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Sensory neurons, also known as afferent neurons, are
neuron A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa ...
s in the
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes ...
, that convert a specific type of stimulus, via their receptors, into
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 graded potentials. This process is called sensory transduction. The cell bodies of the sensory neurons are located in the dorsal ganglia of the
spinal cord The spinal cord is a long, thin, tubular structure made up of nervous tissue, which extends from the medulla oblongata in the brainstem to the lumbar region of the vertebral column (backbone). The backbone encloses the central canal of the sp ...
. The sensory information travels on the afferent nerve fibers in a sensory nerve, to the
brain A brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as Visual perception, vision. I ...
via the
spinal cord The spinal cord is a long, thin, tubular structure made up of nervous tissue, which extends from the medulla oblongata in the brainstem to the lumbar region of the vertebral column (backbone). The backbone encloses the central canal of the sp ...
. The stimulus can come from ''exteroreceptors'' outside the body, for example those that detect light and sound, or from ''interoreceptors'' inside the body, for example those that are responsive to
blood pressure Blood pressure (BP) is the pressure of circulating blood against the walls of blood vessels. Most of this pressure results from the heart pumping blood through the circulatory system. When used without qualification, the term "blood pressure ...
or the sense of body position.


Types and function

Different types of sensory neurons have different sensory receptors that respond to different kinds of stimuli. There are at least six external and two internal sensory receptors:


External receptors

External receptors that respond to stimuli from outside the body are called exteroreceptors. Exteroreceptors include olfactory receptors (smell), taste receptors, photoreceptors (vision), hair cells (hearing), thermoreceptors (temperature), and a number of different mechanoreceptors (stretch, distortion).


Smell

The sensory neurons involved in smell are called olfactory sensory neurons. These neurons contain
receptor Receptor may refer to: *Sensory receptor, in physiology, any structure which, on receiving environmental stimuli, produces an informative nerve impulse *Receptor (biochemistry), in biochemistry, a protein molecule that receives and responds to a n ...
s, called olfactory receptors, that are activated by odor molecules in the air. The molecules in the air are detected by enlarged cilia and
microvilli Microvilli (singular: microvillus) are microscopic cellular membrane protrusions that increase the surface area for diffusion and minimize any increase in volume, and are involved in a wide variety of functions, including absorption, secretion, ...
. These sensory neurons produce action potentials. Their axons form the olfactory nerve, and they synapse directly onto neurons in the cerebral cortex (olfactory bulb). They do not use the same route as other sensory systems, bypassing the brain stem and the thalamus. The neurons in the olfactory bulb that receive direct sensory nerve input, have connections to other parts of the olfactory system and many parts of the limbic system. 9.


Taste

Similar to olfactory receptors, taste receptors (gustatory receptors) in taste buds interact with chemicals in food to produce an
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 ...
.


Vision

Photoreceptor cells are capable of phototransduction, a process which converts light (
electromagnetic radiation In physics, electromagnetic radiation (EMR) consists of waves of the electromagnetic (EM) field, which propagate through space and carry momentum and electromagnetic radiant energy. It includes radio waves, microwaves, infrared, (visib ...
) into electrical signals. These signals are refined and controlled by the interactions with other types of neurons in the retina. The five basic classes of neurons within the retina are photoreceptor cells,
bipolar cells A bipolar neuron, or bipolar cell, is a type of neuron that has two extensions (one axon and one dendrite). Many bipolar cells are specialized sensory neurons for the transmission of sense. As such, they are part of the sensory pathways for smell, ...
, ganglion cells,
horizontal cells Horizontal cells are the laterally interconnecting neurons having cell bodies in the inner nuclear layer of the retina of vertebrate eyes. They help integrate and regulate the input from multiple photoreceptor cells. Among their functions, horizo ...
, and
amacrine cells Amacrine cells are interneurons in the retina. They are named from the Greek roots ''a–'' ("non"), ''makr–'' ("long") and ''in–'' ("fiber"), because of their short neuronal processes. Amacrine cells are inhibitory neurons, and they proje ...
. The basic circuitry of the retina incorporates a three-neuron chain consisting of the photoreceptor (either a rod or cone), bipolar cell, and the ganglion cell. The first action potential occurs in the retinal ganglion cell. This pathway is the most direct way for transmitting visual information to the brain. There are three primary types of photoreceptors: Cones are photoreceptors that respond significantly to
color Color (American English) or colour (British English) is the visual perceptual property deriving from the spectrum of light interacting with the photoreceptor cells of the eyes. Color categories and physical specifications of color are associ ...
. In humans the three different types of cones correspond with a primary response to short wavelength (blue), medium wavelength (green), and long wavelength (yellow/red)."eye, human." Encyclopædia Britannica. Encyclopædia Britannica Ultimate Reference Suite. Chicago: Encyclopædia Britannica, 2010. Rods are photoreceptors that are very sensitive to the intensity of light, allowing for vision in dim lighting. The concentrations and ratio of rods to cones is strongly correlated with whether an animal is diurnal or nocturnal. In humans, rods outnumber cones by approximately 20:1, while in nocturnal animals, such as the tawny owl, the ratio is closer to 1000:1. Retinal ganglion cells are involved in the sympathetic response. Of the ~1.3 million ganglion cells present in the retina, 1-2% are believed to be photosensitive. Issues and decay of sensory neurons associated with vision lead to disorders such as: *
Macular degeneration Macular degeneration, also known as age-related macular degeneration (AMD or ARMD), is a medical condition which may result in blurred or no vision in the center of the visual field. Early on there are often no symptoms. Over time, however, so ...
– degeneration of the central visual field due to either cellular debris or blood vessels accumulating between the retina and the choroid, thereby disturbing and/or destroying the complex interplay of neurons that are present there. *
Glaucoma Glaucoma is a group of eye diseases that result in damage to the optic nerve (or retina) and cause vision loss. The most common type is open-angle (wide angle, chronic simple) glaucoma, in which the drainage angle for aqueous humor, fluid withi ...
– loss of retinal ganglion cells which causes some loss of vision to blindness. * Diabetic retinopathy – poor blood sugar control due to diabetes damages the tiny blood vessels in the retina.


Auditory

The
auditory system The auditory system is the sensory system for the sense of hearing. It includes both the sensory organs (the ears) and the auditory parts of the sensory system. System overview The outer ear funnels sound vibrations to the eardrum, increasin ...
is responsible for converting pressure waves generated by vibrating air molecules or
sound In physics, sound is a vibration that propagates as an acoustic wave, through a transmission medium such as a gas, liquid or solid. In human physiology and psychology, sound is the ''reception'' of such waves and their ''perception'' by ...
into signals that can be interpreted by the brain. This mechanoelectrical transduction is mediated with hair cells within the ear. Depending on the movement, the hair cell can either hyperpolarize or depolarize. When the movement is towards the tallest stereocilia, the Na+ cation channels open allowing Na+ to flow into cell and the resulting depolarization causes the Ca++ channels to open, thus releasing its neurotransmitter into the afferent auditory nerve. There are two types of hair cells: inner and outer. The inner hair cells are the sensory receptors . Problems with sensory neurons associated with the auditory system leads to disorders such as: * Auditory processing disorder – Auditory information in the brain is processed in an abnormal way. Patients with auditory processing disorder can usually gain the information normally, but their brain cannot process it properly, leading to hearing disability. * Auditory verbal agnosia – Comprehension of speech is lost but hearing, speaking, reading, and writing ability is retained. This is caused by damage to the posterior superior
temporal lobes The temporal lobe is one of the four major lobes of the cerebral cortex in the brain of mammals. The temporal lobe is located beneath the lateral fissure on both cerebral hemispheres of the mammalian brain. The temporal lobe is involved in pro ...
, again not allowing the brain to process auditory input correctly.


Temperature

Thermoreceptors are sensory receptors, which respond to varying
temperature Temperature is a physical quantity that expresses quantitatively the perceptions of hotness and coldness. Temperature is measured with a thermometer. Thermometers are calibrated in various temperature scales that historically have relied o ...
s. While the mechanisms through which these receptors operate is unclear, recent discoveries have shown that
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s have at least two distinct types of thermoreceptors.Krantz, John.
Experiencing Sensation and Perception
''. Pearson Education, Limited, 2009. p. 12.3
The
bulboid corpuscle The bulboid corpuscles (end-bulbs of Krause) are cutaneous receptors in the human body. The end-bulbs of Krause were named after the German anatomist Wilhelm Krause (1833–1910). Function The end-bulbs of Krause were thought to be thermorecepto ...
, is a
cutaneous receptor A cutaneous receptor is the type of sensory receptor found in the skin ( the dermis or epidermis). They are a part of the somatosensory system. Cutaneous receptors include mechanoreceptors (pressure or distortion), nociceptors (pain), and thermor ...
a ''cold-sensitive'' receptor, that detects cold temperatures. The other type is a warmth-sensitive receptor.


Mechanoreceptors

Mechanoreceptors are sensory receptors which respond to mechanical forces, such as
pressure Pressure (symbol: ''p'' or ''P'') is the force applied perpendicular to the surface of an object per unit area over which that force is distributed. Gauge pressure (also spelled ''gage'' pressure)The preferred spelling varies by country a ...
or distortion. Specialized sensory receptor cells called mechanoreceptors often encapsulate afferent fibers to help tune the afferent fibers to the different types of somatic stimulation. Mechanoreceptors also help lower thresholds for action potential generation in afferent fibers and thus make them more likely to fire in the presence of sensory stimulation. Some types of mechanoreceptors fire action potentials when their membranes are physically stretched. Proprioceptors are another type of mechanoreceptors which literally means "receptors for self". These receptors provide spatial information about limbs and other body parts. Nociceptors are responsible for processing pain and temperature changes. The burning pain and irritation experienced after eating a chili pepper (due to its main ingredient, capsaicin), the cold sensation experienced after ingesting a chemical such as menthol or icillin, as well as the common sensation of pain are all a result of neurons with these receptors. Problems with mechanoreceptors lead to disorders such as: * Neuropathic pain - a severe pain condition resulting from a damaged sensory nerve * Hyperalgesia - an increased sensitivity to pain caused by sensory ion channel, TRPM8, which is typically responds to temperatures between 23 and 26 degrees, and provides the cooling sensation associated with menthol and icillin *
Phantom limb syndrome A phantom limb is the sensation that an amputated or missing limb is still attached. Approximately 80 to 100% of individuals with an amputation experience sensations in their amputated limb. However, only a small percentage will experience painf ...
- a sensory system disorder where pain or movement is experienced in a limb that does not exist


Internal receptors

Internal receptors that respond to changes inside the body are known as interoceptors.


Blood

The
aortic bodies The aortic bodies are one of several small clusters of peripheral chemoreceptors located along the aortic arch. They are important in measuring partial pressures of oxygen and carbon dioxide in the blood, and blood pH. Structure The aortic bodies ...
and carotid bodies contain clusters of glomus cells – peripheral chemoreceptors that detect changes in chemical properties in the blood such as
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements ...
concentration. These receptors are polymodal responding to a number of different stimuli.


Nociceptors

Nociceptors respond to potentially damaging stimuli by sending signals to the spinal cord and brain. This process, called nociception, usually causes the perception of
pain Pain is a distressing feeling often caused by intense or damaging stimuli. The International Association for the Study of Pain defines pain as "an unpleasant sensory and emotional experience associated with, or resembling that associated with, ...
. They are found in internal organs as well as on the surface of the body to "detect and protect". Nociceptors detect different kinds of noxious stimuli indicating potential for damage, then initiate neural responses to withdraw from the stimulus. * ''Thermal'' nociceptors are activated by noxious heat or cold at various temperatures. * ''Mechanical'' nociceptors respond to excess pressure or mechanical deformation, such as a pinch. * ''Chemical'' nociceptors respond to a wide variety of chemicals, some of which signal a response. They are involved in the detection of some spices in food, such as the pungent ingredients in Brassica and Allium plants, which target the sensory neural receptor to produce acute pain and subsequent pain hypersensitivity.


Connection with the central nervous system

Information coming from the sensory neurons in the head enters the
central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all p ...
(CNS) through cranial nerves. Information from the sensory neurons below the head enters the spinal cord and passes towards the brain through the 31 spinal nerves. The sensory information traveling through the spinal cord follows well-defined pathways. The nervous system codes the differences among the sensations in terms of which cells are active.


Classification


Adequate stimulus

A sensory receptor's
adequate stimulus The adequate stimulus is a property of a sensory receptor that determines the type of energy to which a sensory receptor responds with the initiation of sensory transduction. Sensory receptor are specialized to respond to certain types of stimuli. T ...
is the
stimulus modality 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: ligh ...
for which it possesses the adequate sensory transduction apparatus. Adequate stimulus can be used to classify sensory receptors: *
Baroreceptor Baroreceptors (or archaically, pressoreceptors) are sensors located in the carotid sinus (at the bifurcation of external and internal carotids) and in the aortic arch. They sense the blood pressure and relay the information to the brain, so that ...
s respond to pressure in blood vessels * Chemoreceptors respond to chemical stimuli *
Electromagnetic radiation receptors In physics, electromagnetism is an interaction that occurs between particles with electric charge. It is the second-strongest of the four fundamental interactions, after the strong force, and it is the dominant force in the interactions of a ...
respond to
electromagnetic radiation In physics, electromagnetic radiation (EMR) consists of waves of the electromagnetic (EM) field, which propagate through space and carry momentum and electromagnetic radiant energy. It includes radio waves, microwaves, infrared, (visib ...
** Infrared receptors respond to infrared radiation ** Photoreceptors respond to
visible light Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 t ...
** Ultraviolet receptors respond to ultraviolet radiation *
Electroreceptor Electroreception and electrogenesis are the closely-related biological abilities to perceive electrical stimuli and to generate electric fields. Both are used to locate prey; stronger electric discharges are used in a few groups of fishes to stu ...
s respond to electric fields **
Ampullae of Lorenzini Ampullae of Lorenzini (singular ''Ampulla'') are electroreceptors, sense organs able to detect electric fields. They form a network of mucus-filled pores in the skin of cartilaginous fish (sharks, rays, and chimaeras) and of basal actinopterygi ...
respond to electric fields, salinity, and to temperature, but function primarily as electroreceptors * Hydroreceptors respond to changes in humidity * Magnetoreceptors respond to magnetic fields * Mechanoreceptors respond to mechanical stress or
mechanical strain Mechanical may refer to: Machine * Machine (mechanical), a system of mechanisms that shape the actuator input to achieve a specific application of output forces and movement * Mechanical calculator, a device used to perform the basic operations of ...
* Nociceptors respond to damage, or threat of damage, to body tissues, leading (often but not always) to pain perception * Osmoreceptors respond to the osmolarity of fluids (such as in the hypothalamus) * Proprioceptors provide the sense of position * Thermoreceptors respond to temperature, either heat, cold or both


Location

Sensory receptors can be classified by location: *
Cutaneous receptor A cutaneous receptor is the type of sensory receptor found in the skin ( the dermis or epidermis). They are a part of the somatosensory system. Cutaneous receptors include mechanoreceptors (pressure or distortion), nociceptors (pain), and thermor ...
s are sensory receptors found in the
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided ...
or epidermis. * Muscle spindles contain mechanoreceptors that detect stretch in muscles.


Morphology

Somatic sensory receptors near the surface of the skin can usually be divided into two groups based on morphology: *
Free nerve endings A free nerve ending (FNE) or bare nerve ending, is an unspecialized, afferent nerve fiber sending its signal to a sensory neuron. ''Afferent'' in this case means bringing information from the body's periphery toward the brain. They function as cut ...
characterize the nociceptors and thermoreceptors and are called thus because the terminal branches of the neuron are unmyelinated and spread throughout the
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided ...
and epidermis. * Encapsulated receptors consist of the remaining types of cutaneous receptors. Encapsulation exists for specialized functioning.


Rate of adaptation

* A tonic receptor is a sensory receptor that adapts slowly to a stimulus and continues to produce
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 over the duration of the stimulus.mentor.lscf.ucsb.edu/course/fall/eemb157/lecture/Lectures%2016,%2017%2018.ppt In this way it conveys information about the duration of the stimulus. Some tonic receptors are permanently active and indicate a background level. Examples of such tonic receptors are
pain receptors Nociception (also nocioception, from Latin ''nocere'' 'to harm or 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, ...
, joint capsule, and muscle spindle. * A phasic receptor is a sensory receptor that adapts rapidly to a stimulus. The response of the cell diminishes very quickly and then stops. It does not provide information on the duration of the stimulus; instead some of them convey information on rapid changes in stimulus intensity and rate. An example of a phasic receptor is the Pacinian corpuscle.


Drugs

There are many drugs currently on the market that are used to manipulate or treat sensory system disorders. For instance, Gabapentin is a drug that is used to treat neuropathic pain by interacting with one of the voltage-dependent calcium channels present on non-receptive neurons. Some drugs may be used to combat other health problems, but can have unintended side effects on the sensory system. Ototoxic drugs are drugs which affect the
cochlea The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus. A core component of the cochlea is the Organ of Corti, the sensory o ...
through the use of a toxin like aminoglycoside antibiotics, which poison hair cells. Through the use of these toxins, the K+ pumping hair cells cease their function. Thus, the energy generated by the
endocochlear potential The endocochlear potential (EP; also called endolymphatic potential) is the positive voltage of 80-100mV seen in the cochlear endolymphatic spaces. Within the cochlea the EP varies in the magnitude all along its length. When a sound is presented, th ...
which drives the auditory signal transduction process is lost, leading to hearing loss.


Neuroplasticity

Ever since scientists observed
cortical remapping Cortical remapping, also referred to as cortical reorganization, is the process by which an existing cortical map is affected by a stimulus resulting in the creating of a 'new' cortical map. Every part of the body is connected to a corresponding ...
in the brain of Taub's Silver Spring monkeys, there has been a large amount of research into sensory system plasticity. Huge strides have been made in treating disorders of the sensory system. Techniques such as
constraint-induced movement therapy Constraint-induced movement therapy (CI, CIT, or CIMT) is a form of rehabilitation therapy that improves upper extremity function in stroke and other central nervous system damage patients by increasing the use of their affected upper limb.
developed by Taub have helped patients with paralyzed limbs regain use of their limbs by forcing the sensory system to grow new
neural pathway In neuroanatomy, a neural pathway is the connection formed by axons that project from neurons to make synapses onto neurons in another location, to enable neurotransmission (the sending of a signal from one region of the nervous system to ...
s.
Phantom limb syndrome A phantom limb is the sensation that an amputated or missing limb is still attached. Approximately 80 to 100% of individuals with an amputation experience sensations in their amputated limb. However, only a small percentage will experience painf ...
is a sensory system disorder in which amputees perceive that their amputated limb still exists and they may still be experiencing pain in it. The mirror box developed by V.S. Ramachandran, has enabled patients with
phantom limb syndrome A phantom limb is the sensation that an amputated or missing limb is still attached. Approximately 80 to 100% of individuals with an amputation experience sensations in their amputated limb. However, only a small percentage will experience painf ...
to relieve the perception of paralyzed or painful phantom limbs. It is a simple device which uses a mirror in a box to create an illusion in which the sensory system perceives that it is seeing two hands instead of one, therefore allowing the sensory system to control the "phantom limb". By doing this, the sensory system can gradually get acclimated to the amputated limb, and thus alleviate this syndrome.


Other animals

Hydrodynamic reception Hydrodynamic reception refers to the ability of some animals to sense water movements generated by biotic (conspecifics, predators, or prey) or abiotic sources. This form of mechanoreception is useful for orientation, hunting, predator avoidance, ...
is a form of mechanoreception used in a range of animal species.


Additional images

File:Blausen 0809 Skin TactileReceptors.png, Illustration of Tactile Receptors in the Skin File:Blausen 0804 Skin LamellatedCorpuscle.png, Illustration of Lamellated Corpuscle File:Blausen 0807 Skin RuffiniCorpuscle.png, Illustration of Ruffini Corpuscle File:Blausen 0805 Skin MerkelCell.png, Illustration of Skin Merkel Cell File:Blausen 0808 Skin TactileCorpuscle.png, Illustration of Tactile Corpuscle File:Blausen 0806 Skin RootHairPlexus.png, Illustration of Root Hair Plexus File:Blausen 0803 Skin FreeNerveEndings.png, Illustration of Free Nerve Endings


See also

* Sensory nerve * Pseudounipolar neuron * Efferent nerve * Neural coding * Posterior column * Receptive field * Sensory system


References


External links

*
The major classes of somatic sensory receptors
{{Somatosensory system Afferent neurons Human cells Sensory receptors Receptor cells