Taste
The gustatory system or sense of taste is the
The tongue is covered with thousands of small bumps called
Taste receptors in the mouth sense the five basic tastes:
The basic tastes contribute only partially to the sensation and flavor of food in the mouth—other factors include
As the gustatory system senses both harmful and beneficial things, all basic tastes bring either caution or craving depending upon the effect the things they sense have on the body.[13] Sweetness helps to identify energy-rich foods, while bitterness warns people of poisons.[14]
Among humans, taste perception begins to fade during
Basic tastes
This section needs additional citations for verification. (September 2016) |
The gustatory system allows animals to distinguish between safe and harmful food, and to gauge foods' nutritional value.
The five specific tastes received by taste receptors are saltiness, sweetness, bitterness, sourness, and savoriness (often known by its Japanese name umami 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.[17] By the end of the 20th century, the concept of umami was becoming familiar to Western society.
One study found that salt and sour taste mechanisms both detect, in different ways, the presence of sodium chloride (salt) in the mouth. Acids are also detected and perceived as sour.[18] 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.[citation 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
Sweet taste signals the presence of
The savory taste (known in Japanese as umami), identified by Japanese chemist
Pungency (piquancy or hotness) had traditionally been considered a sixth basic taste.[19] In 2015, researchers suggested a new basic taste of fatty acids called "fat taste",[20] although "oleogustus" and "pinguis" have both been proposed as alternate terms.[21][22]
Sweetness
Sweetness, usually regarded as a pleasurable sensation, is produced by the presence of
Sourness
Sourness is the taste that detects acidity. The sourness of substances is rated relative to dilute hydrochloric acid, which has a sourness index of 1. By comparison, tartaric acid has a sourness index of 0.7, citric acid an index of 0.46, and carbonic acid an index of 0.06.[24][25]
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 (
The most common foods with natural
Saltiness
Saltiness taste seems to have two components: a low-salt signal and a high-salt signal. The low-salt signal causes a sensation of deliciousness, while the high-salt signal typically causes the sensation of "too salty".[30]
The low-salt signal is understood to be caused by the
A number of similar cations also trigger the low salt signal. The size of lithium and potassium ions most closely resemble those of sodium, and thus the saltiness is most similar. In contrast, rubidium and caesium ions are far larger, so their salty taste differs accordingly.[citation needed] The saltiness of substances is rated relative to sodium chloride (NaCl), which has an index of 1.[24][25] Potassium, as potassium chloride (KCl), is the principal ingredient in salt substitutes and has a saltiness index of 0.6.[24][25]
Other
The high-salt signal is still very poorly understood as of 2023. Even in rodents, this signal is not blocked by amiloride. Sour and bitter cells trigger on high chloride levels, but the specific receptor is still being identified.[30]
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
Bitterness is of interest to those who study
The threshold for stimulation of bitter taste by quinine averages a concentration of 8 μ
Research has shown that TAS2Rs (taste receptors, type 2, also known as T2Rs) such as TAS2R38 coupled to the G protein gustducin are responsible for the human ability to taste bitter substances.[38] They are identified not only by their ability to taste for certain "bitter" ligands, but also by the morphology of the receptor itself (surface bound, monomeric).[18] 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.[39] 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.[40] 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.[41] Researchers use two synthetic substances, phenylthiocarbamide (PTC) and 6-n-propylthiouracil (PROP) to study the genetics of bitter perception. These two substances taste bitter to some people, but are virtually tasteless to others. Among the tasters, some are so-called "supertasters" to whom PTC and PROP are extremely bitter. The variation in sensitivity is determined by two common alleles at the TAS2R38 locus.[42] 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.[citation 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.
Umami
Umami, or savoriness, is an
Umami was first studied in 1907 by Ikeda isolating dashi taste, which he identified as the chemical monosodium glutamate (MSG).[17][48] 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.[49]
Some savory taste buds respond specifically to glutamate in the same way that "sweet" ones respond to sugar. Glutamate binds to a variant of
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.[52] Substances are usually measured relative to sucrose,[53] which is usually given an arbitrary index of 1[54][55] or 100.[56] Rebaudioside A is 100 times sweeter than sucrose; fructose is about 1.4 times sweeter; glucose, a sugar found in honey and vegetables, is about three-quarters as sweet; and lactose, a milk sugar, is one-half as sweet.[b][52]
The sourness of a substance can be rated by comparing it to very dilute hydrochloric acid (HCl).[57]
Relative saltiness can be rated by comparison to a dilute salt solution.[58]
Quinine, a bitter medicinal found in tonic water, can be used to subjectively rate the bitterness of a substance.[59] 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.[59] More formal chemical analysis, while possible, is difficult.[59]
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
Functional structure
Taste is a form of
Sweetness
Sweetness is produced by the presence of
Saltiness
Saltiness is a taste produced best by the presence of cations (such as Na+
, K+
or Li+
)[63] and is directly detected by cation influx into glial like cells via leak channels causing depolarisation of the cell.[63]
Other
, ions, in general, elicit a bitter rather than a salty taste even though they, too, can pass directly through ion channels in the tongue.[citation needed
Sourness
Sourness is acidity,[64][65] and, like salt, it is a taste sensed using ion channels.[63] 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 sour.[66]
Bitterness
Research has shown that TAS2Rs (taste receptors, type 2, also known as T2Rs) such as TAS2R38 are responsible for the ability to taste bitter substances in vertebrates.[67] 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).[68]
Savoriness
The
Glutamic acid binds to a variant of the G protein-coupled receptor, producing a savory taste.[50][51]
Further sensations and transmission
The tongue can also feel other sensations not generally included in the basic tastes. These are largely detected by the
The
Pungency (also spiciness or hotness)
Substances such as
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
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 often combine this with chili pepper to produce a 麻辣 málà, "numbing-and-hot", or "mati rasa" flavor.[72] Typical in northern Brazilian cuisine, jambu is an herb used in dishes like tacacá. These sensations, although not taste, fall into a category of chemesthesis.
Astringency
Some foods, such as unripe fruits, contain
Metallicness
A metallic taste may be caused by food and drink, certain medicines or
Fat taste
Recent research reveals a potential
Other possible fat taste receptors have been identified. G protein-coupled receptors free fatty acid receptor 4 (also termed GPR120) and to a much lesser extent Free fatty acid receptor 1 (also termed GPR40)[89] 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.[90]
Monovalent cation channel TRPM5 has been implicated in fat taste as well,[91] 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.[87]
Proposed alternate names to fat taste include oleogustus
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.[95]
Heartiness
Kokumi (/koʊkuːmi/, Japanese: kokumi (コク味)[96] from koku (こく)[96]) 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,
Calcium-sensing receptors (CaSR) are receptors for kokumi substances which, applied around taste pores, induce an increase in the intracellular Ca concentration in a subset of cells.[97] This subset of CaSR-expressing taste cells are independent from the influenced basic taste receptor cells.[99] CaSR agonists directly activate the CaSR on the surface of taste cells and integrated in the brain via the central nervous system. A basal level of calcium, corresponding to the physiological concentration, is necessary for activation of the CaSR to develop the kokumi sensation.[100]
Calcium
The distinctive taste of chalk has been identified as the calcium component of that substance.
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
Starchiness
A 2016 study suggested that humans can taste starch (specifically, a glucose oligomer) independently of other tastes such as sweetness, without suggesting an associated chemical receptor.[104][105][106]
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.[107]
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
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.[108] 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.[109]
Reticular formation (includes Raphe nuclei responsible for serotonin production) is signaled to release serotonin during and after a meal to suppress appetite.[110] 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 pupils.[111]
Spinal ganglion are involved in movement.
The frontal operculum is speculated to be the memory and association hub for taste.[citation needed]
The
Taste in insects
Insects taste using small hair-like structures called taste sensilla, specialized sensory organs located on various body parts such as the mouthparts, legs, and wings. These sensilla contain gustatory receptor neurons (GRNs) sensitive to a wide range of chemical stimuli.
Insects respond to sugar, bitter, acid, and salt tastes. However, their taste spectrum extends to include water, fatty acids, metals, carbonation, RNA, ATP, and pheromones. Detecting these substances is vital for behaviors like feeding, mating, and oviposition.
Invertebrates' ability to taste these compounds is fundamental to their survival and provides insights into the evolution of sensory systems. This knowledge is crucial for understanding insect behavior and has applications in pest control and pollination biology.
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
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 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 sometimes have a hyper-sensitivity to the five primary tastes.[116]
Disorders of taste
- ageusia (complete loss of taste)
- hypogeusia (reduced sense of taste)
- dysgeusia (distortion in sense of taste)
- hypergeusia (abnormally heightened sense of taste)
Viruses can also cause loss of taste. About 50% of patients with
History
In
Research
The receptors for the basic tastes of bitter, sweet and savory have been identified. They are G protein-coupled receptors.[122] The cells that detect sourness have been identified as a subpopulation that express the protein PKD2L1, and The responses are mediated by an influx of protons into the cells.[122] As of 2019, molecular mechanisms for each taste appear to be different, although all taste perception relies on activation of P2X purinoreceptors on sensory nerves.[123]
See also
Notes
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",[124]
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"[125] 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.[57]
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,[24][25] is essentially the same as that of Svrivastava & Rastogi (2003),[126] Guyton & Hall (2006),[57] and Joesten et al. (2007).[54] 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.
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Further reading
- Chandrashekar, Jayaram; Hoon, Mark A.; Ryba; Nicholas, J. P. & Zuker, Charles S. (16 November 2006), "The receptors and cells for mammalian taste" (PDF), S2CID 4431221, archived from the original(PDF) on 22 July 2011, retrieved 13 September 2010
- Chaudhari, Nirupa & Roper, Stephen D. (2010), "The cell biology of taste", PMID 20696704
- The dictionary definition of taste at Wiktionary
- Media related to Taste at Wikimedia Commons