Mechanoreceptor
A mechanoreceptor, also called mechanoceptor, is a
Vertebrate mechanoreceptors
Cutaneous mechanoreceptors
Cutaneous mechanoreceptors respond to mechanical stimuli that result from physical interaction, including pressure and vibration. They are located in the skin, like other
By sensation
- The Slowly Adapting type 1 (SA1) mechanoreceptor, with the Merkel corpuscle end-organ (also known as Merkel discs) detect sustained pressure and underlies the perception of form and roughness on the skin.[1]They have small receptive fields and produce sustained responses to static stimulation.
- The Slowly Adapting type 2 (SA2) mechanoreceptors, with the Ruffini corpuscle end-organ (also known as the bulbous corpuscles), detect tension deep in the skin and fascia and respond to skin stretch, but have not been closely linked to either proprioceptive or mechanoreceptive roles in perception.[2]They also produce sustained responses to static stimulation, but have large receptive fields.
- The Rapidly Adapting (RA) or Meissner corpuscle end-organ mechanoreceptor (also known as the tactile corpuscles) underlies the perception of light touch such as flutter[3] and slip on the skin.[4]It adapts rapidly to changes in texture (vibrations around 50 Hz). They have small receptive fields and produce transient responses to the onset and offset of stimulation.
- The They also produce transient responses, but have large receptive fields.
- Free nerve endings detect touch, pressure, stretching, as well as the tickle and itch sensations. Itch sensations are caused by stimulation of free nerve ending from chemicals.[7]
- Hair follicle receptors called hair root plexuses sense when a hair changes position. Indeed, the most sensitive mechanoreceptors in humans are the hair cells in the cochlea of the inner ear (no relation to the follicular receptors – they are named for the hair-like mechanosensory stereocilia they possess); these receptors transduce sound for the brain.[7]
By rate of adaptation
Cutaneous mechanoreceptors can also be separated into categories based on their rates of adaptation. When a mechanoreceptor receives a stimulus, it begins to fire impulses or action potentials at an elevated frequency (the stronger the stimulus, the higher the frequency). The cell, however, will soon "adapt" to a constant or static stimulus, and the pulses will subside to a normal rate. Receptors that adapt quickly (i.e., quickly return to a normal pulse rate) are referred to as "phasic". Those receptors that are slow to return to their normal firing rate are called tonic. Phasic mechanoreceptors are useful in sensing such things as texture or vibrations, whereas tonic receptors are useful for temperature and proprioception among others.
- Slowly adapting: Slowly adapting mechanoreceptors include free nerve endings.
- Slowly adapting type I mechanoreceptors have multiple Merkel corpuscle end-organs.
- Slowly adapting type II mechanoreceptors have single Ruffini corpuscle end-organs.
- Slowly adapting type I mechanoreceptors have multiple
- Intermediate adapting: Some free nerve endingsare intermediate adapting.
- Rapidly adapting: Rapidly adapting mechanoreceptors include free nerve endings.
- Rapidly adapting type I mechanoreceptors have multiple Meissner corpuscle end-organs.
- Rapidly adapting type II mechanoreceptors (usually called Pacinian) have single Pacinian corpuscle end-organs.
- Rapidly adapting type I mechanoreceptors have multiple
By receptive field
Cutaneous mechanoreceptors with small, accurate
Lamellar corpuscles
Deforming the corpuscle creates a generator potential in the sensory neuron arising within it. This is a graded response: the greater the deformation, the greater the generator potential. If the generator potential reaches threshold, a volley of action potentials (nerve impulses) are triggered at the first node of Ranvier of the sensory neuron.
Once threshold is reached, the magnitude of the stimulus is encoded in the frequency of impulses generated in the neuron. So the more massive or rapid the deformation of a single corpuscle, the higher the frequency of nerve impulses generated in its neuron.
The optimal sensitivity of a lamellar corpuscle is 250 Hz, the frequency range generated upon finger tips by textures made of features smaller than 200 micrometres.[9]
Ligamentous mechanoreceptors
There are four types of mechanoreceptors embedded in ligaments. As all these types of mechanoreceptors are myelinated, they can rapidly transmit sensory information regarding joint positions to the central nervous system.[10]
- Type I: (small) Low threshold, slow adapting in both static and dynamic settings
- Type II: (medium) Low threshold, rapidly adapting in dynamic settings
- Type III: (large) High threshold, slowly adapting in dynamic settings
- Type IV: (very small) High threshold pain receptors that communicate injury
Type II and Type III mechanoreceptors in particular are believed to be linked to one's sense of proprioception.
Other mechanoreceptors
Other mechanoreceptors than cutaneous ones include the
Muscle spindles and the stretch reflex
The
- Some of the branches of the I-a axons synapse directly with alpha motor neurons. These carry impulses back to the same muscle causing it to contract. The leg straightens.
- Some of the branches of the I-a axons synapse with inhibitory interneurons in the spinal cord. These, in turn, synapse with motor neurons leading back to the antagonistic muscle, a flexor in the back of the thigh. By inhibiting the flexor, these interneurons aid contraction of the extensor.
- Still other branches of the I-a axons synapse with interneurons leading to brain centers, e.g., the cerebellum, that coordinate body movements.[11]
Mechanism of sensation
In
More recent work has expanded the role of the cutaneous mechanoreceptors for feedback in
Invertebrate mechanoreceptors
Insect and arthropod mechanoreceptors include:[14]
- Campaniform sensilla: Small domes in the exoskeleton that are distributed all along the insect's body. These cells are thought to detect mechanical load as resistance to muscle contraction, similar to the mammalian Golgi tendon organs.
- Hair plates: Sensory neurons that innervate hairs that are found in the folds of insect joints. These hairs are deflected when one body segment moves relative to an adjoining segment, they have proprioceptive function, and are thought to act as limit detectors encoding the extreme ranges of motion for each joint.[15]
- Chordotonal organs: Internal stretch receptors at the joints, can have both extero- and proprioceptive functions. The neurons in the chordotonal organ in Drosophila melanogaster can be organized into club, claw, and hook neurons. Club neurons are thought to encode vibrational signals while claw and hook neurons can be subdivided into extension and flexion populations that encode joint angle and movement respectively.[16]
- Slit sensilla:Slits in the exoskeleton that detect physical deformation of the animal's exoskeleton, have proprioceptive function
- Bristle sensilla: Bristle neurons are mechanoreceptors that innervate hairs all along the body. Each neuron extends a dendritic process to innervate a single hair and projects its axon to the ventral nerve cord. These neurons are thought to mediate touch sensation by responding to physical deflections of the hair.[17] In line with the fact that many insects exhibit different sized hairs, commonly referred to as macrochaetes (thicker longer hairs) and microchaetes (thinner shorter hairs), previous studies suggest that bristle neurons to these different hairs may have different firing properties such as resting membrane potential and firing threshold.[18][19]
Plant mechanoreceptors
Mechanoreceptors are also present in plant cells where they play an important role in normal growth, development and the sensing of their environment.[20] Mechanoreceptors aid the Venus flytrap (Dionaea muscipula Ellis) in capturing large[21] prey.[22]
Molecular biology
Mechanoreceptor proteins are ion channels whose ion flow is induced by touch. Early research showed that touch transduction in the nematode Caenorhabditis elegans was found to require a two transmembrane, amiloride-sensitive ion channel protein related to epithelial sodium channels (ENaCs).[23] This protein, called MEC-4, forms a heteromeric Na+-selective channel together with MEC-10. Related genes in mammals are expressed in sensory neurons and were shown to be gated by low pH. The first of such receptor was ASIC1a, named so because it is an acid sensing ion channel (ASIC).[24]
See also
References
- PMID 1575442.
- PMID 7234450.
- ^ PMID 4972033.
- S2CID 22450227.
- S2CID 24658742.
- S2CID 15326972.
- ^ OCLC 1059417106.
- .
- S2CID 14459552.
- PMID 7706334.
- ^ Kimball JW (2011). "Mechanoreceptors". Kimball's Biology Pages. Archived from the original on 27 February 2011.
- S2CID 17298704.
- PMID 11744774.
- PMID 27780045.
- S2CID 2634261.
- S2CID 52927792.
- PMID 26919434.
- PMID 2154560.
- S2CID 2187830.
- PMID 23913953.
- OCLC 755641050.
- PMID 24618927.
- S2CID 4334128.
- S2CID 6721868.
External links
- Mechanoreceptors at the U.S. National Library of Medicine Medical Subject Headings (MeSH)