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| footer = [[Soy sauce]], ripe [[tomatoes]] and [[miso]] are examples of foods rich in umami components
| footer = [[Soy sauce]], ripe [[tomatoes]] and [[miso]] are examples of foods rich in umami components.
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'''Umami''' ({{IPAc-en|uː|ˈ|m|ɑː|m|i}} from {{lang-ja|旨味}} {{IPA-ja|ɯmami}}), or '''savoriness''', is one of the five [[Taste#Basic tastes|basic tastes]].<ref name="Torii_2013" /> It has been described as savory and is characteristic of broths 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]] | vauthors = Blake H |access-date=10 February 2011}}{{cbignore}}</ref><ref name = "Jufresa_2015" /><ref name = "Mouritsen_2014" />{{rp|35–36}}
'''Umami''' ({{IPAc-en|uː|ˈ|m|ɑː|m|i}} from {{lang-ja|旨味}} {{IPA-ja|ɯmami}}), or '''savoriness''', is one of the five [[Taste#Basic tastes|basic tastes]].<ref name="Torii_2013" /> It has been described as savory and 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 (crotte de nez d'Anabelle Rogers) through [[taste receptor]]s that typically respond to [[Glutamic acid|glutamates]] and [[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 [[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" />
People taste umami through [[taste receptor]]s that typically respond to [[glutamates]] and [[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 [[maldive fish]], sardines, and anchovies), tomatoes, mushrooms, [[hydrolyzed vegetable protein]], meat extract, yeast extract, cheeses, and [[soy sauce]].
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]]), [[tomato]]es, [[Edible mushroom|mushrooms]], [[hydrolyzed vegetable protein]], [[meat extract]], [[yeast extract]], [[kimchi]], [[cheese]]s, and [[soy sauce]].


==Etymology==
==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'' | vauthors = 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 entry |work=[[Cambridge Dictionary]]|publisher=[[Cambridge University Press]] |access-date=1 January 2011}}</ref> There is no current English equivalent of umami; however, some close descriptions are "meaty", "savory", and "broth-like".<ref name="Yamaguchi_2000"/>
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 current English equivalent of umami; however, some close descriptions are "meaty", "savory", and "broth-like".<ref name="Yamaguchi_2000"/>


==Background==
==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| vauthors = Kean S |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=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.
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.


The sensation of umami is due to the detection of the [[Carboxylate|carboxylate anion]] of [[glutamic acid|glutamate]] in specialized receptor cells present on the human and other animal [[tongue]]s.<ref name="Symposium">{{cite book | title = International Symposium on Olfaction and Taste | volume = 1170 | veditors = Finger TE | work = The Annals of the New York Academy of Sciences | location = Hoboken, NJ |publisher=[[New York Academy of Sciences]]| year = 2009 }}</ref> Some 52 [[peptide]]s may be responsible for detecting umami taste.<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 }}</ref> Its effect is to balance taste and round out the overall flavor of a dish. Umami enhances the palatability of a wide variety of foods.<ref name="Beauchamp_2009">{{cite journal | vauthors = 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> 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>
The sensation of umami is due to the detection of the [[carboxylate anion]] of [[glutamate]] in specialized receptor cells present on human and other animal [[tongue]]s.<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><ref>{{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> Some 52 [[peptide]]s may be responsible for detecting umami taste.<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> Its effect is to balance taste and round out the overall flavor of a dish. 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> 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>
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>


Monosodium [[L-aspartate]] has an umami taste about four times less intense than 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"/>
Monosodium [[L-aspartate]] has an umami taste about four times less intense than 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"/>


==Discovery==
==Discovery==
[[File:Kikunae Ikeda.jpg|thumb|Kikunae Ikeda]]
[[File:Kikunae Ikeda.jpg|thumb|Kikunae Ikeda]]
[[Glutamic acid|Glutamate]] has a long history in cooking.<ref name = "Lehrer_2007">{{cite book | vauthors = 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=http://articles.latimes.com/1998/apr/01/food/fo-34730|title=Rot of Ages| vauthors = Perry C | date = 1 April 1998 |newspaper=[[Los Angeles Times]]|access-date=29 September 2014}}</ref> and fermented [[fish sauce]]s and [[soy sauce]]s have histories going back to the third century in China. To be sure, 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.
[[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=http://articles.latimes.com/1998/apr/01/food/fo-34730|title=Rot of Ages| author = Perry C | date = 1 April 1998 |newspaper=[[Los Angeles Times]]|access-date=29 September 2014}}</ref> and fermented [[fish sauce]]s and [[soy sauce]]s have histories going back to the third century in China. 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 | vauthors = 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 | vauthors = 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 | vauthors = 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"/>
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 | vauthors = 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 | vauthors = 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}}</ref> One of Kuninaka's most important discoveries was the [[Synergy|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.
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.
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==
==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 | vauthors = 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 | vauthors = 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" />
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>
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>
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 ==
== Foods rich in umami components ==
[[File:Dried anchovies.jpg|thumb|[[Anchovies]] are rich in umami]]
[[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 | vauthors = Adams P |publisher=Popular Science, [[Bonnier Corporation]]|date=24 November 2015|access-date=11 December 2015}}</ref>
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>


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]], [[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>
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" />
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" />


==Taste receptors==
==Taste receptors==
{{See also|Taste receptor#Savory or glutamates (Umami)}}
{{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}}
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 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"/>


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. They are [[G protein-coupled receptors]] (GPCRs) with similar signaling molecules that include [[G beta-gamma complex|G proteins beta-gamma]], [[PLCB2]] and [[Inositol triphosphate|PI3]]-mediated release of [[calcium]] (Ca<sup>2+</sup>) from intracellular stores.<ref>{{cite journal | vauthors = 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]] and [[secretion]] of [[neurotransmitter]]s including [[serotonin]].<ref name="Perez">{{cite journal | vauthors = Pérez CA, Huang L, Rong M, Kozak JA, Preuss AK, Zhang H, Max M, Margolskee RF | display-authors = 6 | title = A transient receptor potential channel expressed in taste receptor cells | journal = Nature Neuroscience | volume = 5 | issue = 11 | pages = 1169–76 | date = November 2002 | pmid = 12368808 | doi = 10.1038/nn952 | s2cid = 9010248 }}</ref><ref>{{cite journal | vauthors = Zhang Y, Hoon MA, Chandrashekar J, Mueller KL, Cook B, Wu D, Zuker CS, Ryba NJ | display-authors = 6 | title = Coding of sweet, bitter, and umami tastes: different receptor cells sharing similar signaling pathways | journal = Cell | volume = 112 | issue = 3 | pages = 293–301 | date = February 2003 | pmid = 12581520 | doi = 10.1016/S0092-8674(03)00071-0 | s2cid = 718601 | doi-access = free }}</ref><ref>{{cite journal | vauthors = Dando R, Roper SD | title = Cell-to-cell communication in intact taste buds through ATP signalling from pannexin 1 gap junction hemichannels | journal = The Journal of Physiology | volume = 587 | issue = Pt 24 | pages = 5899–906 | date = December 2009 | pmid = 19884319 | pmc = 2808547 | doi = 10.1113/jphysiol.2009.180083 }}</ref><ref>{{cite journal | vauthors = Roper SD | title = Signal transduction and information processing in mammalian taste buds | journal = Pflügers Archiv | volume = 454 | issue = 5 | pages = 759–76 | date = August 2007 | pmid = 17468883 | pmc = 3723147 | doi = 10.1007/s00424-007-0247-x }}</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. They are [[G protein-coupled receptors]] (GPCRs) with similar signaling molecules that include [[G beta-gamma complex|G proteins beta-gamma]], [[PLCB2]] and [[Inositol triphosphate|PI3]]-mediated release of [[calcium]] (Ca<sup>2+</sup>) from intracellular stores.<ref>{{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]] and [[secretion]] of [[neurotransmitter]]s including [[serotonin]].<ref name="Perez">{{cite journal | vauthors = Pérez CA, Huang L, Rong M, Kozak JA, Preuss AK, Zhang H, Max M, Margolskee RF | display-authors = 6 | title = A transient receptor potential channel expressed in taste receptor cells | journal = Nature Neuroscience | volume = 5 | issue = 11 | pages = 1169–76 | date = November 2002 | pmid = 12368808 | doi = 10.1038/nn952 | s2cid = 9010248}}</ref><ref>{{cite journal | vauthors = Zhang Y, Hoon MA, Chandrashekar J, Mueller KL, Cook B, Wu D, Zuker CS, Ryba NJ | display-authors = 6 | title = Coding of sweet, bitter, and umami tastes: different receptor cells sharing similar signaling pathways | journal = Cell | volume = 112 | issue = 3 | pages = 293–301 | date = February 2003 | pmid = 12581520 | doi = 10.1016/S0092-8674(03)00071-0 | s2cid = 718601 | doi-access = free}}</ref><ref>{{cite journal | vauthors = Dando R, Roper SD | title = Cell-to-cell communication in intact taste buds through ATP signalling from pannexin 1 gap junction hemichannels | journal = The Journal of Physiology | volume = 587 | issue = Pt 24 | pages = 5899–906 | date = December 2009 | pmid = 19884319 | pmc = 2808547 | doi = 10.1113/jphysiol.2009.180083}}</ref><ref>{{cite journal | author = Roper SD | title = Signal transduction and information processing in mammalian taste buds | journal = Pflügers Archiv | volume = 454 | issue = 5 | pages = 759–76 | date = August 2007 | pmid = 17468883 | pmc = 3723147 | doi = 10.1007/s00424-007-0247-x}}</ref>


Cells responding to umami taste stimuli do not possess typical [[synapse]]s, but ATP conveys taste signals to gustatory nerves and in turn to the [[brain]] that interprets and identifies the taste quality via the [[gut-brain axis]].<ref name="Torii_2013"/><ref>{{cite journal | vauthors = Clapp TR, Yang R, Stoick CL, Kinnamon SC, Kinnamon JC | title = Morphologic characterization of rat taste receptor cells that express components of the phospholipase C signaling pathway | journal = The Journal of Comparative Neurology | volume = 468 | issue = 3 | pages = 311–21 | date = January 2004 | pmid = 14681927 | doi = 10.1002/cne.10963 | s2cid = 32048396 }}</ref><ref>{{cite journal | vauthors = Iwatsuki K, Ichikawa R, Hiasa M, Moriyama Y, Torii K, Uneyama H | title = Identification of the vesicular nucleotide transporter (VNUT) in taste cells | journal = Biochemical and Biophysical Research Communications | volume = 388 | issue = 1 | pages = 1–5 | date = October 2009 | pmid = 19619506 | doi = 10.1016/j.bbrc.2009.07.069 }}</ref>
Cells responding to umami taste stimuli do not possess typical [[synapse]]s, but ATP conveys taste signals to gustatory nerves and in turn to the [[brain]] that interprets and identifies the taste quality via the [[gut-brain axis]].<ref name="Torii_2013"/><ref>{{cite journal | vauthors = Clapp TR, Yang R, Stoick CL, Kinnamon SC, Kinnamon JC | title = Morphologic characterization of rat taste receptor cells that express components of the phospholipase C signaling pathway | journal = The Journal of Comparative Neurology | volume = 468 | issue = 3 | pages = 311–21 | date = January 2004 | pmid = 14681927 | doi = 10.1002/cne.10963 | s2cid = 32048396}}</ref><ref>{{cite journal | vauthors = Iwatsuki K, Ichikawa R, Hiasa M, Moriyama Y, Torii K, Uneyama H | title = Identification of the vesicular nucleotide transporter (VNUT) in taste cells | journal = Biochemical and Biophysical Research Communications | volume = 388 | issue = 1 | pages = 1–5 | date = October 2009 | pmid = 19619506 | doi = 10.1016/j.bbrc.2009.07.069}}</ref>


== Consumers and safety ==
== 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|publisher=FoodNavigator.com, William Reed Business Media Ltd., Crawley, England|date=7 October 2015| vauthors = 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 | vauthors = 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|publisher=[[United States Food and Drug Administration]]|location=Silver Spring, Maryland|access-date=19 February 2017}}</ref> There is also no apparent difference in sensitivity to umami when comparing Japanese and Americans.<ref>{{cite journal | vauthors = 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>
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 ==
== 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| vauthors = 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>
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 ==
== See also ==
Line 80: Line 79:
* [[Inosinic acid]]
* [[Inosinic acid]]
* [[Monopotassium glutamate]]
* [[Monopotassium glutamate]]
* [[Tien Chu Ve-Tsin]]
* [[Umami Burger]]
}}
}}


==References==
==References==
{{Reflist|refs=
{{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="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| vauthors = Fleming A |newspaper=The Guardian |location=London, UK|date=9 April 2013|access-date=18 February 2017}}</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 | vauthors = Jufresa L |title=Umami (Mapa de las lenguas) |location=Spain |publisher=[[Penguin Random House]] Grupo Editorial México |language=Spanish |isbn=978-607-31-2817-9 |date=February 16, 2015 |type=Ebook}}</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 = "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="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="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), USA|date=5 November 2007| vauthors = Krulwich R }}</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>
}}
}}


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[[Category:Culinary terminology]]
[[Category:Culinary terminology]]
[[Category:Umami enhancers| ]]
[[Category:Umami enhancers| ]]
[[Category:1908 introductions]]
[[Category:1908 establishments in Japan]]
[[Category:1900s neologisms]]
[[Category:1985 neologisms]]

Latest revision as of 00:25, 9 May 2024

Umami
Soy sauce, ripe tomatoes and miso are examples of foods rich in umami components.

Umami (/ˈmɑːmi/ from Japanese: 旨味 Japanese pronunciation: [ɯmami]), or savoriness, is one of the five basic tastes.[1] It has been described as savory and is characteristic of broths and cooked meats.[2][3][4][5]: 35–36 

People taste umami through taste receptors that typically respond to glutamates and 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).[6][7][8] 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.[1][9]

Foods that have a strong umami flavor include meats, shellfish, fish (including fish sauce and preserved fish such as Maldives fish, Katsuobushi, sardines, and anchovies), tomatoes, mushrooms, hydrolyzed vegetable protein, meat extract, yeast extract, kimchi, cheeses, and soy sauce.

Etymology[edit]

A loanword from Japanese (うま味), umami can be translated as "pleasant savory taste".[10] This neologism was coined in 1908 by Japanese chemist Kikunae Ikeda from a nominalization of umai (うまい) "delicious". The compound 旨味 (with mi () "taste") is used for a more general sense of a food as delicious.[11][12][13] There is no current English equivalent of umami; however, some close descriptions are "meaty", "savory", and "broth-like".[14]

Background[edit]

Scientists have debated whether umami was a basic taste since Kikunae Ikeda first proposed its existence in 1908.[15][16] In 1985, the term umami was recognized as the scientific term to describe the taste of glutamates and nucleotides at the first Umami International Symposium in Hawaii.[17] Umami represents the taste of the amino acid L-glutamate and 5'-ribonucleotides such as guanosine monophosphate (GMP) and inosine monophosphate (IMP).[14] It can be described as a pleasant "brothy" or "meaty" taste with a long-lasting, mouthwatering and coating sensation over the tongue.

The sensation of umami is due to the detection of the carboxylate anion of glutamate in specialized receptor cells present on human and other animal tongues.[18][19] Some 52 peptides may be responsible for detecting umami taste.[20] Its effect is to balance taste and round out the overall flavor of a dish. Umami enhances the palatability of a wide variety of foods.[21] Glutamate in acid form (glutamic acid) imparts little umami taste, whereas the salts of glutamic acid, known as glutamates, give the characteristic umami taste due to their ionized state. GMP and IMP amplify the taste intensity of glutamate.[22] Adding salt to the free acids also enhances the umami taste.[23]

Monosodium L-aspartate has an umami taste about four times less intense than MSG, whereas ibotenic acid and tricholomic acid (likely as their salts or with salt) are claimed to be many times more intense.[23]

Discovery[edit]

Kikunae Ikeda

Glutamate has a long history in cooking.[24] Fermented fish sauces (garum), which are rich in glutamate, were used widely in ancient Rome,[25] fermented barley sauces (murri) rich in glutamate were used in medieval Byzantine and Arab cuisine,[26] and fermented fish sauces and soy sauces have histories going back to the third century in China. In the late 1800s, chef Auguste Escoffier, who opened restaurants in Paris and London, created meals that combined umami with salty, sour, sweet, and bitter tastes.[9] However, he did not know the chemical source of this unique quality.

Umami was first scientifically identified in 1908 by Kikunae Ikeda,[27][28] a professor of the Tokyo Imperial University. He found that 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.[16]

Professor Shintaro Kodama, a disciple of Ikeda, discovered in 1913 that dried bonito flakes (a type of tuna) contained another umami substance.[29] This was the ribonucleotide IMP. In 1957, Akira Kuninaka realized that the ribonucleotide GMP present in shiitake mushrooms also conferred the umami taste.[30] 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.[14]

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[edit]

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.[31][32] By itself, umami is not palatable, but it makes a great variety of foods pleasant, especially in the presence of a matching aroma.[33] Like other basic tastes, umami is pleasant only within a relatively narrow concentration range.[31]

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.[34] 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.[35] 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.[36][37]

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.[38] Some evidence exists to show umami not only stimulates appetite, but also may contribute to satiety.[39]

Foods rich in umami components[edit]

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 (IMP) comes primarily from meats and guanosine (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 adenylate.[5]: 11, 52, 110 [40]

Generally, umami taste is common to foods that contain high levels of L-glutamate, IMP and GMP, most notably in fish, shellfish, cured meats, meat extracts, mushrooms, vegetables (e.g., ripe tomatoes, Chinese cabbage, spinach, celery, etc.), green tea, hydrolyzed vegetable protein, and fermented and aged products involving bacterial or yeast cultures, such as cheeses, shrimp pastes, fish sauce, soy sauce, natto, nutritional yeast, and yeast extracts such as Vegemite and Marmite.[2][41]

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.[42][2][5]

Taste receptors[edit]

Most taste buds 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 receptors responsible for the sense of umami as modified forms of mGluR4, mGluR1, and taste receptor type 1 (TAS1R1 + TAS1R3), all of which have been found in all regions of the tongue bearing taste buds.[8][6][43] These receptors are also found in some regions of the duodenum.[44] 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."[18]

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. They are G protein-coupled receptors (GPCRs) with similar signaling molecules that include G proteins beta-gamma, PLCB2 and PI3-mediated release of calcium (Ca2+) from intracellular stores.[45] Calcium activates a so-called transient-receptor-potential cation channel TRPM5 that leads to membrane depolarization and the consequent release of ATP and secretion of neurotransmitters including serotonin.[46][47][48][49]

Cells responding to umami taste stimuli do not possess typical synapses, but ATP conveys taste signals to gustatory nerves and in turn to the brain that interprets and identifies the taste quality via the gut-brain axis.[1][50][51]

Consumers and safety[edit]

Umami has become popular as a flavor with food manufacturers trying to improve the taste of low sodium offerings.[52] Chefs create "umami bombs", which are dishes made of several umami ingredients like fish sauce.[2][9] Umami may account for the long-term formulation and popularity of ketchup.[53] 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 sensitive to MSG, a study commissioned by the FDA was only able to identify transient, mild symptoms in a few of the subjects, and only when the MSG was consumed in unrealistically large quantities.[54] There is also no apparent difference in sensitivity to umami when comparing Japanese and Americans.[55]

Background of other taste categories[edit]

The five basic tastes (saltiness, sweetness, bitterness, sourness, and savoriness) are detected by specialized taste receptors on the tongue and palate epithelium.[56] The number of taste categories in humans remains under research, with a sixth taste possibly including spicy or pungent.[57]

See also[edit]

References[edit]

  1. ^ a b c Torii K, Uneyama H, Nakamura E (April 2013). "Physiological roles of dietary glutamate signaling via gut-brain axis due to efficient digestion and absorption". Journal of Gastroenterology. 48 (4): 442–51. doi:10.1007/s00535-013-0778-1. PMC 3698427. PMID 23463402.
  2. ^ a b c d Fleming A (9 April 2013). "Umami: why the fifth taste is so important". The Guardian. Retrieved 18 February 2017.
  3. ^ Blake H (9 February 2010). "Umami in a tube: 'fifth taste' goes on sale in supermarkets". The Daily Telegraph. Archived from the original on 11 January 2022. Retrieved 10 February 2011.
  4. ^ Jufresa L (16 February 2015). Umami (Mapa de las lenguas) (ebook) (in Spanish). Spain: Penguin Random House Grupo Editorial México. ISBN 978-607-31-2817-9.
  5. ^ a b c Mouritsen JD, Drotner J, Styrbæk K, Mouritsen OG (April 2014). Umami: Unlocking the Secrets of the Fifth Taste (ebook). United States: Columbia University Press. pp. 35–36. doi:10.7312/mour16890. ISBN 978-0-231-53758-2. JSTOR 10.7312/mour16890.
  6. ^ a b Nelson G, Chandrashekar J, Hoon MA, Feng L, Zhao G, Ryba NJ, Zuker CS (March 2002). "An amino-acid taste receptor". Nature. 416 (6877): 199–202. Bibcode:2002Natur.416..199N. doi:10.1038/nature726. PMID 11894099. S2CID 1730089.
  7. ^ Delay ER, Beaver AJ, Wagner KA, Stapleton JR, Harbaugh JO, Catron KD, Roper SD (October 2000). "Taste preference synergy between glutamate receptor agonists and inosine monophosphate in rats". Chemical Senses. 25 (5): 507–15. doi:10.1093/chemse/25.5.507. PMID 11015322.
  8. ^ a b Chaudhari N, Landin AM, Roper SD (February 2000). "A metabotropic glutamate receptor variant functions as a taste receptor". Nature Neuroscience. 3 (2): 113–9. doi:10.1038/72053. PMID 10649565. S2CID 16650588.
  9. ^ a b c Krulwich R (5 November 2007). "Sweet, Sour, Salty, Bitter ... and Umami". National Public Radio (NPR).
  10. ^ Breen J. "EDICT's entry for umami". Retrieved 5 December 2017.
  11. ^ "うま味 (umami)". Japan Society of Refrigerating and Air Conditioning Engineers. Retrieved 5 June 2017.
  12. ^ "What is umami?". Columbia University Press. Retrieved 5 June 2017.
  13. ^ "Umami". Cambridge Dictionary. Cambridge University Press. Retrieved 1 January 2011.
  14. ^ a b c Yamaguchi S, Ninomiya K (April 2000). "Umami and food palatability". The Journal of Nutrition. 130 (4S Suppl): 921S–6S. doi:10.1093/jn/130.4.921S. PMID 10736353.
  15. ^ Lindemann B, Ogiwara Y, Ninomiya Y (November 2002). "The discovery of umami". Chemical Senses. 27 (9): 843–4. doi:10.1093/chemse/27.9.843. PMID 12438211.
  16. ^ a b Kean, Sam (Fall 2015). "The science of satisfaction". Distillations Magazine. 1 (3). Science History Institute: 5. Retrieved 22 March 2018.
  17. ^ Kawamura Y, Kare MR, eds. (1987). Umami: A basic taste. New York: Marcel Dekker.[page needed]
  18. ^ a b Finger, Thomas E., ed. (2009). International Symposium on Olfaction and Taste. Vol. 1170 (1st ed.). Hoboken, NJ: Wiley-Blackwell. ISBN 978-1573317382.
  19. ^ McGrane, Scott J.; Gibbs, Matthew; Hernangomez de Alvaro, Carlos; Dunlop, Nicola; Winnig, Marcel; Klebansky, Boris; Waller, Daniel (8 August 2023). "Umami taste perception and preferences of the domestic cat (Felis catus), an obligate carnivore". Chemical Senses. 48. doi:10.1093/chemse/bjad026. PMC 10468298. PMID 37551788. Retrieved 21 September 2023.
  20. ^ Zhang Y, Venkitasamy C, Pan Z, Liu W, Zhao L (January 2017). "Novel Umami Ingredients: Umami Peptides and Their Taste". Journal of Food Science. 82 (1): 16–23. doi:10.1111/1750-3841.13576. PMID 27926796.
  21. ^ Beauchamp GK (September 2009). "Sensory and receptor responses to umami: an overview of pioneering work". The American Journal of Clinical Nutrition. 90 (3): 723S–727S. doi:10.3945/ajcn.2009.27462E. PMID 19571221.
  22. ^ Yasuo T, Kusuhara Y, Yasumatsu K, Ninomiya Y (October 2008). "Multiple receptor systems for glutamate detection in the taste organ". Biological & Pharmaceutical Bulletin. 31 (10): 1833–7. doi:10.1248/bpb.31.1833. PMID 18827337.
  23. ^ a b Lioe HN, Selamat J, Yasuda M (April 2010). "Soy sauce and its umami taste: a link from the past to current situation" (PDF). Journal of Food Science. 75 (3): R71-6. doi:10.1111/j.1750-3841.2010.01529.x. PMID 20492309.
  24. ^ Lehrer J (2007). Proust was a Neuroscientist. Mariner Books. ISBN 978-0-547-08590-6.
  25. ^ Smriga M, Mizukoshi T, Iwata D, Sachise E, Miyano H, Kimura T, Curtis R (August 2010). "Amino acids and minerals in ancient remnants of fish sauce (garum) sampled in the "Garum Shop" of Pompeii, Italy". Journal of Food Composition and Analysis. 23 (5): 442–46. doi:10.1016/j.jfca.2010.03.005.
  26. ^ Perry C (1 April 1998), "Rot of Ages", Los Angeles Times, retrieved 29 September 2014
  27. ^ Ikeda K (November 2002). "New Seasonings". Chemical Senses. 27 (9): 847–9. doi:10.1093/chemse/27.9.847. PMID 12438213. (partial translation of Ikeda K (1909). "New Seasonings". Journal of the Chemical Society of Tokyo (in Japanese). 30: 820–36.)
  28. ^ Nakamura E (July 2011). "One hundred years since the discovery of the "Kawamura_2016" taste from seaweed broth by Kikunae Ikeda, who transcended his time". Chemistry: An Asian Journal. 6 (7): 1659–63. doi:10.1002/asia.201000899. PMID 21472994.
  29. ^ Kodama S (1913). "On a procedure for separating inosinic acid". Journal of the Chemical Society of Japan. 34: 751.
  30. ^ Kuninaka A (1960). "Studies on taste of ribonucleic acid derivatives". Journal of the Agricultural Chemical Society of Japan (in Japanese). 34 (6): 487–92. doi:10.1271/nogeikagaku1924.34.6_489.
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Further reading[edit]

  • Barbot P, Matsuhisa N, Mikuni K, Heston B (2009). Dashi and Umami: The Heart of Japanese Cuisine. London: Eat-Japan / Cross Media. ISBN 978-1-897-70193-5.
  • Yamaguchi S, Ninomiya K (1999). "Umami and Food Palatability". In Teranishi R, Wick EL, Hornstein I (eds.). Flavor Chemistry: Thirty Years of Progress. New York: Kluwer Academic/Plenum Publishers. ISBN 978-0-306-46199-6.

External links[edit]