Taste



Taste (also called smatch or gustation; adjectival form: gustatory) is one of the traditional five senses. It refers to the ability to detect the flavor of substances such as food, certain minerals, and poisons, etc.

Humans receive tastes through sensory organs called taste buds, or gustatory calyculi, concentrated on the upper surface of the tongue.

The sensation of taste can be categorized into five basic tastes: sweetness, bitterness, sourness, saltiness, and umami. The recognition and awareness of umami is a relatively recent development in Western cuisine. MSG produces a strong umami taste.

As taste senses are both harmful and beneficial things, all basic tastes are classified as either appetitive or aversive, depending upon the effect the things they sense have on our bodies.

The basic tastes that are here to concontribute only partially to the sensation and flavor of food in the mouth — other factors include smell, detected by the olfactory epithelium of the nose; texture, detected through a variety of mechanoreceptors, muscle nerves, etc.;, temperature, detected by thermoreceptors, and spiciness or piquance also called Chemesthesis.

Introduction
As one of the senses, taste is an essential part of daily life.

History
In The West, Aristotle, who postulated c. 350 BCE that the two most basic tastes were sweet and bitter, was one of the first to develop a list of basic tastes.

Ayurveda, an ancient Indian healing science, has its own tradition of basic tastes, including: astringent, bitter, pungent, salty, sour, and sweet.

Recent discoveries
The receptors for all known basic tastes have been identified. Sour and salty are detected with ion channels, whereas the receptors for sweet, bitter, and umami are G protein-coupled receptors.

There is some evidence for a sixth taste that senses fatty substances.

Taste-map myth
Despite a common misconception that different sections of the tongue specialized in different tastes, all taste sensations come from all regions of the tongue.

Basic tastes
For a long period, it was commonly accepted that there is a finite and small number of "basic tastes" of which all seemingly complex tastes are ultimately composed. Just as with primary colors, the "basic" quality of those sensations derives chiefly from the nature of human perception, in this case the different sorts of tastes the human tongue can identify. Until the twenty-first century, the number of "basic" tastes was considered to be four (bitterness, saltiness, sourness, and sweetness). More recently, a fifth taste, "savory" or "umami", has been proposed by a large number of authorities associated with this field. In Asian countries within the sphere of mainly Chinese, Indian and Japanese cultural influence, Piquance has traditionally been considered a sixth basic taste.

Bitterness
Bitterness is the most sensitive of the tastes, and is perceived by many to be unpleasant, sharp, or disagreeable. Common bitter foods and beverages include coffee, unsweetened cocoa, South American mate, marmalade, bitter melon, beer, bitters, olives, citrus peel, many plants in the Brassicaceae family, dandelion greens, wild chicory, escarole and lemons. Quinine is also known for its bitter taste and is found in tonic water.

Bitterness is of interest to those who study evolution, as well as various health researchers since a large number of natural bitter compounds are known to be toxic. The ability to detect bitter-tasting, toxic compounds at low thresholds is considered to provide an important protective function. Plant leaves often contain toxic compounds, yet even amongst leaf-eating primates, there is a tendency to prefer immature leaves, which tend to be higher in protein and lower in fiber and poisons than mature leaves. Amongst humans, various food processing techniques are used worldwide to detoxify otherwise inedible foods and make them palatable.

The threshold for stimulation of bitter taste by quinine averages a concentration of 0.000008 M. The taste thresholds of other bitter substances are rated relative to quinine, which is thus given a reference index of 1. For example, Brucine has an index of 11, is thus perceived as intensely more bitter than quinine, and is detected at a much lower solution threshold. The most bitter substance known is the synthetic chemical denatonium, which has an index of 1,000. It is used as an aversive agent that is added to toxic substances to prevent accidental ingestion. This was discovered in 1958 during research on lignocaine, a local anesthetic, by Macfarlan Smith of Edinburgh, Scotland.

Research has shown that TAS2Rs (taste receptors, type 2, also known as T2Rs) such as TAS2R38 coupled to the G protein gustducin are responsible for the human ability to taste bitter substances. They are identified not only by their ability to taste for certain "bitter" ligands, but also by the morphology of the receptor itself (surface bound, monomeric). Recently it is speculated that the selective constraints on the TAS2R family have been weakened due to the relatively high rate of mutation and pseudogenization. Researchers use two synthetic substances, phenylthiocarbamide (PTC) and 6-n-propylthiouracil (PROP) to study the genetics of bitter perception. These two substances taste bitter to some people, but are virtually tasteless to others. Among the tasters, some are so-called "supertasters" to whom PTC and PROP are extremely bitter. The variation in sensitivity is determined by two common alleles at the TAS2R38 locus. This genetic variation in the ability to taste a substance has been a source of great interest to those who study genetics.

Saltiness
Saltiness is a taste produced primarily by the presence of sodium ions. Other ions of the alkali metals group also taste salty, but the further from sodium the less salty the sensation is. The size of lithium and potassium ions most closely resemble those of sodium and thus the saltiness is most similar. In contrast rubidium and cesium ions are far larger so their salty taste differs accordingly. The saltiness of substances is rated relative to sodium chloride (NaCl), which has an index of 1. Potassium, as potassium chloride - KCl, is the principal ingredient in salt substitutes, and has a saltiness index of 0.6.

Other monovalent cations, e.g. ammonium, NH4+, and divalent cations of the alkali earth metal group of the periodic table, e.g. calcium, Ca2+, ions generally elicit a bitter rather than a salty taste even though they, too, can pass directly through ion channels in the tongue, generating an action potential.

Sourness
Sourness is the taste that detects acidity. The sourness of substances is rated relative to dilute hydrochloric acid, which has a sourness index of 1. By comparison, tartaric acid has a sourness index of 0.7, citric acid an index of 0.46, and carbonic acid an index of 0.06. The mechanism for detecting sour taste is similar to that which detects salt taste. Hydrogen ion channels detect the concentration of hydronium ions that are formed from acids and water. Additionally, the taste receptor PKD2L1 has been found to be involved in tasting sourness.

Hydrogen ions are capable of permeating the amiloride-sensitive channels, but this is not the only mechanism involved in detecting the quality of sourness. Other channels have also been proposed in the literature. Hydrogen ions also inhibit the potassium channel, which normally functions to hyperpolarize the cell. By a combination of direct intake of hydrogen ions (which itself depolarizes the cell) and the inhibition of the hyperpolarizing channel, sourness causes the taste cell to fire in this specific manner. In addition, it has also been suggested that weak acids, such as CO2 which is converted into the bicarbonate ion by the enzyme carbonic anhydrase, to mediate weak acid transport. The most common food group that contains naturally sour foods is fruit, with examples such as lemon, grape, orange, and sometimes melon. Wine also usually has a sour tinge to its flavor. If not kept correctly, milk can spoil and contain a sour taste. Sour candy is especially popular in North America including Cry Babies, Warheads, Lemon drops, Shock tarts and Sour Skittles and Starburst. Many of these candies contain citric acid.

Sweetness
Sweetness, usually regarded as a pleasurable sensation, is produced by the presence of sugars, some proteins and a few other substances. Sweetness is often connected to aldehydes and ketones, which contain a carbonyl group. Sweetness is detected by a variety of G protein coupled receptors coupled to the G protein gustducin found on the taste buds. At least two different variants of the "sweetness receptors" need to be activated for the brain to register sweetness. The compounds which the brain senses as sweet are thus compounds that can bind with varying bond strength to two different sweetness receptors. These receptors are T1R2+3 (heterodimer) and T1R3 (homodimer), which are shown to be accountable for all sweet sensing in humans and animals. Taste detection thresholds for sweet substances are rated relative to sucrose, which has an index of 1. The average human detection threshold for sucrose is 10 millimoles per litre. For lactose it is 30 millimoles per litre, with a sweetness index of 0.3, and 5-Nitro-2-propoxyaniline 0.002 millimoles per litre.

Umami
Umami is an appetitive taste and is described as a savory or meaty taste. It can be tasted in cheese and soy sauce, and while also found in many other fermented and aged foods, this taste is also present in tomatoes, grains, and beans. Monosodium glutamate (MSG), developed as a food additive in 1908 by Kikunae Ikeda, produces a strong umami taste. A loanword from Japanese meaning "good flavor" or "good taste", Umami (旨味) is considered fundamental to many Eastern cuisines and was first described in 1908, although it was only recently recognized in the West as a basic taste.

Some umami taste buds respond specifically to glutamate in the same way that "sweet" ones respond to sugar. Glutamate binds to a variant of G protein coupled glutamate receptors.

Measuring relative tastes
Measuring the degree by which a substance presents one basic taste can be done in a subjective way by comparing its taste to a reference substance.

Quinine, a bitter medicinal found in tonic water, can be used to subjectively rate the bitterness of a substance. Units of dilute quinine hydrochloride (1 g in 2000 mL of water) can be used to measure the threshold bitterness concentration, the level at which the presence of a dilute bitter substance can be detected by a human taster, of other compounds. More formal chemical analysis, while possible, is difficult.

Relative saltiness can be rated by comparison to a dilute salt solution.

The sourness of a substance can be rated by comparing it to very dilute hydrochloric acid (HCl).

Sweetness is subjectively measured by comparing the threshold values, or level at which the presence of a dilute substance can be detected by a human taster, of different sweet substances. Substances are usually measured relative to sucrose, which is usually given an arbitrary index of 1 or 100. Fructose is about 1.4 times sweeter than sucrose; glucose, a sugar found in honey and vegetables, is about three-quarters as sweet; and lactose, a milk sugar, is one-half as sweet.

Functional structure
Research has shown that TAS2Rs (taste receptors, type 2, also known as T2Rs) such as TAS2R38 are responsible for the human ability to taste bitter substances. They are identified not only by their ability to taste certain bitter ligands, but also by the morphology of the receptor itself (surface bound, monomeric).
 * Bitterness:

Saltiness is a taste produced best by the presence of cations (such as, or ) and, like sour, it is tasted using ion channels.
 * Saltiness:

Other ions of the alkali metals group also taste salty, but the less sodium-like the ion is, the less salty the sensation. As the size of lithium and potassium ions is close to that of sodium, they taste similar to salt. In contrast, the larger rubidium and cesium ions do not taste as salty.

Other monovalent cations, e.g., ammonium,, and divalent cations of the alkali earth metal group of the periodic table, e.g., calcium, , ions, in general, elicit a bitter rather than a salty taste even though they, too, can pass directly through ion channels in the tongue.

Sourness is acidity, and, like salt, it is a taste sensed using ion channels. Hydrogen ion channels detect the concentration of hydronium ions that are formed from acids and water. In addition, the taste receptor PKD2L1 has been found to be involved in tasting sour.
 * Sourness:

Sweetness is produced by the presence of sugars, some proteins, and a few other substances. It is often connected to aldehydes and ketones, which contain a carbonyl group. Sweetness is detected by a variety of G protein-coupled receptors coupled to a G protein that acts as an intermediary in the communication between taste bud and brain, gustducin. These receptors are T1R2+3 (heterodimer) and T1R3 (homodimer), which account for sweet sensing in humans and other animals.
 * Sweetness:

The amino acid glutamic acid is responsible for umami, but some nucleotides (inosinic acid  and guanylic acid ) can act as complements, enhancing the taste.
 * Umami-ness:

Glutamic acid binds to a variant of the G protein-coupled receptor, producing an umami taste.

Further sensations
The tongue can also feel other sensations not generally included in the basic tastes. These are largely detected by the somatosensory system.

In 2008, geneticists discovered a CaSR calcium receptor on the tongues of mice. The CaSR receptor is commonly found in the gastrointestinal tract, kidneys, and brain. Along with the "sweet" T1R3 receptor, the CaSR receptor can detect calcium as a taste. Whether closely related genes in mice and humans means the phenomenon exists in humans as well is unknown.
 * Calcium:

Some substances activate cold trigeminal receptors even when not at low temperatures. This "fresh" or "minty" sensation can be tasted in spearmint, menthol, ethanol, and camphor. Caused by activation of the same mechanism that signals cold, TRPM8 ion channels on nerve cells, unlike the actual change in temperature described for sugar substitutes, this coolness is only a perceived phenomenon.
 * Coolness:

Some foods, such as unripe fruits, contain tannins or calcium oxalate that cause an astringent or rough sensation of the mucous membrane of the mouth. Examples include tea, red wine, rhubarb, and unripe persimmons and bananas.
 * Dryness:

Less exact terms for the astringent sensation are "dry", "rough", "harsh" (especially for wine), "tart" (normally referring to sourness), "rubbery", "hard" or "styptic".

When referring to wine, "dry" is the opposite of "sweet" and does not refer to astringency. Wines that contain tannins and so cause an astringent sensation are not necessarily classified as "dry," and "dry" wines are not necessarily astringent.

In the Indian Ayurvedic tradition one of the 6 tastes is astringency (kasaaya).

Recent research has revealed a potential taste receptor called the CD36 receptor which reacts to fat (to be more specific fatty acids). This receptor was found in mice.
 * Fattiness:

Some Japanese researchers refer to the kokumi of foods laden with alcohol and thiol-groups in their amino acid extracts, and this sensation has also been described as mouthfeel.
 * Heartiness (kokumi):

Both Chinese and Batak Toba cooking include the idea of 麻 (má or mati rasa), a tingling numbness caused by spices such as Sichuan pepper. The cuisines of Sichuan province in China and of the Indonesia province North Sumatra often combine this with chili pepper to produce a 麻辣 málà, "numbing-and-hot", or "mati rasa" flavor. . These sensations although not taste fall into a category of Chemesthesis.
 * Numbness:


 * Spiciness:

Substances such as ethanol and capsaicin cause a burning sensation called Chemesthesis, piquance, spiciness, hotness, or prickliness by inducing a trigeminal nerve reaction together with normal taste reception. The sensation of heat is caused by the food's activating nerves that express TRPV1 and TRPA1 receptors. Two main plant-derived compounds that provide this sensation are capsaicin from chili peppers and piperine from black pepper. The piquant ("hot" or "spicy") sensation provided by chili peppers, black pepper, and other spices like ginger and horseradish plays an important role in a diverse range of cuisines across the world&mdash;especially in equatorial and sub-tropical climates, such as Ethiopian, Peruvian, Hungarian, Indian, Korean, Indonesian, Lao, Malaysian, Mexican, Southwest Chinese (including Szechuan cuisine), and Thai cuisines.

If tissue in the oral cavity has been damaged or sensitized, ethanol may be experienced as pain rather than simply heat. Those having had radiotherapy for oral cancer, thus, find it painful to drink alcohol.

This particular sensation, called Chemesthesis, is not a taste in the technical sense, because the sensation does not arise from taste buds and a different set of nerve fibers carry it to the brain. Foods like chili peppers activate nerve fibers directly; the sensation which is interpreted as "hot" results from the stimulation of somatosensory (pain/temperature) fibers on the tongue. Many parts of the body with exposed membranes but without taste sensors (such as the nasal cavity, under the fingernails, surface of the eye ([cornea]) or a wound) produce a similar sensation of heat when exposed to hotness agents. In Asian countries within the sphere of mainly Chinese, Indian, and Japanese cultural influence, Piquance has traditionally been considered a sixth basic taste.

Temperature can be an essential element of the taste experience. Food and drink that—within a given culture—is considered to be properly served hot is often considered distasteful if cold and vice versa. For example alcoholic beverages, with a few exceptions, are usually best when served cold but soups are usually only eaten hot. A cultural example is soda. In North America it is almost always preferred cold, regardless of season. In South America lukewarm soda is almost exclusively consumed in winter.
 * Temperature:

Supertasters
A supertaster is a person whose sense of taste is significantly more sensitive than average. The cause of this heightened response is thought to be, at least in part, due to an increased number of fungiform papillae.

Aftertaste
Aftertastes arise after food has been swallowed. An aftertaste can differ from the food it follows. Medicines and tablets may also have a lingering aftertaste, as can certain artificial flavor compounds, such as aspartame (artificial sweetener).

Acquired taste
An acquired taste is an appreciation for a food or beverage that is unlikely to be enjoyed at first taste. Many of the world's delicacies are considered to be acquired tastes.

Innervation
Taste is brought to the brainstem by 3 different cranial nerves:


 * Facial Nerve for the anterior 2/3 of the tongue and soft palate.
 * Glossopharyngeal Nerve for the posterior 1/3 of the tongue.
 * Vagus Nerve for the small area on the epiglottis.

Disorders of taste

 * ageusia (complete loss of taste)
 * dysgeusia (persistent abnormal taste)