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Merkel nerve endings are
mechanoreceptor 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, ...
s, a type of sensory receptor, that are found in the basal
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water relea ...
and hair follicles. They are nerve endings and provide information on mechanical pressure, position, and deep static touch features, such as shapes and edges. Merkel cells in the basal epidermis of the skin store
serotonin Serotonin () or 5-hydroxytryptamine (5-HT) is a monoamine neurotransmitter. Its biological function is complex and multifaceted, modulating mood, cognition, reward, learning, memory, and numerous physiological processes such as vomiting and va ...
which they release to associated nerve endings in response to pressure. Each ending consists of a
Merkel cell 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 f ...
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 A sensory nerve, or afferent nerve, is a general anatomic term for a nerve which contains predominantly somatic afferent nerve fibers. Afferent nerve fibers in a sensory nerve carry sensory information toward the central nervous system (CNS) from ...
fibre branches to
innervate A nerve is an enclosed, cable-like bundle of nerve fibers (called axons) in the peripheral nervous system. A nerve transmits electrical impulses. It is the basic unit of the peripheral nervous system. A nerve provides a common pathway for the ...
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 cell 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 f ...
s 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 A mammary gland is an exocrine gland in humans and other mammals that produces milk to feed young offspring. Mammals get their name from the Latin word ''mamma'', "breast". The mammary glands are arranged in organs such as the breasts in prima ...
. Wherever they are found, the
epithelium Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercel ...
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 Proprioception ( ), also referred to as kinaesthesia (or kinesthesia), is the sense of self-movement, force, and body position. It is sometimes described as the "sixth sense". Proprioception is mediated by proprioceptors, mechanosensory neuron ...
. Merkel disks can also respond to light touch. Merkel cells transduce tactile stimuli / mechanical forces into excitatory signals, which trigger vesicular
serotonin Serotonin () or 5-hydroxytryptamine (5-HT) is a monoamine neurotransmitter. Its biological function is complex and multifaceted, modulating mood, cognition, reward, learning, memory, and numerous physiological processes such as vomiting and va ...
release; they have also been called a "serotonergic synapse".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. They have similar functions as the enterochromaffin cell, the mechanosensory cell in the GI epithelium, 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 Mechanosensitive channels, mechanosensitive ion channels or stretch-gated ion channels (not to be confused with mechanoreceptors). They are present in the membranes of organisms from the three domains of life: bacteria, archaea, and eukarya. They a ...
. of which Piezo2 is the Merkel cell's primary mechanosensor.


Electrophysiology

The Merkel cell's somewhat rigid structure, and the fact that they are not encapsulated, causes them to have a sustained response in the form of
action potential An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...
s 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.


Sensitivity and receptive fields

Merkel nerve endings are the most sensitive of the four main types of mechanoreceptors to
vibration Vibration is a mechanical phenomenon whereby oscillations occur about an equilibrium point. The word comes from Latin ''vibrationem'' ("shaking, brandishing"). The oscillations may be periodic, such as the motion of a pendulum—or random, suc ...
s 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 Braille (Pronounced: ) is a tactile writing system used by people who are visually impaired, including people who are blind, deafblind or who have low vision. It can be read either on embossed paper or by using refreshable braille displ ...
). 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.Merkel FS. (1875). Tastzellen und Tastkörperchen bei den Hausthieren und beim Menschen. Archiv für mikroskopische Anatomie, 11: 636-652.


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 Allodynia is a condition in which pain is caused by a stimulus that does not normally elicit pain. For example, bad sunburn can cause temporary allodynia, and touching sunburned skin, or running cold or warm water over it, can be very painful. It i ...
. Recreational drugs acting on serotoninergic synapses can cause exaggerated tactile sensations.


References


External links

* * * {{somatosensory system Sensory receptors Sensory systems