Editing Umami

{{Short description|One of the five basic tastes}}
{{Distinguish|Unami (disambiguation){{!}}Unami}}
{{Use dmy dates|date=November 2017}}
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'''Umami''' ({{IPAc-en|uː|ˈ|m|ɑː|m|i}} from {{langx|ja|うま味}} {{IPA|ja|ɯmami}}), or '''savoriness''', is one of the five [[Taste#Basic tastes|basic tastes]].<ref name="Torii_2013" /> It is characteristic of [[broth]]s and cooked meats.<ref name="Fleming_2013" /><ref>{{cite news|url=https://www.telegraph.co.uk/foodanddrink/foodanddrinknews/7195114/Umami-in-a-tube-fifth-taste-goes-on-sale-in-supermarkets.html |archive-url=https://ghostarchive.org/archive/20220111/https://www.telegraph.co.uk/foodanddrink/foodanddrinknews/7195114/Umami-in-a-tube-fifth-taste-goes-on-sale-in-supermarkets.html |archive-date=11 January 2022 |url-access=subscription |url-status=live |date=9 February 2010 |title=Umami in a tube: 'fifth taste' goes on sale in supermarkets  |newspaper=[[The Daily Telegraph]] | author = Blake H |access-date=10 February 2011}}{{cbignore}}</ref><ref name = "Jufresa_2015" /><ref name = "Mouritsen_2014" />{{rp|35–36}}

People taste umami through [[taste receptor]]s that typically respond to [[Glutamic acid|glutamates]] and  [[nucleotide|nucleotides]], which are widely present in meat broths and fermented products. Glutamates are commonly added to some foods in the form of [[monosodium glutamate]] (MSG), and nucleotides are commonly added in the form of [[disodium guanylate]],  [[inosine monophosphate]] (IMP) or [[guanosine monophosphate]] (GMP).<ref name="Nelson_2002" /><ref name="Delay_2000" /><ref name = "Chaudhari_2000" /> Since umami has its own receptors rather than arising out of a combination of the traditionally recognized taste receptors, scientists now consider umami to be a distinct taste.<ref name="Torii_2013"/><ref name="npr" />

Foods that have a strong umami flavor include meats, [[shellfish]], fish (including [[fish sauce]] and preserved fish such as [[Maldives fish]], ''[[katsuobushi]]'', [[Sardines as food|sardines]], and [[Anchovies as food|anchovies]]), ''[[dashi]]'', [[tomato]]es, [[Edible mushroom|mushrooms]], [[hydrolyzed vegetable protein]], [[meat extract]], [[yeast extract]], [[kimchi]], [[cheese]]s, and [[soy sauce]].

In 1908, Dr. [[Kikunae Ikeda]] of the [[University of Tokyo]] scientifically identified umami as a distinct taste attributed to glutamic acid. As a result, in 1909, Ikeda and Saburōsuke Suzuki founded [[Ajinomoto|Ajinomoto Co., Inc.]] which introduced the world's first umami seasoning: monosodium glutamate (MSG), marketed in Japan under the name "Ajinomoto." MSG subsequently spread worldwide as a seasoning capable of enhancing umami in a wide variety of dishes.<ref name="cnn270125">{{cite web|url=https://edition.cnn.com/travel/article/msg-changing-views-cmd|archive-url=https://web.archive.org/web/20250127164039/https://edition.cnn.com/travel/article/msg-changing-views-cmd|title=MSG is the most misunderstood ingredient of the century. That’s finally changing|publisher=[[CNN]]|date=|archive-date=27 January 2025|access-date=16 May 2025}}</ref><ref name="jt150225">{{cite web|url=https://japantoday.com/category/features/food/what-is-umami-the-mysterious-taste-of-japanese-cooking|archive-url=https://web.archive.org/web/20250215074906/https://japantoday.com/category/features/food/what-is-umami-the-mysterious-taste-of-japanese-cooking|title=What is umami, the mysterious taste of Japanese cooking?|publisher=[[Japan Today]]|date=|archive-date=15 February 2025|access-date=16 May 2025}}</ref><ref name="ym160125">{{cite web|url=https://japannews.yomiuri.co.jp/features/japan-focus/20210413-71775/|archive-url=https://web.archive.org/web/20250116020815/https://japannews.yomiuri.co.jp/features/japan-focus/20210413-71775/|title=Kanagawa: Unmasking science, history behind umami
|publisher=[[Yomiuri Shimbun]]|date=|archive-date=16 January 2025|access-date=16 May 2025}}</ref><ref name="aji210425">{{cite web|url=https://www.ajinomoto.com/aboutus/history|archive-url=https://web.archive.org/web/20250421202224/https://www.ajinomoto.com/aboutus/history|title=History|publisher=[[Ajinomoto]]|date=|archive-date=21 April 2025|access-date=16 May 2025}}</ref>

In 2000, researchers at the [[University of Miami]] discovered the presence of umami receptors on the [[tongue]], and in 2006, Ajinomoto’s research laboratories found similar receptors in the [[stomach]].<ref name="aji210425"/>

==Etymology==
A [[loanword]] from Japanese {{nihongo||うま味}}, ''umami'' can be translated as "pleasant savory taste".<ref>{{cite web |url = http://nihongo.monash.edu/cgi-bin/wwwjdic |title = EDICT's entry for ''umami'' | author = Breen J |access-date = 5 December 2017}}</ref> This [[wikt:neologism|neologism]] was coined in 1908 by Japanese chemist [[Kikunae Ikeda]] from a [[nominalization]] of ''umai'' ({{nihongo2|うまい}}) "delicious". The compound {{nihongo2|旨味}} (with ''mi'' ({{nihongo2|味}}) "taste") is used for a more general sense of a food as delicious.<ref>{{cite web|url=http://www.jsrae.or.jp/annai/yougo/188.html|title=うま味 (umami)|publisher=Japan Society of Refrigerating and Air Conditioning Engineers|access-date=5 June 2017}}</ref><ref>{{cite web|url=http://umamibook.net/what-is-umami|title=What is umami?|publisher=[[Columbia University Press]]|access-date=5 June 2017}}</ref><ref>{{cite web |url=http://dictionary.cambridge.org/dictionary/british/umami |title=Umami |work=[[Cambridge Dictionary]]|publisher=[[Cambridge University Press]] |access-date=1 January 2011}}</ref> There is no English equivalent to ''umami''; however, some close descriptions are "meaty", "savory", and "broth-like".<ref name="Yamaguchi_2000"/>

==Background==
Scientists have debated whether umami was a basic [[taste]] since Kikunae Ikeda first proposed its existence in 1908.<ref>{{cite journal | vauthors = Lindemann B, Ogiwara Y, Ninomiya Y | title = The discovery of umami | journal = Chemical Senses | volume = 27 | issue = 9 | pages = 843–4 | date = November 2002 | pmid = 12438211 | doi = 10.1093/chemse/27.9.843 | doi-access = free}}</ref><ref name="Kean_2015">{{cite journal|last=Kean|first=Sam|title=The science of satisfaction |journal=[[Distillations Magazine]]|publisher=[[Science History Institute]]|date=Fall 2015 |volume=1|issue=3|pages=5 |url=https://www.sciencehistory.org/distillations/magazine/the-science-of-satisfaction|access-date=22 March 2018}}</ref> In 1985, the term ''umami'' was recognized as the scientific term to describe the taste of [[glutamate]]s and [[nucleotide]]s at the first Umami International Symposium in Hawaii.<ref name="Kawamura_2016">{{cite book |title=Umami: A basic taste  | veditors = Kawamura Y, Kare MR |publisher=Marcel Dekker |location=New York| year=1987}}{{page needed|date=August 2016}}</ref> Umami represents the taste of the [[amino acid]] [[glutamic acid|L-glutamate]] and 5'-[[ribonucleotides]] such as [[guanosine monophosphate]] (GMP) and [[inosine monophosphate]] (IMP).<ref name="Yamaguchi_2000">{{cite journal | vauthors = Yamaguchi S, Ninomiya K | title = Umami and food palatability | journal = The Journal of Nutrition | volume = 130 | issue = 4S Suppl | pages = 921S–6S | date = April 2000 | pmid = 10736353 | doi = 10.1093/jn/130.4.921S | doi-access = free}}</ref> It can be described as a pleasant "[[broth]]y" or "[[meat]]y" taste with a long-lasting, mouthwatering and coating sensation over the tongue.   Umami enhances the palatability of a wide variety of foods.<ref name="Beauchamp_2009">{{cite journal | author = Beauchamp GK | title = Sensory and receptor responses to umami: an overview of pioneering work | journal = The American Journal of Clinical Nutrition | volume = 90 | issue = 3 | pages = 723S–727S | date = September 2009 | pmid = 19571221 | doi = 10.3945/ajcn.2009.27462E | doi-access = free}}</ref> 

Specialized taste bud cells detect the chemical species perceived as umami by humans.<ref name="Symposium"/> Glutamate in acid form (glutamic acid) imparts little umami taste, whereas the [[Salt (chemistry)|salts]] of [[glutamic acid]], known as [[glutamate]]s, give the characteristic umami taste due to their ionized state. GMP and IMP amplify the taste intensity of glutamate.<ref name="Yasuo_2008">{{cite journal | vauthors = Yasuo T, Kusuhara Y, Yasumatsu K, Ninomiya Y | title = Multiple receptor systems for glutamate detection in the taste organ | journal = Biological & Pharmaceutical Bulletin | volume = 31 | issue = 10 | pages = 1833–7 | date = October 2008 | pmid = 18827337 | doi = 10.1248/bpb.31.1833 | doi-access = free}}</ref> Adding salt to the free acids also enhances the umami taste.<ref name="Lioe_2010">{{cite journal | vauthors = Lioe HN, Selamat J, Yasuda M | title = Soy sauce and its umami taste: a link from the past to current situation | journal = Journal of Food Science | volume = 75 | issue = 3 | pages = R71-6 | date = April 2010 | pmid = 20492309 | doi = 10.1111/j.1750-3841.2010.01529.x | doi-access = free | url = http://psasir.upm.edu.my/id/eprint/16791/1/Soy%20sauce%20and%20its%20umami%20taste.pdf}}</ref> It is disputed whether umami is truly an independent taste because standalone glutamate without [[table salt]] ions(Na+) is perceived as sour; sweet and umami tastes share a taste receptor subunit, with salty taste blockers reducing discrimination between monosodium glutamate and sucrose; and some people cannot distinguish umami from a salty taste.<ref name="hartley">{{Cite journal |last1=Hartley |first1=Isabella E |last2=Liem |first2=Djin Gie |last3=Keast |first3=Russell |date=2019-01-16 |title=Umami as an 'Alimentary' Taste. A New Perspective on Taste Classification |journal=Nutrients |volume=11 |issue=1 |pages=182 |doi=10.3390/nu11010182 |doi-access=free |issn=2072-6643 |pmc=6356469 |pmid=30654496}}</ref>

Monosodium [[L-aspartate]] has an umami taste about a quarter as intense as MSG, whereas [[ibotenic acid]] and [[tricholomic acid]] (likely as their salts or with salt) are claimed to be many times more intense.<ref name="Lioe_2010"/> Peptides can also generate a umami taste, with 52 of them being known to do so as of 2017 (20 of them are questionable).<ref name="zhang">{{cite journal | vauthors = Zhang Y, Venkitasamy C, Pan Z, Liu W, Zhao L | title = Novel Umami Ingredients: Umami Peptides and Their Taste | journal = Journal of Food Science | volume = 82 | issue = 1 | pages = 16–23 | date = January 2017 | pmid = 27926796 | doi = 10.1111/1750-3841.13576 | doi-access = free}}</ref>

==Discovery==
[[File:Kikunae Ikeda.jpg|thumb|[[Kikunae Ikeda]]]]
[[Glutamate]] has a long history in cooking.<ref name = "Lehrer_2007">{{cite book | author = Lehrer J |author-link = Jonah Lehrer |title = Proust was a Neuroscientist |publisher = [[Mariner Books]] |year =2007|isbn = 978-0-547-08590-6 |ref=Lehrer07|title-link = Proust was a Neuroscientist}}</ref> Fermented fish sauces ([[garum]]), which are rich in glutamate, were used widely in ancient Rome,<ref>{{cite journal |vauthors=Smriga M, Mizukoshi T, Iwata D, Sachise E, Miyano H, Kimura T, Curtis R |title=Amino acids and minerals in ancient remnants of fish sauce (garum) sampled in the "Garum Shop" of Pompeii, Italy |journal=[[Journal of Food Composition and Analysis]] |volume=23 |issue=5 |pages=442–46 |date=August 2010 |doi=10.1016/j.jfca.2010.03.005}}</ref> fermented barley sauces ([[murri (condiment)|murri]]) rich in glutamate were used in medieval [[Byzantine cuisine|Byzantine]] and [[Arab cuisine|Arab]] cuisine,<ref>{{Citation|url=https://www.latimes.com/archives/la-xpm-1998-apr-01-fo-34730-story.html|title=Rot of Ages| author = Perry C | date = 1 April 1998 |newspaper=[[Los Angeles Times]]|access-date=14 August 2024}}</ref> and fermented [[fish sauce]]s and [[soy sauce]]s have histories going back to the third century in China. Cheese varieties are rich in glutamate and umami flavor.<ref>{{Cite journal |last1=Drake |first1=S. L. |last2=Carunchia Whetstine |first2=M. E. |last3=Drake |first3=M. A. |last4=Courtney |first4=P. |last5=Fligner |first5=K. |last6=Jenkins |first6=J. |last7=Pruitt |first7=C. |date=2007 |title=Sources of umami taste in Cheddar and Swiss cheeses |url=https://pubmed.ncbi.nlm.nih.gov/17995691/ |journal=Journal of Food Science |volume=72 |issue=6 |pages=S360–366 |doi=10.1111/j.1750-3841.2007.00402.x |issn=1750-3841 |pmid=17995691}}</ref><ref>{{Cite web |title=Cheese |url=https://www.umamiinfo.com/richfood/foodstuff/cheese.html#:~:text=Mature%20Cheddar%20which%20has%20been,complex%20and%20satisfying%20lingering%20aftertaste. |access-date=2024-09-25 |website=Umami Information Center |language=en}}</ref> In the late 1800s, chef [[Auguste Escoffier]], who opened restaurants in Paris and London, created meals that combined umami with [[Saltiness|salty]], [[sour]], [[Sweetness|sweet]], and [[bitter (taste)|bitter]] tastes.<ref name=npr/> However, he did not know the chemical source of this unique quality. 

Umami was first scientifically identified in 1908 by [[Kikunae Ikeda]],<ref name="Ikeda_2002">{{cite journal | author = Ikeda K | title = New Seasonings | journal = Chemical Senses | volume = 27 | issue = 9 | pages = 847–9 | date = November 2002 | pmid = 12438213 | doi = 10.1093/chemse/27.9.847 | doi-access = free}} (partial translation of {{cite journal | author = Ikeda K | title=New Seasonings | language = Japanese  | journal = Journal of the Chemical Society of Tokyo | year= 1909 | volume=30 | pages= 820–36}})</ref><ref>{{cite journal | author = Nakamura E | title = One hundred years since the discovery of the "Kawamura_2016" taste from seaweed broth by Kikunae Ikeda, who transcended his time | journal = Chemistry: An Asian Journal | volume = 6 | issue = 7 | pages = 1659–63 | date = July 2011 | pmid = 21472994 | doi = 10.1002/asia.201000899}}</ref> a professor of the [[Tokyo Imperial University]]. He found that [[glutamic acid|glutamate]] was responsible for the palatability of the broth from ''[[kombu]]'' seaweed. He noticed that the taste of ''kombu'' ''[[dashi]]'' was distinct from sweet, sour, bitter, and salty and named it ''umami''.<ref name="Kean_2015"/>

Professor Shintaro Kodama, a disciple of Ikeda, discovered in 1913 that [[Katsuobushi|dried bonito flakes]] (a type of tuna) contained another umami substance.<ref>{{cite journal | author = Kodama S |journal=Journal of the Chemical Society of Japan|title=On a procedure for separating inosinic acid|volume=34|page=751|year=1913}}</ref> This was the ribonucleotide [[inosine monophosphate|IMP]]. In 1957, Akira Kuninaka realized that the ribonucleotide GMP present in [[shiitake]] mushrooms also conferred the umami taste.<ref>{{cite journal | author = Kuninaka A |journal= Journal of the Agricultural Chemical Society of Japan |title=Studies on taste of ribonucleic acid derivatives|volume=34|issue=6|pages=487–92|year=1960|doi=10.1271/nogeikagaku1924.34.6_489|doi-access=free|language=JA}}</ref> One of Kuninaka's most important discoveries was the [[synergistic]] effect between ribonucleotides and glutamate. When foods rich in glutamate are combined with ingredients that have ribonucleotides, the resulting taste intensity is higher than would be expected from merely adding the intensity of the individual ingredients.<ref name="Yamaguchi_2000"></ref>

This synergy of umami may help explain various classical food pairings: the Japanese make ''dashi'' with ''[[kombu]]'' seaweed and dried bonito flakes; the Chinese add [[Chinese leek]] and [[Chinese cabbage]] to chicken soup, as do Scots in the similar Scottish dish of [[cock-a-leekie soup]]; and Italians grate the [[Parmigiano-Reggiano]] cheese on a variety of different dishes.

==Properties==
Umami has a mild but lasting aftertaste associated with salivation and a sensation of furriness on the tongue, stimulating the throat, the roof and the back of the mouth.<ref name="Yamaguchi_1998">{{cite journal | author = Yamaguchi, Shizuko |title=Basic properties of umami and its effects on food flavor |journal=Food Reviews International |volume=14 |issue=2&3 |pages=139–76 |year=1998 |doi= 10.1080/87559129809541156}}</ref><ref name="Uneyama_2009">{{cite journal | vauthors = Uneyama H, Kawai M, Sekine-Hayakawa Y, Torii K | title = Contribution of umami taste substances in human salivation during meal | journal = The Journal of Medical Investigation | volume = 56 Suppl | issue = supplement | pages = 197–204 | date = August 2009 | pmid = 20224181 | doi = 10.2152/jmi.56.197 | doi-access = free}}</ref> By itself, umami is not [[palatable]], but it makes a great variety of foods pleasant, especially in the presence of a matching aroma.<ref name="Rolls_2009">{{cite journal | author = Rolls ET | title = Functional neuroimaging of umami taste: what makes umami pleasant? | journal = The American Journal of Clinical Nutrition | volume = 90 | issue = 3 | pages = 804S–813S | date = September 2009 | pmid = 19571217 | doi = 10.3945/ajcn.2009.27462R | doi-access = free}}</ref> Like other basic tastes, umami is pleasant only within a relatively narrow concentration range.<ref name="Yamaguchi_1998" />

The optimum umami taste depends also on the amount of salt, and at the same time, low-salt foods can maintain a satisfactory taste with the appropriate amount of umami.<ref name="Yamaguchi_1984">{{cite journal | vauthors = Yamaguchi S, Takahashi C |title=Interactions of monosodium glutamate and sodium chloride on saltiness and palatability of a clear soup|journal=[[Journal of Food Science]] |volume=49 |pages=82–85 |year=1984 |doi= 10.1111/j.1365-2621.1984.tb13675.x}}</ref> One study showed that ratings of pleasantness, taste intensity, and ideal saltiness of low-salt soups were greater when the soup contained umami, whereas low-salt soups without umami were less pleasant.<ref name="Roininen_1996">{{cite journal | vauthors = Roininen K, Lähteenmäki L, Tuorila H | title = Effect of umami taste on pleasantness of low-salt soups during repeated testing | journal = Physiology & Behavior | volume = 60 | issue = 3 | pages = 953–8 | date = September 1996 | pmid = 8873274 | doi = 10.1016/0031-9384(96)00098-4 | s2cid = 39325526}}</ref> Another study demonstrated that using [[fish sauce]] as a source of umami could reduce the need for salt by 10–25% to flavor such foods as [[chicken broth]], [[tomato sauce]], or coconut [[curry]] while maintaining overall taste intensity.<ref name="Huynh_2016">{{cite journal | vauthors = Huynh HL, Danhi R, Yan SW | title = Using Fish Sauce as a Substitute for Sodium Chloride in Culinary Sauces and Effects on Sensory Properties | journal = Journal of Food Science | volume = 81 | issue = 1 | pages = S150-5 | date = January 2016 | pmid = 26613570 | doi = 10.1111/1750-3841.13171}}</ref><ref>{{cite web|url=https://www.sciencedaily.com/releases/2016/02/160202143938.htm|title=A new alternative to sodium: Fish sauce|work=[[ScienceDaily]]|date=2 February 2016|access-date=20 February 2017}}</ref>

Some population groups, such as the elderly, may benefit from umami taste because their taste and smell sensitivity may be impaired by age and medication. The loss of taste and smell can contribute to poor nutrition, increasing their risk of disease.<ref name="Yamamoto_2009">{{cite journal | vauthors = Yamamoto S, Tomoe M, Toyama K, Kawai M, Uneyama H | title = Can dietary supplementation of monosodium glutamate improve the health of the elderly? | journal = The American Journal of Clinical Nutrition | volume = 90 | issue = 3 | pages = 844S–849S | date = September 2009 | pmid = 19571225 | doi = 10.3945/ajcn.2009.27462X | doi-access = free}}</ref> Some evidence exists to show umami not only stimulates appetite, but also may contribute to [[satiety]].<ref name="masic">{{cite journal | vauthors = Masic U, Yeomans MR | title = Umami flavor enhances appetite but also increases satiety | journal = The American Journal of Clinical Nutrition | volume = 100 | issue = 2 | pages = 532–8 | date = August 2014 | pmid = 24944058 | doi = 10.3945/ajcn.113.080929 | doi-access = free}}</ref>

== Foods rich in umami components ==
[[File:Dried anchovies.jpg|thumb|[[Anchovies]] are rich in umami.]]

Many foods are rich in the amino acids and nucleotides imparting umami. Naturally occurring glutamate can be found in meats and vegetables. [[Inosine]] ([[Inosine monophosphate|IMP]]) comes primarily from meats and [[guanosine]] ([[Guanosine monophosphate|GMP]]) from vegetables. Mushrooms, especially dried [[shiitake]], are rich sources of umami flavor from guanylate. Smoked or fermented fish are high in inosinate, and [[shellfish]] in [[Adenosine monophosphate|adenylate]].<ref name = "Mouritsen_2014" />{{rp|11, 52, 110}}<ref>{{cite web|url=http://www.popsci.com/hidden-secrets-umami|title=Put the science of umami to work for you | author = Adams P |website=[[Popular Science]] |date=24 November 2015|access-date=11 December 2015}}</ref> Protein in food is tasteless, however processes such as fermentation, curing, or heat treatment release glutamate and other amino acids.<ref name=hartley/> 

Generally, umami taste is common to foods that contain high levels of [[glutamic acid|L-glutamate]], [[inosine monophosphate|IMP]] and [[guanosine monophosphate|GMP]], most notably in [[fish]], [[shellfish]], [[Curing (food preservation)|cured meats]], [[meat extract]]s, [[Edible mushroom|mushrooms]], [[vegetable]]s (e.g., ripe [[tomato]]es, [[Chinese cabbage]], [[spinach]], [[celery]], etc.), [[green tea]], [[hydrolyzed vegetable protein]], and fermented and aged products involving bacterial or yeast cultures, such as [[cheese]]s, [[shrimp paste]]s, [[fish sauce]], [[soy sauce]], [[natto]], [[nutritional yeast]], and [[yeast extract]]s such as [[Vegemite]] and [[Marmite]].<ref name="Fleming_2013"/><ref>{{cite journal | vauthors = Hajeb P, Jinap S | title = Umami taste components and their sources in Asian foods | journal = Critical Reviews in Food Science and Nutrition | volume = 55 | issue = 6 | pages = 778–91 | year = 2015 | pmid = 24915349 | doi = 10.1080/10408398.2012.678422 | s2cid = 205690996 | url = http://psasir.upm.edu.my/id/eprint/46789/1/Umami%20taste%20components%20and%20their%20sources%20in%20Asian%20foods.pdf}}</ref>

Studies have shown that the amino acids in [[breast milk]] are often the first encounter humans have with umami. Glutamic acid makes up half of the free amino acids in breast milk.<ref name="Agostoni_2000">{{cite journal | vauthors = Agostoni C, Carratù B, Boniglia C, Riva E, Sanzini E | title = Free amino acid content in standard infant formulas: comparison with human milk | journal = Journal of the American College of Nutrition | volume = 19 | issue = 4 | pages = 434–8 | date = August 2000 | pmid = 10963461 | doi = 10.1080/07315724.2000.10718943 | s2cid = 3141583}}</ref><ref name="Fleming_2013" /><ref name = "Mouritsen_2014" />

== Perceptual independence from salty and sweet taste ==
Since all umami taste compounds are sodium salts, the perceptual differentiation of salty and umami tastes has been difficult in taste tests and studies have found as much as 27% of certain populations may be umami "hypotasters".<ref name=hartley/>

Furthermore single glutamate (glutamic acid) with no table salt ions (Na+) elicits sour taste and in psychophysical tests, sodium or potassium salt cations seem to be required to produce a perceptible umami taste.<ref name=hartley />

Sweet and umami tastes both utilize the taste receptor subunit [[TAS1R3]], with salt taste blockers reducing discrimination between monosodium glutamate and sucrose in rodents.<ref name=hartley />

If umami doesn't have perceptual independence, it could be classified with other tastes like fat, carbohydrate, metallic, and calcium, which can be perceived at high concentrations but may not offer a prominent taste experience.<ref name=hartley/>

==Taste receptors==
{{See also|Taste receptor#Savory or glutamates (Umami)}}
Most [[taste bud]]s on the tongue and other regions of the mouth can detect umami taste, irrespective of their location. (The [[tongue map]] in which different tastes are distributed in different regions of the tongue is a common misconception.) [[Biochemical]] studies have identified the [[taste receptor]]s responsible for the sense of umami as modified forms of [[Metabotropic glutamate receptor 4|mGluR4]], [[Metabotropic glutamate receptor 1|mGluR1]], and [[taste receptor]] type 1 ([[TAS1R1]] + [[TAS1R3]]), all of which have been found in all regions of the tongue bearing taste buds.<ref name = "Chaudhari_2000">{{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}}
*{{cite press release |date=February 2000 |title=Umami taste receptor identified |website=Nature Neuroscience |url=http://www.nature.com/neuro/press_release/nn0200.html |archive-url=https://web.archive.org/web/20130305013817/http://www.nature.com/neuro/press_release/nn0200.html |archive-date=2013-03-05}}</ref><ref name="Nelson_2002"/><ref>{{cite journal | vauthors = San Gabriel A, Uneyama H, Yoshie S, Torii K | title = Cloning and characterization of a novel mGluR1 variant from vallate papillae that functions as a receptor for L-glutamate stimuli | journal = Chemical Senses | volume = 30 Suppl 1 | issue = Suppl | pages = i25-6 | date = January 2005 | pmid = 15738140 | doi = 10.1093/chemse/bjh095 | doi-access = free}}</ref> These receptors are also found in some regions of the [[duodenum]].<ref name="Sasano_2015">{{Cite journal| vauthors = Sasano T, Satoh-Kuriwada S, Shoji N |date=2015-01-26|title=The important role of umami taste in oral and overall health |journal= Flavour|volume=4|issue=1|pages=10|doi=10.1186/2044-7248-4-10|s2cid=14562283|issn=2044-7248|doi-access=free}}</ref> A 2009 review corroborated the acceptance of these receptors, stating, "Recent molecular biological studies have now identified strong candidates for umami receptors, including the heterodimer TAS1R1/TAS1R3, and truncated type 1 and 4 metabotropic glutamate receptors missing most of the N-terminal extracellular domain (taste-mGluR4 and truncated-mGluR1) and brain-mGluR4."<ref name="Symposium">{{cite book |editor-last1=Finger |editor-first1=Thomas E.  |year=2009 |title=International Symposium on Olfaction and Taste |language=English |volume=1170 |edition=1st |location=Hoboken, NJ |publisher=Wiley-Blackwell |isbn=978-1573317382}}</ref>

Receptors mGluR1 and mGluR4 are specific to glutamate whereas TAS1R1 + TAS1R3 are responsible for the synergism already described by Akira Kuninaka in 1957. However, the specific role of each type of receptor in taste bud cells remains unclear.<ref name=Diepeveen/> All three receptors work together to produce the taste sensation.<ref>{{cite journal |last1=Yasumatsu |first1=K |last2=Manabe |first2=T |last3=Yoshida |first3=R |last4=Iwatsuki |first4=K |last5=Uneyama |first5=H |last6=Takahashi |first6=I |last7=Ninomiya |first7=Y |title=Involvement of multiple taste receptors in umami taste: analysis of gustatory nerve responses in metabotropic glutamate receptor 4 knockout mice. |journal=The Journal of Physiology |date=15 February 2015 |volume=593 |issue=4 |pages=1021–34 |doi=10.1113/jphysiol.2014.284703 |pmid=25529865|pmc=4398535 }}</ref>

=== Downstream signaling ===
==== TAS1R1 + TAS1R3 ====
The TAS1R1 + TAS1R3 receptor is a [[G protein-couple receptor]], much like the sweet and bitter receptors. It uses the same downstream signaling molecules, including [[G beta-gamma complex|G proteins beta-gamma]], [[PLCB2]] and [[Inositol triphosphate|IP<sub>3</sub>]], to ultimately cause a release of [[calcium]] (Ca<sup>2+</sup>) from intracellular stores.<ref name=Kinnamon>{{cite journal | author = Kinnamon SC | title = Taste receptor signalling – from tongues to lungs | journal = Acta Physiologica | volume = 204 | issue = 2 | pages = 158–68 | date = February 2012 | pmid = 21481196 | pmc = 3704337 | doi = 10.1111/j.1748-1716.2011.02308.x}}</ref> Calcium activates a so-called transient-receptor-potential cation channel [[TRPM5]] that leads to membrane [[depolarization]] and the consequent release of [[Adenosine triphosphate|ATP]] across a channel of [[CALHM1]] and [[CALHM3]].<ref name=Diepeveen>{{cite journal |last1=Diepeveen |first1=J |last2=Moerdijk-Poortvliet |first2=TCW |last3=van der Leij |first3=FR |title=Molecular insights into human taste perception and umami tastants: A review. |journal=Journal of Food Science |date=April 2022 |volume=87 |issue=4 |pages=1449–1465 |doi=10.1111/1750-3841.16101 |pmid=35301715|pmc=9314127 }}</ref>

The ATP released by the "Type II" cell is detected by P2X receptors on nearby afferent gustatory nerve fibers and P2Y receptors on adjacent taste cells. P2X appears to be indispensable for the transduction of umami, so this is probably the main route for umami signals.<ref name=Kinnamon/> "Type III" cells, which directly connect to the nerve synapses, also respond to the released ATP by releasing neurotransmitters. One of these neurotransmitters, [[serotonin]], regulates the release of ATP by the type II cells.<ref name=Kinnamon/>

==== mGluR1 and mGluR4 ====
Signal from these two receptors are conveyed in a manner independent of TRPM5. Single umami-sensitive fibres in mice are mostly either "sucrose-best" or "glutamate-best". Within each type there are two subtypes: one shows synergistic activation between monopotassium glutamate and inosine monophosphate, the other does not. The TAS1R1/3 + TRPM5 route uses the "sucrose-best" fibers with synergy, while mGluR1 and mGluR4 use both of the "glutamate-best" subtypes.<ref>{{cite journal |last1=Yasumatsu |first1=Keiko |last2=Ogiwara |first2=Yoko |last3=Takai |first3=Shingo |last4=Yoshida |first4=Ryusuke |last5=Iwatsuki |first5=Ken |last6=Torii |first6=Kunio |last7=Margolskee |first7=Robert F. |last8=Ninomiya |first8=Yuzo |title=Umami taste in mice uses multiple receptors and transduction pathways |journal=The Journal of Physiology |date=March 2012 |volume=590 |issue=5 |pages=1155–1170 |doi=10.1113/jphysiol.2011.211920 |pmid=22183726|pmc=3381822 }}</ref>

=== Beyond the tongue ===
The gut has its own umami taste receptors. The ATP taste signals are conveyed to the brain, probably through afferent branches of the [[vagus nerve]] or the afferent sensory nerves in the mouth. The brain uses this information to regulate behaviors and preferences in mice. This is an example of the [[gut-brain axis]].<ref name="Torii_2013"/>

=== In other animals ===
Cats have mutations in Tas1r1 and Tas1r3 that cause their receptor to not perceive glutamate and aspartate as umami. However, their rececptor responds to nucleotide, and some L-amino acids enhance the response to nucleotides. Cats probably perceive tuna as very umami due to it being rich in [[Inosinic acid|inosine monophosphate]] and L-histine.<ref name=McGrane>{{cite journal |last1=McGrane |first1=Scott J. |last2=Gibbs |first2=Matthew |last3=Hernangomez de Alvaro |first3=Carlos |last4=Dunlop |first4=Nicola |last5=Winnig |first5=Marcel |last6=Klebansky |first6=Boris |last7=Waller |first7=Daniel |date=August 8, 2023 |title=Umami taste perception and preferences of the domestic cat (Felis catus), an obligate carnivore |url=https://doi.org/10.1093/chemse/bjad026 |journal=Chemical Senses |volume=48 |doi=10.1093/chemse/bjad026 |pmid=37551788 |pmc=10468298 |access-date=September 21, 2023}}</ref>

The Tas1r1-Tas1r3 receptor of mice is activated by a wide range of free L-amino acids, but not acidic ones such as glutamate.<ref name=McGrane/>

The lineage of aquatic mammals including dolphins and sea lions have no functional Tas1r1, and neither do giant pandas. They cannot generate a functional Tas1r1-Tas1r3 receptor as a result.<ref name=":1">{{cite journal | vauthors = Jiang P, Josue J, Li X, Glaser D, Li W, Brand JG, Margolskee RF, Reed DR, Beauchamp GK |display-authors=3| title = Major taste loss in carnivorous mammals | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 109 | issue = 13 | pages = 4956–61 | date = March 2012 | pmid = 22411809 | pmc = 3324019 | doi = 10.1073/pnas.1118360109 | doi-access = free }}</ref>

== Consumers and safety ==
Umami has become popular as a flavor with food manufacturers trying to improve the taste of low sodium offerings.<ref>{{cite web|url=http://www.foodnavigator.com/Market-Trends/Does-MSG-have-a-future-in-Europe-as-umami-gains-flavour-favour|website=FoodNavigator |date=7 October 2015| author = Michail N |title=Does MSG have a future in Europe as umami gains flavour favour?|access-date=19 February 2017}}</ref> Chefs create "umami bombs", which are dishes made of several umami ingredients like [[fish sauce]].<ref name="Fleming_2013"/><ref name=npr/> Umami may account for the long-term formulation and popularity of [[ketchup]].<ref>{{cite magazine|url=http://www.newyorker.com/archive/2004/09/06/040906fa_fact_gladwell?currentPage=all |title=Taste technologies: The Ketchup Conundrum | author = Gladwell M |date=6 September 2004 |magazine=[[The New Yorker]] |access-date=8 March 2014}}</ref> The United States [[Food and Drug Administration]] has designated the umami enhancer [[monosodium glutamate]] (MSG) as a safe ingredient. While some people identify themselves as [[Glutamate sensitivity|sensitive to MSG]], a study commissioned by the FDA was only able to identify transient, mild [[symptom]]s in a few of the subjects, and only when the MSG was consumed in unrealistically large quantities.<ref>{{cite web|url=https://www.fda.gov/Food/IngredientsPackagingLabeling/FoodAdditivesIngredients/ucm328728.htm|title=Questions and Answers on Monosodium glutamate (MSG)|date=19 November 2012|website=[[United States Food and Drug Administration]] |access-date=19 February 2017}}</ref> There is also no apparent difference in sensitivity to umami when comparing Japanese and Americans.<ref>{{cite journal | author = Yamaguchi S | title = Basic properties of umami and effects on humans | journal = Physiology & Behavior | volume = 49 | issue = 5 | pages = 833–41 | date = May 1991 | pmid = 1679557 | doi = 10.1016/0031-9384(91)90192-Q | series = Umami: Proceedings of the Second International Symposium on Umami | s2cid = 20980527}}</ref>

== Background of other taste categories ==
The five basic tastes (saltiness, sweetness, bitterness, sourness, and savoriness) are detected by specialized taste receptors on the tongue and [[palate]] [[epithelium]].<ref>{{cite journal | vauthors = Chen X, Gabitto M, Peng Y, Ryba NJ, Zuker CS | title = A gustotopic map of taste qualities in the mammalian brain | journal = Science | volume = 333 | issue = 6047 | pages = 1262–6 | date = September 2011 | pmid = 21885776 | pmc = 3523322 | doi = 10.1126/science.1204076 | bibcode = 2011Sci...333.1262C | jstor = 23060168}}</ref> The number of taste categories in humans remains under research, with a sixth taste possibly including spicy or pungent.<ref name="Wertz_2013">{{Cite journal| author = Wertz SK |date=2013|title=The Elements of Taste: How Many Are There? |journal= The Journal of Aesthetic Education |volume=47|issue=1|pages=46–57|doi=10.5406/jaesteduc.47.1.0046|jstor=10.5406/jaesteduc.47.1.0046|s2cid=191458091|issn=0021-8510}}</ref>

== See also ==

{{Portal|Food}}
{{Columns-list|colwidth=16em|
* [[Adenosine monophosphate]]
* [[Ajinomoto]]
* [[Glutamate flavoring]]
* [[Disodium glutamate]]
* [[Disodium inosinate]]
* [[Inosinic acid]]
* [[Monopotassium glutamate]]
}}

==References==
{{Reflist|refs=
<ref name="Torii_2013">{{cite journal | vauthors = Torii K, Uneyama H, Nakamura E | title = Physiological roles of dietary glutamate signaling via gut-brain axis due to efficient digestion and absorption | journal = Journal of Gastroenterology | volume = 48 | issue = 4 | pages = 442–51 | date = April 2013 | pmid = 23463402 | pmc = 3698427 | doi = 10.1007/s00535-013-0778-1}}</ref>
<ref name="Fleming_2013">{{cite web |url= https://www.theguardian.com/lifeandstyle/wordofmouth/2013/apr/09/umami-fifth-taste |title=Umami: why the fifth taste is so important| author = Fleming A |newspaper=[[The Guardian]] |date=9 April 2013|access-date=18 February 2017}}</ref>
<ref name = "Jufresa_2015">{{Cite book | author = Jufresa L |title=Umami (Mapa de las lenguas) |location=Spain |publisher=[[Penguin Random House]] Grupo Editorial México |language=es |isbn=978-607-31-2817-9 |date=February 16, 2015 |type=ebook}}</ref>
<ref name = "Mouritsen_2014">{{cite book|date=April 2014 |type=ebook | vauthors = Mouritsen JD, Drotner J, Styrbæk K, Mouritsen OG |title=Umami: Unlocking the Secrets of the Fifth Taste |location=United States |language=en |publisher=[[Columbia University Press]] |pages=35–36 |doi=10.7312/mour16890 |jstor=10.7312/mour16890 |isbn=978-0-231-53758-2}}</ref>
<ref name="Nelson_2002">{{cite journal | vauthors = Nelson G, Chandrashekar J, Hoon MA, Feng L, Zhao G, Ryba NJ, Zuker CS | title = An amino-acid taste receptor | journal = Nature | volume = 416 | issue = 6877 | pages = 199–202 | date = March 2002 | pmid = 11894099 | doi = 10.1038/nature726 | s2cid = 1730089 | bibcode = 2002Natur.416..199N}}</ref>
<ref name="Delay_2000">{{cite journal | vauthors = Delay ER, Beaver AJ, Wagner KA, Stapleton JR, Harbaugh JO, Catron KD, Roper SD | title = Taste preference synergy between glutamate receptor agonists and inosine monophosphate in rats | journal = Chemical Senses | volume = 25 | issue = 5 | pages = 507–15 | date = October 2000 | pmid = 11015322 | doi = 10.1093/chemse/25.5.507 | doi-access = free}}</ref>
<ref name="npr">{{cite web|url=https://www.npr.org/templates/story/story.php?storyId=15819485|title= Sweet, Sour, Salty, Bitter ... and Umami|publisher=[[National Public Radio]] (NPR) |date=5 November 2007| author = Krulwich R}}</ref>
}}

== Further reading ==
{{Refbegin}}
* {{cite book | vauthors = Barbot P, Matsuhisa N, Mikuni K, Heston B | title = Dashi and Umami: The Heart of Japanese Cuisine | location = London | publisher = Eat-Japan / Cross Media | year = 2009 |isbn=978-1-897-70193-5}}
* {{cite book | title = Flavor Chemistry: Thirty Years of Progress | veditors = Teranishi R, Wick EL, Hornstein I | chapter = Umami and Food Palatability | vauthors = Yamaguchi S, Ninomiya K | isbn = 978-0-306-46199-6 | year = 1999 |location=New York |publisher=Kluwer Academic/[[Plenum Publishers]]}}
{{Refend}}

== External links ==
* [http://www.umamiinfo.com/ Umami Information Center, Tokyo, 2016]

{{Taste}}
{{Japanese food and drink}}
{{Authority control}}

[[Category:Gustation]]
[[Category:Japanese words and phrases]]
[[Category:Culinary terminology]]
[[Category:Umami enhancers| ]]
[[Category:1908 introductions]]
[[Category:1908 establishments in Japan]]
[[Category:1900s neologisms]]
[[Category:20th-century neologisms]]
[[Category:1985 neologisms]]