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The sensory nervous system is a part of 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 th ...
responsible for processing
sensory information. A sensory system consists of
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 potentials. This process is called sensory transduction. The cell ...
s (including the sensory receptor cells),
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 ano ...
s, and parts of the
brain
A brain is an 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 vision. It is the most complex organ in a v ...
involved in sensory
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 system ...
. Commonly recognized sensory systems are those for
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 un ...
,
hearing
Hearing, or auditory perception, is the ability to perceive sounds
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 psycholog ...
,
touch
In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch (haptic perception), as well as temperature (thermoception), body position (proprioception), and pain. It is ...
,
taste
The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
,
smell
Smell may refer to;
* Odor, airborne molecules perceived as a scent or aroma
* Sense of smell, the scent also known scientifically as olfaction
* "Smells" (''Bottom''), an episode of ''Bottom''
* The Smell, a music venue in Los Angeles, Californ ...
, and
balance
Balance or balancing may refer to:
Common meanings
* Balance (ability) in biomechanics
* Balance (accounting)
* Balance or weighing scale
* Balance as in equality or equilibrium
Arts and entertainment Film
* ''Balance'' (1983 film), a Bulgarian ...
. Senses are
transducers
A transducer is a device that converts energy from one form to another. Usually a transducer converts a signal in one form of energy to a signal in another.
Transducers are often employed at the boundaries of automation, measurement, and contro ...
from the physical world to the realm of the mind where people interpret the information, creating their
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 system ...
of the world around them.
The
receptive field
The receptive field, or sensory space, is a delimited medium where some physiological stimuli can evoke a sensory neuronal response in specific organisms.
Complexity of the receptive field ranges from the unidimensional chemical structure of od ...
is the area of the body or environment to which a receptor organ and receptor cells respond. For instance, the part of the world an eye can see, is its receptive field; the light that each
rod or
cone
A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex.
A cone is formed by a set of line segments, half-lines, or lines con ...
can see, is its receptive field.
Receptive field
The receptive field, or sensory space, is a delimited medium where some physiological stimuli can evoke a sensory neuronal response in specific organisms.
Complexity of the receptive field ranges from the unidimensional chemical structure of od ...
s have been identified for the
visual system
The visual system comprises the sensory organ (the eye) and parts of the central nervous system (the retina containing photoreceptor cells, the optic nerve, the optic tract and the visual cortex) which gives organisms the sense of sight (the a ...
,
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 ...
and
somatosensory system
In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch (haptic perception), as well as temperature (thermoception), body position (proprioception), and pain. It ...
.
Stimulus
:Organisms need information to solve at least three kinds of problems: (a) to maintain an appropriate environment, i.e.,
homeostasis
In biology, homeostasis (British English, British also homoeostasis) Help:IPA/English, (/hɒmɪə(ʊ)ˈsteɪsɪs/) is the state of steady internal, physics, physical, and chemistry, chemical conditions maintained by organism, living systems. Thi ...
; (b) to time activities (e.g., seasonal changes in behavior) or synchronize activities with those of
conspecifics
Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species.
Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organis ...
; and (c) to locate and respond to resources or threats (e.g., by moving towards resources or evading or attacking threats). Organisms also need to transmit information in order to influence another's behavior: to identify themselves, warn conspecifics of danger, coordinate activities, or deceive.
Sensory systems code for four aspects of a
stimulus
A stimulus is something that causes a physiological response. It may refer to:
*Stimulation
**Stimulus (physiology), something external that influences an activity
**Stimulus (psychology), a concept in behaviorism and perception
*Stimulus (economi ...
; type (
modality
Modality may refer to:
Humanities
* Modality (theology), the organization and structure of the church, as distinct from sodality or parachurch organizations
* Modality (music), in music, the subject concerning certain diatonic scales
* Modalitie ...
), intensity, location, and duration. Arrival
time
Time is the continued sequence of existence and events that occurs in an apparently irreversible succession from the past, through the present, into the future. It is a component quantity of various measurements used to sequence events, to ...
of a sound
pulse
In medicine, a pulse represents the tactile arterial palpation of the cardiac cycle (heartbeat) by trained fingertips. The pulse may be palpated in any place that allows an artery to be compressed near the surface of the body, such as at the nec ...
and
phase
Phase or phases may refer to:
Science
*State of matter, or phase, one of the distinct forms in which matter can exist
*Phase (matter), a region of space throughout which all physical properties are essentially uniform
* Phase space, a mathematic ...
differences of continuous sound are used for
sound localization
Sound localization is a listener's ability to identify the location or origin of a detected sound in direction and distance.
The sound localization mechanisms of the mammalian auditory system have been extensively studied. The auditory system u ...
. Certain receptors are sensitive to certain types of stimuli (for example, different
mechanoreceptor
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, ...
s respond best to different kinds of touch stimuli, like sharp or blunt objects). Receptors
send impulses in certain patterns to send information about the intensity of a stimulus (for example, how loud a sound is). The location of the receptor that is stimulated gives the brain information about the location of the stimulus (for example, stimulating a mechanoreceptor in a finger will send information to the brain about that finger). The duration of the stimulus (how long it lasts) is conveyed by firing patterns of receptors. These impulses are transmitted to the brain through
afferent neurons
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 potentials. This process is called sensory transduction. The cel ...
.
Quiescent state
Most sensory systems have a quiescent state, that is, the state that a sensory system converges to when there is no input.
This is well-defined for a
linear time-invariant system
In system analysis, among other fields of study, a linear time-invariant (LTI) system is a system that produces an output signal from any input signal subject to the constraints of linearity and time-invariance; these terms are briefly define ...
, whose input space is a vector space, and thus by definition has a point of zero. It is also well-defined for any passive sensory system, that is, a system that operates without needing input power. The quiescent state is the state the system converges to when there is no input power.
It is not always well-defined for nonlinear, nonpassive sensory organs, since they can't function without input energy. For example, a cochlea is not a passive organ, but actively vibrates its own sensory hairs to improve its sensitivity. This manifests as
otoacoustic emission
An otoacoustic emission (OAE) is a sound that is generated from within the inner ear. Having been predicted by Austrian astrophysicist Thomas Gold in 1948, its existence was first demonstrated experimentally by British physicist David Kemp in 1978 ...
s in healthy ears, and
tinnitus
Tinnitus is the perception of sound when no corresponding external sound is present. Nearly everyone experiences a faint "normal tinnitus" in a completely quiet room; but it is of concern only if it is bothersome, interferes with normal hearin ...
in pathological ears. There is still a quiescent state for the cochlea, since there is a well-defined mode of power input that it receives (vibratory energy on the eardrum), which provides an unambiguous definition of "zero input power".
Some sensory systems can have multiple quiescent states depending on its history, like
flip-flops
Flip-flops are a type of light sandal, typically worn as a form of casual footwear. They consist of a flat sole held loosely on the foot by a Y-shaped strap known as a toe thong that passes between the first and second toes and around both side ...
, and magnetic material with
hysteresis
Hysteresis is the dependence of the state of a system on its history. For example, a magnet may have more than one possible magnetic moment in a given magnetic field, depending on how the field changed in the past. Plots of a single component of ...
. It can also adapt to different quiescent states. In complete darkness, the retinal cells become extremely sensitive, and there is noticeable "
visual snow
Visual snow syndrome (VSS) is an uncommon neurological condition in which the primary symptom is that affected individuals see persistent flickering white, black, transparent, or coloured dots across the whole visual field. Other common symptom ...
" caused by the retinal cells firing randomly without any light input. In brighter light, the retinal cells become a lot less sensitive, and consequently visual noise decreases.
Quiescent state is less well-defined when the sensory organ can be controlled by other systems, like a dog's ears that turn towards the front or the sides as the brain commands. Some spiders can use their nets as a large touch-organ, like weaving a skin for themselves. Even in the absence of anything falling on the net, hungry spiders may increase web thread tension, so as to respond promptly even to usually less noticeable, and less profitable prey, such as small fruit flies, creating two different "quiescent states" for the net.
Things become completely ill-defined for a system which connects its output to its own input, thus ever-moving without any external input. The prime example is the brain, with its
default mode network
In neuroscience, the default mode network (DMN), also known as the default network, default state network, or anatomically the medial frontoparietal network (M-FPN), is a large-scale brain network primarily composed of the dorsal medial prefro ...
.
Senses and receptors
While debate exists among neurologists as to the specific number of senses due to differing definitions of what constitutes a
sense
A sense is a biological system used by an organism for sensation, the process of gathering information about the world through the detection of Stimulus (physiology), stimuli. (For example, in the human body, the brain which is part of the cen ...
,
Gautama Buddha
Siddhartha Gautama, most commonly referred to as the Buddha, was a wandering ascetic and religious teacher who lived in South Asia during the 6th or 5th century BCE and founded Buddhism.
According to Buddhist tradition, he was born in Lu ...
and
Aristotle
Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of phil ...
classified five ‘traditional’ human senses which have become universally accepted:
touch
In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch (haptic perception), as well as temperature (thermoception), body position (proprioception), and pain. It is ...
,
taste
The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
,
smell
Smell may refer to;
* Odor, airborne molecules perceived as a scent or aroma
* Sense of smell, the scent also known scientifically as olfaction
* "Smells" (''Bottom''), an episode of ''Bottom''
* The Smell, a music venue in Los Angeles, Californ ...
,
sight
Visual perception is the ability to interpret the surrounding Biophysical environment, environment through photopic vision (daytime vision), color vision, scotopic vision (night vision), and mesopic vision (twilight vision), using light in the ...
, and
hearing
Hearing, or auditory perception, is the ability to perceive sounds
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 psycholog ...
. Other senses that have been well-accepted in most mammals, including humans, include
nociception
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, co ...
,
equilibrioception
The sense of balance or equilibrioception is the perception of balance and spatial orientation. It helps prevent humans and nonhuman animals from falling over when standing or moving. Equilibrioception is the result of a number of sensory systems ...
,
kinaesthesia
Proprioception ( ), also referred to as kinaesthesia (or kinesthesia), is the sense of self-movement, force, and body position. It is sometimes described as the "sixth sense".
Proprioception is mediated by proprioceptors, mechanosensory neurons ...
, and
thermoception
Thermoception or thermoreception is the sensation and perception of temperature, or more accurately, temperature differences inferred from heat flux. It deals with a series of events and processes required for an organism to receive a temperature s ...
. Furthermore, some nonhuman animals have been shown to possess alternate senses, including
magnetoreception
Magnetoreception is a sense which allows an organism to detect the Earth's magnetic field. Animals with this sense include some arthropods, molluscs, and vertebrates (fish, amphibians, reptiles, birds, and mammals, though not humans). The se ...
and
electroreception
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 ...
.
Receptors
The initialization of sensation stems from the response of a specific receptor to a physical stimulus. The receptors which react to the stimulus and initiate the process of sensation are commonly characterized in four distinct categories:
chemoreceptors
A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...
,
photoreceptors,
mechanoreceptors
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, a ...
, and
thermoreceptors
A thermoreceptor is a non-specialised sense 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 s ...
. All receptors receive distinct physical stimuli and transduce the signal into an electrical
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, ...
. This action potential then travels along
afferent neurons
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 potentials. This process is called sensory transduction. The cel ...
to specific brain regions where it is processed and interpreted.
Chemoreceptors
Chemoreceptors, or chemosensors, detect certain chemical stimuli and transduce that signal into an electrical action potential. The two primary types of chemoreceptors are:
* Distance chemoreceptors are integral to receiving stimuli in
gases
Gas is one of the four fundamental states of matter (the others being solid, liquid, and plasma).
A pure gas may be made up of individual atoms (e.g. a noble gas like neon), elemental molecules made from one type of atom (e.g. oxygen), or ...
in the
olfactory system
The olfactory system, or sense of smell, is the sensory system used for smelling (olfaction). Olfaction is one of the special senses, that have directly associated specific organs. Most mammals and reptiles have a main olfactory system and an ac ...
through both olfactory receptor neurons and neurons in the
vomeronasal organ
The vomeronasal organ (VNO), or Jacobson's organ, is the paired auxiliary olfactory (smell) sense organ located in the soft tissue of the nasal septum, in the nasal cavity just above the roof of the mouth (the hard palate) in various tetrapods. ...
.
* Direct chemoreceptors that detect stimuli in
liquids
A liquid is a nearly incompressible fluid that conforms to the shape of its container but retains a (nearly) constant volume independent of pressure. As such, it is one of the four fundamental states of matter (the others being solid, gas, an ...
include the
taste buds
Taste buds contain the taste receptor cells, which are also known as gustatory cells. The taste receptors are located around the small structures known as papillae found on the upper surface of the tongue, soft palate, upper esophagus, the che ...
in the
gustatory system
The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
as well as receptors in 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 ...
which detect changes in
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 as wel ...
concentration.
Photoreceptors
Photoreceptors are capable of
phototransduction Visual phototransduction is the transduction (physiology), sensory transduction process of the visual system by which light is detected to yield Action potential, nerve impulses in the rod cells and cone cells in the retina of the eye in humans and ...
, a process which converts light (
electromagnetic radiation
In physics, electromagnetic radiation (EMR) consists of waves of the electromagnetic field, electromagnetic (EM) field, which propagate through space and carry momentum and electromagnetic radiant energy. It includes radio waves, microwaves, inf ...
) into, among other types of
energy
In physics, energy (from Ancient Greek: ἐνέργεια, ''enérgeia'', “activity”) is the quantitative property that is transferred to a body or to a physical system, recognizable in the performance of work and in the form of heat a ...
, a
membrane potential
Membrane potential (also transmembrane potential or membrane voltage) is the difference in electric potential between the interior and the exterior of a biological cell. That is, there is a difference in the energy required for electric charges ...
. The three primary types of photoreceptors are:
Cones
A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex.
A cone is formed by a set of line segments, half-lines, or lines conn ...
are photoreceptors which 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 which 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
Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite.
Nocturnal creatures generally have highly developed sens ...
. In humans rods outnumber cones by approximately 20:1, while in nocturnal animals, such as the
tawny owl
The tawny owl (''Strix aluco''), also called the brown owl, is commonly found in woodlands across Europe to western Siberia, and has seven recognized subspecies. It is a stocky, medium-sized owl, whose underparts are pale with dark streaks, an ...
, the ratio is closer to 1000:1.
Ganglion Cell
{{stack,
A ganglion cell is a cell found in a ganglion. Examples of ganglion cells include:
* Retinal ganglion cell (RGC) found in the ganglion cell layer of the retina
* Cells that reside in the adrenal medulla, where they are involved in the s ...
s reside in the
adrenal medulla
The adrenal medulla ( la, medulla glandulae suprarenalis) is part of the adrenal gland. It is located at the center of the gland, being surrounded by the adrenal cortex. It is the innermost part of the adrenal gland, consisting of chromaffin cel ...
and
retina
The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
where they are involved in the
sympathetic response
The sympathetic nervous system (SNS) is one of the three divisions of the autonomic nervous system, the others being the parasympathetic nervous system and the enteric nervous system. The enteric nervous system is sometimes considered part of the ...
. Of the ~1.3 million ganglion cells present in the retina, 1-2% are believed to be
photosensitive ganglia. These photosensitive ganglia play a role in conscious vision for some animals,
and are believed to do the same in humans.
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 and e ...
or
distortion
In signal processing, distortion is the alteration of the original shape (or other characteristic) of a signal. In communications and electronics it means the alteration of the waveform of an information-bearing signal, such as an audio signal ...
. While mechanoreceptors are present in
hair cells
Hair cells are the sensory receptors of both the auditory system and the vestibular system in the ears of all vertebrates, and in the lateral line organ of fishes. Through mechanotransduction, hair cells detect movement in their environment.
...
and play an integral role in the
vestibular
The Vestibular (from pt, vestíbulo, "entrance hall") is a competitive examination and is the primary and widespread entrance system used by Brazilian universities to select the students admitted.
The Vestibular usually takes place from Novem ...
and
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 ...
s, the majority of mechanoreceptors are
cutaneous
Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation.
Other animal coverings, such as the arthropod exoskeleton, have different de ...
and are grouped into four categories:
* ''Slowly adapting type 1 receptors'' have small receptive fields and respond to static stimulation. These receptors are primarily used in the sensations of
form
Form is the shape, visual appearance, or configuration of an object. In a wider sense, the form is the way something happens.
Form also refers to:
*Form (document), a document (printed or electronic) with spaces in which to write or enter data
...
and
roughness.
* ''Slowly adapting type 2 receptors'' have large receptive fields and respond to stretch. Similarly to type 1, they produce sustained responses to a continued stimuli.
* ''Rapidly adapting receptors'' have small receptive fields and underlie the perception of slip.
* ''Pacinian receptors'' have large receptive fields and are the predominant receptors for high-frequency vibration.
Thermoreceptors
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 or ...
s have at least two distinct types of thermoreceptors:
[Krantz, John. ]
Experiencing Sensation and Perception
'. Pearson Education, Limited, 2009. p. 12.3
* The end-bulb of Krause, or
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 thermorecept ...
, detects temperatures above body temperature.
*
Ruffini's end organ detects temperatures below body temperature.
TRPV1 is a heat-activated channel that acts as a small heat detecting thermometer in the membrane which begins the polarization of the neural fiber when exposed to changes in temperature. Ultimately, this allows us to detect ambient temperature in the warm/hot range. Similarly, the molecular cousin to TRPV1, TRPM8, is a cold-activated ion channel that responds to cold. Both cold and hot receptors are segregated by distinct subpopulations of sensory nerve fibers, which shows us that the information coming into the spinal cord is originally separate. Each sensory receptor has its own “labeled line” to convey a simple sensation experienced by the recipient. Ultimately, TRP channels act as thermosensors, channels that help us to detect changes in ambient temperatures.
Nociceptors
Nociceptors respond to potentially damaging stimuli by sending signals to the spinal cord and brain. This process, called
nociception
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, co ...
, 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. Nociceptors detect different kinds of damaging stimuli or actual damage. Those that only respond when tissues are damaged are known as "sleeping" or "silent" nociceptors.
* Thermal nociceptors are activated by noxious heat or cold at various temperatures.
* Mechanical nociceptors respond to excess pressure or mechanical deformation.
* Chemical nociceptors respond to a wide variety of chemicals, some of which are signs of tissue damage. They are involved in the detection of some spices in food.
Sensory cortex
All
stimuli
A stimulus is something that causes a physiological response. It may refer to:
*Stimulation
**Stimulus (physiology), something external that influences an activity
**Stimulus (psychology), a concept in behaviorism and perception
*Stimulus (economi ...
received by the
receptors
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 ...
listed above are
transduced to 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, ...
, which is carried along one or more
afferent neurons
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 potentials. This process is called sensory transduction. The cel ...
towards a specific area of the brain. While the term
sensory cortex The sensory cortex can refer informally to the primary somatosensory cortex, or it can be used as a term for the primary and secondary cortices of the different senses (two cortices each, on left and right hemisphere): the visual cortex on the occi ...
is often used informally to refer to the
somatosensory cortex
In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch (haptic perception), as well as temperature (thermoception), body position (proprioception), and pain. It is ...
, the term more accurately refers to the multiple areas of the brain at which
sense
A sense is a biological system used by an organism for sensation, the process of gathering information about the world through the detection of Stimulus (physiology), stimuli. (For example, in the human body, the brain which is part of the cen ...
s are received to be processed. For the five
traditional senses in humans, this includes the primary and secondary
cortices of the different senses: the somatosensory cortex, the
visual cortex
The visual cortex of the brain is the area of the cerebral cortex that processes visual information. It is located in the occipital lobe. Sensory input originating from the eyes travels through the lateral geniculate nucleus in the thalamus and ...
, the
auditory cortex
The auditory cortex is the part of the temporal lobe that processes auditory information in humans and many other vertebrates. It is a part of the auditory system, performing basic and higher functions in hearing, such as possible relations to ...
, the
primary olfactory cortex
The primary olfactory cortex (POC) is a portion of the cerebral cortex. It is found in the inferior part of the temporal lobe of the brain. It receives input from the olfactory tract. It is involved in the sense of smell (olfaction).
Structure ...
, and the
gustatory cortex The primary gustatory cortex is a brain structure responsible for the perception of taste. It consists of two substructures: the anterior insula on the insular lobe and the frontal operculum on the inferior frontal gyrus of the frontal lobe. Be ...
.
[Brynie, F.H. (2009). Brain Sense: The Science of the Senses and How We Process the World Around Us. American Management Association.] Other
modalities have corresponding sensory cortex areas as well, including the
vestibular cortex
Vestibular cortex is the portion of the cerebrum which responds to input from the vestibular system.
The location is not well defined, but some research indicates a Lateralization of brain function, right hemisphere dominance.
Lesions of the vest ...
for the sense of balance.
Somatosensory cortex
Located in the
parietal lobe
The parietal lobe is one of the four major lobes of the cerebral cortex in the brain of mammals. The parietal lobe is positioned above the temporal lobe and behind the frontal lobe and central sulcus.
The parietal lobe integrates sensory informa ...
, the
primary somatosensory cortex
In neuroanatomy, the primary somatosensory cortex is located in the postcentral gyrus of the brain's parietal lobe, and is part of the somatosensory system. It was initially defined from surface stimulation studies of Wilder Penfield, and paralle ...
is the primary receptive area for the sense of touch and
proprioception
Proprioception ( ), also referred to as kinaesthesia (or kinesthesia), is the sense of self-movement, force, and body position. It is sometimes described as the "sixth sense".
Proprioception is mediated by proprioceptors, mechanosensory neurons ...
in the
somatosensory system
In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch (haptic perception), as well as temperature (thermoception), body position (proprioception), and pain. It ...
. This cortex is further divided into
Brodmann areas
A Brodmann area is a region of the cerebral cortex, in the human or other primate brain, defined by its cytoarchitecture, or histological structure and organization of cells.
History
Brodmann areas were originally defined and numbered by the ...
1, 2, and 3.
Brodmann area 3
In neuroanatomy, the postcentral gyrus is a prominent gyrus in the lateral parietal lobe of the human brain. It is the location of the primary somatosensory cortex, the main sensory receptive area for the sense of touch. Like other sensory areas, ...
is considered the primary processing center of the somatosensory cortex as it receives significantly more input from the
thalamus
The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all directions, ...
, has neurons highly responsive to somatosensory stimuli, and can evoke somatic sensations through electrical
stimulation
Stimulation is the encouragement of development or the cause of activity generally. For example, "The press provides stimulation of political discourse." An interesting or fun activity can be described as "stimulating", regardless of its physica ...
. Areas 1 and 2 receive most of their input from area 3. There are also pathways for
proprioception
Proprioception ( ), also referred to as kinaesthesia (or kinesthesia), is the sense of self-movement, force, and body position. It is sometimes described as the "sixth sense".
Proprioception is mediated by proprioceptors, mechanosensory neurons ...
(via the
cerebellum
The cerebellum (Latin for "little brain") is a major feature of the hindbrain of all vertebrates. Although usually smaller than the cerebrum, in some animals such as the mormyrid fishes it may be as large as or even larger. In humans, the cerebel ...
), and
motor
An engine or motor is a machine designed to convert one or more forms of energy into mechanical energy.
Available energy sources include potential energy (e.g. energy of the Earth's gravitational field as exploited in hydroelectric power gen ...
control (via
Brodmann area 4
Brodmann area 4 refers to the primary motor cortex of the human brain. It is located in the posterior portion of the frontal lobe.
Brodmann area 4 is part of the precentral gyrus. The borders of this area are: the precentral sulcus in front ( ...
). See also: S2
Secondary somatosensory cortex
The human secondary somatosensory cortex (S2, SII) is a region of cortex in the parietal operculum on the ceiling of the lateral sulcus.
Region S2 was first described by Adrian in 1940, who found that feeling in cats' feet was not only represente ...
.
Visual cortex
The visual cortex refers to the primary visual cortex, labeled V1 or
Brodmann area 17
The visual cortex of the brain is the area of the cerebral cortex that processes visual information. It is located in the occipital lobe. Sensory input originating from the eyes travels through the lateral geniculate nucleus in the thalamus and ...
, as well as the extrastriate visual cortical areas V2-V5. Located in the
occipital lobe
The occipital lobe is one of the four major lobes of the cerebral cortex in the brain of mammals. The name derives from its position at the back of the head, from the Latin ''ob'', "behind", and ''caput'', "head".
The occipital lobe is the vi ...
, V1 acts as the primary relay station for visual input, transmitting information to two primary pathways labeled the
dorsal
Dorsal (from Latin ''dorsum'' ‘back’) may refer to:
* Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism
* Dorsal, positioned on top of an aircraft's fuselage
* Dorsal co ...
and
ventral stream
The two-streams hypothesis is a model of the neural processing of vision as well as hearing. The hypothesis, given its initial characterisation in a paper by David Milner and Melvyn A. Goodale in 1992, argues that humans possess two distinct visual ...
s. The dorsal stream includes areas V2 and V5, and is used in interpreting visual 'where' and 'how.' The ventral stream includes areas V2 and V4, and is used in interpreting 'what.' Increases in
Task-negative
In neuroscience, the default mode network (DMN), also known as the default network, default state network, or anatomically the medial frontoparietal network (M-FPN), is a large-scale brain network primarily composed of the dorsal medial prefro ...
activity are observed in the ventral attention network, after abrupt changes in sensory stimuli, at the onset and offset of task blocks, and at the end of a completed trial.
Auditory cortex
Located in the
temporal lobe
The temporal lobe is one of the four Lobes of the brain, 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 ...
, the auditory cortex is the primary receptive area for sound information. The auditory cortex is composed of Brodmann areas 41 and 42, also known as the
anterior transverse temporal area 41
The auditory cortex is the part of the temporal lobe that processes auditory information in humans and many other vertebrates. It is a part of the auditory system, performing basic and higher functions in hearing, such as possible relations to ...
and the
posterior transverse temporal area 42
The auditory cortex is the part of the temporal lobe that processes auditory information in humans and many other vertebrates. It is a part of the auditory system, performing basic and higher functions in hearing, such as possible relations to ...
, respectively. Both areas act similarly and are integral in receiving and processing the signals transmitted from
auditory receptors.
Primary olfactory cortex
Located in the temporal lobe, the
primary olfactory cortex
The primary olfactory cortex (POC) is a portion of the cerebral cortex. It is found in the inferior part of the temporal lobe of the brain. It receives input from the olfactory tract. It is involved in the sense of smell (olfaction).
Structure ...
is the primary receptive area for
olfaction
The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste.
In humans, it ...
, or smell. Unique to the olfactory and gustatory systems, at least in
mammals
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 or ...
, is the implementation of both
peripheral
A peripheral or peripheral device is an auxiliary device used to put information into and get information out of a computer. The term ''peripheral device'' refers to all hardware components that are attached to a computer and are controlled by the ...
and
central
Central is an adjective usually referring to being in the center of some place or (mathematical) object.
Central may also refer to:
Directions and generalised locations
* Central Africa, a region in the centre of Africa continent, also known as ...
mechanisms of action. The peripheral mechanisms involve
olfactory receptor neurons
An olfactory receptor neuron (ORN), also called an olfactory sensory neuron (OSN), is a sensory neuron within the olfactory system.
Structure
Humans have between 10 and 20 million olfactory receptor neurons (ORNs). In vertebrates, ORNs are b ...
which
transduce a chemical signal along the
olfactory nerve
The olfactory nerve, also known as the first cranial nerve, cranial nerve I, or simply CN I, is a cranial nerve that contains sensory nerve fibers relating to the sense of smell.
The afferent nerve fibers of the olfactory receptor neurons tr ...
, which terminates in the
olfactory bulb
The olfactory bulb (Latin: ''bulbus olfactorius'') is a grey matter, neural structure of the vertebrate forebrain involved in olfaction, the sense of odor, smell. It sends olfactory information to be further processed in the amygdala, the orbitof ...
. The
chemoreceptors
A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...
in the receptor neurons that start the
signal cascade
Signal transduction is the process by which a chemical or physical signal is transmitted through a cell as a series of molecular events, most commonly protein phosphorylation catalyzed by protein kinases, which ultimately results in a cellular ...
are
G protein-coupled receptors
G protein-coupled receptors (GPCRs), also known as seven-(pass)-transmembrane domain receptors, 7TM receptors, heptahelical receptors, serpentine receptors, and G protein-linked receptors (GPLR), form a large group of evolutionarily-related p ...
. The central mechanisms include the convergence of olfactory nerve
axon
An axon (from Greek ἄξων ''áxōn'', axis), or nerve fiber (or nerve fibre: see spelling differences), is a long, slender projection of a nerve cell, or neuron, in vertebrates, that typically conducts electrical impulses known as action po ...
s into
glomeruli
''Glomerulus'' () is a common term used in anatomy to describe globular structures of entwined vessels, fibers, or neurons. ''Glomerulus'' is the diminutive of the Latin ''glomus'', meaning "ball of yarn".
''Glomerulus'' may refer to:
* the filter ...
in the olfactory bulb, where the signal is then transmitted to the
anterior olfactory nucleus
The anterior olfactory nucleus (AON; also called the anterior olfactory cortex) is a portion of the forebrain of vertebrates.
It is involved in olfaction
and has supposedly strong influence on other olfactory areas like the olfactory bulb and the ...
, the
piriform cortex
The piriform cortex, or pyriform cortex, is a region in the brain, part of the rhinencephalon situated in the cerebrum. The function of the piriform cortex relates to the sense of smell.
Structure
The piriform cortex is part of the rhinencephalo ...
, the medial
amygdala
The amygdala (; plural: amygdalae or amygdalas; also '; Latin from Greek, , ', 'almond', 'tonsil') is one of two almond-shaped clusters of nuclei located deep and medially within the temporal lobes of the brain's cerebrum in complex verteb ...
, and the
entorhinal cortex
The entorhinal cortex (EC) is an area of the brain's allocortex, located in the medial temporal lobe, whose functions include being a widespread network hub for memory, navigation, and the perception of time.Integrating time from experience in the ...
, all of which make up the primary olfactory cortex.
In contrast to vision and hearing, the
olfactory bulb
The olfactory bulb (Latin: ''bulbus olfactorius'') is a grey matter, neural structure of the vertebrate forebrain involved in olfaction, the sense of odor, smell. It sends olfactory information to be further processed in the amygdala, the orbitof ...
s are not cross-hemispheric; the right bulb connects to the right hemisphere and the left bulb connects to the left hemisphere.
Gustatory cortex
The
gustatory cortex The primary gustatory cortex is a brain structure responsible for the perception of taste. It consists of two substructures: the anterior insula on the insular lobe and the frontal operculum on the inferior frontal gyrus of the frontal lobe. Be ...
is the primary receptive area for
taste
The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
. The word ''taste'' is used in a technical sense to refer specifically to sensations coming from taste buds on the tongue. The five qualities of taste detected by the tongue include sourness, bitterness, sweetness, saltiness, and the protein taste quality, called
umami
Umami ( from ja, 旨味 ), or savoriness, is one of the five basic tastes. It has been described as savory and is characteristic of broths and cooked meats.
People taste umami through taste receptors that typically respond to glutamates and ...
. In contrast, the term ''flavor'' refers to the experience generated through integration of taste with smell and tactile information. The gustatory cortex consists of two primary structures: the
anterior insula
The insular cortex (also insula and insular lobe) is a portion of the cerebral cortex folded deep within the lateral sulcus (the fissure separating the temporal lobe from the parietal and frontal lobes) within each hemisphere of the mammalian br ...
, located on the
insular lobe
The insular cortex (also insula and insular lobe) is a portion of the cerebral cortex folded deep within the lateral sulcus (the fissure separating the temporal lobe from the parietal lobe, parietal and frontal lobes) within each brain hemisphere, ...
, and the frontal
operculum, located on the
frontal lobe
The frontal lobe is the largest of the four major lobes of the brain in mammals, and is located at the front of each cerebral hemisphere (in front of the parietal lobe and the temporal lobe). It is parted from the parietal lobe by a groove betwe ...
. Similarly to the olfactory cortex, the gustatory pathway operates through both peripheral and central mechanisms. Peripheral
taste receptor
A taste receptor or tastant is a type of cellular receptor which facilitates the sensation of taste. When food or other substances enter the mouth, molecules interact with saliva and are bound to taste receptors in the oral cavity and other locat ...
s, located on the
tongue
The tongue is a muscular organ (anatomy), organ in the mouth of a typical tetrapod. It manipulates food for mastication and swallowing as part of the digestive system, digestive process, and is the primary organ of taste. The tongue's upper surfa ...
,
soft palate
The soft palate (also known as the velum, palatal velum, or muscular palate) is, in mammals, the soft tissue constituting the back of the roof of the mouth. The soft palate is part of the palate of the mouth; the other part is the hard palate. ...
,
pharynx
The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its struc ...
, and
esophagus
The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
, transmit the received signal to primary sensory axons, where the signal is projected to the
nucleus of the solitary tract
In the human brainstem, the solitary nucleus, also called nucleus of the solitary tract, nucleus solitarius, and nucleus tractus solitarii, (SN or NTS) is a series of purely sensory
nuclei (clusters of nerve cell bodies) forming a vertical column ...
in the
medulla
Medulla or Medullary may refer to:
Science
* Medulla oblongata, a part of the brain stem
* Renal medulla, a part of the kidney
* Adrenal medulla, a part of the adrenal gland
* Medulla of ovary, a stroma in the center of the ovary
* Medulla of th ...
, or the gustatory nucleus of the solitary tract complex. The signal is then transmitted to the
thalamus
The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all directions, ...
, which in turn projects the signal to several regions of the
neocortex
The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, sp ...
, including the gustatory cortex.
The neural processing of taste is affected at nearly every stage of processing by concurrent somatosensory information from the tongue, that is,
mouthfeel
Mouthfeel refers to the physical sensations in the mouth caused by food or drink, making it distinct from taste. It is a fundamental sensory attribute which, along with taste and smell, determines the overall flavor of a food item. Mouthfeel is ...
. Scent, in contrast, is not combined with taste to create flavor until higher cortical processing regions, such as the insula and orbitofrontal cortex.
Human sensory system
The human sensory system consists of the following subsystems:
*
Visual system
The visual system comprises the sensory organ (the eye) and parts of the central nervous system (the retina containing photoreceptor cells, the optic nerve, the optic tract and the visual cortex) which gives organisms the sense of sight (the a ...
*
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 ...
*
Somatosensory system
In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch (haptic perception), as well as temperature (thermoception), body position (proprioception), and pain. It ...
consists of the receptors, transmitters (pathways) leading to S1, and S1 that experiences the sensations labelled as
touch
In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch (haptic perception), as well as temperature (thermoception), body position (proprioception), and pain. It is ...
,
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 and e ...
,
vibration, temperature (warm or cold),
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, ...
(including
itch
Itch (also known as pruritus) is a sensation that causes the desire or reflex to scratch. Itch has resisted many attempts to be classified as any one type of sensory experience. Itch has many similarities to pain, and while both are unpleasant ...
and tickle), and the sensations of muscle movement and joint position including
posture
Posture or posturing may refer to:
Medicine
* Human position
** Abnormal posturing, in neurotrauma
**Spinal posture
** List of human positions
* Posturography, in neurology
Other uses
* Posture (psychology)
In humans, posture can provide a s ...
,
movement
Movement may refer to:
Common uses
* Movement (clockwork), the internal mechanism of a timepiece
* Motion, commonly referred to as movement
Arts, entertainment, and media
Literature
* "Movement" (short story), a short story by Nancy Fu ...
, and facial expression (collectively also called proprioception)
*
Gustatory system
The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
*
Olfactory system
The olfactory system, or sense of smell, is the sensory system used for smelling (olfaction). Olfaction is one of the special senses, that have directly associated specific organs. Most mammals and reptiles have a main olfactory system and an ac ...
*
Vestibular system
The vestibular system, in vertebrates, is a sensory system that creates the sense of balance and spatial orientation for the purpose of coordinating movement with balance. Together with the cochlea, a part of the auditory system, it constitutes ...
*
Interoceptive system
Diseases
![Sense organ diseases world map - DALY - WHO2002](https://upload.wikimedia.org/wikipedia/commons/0/08/Sense_organ_diseases_world_map_-_DALY_-_WHO2002.svg)
*
Amblyopia
Amblyopia, also called lazy eye, is a disorder of sight in which the brain fails to fully process input from one eye and over time favors the other eye. It results in decreased vision in an eye that typically appears normal in other aspects. Amb ...
*
Anacusis
Hearing loss is a partial or total inability to hear. Hearing loss may be present at birth or acquired at any time afterwards. Hearing loss may occur in one or both ears. In children, hearing problems can affect the ability to acquire spoken la ...
*
Color blindness
Color blindness or color vision deficiency (CVD) is the decreased ability to color vision, see color or differences in color. It can impair tasks such as selecting ripe fruit, choosing clothing, and reading traffic lights. Color blindness may ...
*
Deafness
Deafness has varying definitions in cultural and medical contexts. In medical contexts, the meaning of deafness is hearing loss that precludes a person from understanding spoken language, an audiological condition. In this context it is written ...
See also
*
Multisensory integration
Multisensory integration, also known as multimodal integration, is the study of how information from the different sensory modalities (such as sight, sound, touch, smell, self-motion, and taste) may be integrated by the nervous system. A coherent r ...
*
Neural adaptation
Neural adaptation or sensory adaptation is a gradual decrease over time in the responsiveness of the sensory system to a constant stimulus. It is usually experienced as a change in the stimulus. For example, if a hand is rested on a table, the tabl ...
*
Neural coding
Neural coding (or Neural representation) is a neuroscience field concerned with characterising the hypothetical relationship between the stimulus and the individual or ensemble neuronal responses and the relationship among the electrical activity o ...
*
Sensor
A sensor is a device that produces an output signal for the purpose of sensing a physical phenomenon.
In the broadest definition, a sensor is a device, module, machine, or subsystem that detects events or changes in its environment and sends ...
*
Sensory augmentation
Sensory may refer to:
Biology
* Sensory ecology, how organisms obtain information about their environment
* Sensory neuron, nerve cell responsible for transmitting information about external stimuli
* Sensory perception, the process of acquiri ...
*
Sensory neuroscience
Sensory neuroscience is a subfield of neuroscience which explores the anatomy and physiology of neurons that are part of sensory systems such as vision, hearing, and olfaction. Neurons in sensory regions of the brain respond to stimuli by firing o ...
*
Sensory systems in fish
Most fish possess highly developed sense organs. Nearly all daylight fish have color vision that is at least as good as a human's (see vision in fishes). Many fish also have chemoreceptors that are responsible for extraordinary senses of taste an ...
References
External links
{{DEFAULTSORT:Sensory System
Nervous system
Sensory systems
Sensory organs