrease in discharge rate occurred at frequencies of 120 and 240 Hz, with the more pronounced effect at 240 Hz; excitation occurred at 40 Hz and there was no statistically significant effect at other frequencies. Amplitude- and frequency-response relationships for the depressant and the biphasic responses were analyzed separately. In the case of depressant responses, the magnitude was monotonically related to the amplitude of stimulation and depression occurred only at frequencies of 80 Hz or greater, with higher frequencies being more effective. The biphasic responses appeared to consist of 2 subtypes termed biphasic-1 and biphasic-2 responses. For biphasic-1 responses, the amplitude- and frequency-response curves were similar to those of depressant responses. Biphasic-2 responses differed in that the response was biphasic when the stimulation frequency was 80 Hz or greater and the amplitude was 0.3 mm or more, yet, at lower frequencies and/or amplitudes vibration evoked excitation. The similarities in the amplitude- and frequency-response relationships of depressant and biphasic-1 responses raise the possibility that these responses might be mediated by a single class of primary afferent. Both depressant and biphasic responses were evoked when stimulation parameters (2 microns, 240 Hz) were used which selectively activate Pacinian corpuscle afferents. Depression with 240-Hz stimulation was attenuated by administration of caffeine (60 mg/kg i.v.) suggesting that the depressant and biphasic-1 responses may be mediated by afferents from Pacinian corpuscles.

Action Potentials↗
PubMed1975

Quantitative evaluation of sensory nerve endings in hypertrophy of labia minora pudendi in women.

The authors studied the types of sensory nerve endings in hypertrophic labia minora pudendi in women aged between 18 and 69. The nervous tissue was impregnated with silver nitrate by Lavrentyev's modification of the Bielschowski-Gross method. The sensory nerve endings were divided into the following groups: free endings and arborizations, spray-like endings, seven types of clew-like nerve endings, and Pacinian corpuscles. It is the clew-like endings that absolutely predominate, they were 2,027 in number. The arborizations and spray-like endings found were 105 in number and only 4 of Pacinian corpuscles were found. As compared with normal labia there are no principal differences among the types of sensory nerve endings. The main difference consists in the rate of occurrence (89.6 as against 57.8) and in the thickness of the capsule, while the nerve clew proper does not grow in diameter. Therefore, we are of theopinion that a more frequent occurrence and larger size of the capsule is associated with local changes in the organ (especially with hypertrophy of the connective tissue). We observed no differences due to age either in the occurrence of the individual types or in the thickness of the capsule. Great individual variability in both directions was seen. The findings corroborated again the rare occurrence of meissner's endings in the labia minora in woman. Therefore it may be assumed that it is rather a random finding and that it is a type of clew-like nerve ending. Finally, we presume that the individually varying rate of occurrence of the nerve endings in the external genitals could also affect general sensitivity in this region, including sexual sensitivity.

Adolescent↗
PubMed2004

Absence of Meissner corpuscles in the digital pads of mice lacking functional TrkB.

The TrkB-expressing sensory neurons seem to be involved in touch and other discriminative sensibilities. Thus, several slowly and rapidly adapting cutaneous mechanoreceptors, as well as muscle spindles, are reduced or absent in the territory of the trigeminal nerve in functionally TrkB-deficient mice. Whether this also occurs in the cutaneous or muscular territories of dorsal root ganglia has not been analyzed. Here we used immunohistochemistry and transmission-electron microscopy to analyze the impact of a mutation in the gene coding for TrkB on Meissner and Pacinian corpuscles, and muscle spindles. The animals were studied at the post-natal days 15 and 25, because at this time all the mechanoreceptors examined are fully developed. Typical Meissner's corpuscles, displaying S-100 protein immunoreactivity, were found in the digital pads of wild-type and TrkB+/- mice whereas they were absent in the TrkB-/- animals. Regarding Pacinian corpuscles, the mutation in the trkB gene does not alter either the immunohistochemical or the ultrastructural characteristics. Finally, in muscle spindles the arrangement of the intrafusal muscle fibers and nerve fibers was unchanged in the mutated animals. Nevertheless, about 10% of muscle spindles showed increased number of the intrafusal cells (between 6 and 12) and were supplied by more than one large myelinic nerve fiber. The present results strongly suggest that TrkB-expressing sensory neurons in dorsal root ganglia, like those of the trigeminal ganglion, are responsible for the development and maintenance of several rapidly adapting cutaneous mechanoreceptors, i.e. Meissner's corpuscles.

Animals↗
PubMed1988

The structure and function of cutaneous sensory receptors.

The present review of cutaneous sensory receptors begins with a consideration of free nerve endings (FNEs) that can be considered as sensory terminals evidencing the least structural specialization of the axon and associated cells. Using the criteria established by Kruger et al (1981), FNEs of both A delta and C fibers can be identified on the basis of ultrastructural characteristics that include an intimate relationship between axons and the associated epithelium, the lack of a complete Schwann cell investment, the accumulation of numerous vesicles and other cytoplasmic organelles, and for A delta terminals a 1:1 relationship between axon and investing Schwann cell. Using these criteria, the so-called genital end bulbs of the human glans penis are merely a skein of FNEs based on the ultrastructural study of Halata and Munger (1986). Hair follicles of most species studied to date (the exception being the rabbit and to some extent the guinea pig) are multiply innervated with lanceolate, Ruffini and FNEs. The lanceolate terminals are the rapidly adapting terminals that are numerous in guard hairs. Ruffini terminals of hairs resemble those of the periodontal ligament or joint capsules and both are remarkably similar to Golgi tendon organs in terms of ultrastructural characteristics. The key ultrastructural characteristic is the encircling of collagen bundles by axons and associated Schwann and connective tissue cells. Axons frequently enter the epidermis either to terminate as FNEs or become associated with Merkel cells in glabrous skin at the base of the papillary ridges or in clusters of Merkel cells in hairy skin in touch domes or Haarscheiben. Merkel cells have clusters of apparent secretory granules polarized toward the axon and the axon is typically a slowly adapting mechanoreceptor. The function of the granules is not known. Pacinian corpuscles are the largest of the corpuscular receptors of the dermis and are characterized by an elaborate inner core of stacks of numerous thin lamellae arranged in a bilaterally symmetrical manner. Based on the fact that the lamellae are coupled with gap junctions and the outer core lamellae isolated by numerous tight junctions, the authors have proposed that the unique ionic environment may be in part responsible for the remarkable sensitivity of Pacinian corpuscles (Munger and Ide, 1987). Meissner corpuscles are a typical corpuscular receptor of murine (Ide, 1976, 1977), marsupial and primate glabrous skin (Munger, 1971). The axons typically weave back and forth between stacks of lamellae.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1975

Nervous system degeneration produced by acrylamide monomer.

Acrylamide, widely employed as a vinyl monomer in the polymer industry, is a potent neurotoxin to man and to animals. The cumulative effect of prolonged, low-level exposure to acrylamide monomer is the insidious development of a progressive peripheral neuropathy. Sensory symptoms begin in the hands and feet (numbness, pins and needles), certain reflexes are lost and, with severe exposure, muscle weakness and atrophy occur in the extremities. The peripheral neuropathy may be supplemented by symptoms indicative of central nervous system damage (ataxia, tremor, somnolence and mental changes). The neuropathologic basis for this clinical picture has been determined in cats. Here, chronic acrylamide intoxication produces selective peripheral and central nerve fiber degeneration. Degeneration first occurs in the extremities of long and large nerve fibers which later undergo a progressive, seriate proximal axonal degeneration known as dying-back. Especially vulnerable are sensory axons supplying Pacinian corpuscles and muscle spindles in the hindfoot toepads, while adjacent motor nerve axons die back later. Distal central nerve fiber degeneration is seen in the medulla and the cerebellum. The neurotoxic property of acrylamide is of practical concern in two areas. One major problem is the protection of factory workers engaged in the manufacture of acrylamide. A sensitive test of neurologic function in these individuals, i.e., touch sensation, based on the experimental observation of the exquisite vulnerability of Pacinian corpuscles in acrylamide intoxicated cats, is presently under consideration. The second area for concern is the exposure of the populace to minute amounts of neurotoxic acrylamide monomer which contaminate acrylamide polymers currently deployed in the environment. Federal restrictions on the maximum permitted exposure to acrylamide, based on a largely clinical study of acrylamide neurotoxicity conducted ten years ago, may require a re-evaluation in the light of recent advances which have pinpointed the initial sites of nerve fiber degeneration.

Acrylamides↗
PubMed1984

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.

Afferent Pathways↗
PubMed1978

[Degeneration of the laminated corpuscles (of Vater--Pacini) following colchicine application to a nerve].

Laminated pacinian corpuscles from the cat mesentery have been studied morphologically and morphometrically after nerve section and colchicine application to the nerve and the results obtained are represented. Similar interventions in the nerve produce changes in the receptors resembling those of wallerian type degeneration, degeneration rate after sectioning being higher than after colchicine application. At early stages after colchicine application the internal cone and its nuclei increase in size. The data obtained suggest the nuclei of the internal cone to be under neurotrophic control of the sensory neuron that might be realized via axoplasmic transport of substances.

Animals↗
PubMed2001

Response of muscle proprioceptors to spinal manipulative-like loads in the anesthetized cat.

OBJECTIVE: The mechanisms underlying the benefits of spinal manipulation are not well understood. Neurophysiological mechanisms likely mediate its effects, at least in part, yet we know little about how the nervous system is affected by spinal manipulation. The purpose of the present study was to determine whether muscle spindles and Golgi tendon organs in paraspinal muscles respond to a mechanical load whose force-time profile is similar to that of a spinal manipulation. METHODS: Experiments were performed on 10 anesthetized adult cats. The L6 dorsal root was isolated for electrophysiological recordings while the L6-L7 vertebrae and associated paraspinal tissues on one side of the vertebral column were left intact. Single unit recordings were obtained from 5 muscle spindles, 4 Golgi tendon organs, and 1 presumed Pacinian corpuscle afferent with receptive fields in paraspinal muscles. Loads were applied at the spinous process of the L6 vertebra through use of an electronic feedback control system. The load simulated the force-time profile of a spinal manipulation. Loads were applied in compressive and distractive directions and at 2 different angles (0 degrees and 45 degrees) with respect to the long axis of the vertebral column. RESULTS: Golgi tendon organ afferent discharge frequency increased more to the impulse than to the preload during 13 of 15 spinal manipulations. Generally, the 4 Golgi tendon organ afferents became silent immediately at the end of each impulse. Similarly, muscle spindle discharge frequency increased more to the impulse than to the preload during 10 of 16 manipulations. Distractive manipulations loaded the spindles more effectively than compressive manipulations. After 7 of these 10 manipulations, muscle spindles became silent for 1.3 +/- 0.6 seconds (range, 0.1-4.3 seconds). Six of the 16 manipulations unloaded the muscle spindles. A presumed Pacinian corpuscle responded to the impulse of a manipulative-like load but not to loads with a slower force-time profile. CONCLUSION: The data suggest that the high-velocity, short-duration load delivered during the impulse of a spinal manipulation can stimulate muscle spindles and Golgi tendon organs more than the preload. The physiologically relevant portion of the manipulation may relate to its ability to increase as well as decrease the discharge of muscle proprioceptors. In addition, the preload, even in the absence of the impulse, can change the discharge of paraspinal muscle spindles. Loading of the vertebral column during a sham manipulation may affect the discharge of paraspinal proprioceptors.

Anesthesia, Inhalation↗
PubMed1989

Simulation of motion on the skin. II. Cutaneous mechanoreceptor coding of the width and texture of bar patterns displaced across the OPTACON.

1. These experiments assay the functional significance of receptive-field architecture for information processing. Rapidly adapting (RA) afferents have been previously shown to converge information from clusters of 14-25 Meissner's corpuscles, whereas afferents innervating Pacinian corpuscles (PCs) have only a single, large receptor terminal. We tested two opposing hypotheses of functional architecture: 1) summation models, in which an afferent integrates signals from all of its terminals, showing monotonic increases in activity as a function of contact area, and 2) winner-take-all models, in which the most strongly activated receptor in the cluster dominates axonal output by cancellation of signals from other branches. 2. Bar and stripe patterns have been swept across the finger or palm of the monkey's hand at speeds of 30-120 mm/s with the use of a computer-controlled grid of sequentially activated miniature probes (OPTACON stimulator). The dense packing of OPTACON probes permits placement of up to five groups of stimulators within an individual receptive field, allowing us to activate one or more clusters of Meissner's corpuscles simultaneously and to stimulate the bulbar corpuscle of PC afferents at different orientations through the skin. Integration of information from moving bar patterns has been tested with two protocols. In the variable width protocol, the total number of activated rows in the pattern is varied from one to five, with a constant spacing of 1.2 mm between pulsed rows. In the variable density protocol, the length of skin stimulated is held constant at 5 mm and the spacing of stimuli varied. 3. RA afferents show no evidence of summation of inputs within their receptive fields. Motion of wide bars across the field increases the duration of firing but not the total spikes evoked by each pulse. Responses to the leading edge of wide bars were found to be identical to those evoked by a single-row bar. Simultaneous activation of two to five rows evokes the same or fewer spikes per pulse than the most effective individual row tested alone. When broad-bar patterns are centered over the field, contacting the maximum number of receptors, RAs follow activity in the dominant branch or terminus, suppressing additional inputs. Lack of summation is observed at all pulse frequencies tested (25-100 Hz). 4. Moving bar patterns evoke responses as long as at least one row stimulates the receptive field; broader patterns evoke longer spike trains whose total number of impulses is proportional to bar width.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗
PubMed1996

Study of human cutaneous sensory corpuscles using double immunolabelling and confocal laser scanning microscopy.

BACKGROUND: The main constituents of sensory corpuscles, i.e., the central axon, the periaxonic Schwann-related, cells, and the perineurial-related cells, can be identified light microscopically by simple immunohistochemistry using specific antibodies. This paper demonstrates the usefulness of double immunolabelling for light and confocal laser-scanning microscopy (CLSM) in the study of human cutaneous sensory corpuscles. MATERIALS AND METHODS: Antibodies directed against neurofilament proteins (NFPs) and S-100 protein were used to label the central axon and the lamellar cells of Meissner corpuscles or the inner-core lamellae of digital cutaneous Pacinian corpuscles, respectively. Samples were obtained from subjects with normal sensitivity and from patients with paresthesia or absence of clinical sensitivity. Single and double immunolabelling was performed, and the sections were studied by light or CLSM microscopy. RESULTS: Double immunolabelling was effective for simultaneous observation of the central axon (NFP-positive) and periaxonic Schwann-related (S-100 protein-positive) cells in sensory corpuscles from normal human digital skin. The images that were obtained with both methods were comparable, but the axonic profiles were sharper with diaminobenzidine (DAB) used as a chromogen rather than with Texas-red used as a fluorochrome. Nevertheless, the ability to manipulate the focal plane by using CLSM permits one to better analyze the intracorpuscular relationships of the axon. Double immunolabelling in sensory corpuscles from the skin of patients with nerve compression showed the presence of a central axon in the corpuscles, whereas it was absent in the sensory corpuscles of clinically denervated skin. CONCLUSIONS: Double immunolabelling is a useful method with which to analyze simultaneously two of the corpuscular constituents, and it may be used in the study of denervated and reinnervated sensory corpuscles.

Biopsy↗
PubMed1998

S100alpha and S100beta proteins in human cutaneous sensory corpuscles: effects of nerve and spinal cord injury.

S100 protein in the vertebrate peripheral nervous system consists of homo- or heterodimers of S100alpha and S100beta proteins, the first predominating in neurons and the second in glial cells. Recently, however, occurrence of S100beta protein in neurons has been reported. The expression of S100 protein by Schwann cells, as well as their derivatives in sensory corpuscles, depends on the sensory axon (i.e., the Schwann cell-axon contact). The present study analyzed the distribution of S100alpha and S100beta proteins in human cutaneous sensory corpuscles and the effects of peripheral or central sensory axon severance in the expression of these proteins. Simple or double immunohistochemistry was carried out using a panel of antibodies against S100alpha, S100beta or S100alpha+beta proteins, and the sections were examined by light or laser confocal scanning microscopy. Skin samples were obtained from normal subjects and patients with spinal cord injury, nerve entrapment, and nerve sections plus graft. The lamellar cells of Meissner corpuscles as well as the inner-core lamellae of the Pacinian corpuscles displayed strong immunoreactivity (IR) for all antigens examined, the most intense labeling being obtained for S100beta protein. The pattern of immunostaining was unchanged after spinal cord injury, whereas the number of stained corpuscles as well as the intensity of IR for each antigen decreased in cutaneous sensory corpuscles after nerve injury, both entrapment and section plus graft. No evidence was found of axonal labeling. The present results provide evidence that Schwann-related cells in human cutaneous sensory corpuscles contain both S100alpha and S100beta and that the expression of these proteins is dependent on the functional and structural integrity of sensory fibers.

Adult↗
PubMed2006

Pacinian neuroma in adipose herniation of the buccal mucosa.

An interesting case of a trauma-induced tender mass of the buccal mucosa in a 45-year-old man was presented. Following surgery, the patient was relieved from pain. Microscopically, the mature adipose tissue is unique in that it contained a single enlarged Pacinian corpuscle near the deep margin. This is the hitherto undescribed intraoral lesion of Pacinian neuroma in the herniated buccal fat pad.

Adipose Tissue↗
PubMed1994

Expression of epidermal growth factor receptor (EGFr) immunoreactivity in human cutaneous nerves and sensory corpuscles.

BACKGROUND: The epidermal growth factor receptor (EGFr) binds both epidermal growth factor (EGF) and transforming growth factor alpha (TGF alpha), which are currently considered among putative growth factors playing a role in the nervous system. EGFr and their ligands have been localized in the mammalian peripheral nervous system. The present study was undertaken to investigate whether nerves and sensory corpuscles supplying human glabrous skin express EGFr. METHODS: Formaldehyde fixed, paraffin embedded samples of finger-tip digital skin obtained from adult healthy subjects were processed for indirect PAP immunohistochemistry using a monoclonal antibody against an epitope of the intracellular domain of EGFr. To ascertain the localization of EGFr immunoreactivity, neurofilament proteins (NFP), S100 protein (S100P), and epithelial membrane antigen (EMA) were studied in parallel to label axons, Schwann cells, and perineurial cells, respectively, as well as their corpuscular derivatives. RESULTS: A variable intensity of EGFr immunostaining was regularly observed in the perineurium and Schwann cells, and occasionally in the axons of nerve bundles. EGFr immunoreactivity was also present in the axon and lamellar cells of Meissner corpuscles, and within the axon, inner-core, outer-core, and capsule of Pacinian corpuscles. CONCLUSIONS: Present results demonstrate that human cutaneous nerves and sensory corpuscles express EGFr suggesting a role for peptides able to bind EGFr, i.e., EGF and TGF alpha, in the human peripheral nervous sensory system.

Adult↗
PubMed1996

[Regeneration of free nerve endings and sensory corpuscles after implantation of nerve into the denervated skin in the monkey].

Two rhesus monkeys were used. Their fingers were denervated first and then the nerve was implanted into the denervated skin. The results showed that: (1) the process of regeneration of nerve fibers was similar to that of previous experiments in rabbits; (2) regeneration of nervous tissue included 2 kinds of sensory receptors with capsule-Meissner corpuscles and Merkel cell-dendrite complex; (3) the regenerative rate of Meissner corpuscles increased progressively and their morphology and structure became matured with elapse of time; (4) no regeneration of pacinian corpuscle was found in 6 months after nerve implantation; (5) Meissner corpuscles also regenerated in the transplanted glabrous skin.

Animals↗
PubMed1987

Carbonic anhydrase activity in axon terminals of sensory corpuscles.

The distribution of carbonic anhydrase (CA) activity was studied by electron microscopic histochemistry in rat Pacinian corpuscles, Meissner corpuscles and Merkel cell-neurite complexes using the cobalt bicarbonate method. The distribution of CA activity in these axon terminals was compared to the activity in sciatic nerve axons. An intense enzymatic CA activity was demonstrated in axon terminals of both Pacinian and Meissner corpuscles, while a weak activity was found within the axoplasm of terminals abutting Merkel cells. Some large- and medium-sized axons in sciatic nerves exhibited an intense activity. These findings indicate that large- or medium-diameter sensory axons innervating corpuscular endings have an intense CA activity extending from their somata to their sensory terminals. Axons to Merkel-neurite complexes differ in CA activity from those innervating Meissner and Pacinian corpuscular endings.

Animals↗
PubMed1976

Pacinian neurofibroma.

A 40-year-old woman developed a tender, greyish-white nodule of the index finger. Histologic examination disclosed mature and immature pacinian corpuscles.

Adult↗
PubMed1979

[Histology of a plexiform neuroma with Vater-Pacini-lamellar-corpuscle-like structures].

From the finger of a 11 year old boy a tumor that had been present since birth was removed. Histological examination showed a great number of immature nerve bundles, and structures resembling Pacinian corpuscles. It could be shown that the organoid structures are a result of a proliferation of the so-called perineurial cells. These formations do not correspond with real tactile corpuscles, rather they can be considered as neoplastic structures of the perineurium.

Child↗
PubMed2002

Distribution and terminal arborizations of cutaneous mechanoreceptors in the glabrous finger pads of the monkey.

Recent electrophysiological studies demonstrated that neurons in the somatosensory cortex of monkeys respond to tangential forces applied to glabrous skin. To unravel the peripheral basis for this cortical response, we determined the distribution of presumptive low-threshold mechanoreceptors innervating the distal finger pads of monkeys. Endings were reconstructed in immunolabeled serial sections imaged by epifluorescence and confocal microscopy. Although classically implicated as cutaneous stretch receptors, no Ruffini corpuscles were found in the glabrous skin. Ruffini-like endings were only detected at the base of the finger nails. Pacinian corpuscles were sparsely distributed in the deep dermis. Meissner corpuscles (MCs) in dermal papillary ridges had a comparably high density in the thumb, index, and fifth fingers. Each MC was innervated by several large-caliber axons. Within the limits of our reconstructions, some of these axons terminated in only one MC, whereas others innervated several MCs. Merkel endings covered about 80% of the base of the intermediate epidermal ridges that form the pattern of fingerprints. In some cases, the distal tip of a Merkel-related axon gave rise to a several terminal branches that supplied endings to tightly circumscribed (30-70 microm) clusters of Merkel cells. In other cases, the nodes of axons gave rise to en passant branches that formed extended chains of endings among Merkel cells spread over territories up to 300 microm long. Based on their relatively diffuse distributions, the axons that innervate multiple MCs or the axons with en passant Merkel terminations seem most suited to transduce tangential forces.

Animals↗