Sense

This article is about the empirical or physical senses of living organisms (vision, taste, etc.). For other uses, see Sense (disambiguation).

Senses are the physiological capacities within organisms that provide inputs for 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.

Human beings have a multitude of senses. In addition to the traditional five senses of sight (ophthalmoception), hearing (audioception), taste (gustaoception), smell (olfacoception or olfacception), and touch (tactioception); other senses include temperature (thermoception), kinesthetic sense (proprioception), pain (nociception), direction (magetoception), balance (equilibrioception) and acceleration (kinesthesioception). What constitutes a sense is a matter of some debate, leading to difficulties in defining what exactly a sense is.

Non-human animals also have receptors to sense the world around them, with degrees of capability varying greatly between species. Humans have a comparatively weak sense of smell, whilst other species may lack one of more of the traditional five senses. Other species may also intake and interpret senses in very different ways. Some species of animals are able to sense the world in a way that humans cannot, with some species able to sense electrical and magnetic fields, and detect water pressure and currents.

Definition

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 human olfactory system contains about 10,000,000 olfactory receptors. The traditional five senses are sight, hearing, touch, smell and taste, a classification attributed to Aristotle. Humans are considered to have at least five additional senses that include: nociception (pain); equilibrioception (balance); proprioception and kinaesthesia (joint motion and acceleration); sense of time; thermoception (temperature differences); and possibly an additional weak magnetoception (direction), and six more if interoceptive senses (see other internal senses below) are also considered.

One commonly recognized categorisation for human senses is as follows: chemoreception; photoreception; mechanoreception; and thermoception. This categorisation has been criticized as too restrictive, however, as it does not include categories for accepted senses such as the sense of time and sense of pain.

Non-human animals may possess senses that are absent in humans, such as electroreception and detection of polarized light.

A broadly acceptable definition of a sense would be "A system that consists of a group of 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 typically arise around 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 eye(s) to focus and detect images of visible light on the retina in each eye, and determine varying colors, hues, and brightness detected by each retinal receptor. There is some 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 color and brightness. Some argue that stereopsis, the perception of depth using both eyes, also constitutes a sense, but it is generally regarded as a cognitive (that is, post-sensory) function of the visual cortex of the brain where patterns and objects in images are recognized and interpreted based on previously learned information. The inability to see is called blindness.

Blindness may result from damage to the eyeball, especially to the retina, damage to the optic nerve that connects each eye to the brain, and/or from stroke (infarcts in the brain). Temporary or permanent blindness can be caused by poisons or medications.

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 because these vibrations are mechanically conducted from the eardrum through a series of tiny bones to hair-like fibers in the inner ear which detect mechanical motion of the fibers within a range of about 20 to 20,000 hertz, with substantial variation between individuals. Hearing at high frequencies declines with age. Sound can also be detected as vibrations conducted through the body by tactition. Lower frequencies than that can be heard are detected this way. 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, salty, 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 neurotransmitter commonly found in meat and in artificial flavourings containing monosodium glutamate. Another taste sense for ginger, mild chilli, and olive oil is perceived as a peppery tickle at the back of the throat (which senses polyphenols in unprocessed olive oil).

The University of Maryland School of Medicine announced in 2010 that bitter taste receptors had been discovered on smooth muscle in human lung bronchi. These smooth muscles control airway contraction and dilation - contrary to expectation, bitter substances such as quinine or chloroquine opened contracted airways, offering new insight into asthma.

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. Odor 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. Some neurons in the nose are specialized to detect pheromones.

Touch

Touch, also called tactition or mechanoreception, is a perception resulting from activation of neural receptors, generally in the skin including hair follicles, but also in the tongue, throat, and mucosa. A variety of pressure receptors respond to variations in pressure (firm, brushing, sustained, etc.). The touch sense of itching caused by insect bites or allergies involves special itch-specific neurons in the skin and spinal cord. The loss or impairment of the ability to feel anything touched is called tactile anesthesia. Paresthesia is a sensation of tingling, pricking, or numbness of the skin that may result from nerve damage and may be permanent or temporary.

Balance and acceleration

Main article: Vestibular system

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 of angular momentum acceleration and linear acceleration (which also senses gravity), but they are known together as equilibrioception.

The vestibular nerve conducts information from sensory receptors in three ampulla that sense motion of fluid in three semicircular canals caused by three-dimensional rotation of the head. The vestibular nerve also conducts information from the utricle and the saccule which contain hair-like sensory receptors that bend under the weight of otoliths (which are small crystals of calcium carbonate) that provide the inertia needed to detect head rotation, linear acceleration, and the direction of gravitational force.

Temperature

Thermoception is the sense of heat and the absence of heat (cold) by the skin and including internal skin passages, or rather, the heat flux (the rate of heat flow) in these areas. There are specialized receptors for cold (declining temperature) and to heat. The cold receptors play an important part in the dog's sense of smell, telling wind direction. The heat receptors are sensitive to infrared radiation and can occur in specialized organs for instance in pit vipers. 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 their own nose with the tip of a finger. 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 nerve-damage or damage to tissue. The three types of pain receptors are cutaneous (skin), somatic (joints and bones) and visceral (body organs). It was previously 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. The main function of pain is to warn us about dangers. For example, humans avoid touching a sharp needle or hot object or extending an arm beyond a safe limit because it hurts, and thus is dangerous. Without pain, people could do many dangerous things without realizing it.

Direction

Magnetoception (or magnetoreception) is the ability to detect the direction one is facing based on the Earth's magnetic field. Directional awareness is most commonly observed in birds. 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.

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.
• The chemoreceptor trigger zone is an area of the medulla in the brain that receives inputs from blood-borne drugs or hormones, and communicates with the vomiting center.
• 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 gag 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.

Echolocation

Main article: Animal echolocation

Certain animals, including bats and cetaceans, have the ability to determine orientation to other objects through interpretation of reflected sound (like sonar). They most often use this 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.

Smell

Most non-human mammals have a much keener sense of smell than humans, although the mechanism is similar. Sharks combine their keen sense of smell with timing to determine the direction of a smell. They follow the nostril that first detected the smell. Insects have olfactory receptors on their antennae.

Vomeronasal organ

Many animals (salamanders, reptiles, mammals) have a vomeronasal organ that is connected with the mouth cavity. In mammals it is mainly used to detect pheromones to mark their territory, trails, and sexual state. Reptiles like snakes and monitor lizards make extensive use of it as a smelling organ by transferring scent molecules to the vomeronasal organ with the tips of the forked tongue. In mammals it is often associated with a special behavior called flehmen characterized by uplifting of the lips. The organ is vestigial in humans, because associated neurons have not been found that give any sensory input in humans.

Taste

Flies and butterflies have taste organs on their feet, allowing them to taste anything they land on. Catfish have taste organs across their entire bodies, and can taste anything they touch, including chemicals in the water.

Vision

Cats have the ability to see in low light 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 and dragonflies are also able to see in the ultraviolet.

Balance

Ctenophora 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:

• Electroreception (or electroception) 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.

• Magnetoreception (or magnetoception) is the ability to detect a magnetic field to perceive direction, altitude or location. This sense plays a role in the navigational abilities of several animal species and has been postulated as a method for animals to develop regional maps.

• Pressure detection uses the organ of Weber, a system consisting of three appendages of vertebrae transferring changes in shape of the gas bladder to the middle ear. It can be used to regulate the buoyancy of the fish. Fish like the weather fish and other loaches are also known to respond to low pressure areas but they lack a swim bladder.

• Current detection The lateral line in fish and aquatic forms of amphibians is a detection system of water currents, mostly consisting of vortices. The lateral line is also sensitive to low frequency vibrations. The mechanoreceptors are hair cells, the same mechanoreceptors for vestibular sense and hearing. It is used primarily for navigation, hunting, and schooling. The receptors of the electrical sense are modified hair cells of the lateral line system.

• Polarized light direction/detection is used by bees to orient themselves, especially on cloudy days. Cuttlefish can also perceive the polarization of light. Most sighted humans can in fact learn to roughly detect large areas of polarization by an effect called Haidinger's brush, however this is considered an entoptic phenomenon rather than a separate sense.

• Slit sensillae of spiders detect mechanical strain in the exoskeleton, providing information on force and vibrations.

Plant senses

Some plants have sensory organs, for example the Venus fly trap, that respond to vibration, light, water, scents, or other specific chemicals. Some plants sense the location of other plants and attack and eat part of them.

Culture

Further information: Ṣaḍāyatana, Ayatana, and Indriya

The traditional five senses are enumerated as the "five material faculties" (pañcannaṃ indriyānaṃ avakanti) in Buddhist literature. They appear in allegorical representation as early as in the Katha Upanishad (roughly 6th century BC), as five horses drawing the "chariot" of the body, guided by the mind as "chariot driver".

Depictions of the five traditional senses as allegory became a popular subject for seventeenth-century artists, especially among Dutch and Flemish Baroque painters. A typical example is Gérard de Lairesse's Allegory of the Five Senses (1668), in which each of the figures in the main group allude to a sense: sight is the reclining boy with a convex mirror, hearing is the cupid-like boy with a triangle, smell is represented by the girl with flowers, taste by the woman with the fruit and touch by the woman holding the bird.

See also

• Attention
• Apperception
• Āyatana (Sense Bases in Theravada Buddhism)
• Basic tastes
• Communication
• Empiricism
• Extrasensory perception
• Hypersensitivity (people with unusual sense abilities)
  • Human echolocation
  • Supertaster
  • Vision-related:
    • Haidinger's brush (ordinary people sensing light polarisation)
    • Tetrachromat (increased colour perception)
• Illusions
  • Auditory illusion
  • Optical illusion
  • Touch illusion
• Intuition
• Multimodal integration
• Perception
• Phantom limb
• Saḷāyatana and Ayatana (the six senses as a concept in Buddhism)
• Sensation and perception psychology
• Sense of time
• Sensitivity (human)
• Sensorium
• Sensory system
• Synesthesia

Research centers

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

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.
• HHMI - World renowned centre of medical excellence
• 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.