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The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the
mouth In animal anatomy, the mouth, also known as the oral cavity, or in Latin cavum oris, is the opening through which many animals take in food and issue vocal sounds. It is also the cavity lying at the upper end of the alimentary canal, bounded on ...
reacts chemically with taste receptor cells located on taste buds in the oral cavity, mostly on the tongue. Taste, along with
olfaction The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
and trigeminal nerve stimulation (registering texture, pain, and temperature), determines
flavor Flavor or flavour is either the sensory perception of taste or smell, or a flavoring in food that produces such perception. Flavor or flavour may also refer to: Science *Flavors (programming language), an early object-oriented extension to Lis ...
s of
food Food is any substance consumed by an organism for nutritional support. Food is usually of plant, animal, or fungal origin, and contains essential nutrients, such as carbohydrates, fats, proteins, vitamins, or minerals. The substance is inge ...
and other substances. Humans have taste receptors on taste buds and other areas, including the upper surface of the tongue and the epiglottis. The gustatory cortex is responsible for the perception of taste. The tongue is covered with thousands of small bumps called
papillae Papilla (Latin, 'nipple') or papillae may refer to: In animals * Papilla (fish anatomy), in the mouth of fish * Basilar papilla, a sensory organ of lizards, amphibians and fish * Dental papilla, in a developing tooth * Dermal papillae, part of ...
, which are visible to the naked eye. Within each papilla are hundreds of taste buds. The exception to this is the filiform papillae that do not contain taste buds. There are between 2000 and 5000Boron, W.F., E.L. Boulpaep. 2003. Medical Physiology. 1st ed. Elsevier Science USA. 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. Taste receptors in the mouth sense the five taste modalities:
sweetness Sweetness is a Taste#Basic tastes, basic taste most commonly Perception, perceived when eating foods rich in sugars. Sweet tastes are generally regarded as pleasure, pleasurable. In addition to sugars like sucrose, many other chemical compounds ...
, sourness, saltiness, bitterness, and savoriness (also known as ''savory'' or ''umami''). Scientific experiments have demonstrated that these five tastes exist and are distinct from one another. Taste buds are able to distinguish between different tastes through detecting interaction with different molecules or ions. Sweet, savoriness, 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 The alkali metals consist of the chemical elements lithium (Li), sodium (Na), potassium (K),The symbols Na and K for sodium and potassium are derived from their Latin names, ''natrium'' and ''kalium''; these are still the origins of the names ...
or hydrogen ions enter taste buds, respectively. The basic taste modalities contribute only partially to the sensation and flavor of food in the mouth—other factors include
smell Smell may refer to; * Odor, airborne molecules perceived as a scent or aroma * Sense of smell, the scent also known scientifically as olfaction * "Smells" (''Bottom''), an episode of ''Bottom'' * The Smell, a music venue in Los Angeles, Californ ...
, detected by the olfactory epithelium of the nose; texture, detected through a variety of
mechanoreceptor A mechanoreceptor, also called mechanoceptor, is a sensory receptor that responds to mechanical pressure or distortion. Mechanoreceptors are innervated by sensory neurons that convert mechanical pressure into electrical signals that, in animals, ...
s, muscle nerves, etc.; temperature, detected by
thermoreceptor A thermoreceptor is a non-specialised sense receptor, or more accurately the receptive portion of a sensory neuron, that codes absolute and relative changes in temperature, primarily within the innocuous range. In the mammalian peripheral nervous s ...
s; and "coolness" (such as of
menthol Menthol is an organic compound, more specifically a monoterpenoid, made synthetically or obtained from the oils of corn mint, peppermint, or other mints. It is a waxy, clear or white crystalline substance, which is solid at room temperature and ...
) and "hotness" ( pungency), through chemesthesis. As the gustatory system senses both harmful and beneficial things, all basic taste modalities are classified as either aversive or appetitive, depending upon the effect the things they sense have on the body.Why do two great tastes sometimes not taste great together?
scientificamerican.com. Dr. Tim Jacob, Cardiff University. 22 May 2009.
Sweetness helps to identify energy-rich foods, while bitterness serves as a warning sign of poisons. Among humans, taste perception begins to fade at an older age because of loss of tongue papillae and a general decrease in
saliva Saliva (commonly referred to as spit) is an extracellular fluid produced and secreted by salivary glands in the mouth. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells (from which DNA can be ...
production. Humans can also have distortion of tastes ( dysgeusia). Not all
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s share the same taste modalities: some rodents can taste
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...
(which humans cannot), cats cannot taste sweetness, and several other carnivores including
hyena Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the clas ...
s, dolphins, and sea lions, have lost the ability to sense up to four of their ancestral five taste modalities.


Basic tastes

The gustatory system allows animals 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 Papilla (Latin, 'nipple') or papillae may refer to: In animals * Papilla (fish anatomy), in the mouth of fish * Basilar papilla, a sensory organ of lizards, amphibians and fish * Dental papilla, in a developing tooth * Dermal papillae, part of ...
, which are visible to the naked eye. Within each papilla are hundreds of taste buds. The exception to this are the filiform papillae that do not contain taste buds. There are between 2000 and 5000 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 five specific tastes received by taste receptors are saltiness,
sweetness Sweetness is a Taste#Basic tastes, basic taste most commonly Perception, perceived when eating foods rich in sugars. Sweet tastes are generally regarded as pleasure, pleasurable. In addition to sugars like sucrose, many other chemical compounds ...
, bitterness, sourness, and savoriness, often known by its Japanese name which translates to 'deliciousness'. As of the early 20th century, Western physiologists and psychologists believed there were four basic tastes: sweetness, sourness, saltiness, and bitterness. The concept of a "savory" taste was not present in Western science at that time, but was postulated in Japanese research.; a partial translation from By the end of the 20th century, the concept of umami was becoming familiar to Western society. One study found that both salt and sour taste mechanisms detect, in different ways, the presence of
sodium chloride Sodium chloride , commonly known as salt (although sea salt also contains other chemical salts), is an ionic compound with the chemical formula NaCl, representing a 1:1 ratio of sodium and chloride ions. With molar masses of 22.99 and 35.45 g ...
(salt) in the mouth. However, acids are also detected and perceived as sour. 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 blood. 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. Sweet taste signals the presence of
carbohydrates In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or may ...
in solution. Since carbohydrates have a very high
calorie The calorie is a unit of energy. For historical reasons, two main definitions of "calorie" are in wide use. The large calorie, food calorie, or kilogram calorie was originally defined as the amount of heat needed to raise the temperature of on ...
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 Glycogen is a multibranched polysaccharide of glucose that serves as a form of energy storage in animals, fungi, and bacteria. The polysaccharide structure represents the main storage form of glucose in the body. Glycogen functions as one o ...
). However, there are many non-carbohydrate molecules that trigger a sweet response, leading to the development of many artificial sweeteners, including saccharin,
sucralose Sucralose is an artificial sweetener and sugar substitute. The majority of ingested sucralose is not broken down by the body, so it is noncaloric. In the European Union, it is also known under the E number E955. It is produced by chlorination of ...
, and
aspartame Aspartame is an artificial non-saccharide sweetener 200 times sweeter than sucrose and is commonly used as a sugar substitute in foods and beverages. It is a methyl ester of the aspartic acid/phenylalanine dipeptide with the trade names ...
. It is still unclear how these substances activate the sweet receptors and what adaptational significance this has had. The savory taste (known in Japanese as ) was identified by Japanese chemist Kikunae Ikeda, which signals the presence of the amino acid
L-glutamate Glutamic acid (symbol Glu or E; the ionic form is known as glutamate) is an α-amino acid that is used by almost all living beings in the biosynthesis of proteins. It is a non-essential nutrient for humans, meaning that the human body can synt ...
, triggers a pleasurable response and thus encourages the intake of
peptides Peptides (, ) are short chains of amino acids linked by peptide bonds. Long chains of amino acids are called proteins. Chains of fewer than twenty amino acids are called oligopeptides, and include dipeptides, tripeptides, and tetrapeptides. A p ...
and
proteins Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
. The amino acids in proteins are used in the body to build muscles and organs, transport molecules ( hemoglobin),
antibodies An antibody (Ab), also known as an immunoglobulin (Ig), is a large, Y-shaped protein used by the immune system to identify and neutralize foreign objects such as pathogenic bacteria and viruses. The antibody recognizes a unique molecule of the ...
, and the organic catalysts known as
enzymes Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different molecule ...
. 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. Pungency (piquancy or hotness) had traditionally been considered a sixth basic taste. In 2015, researchers suggested a new basic taste of fatty acids called 'fat taste', although 'oleogustus' and 'pinguis' have both been proposed as alternate terms.


Sweetness

Sweetness, usually regarded as a pleasurable sensation, is produced by the presence of
sugar Sugar is the generic name for sweet-tasting, soluble carbohydrates, many of which are used in food. Simple sugars, also called monosaccharides, include glucose, fructose, and galactose. Compound sugars, also called disaccharides or double ...
s and substances that mimic sugar. Sweetness may be connected to aldehydes and
ketone In organic chemistry, a ketone is a functional group with the structure R–C(=O)–R', where R and R' can be a variety of carbon-containing substituents. Ketones contain a carbonyl group –C(=O)– (which contains a carbon-oxygen double bo ...
s, which contain a carbonyl group. Sweetness is detected by a variety of G protein coupled receptors (GPCR) coupled to the G protein
gustducin Gustducin is a G protein associated with taste and the gustatory system, found in some taste receptor cells. Research on the discovery and isolation of gustducin is recent. It is known to play a large role in the transduction of bitter, sweet a ...
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 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 animals.
Taste detection threshold Taste detection threshold is the minimum concentration of a flavoured substance detectable by the sense of taste. Sweetness detection thresholds are usually measured relative to that of sucrose, sourness relative to dilute hydrochloric acid, salti ...
s for sweet substances are rated relative to
sucrose Sucrose, a disaccharide, is a sugar composed of glucose and fructose subunits. It is produced naturally in plants and is the main constituent of white sugar. It has the molecular formula . For human consumption, sucrose is extracted and refined ...
, which has an index of 1. The average human detection threshold for sucrose is 10 millimoles per liter. For
lactose Lactose is a disaccharide sugar synthesized by galactose and glucose subunits and has the molecular formula C12H22O11. Lactose makes up around 2–8% of milk (by mass). The name comes from ' (gen. '), the Latin word for milk, plus the suffix '' - ...
it is 30 millimoles per liter, with a sweetness index of 0.3, and
5-nitro-2-propoxyaniline 5-Nitro-2-propoxyaniline, also known as P-4000 and Ultrasüss, is about 4,000 times the intensity of sucrose (hence its alternate name, P-4000). It is an orange solid that is only slightly soluble in water. It is stable in boiling water and dilu ...
0.002 millimoles per liter. “Natural” sweeteners such as saccharides activate the GPCR, which releases
gustducin Gustducin is a G protein associated with taste and the gustatory system, found in some taste receptor cells. Research on the discovery and isolation of gustducin is recent. It is known to play a large role in the transduction of bitter, sweet a ...
. The gustducin then activates the molecule adenylate cyclase, which catalyzes the production of the molecule
cAMP Camp may refer to: Outdoor accommodation and recreation * Campsite or campground, a recreational outdoor sleeping and eating site * a temporary settlement for nomads * Camp, a term used in New England, Northern Ontario and New Brunswick to descri ...
, 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.


Sourness

Sourness is the taste that detects
acid In computer science, ACID ( atomicity, consistency, isolation, durability) is a set of properties of database transactions intended to guarantee data validity despite errors, power failures, and other mishaps. In the context of databases, a sequ ...
ity. 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. Sour taste is detected by a small subset of cells that are distributed across all taste buds called Type III taste receptor cells. H+ ions (
proton A proton is a stable subatomic particle, symbol , H+, or 1H+ with a positive electric charge of +1 ''e'' elementary charge. Its mass is slightly less than that of a neutron and 1,836 times the mass of an electron (the proton–electron mass ...
s) that are abundant in sour substances can directly enter the Type III taste cells through a proton channel. This channel was identified in 2018 as otopetrin 1 (OTOP1). The transfer of positive charge into the cell can itself trigger an electrical response. Some weak acids such as acetic acid, can also penetrate taste cells; intracellular hydrogen ions inhibit potassium channels, which normally function to hyperpolarize the cell. By a combination of direct intake of hydrogen ions through OTOP1 ion channels (which itself depolarizes the cell) and the inhibition of the hyperpolarizing channel, sourness causes the taste cell to fire action potentials and release neurotransmitter. The most common foods with natural sourness are fruits, such as lemon, lime, grape, orange, tamarind, and bitter melon. Fermented foods, such as wine, vinegar or yogurt, may have sour taste. Children show a greater enjoyment of sour flavors than adults, and sour candy containing citric acid or malic acid is common.


Saltiness

The simplest receptor found in the mouth is the
sodium chloride Sodium chloride , commonly known as salt (although sea salt also contains other chemical salts), is an ionic compound with the chemical formula NaCl, representing a 1:1 ratio of sodium and chloride ions. With molar masses of 22.99 and 35.45 g ...
(salt) receptor. Saltiness is a taste produced primarily by the presence of
sodium ion Sodium is a chemical element with the symbol Na (from Latin ''natrium'') and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 of the periodic table. Its only stable is ...
s. 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
cation An ion () is an atom or molecule with a net electrical charge. The charge of an electron is considered to be negative by convention and this charge is equal and opposite to the charge of a proton, which is considered to be positive by convent ...
s 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 A neurotransmitter is a signaling molecule secreted by a neuron to affect another cell across a synapse. The cell receiving the signal, any main body part or target cell, may be another neuron, but could also be a gland or muscle cell. Neuro ...
release. This sodium channel is known as an
epithelial sodium channel The epithelial sodium channel (ENaC), (also known as amiloride-sensitive sodium channel) is a membrane-bound ion channel that is selectively permeable to sodium ions (). It is assembled as a heterotrimer composed of three homologous subunits α ...
(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 Rubidium is the chemical element with the symbol Rb and atomic number 37. It is a very soft, whitish-grey solid in the alkali metal group, similar to potassium and caesium. Rubidium is the first alkali metal in the group to have a density higher ...
and
caesium Caesium (IUPAC spelling) (or cesium in American English) is a chemical element with the symbol Cs and atomic number 55. It is a soft, silvery-golden alkali metal with a melting point of , which makes it one of only five elemental metals that a ...
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 Potassium chloride (KCl, or potassium salt) is a metal halide salt composed of potassium and chlorine. It is odorless and has a white or colorless vitreous crystal appearance. The solid dissolves readily in water, and its solutions have a salt ...
(KCl), is the principal ingredient in salt substitutes and has a saltiness index of 0.6. Other monovalent cations, e.g.
ammonium The ammonium cation is a positively-charged polyatomic ion with the chemical formula or . It is formed by the protonation of ammonia (). Ammonium is also a general name for positively charged or protonated substituted amines and quaternary a ...
(NH4+), and divalent cations of the alkali earth metal group of the
periodic table The periodic table, also known as the periodic table of the (chemical) elements, is a rows and columns arrangement of the chemical elements. It is widely used in chemistry, physics, and other sciences, and is generally seen as an icon of ch ...
, 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. But the chloride of calcium is saltier and less bitter than potassium chloride, and is commonly used in pickle brine instead of KCl.


Bitterness

Bitterness is one of 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 Cocoa may refer to: Chocolate * Chocolate * ''Theobroma cacao'', the cocoa tree * Cocoa bean, seed of ''Theobroma cacao'' * Chocolate liquor, or cocoa liquor, pure, liquid chocolate extracted from the cocoa bean, including both cocoa butter and ...
, South American
mate Mate may refer to: Science * Mate, one of a pair of animals involved in: ** Mate choice, intersexual selection ** Mating * Multi-antimicrobial extrusion protein, or MATE, an efflux transporter family of proteins Person or title * Friendship ...
,
coca tea Coca tea, also called mate de coca, is an herbal tea (infusion) made using the raw or dried leaves of the coca plant, which is native to South America. It is made either by submerging the coca leaf or dipping a tea bag in hot water. The tea is mo ...
, bitter gourd, uncured olives, citrus peel, some varieties of
cheese Cheese is a dairy product produced in wide ranges of flavors, textures, and forms by coagulation of the milk protein casein. It comprises proteins and fat from milk, usually the milk of cows, buffalo, goats, or sheep. During production, ...
, many plants in the family Brassicaceae, dandelion greens, horehound, wild chicory, and
escarole Endive () is a leaf vegetable belonging to the genus ''Cichorium'', which includes several similar bitter-leafed vegetables. Species include '' Cichorium endivia'' (also called endive), ''Cichorium pumilum'' (also called wild endive), and ''Cich ...
. The ethanol in alcoholic beverages tastes bitter, as do the additional bitter ingredients found in some alcoholic beverages including
hops Hops are the flowers (also called seed cones or strobiles) of the hop plant ''Humulus lupulus'', a member of the Cannabaceae family of flowering plants. They are used primarily as a bittering, flavouring, and stability agent in beer, to whi ...
in beer and gentian 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 researchersLogue, A.W. (1986) ''The Psychology of Eating and Drinking''. New York: W.H. Freeman & Co. 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, and among 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.Jones, S., Martin, R., & Pilbeam, D. (1994) ''The Cambridge Encyclopedia of Human Evolution''. Cambridge: Cambridge University Press Amongst humans, various
food processing Food processing is the transformation of agricultural products into food, or of one form of food into other forms. Food processing includes many forms of processing foods, from grinding grain to make raw flour to home cooking to complex industr ...
techniques are used worldwide to detoxify otherwise inedible foods and make them palatable. 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 species. The threshold for stimulation of bitter taste by quinine averages a concentration of 8 μ M (8 micromolar).Guyton, Arthur C. (1991) ''Textbook of Medical Physiology''. (8th ed). Philadelphia: W.B. Saunders The taste thresholds of other bitter substances are rated relative to quinine, which is thus given a reference index of 1. For example,
brucine Brucine, is an alkaloid closely related to strychnine, most commonly found in the ''Strychnos nux-vomica'' tree. Brucine poisoning is rare, since it is usually ingested with strychnine, and strychnine is more toxic than brucine. In synthetic ch ...
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 natural substance is amarogentin, a compound present in the roots of the plant '' Gentiana lutea'', and the most bitter substance known is the synthetic chemical denatonium, which has an index of 1,000. It is used as an
aversive agent Aversive agents are unpleasantly flavored substances added to poisonous household goods to discourage children and animals from consuming them. Aversive agents are not intended to be harmful, only unpleasant. For example, to prevent children from co ...
(a bitterant) that is added to toxic substances to prevent accidental ingestion. It was discovered accidentally in 1958 during research on a local anesthetic, by MacFarlan Smith of Gorgie, Edinburgh, Scotland. Research has shown that TAS2Rs (taste receptors, type 2, also known as T2Rs) such as
TAS2R38 Taste receptor 2 member 38 is a protein that in humans is encoded by the ''TAS2R38'' gene. TAS2R38 is a bitter taste receptor; varying genotypes of ''TAS2R38'' influence the ability to taste both 6-''n''-propylthiouracil (PROP) and phenylthiocar ...
coupled to the G protein
gustducin Gustducin is a G protein associated with taste and the gustatory system, found in some taste receptor cells. Research on the discovery and isolation of gustducin is recent. It is known to play a large role in the transduction of bitter, sweet a ...
are responsible for the human ability to taste bitter substances. They are identified not only by their ability to taste for certain "bitter"
ligands In coordination chemistry, a ligand is an ion or molecule (functional group) that binds to a central metal atom to form a coordination complex. The bonding with the metal generally involves formal donation of one or more of the ligand's electro ...
, but also by the morphology of the receptor itself (surface bound, monomeric). 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 compounds. Over 670 bitter-tasting compounds have been identified, on a bitter database, of which over 200 have been assigned to one or more specific receptors. 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 "
supertaster A supertaster is a person whose sense of taste is of far greater intensity than the average person. Some studies also show that an increased sensitivity to bitter tastes may be a cause of selective eating. History The term originated with ...
s" 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. 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 channels. Also, this secondary messenger can stimulate the
endoplasmic reticulum The endoplasmic reticulum (ER) is, in essence, the transportation system of the eukaryotic cell, and has many other important functions such as protein folding. It is a type of organelle made up of two subunits – rough 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.


Umami

Umami, or savory, is an appetitive taste. It can be tasted in
soy sauce Soy sauce (also called simply soy in American English and soya sauce in British English) is a liquid condiment of Chinese origin, traditionally made from a fermented paste of soybeans, roasted grain, brine, and '' Aspergillus oryzae'' or ''Asp ...
,
meat Meat is animal flesh that is eaten as food. Humans have hunted, farmed, and scavenged animals for meat since prehistoric times. The establishment of settlements in the Neolithic Revolution allowed the domestication of animals such as chic ...
, dashi and consomme. A loanword from Japanese meaning "good flavor" or "good taste", is considered fundamental to many East Asian cuisines, such as Japanese cuisine. It dates back to the use of fermented fish sauce: '' garum'' in ancient Rome and or in ancient China. Umami was first studied in 1907 by Ikeda isolating dashi taste, which he identified as the chemical monosodium glutamate (MSG). MSG is a sodium salt that produces a strong savory taste, especially combined with foods rich in nucleotides such as meats, fish, nuts, and mushrooms. Some savory 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. L-glutamate may bond to a type of GPCR known as a metabotropic glutamate receptor (
mGluR4 Metabotropic glutamate receptor 4 is a protein that in humans is encoded by the ''GRM4'' gene. Together with GRM6, GRM7 and GRM8 it belongs to Metabotropic glutamate receptor#Group II & Group III, group III of the metabotropic glutamate receptor ...
) which causes the G-protein complex to activate the sensation of umami.


Measuring relative tastes

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 substances. Substances are usually measured relative to
sucrose Sucrose, a disaccharide, is a sugar composed of glucose and fructose subunits. It is produced naturally in plants and is the main constituent of white sugar. It has the molecular formula . For human consumption, sucrose is extracted and refined ...
, which is usually given an arbitrary index of 1 or 100. Rebaudioside A is 100 times sweeter than sucrose;
fructose Fructose, or fruit sugar, is a Ketose, ketonic monosaccharide, simple sugar found in many plants, where it is often bonded to glucose to form the disaccharide sucrose. It is one of the three dietary monosaccharides, along with glucose and galacto ...
is about 1.4 times sweeter; glucose, a sugar found in honey and vegetables, is about three-quarters as sweet; and
lactose Lactose is a disaccharide sugar synthesized by galactose and glucose subunits and has the molecular formula C12H22O11. Lactose makes up around 2–8% of milk (by mass). The name comes from ' (gen. '), the Latin word for milk, plus the suffix '' - ...
, a milk sugar, is one-half as sweet. The sourness of a substance can be rated by comparing it to very dilute hydrochloric acid (HCl). Relative saltiness can be rated by comparison to a dilute salt solution. 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.Quality control methods for medicinal plant materials, Pg. 38
World Health Organization, 1998.
More formal chemical analysis, while possible, is difficult. There may not be an absolute measure for pungency, though there are tests for measuring the subjective presence of a given pungent substance in food, such as the Scoville scale for capsaicine in peppers or the
Pyruvate scale The pyruvate scale measures pungency in onions and garlic with units of μmol/gfw (micromoles per gram fresh weight). It is named after pyruvic acid, the alpha-keto acid co-product created in the biochemical pathway that forms ''syn''-propanethial ...
for
pyruvate Pyruvic acid (CH3COCOOH) is the simplest of the alpha-keto acids, with a carboxylic acid and a ketone functional group. Pyruvate, the conjugate base, CH3COCOO−, is an intermediate in several metabolic pathways throughout the cell. Pyruvic aci ...
s in garlics and onions.


Functional structure

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 these receptors can detect which are recognized: salt, sweet, sour, bitter, and umami. Each type of 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, though each neuron responds most strongly to one tastant". Researchers 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." As well,
serotonin Serotonin () or 5-hydroxytryptamine (5-HT) is a monoamine neurotransmitter. Its biological function is complex and multifaceted, modulating mood, cognition, reward, learning, memory, and numerous physiological processes such as vomiting and vas ...
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 Microvilli (singular: microvillus) are microscopic cellular membrane protrusions that increase the surface area for diffusion and minimize any increase in volume, and are involved in a wide variety of functions, including absorption, secretion, ...
of the taste cells. ;Sweetness Sweetness is produced by the presence of
sugar Sugar is the generic name for sweet-tasting, soluble carbohydrates, many of which are used in food. Simple sugars, also called monosaccharides, include glucose, fructose, and galactose. Compound sugars, also called disaccharides or double ...
s, some proteins, and other substances such as alcohols like
anethol Anethole (also known as anise camphor) is an organic compound that is widely used as a flavoring substance. It is a derivative of phenylpropene, a type of aromatic compound that occurs widely in nature, in essential oils. It is in the class of phe ...
, glycerol and propylene glycol,
saponins Saponins (Latin "sapon", soap + "-in", one of), also selectively referred to as triterpene glycosides, are bitter-tasting usually toxic plant-derived organic chemicals that have a foamy quality when agitated in water. They are widely distributed ...
such as
glycyrrhizin Glycyrrhizin (or glycyrrhizic acid or glycyrrhizinic acid) is the chief sweet-tasting constituent of ''Glycyrrhiza glabra'' ( liquorice) root. Structurally, it is a saponin used as an emulsifier and gel-forming agent in foodstuffs and cosmet ...
, artificial sweeteners (organic compounds with a variety of structures), and lead compounds such as lead acetate. It is often connected to aldehydes and
ketone In organic chemistry, a ketone is a functional group with the structure R–C(=O)–R', where R and R' can be a variety of carbon-containing substituents. Ketones contain a carbonyl group –C(=O)– (which contains a carbon-oxygen double bo ...
s, which contain a carbonyl group. Many foods can be perceived as sweet regardless of their actual sugar content. For example, some plants such as
liquorice Liquorice (British English) or licorice (American English) ( ; also ) is the common name of ''Glycyrrhiza glabra'', a flowering plant of the bean family Fabaceae, from the root of which a sweet, aromatic flavouring can be extracted. The liqu ...
,
anise Anise (; '), also called aniseed or rarely anix is a flowering plant in the family Apiaceae native to Eurasia. The flavor and aroma of its seeds have similarities with some other spices and herbs, such as star anise, fennel, licorice, and ta ...
or
stevia Stevia () is a natural sweetener and sugar substitute derived from the leaves of the plant species ''Stevia rebaudiana'', native to Paraguay and Brazil. The active compounds are steviol glycosides (mainly stevioside and rebaudioside), which h ...
can be used as sweeteners. Rebaudioside A is a steviol glycoside coming from stevia that is 200 times sweeter than sugar. Lead acetate and other lead compounds were used as sweeteners, mostly for wine, until
lead poisoning Lead poisoning, also known as plumbism and saturnism, is a type of metal poisoning caused by lead in the body. The brain is the most sensitive. Symptoms may include abdominal pain, constipation, headaches, irritability, memory problems, inferti ...
became known. Romans used to deliberately boil the must inside of lead vessels to make a sweeter wine. 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 Gustducin is a G protein associated with taste and the gustatory system, found in some taste receptor cells. Research on the discovery and isolation of gustducin is recent. It is known to play a large role in the transduction of bitter, sweet a ...
. These receptors are T1R2+3 (heterodimer) and T1R3 (homodimer), which account for sweet sensing in humans and other animals. ;Saltiness Saltiness is a taste produced best by the presence of cations (such as , or ) and is directly detected by cation influx into glial like cells via leak channels causing depolarisation of the cell. Other monovalent cations, e.g.,
ammonium The ammonium cation is a positively-charged polyatomic ion with the chemical formula or . It is formed by the protonation of ammonia (). Ammonium is also a general name for positively charged or protonated substituted amines and quaternary a ...
, , and divalent cations of the alkali earth metal group of the
periodic table The periodic table, also known as the periodic table of the (chemical) elements, is a rows and columns arrangement of the chemical elements. It is widely used in chemistry, physics, and other sciences, and is generally seen as an icon of ch ...
, e.g., calcium, , ions, in general, elicit a bitter rather than a salty taste even though they, too, can pass directly through
ion channel Ion channels are pore-forming membrane proteins that allow ions to pass through the channel pore. Their functions include establishing a resting membrane potential, shaping action potentials and other electrical signals by gating the flow of io ...
s in the tongue. ;Sourness Sourness is
acid In computer science, ACID ( atomicity, consistency, isolation, durability) is a set of properties of database transactions intended to guarantee data validity despite errors, power failures, and other mishaps. In the context of databases, a sequ ...
ity, and, like salt, it is a taste sensed using
ion channel Ion channels are pore-forming membrane proteins that allow ions to pass through the channel pore. Their functions include establishing a resting membrane potential, shaping action potentials and other electrical signals by gating the flow of io ...
s.channels in sensory cells (Page 155/304)
Stephan Frings, Jonathan Bradley. Wiley-VCH, 2004.
Undissociated acid diffuses across the plasma membrane of a presynaptic cell, where it dissociates in accordance with
Le Chatelier's principle Le Chatelier's principle (pronounced or ), also called Chatelier's principle (or the Equilibrium Law), is a principle of chemistry used to predict the effect of a change in conditions on chemical equilibria. The principle is named after French c ...
. 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 sour. ;Bitterness Research has shown that TAS2Rs (taste receptors, type 2, also known as T2Rs) such as
TAS2R38 Taste receptor 2 member 38 is a protein that in humans is encoded by the ''TAS2R38'' gene. TAS2R38 is a bitter taste receptor; varying genotypes of ''TAS2R38'' influence the ability to taste both 6-''n''-propylthiouracil (PROP) and phenylthiocar ...
are responsible for the ability to taste bitter substances in vertebrates. 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). ;Savoriness The amino acid
glutamic acid Glutamic acid (symbol Glu or E; the ionic form is known as glutamate) is an α-amino acid that is used by almost all living beings in the biosynthesis of proteins. It is a non-essential nutrient for humans, meaning that the human body can synt ...
is responsible for savoriness,What Is Umami?: What Exactly is Umami?
Umami Information Center
but some nucleotides (
inosinic acid Inosinic acid or inosine monophosphate (IMP) is a nucleotide (that is, a nucleoside monophosphate). Widely used as a flavor enhancer, it is typically obtained from chicken byproducts or other meat industry waste. Inosinic acid is important in met ...
and
guanylic acid Guanosine monophosphate (GMP), also known as 5′-guanidylic acid or guanylic acid (conjugate base guanylate), is a nucleotide that is used as a monomer in RNA. It is an ester of phosphoric acid with the nucleoside guanosine. GMP consists of the ...
) can act as complements, enhancing the taste.What Is Umami?: The Composition of Umami
Umami Information Center
Glutamic acid binds to a variant of the G protein-coupled receptor, producing a
savory Savory or Savoury may refer to: Common usage * Herbs of the genus ''Satureja'', particularly: ** Summer savory (''Satureja hortensis''), an annual herb, used to flavor food ** Winter savory (''Satureja montana''), a perennial herb, also used to ...
taste.


Further sensations and transmission

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 hotness)

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 Ginger (''Zingiber officinale'') is a flowering plant whose rhizome, ginger root or ginger, is widely used as a spice and a folk medicine. It is a herbaceous perennial which grows annual pseudostems (false stems made of the rolled bases of lea ...
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 Indonesian is anything of, from, or related to Indonesia, an archipelagic country in Southeast Asia. It may refer to: * Indonesians, citizens of Indonesia ** Native Indonesians, diverse groups of local inhabitants of the archipelago ** Indonesian ...
, Lao, Malaysian,
Mexican Mexican may refer to: Mexico and its culture *Being related to, from, or connected to the country of Mexico, in North America ** People *** Mexicans, inhabitants of the country Mexico and their descendants *** Mexica, ancient indigenous people ...
, New Mexican, Pakistani, Singaporean, Southwest Chinese (including
Sichuan cuisine Sichuan cuisine, alternatively romanized as Szechwan cuisine or Szechuan cuisine (, Standard Mandarin pronunciation: ), is a style of Chinese cuisine originating from Sichuan Province. It has bold flavours, particularly the pungency and spicin ...
), Vietnamese, and
Thai Thai or THAI may refer to: * Of or from Thailand, a country in Southeast Asia ** Thai people, the dominant ethnic group of Thailand ** Thai language, a Tai-Kadai language spoken mainly in and around Thailand *** Thai script *** Thai (Unicode block ...
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.


Coolness

Some substances activate cold trigeminal receptors even when not at low temperatures. This "fresh" or "minty" sensation can be tasted in peppermint, spearmint and is triggered by substances such as
menthol Menthol is an organic compound, more specifically a monoterpenoid, made synthetically or obtained from the oils of corn mint, peppermint, or other mints. It is a waxy, clear or white crystalline substance, which is solid at room temperature and ...
,
anethol Anethole (also known as anise camphor) is an organic compound that is widely used as a flavoring substance. It is a derivative of phenylpropene, a type of aromatic compound that occurs widely in nature, in essential oils. It is in the class of phe ...
, ethanol, and
camphor Camphor () is a waxy, colorless solid with a strong aroma. It is classified as a terpenoid and a cyclic ketone. It is found in the wood of the camphor laurel ('' Cinnamomum camphora''), a large evergreen tree found in East Asia; and in the k ...
. 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.


Numbness

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 North Sumatra ( id, Sumatra Utara) is a province of Indonesia located on the northern part of the island of Sumatra. Its capital and largest city is Medan. North Sumatra is Indonesia's fourth most populous province after West Java, East Java and ...
often combine this with chili pepper to produce a 麻辣 ''málà'', "numbing-and-hot", or "mati rasa" flavor. Typical in northern Brazilian cuisine, jambu is an herb used in dishes like
tacacá Tacacá () is a typical dish of Northern Brazil (mostly consumed in Pará, Amazonas, Acre, Amapá and Roraima). It is made with ''jambu'' (a native variety of paracress), and ''tucupi'' (a broth made with wild manioc), cooked tapioca starch ...
. These sensations, although not taste, fall into a category of chemesthesis.


Astringency

Some foods, such as unripe fruits, contain tannins or
calcium oxalate Calcium oxalate (in archaic terminology, oxalate of lime) is a calcium salt of oxalic acid with the chemical formula . It forms hydrates , where ''n'' varies from 1 to 3. Anhydrous and all hydrated forms are colorless or white. The monohydrate ...
that cause an astringent or puckering sensation of the mucous membrane of the mouth. Examples include tea,
red wine Red wine is a type of wine made from dark-colored grape varieties. The color of the wine can range from intense violet, typical of young wines, through to brick red for mature wines and brown for older red wines. The juice from most purple grap ...
, or
rhubarb Rhubarb is the fleshy, edible stalks ( petioles) of species and hybrids (culinary rhubarb) of ''Rheum'' in the family Polygonaceae, which are cooked and used for food. The whole plant – a herbaceous perennial growing from short, thick rhizo ...
. Other terms for the astringent sensation are "dry", "rough", "harsh" (especially for wine), "tart" (normally referring to sourness), "rubbery", "hard" or "styptic".


Metallicness

A metallic taste may be caused by food and drink, certain medicines or
amalgam Amalgam most commonly refers to: * Amalgam (chemistry), mercury alloy * Amalgam (dentistry), material of silver tooth fillings ** Bonded amalgam, used in dentistry Amalgam may also refer to: * Amalgam Comics, a publisher * Amalgam Digital, an in ...
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 current. Some artificial sweeteners are perceived to have a metallic taste, which is detected by the TRPV1 receptors. Many people consider blood to have a metallic taste. 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 Dysgeusia, also known as parageusia, is a distortion of the sense of taste. Dysgeusia is also often associated with ageusia, which is the complete lack of taste, and hypogeusia, which is a decrease in taste sensitivity. An alteration in taste or ...
, referring to distortions of the sense of taste, and can be caused by medication, including saquinavir, zonisamide, and various kinds of chemotherapy, as well as occupational hazards, such as working with pesticides.


Fat taste

Recent research reveals a potential taste receptor called the CD36 receptor. CD36 was targeted as a possible lipid taste receptor because it binds to fat molecules (more specifically, long-chain fatty acids), and it has been localized to taste bud cells (specifically, the circumvallate and foliate
papillae Papilla (Latin, 'nipple') or papillae may refer to: In animals * Papilla (fish anatomy), in the mouth of fish * Basilar papilla, a sensory organ of lizards, amphibians and fish * Dental papilla, in a developing tooth * Dermal papillae, part of ...
). 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 fat. Although CD36 has been studied primarily in
mice A mouse ( : mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
, 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; 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 acids. Monovalent cation channel
TRPM5 Transient receptor potential cation channel subfamily M member 5 (TRPM5), also known as long transient receptor potential channel 5 is a protein that in humans is encoded by the ''TRPM5'' gene. Function TRPM5 is a calcium-activated non-selecti ...
has been implicated in fat taste as well, 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 savory. Proposed alternate names to fat taste include oleogustus and pinguis, although these terms are not widely accepted. 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 hydrolysed into fatty acids via lipases. 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 inedible. To demonstrate that individuals can distinguish fat taste from other tastes, 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 savory 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 eaten. There are few regularly consumed foods rich in fat taste, due to the negative flavor that is evoked in large quantities. Foods whose flavor to which fat taste makes a small contribution include olive oil and fresh butter, along with various kinds of vegetable and nut oils.


Heartiness

Kokumi (, Japanese: from ) is translated as "heartiness", "full flavor" or "rich" and describes compounds in food that do not have their own taste, but enhance the characteristics when combined. Alongside the five basic tastes of sweet, sour, salt, bitter and savory, ''
kokumi The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth biochemistry, reacts chemically with ta ...
'' has been described as something that may enhance the other five tastes by magnifying and lengthening the other tastes, or "mouthfulness". Garlic is a common ingredient to add flavor used to help define the characteristic ''kokumi'' flavors. Calcium-sensing receptors (CaSR) are receptors for "''kokumi''" substances. ''Kokumi'' substances, applied around taste pores, induce an increase in the intracellular Ca concentration in a subset of cells. This subset of CaSR-expressing taste cells are independent from the influenced basic taste receptor cells. CaSR agonists directly activate the CaSR on the surface of taste cells and integrated in the brain via the central nervous system. However, a basal level of calcium, corresponding to the physiological concentration, is necessary for activation of the CaSR to develop the ''kokumi'' sensation.


Calcium

The distinctive taste of chalk has been identified as the calcium component of that substance. In 2008, geneticists discovered a calcium receptor on the tongues of
mice A mouse ( : mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
. The CaSR receptor is commonly found in the
gastrointestinal tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organ (biology), organs of the digestive syste ...
, kidneys, and brain. Along with the "sweet" T1R3 receptor, the CaSR receptor can detect calcium as a taste. Whether the perception exists or not in humans is unknown.


Temperature

Temperature can be an essential element of the taste experience. Heat can accentuate some flavors and decrease others by varying the density and phase equilibrium of a substance. 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 A soft drink (see § Terminology for other names) is a drink that usually contains water (often carbonated), a sweetener, and a natural and/or artificial flavoring. The sweetener may be a sugar, high-fructose corn syrup, fruit juice, a s ...
. In North America it is almost always preferred cold, regardless of season.


Starchiness

A 2016 study suggested that humans can taste
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...
(specifically, a glucose
oligomer In chemistry and biochemistry, an oligomer () is a molecule that consists of a few repeating units which could be derived, actually or conceptually, from smaller molecules, monomers.Quote: ''Oligomer molecule: A molecule of intermediate relativ ...
) independently of other tastes such as sweetness. However, no specific chemical receptor has yet been found for this taste.


Nerve supply and neural connections

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 tympani. 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 afferent A Special visceral afferent fibers (SVA) is a afferent fiber that develop in association with the gastrointestinal tract. They carry the special senses of smell (olfaction) and taste (gustation). The cranial nerves containing SVA fibers are the olfa ...
s of the vagus nerve carry taste from the
epiglottal A pharyngeal consonant is a consonant that is articulated primarily in the pharynx. Some phoneticians distinguish upper pharyngeal consonants, or "high" pharyngeals, pronounced by retracting the root of the tongue in the mid to upper pharynx, ...
region of the tongue. The lingual nerve (trigeminal, not shown in diagram) is deeply interconnected with the chorda tympani in that it provides all other sensory info from the anterior ⅔ of the tongue. This info is processed separately (nearby) in the rostal lateral subdivision of the 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.) info. Reticular formation (includes Raphe nuclei responsible for serotonin production) is signaled to release serotonin during and after a meal to suppress appetite. Similarly, salivary nuclei are signaled to decrease saliva secretion.
Hypoglossal The hypoglossal nerve, also known as the twelfth cranial nerve, cranial nerve XII, or simply CN XII, is a cranial nerve that innervates all the extrinsic and intrinsic muscles of the tongue except for the palatoglossus, which is innervated by ...
and
thalamic The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all directions, ...
connections aid in oral-related movements. Hypothalamus connections hormonally regulate hunger and the digestive system.
Substantia innominata The substantia innominata also innominate substance, or substantia innominata of Meynert (Latin for unnamed substance) is a series of layers in the human brain consisting partly of gray and partly of white matter, which lies below the anterior part ...
connects the thalamus, temporal lobe, and insula. Edinger-Westphal nucleus reacts to taste stimuli by dilating and constricting the pupils. Spinal ganglion are involved in movement. The frontal operculum is speculated to be the memory and association hub for taste. The
insula cortex The insular cortex (also insula and insular lobe) is a portion of the cerebral cortex folded deep within the lateral sulcus (the fissure separating the temporal lobe from the parietal and frontal lobes) within each hemisphere of the mammalian bra ...
aids in swallowing and gastric motility.


Other concepts


Supertasters

A supertaster is a person whose sense of taste is significantly more sensitive than most. The cause of this heightened response is likely, at least in part, due to an increased number of
fungiform papillae Lingual papillae (singular papilla) are small structures on the upper surface of the tongue that give it its characteristic rough texture. The four types of papillae on the human tongue have different structures and are accordingly classified as c ...
. 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 most people. 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-salted.)


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 Aspartame is an artificial non-saccharide sweetener 200 times sweeter than sucrose and is commonly used as a sugar substitute in foods and beverages. It is a methyl ester of the aspartic acid/phenylalanine dipeptide with the trade names ...
(artificial sweetener).


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 bitterness, a strong or strange odor, taste, or appearance.


Clinical significance

Patients with Addison's disease, pituitary insufficiency, or
cystic fibrosis Cystic fibrosis (CF) is a rare genetic disorder that affects mostly the lungs, but also the pancreas, liver, kidneys, and intestine. Long-term issues include difficulty breathing and coughing up mucus as a result of frequent lung infections. O ...
sometimes have a hyper-sensitivity to the five primary tastes.


Disorders of taste

* ageusia (complete loss of taste) * hypogeusia (reduced sense of taste) * dysgeusia (distortion in sense of taste) *
hypergeusia Hypergeusia is a taste disorder where the sense is abnormally heightened. It can be associated with a lesion of the posterior fossa and Addison's disease; where a patient will crave for salty and sour taste due to the abnormal loss of ions with ur ...
(abnormally heightened sense of taste) Viruses can also cause loss of taste. About 50% of patients with SARS-CoV-2 (causing COVID-19) experience some type of disorder associated with their sense of smell or taste, including ageusia and dysgeusia.
SARS-CoV-1 Severe acute respiratory syndrome coronavirus 1 (SARS-CoV-1; or Severe acute respiratory syndrome coronavirus, SARS-CoV) is a strain of coronavirus that causes severe acute respiratory syndrome (SARS), the respiratory illness responsible for th ...
, MERS-CoV and even the flu ( influenza virus) can also disrupt olfaction.


History

Ayurveda, an ancient Indian healing science, has its own tradition of basic tastes, comprising sweet, salty, sour, pungent, bitter & astringent.Ayurvedic balancing: an integration of Western fitness with Eastern wellness (Pages 25-26/188)
Joyce Bueker. Llewellyn Worldwide, 2002.
In the West, Aristotle postulated in BC that the two most basic tastes were sweet and bitter. He was one of the first identified persons to develop a list of basic tastes. The Ancient Chinese regarded
spiciness Pungency () refers to the taste of food commonly referred to as spiciness, hotness or heat, found in foods such as chili peppers. Highly pungent tastes may be experienced as unpleasant. The term piquancy () is sometimes applied to foods with a l ...
as a basic taste.


Research

The receptors for the basic tastes of bitter, sweet and savory have been identified. They are G protein-coupled receptors. 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 channel. There is some evidence for a sixth taste that senses fatty substances. 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 disease.


See also

*
Beefy meaty peptide Beefy meaty peptide, also known as delicious peptide and abbreviated as BMP, is an 8-amino acid long peptide that has been identified as giving a beefy flavour to foods in which it is present. It was isolated from beef soup by Yamasaki and Maekaw ...
* Digital lollipop * Optimal foraging theory *
Palatability Palatability (or palatableness) is the hedonic reward (i.e., pleasure) provided by foods or fluids that are agreeable to the "palate", which often varies relative to the homeostatic satisfaction of nutritional, water, or energy needs. The palatabil ...
* Vomeronasal organ * Sensory analysis *
Tea tasting Tea tasting is the process in which a trained taster determines the quality of a particular tea. Due to climatic conditions, topography, manufacturing process, and different cultivars of the '' Camellia sinensis'' plant (tea), the final product m ...
* Wine tasting


Notes

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 sensations...However, there might be other less conspicuous classes or subclasses of primary sensations",
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" 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 purposes. 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, is essentially the same as that of Svrivastava & Rastogi (2003), Guyton & Hall (2006), and Joesten ''et al.'' (2007). 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 0.75.


References


Further reading

* * {{Authority control Sensory systems Gustatory system