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Taste

taste of home, tastefully simple
Taste, gustatory perception, or gustation1 is one of the five traditional senses that belongs to the gustatory system

Taste is the sensation produced when a substance in the mouth reacts chemically with taste receptor cells located on taste buds in the oral cavity, mostly on the tongue Taste, along with smell olfaction and trigeminal nerve stimulation registering texture, pain, and temperature, determines flavors of food or other substances Humans have taste receptors on taste buds gustatory calyculi and other areas including the upper surface of the tongue and the epiglottis23 The gustatory cortex is responsible for the perception of taste

The tongue is covered with thousands of small bumps called papillae, which are visible to the naked eye Within each papilla are hundreds of taste buds4 The exception to this is the filiform papillae that do not contain taste buds There are between 2000 and 50005 taste buds that are located on the back and front of the tongue Others are located on the roof, sides and back of the mouth, and in the throat Each taste bud contains 50 to 100 taste receptor cells

The sensation of taste includes five established basic tastes: sweetness, sourness, saltiness, bitterness, and umami67 Scientific experiments have proven that these five tastes exist and are distinct from one anothercitation needed Taste buds are able to differentiate among different tastes through detecting interaction with different molecules or ions Sweet, umami, and bitter tastes are triggered by the binding of molecules to G protein-coupled receptors on the cell membranes of taste buds Saltiness and sourness are perceived when alkali metal or hydrogen ions enter taste buds, respectively8

The basic tastes contribute only partially to the sensation and flavor of food in the mouth—other factors include smell,2 detected by the olfactory epithelium of the nose;9 texture,10 detected through a variety of mechanoreceptors, muscle nerves, etc;11 temperature, detected by thermoreceptors; and "coolness" such as of menthol and "hotness" pungency, through chemesthesis

As taste senses both harmful and beneficial things, all basic tastes are classified as either aversive or appetitive, depending upon the effect the things they sense have on our bodies12 Sweetness helps to identify energy-rich foods, while bitterness serves as a warning sign of poisons13

Among humans, taste perception begins to fade around 50 years of age because of loss of tongue papillae and a general decrease in saliva production14 Also, not all mammals share the same taste senses: some rodents can taste starch which humans cannot, cats cannot taste sweetness, and several other carnivores including hyenas, dolphins, and sea lions, have lost the ability to sense up to four of their ancestral five taste senses15

Contents

  • 1 Basic tastes
    • 11 Sweetness
    • 12 Sourness
    • 13 Saltiness
    • 14 Bitterness
    • 15 Umami
  • 2 Measuring relative tastes
  • 3 Functional structure
  • 4 Further sensations and transmission
    • 41 Pungency also spiciness or hotness
    • 42 Coolness
    • 43 Numbness
    • 44 Astringency
    • 45 Metallicness
    • 46 Calcium
    • 47 Fattiness oleogustus
    • 48 Heartiness kokumi
    • 49 Temperature
    • 410 Starchiness
  • 5 Nerve supply and neural connections
  • 6 Other concepts
    • 61 Taste as a philosophical concept
    • 62 Supertasters
    • 63 Aftertaste
    • 64 Acquired taste
  • 7 Clinical significance
    • 71 Disorders of taste
  • 8 History
  • 9 Research
  • 10 See also
  • 11 Notes
    • 111 Footnotes
    • 112 Citations
  • 12 Further reading
  • 13 External links

Basic tastesedit

Taste in the gustatory system allows humans to distinguish between safe and harmful food, and to gauge foods’ nutritional value Digestive enzymes in saliva begin to dissolve food into base chemicals that are washed over the papillae and detected as tastes by the taste buds The tongue is covered with thousands of small bumps called papillae, which are visible to the naked eye Within each papilla are hundreds of taste buds4 The exception to this are the filiform papillae that do not contain taste buds There are between 2000 and 50005 taste buds that are located on the back and front of the tongue Others are located on the roof, sides and back of the mouth, and in the throat Each taste bud contains 50 to 100 taste receptor cells

Bitter foods are generally found unpleasant, while sour, salty, sweet, and meaty tasting foods generally provide a pleasurable sensation The five specific tastes received by taste receptors are saltiness, sweetness, bitterness, sourness, and umami, which means ‘delicious’ in Japanese and is sometimes translated as “savory” in English As of the early twentieth century, physiologists and psychologists believed there were four basic tastes: sweetness, sourness, saltiness, and bitterness At that time umami was not identified16 but now a large number of authorities recognize it as the fifth taste

One study found that both salt and sour taste mechanisms detect, in different ways, the presence of sodium chloride salt in the mouth, however, acids are also detected and perceived as sour17 The detection of salt is important to many organisms, but specifically mammals, as it serves a critical role in ion and water homeostasis in the body It is specifically needed in the mammalian kidney as an osmotically active compound which facilitates passive re-uptake of water into the bloodcitation needed Because of this, salt elicits a pleasant taste in most humans

Sour and salt tastes can be pleasant in small quantities, but in larger quantities become more and more unpleasant to taste For sour taste this is presumably because the sour taste can signal under-ripe fruit, rotten meat, and other spoiled foods, which can be dangerous to the body because of bacteria which grow in such media Additionally, sour taste signals acids, which can cause serious tissue damage

The bitter taste is almost universally unpleasant to humans This is because many nitrogenous organic molecules which have a pharmacological effect on humans taste bitter These include caffeine, nicotine, and strychnine, which respectively compose the stimulant in coffee, addictive agent in cigarettes, and active compound in many pesticides It appears that some psychological process allows humans to overcome their innate aversion to bitter taste, as caffeinated drinks are widely consumed and enjoyed around the world It is also interesting to note that many common medicines have a bitter taste if chewed; the gustatory system apparently interprets these compounds as poisons In this manner, the unpleasant reaction to the bitter taste is a last-line warning system before the compound is ingested and can do damage

Sweet taste signals the presence of carbohydrates in solution Since carbohydrates have a very high calorie count saccharides have many bonds, therefore much energy, they are desirable to the human body, which evolved to seek out the highest calorie intake foods They are used as direct energy sugars and storage of energy glycogen However, there are many non-carbohydrate molecules that trigger a sweet response, leading to the development of many artificial sweeteners, including saccharin, sucralose, and aspartame It is still unclear how these substances activate the sweet receptors and what adaptational significance this has had

The umami taste, identified by Japanese chemist Kikunae Ikeda of Tokyo Imperial University, which signals the presence of the amino acid L-glutamate, triggers a pleasurable response and thus encourages the intake of peptides and proteins The amino acids in proteins are used in the body to build muscles and organs, transport molecules hemoglobin, antibodies, and the organic catalysts known as enzymes These are all critical molecules, and as such it is important to have a steady supply of amino acids, hence the pleasurable response to their presence in the mouth

In Asian countries within the sphere of mainly Chinese and Indian cultural influence, pungency piquancy or hotness had traditionally been considered a sixth basic taste18 In 2015, researchers at Purdue University suggested a new basic taste of fats called oleogustus19

Sweetnessedit

The diagram above depicts the signal transduction pathway of the sweet taste Object A is a taste bud, object B is one taste cell of the taste bud, and object C is the neuron attached to the taste cell I Part I shows the reception of a molecule 1 Sugar, the first messenger, binds to a protein receptor on the cell membrane II Part II shows the transduction of the relay molecules 2 G Protein-coupled receptors, second messengers, are activated 3 G Proteins activate adenylate cyclase, an enzyme, which increases the cAMP concentration Depolarization occurs 4 The energy, from step 3, is given to activate the K+, potassium, protein channelsIII Part III shows the response of the taste cell 5 Ca+, calcium, protein channels is activated6 The increased Ca+ concentration activates neurotransmitter vesicles 7 The neuron connected to the taste bud is stimulated by the neurotransmitters Main article: Sweetness

Sweetness, usually regarded as a pleasurable sensation, is produced by the presence of sugars 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" must be activated for the brain to register sweetness Compounds 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 account for all sweet sensing in humans and animals20 Taste detection thresholds for sweet substances are rated relative to sucrose, which has an index of 12122 The average human detection threshold for sucrose is 10 millimoles per liter For lactose it is 30 millimoles per liter, with a sweetness index of 03,21 and 5-Nitro-2-propoxyaniline 0002 millimoles per liter “Natural” sweeteners such as saccharides activate the GPCR, which releases gustducin The gustducin then activates the molecule adenylate cyclase, which catalyzes the production of the molecule cAMP, or adenosine 3', 5'-cyclic monophosphate This molecule closes potassium ion channels, leading to depolarization and neurotransmitter release Synthetic sweeteners such as saccharin activate different GPCRs and induce taste receptor cell depolarization by an alternate pathway

Sournessedit

"Sour" redirects here For other uses, see Sour disambiguation The diagram depicts the signal transduction pathway of the sour or salty taste Object A is a taste bud, object B is a taste receptor cell within object A, and object C is the neuron attached to object B I Part I is the reception of hydrogen ions or sodium ions 1 If the taste is sour, H+ ions, from an acidic substances, pass through their specific ion channel Some can go through the Na+ channels If the taste is salty Na+, sodium, molecules pass through the Na+ channels Depolarization takes place II Part II is the transduction pathway of the relay molecules2 Cation, such as K+, channels are opened III Part III is the response of the cell 3 An influx of Ca+ ions is activated4 The Ca+ activates neurotransmitters 5 A signal is sent to the neuron attached to the taste bud

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 07, citric acid an index of 046, and carbonic acid an index of 0062122

Sour taste is detected by a small subset of cells that are distributed across all taste buds in the tongue Sour taste cells can be identified by expression of the protein PKD2L1,23 although this gene is not required for sour responses There is evidence that the protons that are abundant in sour substances can directly enter the sour taste cells through apically located ion channels24 This transfer of positive charge into the cell can itself trigger an electrical response It has also been proposed that weak acids such as acetic acid, which are not fully dissociated at physiological pH values, can penetrate taste cells and thereby elicit an electrical response According to this mechanism, intracellular hydrogen ions inhibit potassium channels, which normally function 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 action potentials and release neurotransmitter25

The most common food group that contains naturally sour foods is fruit, such as lemon, grape, orange, tamarind, and sometimes melon Wine also usually has a sour tinge to its flavor, and if not kept correctly, milk can spoil and develop a sour taste Children in the US and UK show a greater enjoyment of sour flavors than adults,26 and sour candy is popular in North America27 including Cry Babies, Warheads, Lemon drops, Shock Tarts and sour versions of Skittles and Starburst Many of these candies contain citric acid

Saltinessedit

"Saltiness" redirects here For the saltiness in the water, see Salinity

The simplest receptor found in the mouth is the sodium chloride salt receptor 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 A sodium channel in the taste cell wall allows sodium cations to enter the cell This on its own depolarizes the cell, and opens voltage-dependent calcium channels, flooding the cell with positive calcium ions and leading to neurotransmitter release This sodium channel is known as an epithelial sodium channel ENaC and is composed of three subunits An ENaC can be blocked by the drug amiloride in many mammals, especially rats The sensitivity of the salt taste to amiloride in humans, however, is much less pronounced, leading to conjecture that there may be additional receptor proteins besides ENaC to be discovered

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 accordinglycitation needed The saltiness of substances is rated relative to sodium chloride NaCl, which has an index of 12122 Potassium, as potassium chloride KCl, is the principal ingredient in salt substitutes and has a saltiness index of 062122

Other monovalent cations, eg ammonium, NH4+, and divalent cations of the alkali earth metal group of the periodic table, eg 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

Bitternessedit

See also: Bitter taste evolution The diagram depicted above shows the signal transduction pathway of the bitter taste Bitter taste has many different receptors and signal transduction pathways Bitter indicates poison to animals It is most similar to sweet Object A is a taste bud, object B is one taste cell, and object C is a neuron attached to object B I Part I is the reception of a molecule1 A bitter substance such as quinine, is consumed and binds to G Protein-coupled receptorsII Part II is the transduction pathway 2 Gustducin, a G protein second messenger, is activated 3 Phosphodiesterase, an enzyme, is then activated 4 Cyclic nucleotide, cNMP, is used, lowering the concentration 5 Channels such as the K+, potassium, channels, closeIII Part III is the response of the taste cell 6 This leads to increased levels of Ca+ 7 The neurotransmitters are activated 8 The signal is sent to the neuron

Bitterness is the most sensitive of the tastes, and many perceive it as unpleasant, sharp, or disagreeable, but it is sometimes desirable and intentionally added via various bittering agents Common bitter foods and beverages include coffee, unsweetened cocoa, South American mate, bitter gourd, olives, citrus peel, many plants in the Brassicaceae family, dandelion greens, wild chicory, and escarole The ethanol in alcoholic beverages tastes bitter,28 as do the additional bitter ingredients found in some alcoholic beverages including hops in beer and orange in bitters 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 researchers2129 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 function212930 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 leaves31 Amongst humans, various food processing techniques are used worldwide to detoxify otherwise inedible foods and make them palatable32 Furthermore, the use of fire, changes in diet, and avoidance of toxins has led to neutral evolution in human bitter sensitivity This has allowed several loss of function mutations that has led to a reduced sensory capacity towards bitterness in humans when compared to other species33

The threshold for stimulation of bitter taste by quinine averages a concentration of 8 μM 8 micromolar21 The taste thresholds of other bitter substances are rated relative to quinine, which is thus given a reference index of 12122 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 threshold21 The most bitter substance known is the synthetic chemical denatonium, which has an index of 1,00022 It is used as an aversive agent a bitterant 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 Gorgie, Edinburgh, Scotlandcitation needed

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 substances34 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, monomeric35 The TAS2R family in humans is thought to comprise about 25 different taste receptors, some of which can recognize a wide variety of bitter-tasting compounds36 Over 550 bitter-tasting compounds have been identified, on a bitter database, of which about 100 have been assigned to one or more specific receptors37 Recently it is speculated that the selective constraints on the TAS2R family have been weakened due to the relatively high rate of mutation and pseudogenization38 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 locus39 This genetic variation in the ability to taste a substance has been a source of great interest to those who study genetics

Gustducin is made of three subunits When it is activated by the GPCR, its subunits break apart and activate phosphodiesterase, a nearby enzyme, which in turn converts a precursor within the cell into a secondary messenger, which closes potassium ion channelscitation needed Also, this secondary messenger can stimulate the endoplasmic reticulum to release Ca2+ which contributes to depolarization This leads to a build-up of potassium ions in the cell, depolarization, and neurotransmitter release It is also possible for some bitter tastants to interact directly with the G protein, because of a structural similarity to the relevant GPCR

Umamiedit

Main article: Umami

Umami is an appetitive taste12 and is described as savory4041 or meaty4142 It can be tasted in cheese43 and soy sauce,44 and is also found in many other fermented and aged foods This taste is also present in tomatoes, grains, and beans43

A loanword from Japanese meaning "good flavor" or "good taste",45 umami 旨味 is considered fundamental to many Eastern cuisines;46 and other cuisines have long operated under principles that sought to combine foods to produce umami flavors, such as in the emphasis on veal stock by Auguste Escoffier, the pre-eminent chef of 19th century French cuisine,47 and in the Romans' deliberate use of fermented fish sauce48 However, it was only recently recognized in modern science as a basic taste; well after the other basic tastes have been recognized by scientists, in part due to their correspondence with the four tastes of ancient Greek philosophy4449 Umami, or “scrumptiousness”, was first studied with the scientific method and identified by Kikunae Ikeda, who began to analyze kombu in 1907, attempting to isolate its dashi taste He isolated a substance he called ajinomoto, Japanese for “at the origin of flavor” Later identified as the chemical monosodium glutamate MSG, and increasingly used independently as a food additive,650 it is a sodium salt that produces a strong umami taste, especially combined with foods rich in nucleotides such as meats, fish, nuts, and mushrooms4451

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 receptors5253 It is thought that the amino acid L-glutamate bonds to a type of GPCR known as a metabotropic glutamate receptor mGluR4 This causes the G-protein complex to activate a secondary receptor, which ultimately leads to neurotransmitter release The intermediate steps are not known See TAS1R1 and TAS1R3 pages for a further explanation of the amino-acid taste receptor

Measuring relative tastesedit

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

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 substances54 Substances are usually measured relative to sucrose,55 which is usually given an arbitrary index of 15657 or 10058 Fructose is about 14 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 sweetb54

The sourness of a substance can be rated by comparing it to very dilute hydrochloric acid HCl59

Relative saltiness can be rated by comparison to a dilute salt solution60

Quinine, a bitter medicinal found in tonic water, can be used to subjectively rate the bitterness of a substance61 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 compounds61 More formal chemical analysis, while possible, is difficult61

Functional structureedit

Taste buds and papillae of the tongue

In the human body a stimulus refers to a form of energy which elicits a physiological or psychological action or response Sensory receptors are the structures in the body which change the stimulus from one form of energy to another This can mean changing the presence of a chemical, sound wave, source of heat, or touch to the skin into an electrical action potential which can be understood by the brain, the body’s control center Sensory receptors are modified ends of sensory neurons; modified to deal with specific types of stimulus, thus there are many different types of sensory receptors in the body The neuron is the primary component of the nervous system, which transmits messages from sensory receptors all over the body

Taste is a form of chemoreception which occurs in the specialised taste receptors in the mouth To date, there are five different types of taste receptors known: salt, sweet, sour, bitter, and umami Each receptor has a different manner of sensory transduction: that is, of detecting the presence of a certain compound and starting an action potential which alerts the brain It is a matter of debate whether each taste cell is tuned to one specific tastant or to several; Smith and Margolskee claim that "gustatory neurons typically respond to more than one kind of stimulus, although each neuron responds most strongly to one tastant" Researcherswho believe that the brain interprets complex tastes by examining patterns from a large set of neuron responses This enables the body to make "keep or spit out" decisions when there is more than one tastant present "No single neuron type alone is capable of discriminating among stimuli or different qualities, because a given cell can respond the same way to disparate stimuli"citation needed As well, serotonin is thought to act as an intermediary hormone which communicates with taste cells within a taste bud, mediating the signals being sent to the brain Receptor molecules are found on the top of microvilli of the taste cells

Sweetness

Sweetness is produced by the presence of sugars, some proteins, and a few other substancescitation needed It is often connected to aldehydes and ketones, which contain a carbonyl groupcitation needed 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, gustducin62 These receptors are T1R2+3 heterodimer and T1R3 homodimer, which account for sweet sensing in humans and other animals63

Sourness

Sourness is acidity,6465 and, like salt, it is a taste sensed using ion channels66 Undissociated acid diffuses across the plasma membrane of a presynaptic cell, where it dissociates in accordance with Le Chatelier's principle The protons that are released then block potassium channels, which depolarise the cell and cause calcium influx In addition, the taste receptor PKD2L1 has been found to be involved in tasting sour67

Saltiness

Saltiness is a taste produced best by the presence of cations such as Na+
, K+
or Li+
66 and is directly detected by cation influx into glial like cells via leak channels causing depolarisation of the cell66

Other monovalent cations, eg, ammonium, NH+
4, and divalent cations of the alkali earth metal group of the periodic table, eg, calcium, Ca2+
, ions, in general, elicit a bitter rather than a salty taste even though they, too, can pass directly through ion channels in the tonguecitation needed

Bitterness

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 substances68 They are identified not only by their ability to taste certain bitter ligands, but also by the morphology of the receptor itself surface bound, monomeric69

Umami

The amino acid glutamic acid is responsible for umami,7071 but some nucleotides inosinic acid4672 and guanylic acid70 can act as complements, enhancing the taste4672

Glutamic acid binds to a variant of the G protein-coupled receptor, producing an umami taste5253

Further sensations and transmissionedit

The tongue can also feel other sensations not generally included in the basic tastes These are largely detected by the somatosensory system In humans, the sense of taste is conveyed via three of the twelve cranial nerves The facial nerve VII carries taste sensations from the anterior two thirds of the tongue, the glossopharyngeal nerve IX carries taste sensations from the posterior one third of the tongue while a branch of the vagus nerve X carries some taste sensations from the back of the oral cavity

The trigeminal nerve cranial nerve V provides information concerning the general texture of food as well as the taste-related sensations of peppery or hot from spices

Pungency also spiciness or hotnessedit

Main articles: Pungency and 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's activating nerves that express TRPV1 and TRPA1 receptors Some such plant-derived compounds that provide this sensation are capsaicin from chili peppers, piperine from black pepper, gingerol from ginger root and allyl isothiocyanate from horseradish The piquant "hot" or "spicy" sensation provided by such foods and spices plays an important role in a diverse range of cuisines across the world—especially in equatorial and sub-tropical climates, such as Ethiopian, Peruvian, Hungarian, Indian, Korean, Indonesian, Lao, Malaysian, Mexican, New Mexican, Singaporean, Southwest Chinese including Szechuan cuisine, Vietnamese, and Thai cuisines

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 interpreted as "hot" results from the stimulation of somatosensory pain/temperature fibers on the tongue Many parts of the body with exposed membranes but no taste sensors such as the nasal cavity, under the fingernails, surface of the eye or a wound produce a similar sensation of heat when exposed to hotness agents Asian countries within the sphere of, mainly, Chinese, Indian, and Japanese cultural influence, often wrote of pungency as a fifth or sixth taste

Coolnessedit

Some substances activate cold trigeminal receptors even when not at low temperatures This "fresh" or "minty" sensation can be tasted in peppermint, 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

Numbnessedit

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 Indonesian province of North Sumatra often combine this with chili pepper to produce a 麻辣 málà, "numbing-and-hot", or "mati rasa" flavor73 These sensations although not taste fall into a category of Chemesthesis

Astringencyedit

Some foods, such as unripe fruits, contain tannins or calcium oxalate that cause an astringent or puckering sensation of the mucous membrane of the mouth Examples include tea, red wine, rhubarb, some fruits of the Syzygium genus, and unripe persimmons and bananas

Less exact terms for the astringent sensation are "dry", "rough", "harsh" especially for wine, "tart" normally referring to sourness, "rubbery", "hard" or "styptic"74

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 six tastes is astringency kasaaya75 In Sinhala and Sri Lankan English it is referred to as kahata76

Metallicnessedit

A metallic taste may be caused by food and drink, certain medicines or amalgam dental fillings It is generally considered an off flavor when present in food and drink A metallic taste may be caused by galvanic reactions in the mouth In the case where it is caused by dental work, the dissimilar metals used may produce a measurable current77 Some artificial sweeteners are perceived to have a metallic taste, which is detected by the TRPV1 receptors78 Blood is considered by many people to have a metallic taste7980 A metallic taste in the mouth is also a symptom of various medical conditions, in which case it may be classified under the symptoms dysgeusia or parageusia, referring to distortions of the sense of taste,81 and can be caused by various kinds of medication, including saquinavir81 and zonisamide,82 and occupational hazards, such as working with pesticides83

Calciumedit

The distinctive taste of chalk has been identified as the calcium component of that substance84 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 unknown8586

Fattiness oleogustusedit

Recent research reveals a potential taste receptor called the CD36 receptor878889 CD36 was targeted as a possible lipid taste receptor because it binds to fat molecules more specifically, long-chain fatty acids,90 and it has been localized to taste bud cells specifically, the circumvallate and foliate papillae91 There is a debate over whether we can truly taste fats, and supporters of our ability to taste free fatty acids FFAs have based the argument on a few main points: there is an evolutionary advantage to oral fat detection; a potential fat receptor has been located on taste bud cells; fatty acids evoke specific responses that activate gustatory neurons, similar to other currently accepted tastes; and, there is a physiological response to the presence of oral fat92 Although CD36 has been studied primarily in mice, research examining human subjects' ability to taste fats found that those with high levels of CD36 expression were more sensitive to tasting fat than were those with low levels of CD36 expression;93 this study points to a clear association between CD36 receptor quantity and the ability to taste fat

Other possible fat taste receptors have been identified G protein-coupled receptors GPR120 and GPR40 have been linked to fat taste, because their absence resulted in reduced preference to two types of fatty acid linoleic acid and oleic acid, as well as decreased neuronal response to oral fatty acids94

Monovalent cation channel TRPM5 has been implicated in fattiness taste as well,95 but it is thought to be involved primarily in downstream processing of the taste rather than primary reception, as it is with other tastes such as bitter, sweet, and umami92

A 2015 study, proposed naming the taste of fat as "oleogustus"1996 The main form of fat that is commonly ingested is triglycerides, which are composed of three fatty acids bound together In this state, triglycerides are able to give fatty foods unique textures that are often described as creaminess But this texture is not an actual taste It is only during ingestion that the fatty acids that make up triglycerides are broken apart and the taste of fat is revealed The taste is commonly related to other, more negative, tastes such as bitter and sour due to how unpleasant the taste is for humans Richard Mattes, a co-author of the study, explained that low concentrations of these fatty acids can create an overall better flavor in a food, much like how small uses of bitterness can make certain foods more rounded However, a high concentration of fatty acids in certain foods is generally considered inedible97 To demonstrate that individuals can distinguish oleogustus from other flavors, the researchers separated volunteers into groups and had them try samples that also contained the other basic tastes Volunteers were able to separate the taste of fatty acids into their own category, with some overlap with umami samples, which the researchers hypothesized was due to poor familiarity with both The researchers note that the usual "creaminess and viscosity we associate with fatty foods is largely due to triglycerides", unrelated to the taste; while the actual taste of fatty acids is not pleasant Mattes described the taste as "more of a warning system" that a certain food should not be eaten19

There are few regularly consumed foods rich in oleogustus, due to the negative flavor that is evoked in large quantities Foods whose flavor to which oleogustus makes a small contribution include olive oil and fresh butter, along with various kinds of vegetable and nut oils98

Heartiness kokumiedit

Some Japanese researchers refer to the kokumi of foods This sensation has also been described as mouthfulness,99:290 and appears to be related to a number of γ-L-glutamyl peptides, which activate a calcium-sensing receptor which is also sensitive to glutathione99

Temperatureedit

Temperature can be an essential element of the taste experience Food and drink that—in a given culture—is traditionally served hot is often considered distasteful if cold, and vice versa For example, alcoholic beverages, with a few exceptions, are usually thought best when served at room temperature or chilled to varying degrees, but soups—again, with exceptions—are usually only eaten hot A cultural example are soft drinks In North America it is almost always preferred cold, regardless of season In South America soda is almost exclusively consumed lukewarm in wintercitation needed, though not in Brazil, where it is only consumed cold

Starchinessedit

A 2016 study suggested that humans can taste starch specifically, a glucose oligomer independently of other tastes such as sweetness However, no specific chemical receptor has been yet been found for this taste100101

Nerve supply and neural connectionsedit

This diagram linearly unless otherwise mentioned tracks the projections of all known structures that allow for taste to their relevant endpoints in the human brain

The glossopharyngeal nerve innervates a third of the tongue including the circumvallate papillae The facial nerve innervates the other two thirds of the tongue and the cheek via the chorda tympani102

The pterygopalatine ganglia are ganglia one on each side of the soft palate The greater petrosal, lesser palatine and zygomatic nerves all synapse here The greater petrosal, carries soft palate taste signals to the facial nerve The lesser palatine sends signals to the nasal cavity; which is why spicy foods cause nasal drip The zygomatic sends signals to the lacrimal nerve that activate the lacrimal gland; which is the reason that spicy foods can cause tears Both the lesser palatine and the zygomatic are maxillary nerves from the trigeminal nerve

The special visceral afferents of the vagus nerve carry taste from the epiglottal region of the tongue

The lingual nerve trigeminal, not shown in diagram is deeply interconnected with chorda tympani in that it provides all other sensory info from the ⅔ of the tongue103 This info is processed separately nearby in rostal lateral subdivision of nucleus of the solitary tract NST

NST receives input from the amygdala regulates oculomotor nuclei output, bed nuclei of stria terminalis, hypothalamus, and prefrontal cortex NST is the topographical map that processes gustatory and sensory temp, texture, etc info104

Reticular formation includes Raphe nuclei responsible for serotonin production is signaled to release serotonin during and after a meal to suppress appetite105 Similarly, salivary nuclei are signaled to decrease saliva secretion

Hypoglossal and thalamic connections aid in oral-related movements

Hypothalamus connections hormonally regulate hunger and the digestive system

Substantia innominata connects the thalamus, temporal lobe, and insula

Edinger-Westphal nucleus reacts to taste stimuli by dilating and constricting the pupils106

Spinal ganglion are involved in movement

The frontal operculum is speculated to be the memory and association hub for tastecitation needed

The insula cortex aids in swallowing and gastric motility107108

Other conceptsedit

Taste as a philosophical conceptedit

See also: Philosophy of perception

Taste can be objective in terms of the five tastes sweet, salt, sour, bitter, and umami but it can also be subjective in terms of what we deem "good" and "bad" Taste is "subjective, objective, and qualitative"109 In terms of it being a philosophical concept, taste is hard to define because it is essentially subjective when pertaining to the personal preferences of individuals ie "'de gustibus non est disputandum' there is no disputing taste"109 We cannot tell someone they do not think something tastes good because we do not agree, and vice versa In order to evaluate taste in this context, we must explore all the ways in which taste can be defined According to Alan Weiss, taste fulfills the purpose of six functions: taste is the tool in which we use to define flavor; it is also flavor and how we categorize flavor sweet or salty; it is the preference, we as the tastemakers, place on specific flavors and our demand for those flavors; it is whether we choose to like or dislike a certain taste and therefore allow it into our general society of acceptable tastes or exile it; it is the value in which we place on certain taste one might believe one's taste in Bach or Rothko earns one capital; and lastly, with good judgement comes good taste and therefore, one with expressively good taste are expected to have good judgement, just as those in bad taste are expected to be in bad judgement 110

Supertastersedit

Main article: Supertaster

A supertaster is a person whose sense of taste is significantly more sensitive than average The cause of this heightened response is likely, at least in part, due to an increased number of fungiform papillae111 Studies have shown that supertasters require less fat and sugar in their food to get the same satisfying effects However, contrary to what one might think, these people actually tend to consume more salt than the average person This is due to their heightened sense of the taste of bitterness, and the presence of salt drowns out the taste of bitterness This also explains why supertasters prefer salted cheddar cheese over non-salted112

Aftertasteedit

Main article: 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 they can contain certain artificial flavor compounds, such as aspartame artificial sweetener

Acquired tasteedit

Main article: Acquired taste

An acquired taste often refers to an appreciation for a food or beverage that is unlikely to be enjoyed 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

Clinical significanceedit

Patients with Addison's disease, pituitary insufficiency, or cystic fibrosis sometimes have a hyper-sensitivity to the five primary tastes113

Disorders of tasteedit

  • ageusia complete loss of taste
  • hypogeusia reduced sense of taste
  • dysgeusia distortion in sense of taste
  • hypergeusia abnormally heightened sense of taste

Historyedit

In the West, Aristotle postulated in c 350 BCE114 that the two most basic tastes were sweet and bitter115 He was one of the first to develop a list of basic tastes116

Ayurveda, an ancient Indian healing science, has its own tradition of basic tastes, comprising sweet, salty, sour, pungent, bitter & astringent18

Similarly, the Ancient Chinese regarded spiciness as a basic taste

Researchedit

The receptors for the basic tastes of bitter, sweet and umami have been identified They are G protein-coupled receptors117 The cells that detect sourness have been identified as a subpopulation that express the protein PKD2L1 The responses are mediated by an influx of protons into the cells but the receptor for sour is still unknown The receptor for amiloride-sensitive attractive salty taste in mice has been shown to be a sodium channel118 There is some evidence for a sixth taste that senses fatty substances119

In 2010, researchers found bitter taste receptors in lung tissue, which cause airways to relax when a bitter substance is encountered They believe this mechanism is evolutionarily adaptive because it helps clear lung infections, but could also be exploited to treat asthma and chronic obstructive pulmonary disease120

See alsoedit

  • Food portal
  • Beefy meaty peptide
  • Digital lollipop
  • Optimal foraging theory
  • Palatability
  • Vomeronasal organ
  • Sensory analysis
  • Tea tasting
  • Wine tasting

Notesedit

Footnotesedit

a ^ It has been known for some time that these categories may not be comprehensive In Guyton's 1976 edition of Textbook of Medical Physiology, he wrote:

On the basis of physiologic studies, there are generally believed to be at least four primary sensations of taste: sour, salty, sweet, and bitter Yet we know that a person can perceive literally hundreds of different tastes These are all supposed to be combinations of the four primary sensationsHowever, there might be other less conspicuous classes or subclasses of primary sensations",121

b ^ Some variation in values is not uncommon between various studies Such variations may arise from a range of methodological variables, from sampling to analysis and interpretation In fact there is a "plethora of methods"122 Indeed, the taste index of 1, assigned to reference substances such as sucrose for sweetness, hydrochloric acid for sourness, quinine for bitterness, and sodium chloride for saltiness, is itself arbitrary for practical purposes59

Some values, such as those for maltose and glucose, vary little Others, such as aspartame and sodium saccharin, have much larger variation Regardless of variation, the perceived intensity of substances relative to each reference substance remains consistent for taste ranking purposes The indices table for McLaughlin & Margolskee 1994 for example,21123 is essentially the same as that of Svrivastava & Rastogi 2003,124 Guyton & Hall 2006,59 and Joesten et al 200756 The rankings are all the same, with any differences, where they exist, being in the values assigned from the studies from which they derive

As for the assignment of 1 or 100 to the index substances, this makes no difference to the rankings themselves, only to whether the values are displayed as whole numbers or decimal points Glucose remains about three-quarters as sweet as sucrose whether displayed as 75 or 075

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  122. ^ Macbeth, Helen M; MacClancy, Jeremy, eds 2004, "plethora of methods characterising human taste perception", Researching Food Habits: Methods and Problems, The anthropology of food and nutrition, Vol 5, New York: Berghahn Books, pp 87–88, ISBN 1-57181-544-9, retrieved 15 September 2010=  Paperback ISBN 1-57181-545-7 
  123. ^ McLaughlin, Susan, & Margolskee, Rorbert F November–December 1994, The Sense of Taste American Scientist, 82 6, pp 538–545 CS1 maint: Multiple names: authors list link
  124. ^ Svrivastava, RC & Rastogi, RP 2003, "Relative taste indices of some substances", in , Transport Mediated by Electrical Interfaces, Studies in interface science, vol18, Amsterdam, Netherlands: Elsevier Science, ISBN 0-444-51453-8, retrieved 12 September 2010  Taste indices of table 9, p274 are select sample taken from table in Guyton's Textbook of Medical Physiology present in all editions 

Further readingedit

  • The Science of taste at Kitchen Geekery An informative article about the science behind taste Written from a culinary science perspective
  • Bartoshuk, Linda M June 1978, "The Psychophysics of Taste" PDF, American Journal of Clinical Nutrition, 31 6: 1068–1077, PMID 352127, retrieved 12 September 2010 
  • Chandrashekar, Jayaram; Hoon, Mark A; Ryba , Nicholas J P & Zuker, Charles S 16 November 2006, "The receptors and cells for mammalian taste" PDF, Nature, 444 7117: 288–294, doi:101038/nature05401, PMID 17108952, retrieved 13 September 2010 CS1 maint: Multiple names: authors list link
  • Chaudhari, Nirupa & Roper, Stephen D 2010, "The cell biology of taste" PDF, Journal of Cell Biology, 190 3: 285–296, doi:101083/jcb201003144, PMC 2922655 , PMID 20696704, retrieved 13 September 2010 
  • Danker, WH 1968, Basic Principles of Sensory Evaluation, Philadelphia: American Society for Testing and Materials, ISBN 978-0-8031-4572-6, retrieved 13 September 2010 
  • Dulac, Catherine 17 March 2000, "The Physiology of Taste, Vintage 2000" PDF, Cell, 100 6: 607–610, doi:101016/S0092-86740080697-2, PMID 10761926, retrieved 13 September 2010 
  • Finger, Thomas E, ed 2009, International Symposium on Olfaction and Taste, Boston: Blackwell, for the New York Academy of Sciences, ISBN 1-57331-738-1, retrieved 12 September 2010  Alternative ISBN 978-1-57331-738-2
  • Hui, YH, ed 2010, Handbook of Fruit and Vegetable Flavors, Hoboken, New Jersey: John Wiley & Sons, ISBN 978-0-470-22721-3, retrieved 13 September 2010  See especially comments and key references in regards taste 
  • Thomas Hummel; Antje Welge-Lüssen, eds 2006, Taste and Smell: An Update, Advances in Oto-Rhino-Laryngolog, Vol63, Basel, Switzerland: Karger, ISBN 3-8055-8123-8, retrieved 12 September 2010 
  • Lawless, Harry T, & Heymann, Hildegarde 1998, Sensory Evaluation of Food: Principles and Practices, New York: Kluwer Academic/Plenum Publishers, ISBN 0-8342-1752-X, retrieved 13 September 2010 CS1 maint: Multiple names: authors list link
  • Macbeth, Helen, ed 2006, Food Preferences and Taste: Continuity and Change, The Anthropology of Food and Nutrition, Vol2, Providence, Rhode Island: Berghahn Books, ISBN 1-57181-958-4, retrieved 12 September 2010  Paperback ISBN 1-57181-970-3
  • PATTON HD 1950 "Physiology of smell and taste" Annual Review of Physiology 12 1: 469–84 doi:101146/annurevph12030150002345 PMID 15411178 
  • Reed, Danielle R; Tanaka, Toshiko & McDaniel, Amanda H 30 June 2006, "Diverse tastes: Genetics of sweet and bitter perception", Physiology & Behavior, 88 3: 215–226, doi:101016/jphysbeh200605033, PMC 1698869 , PMID 16782140 
  • Reineccius, Gary, ed 1999, Source Book of Flavours 2nd ed, Gaithersburg, Maryland: Aspen, ISBN 0-8342-1307-9, retrieved 12 September 2010  Previously published 1994 by Chapman & Hall, New York ISBN 0-442-00376-5 
  • Schiffman SS May 1983 "Taste and smell in disease first of two parts" The New England Journal of Medicine 308 21: 1275–9 doi:101056/NEJM198305263082107 PMID 6341841 
  • Schiffman SS June 1983 "Taste and smell in disease second of two parts" The New England Journal of Medicine 308 22: 1337–43 doi:101056/NEJM198306023082207 PMID 6341845 
  • Schiffman SS, Graham BG June 2000 "Taste and smell perception affect appetite and immunity in the elderly" European Journal of Clinical Nutrition 54 Suppl 3: S54–63 doi:101038/sjejcn1601026 PMID 11041076 
  • Seiden, Allen M, ed 1997, Taste and Smell Disorders, Rhinology and Sinusology, New York: Thieme, ISBN 0-86577-533-8, retrieved 12 September 2010  Alternative ISBN 3-13-107261-X
  • Shallenberger, RS 1993, Taste Chemistry, London & New York: Blackie Academic & Professional imprint of Chapman & Hall, ISBN 0-7514-0150-1, retrieved 12 September 2010 
  • Svrivastava, RC & Rastogi, RP 2003, "Relative taste indices of some substances", in , Transport Mediated by Electrical Interfaces, Studies in interface science, vol18, Amsterdam, Netherlands: Elsevier Science, ISBN 0-444-51453-8, retrieved 12 September 2010  Taste indices of table 9, p274 are select sample taken from table in Guyton's Textbook of Medical Physiology present in all editions 
  • Li X, Staszewski L, Xu H, Durick K, Zoller M, Adler E April 2002 "Human receptors for sweet and umami taste" Proceedings of the National Academy of Sciences of the United States of America 99 7: 4692–6 doi:101073/pnas072090199 PMC 123709  PMID 11917125 

External linksedit

  • Researchers Define Molecular Basis of Human "Sweet Tooth" and Umami Taste
  • Statistics on Taste at National Institute on Deafness and Other Communication Disorders An informative overview with good list of references
  • The Science of taste at Kitchen Geekery An informative article about the science behind taste Written from a culinary science perspective

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