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Taste (or more formally, gustation) is a form of direct chemoreception and is one of the traditional five senses. It refers to the ability to detect the flavor of substances such as food and poisons. In humans and many other vertebrate animals the sense of taste partners with the less direct sense of smell, in the brain's perception of flavor. Classical taste sensations include sweet, salty, sour, and bitter. More recently, psychophysicists and neuroscientists have suggested other taste categories (umami and fatty acid taste most prominently.)
Taste is a sensory function of the central nervous system. The receptor cells for taste in humans are found on the surface of the tongue, along the soft palate, and in the epithelium of the pharynx and epiglottis.
- 1 Basic taste
- 2 ==
- 2.1 Coolness
- 2.2 Spiciness or hotness
- 2.3 Numbness
- 2.4 Fat
- 2.5 Kokumi
- 2.6 Kasaaya
- 2.7 Astringency
- 2.8 Metallic taste
- 2.9 Supertasters
- 2.10 Aftertaste
- 2.11 Acquired taste
- 2.12 Factors affecting taste perception
- 2.13 Disorders of taste
- 2.14 Developmental aspects of gustation
- 2.15 Neurobiology of taste perception
- 2.16 Genetics of taste perception
- 2.17 See also
- 2.18 References
- 2.19 Further reading
Psychophysicists have long suggested the existence of four taste 'primaries', referred to as the basic tastes: sweetness, sourness, bitterness, and saltiness. Umami, or savoriness, has been suggested as a fifth basic taste, exemplified by the non-salty sensations evoked by Monosodium glutamate.
Other possible categories have been suggested, such as a taste exemplified by certain fatty acids such as linoleic acid.   Some researchers still argue against the notion of primaries at all and instead favor a continuum of percepts, in sharp contrast to color vision.
All of these taste sensations arise from all regions of the oral cavity, despite the common misperception of a "taste map" of sensitivity to different tastes thought to correspond to specific areas of the tongue. The "mouth map" is a myth, generally attributed to the mis-translation of a German text, and perpetuated in North American schools since the early twentieth century . Very slight regional differences in sensitivity to compounds exist, though these regional differences are subtle and do not conform exactly to the mythical tongue map. Individual taste buds (which contain approximately 100 taste receptor cells), in fact, typically respond to compounds evoking each of the four basic tastes.
The basic tastes are those commonly recognized types of taste sensed by humans. Humans receive tastes through sensory organs called taste buds or gustatory calyculi, concentrated on the upper surface of the tongue. Scientists describe five basic tastes: bitter, salty, sour, sweet, and umami (described as savoury, meaty, or brothy). Taste and smell are subsumed under the term flavor. The basic tastes are only one component that contributes to the sensation of food in the mouth — other factors include the food's smell, detected by the olfactory epithelium of the nose, its texture, detected by mechanoreceptors, and its temperature, detected by thermoreceptors.
In Western culture, the concept of basic tastes can be traced back at least to Aristotle, who cited "sweet" and "bitter," with "succulent," "salt," "pungent," "harsh," "astringent" and "acid" as elaborations of those two basics. The ancient Chinese Five Elements philosophy lists slightly different five basic tastes: bitter, salty, sour, sweet, and spicy. Japanese and Indian cultures each add their own sixth taste to the basic five.
For many years, books on the physiology of human taste contained diagrams of the tongue showing levels of sensitivity to different tastes in different regions. In fact, taste qualities are found in all areas of the tongue, in contrast with the popular view that different tastes map to different areas of the tongue.
The receptors for all known basic tastes have been identified. The receptors for sour and salty are ion channels while the receptors for sweet, bitter, and umami belong to the class of G protein coupled receptors.
In November 2005, a team of French researchers experimenting on rodents claimed to have evidence for a sixth taste, for fatty substances. It is speculated that humans may also have the same receptors. Fat has occasionally been raised as a possible basic taste in the past (Bravo 1592, Linnaeus 1751) but later classifications abandoned fat as a separate taste (Haller 1751 and 1763). 
Five basic tastes
For a long period, it has been commonly accepted that there are a finite number of "basic tastes" by which all foods and tastes can be grouped. For the past decades, this was considered to be a group of four basic tastes. Around 2007, a fifth taste, umami, was added by a wide number of authorities in this field.
Saltiness is a taste produced primarily by the presence of sodium ions. They can pass directly through ion channels in the tongue, generating an action potential. Calcium (Ca2+) ions can also easily activate the taste, but potassium and magnesium ions do not do so nearly as effectively, instead activating the bitter taste[How to reference and link to summary or text].
- "Sour" redirects here. For other uses, see Sour (disambiguation).
Sourness is the taste that detects acidity. The mechanism for detecting sour taste is similar to that which detects salt taste. Hydrogen ion channels detect the concentration of hydronium ions (H3O+ ions) that are formed from acids and water.
Hydrogen ions are capable of permeating the amiloride-sensitive sodium channels, but this is not the only mechanism involved in detecting the quality of sourness. Hydrogen ions also inhibit the potassium channel, which normally functions to hyperpolarize the cell. Thus, 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.
- Main article: Sweetness
Sweetness 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 (8). The average human detection threshold for sucrose is 10 millimoles per litre. For lactose it is 30 millimoles per litre, and 5-Nitro-2-propoxyaniline 0.002 millimoles per litre.
- See also: Miraculin and Curculin
The bitter taste is perceived by many to be unpleasant, sharp, or disagreeable. Evolutionary biologists have suggested that a distaste for bitter substances may have evolved as a defense mechanism against accidental poisoning. Common bitter foods and beverages include coffee, unsweetened chocolate, bitter melon, beer, uncured olives, citrus peel, many plants in the Brassicaceae family, dandelion greens and escarole. Quinine is also known for its bitter taste and is found in tonic water. (Although it must be stated that "coffee" has different types such as Robusta and Arabica; and can be quite sweet indeed regarding the region it grew.)
Research has shown that TAS2Rs (taste receptors, type 2) such as TAS2R16 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). 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. This genetic variation in the ability to taste a substance has been a source of great interest to those who study genetics. In addition, it is of interest to those who study evolution since PTC-tasting is associated with the ability to taste numerous natural bitter compounds, a large number of which are known to be toxic.
- Main article: umami
Umami is the name for the taste sensation produced by compounds such as glutamate are commonly found in fermented and aged foods. In English, it is sometimes described as "meaty" or "savoury". The word is from the Japanese word 旨味, うまみ whose characters literally mean "delicious flavour." Umami is now the commonly used term by taste scientists. The same taste is referred to as xiānwèi (鮮味 or 鲜味) in Chinese cooking. Umami is considered a fundamental taste in Chinese and Japanese cooking, but is not discussed as much in Western cuisine.
Examples of food containing glutamate (and thus strong in the umami taste) are parmesan and roquefort cheese as well as soy sauce and fish sauce. It is also found in significant amounts in various unfermented foods such as walnuts, grapes, broccoli, tomatoes, and mushrooms, and to a lesser degree in meat. The glutamate taste sensation is most intense in combination with sodium chloride (table salt). This is one reason why tomatoes exhibit a stronger taste after adding salt. Sauces with umami and salty tastes are very popular for cooking, such as tomato sauces and ketchup for Western cuisines and soy sauce and fish sauce for East Asian and Southeast Asian cuisines.
The additive monosodium glutamate (MSG), which was developed as a food additive in 1907 by Kikunae Ikeda, produces a strong umami taste. Umami is also provided by the nucleotides 5’-inosine monophosphate (IMP) and 5’-guanosine monophosphate (GMP). These are naturally present in many protein-rich foods. IMP is present in high concentrations in many foods, including dried skipjack tuna flakes used to make dashi, a Japanese broth. GMP is present in high concentration in dried shiitake mushrooms, used in much of the cuisine of Asia. There is a synergistic effect between MSG, IMP and GMP which together in certain ratios produce a strong umami taste.
Some umami taste buds responds specifically to glutamate in the same way that sweet ones respond to sugar. Glutamate binds to a variant of G protein coupled glutamate receptors. Earlier reports had postulated that a metabotropic glutamate receptor (mGluR4) and the NMDA receptor might play a role in umami perception.
Some substances activate cold trigeminal receptors. One can sense a cool sensation (also known as "fresh" or "minty") from, e.g., spearmint, menthol, ethanol or camphor, which is caused by the food activating the TRP-M8 ion channel on nerve cells that also signal cold. Unlike the actual change in temperature described for sugar substitutes, coolness is only a perceived phenomenon.
Spiciness or hotness
- See also: Scoville scale
Substances such as ethanol and capsaicin cause a burning sensation by inducing a trigeminal nerve reaction together with normal taste reception. The sensation of heat is caused by the food activating a nerve cell called TRP-V1, which is also activated by hot temperatures. The sensation, usually referred to as being "hot" or "spicy", is a notable feature of Mexican, Indian, Szechuan, Korean, Indonesian, central Vietnamese, and Thai cuisines.
The two main plants providing this sensation are chili peppers (those fruits of the Capsicum plant that contain capsaicin) and black pepper.
If tissue in the oral cavity has been damaged or sensitised ethanol may be experienced as pain rather than simply heat. Those who have had radiotherapy for oral cancer thus find it painful to drink alcohol.[How to reference and link to summary or text]
Chinese cooking includes the idea of 麻 má, the sensation of tingling numbness caused by spices such as Sichuan pepper. The cuisine of Sichuan province often combines this with chili pepper to produce a 麻辣 málà, "numbing-and-hot", flavour.
Recent research has revealed a potential taste receptor called the CD36 receptor to be reacting to fat, more specifically, fatty acids. This receptor was found in mice, but probably exists among other mammals as well. In experiments, mice with a genetic defect that blocked this receptor didn't show the same urge to consume fatty acids as normal mice, and failed to prepare gastric juices in their digestive tracts to digest fat. This discovery may lead to a better understanding of the biochemical reasons behind this behaviour, although more research is still necessary to confirm the relationship of CD36 and the cravings of fat.
Some Japanese researchers refer to a flavour called kokumi which has been described variously as continuity, "mouthfulness", mouthfeel and thickness.
In Indian tradition, the tastes are referred to as 'Arusuvai' or six tastes . These tastes are normally referred to as the following: sweet, sour, salty, bitter, hot / spicy and astringent. Some people call the sixth taste as neutral or tasteless. A typical example of a neutral tasting substance is water. Certain others say the astringent or the sixth taste is a mix of varied tastes and is termed Kasaaya, in India. That is more in line with the Japanese approach to umami.
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 or the teeth. Examples include tea, rhubarb, grapes and unripe persimmons and bananas.
Less exact terms for the astringent sensation include: "rubbery", "hard", "styptic", "dry", "rough", "harsh" (especially for wine) and "tart" (normally referring to sourness) . The Chinese have a term for this: 澀 (sè), the Korean have 떫다 (tteolda), the Japanese call it 渋い (shibui), while Thai have ฝาด (fard), the Malay use kelat, and in Russian there is вяжущий (vyazhuschiy) or тёрпкий (tjorpky).
Most people know this taste (e.g. Cu2+, FeSO4, or blood in mouth), but it is not only taste but olfactory receptors worked in this case (Guth and Grosch, 1990).
- Some diseases cause a "metallic taste"
- Some substances (tetracycline, H2S) can stimulate "metallic taste" 
- Main article: Supertaster
A supertaster is a person whose sense of taste is significantly more acute than average. Women are more likely to be supertasters, as are Asians, Africans and South Americans. Among individuals of European descent, it is estimated that about 25% of the population are supertasters. The cause of this heightened response is currently unknown, although it is thought to be, at least in part, due to an increased number of fungiform papillae. The evolutionary advantage to supertasting is unclear. In some environments, heightened taste response, particularly to bitterness, would represent an important advantage in avoiding potentially toxic plant alkaloids. However, in other environments, increased response to bitter may have limited the range of palatable foods. In our modern, energy-rich environment, supertasting may be cardioprotective, due to decreased liking and intake of fat, but may increase cancer risk via decreased vegetable intake. It may be a cause of picky eating, but picky eaters are not necessarily supertasters, and vice versa.
- Main article: Aftertaste
Aftertaste is the persistence of a sensation of flavor after the stimulating substance has passed out of contact with the sensory end organs for taste. Some aftertastes may be pleasant, others unpleasant.
Alcoholic beverages such as wine, beer and whiskey are noted for having particularly strong aftertastes. Foodstuffs with notable aftertastes include spicy food, such as Mexican food (e.g. chili pepper), or Indian food (such as curry).
Medicines and tablets may also have a lingering aftertaste.
An acquired taste is an appreciation for a food or beverage that is unlikely to be enjoyed, in part or in full, by a person who has not had substantial exposure to it, usually because of some unfamiliar aspect of the food or beverage, including a strong or strange odor, taste, or appearance. The process of “acquiring” a taste involves consuming a food or beverage in the hope of learning to enjoy it. In most cases, this introductory period is considered worthwhile, as many of the world's delicacies are considered to be acquired tastes. A connoisseur is one who is held to have an expert judgment of taste.
Factors affecting taste perception
Many factors affect taste perception, including:
- Color/vision impairments
- Hormonal influences
- Genetic variations - See Phenylthiocarbamide
- Oral temperature
- Drugs and chemicals
- CNS Tumors (esp. Temporal lobe lesions)
- Plugged noses
It is also important to consider that flavor is the overall, total sensation induced during mastication (e.g. taste, touch, pain and smell). Smell (olfactory stimulation) plays a major role in flavor perception.
Disorders of taste
- ageusia (complete loss)
- hypogeusia (partial loss)
- parageusia (unpleasant taste)
- dysgeusia (inaccurate taste)
Developmental aspects of gustation
Everyone knows that children have different tastes than adults.
- Main article: Developmental aspects of gustation
Neurobiology of taste perception
- Main article: Neurobiology of taste perception
Genetics of taste perception
- Lexical-gustatory synesthesia
- Olfactory perception
- Optimal foraging theory
- Taste buds
- Taste disorders
- Taste stimulation
- Vomeronasal organ
- Journal of the Chemical Society of Tokyo, 30, 820-836, (1909)
- Nelson G, Chandrashekar J, Hoon MA, et al (2002). An amino-acid taste receptor. Nature 416 (6877): 199-202.
- Lindemann, Bernd (1999). Receptor seeks ligand: On the way to cloning the molecular receptors for sweet and bitter taste. Nature Medicine 5 (4): 381.
- Huang A. L., et al. "The cells and logic for mammalian sour taste detection" (no free access). Nature, 442. 934 - 938 (2006).
- Scenta. "How sour taste buds grow". August 25, 2006.
- Laugerette, Fabienne, Patricia Passilly-Degrace, Bruno Patris, Isabelle Niot, Maria Febbraio, Jean-Pierre Montmayeur, Philippe Besnard (November 2005). CD36 involvement in orosensory detection of dietary lipids, spontaneous fat preference, and digestive secretions. The Journal of Clinical Investigation 115 (11): 3177-3184.
- Abumrad, Nada A. (November 2005). CD36 may determine our desire for dietary fats. The Journal of Clinical Investigation 115 (11): 2965-2967.
- Boring, Edwin G. (1942). Sensation and Perception in the History of Experimental Psychology, p. 453, Appleton Century Crofts.
- Lindemann "Receptors and transduction in taste." Nature 2001
- Lindemann, Bernd (2000). A taste for umami. Nature Neuroscience.
- Christian Murray MD, Nowell Solish MD, FRCPC (2003) Metallic Taste: An Unusual Reaction to Botulinum Toxin A / Dermatologic Surgery 29 (5), 562–563.
- Long-term effects on the olfactory system of exposure to hydrogen sulphide / AR Hirsch and G Zavala / Occupational and Environmental Medicine, Vol 56, 284-287
- Bartoshuk, L. M., V. B. Duffy, et al. (1994). "PTC/PROP tasting: anatomy, psychophysics, and sex effects." 1994. Physiol Behav 56(6): 1165-71.
- Doty, R.L. (2003). Handbook of Gustation and Olfaction.Marcel Dekker, Inc.:NY
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