Thermoreceptor
Encyclopedia
A thermoreceptor is a sensory receptor
, or more accurately the receptive portion of a sensory neuron, that codes absolute and relative changes in temperature, primarily within the innocuous range. In the mammalian peripheral nervous system warmth receptors are thought to be unmyelinated
C-fibres (low conduction velocity), while those responding to cold have both C-fibers and thinly myelinated A delta fiber
s (faster conduction velocity). The adequate stimulus for a warm receptor is warming, which results in an increase in their action potential discharge rate. Cooling results in a decrease in warm receptor discharge rate. For cold receptors their firing rate increases during cooling and decreases during warming. Some cold receptors also respond with a brief action potential discharge to high temperatures, i.e. typically above 45°C, and this is known as a paradoxical response to heat. The mechanism responsible for this behavior has not been determined.
A special form of thermoreceptor is found in some snakes, the viper pit organ and this specialized structure is sensitive to energy in the infrared
part of the spectrum.
and urinary bladder
. Neurons from the pre-optic and hypothalamic regions of the brain that respond to small changes in temperature have also been described, providing information on core temperature. The hypothalamus is involved in thermoregulation
, the thermoreceptors allowing feed-forward
responses to a predicted change in core body temperature in response to changing environmental conditions.
; the mechanism of activation in response to temperature changes is not completely understood.
In the cornea cold receptors are thought to respond with an increase in firing rate to cooling produced by evaporation of lacrimal fluid 'tears' and thereby to elicit a reflex blink.
s that may respond to noxious cold, noxious heat or more than one noxious stimulus modality (i.e., they are polymodal). The nerve endings of sensory neurons that respond preferentially to cooling are found in moderate density in the skin but also occur in relatively high spatial density in the cornea, tongue, bladder, and facial skin. The speculation is that lingual cold receptors deliver information that modulates the sense of taste; i.e. some foods taste good when cold, while others do not.
(TRP) family of proteins. The transduction of temperature in cold receptors is mediated in part by the TRPM8 channel. This channel passes a mixed inward cationic (predominantly carried by Na+ ions although the channel is also permeable to Ca2+) current of a magnitude that is inversely proportional to temperature. The channel is sensitive over a temperature range spanning about 10-35°C. TRPM8 can also be activated by the binding of an extracellular ligand. Menthol can activate the TRPM8 channel in this way. Since the TRPM8 is expressed in neurons whose physiological role is to signal cooling, menthol applied to various bodily surfaces evokes a sensation of cooling. The feeling of freshness associated with the activation of cold receptors by menthol, particularly those in facial areas with axons in the trigeminal (V) nerve, accounts for its use in numerous toiletries including toothpaste, shaving lotions, facial creams and the like.
Another molecular component of cold transduction is the temperature dependence of so-called leak channels which pass an outward current carried by potassium ions. Some leak channels derive from the family of two-pore (2P) domain potassium channels. Amongst the various members of the 2P-domain channels, some close quite promptly at temperatures less than about 28°C (eg. TRAAK, TREK). Temperature also modulates the activity of the Na+/K+-ATPase. The Na+/K+-ATPase is a P-type pump that extrudes 3Na+ ions in exchange for 2K+ ions for each hydrolytic cleavage of ATP. This results in a net movement of positive charge out of the cell, i.e. a hyperpolarizing current. The magnitude of this current is proportional to the rate of pump activity.
It has been suggested that it is the constellation of various thermally sensitive proteins together in a neuron that gives rise to a cold receptor. This emergent property of the neuron is thought to comprise, the expression of the aforementioned proteins as well as various voltage-sensitive channels including the hyperpolarization-activated, cyclic nucleotide-gated (HCN) channel and the rapidly activating and inactivating transient potassium channel (IKA).
Sensory receptor
In a sensory system, a sensory receptor is a sensory nerve ending that responds to a stimulus in the internal or external environment of an organism...
, or more accurately the receptive portion of a sensory neuron, that codes absolute and relative changes in temperature, primarily within the innocuous range. In the mammalian peripheral nervous system warmth receptors are thought to be unmyelinated
Myelin
Myelin is a dielectric material that forms a layer, the myelin sheath, usually around only the axon of a neuron. It is essential for the proper functioning of the nervous system. Myelin is an outgrowth of a type of glial cell. The production of the myelin sheath is called myelination...
C-fibres (low conduction velocity), while those responding to cold have both C-fibers and thinly myelinated A delta fiber
A delta fiber
A delta fibers, or Aδ fibers, are a type of sensory fiber.They are associated with cold and pressure, and as nociceptors stimulation of them is interpreted as fast/first pain information....
s (faster conduction velocity). The adequate stimulus for a warm receptor is warming, which results in an increase in their action potential discharge rate. Cooling results in a decrease in warm receptor discharge rate. For cold receptors their firing rate increases during cooling and decreases during warming. Some cold receptors also respond with a brief action potential discharge to high temperatures, i.e. typically above 45°C, and this is known as a paradoxical response to heat. The mechanism responsible for this behavior has not been determined.
A special form of thermoreceptor is found in some snakes, the viper pit organ and this specialized structure is sensitive to energy in the infrared
Infrared
Infrared light is electromagnetic radiation with a wavelength longer than that of visible light, measured from the nominal edge of visible red light at 0.74 micrometres , and extending conventionally to 300 µm...
part of the spectrum.
Location
In mammals, temperature receptors innervate various tissues including the skin (as cutaneous receptors), corneaCornea
The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Together with the lens, the cornea refracts light, with the cornea accounting for approximately two-thirds of the eye's total optical power. In humans, the refractive power of the cornea is...
and urinary bladder
Urinary bladder
The urinary bladder is the organ that collects urine excreted by the kidneys before disposal by urination. A hollow muscular, and distensible organ, the bladder sits on the pelvic floor...
. Neurons from the pre-optic and hypothalamic regions of the brain that respond to small changes in temperature have also been described, providing information on core temperature. The hypothalamus is involved in thermoregulation
Thermoregulation
Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different...
, the thermoreceptors allowing feed-forward
Feed-forward
Feed-forward is a term describing an element or pathway within a control system which passes a controlling signal from a source in the control system's external environment, often a command signal from an external operator, to a load elsewhere in its external environment...
responses to a predicted change in core body temperature in response to changing environmental conditions.
Structure
Thermoreceptors have been classically described as having 'free' non-specialised endingsFree nerve ending
A free nerve ending is an unspecialized, afferent nerve ending, meaning it brings information from the body's periphery toward the brain. They function as cutaneous receptors and are essentially used by vertebrates to detect pain.-Structure:...
; the mechanism of activation in response to temperature changes is not completely understood.
Function
Cold-sensitive thermoreceptors give rise to the sensations of cooling, cold and freshness.In the cornea cold receptors are thought to respond with an increase in firing rate to cooling produced by evaporation of lacrimal fluid 'tears' and thereby to elicit a reflex blink.
Distribution
Warm and cold receptors play a part in sensing innocuous environmental temperature. Temperatures likely to damage an organism are sensed by sub-categories of nociceptorNociceptor
A nociceptor is a sensory receptor that responds to potentially damaging stimuli by sending nerve signals to the spinal cord and brain. This process, called nociception, usually causes the perception of pain.-History:...
s that may respond to noxious cold, noxious heat or more than one noxious stimulus modality (i.e., they are polymodal). The nerve endings of sensory neurons that respond preferentially to cooling are found in moderate density in the skin but also occur in relatively high spatial density in the cornea, tongue, bladder, and facial skin. The speculation is that lingual cold receptors deliver information that modulates the sense of taste; i.e. some foods taste good when cold, while others do not.
Mechanism of transduction
This area of research has recently received considerable attention with the identification and cloning of the Transient Receptor PotentialTransient receptor potential
Transient receptor potential channels are a group of ion channels located mostly on the plasma membrane of numerous human and animal cell types. There are about 28 TRP channels that share some structural similarity to each other...
(TRP) family of proteins. The transduction of temperature in cold receptors is mediated in part by the TRPM8 channel. This channel passes a mixed inward cationic (predominantly carried by Na+ ions although the channel is also permeable to Ca2+) current of a magnitude that is inversely proportional to temperature. The channel is sensitive over a temperature range spanning about 10-35°C. TRPM8 can also be activated by the binding of an extracellular ligand. Menthol can activate the TRPM8 channel in this way. Since the TRPM8 is expressed in neurons whose physiological role is to signal cooling, menthol applied to various bodily surfaces evokes a sensation of cooling. The feeling of freshness associated with the activation of cold receptors by menthol, particularly those in facial areas with axons in the trigeminal (V) nerve, accounts for its use in numerous toiletries including toothpaste, shaving lotions, facial creams and the like.
Another molecular component of cold transduction is the temperature dependence of so-called leak channels which pass an outward current carried by potassium ions. Some leak channels derive from the family of two-pore (2P) domain potassium channels. Amongst the various members of the 2P-domain channels, some close quite promptly at temperatures less than about 28°C (eg. TRAAK, TREK). Temperature also modulates the activity of the Na+/K+-ATPase. The Na+/K+-ATPase is a P-type pump that extrudes 3Na+ ions in exchange for 2K+ ions for each hydrolytic cleavage of ATP. This results in a net movement of positive charge out of the cell, i.e. a hyperpolarizing current. The magnitude of this current is proportional to the rate of pump activity.
It has been suggested that it is the constellation of various thermally sensitive proteins together in a neuron that gives rise to a cold receptor. This emergent property of the neuron is thought to comprise, the expression of the aforementioned proteins as well as various voltage-sensitive channels including the hyperpolarization-activated, cyclic nucleotide-gated (HCN) channel and the rapidly activating and inactivating transient potassium channel (IKA).