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{{short description|Sense of chemicals on the tongue}}
{{about|the sense|the social and aesthetic aspects of "taste"|Taste (sociology)|other uses}}
{{Use dmy dates|date=December 2017}}
[[File:Taste bud.svg|thumb|right|[[Taste bud]]]]
The '''gustatory system''' or '''sense of taste''' is the [[sensory system]] that is partially responsible for the [[perception]] of taste (flavor).<ref name="Trivedi">{{cite journal | last=Trivedi | first=Bijal P. | title=Gustatory system: The finer points of taste | journal=Nature| volume=486 | issue=7403 | year=2012 | issn=0028-0836 | doi=10.1038/486s2a | pages=S2–S3|pmid=22717400| bibcode=2012Natur.486S...2T | s2cid=4325945 | doi-access=free }}</ref> Taste is the perception stimulated when a substance in the [[mouth]] [[biochemistry|reacts chemically]] with [[taste receptor]] cells located on [[taste bud]]s in the [[oral cavity]], mostly on the [[tongue]]. Taste, along with [[olfaction|the sense of smell]] and [[trigeminal nerve]] stimulation (registering texture, pain, and temperature), determines [[Flavoring|flavor]]s of [[food]] and other substances. [[Human]]s have taste receptors on taste buds and other areas, including the upper surface of the [[tongue]] and the [[epiglottis]].<ref name="Witt">{{cite book | last=Witt | first=Martin | title=Smell and Taste | chapter=Anatomy and development of the human taste system | series=Handbook of Clinical Neurology | volume=164 | year=2019 | isbn=978-0-444-63855-7 | issn=0072-9752 | pmid=31604544 | doi=10.1016/b978-0-444-63855-7.00010-1 | pages=147–171| s2cid=204332286 }}</ref><ref>[https://books.google.com/books?id=dNhFLnc6NRkC&dq=taste+bud+concentrated&pg=PA201 Human biology (Page 201/464)] {{Webarchive|url=https://web.archive.org/web/20230326054409/https://books.google.com/books?id=dNhFLnc6NRkC&dq=taste+bud+concentrated&pg=PA201 |date=26 March 2023 }} Daniel D. Chiras. Jones & Bartlett Learning, 2005.</ref> The [[gustatory cortex]] is responsible for the perception of taste.

The tongue is covered with thousands of small bumps called [[lingual papillae|papillae]], which are visible to the [[naked eye]].<ref name=Witt/> Within each papilla are hundreds of taste buds.<ref name=Trivedi/><ref name = Schacter169>{{cite book|last=Schacter|first=Daniel|title=Psychology Second Edition|year=2009|publisher=Worth Publishers|location=United States of America|isbn=978-1-4292-3719-2|page=[https://archive.org/details/psychology0000scha/page/169 169]|url=https://archive.org/details/psychology0000scha/page/169}}</ref> The exceptions to this is the [[filiform papillae]] that do not contain taste buds. There are between 2000 and 5000<ref name="Boron, W.F. 2003">Boron, W.F., E.L. Boulpaep. 2003. Medical Physiology. 1st ed. Elsevier Science USA.</ref> 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 basic tastes: [[#Sweetness|sweetness]], [[#Sourness|sourness]], [[#Saltiness|saltiness]], [[Bitter (taste)#Bitterness|bitterness]], and [[savoriness]] (also known as ''savory'' or ''umami'').<ref name=Trivedi/><ref name=Witt/><ref name=Kean>{{cite journal|last1=Kean|first1=Sam|title=The science of satisfaction|journal=Distillations Magazine|date=Fall 2015|volume=1|issue=3|pages=5|url=https://www.sciencehistory.org/distillations/magazine/the-science-of-satisfaction|access-date=20 March 2018|archive-date=17 November 2019|archive-url=https://web.archive.org/web/20191117095930/https://www.sciencehistory.org/distillations/magazine/the-science-of-satisfaction|url-status=live}}</ref><ref>{{cite web|url= https://www.ncbi.nlm.nih.gov/pubmedhealth/PMH0072592/|title= How does our sense of taste work?|date= 6 January 2012|website= PubMed|access-date= 5 April 2016|archive-date= 9 March 2015|archive-url= https://web.archive.org/web/20150309010345/https://www.ncbi.nlm.nih.gov/pubmedhealth/PMH0072592/|url-status= live}}</ref> Scientific experiments have demonstrated that these five tastes exist and are distinct from one another. Taste buds are able to tell different tastes apart when they interact with different molecules or ions. Sweetness, savoriness, and bitter tastes are triggered by the binding of molecules to [[G protein-coupled receptors]] on the [[cell membrane]]s of taste buds. Saltiness and sourness are perceived when [[alkali metal]]s or [[hydrogen]] [[ions]] meet taste buds, respectively.<ref>Human Physiology: An integrated approach 5th Edition -Silverthorn, Chapter-10, Page-354</ref><ref>{{Cite journal |last1=Turner |first1=Heather N. |last2=Liman |first2=Emily R. |date=2022-02-10 |title=The Cellular and Molecular Basis of Sour Taste |journal=Annual Review of Physiology |language=en |volume=84 |issue=1 |pages=41–58 |doi=10.1146/annurev-physiol-060121-041637 |pmid=34752707 |s2cid=243940546 |issn=0066-4278 |pmc=10191257 }}</ref>

The basic tastes contribute only partially to the sensation and flavor of food in the mouth—other factors include [[Odor|smell]],<ref name=Trivedi/> detected by the [[olfactory epithelium]] of the nose;<ref>[http://faculty.washington.edu/chudler/nosek.html Smell – The Nose Knows] {{Webarchive|url=https://web.archive.org/web/20170913090326/http://faculty.washington.edu/chudler/nosek.html |date=13 September 2017 }} washington.edu, Eric H. Chudler.</ref> [[Texture (food)|texture]],<ref>
* [https://books.google.com/books?id=aJBIbvClWfcC&dq=Food%20texture&pg=PA36 Food texture: measurement and perception (page 36/311)] Andrew J. Rosenthal. Springer, 1999.
* [https://books.google.com/books?id=aJBIbvClWfcC&dq=Food%20texture&pg=PA3 Food texture: measurement and perception (page 3/311)] Andrew J. Rosenthal. Springer, 1999.</ref> detected through a variety of [[mechanoreceptor]]s, muscle nerves, etc.;<ref>[https://books.google.com/books?id=aJBIbvClWfcC&dq=Food%20texture&pg=PA4 Food texture: measurement and perception (page 4/311)] {{Webarchive|url=https://web.archive.org/web/20230326054409/https://books.google.com/books?id=aJBIbvClWfcC&dq=Food%20texture&pg=PA4 |date=26 March 2023 }} Andrew J. Rosenthal. Springer, 1999.</ref> temperature, detected by [[thermoreceptor|temperature receptor]]s; and "coolness" (such as of [[menthol]]) and "hotness" ([[pungency]]), by [[chemesthesis]].

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.<ref name=aa>[http://www.scientificamerican.com/article.cfm?id=two-great-tastes-not-great-together Why do two great tastes sometimes not taste great together?] {{Webarchive|url=https://web.archive.org/web/20111128175618/http://www.scientificamerican.com/article.cfm?id=two-great-tastes-not-great-together |date=28 November 2011 }} scientificamerican.com. Dr. Tim Jacob, Cardiff University. 22 May 2009.</ref> Sweetness helps to identify energy-rich foods, while bitterness warns people of poisons.<ref>{{cite journal|last=Miller|first=Greg|title=Sweet here, salty there: Evidence of a taste map in the mammilian brain.|journal=Science|date=2 September 2011|volume=333|issue=6047|page=1213|doi=10.1126/science.333.6047.1213|pmid=21885750|bibcode=2011Sci...333.1213M}}</ref>

Among humans, taste perception begins to fade during [[ageing]], tongue papillae are lost, and [[saliva]] production slowly decreases.<ref name="SeidelBall2010">{{cite book|author1=Henry M Seidel|author2=Jane W Ball|author3=Joyce E Dains|title=Mosby's Guide to Physical Examination|url=https://books.google.com/books?id=j7HSCQAAQBAJ&pg=PA303|date=1 February 2010|publisher=Elsevier Health Sciences|isbn=978-0-323-07357-8|page=303}}</ref> Humans can also have distortion of tastes ([[dysgeusia]]). Not all [[mammal]]s share the same tastes: some [[rodent]]s can taste [[starch]] (which humans cannot), [[cat]]s cannot taste sweetness, and several other [[carnivores]], including [[hyena]]s, [[dolphin]]s, and [[sea lion]]s, have lost the ability to sense up to four of their ancestral five basic tastes.<ref>{{cite web|last1=Scully|first1=Simone M.|title=The Animals That Taste Only Saltiness|url=http://nautil.us/blog/the-animals-that-taste-only-saltiness|website=Nautilus|access-date=8 August 2014|date=2014-06-09|archive-date=14 June 2014|archive-url=https://web.archive.org/web/20140614013146/http://nautil.us/blog/the-animals-that-taste-only-saltiness|url-status=dead}}</ref>

==Basic tastes==
{{More citations needed section|date=September 2016}}
The gustatory system allows animals to distinguish between safe and harmful food and to gauge different foods' nutritional value. [[Digestive enzyme]]s 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 [[lingual papillae|papillae]], which are visible to the naked eye. Within each papilla are hundreds of taste buds.<ref name="Schacter169"/> The exception to this are the [[filiform papillae]], which do not contain taste buds. There are between 2,000 and 5,000<ref name="Boron, W.F. 2003"/> 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 receptor]]s are saltiness, [[sweetness]], bitterness, sourness, and [[umami|savoriness]] (often known by its Japanese name {{transliteration|ja|umami}}, which translates to 'deliciousness').

As of the early 20th century, Western physiologists and psychologists believed that 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.<ref name="Ikeda1909">{{cite journal | last=Ikeda | first=Kikunae | title=New Seasonings | journal=Chemical Senses |orig-year=1909 | year= 2002 | volume=27 | issue=9 | pages= 847–849 | doi=10.1093/chemse/27.9.847 | pmid=12438213| doi-access=free }}; a partial translation from {{cite journal|last=Ikeda|first=Kikunae|title=New Seasonings|journal=Journal of the Chemical Society of Tokyo|language=ja|volume=30|issue=8|pages=820–836|year=1909|doi=10.1246/nikkashi1880.30.820 |pmid=12438213|doi-access=free}}</ref> 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.<ref name="auto">{{cite journal | last=Lindemann | first=Bernd | title=Receptors and transduction in taste| journal=Nature |date=13 September 2001 | volume=413 |pages= 219–225| doi=10.1038/35093032| pmid=11557991 | issue=6852| bibcode=2001Natur.413..219L | s2cid=4385513 }}</ref> The detection of salt is important to many organisms, but especially 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 that facilitates passive re-uptake of water into the blood.{{citation needed|reason=Which studies purport this?|date=September 2016}} 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 presumably is 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 that grow in such media. Additionally, sour taste signals [[acids]], which can cause serious tissue damage.

Sweet taste signals the presence of [[carbohydrates]] in solution.{{citation needed|date=February 2023}} Since carbohydrates have a very high [[calorie]] count (saccharides have many bonds, therefore much energy),{{citation needed|date=April 2020}} they are desirable to the human body, which evolved to seek out the highest-calorie-intake foods.{{citation needed|date=February 2023}} They are used as direct energy ([[sugars]]) and storage of energy ([[glycogen]]). Many non-carbohydrate molecules trigger a sweet response, leading to the development of many artificial sweeteners, including [[saccharin]], [[sucralose]], and [[aspartame]]. It is still unclear how these substances activate the sweet receptors and what adaptative significance this has had.

The savory taste (known in Japanese as {{transliteration|ja|umami}}), identified by Japanese chemist [[Kikunae Ikeda]], signals the presence of the [[amino acid]] [[L-glutamate]]. The amino acids in proteins are used in the body to build muscles and organs, and to transport molecules ([[hemoglobin]]), [[antibodies]], and the organic catalysts known as [[enzymes]]. These are all critical molecules, and it is important to have a steady supply of amino acids; consequently, savory tastes trigger a pleasurable response, encouraging the intake of [[peptides]] and [[proteins]].

[[Pungency]] (piquancy or hotness) had traditionally been considered a sixth basic taste.<ref name="books.google.com">[https://books.google.com/books?id=s2hsBJAp5fYC&q=the+six+tastes&pg=PA25 Ayurvedic balancing: an integration of Western fitness with Eastern wellness (Pages 25-26/188)] Joyce Bueker. Llewellyn Worldwide, 2002.</ref> In 2015, researchers suggested a new basic taste of [[fatty acid]]s called "fat taste",<ref>{{Cite journal|last1=Keast|first1=Russell SJ|last2=Costanzo|first2=Andrew|date=2015-02-03|title=Is fat the sixth taste primary? Evidence and implications|journal=Flavour|volume=4|pages=5|doi=10.1186/2044-7248-4-5|issn=2044-7248|doi-access=free|hdl=10536/DRO/DU:30069796|hdl-access=free}}</ref> although "oleogustus" and "pinguis" have both been proposed as alternate terms.<ref>{{Cite journal|last1=Running|first1=Cordelia A.|last2=Craig|first2=Bruce A.|last3=Mattes|first3=Richard D.|date=2015-09-01|title=Oleogustus: The Unique Taste of Fat|journal=Chemical Senses|language=en|volume=40|issue=7|pages=507–516|doi=10.1093/chemse/bjv036|pmid=26142421|issn=0379-864X|doi-access=free}}</ref><ref name="Reed 353S–360S">{{Cite journal|last1=Reed|first1=Danielle R.|last2=Xia|first2=Mary B.|date=2015-05-01|title=Recent Advances in Fatty Acid Perception and Genetics|journal=Advances in Nutrition|language=en|volume=6|issue=3|pages=353S–360S|doi=10.3945/an.114.007005|issn=2156-5376|pmid=25979508|pmc=4424773}}</ref>

===Sweetness===
{{Main|Sweetness}}
[[File:Signal Transaction of the Sweet Taste.svg|thumb|The diagram above depicts the signal transduction pathway of the sweet taste. Object A is a taste bud, object B is one taste cell of the taste bud, and object C is the neuron attached to the taste cell. I. Part I shows the reception of a molecule. 1. Sugar, the first messenger, binds to a protein receptor on the cell membrane. II. Part II shows the transduction of the relay molecules. 2. G Protein-coupled receptors, second messengers, are activated. 3. G Proteins activate adenylate cyclase, an enzyme, which increases the cAMP concentration. Depolarization occurs. 4. The energy, from step 3, is given to activate the K+, potassium, protein channels.III. Part III shows the response of the taste cell. 5. Ca+, calcium, protein channels is activated.6. The increased Ca+ concentration activates neurotransmitter vesicles. 7. The neuron connected to the taste bud is stimulated by the neurotransmitters.]]

Sweetness, usually regarded as a pleasurable sensation, is produced by the presence of [[sugar]]s and substances that mimic sugar. Sweetness may be connected to [[aldehyde]]s and [[ketone]]s, which contain a [[carbonyl group]]. Sweetness is detected by a variety of [[G protein coupled receptor]]s (GPCR) coupled to the [[G protein]] [[gustducin]] found on the [[taste bud]]s. 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.<ref>{{cite journal |last=Zhao |first=Grace Q. |author2=Yifeng Zhang |author3=Mark A. Hoon |author4=Jayaram Chandrashekar |author5=Isolde Erlenbach |author6=Nicholas J.P. Ryba |author7=Charles S. Zuker |title=The Receptors for Mammalian Sweet and Savory taste |journal=Cell |date=October 2003 |volume=115 |issue=3 |pages=255–266 |doi=10.1016/S0092-8674(03)00844-4 |pmid=14636554 |s2cid=11773362 |df=dmy-all |doi-access=free }}</ref>

[[Taste detection threshold]]s for sweet substances are rated relative to [[sucrose]], which has an index of 1.<ref name="textbookofmedicalphysiology8thed"/><ref name=McLaughlin&Margolskee/> The average human detection threshold for sucrose is 10 millimoles per liter. For [[lactose]] it is 30 millimoles per liter, with a sweetness index of 0.3,<ref name="textbookofmedicalphysiology8thed"/> and [[5-Nitro-2-propoxyaniline|5-nitro-2-propoxyaniline]] 0.002 millimoles per liter. "Natural" sweeteners such as [[saccharide]]s activate the GPCR, which releases [[gustducin]]. The gustducin then activates the molecule [[adenylate cyclase]], which catalyzes the production of the molecule [[cyclic adenosine monophosphate|cAMP]], or adenosine 3', 5'-cyclic monophosphate. This molecule closes potassium ion channels, leading to depolarization and neurotransmitter release. Synthetic sweeteners such as [[saccharin]] activate different GPCRs and induce taste receptor cell depolarization by an alternate pathway.

===Sourness===
{{Redirect|Sour}}[[File:Signal Transaction of Taste; Sour & Salty.svg|thumb|The diagram depicts the signal transduction pathway of the sour or salty taste. Object A is a taste bud, object B is a taste receptor cell within object A, and object C is the neuron attached to object B.

I. Part I is the reception of hydrogen ions or sodium ions.

1. If the taste is sour, H+ ions, from acidic substances, pass through H+ channels. Depolarization takes place

II. Part II is the transduction pathway of the relay molecules. 2. Cation, such as K+, channels are opened.

III. Part III is the response of the cell.

3. An influx of Ca+ ions is activated.

4. The Ca+ activates neurotransmitters.

5. A signal is sent to the neuron attached to the taste bud.]]

Sourness is the taste that detects [[acid]]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.<ref name="textbookofmedicalphysiology8thed"/><ref name=McLaughlin&Margolskee/>

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]]s) that are abundant in sour substances can directly enter the Type III taste cells through a proton channel.<ref>{{cite journal | title= A proton current drives action potentials in genetically identified sour taste cells|author1=Rui Chang, Hang Waters |author2=Emily Liman |name-list-style=amp | journal= Proc Natl Acad Sci U S A|date= 2010 | volume=107 | issue=51 | pages=22320–22325 | pmc= 3009759
 | pmid=21098668 | doi=10.1073/pnas.1013664107|bibcode=2010PNAS..10722320C|doi-access=free }}</ref> This channel was identified in 2018 as [[Otopetrin|otopetrin 1 (OTOP1)]].<ref>{{Cite journal|last=Tu|first=YH|date=2018|title=An evolutionarily conserved gene family encodes proton-selective ion channels.|journal=Science|volume=359|issue=6379|pages=1047–1050|doi=10.1126/science.aao3264|pmid=29371428|pmc=5845439|bibcode=2018Sci...359.1047T}}</ref> 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.<ref>{{cite journal | title=The K+ channel KIR2.1 functions in tandem with proton influx to mediate sour taste transduction | vauthors=Ye W, Chang RB, Bushman JD, Tu YH, Mulhall EM, Wilson CE, Cooper AJ, Chick WS, Hill-Eubanks DC, Nelson MT, Kinnamon SC, Liman ER | journal=Proc Natl Acad Sci U S A|date= 2016 | volume=113 | issue=2 | pages=E229–238 | pmid=26627720| pmc= 4720319 | doi=10.1073/pnas.1514282112| bibcode=2016PNAS..113E.229Y| doi-access=free }}</ref>

The most common foods with natural [[sourness]] are [[fruit]]s, such as [[lemon]], [[Lime (fruit)|lime]], [[grape]], [[Orange (fruit)|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,<ref>{{cite journal | title=Heightened Sour Preferences During Childhood |author1=Djin Gie Liem |author2=Julie A. Mennella |name-list-style=amp | journal=Chem Senses |date=February 2003 | volume=28 | issue=2 | pages=173–180 | pmc=2789429 | doi=10.1093/chemse/28.2.173 | pmid=12588738}}</ref> and [[Sour sanding|sour candy]] containing citric acid or [[malic acid]] is common.

===Saltiness===
{{Redirect|Saltiness|saltiness in water|Salinity}}
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".<ref name=Taruno2023>{{cite journal |last1=Taruno |first1=Akiyuki |last2=Gordon |first2=Michael D. |title=Molecular and Cellular Mechanisms of Salt Taste |journal=Annual Review of Physiology |date=10 February 2023 |volume=85 |issue=1 |pages=25–45 |doi=10.1146/annurev-physiol-031522-075853 |doi-access=free}} {{cite web |last1=Elahi |first1=Tasnuva |title=Salt Taste Is Surprisingly Mysterious |url=https://nautil.us/salt-taste-is-surprisingly-mysterious-382855/ |website=Nautilus |date=15 September 2023}}</ref>

The low-salt signal is understood to be caused by the [[epithelial sodium channel]] (ENaC), which is composed of three subunits. ENaC in the taste cells allow sodium [[cation]]s to enter the cell. This on its own depolarizes the cell, and opens [[voltage-dependent calcium channel]]s, flooding the cell with positive calcium ions and leading to [[neurotransmitter]] release. ENaC can be blocked by the drug [[amiloride]] in many mammals, especially rats. The sensitivity of the low-salt taste to amiloride in humans is much less pronounced, leading to conjecture that there may be additional low-salt receptors besides ENaC to be discovered.<ref name=Taruno2023/>

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|date=November 2008}} The saltiness of substances is rated relative to sodium chloride (NaCl), which has an index of 1.<ref name="textbookofmedicalphysiology8thed"/><ref name=McLaughlin&Margolskee/> Potassium, as [[potassium chloride]] (KCl), is the principal ingredient in [[salt substitute]]s and has a saltiness index of 0.6.<ref name="textbookofmedicalphysiology8thed"/><ref name=McLaughlin&Margolskee/>

Other [[Valence (chemistry)|monovalent]] [[cations]], e.g. [[ammonium]] (NH4+), and [[divalent]] cations of the [[alkali earth metal]] group of the [[periodic table]], e.g. calcium (Ca2+), ions generally elicit a bitter rather than a salty taste even though they, too, can pass directly through ion channels in the tongue, generating an [[action potential]]. But the chloride of calcium is saltier and less bitter than potassium chloride, and is commonly used in pickle brine instead of KCl.{{citation needed|date=September 2023}}

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.<ref name=Taruno2023/>

===Bitterness===
{{See also|Bitter taste evolution}}
[[File:Signal Transaction of Taste; Bitter.svg|thumb|The diagram depicted above shows the signal transduction pathway of the bitter taste. Bitter taste has many different receptors and signal transduction pathways. Bitter indicates poison to animals. It is most similar to sweet. Object A is a taste bud, object B is one taste cell, and object C is a neuron attached to object B. I. Part I is the reception of a molecule.1. A bitter substance such as quinine, is consumed and binds to G Protein-coupled receptors.II. Part II is the transduction pathway 2. Gustducin, a G protein second messenger, is activated. 3. Phosphodiesterase, an enzyme, is then activated. 4. Cyclic nucleotide, cNMP, is used, lowering the concentration 5. Channels such as the K+, potassium, channels, close. III. Part III is the response of the taste cell. 6. This leads to increased levels of Ca+. 7. The neurotransmitters are activated. 8. The signal is sent to the neuron.]]

Bitterness is 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 agent]]s. Common bitter foods and beverages include [[coffee]], unsweetened [[Cocoa solids|cocoa]], South American [[Mate (beverage)|mate]], [[coca tea]], [[bitter gourd]], uncured [[Olive (fruit)|olives]], [[Peel (fruit)|citrus peel]], some varieties of [[cheese]], many plants in the family [[Brassicaceae]], [[dandelion]] greens, [[marrubium vulgare|horehound]], wild [[chicory]], and [[escarole]]. The ethanol in [[alcoholic beverages]] tastes bitter,<ref>{{cite journal |vauthors=Scinska A, Koros E, Habrat B, Kukwa A, Kostowski W, Bienkowski P |title=Bitter and sweet components of ethanol taste in humans |journal=Drug and Alcohol Dependence |volume=60 |issue=2 |pages=199–206 |date=August 2000 |pmid=10940547 |doi=10.1016/S0376-8716(99)00149-0}}</ref> as do the additional bitter ingredients found in some alcoholic beverages including [[hops]] in [[beer]] and [[Gentiana lutea|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 researchers<ref name="textbookofmedicalphysiology8thed" /><ref name="psychologyofeating&drinking">{{cite book|last=Logue|first=Alexandra W.|year=1986|title=The Psychology of Eating and Drinking|location=New York|publisher=W.H. Freeman & Co.|isbn=978-0-415-81708-0}}{{page needed|date=August 2014}}</ref> 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.<ref name="textbookofmedicalphysiology8thed" /><ref name= psychologyofeating&drinking/><ref>{{cite journal |last=Glendinning|first=J. I. |title=Is the bitter rejection response always adaptive? |journal=Physiol Behav |volume=56 |year=1994 |pages=1217–1227|doi=10.1016/0031-9384(94)90369-7 |pmid=7878094 |issue=6 |s2cid=22945002 }}</ref> 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.<ref name="encylopediahumanevolution">Jones, S., Martin, R., & Pilbeam, D. (1994) ''The Cambridge Encyclopedia of Human Evolution''. Cambridge: Cambridge University Press{{page needed|date=August 2014}}</ref> Amongst humans, various [[food processing]] techniques are used worldwide to detoxify otherwise inedible foods and make them palatable.<ref>Johns, T. (1990). ''With Bitter Herbs They Shall Eat It: Chemical ecology and the origins of human diet and medicine''. Tucson: University of Arizona Press{{page needed|date=August 2014}}</ref> 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.<ref>{{cite journal | last1 = Wang | first1 = X. | year = 2004 | title = Relaxation Of Selective Constraint And Loss Of Function In The Evolution Of Human Bitter Taste Receptor Genes | journal = Human Molecular Genetics | volume = 13 | issue = 21| pages = 2671–2678 | doi=10.1093/hmg/ddh289 | pmid=15367488| doi-access = free }}</ref>

The threshold for stimulation of bitter taste by quinine averages a concentration of 8 μ[[Molarity|M]] (8 micromolar).<ref name="textbookofmedicalphysiology8thed">Guyton, Arthur C. (1991) ''Textbook of Medical Physiology''. (8th ed). Philadelphia: W.B. Saunders</ref> The taste thresholds of other bitter substances are rated relative to quinine, which is thus given a reference index of 1.<ref name="textbookofmedicalphysiology8thed"/><ref name="McLaughlin&Margolskee">{{cite journal |date=November–December 1994 |author1= McLaughlin, Susan |author2= Margolskee, Rorbert F. |title= The Sense of Taste |journal= [[American Scientist]] |volume= 82 |issue= 6 |pages= 538–545}}</ref> For example, [[brucine]] has an index of 11, is thus perceived as intensely more bitter than quinine, and is detected at a much lower solution threshold.<ref name="textbookofmedicalphysiology8thed"/> 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.<ref name=McLaughlin&Margolskee/> It is used as an [[aversive agent]] (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.<ref>{{Cite web|url=https://www.bitrex.com/about-bitrex/what-is-bitrex|title=What is Bitrex?|date=21 December 2015|website=Bitrex – Keeping children safe|access-date=20 May 2020|archive-date=20 May 2020|archive-url=https://web.archive.org/web/20200520034844/https://www.bitrex.com/about-bitrex/what-is-bitrex|url-status=live}}</ref>

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.<ref>{{cite journal |last1=Maehashi|first1=K.|first2=M.|last2= Matano|first3= H. |last3=Wang|first4= L. A. |last4=Vo|first5=Y. |last5=Yamamoto|first6= L. |last6=Huang |title=Bitter peptides activate hTAS2Rs, the human bitter receptors |journal=Biochem Biophys Res Commun |volume=365 |year=2008 |pages=851–855 |doi=10.1016/j.bbrc.2007.11.070 |pmid=18037373 |issue=4|pmc=2692459}}</ref> They are identified not only by their ability to taste for certain "bitter" [[Ligand (biochemistry)|ligands]], but also by the morphology of the receptor itself (surface bound, monomeric).<ref name="auto"/> 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.<ref>{{cite journal|last=Meyerhof|year=2010|doi=10.1093/chemse/bjp092|pmid=20022913|volume=35|issue=2|title=The molecular receptive ranges of human TAS2R bitter taste receptors.|journal=Chem Senses|pages=157–70|doi-access=free}}</ref> Over 670 bitter-tasting compounds have been identified, on a [[BitterDB|bitter database]], of which over 200 have been assigned to one or more specific receptors.<ref>{{cite journal|last=Wiener|year=2012|doi=10.1093/nar/gkr755|url= |pmid=21940398|pmc=3245057|volume=40|issue=Database issue|title=BitterDB: a database of bitter compounds|journal=Nucleic Acids Res.|pages=D413–9}}</ref> It is speculated that the selective constraints on the TAS2R family have been weakened due to the relatively high rate of mutation and pseudogenization.<ref>{{cite journal |last1=Wang|first1=X.|first2=S. D. |last2=Thomas|first3= J. |last3=Zhang |title=Relaxation of selective constraint and loss of function in the evolution of human bitter taste receptor genes |journal=Hum Mol Genet |volume=13 |year=2004 |pages=2671–2678|doi=10.1093/hmg/ddh289 |pmid=15367488 |issue=21|doi-access=free }}</ref> Researchers use two synthetic substances, [[phenylthiocarbamide]] (PTC) and [[propylthiouracil|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]]s" to whom PTC and PROP are extremely bitter. The variation in sensitivity is determined by two common alleles at the TAS2R38 locus.<ref>{{cite journal|last1=Wooding|first1=S.|first2=U. K. |last2=Kim|first3=M. J. |last3=Bamshad|first4=J.|last4=Larsen|first5= L. B.|last5= Jorde|first6= D. |last6=Drayna |title=Natural selection and molecular evolution in PTC, a bitter-taste receptor gene |journal=Am J Hum Genet |volume=74 |year=2004 |pages=637–646 |doi=10.1086/383092|pmid=14997422 |issue=4 |pmc=1181941}}</ref> 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|date=March 2016}} 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===
{{Main|Umami}} {{See also|Glutamate flavoring}}

Umami, or savoriness, is an [[appetite|appetitive]] taste.<ref name=aa/><ref name="Ikeda1909" /> It can be tasted in [[soy sauce]], [[meat]], [[dashi]] and [[Consommé|consomme]]. A [[loanword]] from [[Japanese language|Japanese]] meaning "good flavor" or "good taste",<ref>[http://jisho.org/words?jap=%E6%97%A8%E5%91%B3&eng=&dict=edict 旨味 definition in English] {{Webarchive|url=https://web.archive.org/web/20110808195413/http://jisho.org/words?jap=%E6%97%A8%E5%91%B3&eng=&dict=edict |date=8 August 2011 }} Denshi Jisho—Online Japanese dictionary</ref> {{nihongo|''umami''|旨味}} is considered fundamental to many [[East Asian cuisine]]s,<ref>{{Cite web |date=2015 |title=Umami Taste Components and Their Sources in Asian Foods |url=https://www.researchgate.net/publication/262976927 |website=researchgate.net}}</ref> such as [[Japanese cuisine]].<ref name=jmin/> It dates back to the use of fermented [[fish sauce]]: ''[[garum]]'' in ancient Rome<ref>{{cite web|url=https://www.npr.org/sections/thesalt/2013/10/26/240237774/fish-sauce-an-ancient-roman-condiment-rises-again|title=Fish sauce: An ancient Roman condiment rises again|first=Deena|last=Prichep|date=26 October 2013|publisher=US National Public Radio|access-date=5 April 2018|archive-date=16 June 2018|archive-url=https://web.archive.org/web/20180616204216/https://www.npr.org/sections/thesalt/2013/10/26/240237774/fish-sauce-an-ancient-roman-condiment-rises-again|url-status=live}}</ref> and {{transliteration|zh|ge-thcup}} or {{transliteration|zh|[[Ketchup#Etymology|koe-cheup]]}} in ancient China.<ref>{{Cite web |last=Butler |first=Stephanie |date=2012-07-20 |title=The Surprisingly Ancient History of Ketchup |url=https://www.history.com/news/ketchup-surprising-ancient-history |access-date=2022-04-19 |website=HISTORY |language=en |archive-date=19 April 2022 |archive-url=https://web.archive.org/web/20220419033206/https://www.history.com/news/ketchup-surprising-ancient-history |url-status=live }}</ref>

Umami was first studied in 1907 by [[Kikunae Ikeda|Ikeda]] isolating [[dashi]] taste, which he identified as the chemical [[monosodium glutamate]] (MSG).<ref name="Ikeda1909" /><ref>{{cite journal |vauthors=Nelson G, Chandrashekar J, Hoon MA, etal |title=An amino-acid taste receptor |journal=Nature |volume=416 |issue=6877 |pages=199–202 |date=March 2002 |pmid=11894099 |doi=10.1038/nature726|bibcode=2002Natur.416..199N |s2cid=1730089 }}</ref> MSG is a sodium salt that produces a strong savory taste, especially combined with foods rich in [[nucleotide]]s such as meats, fish, nuts, and mushrooms.<ref name="times">{{cite news |last1=O'Connor |first1=Anahad |date=10 November 2008 |title=The Claim: The tongue is mapped into four areas of taste |work=The New York Times |url=https://www.nytimes.com/2008/11/11/health/11real.html?_r=1 |access-date=13 September 2010 |archive-date=16 December 2017 |archive-url=https://web.archive.org/web/20171216054031/http://www.nytimes.com/2008/11/11/health/11real.html?_r=1 |url-status=live }}</ref>

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 receptor|G protein coupled glutamate receptors]].<ref name="Lindemann B 99–100">{{cite journal |last=Lindemann |first=B |title=A taste for umami |journal=Nature Neuroscience |volume=3 |issue=2 |pages=99–100 |date=February 2000 |pmid=10649560 |doi=10.1038/72153|s2cid=10885181 }}</ref><ref name="Chaud">{{cite journal |vauthors=Chaudhari N, Landin AM, Roper SD |title=A metabotropic glutamate receptor variant functions as a taste receptor |journal=Nature Neuroscience |volume=3 |issue=2 |pages=113–9 |date=February 2000 |pmid=10649565 |doi=10.1038/72053|s2cid=16650588 }}</ref> L-glutamate may bond to a type of GPCR known as a metabotropic glutamate receptor ([[Metabotropic glutamate receptor 4|mGluR4]]) which causes the G-protein complex to activate the sensation of umami.<ref name=Chaud/>

==Measuring relative tastes==
{{Further|Taste confusion matrix}}

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.<ref name=Tsai2007>{{citation |date=14 May 2007 |last=Tsai |first=Michelle |title=How Sweet It Is? Measuring the intensity of sugar substitutes |url=http://www.slate.com/id/2165999/ |work=[[Slate (magazine)|Slate]] |publisher=[[The Washington Post Company]] |access-date=14 September 2010 |archive-date=13 August 2010 |archive-url=https://web.archive.org/web/20100813223327/http://www.slate.com/id/2165999/ |url-status=live }}</ref> Substances are usually measured relative to [[sucrose]],<ref name=Walters>{{citation |date=13 May 2008 |last=Walters |first=D. Eric |chapter=How is Sweetness Measured? |title=All About Sweeteners |chapter-url=http://www.sweetenerbook.com/measure.html |access-date=15 September 2010 |archive-date=24 December 2010 |archive-url=https://web.archive.org/web/20101224211700/http://www.sweetenerbook.com/measure.html |url-status=live }}</ref> which is usually given an arbitrary index of 1<ref name=JoestenEal2007>{{citation |year=2007 |author1=Joesten, Melvin D |author2=Hogg, John L |author3=Castellion, Mary E |chapter=Sweeteness Relative to Sucrose (table) |page=359 |chapter-url=https://books.google.com/books?id=8hIoN3Q_zOkC&q=%22relative+to+sucrose%22&pg=PA359 |title=The World of Chemistry: Essentials |edition=4th |place=Belmont, California|publisher=Thomson Brooks/Cole |isbn=978-0-495-01213-9 |url=https://books.google.com/books?id=8hIoN3Q_zOkC&q=world+chemistry |access-date=14 September 2010}}</ref><ref name=Coultate>{{citation |year=2009 |last=Coultate|first=Tom P |chapter=Sweetness relative to sucrose as an arbitrary standard |chapter-url=https://books.google.com/books?id=KF2A8Cz7B-cC&q=%22relative+to+sucrose%22&pg=PA268 |title=Food: The Chemistry of its Components |edition=5th |url=https://books.google.com/books?id=KF2A8Cz7B-cC&q=food+chemistry+components |pages=268–269 |place=Cambridge, UK |publisher=[[Royal Society of Chemistry]] |isbn=978-0-85404-111-4 |access-date=15 September 2010}}</ref> or 100.<ref name=Mehta2005>{{citation |year=2005 |author1=Mehta, Bhupinder |author2=Mehta, Manju |name-list-style=amp |chapter=Sweetness of sugars |title=Organic Chemistry |page=956 |chapter-url=https://books.google.com/books?id=QV6cwXA9XkEC&pg=PA956 |url=https://books.google.com/books?id=QV6cwXA9XkEC |place=India |publisher=Prentice-Hall |access-date=15 September 2010|isbn=978-81-203-2441-1}}</ref> [[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.{{Ref label|b|b|none}}<ref name=Tsai2007/>

The sourness of a substance can be rated by comparing it to very dilute [[hydrochloric acid]] (HCl).<ref name=Guyton&Hall2006/>

Relative saltiness can be rated by comparison to a dilute salt solution.<ref>[https://books.google.com/books?id=xteiARU46SQC&dq=rating%20a%20salty%20taste&pg=PA38 Food Chemistry (Page 38/1070)] H. D. Belitz, Werner Grosch, Peter Schieberle. Springer, 2009.</ref>

[[Quinine]], a bitter medicinal found in [[tonic water]], can be used to subjectively rate the bitterness of a substance.<ref name=qn/> 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.<ref name=qn>[https://books.google.com/books?id=4LazhtBDub0C&dq=measure+of+bitter+quinine&pg=PA38 Quality control methods for medicinal plant materials, Pg. 38] World Health Organization, 1998.</ref> More formal chemical analysis, while possible, is difficult.<ref name=qn/>

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]] for [[pyruvate]]s in garlics and onions.

==Functional structure==
[[File:1402 The Tongue.jpg|thumb|350px|Taste buds and papillae of the human tongue]]
[[File:Jaime Lara 2023 PMID 36409650 Figure 3.jpg|thumb|Taste receptors of the human tongue]]
[[File:Jaime Lara 2023 PMID 36409650 Figure 6.jpg|thumb|[[Signal transduction]] of taste receptors]]
Taste is a form of [[chemoreception]] which occurs in the specialised [[taste receptor]]s 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, [a]lthough 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."<ref>David V. Smith, Robert F. Margolskee: [https://www.scientificamerican.com/article/making-sense-of-taste-2006-09/ Making Sense of Taste] {{Webarchive|url=https://web.archive.org/web/20201029233717/https://www.scientificamerican.com/article/making-sense-of-taste-2006-09/ |date=29 October 2020 }} (Scientific American, September 1, 2006)</ref> As well, [[serotonin]] is thought to act as an intermediary hormone which communicates with taste cells within a taste bud, mediating the signals being sent to the brain. Receptor molecules are found on the top of [[microvilli]] of the taste cells.

===Sweetness===
Sweetness is produced by the presence of [[sugar]]s, some proteins, and other substances such as alcohols like [[anethol]], [[glycerol]] and [[propylene glycol]], [[saponins]] such as [[glycyrrhizin]], [[artificial sweetener]]s (organic compounds with a variety of structures), and [[lead]] compounds such as [[Lead(II) acetate|lead acetate]].{{Citation needed|date=October 2010}} It is often connected to [[aldehyde]]s and [[ketone]]s, which contain a [[carbonyl group]].{{Citation needed|date=October 2010}} Many foods can be perceived as sweet regardless of their actual sugar content. For example, some plants such as [[liquorice]], [[anise]] or [[stevia]] 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]] 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 receptor]]s coupled to a [[G protein]] that acts as an intermediary in the communication between taste bud and brain, [[gustducin]].<ref>[https://www.nytimes.com/1992/08/04/science/how-the-taste-bud-translates-between-tongue-and-brain.html?pagewanted=all How the Taste Bud Translates Between Tongue and Brain] {{Webarchive|url=https://web.archive.org/web/20170305103715/http://www.nytimes.com/1992/08/04/science/how-the-taste-bud-translates-between-tongue-and-brain.html?pagewanted=all |date=5 March 2017 }} nytimes.com, 4 August 1992.</ref> These receptors are T1R2+3 (heterodimer) and T1R3 (homodimer), which account for sweet sensing in humans and other animals.<ref>{{cite journal |vauthors=Zhao GQ, Zhang Y, Hoon MA, etal |title=The receptors for mammalian sweet and umami taste |journal=Cell |volume=115 |issue=3 |pages=255–66 |date=October 2003 |pmid=14636554 |doi=10.1016/S0092-8674(03)00844-4|s2cid=11773362 |doi-access=free }}</ref>

===Saltiness===
Saltiness is a taste produced best by the presence of [[ion|cations]] (such as {{chem|Na|+}}, {{chem|K|+}} or {{chem|Li|+}})<ref name=ion/> and is directly detected by cation influx into glial like cells via leak channels causing depolarisation of the cell.<ref name=ion/>

Other [[Valence (chemistry)|monovalent]] cations, e.g., [[ammonium]], {{chem|NH|4|+}}, and [[divalent]] cations of the [[alkali earth metal]] group of the [[periodic table]], e.g., calcium, {{chem|Ca|2+}}, ions, in general, elicit a bitter rather than a salty taste even though they, too, can pass directly through [[ion channel]]s in the tongue.{{Citation needed|date=September 2010}}

===Sourness===
Sourness is [[acid]]ity,<ref>[https://books.google.com/books?id=HL88AAAAIAAJ&dq=dilute%20hcl%20sourness&pg=PA241 outlines of chemistry with practical work (Page 241)] Henry John Horstman Fenton. CUP Archive.</ref><ref>[https://books.google.com/books?id=-liIkg49at0C&dq=dilute%20hcl%20sourness&pg=PA242 Focus Ace Pmr 2009 Science (Page 242/522)] Chang See Leong, Chong Kum Ying, Choo Yan Tong & Low Swee Neo. Focus Ace Pmr 2009 Science.</ref> and, like salt, it is a taste sensed using [[ion channel]]s.<ref name=ion>[https://books.google.com/books?id=7r4NFLOBSmsC&q=salty+sodium+ion+channelTransduction&pg=PA155 channels in sensory cells (Page 155/304)] Stephan Frings, Jonathan Bradley. Wiley-VCH, 2004.</ref> 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.<ref name=ScienceDaily24Aug06Sour>{{citation |date=24 August 2006 |title=Biologists Discover How We Detect Sour Taste |work=[[Science Daily]] |url=https://www.sciencedaily.com/releases/2006/08/060823184824.htm |access-date=12 September 2010 |archive-date=30 October 2009 |archive-url=https://web.archive.org/web/20091030174710/http://www.sciencedaily.com/releases/2006/08/060823184824.htm |url-status=live }}</ref>

===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.<ref>{{cite journal |vauthors=Maehashi K, Matano M, Wang H, Vo LA, Yamamoto Y, Huang L |title=Bitter peptides activate hTAS2Rs, the human bitter receptors |journal=Biochemical and Biophysical Research Communications |volume=365 |issue=4 |pages=851–5 |date=January 2008 |pmid=18037373 |pmc=2692459 |doi=10.1016/j.bbrc.2007.11.070}}</ref> 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).<ref>{{cite journal |last=Lindemann |first=B |title=Receptors and transduction in taste |journal=Nature |volume=413 |issue=6852 |pages=219–25 |date=September 2001 |pmid=11557991 |doi=10.1038/35093032|bibcode=2001Natur.413..219L |s2cid=4385513 }}</ref>

===Savoriness===
The [[amino acid]] [[glutamic acid]] is responsible for savoriness,<ref name=umami2>[http://www.umamiinfo.com/what_exactly_is_umami?/ What Is Umami?: What Exactly is Umami?] {{Webarchive|url=https://web.archive.org/web/20110423145431/http://www.umamiinfo.com/what_exactly_is_umami?%2F |date=23 April 2011 }} Umami Information Center</ref><ref name=ChandrashekarHoonEtal2006>{{Citation |date=16 November 2006 |last1=Chandrashekar|first1= Jayaram|last2=Hoon|first2=Mark A|last3= Ryba|first3=Nicholas J. P. |last4=Zuker|first4= Charles S |name-list-style=amp |title=The receptors and cells for mammalian taste |journal=[[Nature (journal)|Nature]] |pmid=17108952 |volume=444 |issue=7117 |pages=288–294 |doi=10.1038/nature05401 |url=https://wiki.brown.edu/confluence/download/attachments/1444406/nature05401.pdf |access-date=13 September 2010 |bibcode=2006Natur.444..288C |s2cid=4431221|archive-url=https://web.archive.org/web/20110722041026/https://wiki.brown.edu/confluence/download/attachments/1444406/nature05401.pdf |archive-date=22 July 2011 |url-status=dead }}</ref> but some [[nucleotide]]s ([[inosinic acid]]<ref name=jmin/><ref name=umami3/> and [[guanylic acid]]<ref name=umami2/>) can act as complements, enhancing the taste.<ref name=jmin>{{Cite web |url=https://tasteofjapan.maff.go.jp/en/ingredient/umami/ |title=Essiential Ingredients of Japanese Food – Umami |access-date=2022-04-20 |website=Taste of Japan |archive-url=https://web.archive.org/web/20210516051532/https://tasteofjapan.maff.go.jp/en/ingredient/umami/ |archive-date=2021-05-16 |url-status=live |publisher=[[Ministry of Agriculture, Forestry and Fisheries (Japan)]]}}</ref><ref name=umami3>[http://www.umamiinfo.com/what_is_umami?/what_is_umami?/the_composition_of_umami/ What Is Umami?: The Composition of Umami] {{Webarchive|url=https://web.archive.org/web/20090527004141/http://www.umamiinfo.com/what_is_umami?%2Fwhat_is_umami%3F%2Fthe_composition_of_umami%2F |date=27 May 2009 }} Umami Information Center</ref>

Glutamic acid binds to a variant of the G protein-coupled receptor, producing a [[umami|savory]] taste.<ref name="Lindemann B 99–100"/><ref name="Chaud" />

==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)===
{{Main|Pungency|Scoville scale}}

Substances such as [[ethanol]] and [[capsaicin]] cause a burning sensation by inducing a trigeminal nerve reaction together with normal taste reception. The sensation of heat is caused by the food's activating nerves that express [[TRPV1]] and [[TRPA1]] receptors. Some such plant-derived compounds that provide this sensation are capsaicin from [[chili pepper]]s, [[piperine]] from [[black pepper]], [[gingerol]] from [[ginger root]] and [[allyl isothiocyanate]] from [[horseradish]]. The [[pungency|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 [[Cuisine of Ethiopia|Ethiopian]], [[peruvian cuisine|Peruvian]], [[Hungarian cuisine|Hungarian]], [[Indian cuisine|Indian]], [[Cuisine of Korea|Korean]], [[Indonesian cuisine|Indonesian]], [[Cuisine of Laos|Lao]], [[Malaysian cuisine|Malaysian]], [[Mexican cuisine|Mexican]], [[New Mexican cuisine|New Mexican]], [[Pakistani cuisine|Pakistani]], [[Singaporean cuisine|Singaporean]], [[Southwest China|Southwest Chinese]] (including [[Sichuan cuisine]]), [[Vietnamese cuisine|Vietnamese]], and [[Cuisine of Thailand|Thai]] cuisines.

This particular sensation, called [[chemesthesis]], is not a taste in the technical sense, because the sensation does not arise from taste buds, and a different set of nerve fibers carry it to the brain. Foods like chili peppers activate nerve fibers directly; the sensation interpreted as "hot" results from the stimulation of somatosensory (pain/temperature) fibers on the tongue. Many parts of the body with exposed membranes but no taste sensors (such as the nasal cavity, under the fingernails, [[Cornea|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]] and [[spearmint]] and is triggered by substances such as [[menthol]], [[anethol]], ethanol, and [[camphor]]. Caused by activation of the same mechanism that signals cold, [[TRPM8]] ion channels on [[neuron|nerve cells]], unlike the actual change in temperature described for sugar substitutes, this coolness is only a perceived phenomenon.

===Numbness===
Both Chinese and [[Toba Batak people|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 cuisine|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.<ref>{{cite web|url=http://gernot-katzers-spice-pages.com/engl/Zant_pip.html|title=Spice Pages: Sichuan Pepper (Zanthoxylum, Szechwan peppercorn, fagara, hua jiao, sansho 山椒, timur, andaliman, tirphal)|first=Gernot|last=Katzer|website=gernot-katzers-spice-pages.com|access-date=16 May 2013|archive-date=19 November 2012|archive-url=https://web.archive.org/web/20121119072929/http://gernot-katzers-spice-pages.com/engl/Zant_pip.html|url-status=live}}</ref>
Typical in northern Brazilian cuisine, [[Acmella oleracea|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 [[tannins]] or [[calcium oxalate]] that cause an astringent or puckering sensation of the mucous membrane of the mouth. Examples include [[tea]], [[red wine]], or [[rhubarb]].{{citation needed|date=April 2021}} Other terms for the astringent sensation are "dry", "rough", "harsh" (especially for wine), "tart" (normally referring to sourness), "rubbery", "hard" or "styptic".<ref>{{cite journal|last1=Peleg|first1=Hanna|last2=Gacon|first2=Karine|last3=Schlich|first3=Pascal|last4=Noble|first4=Ann C|title=Bitterness and astringency of flavan-3-ol monomers, dimers and trimers|journal=Journal of the Science of Food and Agriculture|date=June 1999|volume=79|issue=8|pages=1123–1128|doi=10.1002/(SICI)1097-0010(199906)79:8<1123::AID-JSFA336>3.0.CO;2-D}}</ref>

===Metallicness===
A metallic taste may be caused by food and drink, certain medicines or [[amalgam (dentistry)|amalgam]] dental fillings. It is generally considered an off flavor when present in food and drink. A metallic taste may be caused by [[Galvanic cell|galvanic]] reactions in the mouth. In the case where it is caused by dental work, the dissimilar metals used may produce a measurable current.<ref>{{cite web|url=http://www.kcdentalworks.com/blog/2019/4/24/could-your-mouth-charge-your-iphone|title=Could your mouth charge your iPhone?|date=April 24, 2019|publisher=kcdentalworks.com|access-date=3 May 2019|archive-date=3 May 2019|archive-url=https://web.archive.org/web/20190503145204/http://www.kcdentalworks.com/blog/2019/4/24/could-your-mouth-charge-your-iphone|url-status=live}}</ref> Some artificial sweeteners are perceived to have a metallic taste, which is detected by the [[TRPV1]] receptors.<ref>{{cite journal |last1=Riera |first1=Céline E. |last2=Vogel |first2=Horst |last3=Simon |first3=Sidney A. |last4=le Coutre |first4=Johannes |title=Artificial sweeteners and salts producing a metallic taste sensation activate TRPV1 receptors. |date=2007 |journal=American Journal of Physiology |volume=293 |number=2 |pages=R626–R634 |doi=10.1152/ajpregu.00286.2007 |pmid=17567713 }}</ref> Many people consider [[blood]] to have a metallic taste.<ref>{{cite journal|last=Willard|first=James P.|year=1905|title=Current Events|journal=Progress: A Monthly Journal Devoted to Medicine and Surgery|volume=4|pages=861–68|url=https://books.google.com/books?id=6xygAAAAMAAJ&pg=PA862}}</ref><ref>{{cite book|last=Monosson|first=Emily|title=Evolution in a Toxic World: How Life Responds to Chemical Threats|url=https://books.google.com/books?id=vqtrn8iwtecC&pg=PA49|year=2012|publisher=Island Press|isbn=9781597269766|page=49}}</ref> A metallic taste in the mouth is also a symptom of various medical conditions, in which case it may be classified under the symptoms [[dysgeusia]] or [[parageusia]], referring to distortions of the sense of taste,<ref name=goldstein>{{cite book|last=Goldstein|first=E. Bruce|title=Encyclopedia of Perception|url=https://books.google.com/books?id=6M3NSNm6MlkC&pg=PA959|volume=2|year=2010|publisher=SAGE|isbn=9781412940818|pages=958–59}}</ref> and can be caused by medication, including [[saquinavir]],<ref name=goldstein/> [[zonisamide]],<ref>{{cite book|last=Levy|first=René H.|title=Antiepileptic Drugs|url=https://books.google.com/books?id=HAOY0qG-vAYC&pg=PA875|year=2002|publisher=Lippincott Williams & Wilkins|isbn=9780781723213|page=875}}</ref> and various kinds of [[chemotherapy]],<ref>{{cite journal |last1=Reith |first1=Alastair J. M. |last2=Spence |first2=Charles |title=The mystery of "metal mouth" in chemotherapy. |date=2020 |journal=Chemical Senses |volume=45 |number=2 |pages=73–84 |doi=10.1093/chemse/bjz076 |pmid=32211901 |url=https://ora.ox.ac.uk/objects/uuid:bf56172d-ea67-4880-8b45-d13fde173cd9 |doi-access=free |access-date=15 October 2020 |archive-date=14 April 2021 |archive-url=https://web.archive.org/web/20210414173945/https://ora.ox.ac.uk/objects/uuid:bf56172d-ea67-4880-8b45-d13fde173cd9 |url-status=live }}</ref> as well as occupational hazards, such as working with [[pesticide]]s.<ref>{{cite book|last=Stellman|first=Jeanne Mager|title=Encyclopaedia of Occupational Health and Safety: The body, health care, management and policy, tools and approaches|url=https://books.google.com/books?id=vW6rXFvm4sQC&pg=PT299|year=1998|publisher=International Labour Organization|isbn=9789221098140|page=299}}</ref>

===Fat taste===
Recent research reveals a potential [[taste receptor]] called the [[CD36|CD36 receptor]].<ref>{{cite web|url=http://www.scientificamerican.com/article/potential-taste-receptor/|title=Potential Taste Receptor for Fat Identified|first=David|last=Biello|website=[[Scientific American]]|access-date=20 January 2015|archive-date=9 December 2014|archive-url=https://web.archive.org/web/20141209122639/http://www.scientificamerican.com/article/potential-taste-receptor/|url-status=live}}</ref><ref>{{Cite journal
| pmid = 16276419
| pmc = 1265871
| year = 2005
| last1 = Laugerette
| first1 = F
| title = CD36 involvement in orosensory detection of dietary lipids, spontaneous fat preference, and digestive secretions
| journal = Journal of Clinical Investigation
| volume = 115
| issue = 11
| pages = 3177–84
| last2 = Passilly-Degrace
| first2 = P
| last3 = Patris
| first3 = B
| last4 = Niot
| first4 = I
| last5 = Febbraio
| first5 = M
| last6 = Montmayeur
| first6 = J. P.
| last7 = Besnard
| first7 = P
| doi = 10.1172/JCI25299
}}</ref><ref>{{Cite journal
| pmid = 24631296
| year = 2014
| last1 = Dipatrizio
| first1 = N. V.
| title = Is fat taste ready for primetime?
| journal = Physiology & Behavior
| volume = 136C
| pages = 145–154
| doi = 10.1016/j.physbeh.2014.03.002
| pmc=4162865
}}</ref> CD36 was targeted as a possible lipid taste receptor because it binds to [[fat]] molecules (more specifically, long-chain [[fatty acid]]s),<ref>{{Cite journal
| pmid = 8694909
| year = 1996
| last1 = Baillie
| first1 = A. G.
| title = Reversible binding of long-chain fatty acids to purified FAT, the adipose CD36 homolog
| journal = The Journal of Membrane Biology
| volume = 153
| issue = 1
| pages = 75–81
| last2 = Coburn
| first2 = C. T.
| last3 = Abumrad
| first3 = N. A.
| doi=10.1007/s002329900111
| s2cid = 5911289
}}</ref> and it has been localized to [[taste bud]] cells (specifically, the circumvallate and foliate [[lingual papillae|papillae]]).<ref>{{Cite journal
| pmid = 20950842
| year = 2011
| last1 = Simons
| first1 = P. J.
| title = Apical CD36 immunolocalization in human and porcine taste buds from circumvallate and foliate papillae
| journal = Acta Histochemica
| volume = 113
| issue = 8
| pages = 839–43
| last2 = Kummer
| first2 = J. A.
| last3 = Luiken
| first3 = J. J.
| last4 = Boon
| first4 = L
| doi = 10.1016/j.acthis.2010.08.006
}}</ref> There is a debate over whether we can truly taste fats, and supporters of human 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 system|gustatory]] neurons, similar to other currently accepted tastes; and, there is a physiological response to the presence of oral fat.<ref name="pmid21557960">{{Cite journal
| pmid = 21557960
| pmc = 3139746
| year = 2011
| last1 = Mattes
| first1 = R. D.
| title = Accumulating evidence supports a taste component for free fatty acids in humans
| journal = Physiology & Behavior
| volume = 104
| issue = 4
| pages = 624–31
| doi = 10.1016/j.physbeh.2011.05.002
}}</ref> Although CD36 has been studied primarily in [[house mouse|mice]], research examining human subjects' ability to taste fats found that those with high levels of CD36 [[gene expression|expression]] were more sensitive to tasting fat than were those with low levels of CD36 expression;<ref>{{Cite journal
| pmid = 22210925
| pmc = 3276480
| year = 2012
| last1 = Pepino
| first1 = M. Y.
| title = The fatty acid translocase gene CD36 and lingual lipase influence oral sensitivity to fat in obese subjects
| journal = The Journal of Lipid Research
| volume = 53
| issue = 3
| pages = 561–6
| last2 = Love-Gregory
| first2 = L
| last3 = Klein
| first3 = S
| last4 = Abumrad
| first4 = N. A.
| doi = 10.1194/jlr.M021873
 |doi-access=free }}</ref> 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 receptor]]s [[free fatty acid receptor 4]] (also termed GPR120) and to a much lesser extent [[Free fatty acid receptor 1]] (also termed GPR40)<ref name="pmid31487233">{{cite journal | vauthors = Kimura I, Ichimura A, Ohue-Kitano R, Igarashi M | title = Free Fatty Acid Receptors in Health and Disease | journal = Physiological Reviews | volume = 100 | issue = 1 | pages = 171–210 | date = January 2020 | pmid = 31487233 | doi = 10.1152/physrev.00041.2018 | url = | doi-access = free }}</ref> 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.<ref>{{Cite journal
| pmid = 20573884
| year = 2010
| last1 = Cartoni
| first1 = C
| title = Taste preference for fatty acids is mediated by GPR40 and GPR120
| journal = Journal of Neuroscience
| volume = 30
| issue = 25
| pages = 8376–82
| last2 = Yasumatsu
| first2 = K
| last3 = Ohkuri
| first3 = T
| last4 = Shigemura
| first4 = N
| last5 = Yoshida
| first5 = R
| last6 = Godinot
| first6 = N
| last7 = Le Coutre
| first7 = J
| last8 = Ninomiya
| first8 = Y
| last9 = Damak
| first9 = S
| doi = 10.1523/JNEUROSCI.0496-10.2010
| pmc = 6634626
}}</ref>

Monovalent cation channel [[TRPM5]] has been implicated in fat taste as well,<ref>{{Cite journal
| pmid = 21653867
| pmc = 3125678
| year = 2011
| last1 = Liu
| first1 = P
| title = Transient receptor potential channel type M5 is essential for fat taste
| journal = Journal of Neuroscience
| volume = 31
| issue = 23
| pages = 8634–42
| last2 = Shah
| first2 = B. P.
| last3 = Croasdell
| first3 = S
| last4 = Gilbertson
| first4 = T. A.
| doi = 10.1523/JNEUROSCI.6273-10.2011
}}</ref> 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.<ref name="pmid21557960"/>

Proposed alternate names to fat taste include oleogustus<ref>{{cite journal |last1=Running |first1=Cordelia A. |last2=Craig |first2=Bruce A. |last3=Mattes |first3=Richard D. |date=3 July 2015 |title=Oleogustus: The Unique Taste of Fat |journal=Chemical Senses |volume=40 |issue=6 |pages= 507–516|doi=10.1093/chemse/bjv036 |pmid=26142421 |doi-access=free }}</ref> and pinguis,<ref name="Reed 353S–360S"/> 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. A high concentration of fatty acids in certain foods is generally considered inedible.<ref name="Purdue">{{cite web |url=http://www.purdue.edu/newsroom/releases/2015/Q3/research-confirms-fat-is-sixth-taste-names-it-oleogustus.html |title=Research confirms fat is sixth taste; names it oleogustus |last1=Neubert |first1=Amy Patterson |date=23 July 2015 |website=Purdue News |publisher=[[Purdue University]] |access-date=4 August 2015 |archive-date=8 August 2015 |archive-url=https://web.archive.org/web/20150808232426/http://www.purdue.edu/newsroom/releases/2015/Q3/research-confirms-fat-is-sixth-taste-names-it-oleogustus.html |url-status=live }}</ref> 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.<ref name="oleogustus">{{cite news|title=Is fat the sixth taste primary? Evidence and implications|last=Keast|first=Russell|journal=Flavour |date=3 February 2015|volume=4 |doi = 10.1186/2044-7248-4-5 |doi-access=free }}</ref>

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.<ref>{{cite news |last=Feldhausen |first=Teresa Shipley |date=31 July 2015 |title=The five basic tastes have sixth sibling: oleogustus |url=https://www.sciencenews.org/article/five-basic-tastes-have-sixth-sibling-oleogustus |newspaper=[[Science News]] |access-date=4 August 2015 |archive-date=16 August 2015 |archive-url=https://web.archive.org/web/20150816230541/https://www.sciencenews.org/article/five-basic-tastes-have-sixth-sibling-oleogustus |url-status=live }}</ref>

=== Heartiness ===
''Kokumi'' ({{IPAc-en|k|oʊ|k|uː|m|i}}, Japanese: {{nihongo||コク味|kokumi}}<ref name="Nishimura2016">{{Cite magazine|title="Koku" Involved in Food Palatability: An Overview of Pioneering Work and Outstanding Questions|date=2016-01-20|magazine=Kagaku to Seibutsu|url=https://katosei.jsbba.or.jp/view_html.php?aid=527|last1=Nishimura|first1=Toshihide|doi=10.1271/kagakutoseibutsu.54.102|pages=102–108|access-date=2020-08-11|last2=Egusa|first2=Ai|publisher=Japan Society for Bioscience, Biotechnology, and Agrochemistry (JSBBA)|issue=54|volume=2|language=ja|script-title=ja:食べ物の「こく」を科学するその現状と展望|quote=「こく」appears in abstract. 「コク味物質」appears in p106 1.b}}</ref> from {{nihongo||こく|koku}}<ref name="Nishimura2016" />) 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'' has been described as something that may enhance the other five tastes by magnifying and lengthening the other tastes, or "mouthfulness".<ref name="foodprot">{{cite book |editor1-last=Hettiarachchy |editor1-first=Navam S. |editor2-last=Sato |editor2-first=Kenji |editor3-last=Marshall |editor3-first=Maurice R. |title=Food proteins and peptides: chemistry, functionality interactions, and commercialization |date=2010 |publisher=CRC |location=Boca Raton, Fla. |isbn=9781420093414 |url=https://books.google.com/books?id=-h8UEImN7SAC&q=kokumi&pg=PA290 |access-date=26 June 2014 }}</ref>{{rp|290}}<ref name=":0">{{cite journal | last1 = Ueda | first1 = Yoichi | last2 = Sakaguchi | first2 = Makoto | last3 = Hirayama | first3 = Kazuo | last4 = Miyajima | first4 = Ryuichi | last5 = Kimizuka | first5 = Akimitsu | year = 1990 | title = Characteristic Flavor Constituents in Water Extract of Garlic | journal = Agricultural and Biological Chemistry | volume = 54 | issue = 1| pages = 163–169 | doi = 10.1080/00021369.1990.10869909 }}</ref> Garlic is a common ingredient to add flavor used to help define the characteristic ''kokumi'' flavors.<ref name=":0" />

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.<ref name=foodprot/> This subset of CaSR-expressing taste cells are independent from the influenced basic taste receptor cells.<ref>{{Cite journal|last1=Eto|first1=Yuzuru|last2=Kuroda|first2=Motonaka|last3=Yasuda|first3=Reiko|last4=Maruyama|first4=Yutaka|date=2012-04-12|title=Kokumi Substances, Enhancers of Basic Tastes, Induce Responses in Calcium-Sensing Receptor Expressing Taste Cells|journal=PLOS ONE|language=en|volume=7|issue=4|pages=e34489|doi=10.1371/journal.pone.0034489|pmid=22511946|pmc=3325276|bibcode=2012PLoSO...734489M|issn=1932-6203|doi-access=free}}</ref> 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.<ref>{{Cite journal|last1=Eto|first1=Yuzuru|last2=Miyamura|first2=Naohiro|last3=Maruyama|first3=Yutaka|last4=Hatanaka|first4=Toshihiro|last5=Takeshita|first5=Sen|last6=Yamanaka|first6=Tomohiko|last7=Nagasaki|first7=Hiroaki|last8=Amino|first8=Yusuke|last9=Ohsu|first9=Takeaki|date=2010-01-08|title=Involvement of the Calcium-sensing Receptor in Human Taste Perception|journal=Journal of Biological Chemistry|language=en|volume=285|issue=2|pages=1016–1022|doi=10.1074/jbc.M109.029165|issn=0021-9258|pmid=19892707|pmc=2801228|doi-access=free}}</ref>

===Calcium===
The distinctive taste of chalk has been identified as the calcium component of that substance.<ref>{{cite web|url=http://www.scientificamerican.com/article/osteoporosis-calcium-taste-chalk/|title=Like the Taste of Chalk? You're in Luck—Humans May Be Able to Taste Calcium.|publisher=Scientific American|date=20 August 2008|access-date=14 March 2014|archive-date=28 March 2014|archive-url=https://web.archive.org/web/20140328072729/http://www.scientificamerican.com/article/osteoporosis-calcium-taste-chalk/|url-status=live}}</ref> In 2008, geneticists discovered a [[calcium receptor]] on the tongues of [[mice]]. The CaSR receptor is commonly found in the [[gastrointestinal tract]], [[kidneys]], and [[brain]]. Along with the "sweet" T1R3 receptor, the CaSR receptor can detect calcium as a taste. Whether the perception exists or not in humans is unknown.<ref>{{Citation|first = Michael G.|last = Tordorf|contribution = Chemosensation of Calcium|title = American Chemical Society National Meeting, Fall 2008, 236th|year = 2008|pages = AGFD 207|place = Philadelphia, PA|publisher = American Chemical Society|contribution-url = http://portal.acs.org/portal/acs/corg/content?_nfpb=true&_pageLabel=PP_TRANSITIONMAIN&node_id=859&use_sec=false&sec_url_var=region1|no-pp = true|author-link = Michael Tordoff|title-link = American Chemical Society|access-date = 27 August 2008|archive-date = 25 August 2009|archive-url = https://web.archive.org/web/20090825091613/http://portal.acs.org/portal/acs/corg/content?_nfpb=true&_pageLabel=PP_TRANSITIONMAIN&node_id=859&use_sec=false&sec_url_var=region1|url-status = live}}</ref><ref name="ScienceDaily21Aug08Calcium">{{citation |date=21 August 2008 |title=That Tastes ... Sweet? Sour? No, It's Definitely Calcium! |url=https://www.sciencedaily.com/releases/2008/08/080820163008.htm |work=Science Daily |access-date=14 September 2010 |archive-date=18 October 2009 |archive-url=https://web.archive.org/web/20091018075828/http://www.sciencedaily.com/releases/2008/08/080820163008.htm |url-status=live }}</ref>

===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]]. In North America it is almost always preferred cold, regardless of season.

===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.<ref>{{Cite journal|last1=Lapis|first1=Trina J.|last2=Penner|first2=Michael H.|last3=Lim|first3=Juyun|date=23 August 2016|title=Humans Can Taste Glucose Oligomers Independent of the hT1R2/hT1R3 Sweet Taste Receptor|url=https://nature.berkeley.edu/garbelottoat/wp-content/uploads/lapis-etal-2106.pdf|journal=Chemical Senses|language=en|pages=755–762|doi=10.1093/chemse/bjw088|issn=0379-864X|pmid=27553043|volume=41|issue=9|doi-access=free|access-date=26 September 2017|archive-date=26 September 2017|archive-url=https://web.archive.org/web/20170926144219/https://nature.berkeley.edu/garbelottoat/wp-content/uploads/lapis-etal-2106.pdf|url-status=live}}</ref><ref>{{cite journal|first1=Alexa J.|last1=Pullicin|first2=Michael H.|last2=Penner|first3=Juyun|last3=Lim|title=Human taste detection of glucose oligomers with low degree of polymerization|journal=PLOS ONE|date=29 August 2017|issn=1932-6203|pages=e0183008|volume=12|issue=8|doi=10.1371/journal.pone.0183008|pmid=28850567|pmc=5574539|bibcode=2017PLoSO..1283008P|doi-access=free}}</ref><ref>{{Cite web|url=https://www.newscientist.com/article/2104244-there-is-now-a-sixth-taste-and-it-explains-why-we-love-carbs/|title=There is now a sixth taste – and it explains why we love carbs|last=Hamzelou|first=Jessica|date=2 September 2016|website=[[New Scientist]]|language=en-US|access-date=14 September 2016|archive-date=16 September 2016|archive-url=https://web.archive.org/web/20160916071519/https://www.newscientist.com/article/2104244-there-is-now-a-sixth-taste-and-it-explains-why-we-love-carbs/|url-status=live}}</ref>

==Nerve supply and neural connections==
[[File:Jaime Lara 2023 PMID 36409650 Figure 8.jpg|thumb|Active brain areas in taste perception]]
[[File:Comprehensive List of Relevant Pathways for the Gustatory System.png|thumb|This diagram linearly (unless otherwise mentioned) tracks the projections of all known structures that allow for taste to their relevant endpoints in the human brain.]]

The [[glossopharyngeal nerve#Overview of special sensory component|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]].<ref>Eliav, Eli, and Batya Kamran. "Evidence of Chorda Tympani Dysfunction in Patients with Burning Mouth Syndrome." [[ScienceDirect|Science Direct]]. May 2007. Web. 27 March 2016.</ref>

The [[Pterygopalatine ganglion|pterygopalatine ganglia]] are ganglia (one on each side) of the [[soft palate]]. The [[greater petrosal nerve|greater petrosal]], [[lesser palatine nerve|lesser palatine]]  and [[zygomatic nerve]]s 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 nerve]]s (from the [[trigeminal nerve]]).

The [[special visceral afferent]]s of the [[vagus nerve]] carry taste from the [[epiglottis|epiglottal]] 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.<ref>Mu, Liancai, and Ira Sanders. "Human Tongue Neuroanatomy: Nerve Supply and Motor Endplates." Wiley Online Library. Oct. 2010. Web. 27 March 2016.</ref> 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.<ref>King, Camillae T., and Susan P. Travers. "Glossopharyngeal Nerve Transection Eliminates Quinine-Stimulated Fos-Like Immunoreactivity in the Nucleus of the Solitary Tract: Implications for a Functional Topography of Gustatory Nerve Input in Rats." JNeurosci. 15 April 1999. Web. 27 March 2016.</ref>

Reticular formation (includes Raphe nuclei responsible for serotonin production) is signaled to release serotonin during and after a meal to suppress appetite.<ref>Hornung, Jean-Pierre. "The Human Raphe Nuclei and the Serotonergic System."Science Direct. Dec. 2003. Web. 27 March 2016.</ref> Similarly, salivary nuclei are signaled to decrease saliva secretion.

[[Hypoglossal nerve|Hypoglossal]] and [[thalamus|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|Edinger-Westphal nucleus]] reacts to taste stimuli by dilating and constricting the pupils.<ref>Reiner, Anton, and Harvey J. Karten. "Parasympathetic Ocular Control — Functional Subdivisions and Circuitry of the Avian Nucleus of Edinger-Westphal."Science Direct. 1983. Web. 27 March 2016.</ref>

Spinal ganglion are involved in movement.

The [[Operculum (brain)|frontal operculum]] is speculated to be the memory and association hub for taste.{{citation needed|date=April 2016}}

The [[insula cortex]] aids in swallowing and gastric motility.<ref>Wright, Christopher I., and Brain Martis. "Novelty Responses and Differential Effects of Order in the Amygdala, Substantia Innominata, and Inferior Temporal Cortex." Science Direct. Mar. 2003. Web. 27 March 2016.</ref><ref>Menon, Vinod, and Lucina Q. Uddin. "Saliency, Switching, Attention and Control: A Network Model of Insula." Springer. 29 May 2010. Web. 28 March 2016.</ref>

==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===
{{Main|Supertaster}}

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]].<ref>{{cite journal |author1=Bartoshuk L. M. |author2=Duffy V. B. | year = 1994 | title = PTC/PROP tasting: anatomy, psychophysics, and sex effects." 1994 | journal = Physiol Behav | volume = 56 | issue = 6| pages = 1165–71 | doi=10.1016/0031-9384(94)90361-1 | pmid=7878086|s2cid=40598794 |display-authors=etal| doi-access=free }}</ref> Studies have shown that supertasters require less fat and sugar in their food to get the same satisfying effects. These people tend to consume more salt than others. 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.)<ref>{{cite news|last=Gardner|first=Amanda|title=Love salt? You might be a 'supertaster'|url=http://www.cnn.com/2010/HEALTH/06/16/salt.taste/index.html|publisher=CNN Health|access-date=9 April 2012|date=16 June 2010|archive-date=9 April 2012|archive-url=https://web.archive.org/web/20120409095146/http://www.cnn.com/2010/HEALTH/06/16/salt.taste/index.html|url-status=live}}</ref>

===Aftertaste===
{{Main|Aftertaste}}

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

===Acquired taste===
{{Main|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.<ref>{{cite book|chapter-url=https://www.ncbi.nlm.nih.gov/books/NBK385/|title=Clinical Methods: The History, Physical, and Laboratory Examinations|last=Walker|first=H. Kenneth|year=1990|access-date=1 May 2014|isbn=9780409900774|chapter=Cranial Nerve VII: The Facial Nerve and Taste|publisher=Butterworths|archive-date=26 January 2016|archive-url=https://web.archive.org/web/20160126215348/http://www.ncbi.nlm.nih.gov/books/NBK385/|url-status=live}}</ref>

===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 [[Impact of COVID-19 on neurological, psychological and other mental health outcomes|SARS-CoV-2]] (causing COVID-19) experience some type of [[Impact of COVID-19 on neurological, psychological and other mental health outcomes|disorder associated with their sense of smell or taste]], including [[ageusia]] and [[dysgeusia]]. [[SARS-CoV-1]], [[Middle East respiratory syndrome–related coronavirus|MERS-CoV]] and even the flu ([[Orthomyxoviridae|influenza virus]]) can also disrupt olfaction.<ref>{{Cite journal|last1=Meunier|first1=Nicolas|last2=Briand|first2=Loïc|last3=Jacquin-Piques|first3=Agnès|last4=Brondel|first4=Laurent|last5=Pénicaud|first5=Luc|date=2020|title=COVID 19-Induced Smell and Taste Impairments: Putative Impact on Physiology|journal=Frontiers in Physiology|volume=11|pages=625110|doi=10.3389/fphys.2020.625110|issn=1664-042X|pmc=7870487|pmid=33574768|doi-access=free}}</ref><ref>{{Cite journal|last1=Veronese|first1=Sheila|last2=Sbarbati|first2=Andrea|date=2021-03-03|title=Chemosensory Systems in COVID-19: Evolution of Scientific Research|journal=ACS Chemical Neuroscience|volume=12|issue=5|pages=813–824|doi=10.1021/acschemneuro.0c00788|issn=1948-7193|pmc=7885804|pmid=33559466}}</ref>

==History==

In [[The Western world|the West]], [[Aristotle]] postulated in {{circa|350}} [[Before Christ|BC]]<ref>[http://classics.mit.edu/Aristotle/soul.html On the Soul] {{Webarchive|url=https://web.archive.org/web/20110106095759/http://classics.mit.edu/Aristotle/soul.html |date=6 January 2011 }} Aristotle. Translated by J. A. Smith. The Internet Classics Archive.</ref> that the two most basic tastes were sweet and bitter.<ref>[https://books.google.com/books?id=QPnxaraJ7LQC&q=succulent&pg=PA193 Aristotle's De anima (422b10-16)] {{Webarchive|url=https://web.archive.org/web/20230326070951/https://books.google.com/books?id=QPnxaraJ7LQC&q=succulent&pg=PA193 |date=26 March 2023 }} Ronald M. Polansky. Cambridge University Press, 2007.</ref> He was one of the first persons to develop a list of basic tastes.<ref>[https://books.google.com/books?id=_GMeW9E1IB4C&dq=Aristotle+basic+taste&pg=PA165 Origins of neuroscience: a history of explorations into brain function (Page 165/480)] {{Webarchive|url=https://web.archive.org/web/20230326070950/https://books.google.com/books?id=_GMeW9E1IB4C&dq=Aristotle+basic+taste&pg=PA165 |date=26 March 2023 }} Stanley Finger. Oxford University Press US, 2001.</ref>

==Research==
The [[Receptor (biochemistry)|receptors]] for the basic tastes of bitter, sweet and savory have been identified. They are [[G protein-coupled receptor]]s.<ref name="bach">{{Cite journal | last1 = Bachmanov | first1 = AA. | last2 = Beauchamp | first2 = GK. | title = Taste receptor genes. | journal = Annu Rev Nutr | volume = 27 | issue = 1| pages = 389–414 | year = 2007 | doi = 10.1146/annurev.nutr.26.061505.111329 | pmid = 17444812 | pmc=2721271}}</ref> 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.<ref name=bach/> As of 2019, molecular mechanisms for each taste appear to be different, although all taste perception relies on activation of [[P2X purinoreceptor]]s on [[sensory nerve]]s.<ref name="kinn">{{cite journal |vauthors=Kinnamon SC, Finger TE |title=Recent advances in taste transduction and signaling |journal=F1000Research |volume=8 |date=2019 |page=2117 |pmid=32185015 |pmc=7059786 |doi=10.12688/f1000research.21099.1 |doi-access=free }}</ref>

==See also==
{{Portal|Food}}
<div col||20em|small=yes>
* [[Beefy meaty peptide]]
* [[Digital lollipop]]
* [[Optimal foraging theory]]
* [[Palatability]]
* [[Mouthfeel]]
* [[Vomeronasal organ]]
* [[Sensory analysis]]
* [[Tea tasting]]
* [[Wine tasting]]
</div col>

==Notes==
{{refbegin}}

'''a.''' {{Note label|a|a|none}} 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:<blockquote>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",<ref name=Guyton1976>{{citation |title=Textbook of Medical Physiology |last=Guyton |first=Arthur C. |author-link=Arthur Guyton |year=1976 |edition=5th |page=[https://archive.org/details/textbookofmedica0005guyt/page/839 839] |publisher=W.B. Saunders |location=Philadelphia |isbn=978-0-7216-4393-9 |url=https://archive.org/details/textbookofmedica0005guyt/page/839 }}</ref></blockquote>

'''b.''' {{Note label|b|b|none}} 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"<ref name=Macbeth&MacClancy2004>{{citation |year=2004 |editor=Macbeth, Helen M. |editor2= MacClancy, Jeremy |chapter=plethora of methods characterising human taste perception |pages=87–88|chapter-url=https://books.google.com/books?id=ZLQTSqfB5igC&pg=PA88 |title=Researching Food Habits: Methods and Problems |series=The anthropology of food and nutrition |volume=5 |url=https://books.google.com/books?id=ZLQTSqfB5igC |place=New York |publisher=Berghahn Books |isbn=9781571815446|access-date=15 September 2010}}</ref> 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.<ref name=Guyton&Hall2006>{{citation |year=2006 |last1=Guyton |first1=Arthur C |author-link=Arthur Guyton |last2=Hall |first2=John E. |title=Guyton and Hall Textbook of Medical Physiology |page=664 |edition=11th |publisher=Elsevier Saunders |location=Philadelphia |isbn=978-0-7216-0240-0}}</ref>

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,<ref name="textbookofmedicalphysiology8thed"/><ref name=McLaughlin&Margolskee/> is essentially the same as that of Svrivastava & Rastogi (2003),<ref name=Svrivastava&Rastogi2003>{{Citation |year=2003 |author1=Svrivastava, R.C. |author2=Rastogi, R.P |name-list-style=amp |chapter=Relative taste indices of some substances |chapter-url=https://books.google.com/books?id=hIyM_o4YFZ4C&pg=PA274 |title=Transport Mediated by Electrical Interfaces |series=Studies in interface science |volume=18 |place=Amsterdam, Netherlands |publisher=Elsevier Science |isbn=978-0-444-51453-0|url=https://books.google.com/books?id=hIyM_o4YFZ4C |access-date=12 September 2010 |postscript=Taste indices of table 9, p.274 are select sample taken from table in Guyton's ''Textbook of Medical Physiology'' (present in all editions)}}</ref> Guyton & Hall (2006),<ref name=Guyton&Hall2006/> and Joesten ''et al.'' (2007).<ref name=JoestenEal2007/> 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.
{{refend}}

==References==
{{Reflist}}

==Further reading==
*{{Citation |date=16 November 2006 |last1=Chandrashekar|first1=Jayaram|last2=Hoon|first2=Mark A.|last3=Ryba|last4=Nicholas |first4=J. P. |last5=Zuker|first5=Charles S. |name-list-style=amp |title=The receptors and cells for mammalian taste |journal=[[Nature (journal)|Nature]] |pmid=17108952 |volume=444 |issue=7117 |pages=288–294 |doi=10.1038/nature05401 |url=https://wiki.brown.edu/confluence/download/attachments/1444406/nature05401.pdf |access-date=13 September 2010 |bibcode=2006Natur.444..288C |s2cid=4431221|archive-url=https://web.archive.org/web/20110722041026/https://wiki.brown.edu/confluence/download/attachments/1444406/nature05401.pdf |archive-date=22 July 2011 |url-status=dead }}
*{{Citation |year=2010 |author1=Chaudhari, Nirupa  |author2=Roper, Stephen D. |name-list-style=amp |title=The cell biology of taste |journal=[[Journal of Cell Biology]] |pmid=20696704 |volume=190 |issue=3 |pmc=2922655 |pages=285–296 |doi=10.1083/jcb.201003144 }}
* {{Wiktionary-inline|taste}}
* {{Commons category-inline|Taste}}
{{Taste}}
{{Sensation and perception}}

[[Category:Sensory systems]]
[[Category:Gustation| ]]
[[Category:Gustatory system]]

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Greek: Ά ά Έ έ Ή ή Ί ί Ό ό Ύ ύ Ώ ώ Α α Β β Γ γ Δ δ Ε ε Ζ ζ Η η Θ θ Ι ι Κ κ Λ λ Μ μ Ν ν Ξ ξ Ο ο Π π Ρ ρ Σ σ ς Τ τ Υ υ Φ φ Χ χ Ψ ψ Ω ω {{Polytonic|}}
Cyrillic: А а Б б В в Г г Ґ ґ Ѓ ѓ Д д Ђ ђ Е е Ё ё Є є Ж ж З з Ѕ ѕ И и І і Ї ї Й й Ј ј К к Ќ ќ Л л Љ љ М м Н н Њ њ О о П п Р р С с Т т Ћ ћ У у Ў ў Ф ф Х х Ц ц Ч ч Џ џ Ш ш Щ щ Ъ ъ Ы ы Ь ь Э э Ю ю Я я ́
IPA: t̪ d̪ ʈ ɖ ɟ ɡ ɢ ʡ ʔ ɸ β θ ð ʃ ʒ ɕ ʑ ʂ ʐ ç ʝ ɣ χ ʁ ħ ʕ ʜ ʢ ɦ ɱ ɳ ɲ ŋ ɴ ʋ ɹ ɻ ɰ ʙ ⱱ ʀ ɾ ɽ ɫ ɬ ɮ ɺ ɭ ʎ ʟ ɥ ʍ ɧ ʼ ɓ ɗ ʄ ɠ ʛ ʘ ǀ ǃ ǂ ǁ ɨ ʉ ɯ ɪ ʏ ʊ ø ɘ ɵ ɤ ə ɚ ɛ œ ɜ ɝ ɞ ʌ ɔ æ ɐ ɶ ɑ ɒ ʰ ʱ ʷ ʲ ˠ ˤ ⁿ ˡ ˈ ˌ ː ˑ ̪ {{IPA|}}

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taste: Sitelink, Title, Some statements, Description: en, Miscellaneous (e.g. aliases, entity existence)
Category:Gustation: Sitelink

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