Merkel nerve ending

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Merkel nerve endings are mechanoreceptors, a type of sensory receptor, that are found in the basal epidermis and hair follicles. They are nerve endings and provide information on mechanical pressure, position, and deep static touch features, such as shapes and edges.

Contents

Merkel cells in the basal epidermis of the skin store serotonin which they release to associated nerve endings in response to pressure. Each ending consists of a Merkel cell in close apposition with an enlarged nerve terminal. This is sometimes referred to as a Merkel cell–neurite complex, or a Merkel disc receptor. A single afferent nerve fibre branches to innervate up to 90 such endings.

Location

In mammals, Merkel nerve endings have a wide distribution and are found in the basal layer of glabrous and hairy skin, in hair follicles, and in oral and anal mucosa. Microscopically they are relatively large, myelinated nerve endings.

In humans, Merkel cells along with Meissner's corpuscles occur in the superficial skin layers, and are most densely clustered beneath the ridges of the highly sensitive fingertips which make up fingerprints, and less so in the palms and forearm. In hairy skin, Merkel nerve endings are clustered into specialized epithelial structures called "touch domes" or "hair disks". Merkel receptors are also located in the mammary glands. Wherever they are found, the epithelium is arranged to optimize the transfer of pressure to the ending.

Functions

Merkel cells provide information on pressure, position, and deep static touch features such as shapes and edges. They are tactile sensors in the business of mechanotransduction. They encode surface features of touched objects into perception, but also have to do with proprioception. [1]

Merkel cells transduce tactile stimuli / mechanical forces into excitatory signals, which trigger vesicular serotonin release; they have also been called a "serotonergic synapse". [2] They have similar functions as the enterochromaffin cell, the mechanosensory cell in the GI epithelium, [2] which synthesizes 95% of the body's total serotonin or 5-HT. Like the cells responsible for the mechanotransduction in hearing, Merkel cells transduce mechanical forces into excitatory signals via ion conductance on mechanosensitive channels. [3] of which Piezo2 is the Merkel cell's primary mechanosensor. [4]

Electrophysiology

The Merkel cell's somewhat rigid structure, and the fact that they are not encapsulated,[ clarification needed ] causes them to have a sustained response in the form of action potentials (or spikes) to mechanical deflection of the tissue. Because of their sustained response to pressure, Merkel nerve endings are classified as slowly adapting in contrast to rapidly adapting receptors by Pacinian and Meissner's corpuscles, which respond only to the onset and offset of mechanical deflection. In mammals, electrical recordings from single afferent nerve fibres have shown that the responses of Merkel nerve endings are characterized by a vigorous response to the onset of a mechanical ramp stimulus (dynamic), and then continued firing during the plateau phase (static). Firing during the static phase can continue for more than 30 minutes. The inter-spike intervals during sustained firing are irregular, in contrast to the highly regular pattern of inter-spike intervals obtained from slowly adapting type II mechanoreceptors.

They fire fastest, when small points indent the skin, and fire at a low rate on slow curves or flat surfaces. Convexities reduce their rate of firing further still. [5]

Sensitivity and receptive fields

Merkel nerve endings are the most sensitive of the four main types of mechanoreceptors to vibrations at low frequencies, around 5 to 15 Hz. Merkel nerve endings are extremely sensitive to tissue displacement, and may respond to displacements of less than 1 μm. A mechanoreceptor's receptive field is the area within which a stimulus can excite the cell. If the skin is touched in two separate points within a single receptive field, the person will be unable to feel the two separate points. If the two points touched span more than a single receptive field then both will be felt. The size of mechanoreceptors' receptive fields in a given area determines the degree to which detailed stimuli can be resolved: the smaller and more densely clustered the receptive fields, the higher the resolution.

Type I afferent fibres have smaller receptive fields than type II fibres. Several studies indicate that type I fibres mediate high resolution tactile discrimination, and are responsible for the ability of our finger tips to feel fine detailed surface patterns (e.g. for reading Braille). Merkel's discs have small receptive fields which allow for them to detect fine spatial separation. They also have two point discrimination.

Eponym

Merkel's discs are named after German anatomist Friedrich Merkel (1845–1919), who was 30 years old when he described them. [6]

Diseases

In burns, Merkel endings are most commonly lost.

People who have diabetes, inflammatory diseases, or undergo chemotherapy can lose tactile sensitivity and develop tactile allodynia. Recreational drugs acting on serotoninergic synapses can cause exaggerated tactile sensations. [2]

Related Research Articles

<span class="mw-page-title-main">Sensory nervous system</span> Part of the nervous system responsible for processing sensory information

The sensory nervous system is a part of the nervous system responsible for processing sensory information. A sensory system consists of sensory neurons, neural pathways, and parts of the brain involved in sensory perception and interoception. Commonly recognized sensory systems are those for vision, hearing, touch, taste, smell, balance and visceral sensation. Sense organs are transducers that convert data from the outer physical world to the realm of the mind where people interpret the information, creating their perception of the world around them.

Stimulus modality, also called sensory modality, is one aspect of a stimulus or what is perceived after a stimulus. For example, the temperature modality is registered after heat or cold stimulate a receptor. Some sensory modalities include: light, sound, temperature, taste, pressure, and smell. The type and location of the sensory receptor activated by the stimulus plays the primary role in coding the sensation. All sensory modalities work together to heighten stimuli sensation when necessary.

A cutaneous receptor is the type of sensory receptor found in the skin. They are a part of the somatosensory system. Cutaneous receptors include mechanoreceptors, nociceptors (pain), and thermoreceptors (temperature).

A mechanoreceptor, also called mechanoceptor, is a sensory receptor that responds to mechanical pressure or distortion. Mechanoreceptors are innervated by sensory neurons that convert mechanical pressure into electrical signals that, in animals, are sent to the central nervous system.

<span class="mw-page-title-main">Dorsal column–medial lemniscus pathway</span> Sensory spinal pathway

The dorsal column–medial lemniscus pathway (DCML) is a sensory pathway of the central nervous system that conveys sensations of fine touch, vibration, two-point discrimination, and proprioception from the skin and joints. It transmits information from the body to the primary somatosensory cortex in the postcentral gyrus of the parietal lobe of the brain. The pathway receives information from sensory receptors throughout the body, and carries this in nerve tracts in the white matter of the dorsal column of the spinal cord to the medulla, where it is continued in the medial lemniscus, on to the thalamus and relayed from there through the internal capsule and transmitted to the somatosensory cortex. The name dorsal-column medial lemniscus comes from the two structures that carry the sensory information: the dorsal columns of the spinal cord, and the medial lemniscus in the brainstem.

<span class="mw-page-title-main">Tactile corpuscle</span> Type of mechanoreceptor that detects light touch

Tactile corpuscles or Meissner's corpuscles are a type of mechanoreceptor discovered by anatomist Georg Meissner (1829–1905) and Rudolf Wagner. This corpuscle is a type of nerve ending in the skin that is responsible for sensitivity to pressure. In particular, they have their highest sensitivity when sensing vibrations between 10 and 50 hertz. They are rapidly adaptive receptors. They are most concentrated in thick hairless skin, especially at the finger pads.

<span class="mw-page-title-main">Sensory neuron</span> Nerve cell that converts environmental stimuli into corresponding internal stimuli

Sensory neurons, also known as afferent neurons, are neurons in the nervous system, that convert a specific type of stimulus, via their receptors, into action potentials or graded receptor potentials. This process is called sensory transduction. The cell bodies of the sensory neurons are located in the dorsal ganglia of the spinal cord.

<span class="mw-page-title-main">Dorsal root ganglion</span> Cluster of neurons in a dorsal root of a spinal nerve

A dorsal root ganglion is a cluster of neurons in a dorsal root of a spinal nerve. The cell bodies of sensory neurons known as first-order neurons are located in the dorsal root ganglia.

<span class="mw-page-title-main">Pacinian corpuscle</span> Type of mechanoreceptor cell in hairless mammals

The Pacinian corpuscle, lamellar corpuscle or Vater-Pacini corpuscle is one of the four major types of mechanoreceptors found in mammalian skin. This type of mechanoreceptor is found in both hairy, and hairless skin, viscera, joints, and attached to periosteum of bone, primarily responsible for sensitivity to vibration. Few of them are also sensitive to quasi-static or low frequency pressure stimulus. Most of them respond only to sudden disturbances and are especially sensitive to vibration of few hundreds of Hz. The vibrational role may be used for detecting surface texture, e.g., rough vs. smooth. Most of the Pacinian corpuscles act as rapidly adapting mechanoreceptors. Groups of corpuscles respond to pressure changes, e.g. on grasping or releasing an object.

<span class="mw-page-title-main">Mechanotransduction</span> Conversion of mechanical stimulus of a cell into electrochemical activity

In cellular biology, mechanotransduction is any of various mechanisms by which cells convert mechanical stimulus into electrochemical activity. This form of sensory transduction is responsible for a number of senses and physiological processes in the body, including proprioception, touch, balance, and hearing. The basic mechanism of mechanotransduction involves converting mechanical signals into electrical or chemical signals.

<span class="mw-page-title-main">Merkel cell</span> Receptors in the skin of vertebrates

Merkel cells, also known as Merkel-Ranvier cells or tactile epithelial cells, are oval-shaped mechanoreceptors essential for light touch sensation and found in the skin of vertebrates. They are abundant in highly sensitive skin like that of the fingertips in humans, and make synaptic contacts with somatosensory afferent nerve fibers. It has been reported that Merkel cells are derived from neural crest cells, though more recent experiments in mammals have indicated that they are epithelial in origin.

<span class="mw-page-title-main">Tactile corpuscles of Grandry</span>

The tactile corpuscles of Grandry or Grandry corpuscles are mechanoreceptors found in the beak skin and oral mucosa of aquatic birds. They were first described by Grandry in 1869 in the bill skin of ducks and geese. Their general structure includes the flattened endings of an afferent nerve fiber sandwiched between two or more somewhat flattened sensory cells called Grandry cells, all surrounded by a layer of satellite cells and a partial capsule of collagen protein. Electrophysiological studies have shown that Grandry corpuscles function as rapidly adapting velocity detectors. In birds, Grandry and Merkel corpuscles share many morphological similarities, which has led to some confusion in the literature over their classification.

<span class="mw-page-title-main">Microneurography</span>

Microneurography is a neurophysiological method employed to visualize and record the traffic of nerve impulses that are conducted in peripheral nerves of waking human subjects. It can also be used in animal recordings. The method has been successfully employed to reveal functional properties of a number of neural systems, e.g. sensory systems related to touch, pain, and muscle sense as well as sympathetic activity controlling the constriction state of blood vessels. To study nerve impulses of an identified nerve, a fine tungsten needle microelectrode is inserted into the nerve and connected to a high input impedance differential amplifier. The exact position of the electrode tip within the nerve is then adjusted in minute steps until the electrode discriminates nerve impulses of interest. A unique feature and a significant strength of the microneurography method is that subjects are fully awake and able to cooperate in tests requiring mental attention, while impulses in a representative nerve fibre or set of nerve fibres are recorded, e.g. when cutaneous sense organs are stimulated or subjects perform voluntary precision movements.

Cutaneous innervation refers to an area of the skin which is supplied by a specific cutaneous nerve.

Eimer's organs are sensory organs in which the epidermis is modified to form bulbous papillae. First isolated by Theodor Eimer from the European mole in 1871, these organs are present in many moles, and are particularly common in the star-nosed mole, which bears 25,000 of them on its unique tentacled snout. The organs are formed from a stack of epidermal cells, which is innervated by nerve processes from myelinated fibers in the dermis, which form terminal swellings just below the outer keratinized layer of epidermis. They contain a Merkel cell-neurite complex in the epidermis and a lamellated corpuscle in the dermal connective tissue.

Pallesthesia, or vibratory sensation, is the ability to perceive vibration. This sensation, often conducted through skin and bone, is usually generated by mechanoreceptors such as Pacinian corpuscles, Merkel disk receptors, and tactile corpuscles. All of these receptors stimulate an action potential in afferent nerves found in various layers of the skin and body. The afferent neuron travels to the spinal column and then to the brain where the information is processed. Damage to the peripheral nervous system or central nervous system can result in a decline or loss of pallesthesia.

Mechanosensation is the transduction of mechanical stimuli into neural signals. Mechanosensation provides the basis for the senses of light touch, hearing, proprioception, and pain. Mechanoreceptors found in the skin, called cutaneous mechanoreceptors, are responsible for the sense of touch. Tiny cells in the inner ear, called hair cells, are responsible for hearing and balance. States of neuropathic pain, such as hyperalgesia and allodynia, are also directly related to mechanosensation. A wide array of elements are involved in the process of mechanosensation, many of which are still not fully understood.

<span class="mw-page-title-main">Somatosensory system</span> Nerve system for sensing touch, temperature, body position, and pain

In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch, as well as temperature (thermoception), body position (proprioception), and pain. It is a subset of the sensory nervous system, which also represents visual, auditory, olfactory, and gustatory stimuli.

Tactile induced analgesia is the phenomenon where concurrent touch and pain on the skin reduces the intensity of pain that is felt.

C tactile afferents are nerve receptors in mammalian skin that generally respond to nonpainful stimulation such as light touch. For this reason they are classified as ‘low-threshold mechanoreceptors’. As group C nerve fibers, they are unmyelinated and have slow conduction velocities. They are mostly associated with the sensation of pleasant touch, though they may also mediate some forms of pain. CT afferents were discovered by Åke Vallbo using the technique of microneurography.

References

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  2. 1 2 3 Chang W, Kanda H, Ikeda R, Ling J, DeBerry JJ, Gu JG. Merkel disc is a serotonergic synapse in the epidermis for transmitting tactile signals in mammals. Proc Natl Acad Sci U S A. 2016 Sep 13;113(37): E5491-500. doi: 10.1073/pnas.1610176113.
  3. Arnadóttir J., Chalfie M. Eukaryotic mechanosensitive channels. Annu. Rev. Biophys. 39, 111–137. (2010) DOI 10.1146/annurev.biophys.37.032807.125836
  4. Woo S. H., Ranade S., Weyer A. D., Dubin A. E., Baba Y., Qiu Z., et al. Piezo2 is required for Merkel-cell mechanotransduction. Nature (2014)509, 622–626. DOI 10.1038/nature13251
  5. Kandel E.R., Schwartz, J.H., Jessell, T.M. (2000). Principles of Neural Science , 4th ed., p.433. McGraw-Hill, New York.
  6. Merkel FS. (1875). Tastzellen und Tastkörperchen bei den Hausthieren und beim Menschen. Archiv für mikroskopische Anatomie, 11: 636-652.