Merkel cell | |
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![]() Merkel cells (shown in blue) are located in the basal epidermal layer of the skin. | |
Details | |
Location | Skin of vertebrates |
Function | Light touch sensation |
Identifiers | |
MeSH | D018862 |
NeuroLex ID | nifext_87 |
Anatomical terms of neuroanatomy |
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.
Merkel cells functionally resemble the enterochromaffin cell, the mechanosensory cell of the gastrointestinal epithelium. [1]
Merkel cells are found in the skin and some parts of the mucosa of all vertebrates. In mammalian skin, they are clear cells found in the stratum basale [2] [3] (at the bottom of sweat duct ridges) of the epidermis approximately 10 μm in diameter. They 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. [4] They also occur in epidermal invaginations of the plantar foot surface called rete ridges. [5]
Most often, they are associated with sensory nerve endings, when they are known as Merkel nerve endings (also called a Merkel cell-neurite complex). They are associated with slowly adapting (SA1) somatosensory nerve fibers. They react to low vibrations (5–15 Hz) and deep static touch such as shapes and edges. Due to a small receptive field (extremely detailed info) they are densely present in areas like fingertips; they are not covered (shelled) and thus respond to pressures over long periods.
The origin of Merkel cells has been debated for over 20 years. Evidence from skin graft experiments in birds implies that they are neural crest derived, [6] but experiments in mammals now demonstrate an epidermal origin. [7] [8]
The German anatomist Friedrich Sigmund Merkel referred to Merkel cells as Tastzellen or "touch cells". Until recently this proposed function was controversial and hard to prove, due to the close physical association of Merkel cells with sensory nerve endings. However, recent work in mice and other model organisms demonstrates that Merkel cells intrinsically transform touch into electrical signals that are transmitted to the nervous system. [9] Merkel cells express PIEZO2, a mechanosensitive ion channel that responds to mechanical forces. Mice in which Piezo2 is knocked-out specifically in skin cells, but not sensory neurons, show decreased behavioral responses to gentle touch. [10]
Merkel cells are sometimes considered APUD cells (an older definition, more commonly classified as a part of dispersed neuroendocrine system) because they contain dense core granules, and thus may also have a neuroendocrine function. [11]
Although uncommon, these cells may become malignant and form a Merkel cell carcinoma—an aggressive and difficult to treat skin cancer. [12] [13]
In physiology, nociception, also nocioception; from Latin nocere 'to harm/hurt') is the sensory nervous system's process of encoding noxious stimuli. It deals with a series of events and processes required for an organism to receive a painful stimulus, convert it to a molecular signal, and recognize and characterize the signal to trigger an appropriate defensive response.
In neuroanatomy, the trigeminal nerve (lit. triplet nerve), also known as the fifth cranial nerve, cranial nerve V, or simply CN V, is a cranial nerve responsible for sensation in the face and motor functions such as biting and chewing; it is the most complex of the cranial nerves. Its name (trigeminal, from Latin tri- 'three' and -geminus 'twin') derives from each of the two nerves (one on each side of the pons) having three major branches: the ophthalmic nerve (V1), the maxillary nerve (V2), and the mandibular nerve (V3). The ophthalmic and maxillary nerves are purely sensory, whereas the mandibular nerve supplies motor as well as sensory (or "cutaneous") functions. Adding to the complexity of this nerve is that autonomic nerve fibers as well as special sensory fibers (taste) are contained within it.
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 released from the body into the atmosphere through transepidermal water loss.
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 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.
A nociceptor is a sensory neuron that responds to damaging or potentially damaging stimuli by sending "possible threat" signals to the spinal cord and the brain. The brain creates the sensation of pain to direct attention to the body part, so the threat can be mitigated; this process is called nociception.
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.
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 root ganglia of the spinal cord.
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.
Merkel nerve endings are mechanoreceptors situated in the basal epidermis as well as around the apical ends or some hair follicles. They are slowly adapting They have small receptive fields measuring some milimeters in diameter. Most are associated with fast-conducting large myelinated axons. A single afferent nerve fibre branches to innervate up to 90 such endings. Merkel nerve endings respond to light touch. They respond to sustained pressure, and are sensitive to edges of objects. Their exact functions remain controversial.
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.
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.
Group Aβ of the type II sensory fiber is a type of sensory fiber, the second of the two main groups of touch receptors. The responses of different type Aβ fibers to these stimuli can be subdivided based on their adaptation properties, traditionally into rapidly adapting (RA) or slowly adapting (SA) neurons. Type II sensory fibers are slowly-adapting (SA), meaning that even when there is no change in touch, they keep respond to stimuli and fire action potentials. In the body, Type II sensory fibers belong to pseudounipolar neurons. The most notable example are neurons with Merkel cell-neurite complexes on their dendrites and Ruffini endings. Under pathological conditions they may become hyper-excitable leading to stimuli that would usually elicit sensations of tactile touch causing pain. These changes are in part induced by PGE2 which is produced by COX1, and type II fibers with free nerve endings are likely to be the subdivision of fibers that carry out this function.
Eimer's organs are organs for the sense of touch, shaped like bulbous papillae, formed from modified epidermis. First isolated by Theodor Eimer from the European mole in 1871, these organs are present in many moles, and are particularly dense on 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 myelinated fibers from the dermis, which form terminal swellings just below the keratinized outer surface of the epidermis. They contain a complex of Merkel cell and neurite in the epidermis, and a lamellated corpuscle in the dermal connective tissue.
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.
The somatosensory system, or somatic sensory system is a subset of the sensory nervous system. It has two subdivisions, one for the detection of mechanosensory information related to touch, and the other for the nociception detection of pain and temperature. The main functions of the somatosensory system are the perception of external stimuli, the perception of internal stimuli, and the regulation of body position and balance (proprioception).
Group A nerve fibers are one of the three classes of nerve fiber as generally classified by Erlanger and Gasser. The other two classes are the group B nerve fibers, and the group C nerve fibers. Group A are heavily myelinated, group B are moderately myelinated, and group C are unmyelinated.
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.
Ellen Lumpkin is an American neuroscientist and professor of cell and developmental biology and neurobiology at the Helen Wills Neuroscience Institute at the University of California, Berkeley. She is also co-director of the MBL Advanced Training Course in Neurobiology, and adjunct associate professor of physiology and cellular biophysics and co-director of the Thompson Family Foundation Initiative in CIPN and Sensory Neuroscience at Columbia University. Lumpkin's group studies genes, cells and signals that mediate the sensation of touch. Lumpkin is most interested in the somatosensory system and how it gives feedback to the brain on sensations such as pain or touch. She is known for her significant contributions in somatosensory system research.