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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, a ...
s, a type of
sensory receptor 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 potentials. This process is called sensory transduction. The ...
, 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 rel ...
and
hair follicle The hair follicle is an organ found in mammalian skin. It resides in the dermal layer of the skin and is made up of 20 different cell types, each with distinct functions. The hair follicle regulates hair growth via a complex interaction between ...
s. 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 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 ...
. 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) fro ...
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 The ''stratum basale'' (basal layer, sometimes referred to as ''stratum germinativum'') is the deepest layer of the five layers of the epidermis, the external covering of skin in mammals. The ''stratum basale'' is a single layer of columnar or ...
of
glabrous Glabrousness (from the Latin '' glaber'' meaning "bald", "hairless", "shaved", "smooth") is the technical term for a lack of hair, down, setae, trichomes or other such covering. A glabrous surface may be a natural characteristic of all or part of ...
and hairy skin, in
hair follicle The hair follicle is an organ found in mammalian skin. It resides in the dermal layer of the skin and is made up of 20 different cell types, each with distinct functions. The hair follicle regulates hair growth via a complex interaction between ...
s, 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 corpuscle 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 pressur ...
s 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 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 intercellul ...
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 sensor A tactile sensor is a device that measures information arising from physical interaction with its environment. Tactile sensors are generally modeled after the biological sense of cutaneous touch which is capable of detecting stimuli resultin ...
s 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 neurons ...
. 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 Enterochromaffin (EC) cells (also known as Kulchitsky cells) are a type of enteroendocrine cell, and neuroendocrine cell. They reside alongside the epithelium lining the lumen of the digestive tract and play a crucial role in gastrointestinal reg ...
, 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. of which
Piezo2 Piezo-type mechanosensitive ion channel component 2 is a protein that in humans is encoded by the PIEZO2 gene. It has a homotrimeric structure, with three blades curving into a nano-dome, with a diameter of 28 nanometers. Function Piezos are ...
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 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, a ...
in contrast to rapidly adapting receptors by
Pacinian Pacinian corpuscle or lamellar corpuscle or Vater-Pacini corpuscle; is one of the four major types of mechanoreceptors (specialized nerve ending with adventitious tissue for mechanical sensation) found in mammalian skin. This type of mechanorece ...
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, su ...
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 The receptive field, or sensory space, is a delimited medium where some physiological stimuli can evoke a sensory neuronal response in specific organisms. Complexity of the receptive field ranges from the unidimensional chemical structure of o ...
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