Thermoreception
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physiology Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
, thermoception or thermoreception is the
sensation Sensation (psychology) refers to the processing of the senses by the sensory system. Sensation or sensations may also refer to: In arts and entertainment In literature *Sensation (fiction), a fiction writing mode *Sensation novel, a British ...
and
perception Perception () is the organization, identification, and interpretation of sensory information in order to represent and understand the presented information or environment. All perception involves signals that go through the nervous syste ...
of
temperature Temperature is a physical quantity that quantitatively expresses the attribute of hotness or coldness. Temperature is measurement, measured with a thermometer. It reflects the average kinetic energy of the vibrating and colliding atoms making ...
, or more accurately, temperature differences inferred from
heat flux In physics and engineering, heat flux or thermal flux, sometimes also referred to as heat flux density, heat-flow density or heat-flow rate intensity, is a flow of energy per unit area per unit time (physics), time. Its SI units are watts per sq ...
. It deals with a series of events and processes required for an organism to receive a temperature stimulus, convert it to a molecular signal, and recognize and characterize the signal in order to trigger an appropriate response. Thermal stimuli may be noxious (posing a threat to the subject) or innocuous (no threat). The temperature sensitive proteins in
thermoreceptor A thermoreceptor is a non-specialised sense Cutaneous receptor, 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 ...
s may also be activated by menthol or capsaicin, hence why these molecules evoke cooling and burning sensations, respectively. A
thermoreceptor A thermoreceptor is a non-specialised sense Cutaneous receptor, 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 ...
may absorb heat via
conduction Conductor or conduction may refer to: Biology and medicine * Bone conduction, the conduction of sound to the inner ear * Conduction aphasia, a language disorder Mathematics * Conductor (ring theory) * Conductor of an abelian variety * Condu ...
,
convection Convection is single or Multiphase flow, multiphase fluid flow that occurs Spontaneous process, spontaneously through the combined effects of material property heterogeneity and body forces on a fluid, most commonly density and gravity (see buoy ...
or
radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or a material medium. This includes: * ''electromagnetic radiation'' consisting of photons, such as radio waves, microwaves, infr ...
. However, the type of
heat transfer Heat transfer is a discipline of thermal engineering that concerns the generation, use, conversion, and exchange of thermal energy (heat) between physical systems. Heat transfer is classified into various mechanisms, such as thermal conduction, ...
is usually irrelevant to the functioning of a thermoceptor.
Transient receptor potential channel Transient receptor potential channels (TRP channels) are a group of ion channels located mostly on the plasma membrane of numerous animal cell types. Most of these are grouped into two broad groups: Group 1 includes TRPC ( "C" for canonical), TRP ...
s (TRP channels) are believed to play a role in many species in sensation of hot, cold, and pain.
Vertebrates Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
have at least two types of thermoreceptors: those that detect heat and those that detect cold.


Heat transfer

Thermoception depends on the transfer of heat from the environment to the thermoreceptor. The transfer may be
conductive In physics and electrical engineering, a conductor is an object or type of material that allows the flow of Electric charge, charge (electric current) in one or more directions. Materials made of metal are common electrical conductors. The flow ...
,
convective Convection is single or multiphase fluid flow that occurs spontaneously through the combined effects of material property heterogeneity and body forces on a fluid, most commonly density and gravity (see buoyancy). When the cause of the convec ...
or radiative, but the method is irrelevant to the thermoceptor, which simply detects its own temperature, not that of the environment. The temperature of a thermoreceptor is the result of an energy balance between the heat flux from the environment and the heat dissipation to the rest of the body (or vice versa for cold detection). For example, a low-temperature metal, with high
thermal conductivity The thermal conductivity of a material is a measure of its ability to heat conduction, conduct heat. It is commonly denoted by k, \lambda, or \kappa and is measured in W·m−1·K−1. Heat transfer occurs at a lower rate in materials of low ...
may feel warmer than a high-temperature ceramic, with low thermal conductivity, because touching the metal results in a higher temperature of the thermoreceptor itself.


Thermoreceptors

A
thermoreceptor A thermoreceptor is a non-specialised sense Cutaneous receptor, 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 ...
is a non-specialised
sense A sense is a biological system used by an organism for sensation, the process of gathering information about the surroundings through the detection of Stimulus (physiology), stimuli. Although, in some cultures, five human senses were traditio ...
receptor Receptor may refer to: * Sensory receptor, in physiology, any neurite structure that, on receiving environmental stimuli, produces an informative nerve impulse *Receptor (biochemistry), in biochemistry, a protein molecule that receives and respond ...
, or more accurately the receptive portion of a
sensory neuron 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 transduc ...
, that detects absolute and relative changes in
temperature Temperature is a physical quantity that quantitatively expresses the attribute of hotness or coldness. Temperature is measurement, measured with a thermometer. It reflects the average kinetic energy of the vibrating and colliding atoms making ...
In mammals, temperature receptors innervate various tissues including the skin (as
cutaneous receptors A cutaneous receptor is a sensory receptor found in the skin that provides information about temperature, touch (including vibration and pain), spatial orientation,pressure (stretching or squeezing), and metabolic circumstances (including those i ...
),
cornea The cornea is the transparency (optics), transparent front part of the eyeball which covers the Iris (anatomy), iris, pupil, and Anterior chamber of eyeball, anterior chamber. Along with the anterior chamber and Lens (anatomy), lens, the cornea ...
and
urinary bladder The bladder () is a hollow organ in humans and other vertebrates that stores urine from the Kidney (vertebrates), kidneys. In placental mammals, urine enters the bladder via the ureters and exits via the urethra during urination. In humans, the ...
. In warm receptors, warming results in an increase in their action potential discharge rate, while cooling results in a decrease in discharge rate. In cold receptors, their firing rate increases during cooling and decreases during warming.


Molecular basis

This area of research has recently received considerable attention with the identification and cloning of the
Transient Receptor Potential Transient receptor potential channels (TRP channels) are a group of ion channels located mostly on the plasma membrane of numerous animal cell types. Most of these are grouped into two broad groups: Group 1 includes TRPC ( "C" for canonical), TRP ...
(TRP) family of proteins. A number of
ion channels Ion channels are pore-forming membrane proteins that allow ions to pass through the channel pore. Their functions include establishing a resting membrane potential, shaping action potentials and other electrical signals by gating the flow of ...
are responsible for thermoception and activate at various temperatures so are therefore responsible for different types of thermal stimuli.
TRPV1 The transient receptor potential cation channel subfamily V member 1 (TRPV1), also known as the capsaicin receptor and the vanilloid receptor 1, is a protein that, in humans, is encoded by the ''TRPV1'' gene. It was the first isolated member of ...
is the primary channel associated with noxious heat sensing, as well as the detection of
capsaicin Capsaicin (8-methyl-''N''-vanillyl-6-nonenamide) (, rarely ) is an active component of chili peppers, which are plants belonging to the genus ''Capsicum''. It is a potent Irritation, irritant for Mammal, mammals, including humans, and produces ...
. Innocuous warm sensation is mediated by activation of
TRPM2 Transient receptor potential cation channel, subfamily M, member 2, also known as TRPM2, is a protein that in humans is encoded by the ''TRPM2'' gene. Structure The protein encoded by this gene is a non-selective calcium-permeable cation channe ...
. Innocuous cool sensation is mediated by activation of
TRPM8 Transient receptor potential cation channel subfamily M (melastatin) member 8 (TRPM8), also known as the cold and menthol receptor 1 (CMR1), is a protein that in humans is encoded by the ''TRPM8'' gene. The TRPM8 channel is the primary molecular ...
.
TRPA1 Transient receptor potential cation channel, subfamily A, member 1, also known as transient receptor potential ankyrin 1, TRPA1, or The Mustard and Wasabi Receptor, is a protein that in humans is encoded by the ''TRPA1'' (and in mice and rats by ...
is sometimes sensitive to
menthol Menthol is an organic compound, specifically a Monoterpene, monoterpenoid, that occurs naturally in the oils of several plants in the Mentha, mint family, such as Mentha arvensis, corn mint and peppermint. It is a white or clear waxy crystallin ...
and considered to be related to noxious cool sensation, but the mechanism is unclear. The
Nobel Prize in Physiology or Medicine The Nobel Prize in Physiology or Medicine () is awarded yearly by the Nobel Assembly at the Karolinska Institute for outstanding discoveries in physiology or medicine. The Nobel Prize is not a single prize, but five separate prizes that, acco ...
in 2021 was attributed to David Julius (professor at the
University of California, San Francisco The University of California, San Francisco (UCSF) is a Public university, public Land-grant university, land-grant research university in San Francisco, California, United States. It is part of the University of California system and is dedic ...
, USA) and
Ardem Patapoutian Ardem Patapoutian (born 1 October 1967) is a Lebanese-American molecular biologist, neuroscientist, and Nobel Prize laureate of Armenian descent. He is known for his work in characterizing the PIEZO1, PIEZO2, and TRPM8 receptors that detect pres ...
(neuroscience professor at
Scripps Research Scripps Research is a nonprofit American medical research facility that focuses on research and education in the biomedical sciences. Headquartered in San Diego, California, the institute has over 170 laboratories employing 2,100 scientists, tec ...
in
La Jolla, California La Jolla ( , ) is a hilly, seaside neighborhood in San Diego, California, occupying of curving coastline along the Pacific Ocean. The population reported in the 2010 census was 46,781. The climate is mild, with an average daily temperature o ...
, USA) "for their discovery of receptors for temperature and touch".


In humans

In humans, temperature sensation from
thermoreceptor A thermoreceptor is a non-specialised sense Cutaneous receptor, 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 ...
s enters the spinal cord along the axons of
Lissauer's tract The posterolateral tract (fasciculus of Lissauer, Lissauer's tract, tract of Lissauer, dorsolateral fasciculus, dorsolateral tract, zone of Lissauer) is a small strand situated in relation to the tip of the posterior column close to the entrance of ...
that synapse on second order neurons in
grey matter Grey matter, or gray matter in American English, is a major component of the central nervous system, consisting of neuronal cell bodies, neuropil ( dendrites and unmyelinated axons), glial cells ( astrocytes and oligodendrocytes), synapses, ...
of the dorsal horn. The axons of these second order neurons then
decussate Decussation is used in biological contexts to describe a crossing (due to the shape of the Roman numeral for ten, an uppercase 'X' (), ). In Latin anatomical terms, the form is used, e.g. . Similarly, the anatomical term chiasma is named aft ...
, joining the
spinothalamic tract The spinothalamic tract is a nerve tract in the anterolateral system in the spinal cord. This tract is an ascending sensory pathway to the thalamus. From the ventral posterolateral nucleus in the thalamus, sensory information is relayed upwar ...
as they ascend to neurons in the
ventral posterolateral nucleus The ventral posterolateral nucleus (VPL) is one of the subdivisions of the ventral posterior nucleus in the ventral nuclear group of the thalamus. It relays sensory information from the second-order neurons of the neospinothalamic tract and me ...
of the
thalamus The thalamus (: thalami; from Greek language, Greek Wikt:θάλαμος, θάλαμος, "chamber") is a large mass of gray matter on the lateral wall of the third ventricle forming the wikt:dorsal, dorsal part of the diencephalon (a division of ...
. A study in 2017 shows that the thermosensory information passes to the lateral parabrachial nucleus rather than to the thalamus and this drives thermoregulatory behaviour.


Thermal vision

Thermal vision is the ability to detect heat through radiative means.
Vision Vision, Visions, or The Vision may refer to: Perception Optical perception * Visual perception, the sense of sight * Visual system, the physical mechanism of eyesight * Computer vision, a field dealing with how computers can be made to gain und ...
specifically denotes the ability to not only detect heat but also form an image with that information. However, given the lack of knowledge or uncertainty of how an organism may interpret their thermoreceptor signals, any organism with organs specifically evolved for radiative thermoception are generally classed as thermal vision.


In snakes

Crotalinae The Crotalinae, commonly known as pit vipers,Mehrtens JM (1987). ''Living Snakes of the World in Color''. New York: Sterling Publishers. 480 pp. . or pit adders, are a subfamily of vipers found in Asia and the Americas. Like all other vipers ...
(pit viper) and
Boidae The Boidae, commonly known as boas or boids, are a family of nonvenomous snakes primarily found in the Americas, as well as Africa, Europe, Asia, and some Pacific islands. Boas include some of the world's largest snakes, with the green anaconda ...
(boa) snakes can effectively see the infrared radiation emitted by hot objects. The snakes' face has a pair of holes, or pits, lined with temperature sensors. The sensors indirectly detect infrared radiation by its heating effect on the skin inside the pit. They can work out which part of the pit is hottest, and therefore the direction of the heat source, which could be a warm-blooded prey animal. By combining information from both pits, the snake can also estimate the distance of the object.


In vampire bats

The
Common vampire bat The common vampire bat (''Desmodus rotundus'') is a small, leaf-nosed bat native to the Americas. It is one of three extant species of vampire bats, the other two being the Hairy-legged vampire bat, hairy-legged and the white-winged vampire bats ...
has specialized infrared sensors in its nose-leaf. Vampire bats are the only mammals that feed exclusively on blood. The infrared sense enables Desmodus to localize homeothermic (warm-blooded) animals (
cattle Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are calle ...
,
horses The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 milli ...
, wild mammals) within a range of about 10 to 15 cm. This infrared perception is possibly used in detecting regions of maximal blood flow on targeted prey.


In other mammals

Dogs, like vampire bats, can detect weak thermal radiation with their rhinaria (noses). On February 14, 2013 researchers developed a
neural implant Brain implants, often referred to as neural implants, are technological devices that connect directly to a biological subject's brain – usually placed on the surface of the brain, or attached to the brain's cortex. A common purpose of modern bra ...
that gives
rat Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus ''Rattus''. Other rat genera include '' Neotoma'' (pack rats), '' Bandicota'' (bandicoo ...
s the ability to sense
infrared Infrared (IR; sometimes called infrared light) is electromagnetic radiation (EMR) with wavelengths longer than that of visible light but shorter than microwaves. The infrared spectral band begins with the waves that are just longer than those ...
light which for the first time provides living creatures with new abilities, instead of simply replacing or augmenting existing abilities.


In invertebrates

Other animals with specialized heat detectors are forest fire seeking beetles ('' Melanophila acuminata''), which lay their eggs in
conifers Conifers () are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All e ...
freshly killed by forest fires. Darkly pigmented butterflies ''
Pachliopta aristolochiae ''Pachliopta aristolochiae'', the common rose, is a species of swallowtail butterfly belonging to the genus ''Pachliopta'', the roses, or red-bodied swallowtails. It is a common butterfly which is extensively distributed across South Asia, south ...
'' and '' Troides rhadamantus'' use specialized heat detectors to avoid damage while basking. The blood sucking bugs ''
Triatoma infestans ''Triatoma infestans'', commonly called winchuka or vinchuca in Argentina, Bolivia, Uruguay and Chile, barbeiro in Brazil, chipo in Venezuela and also known as "kissing bug" or "barber bug" in English, is a blood-sucking bug (like virtually a ...
'' may also have a specialised thermoception organ.


See also

*
Infrared sensing in snakes The ability to sense infrared thermal radiation evolved independently in three different groups of snakes, consisting of the families of Boidae (boas), Pythonidae (pythons), and the subfamily Crotalinae (pit vipers). What is commonly called a p ...
* Infrared sensing in vampire bats * List of Nobel laureates in Physiology or Medicine#Laureates * * * * *
Thermoregulation Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...


Notes


References


Further reading

* {{Authority control Perception Sensory systems Temperature