sense, faculty by which external or internal stimuli are conveyed to the brain centers, where they are registered as sensations. Sensory reception occurs in higher animals through a process known as transduction, in which stimuli are converted into nerve impulses and relayed to the brain. The four commonly known special senses (sight, hearing, smell, and taste) are concerned with the outer world, and external stimuli are received and conducted by sensory receptors concentrated in the eye, ear, olfactory organ, and the taste buds. The so-called somatic senses respond to both external and internal stimuli. Although most of the somatic receptors are located in the skin (conveying the external sensations of touch, heat, cold, pressure, and pain), others are located in internal organs (e.g., the heart and the stomach). Somatic sensations such as hunger, thirst, and fatigue are thought to originate in specific areas of the nervous system. The sense of balance, or equilibrium, is related to the flow of endolymph, a fluid found in the inner ear.

humour(Latin; “fluid”)

In early Western physiological theory, one of the four body fluids thought to determine a person's temperament and features. As hypothesized by Galen, the four cardinal humours were blood, phlegm, choler (yellow bile), and melancholy (black bile). The variant mixture of these humours in each person determined his “complexion” or temperament and his mental and physical qualities. The ideal person had the perfectly proportioned mixture of the four fluids; a disproportionate amount of one humour created a personality dominated by one set of related emotions (e.g., a choleric man was easily angered, proud, ambitious, and vengeful).

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Humour marked by the use of morbid, ironic, or grotesquely comic episodes that ridicule human folly. The term came into common use in the 1960s to describe the work of novelists such as Joseph Heller, whose Catch-22 (1961) is an outstanding example; Kurt Vonnegut, particularly in Slaughterhouse Five (1969); and Thomas Pynchon, in V (1963) and Gravity's Rainbow (1973). A film exemplar is Stanley Kubrick's Dr. Strangelove (1963). The term black comedy has been applied to some playwrights in the Theatre of the Absurd, especially Eugène Ionesco.

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or sensory reception or sense perception

Mechanism by which information is received about one's external or internal environment. Stimuli received by nerves, in some cases through specialized organs with receptor cells sensitive to one type of stimulus, are converted into impulses that travel to specialized areas of the brain, where they are analyzed. In addition to the “five senses”—sight, hearing, smell, taste, and touch—humans have senses of motion (kinesthetic sense), heat, cold, pressure, pain, and balance. Temperature, pressure, and pain are cutaneous (skin) senses; different points on the skin are particularly sensitive to each. Seealso chemoreception, ear, eye, inner ear, mechanoreception, nose, photoreception, proprioception, taste, thermoreception, tongue.

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Senses are the physiological methods of perception. The senses and their operation, classification, and theory are overlapping topics studied by a variety of fields, most notably neuroscience, cognitive psychology (or cognitive science), and philosophy of perception. The nervous system has a specific sensory system, or organ, dedicated to each sense.

Definition of sense

There is no firm agreement among neurologists as to the number of senses because of differing definitions of what constitutes a sense. One definition states that an exteroceptive sense is a faculty by which outside stimuli are perceived. The traditional five senses are sight, hearing, touch, smell, taste: a classification attributed to Aristotle. Humans also have at least six additional senses (a total of eleven including interoceptive senses) that include: nociception (pain), equilibrioception (balance), proprioception & kinesthesia (joint motion and acceleration), sense of time, thermoception (temperature differences), and in some a weak magnetoception (direction).

One commonly recognized catagorisation for human senses is as follows: chemoreception; photoreception; mechanoreception; and thermoception. Indeed, all human senses fit into one of these four categories.

Different senses also exist in other organisms, for example electroreception.

A broadly acceptable definition of a sense would be "a system that consists of a group sensory cell types that responds to a specific physical phenomenon, and that corresponds to a particular group of regions within the brain where the signals are received and interpreted." Disputes about the number of senses arise typically regarding the classification of the various cell types and their mapping to regions of the brain.

Senses

Sight

Sight or vision is the ability of the brain and eye to detect electromagnetic waves within the visible range (light) interpreting the image as "sight." There is disagreement as to whether this constitutes one, two or three senses. Neuroanatomists generally regard it as two senses, given that different receptors are responsible for the perception of colour (the frequency of photons of light) and brightness (amplitude/intensity - number of photons of light). Some argue that stereopsis, the perception of depth, also constitutes a sense, but it is generally regarded as a cognitive (that is, post-sensory) function of brain to interpret sensory input and to derive new information. The inability to see is called blindness.

Hearing

Hearing or audition is the sense of sound perception. Since sound is vibrations propagating through a medium such as air, the detection of these vibrations, that is the sense of the hearing, is a mechanical sense akin to a sense of touch, albeit a very specialized one. In humans, this perception is executed by tiny hair fibres in the inner ear which detect the motion of a membrane which vibrates in response to changes in the pressure exerted by atmospheric particles within a range of 20 to 22000 Hz, with substantial variation between individuals. Sound can also be detected as vibrations conducted through the body by tactition. Lower and higher frequencies than that can be heard are detected this way only. The inability to hear is called deafness.

Taste

Taste or gustation is one of the two main "chemical" senses. There are at least four types of tastes that "buds" (receptors) on the tongue detect, and hence there are anatomists who argue that these constitute five or more different senses, given that each receptor conveys information to a slightly different region of the brain. The inability to taste is called ageusia.

The four well-known receptors detect sweet, salt, sour, and bitter, although the receptors for sweet and bitter have not been conclusively identified. A fifth receptor, for a sensation called umami, was first theorised in 1908 and its existence confirmed in 2000. The umami receptor detects the amino acid glutamate, a flavour commonly found in meat and in artificial flavourings such as monosodium glutamate.

Note that taste is not the same as flavour; flavour includes the smell of a food as well as its taste.

Smell

Smell or olfaction is the other "chemical" sense. Unlike taste, there are hundreds of olfactory receptors, each binding to a particular molecular feature. Odour molecules possess a variety of features and thus excite specific receptors more or less strongly. This combination of excitatory signals from different receptors makes up what we perceive as the molecule's smell. In the brain, olfaction is processed by the olfactory system. Olfactory receptor neurons in the nose differ from most other neurons in that they die and regenerate on a regular basis. The inability to smell is called anosmia.

Touch

Touch, also called tactition, mechanoreception or somatic sensation, is the sense of pressure perception, generally in the skin. There are a variety of nerve endings that respond to variations in pressure (e.g., firm, brushing, and sustained). The inability to feel anything or almost anything is called anesthesia. Paresthesia is a sensation of tingling, pricking, or numbness of a person's skin with no apparent long term physical effect.

Balance and acceleration

Balance, Equilibrioception, or vestibular sense, is the sense which allows an organism to sense body movement, direction, and acceleration, and to attain and maintain postural equilibrium and balance. The organ of equilibrioception is the vestibular labyrinthine system found in both of the inner ears. Technically this organ is responsible for two senses, angular momentum and linear acceleration (which also senses gravity), but they are known together as equilibrioception.

The vestibular nerve conducts information from the three semicircular canals, corresponding to the three spatial planes, the utricle, and the saccule. The ampulla, or base, portion of the three semicircular canals each contain a structure called a crista. These bend in response to angular momentum or spinning. The saccule and utricle, also called the "otolith organs", sense linear acceleration and thus gravity. Otoliths are small crystals of calcium carbonate that provide the inertia needed to detect changes in acceleration or gravity.

Temperature

Thermoception is the sense of heat and the absence of heat (cold) by the skin and including internal skin passages. The thermoceptors in the skin are quite different from the homeostatic thermoceptors in the brain (hypothalamus) which provide feedback on internal body temperature.

Kinesthetic sense

Proprioception, the kinesthetic sense, provides the parietal cortex of the brain with information on the relative positions of the parts of the body. Neurologists test this sense by telling patients to close their eyes and touch the tip of a finger to their nose. Assuming proper proprioceptive function, at no time will the person lose awareness of where the hand actually is, even though it is not being detected by any of the other senses. Proprioception and touch are related in subtle ways, and their impairment results in surprising and deep deficits in perception and action.

Pain

Nociception (physiological pain) signals near-damage or damage to tissue. The three types of pain receptors are cutaneous (skin), somatic (joints and bones) and visceral (body organs). It was believed that pain was simply the overloading of pressure receptors, but research in the first half of the 20th century indicated that pain is a distinct phenomenon that intertwines with all of the other senses, including touch. Pain was once considered an entirely subjective experience, but recent studies show that pain is registered in the anterior cingulate gyrus of the brain.

Other internal senses

An internal sense or interoception is "any sense that is normally stimulated from within the body. These involve numerous sensory receptors in internal organs, such as stretch receptors that are neurologically linked to the brain.

• Pulmonary stretch receptors are found in the lungs and control the respiratory rate.
• Cutaneous receptors in the skin not only respond to touch, pressure, and temperature, but also respond to vasodilation in the skin such as blushing.
• Stretch receptors in the gastrointestinal tract sense gas distension that may result in colic pain.
• Stimulation of sensory receptors in the esophagus result in sensations felt in the throat when swallowing, vomiting, or during acid reflux.
• Sensory receptors in pharynx mucosa, similar to touch receptors in the skin, sense foreign objects such as food that may result in a gagging reflex and corresponding gagging sensation.
• Stimulation of sensory receptors in the urinary bladder and rectum may result in sensations of fullness.
• Stimulation of stretch sensors that sense dilation of various blood vessels may result in pain, for example headache caused by vasodilation of brain arteries.

Non-human senses

Analogous to human senses

Other living organisms have receptors to sense the world around them, including many of the senses listed above for humans. However, the mechanisms and capabilities vary widely.

Smell

Among non-human species, dogs have a much keener sense of smell than humans, although the mechanism is similar. Insects have olfactory receptors on their antennae.

Vision

Cats have the ability to see in the dark due to muscles surrounding their irises to contract and expand pupils as well as the tapetum lucidum, a reflective membrane that optimizes the image. Pitvipers, pythons and some boas have organs that allow them to detect infrared light, such that these snakes are able to sense the body heat of their prey. The common vampire bat may also have an infrared sensor on its nose. It has been found that birds and some other animals are tetrachromats and have the ability to see in the ultraviolet down to 300 nanometers. Bees are also able to see in the ultraviolet.

Balance

Ctenophores have a balance receptor (a statocyst) that works very differently from the mammalian's semi-circular canals.

Not analogous to human senses

In addition, some animals have senses that humans do not, including the following:

• Electroception (or "electroreception"), the most significant of the non-human senses, is the ability to detect electric fields. Several species of fish, sharks and rays have the capacity to sense changes in electric fields in their immediate vicinity. Some fish passively sense changing nearby electric fields; some generate their own weak electric fields, and sense the pattern of field potentials over their body surface; and some use these electric field generating and sensing capacities for social communication. The mechanisms by which electroceptive fish construct a spatial representation from very small differences in field potentials involve comparisons of spike latencies from different parts of the fish's body.

The only order of mammals that is known to demonstrate electroception is the monotreme order. Among these mammals, the platypus has the most acute sense of electroception.

Body modification enthusiasts have experimented with magnetic implants to attempt to replicate this sense, however in general humans (and probably other mammals) can detect electric fields only indirectly by detecting the effect they have on hairs. An electrically charged balloon, for instance, will exert a force on human arm hairs, which can be felt through tactition and identified as coming from a static charge (and not from wind or the like). This is however not electroception as it is a post-sensory cognitive action.

• Echolocation is the ability to determine orientation to other objects through interpretation of reflected sound (like sonar). Bats and cetaceans are noted for this ability, though some other animals use it, as well. It is most often used to navigate through poor lighting conditions or to identify and track prey. There is currently an uncertainty whether this is simply an extremely developed post-sensory interpretation of auditory perceptions or it actually constitutes a separate sense. Resolution of the issue will require brain scans of animals while they actually perform echolocation, a task that has proven difficult in practice. Blind people report they are able to navigate by interpreting reflected sounds (esp. their own footsteps), a phenomenon which is known as Human echolocation.
• Magnetoception (or "magnetoreception") is the ability to detect fluctuations in magnetic fields and is most commonly observed in birds, though it has also been observed in insects such as bees. Although there is no dispute that this sense exists in many avians (it is essential to the navigational abilities of migratory birds), it is not a well-understood phenomenon. One study has found that cattle make use of magnetoception, as they tend to align themselves in a North-South direction. Magnetotactic bacteria build miniature magnets inside themselves and use them to determine their orientation relative to the Earth's magnetic field.
• Pressure detection uses the lateral line, which is a pressure-sensing system of hairs found in fish and some aquatic amphibians. It is used primarily for navigation, hunting, and schooling. Humans have a basic relative-pressure detection ability when eustachian tube(s) are blocked, as demonstrated in the ear's response to changes in altitude.
• Polarized light direction / detection is used by bees to orient themselves, especially on cloudy days. Cuttlefish can also perceive the polarization of light.

See also

• Attention
• Auditory illusion, Optical illusion, Touch illusion
• Basic tastes
• Communication
• Empiricism
• Intuition
• Sensation
• Multimodal integration
• Sensitivity (human)
• Sense of time
• Sensorium
• Synesthesia

Research Centers

• Howard Hughes Medical Institute (HHMI)
• Institute for Advanced Science & Engineering (IASE)

References

External links

• The 2004 Nobel Prize in Physiology or Medicine (announced 4 October 2004) was won by Richard Axel and Linda Buck for their work explaining olfaction, published first in a joint paper in 1991 that described the very large family of about one thousand genes for odorant receptors and how the receptors link to the brain.
• [[Howard Hughes Medical Institute|HHMI] - World renowned centre of medical excellence]
• [[Institute for Advanced Science & Engineering|IASE] - senses.info] - A research center that focuses on the basis of senses.
• Answers to several questions related to senses and human feeling from curious kids
• The Physiology of the Senses tutorial 12 animated chapters on vision, hearing, touch, balance and memory.