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<p><span style="color: #000000"><b>Umami</b>, also referred to as <b>savoriness</b>, is one of the basic tastes sensed by specialized receptor cells present on the human and animal tongue. <i>Umami</i> <span style="font-weight: normal">(<span lang="ja" class="t_nihongo_kanji" xml:lang="ja">旨味</span><span class="t_nihongo_help noprint"><sup><span style="padding-bottom: 0px; padding-left: 0.1em; padding-right: 0.1em; font: bold 80% sans-serif; text-decoration: none; padding-top: 0px" class="t_nihongo_icon"><font size="1">?</font></span></sup></span>)</span> is a loanword from Japanese meaning "good flavor" or "good taste" (noun).<sup id="cite_ref-0" class="reference"><font size="2">[1]</font></sup> "Brothy", "meaty", or "savory" have been proposed as alternative translations, however.<sup id="cite_ref-nature06_1-0" class="reference"><font size="2">[2]</font></sup><sup id="cite_ref-ikeda02_2-0" class="reference"><font size="2">[3]</font></sup> In as much as it describes the flavor common to savory products such as meat, cheese, and mushrooms, umami is similar to Brillat-Savarin's concept of <i>osmazome</i>, an early Western attempt to describe the main flavoring component of meat as extracted in the process of making stock.</span></p>
<p><span style="color: #000000">The umami taste is due to the detection of the carboxylate anion of glutamic acid, a naturally occurring amino acid common in meat (particularly bacon), cheese, broth, stock, and other protein-heavy foods. Salts of glutamic acid, known as glutamates, easily ionize to give the same carboxylate form and therefore the same taste. For this reason, they are used as flavor enhancers. The most commonly used of these is monosodium glutamate (MSG). While the umami taste is due to glutamates, 5'-ribonucleotides such as guanosine monophosphate (GMP) and inosine monophosphate (IMP) greatly enhance its perceived intensity. Since these ribonucleotides are also acids, their salts are sometimes added together with glutamates to obtain a synergistic flavor enhancement effect.<sup id="cite_ref-nature06_1-1" class="reference"><font size="2">[2]</font></sup><sup id="cite_ref-pmid18827337_3-0" class="reference"><font size="2">[4]</font></sup></span></p>
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<h2><span style="color: #000000">Contents</span></h2>
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<li class="toclevel-1 tocsection-1"><span style="color: #000000"><span class="tocnumber">1</span> <span class="toctext">Excitatory substances</span></span></li>
<li class="toclevel-1 tocsection-2"><span style="color: #000000"><span class="tocnumber">2</span> <span class="toctext">Taste receptors</span></span></li>
<li class="toclevel-1 tocsection-3"><span style="color: #000000"><span class="tocnumber">3</span> <span class="toctext">Bacon</span></span></li>
<li class="toclevel-1 tocsection-4"><span style="color: #000000"><span class="tocnumber">4</span> <span class="toctext">See also</span></span></li>
<li class="toclevel-1 tocsection-5"><span style="color: #000000"><span class="tocnumber">5</span> <span class="toctext">Notes</span></span></li>
<li class="toclevel-1 tocsection-6"><span style="color: #000000"><span class="tocnumber">6</span> <span class="toctext">References</span></span></li>
<li class="toclevel-1 tocsection-7"> </li>
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<h2><span style="color: #000000"><span id="Excitatory_substances" class="mw-headline">Excitatory substances</span></span></h2>
<div class="rellink relarticle mainarticle"><span style="color: #000000">Main article: Glutamic acid (flavor)</span></div>
<div class="rellink boilerplate seealso"><span style="color: #000000">See also: Guanosine monophosphate and Inosine monophosphate</span></div>
<p><span style="color: #000000">Umami as a separate taste was first identified in 1908 by Kikunae Ikeda of the Tokyo Imperial University while researching the strong flavor in seaweed broth.<sup id="cite_ref-ikeda02_2-1" class="reference"><font size="2">[3]</font></sup> Ikeda isolated monosodium glutamate (MSG) as the chemical responsible and, with the help of the Ajinomoto company, began commercial distribution of MSG products.</span></p>
<p><span style="color: #000000">Glutamate has a long history in cooking.<sup id="cite_ref-Lehrer07_4-0" class="reference"><font size="2">[5]</font></sup> It is naturally found in East and Southeast Asian foods, such as soy sauce and fish sauce, and in Italian foods like Parmesan cheese, anchovies and ripe tomatoes (though tomatoes did not originate in Europe). It is also prevalent in seafood, such as lobster, crabs, and shrimp.<sup id="cite_ref-5" class="reference"><font size="2">[6]</font></sup></span></p>
<h2><span style="color: #000000"><span id="Taste_receptors" class="mw-headline">Taste receptors</span></span></h2>
<p><span style="color: #000000">Umami has been described in biochemical studies identifying the actual taste receptor responsible for the sense of umami, a modified form of mGluR4<sup id="cite_ref-6" class="reference"><font size="2">[7]</font></sup> named "taste-mGluR4."</span></p>
<p><span style="color: #000000">Umami tastes are initiated by these specialized receptors, with subsequent steps involving secretion of neurotransmitters including serotonin.<sup id="cite_ref-7" class="reference"><font size="2">[8]</font></sup> Other evidence indicates guanosine derivatives may interact with and boost the initial umami signal.<sup id="cite_ref-8" class="reference"><font size="2">[9]</font></sup></span></p>
<p><span style="color: #000000">Cells responding to umami taste stimuli do not possess typical synapses, but instead secrete the neurotransmitter ATP in a mechanism exciting sensory fibers that convey taste signals to the brain.</span></p>
<p><span style="color: #000000">In monkey studies, most umami signals from taste buds excite neurons in the orbitofrontal cortex of the brain, showing spatially specific characteristics:<sup id="cite_ref-9" class="reference"><font size="2">[10]</font></sup></span></p>
<ul>
<li><span style="color: #000000">Single neurons having vigorous responses to sodium glutamate also respond to glutamic acid</span></li>
<li><span style="color: #000000">Some neurons display a mechanism of satiety.</span></li>
</ul>
<p><span style="color: #000000">The stomach can "taste" sodium glutamate using glumate receptors<sup id="cite_ref-10" class="reference"><font size="2">[11]</font></sup> and this information is passed to the lateral hypothalamus and limbic system in the brain as a palatability signal through the vagus nerve.<sup id="cite_ref-11" class="reference"><font size="2">[12]</font></sup></span></p>
<h2><span style="color: #000000"><span id="Bacon" class="mw-headline">Bacon</span></span></h2>
<p><span style="color: #000000">Arun Gupta of <i>The Indypendent</i> has pointed out how bacon in particular possesses six ingredient types of umami, which elicits an addictive neurochemical response.<sup id="cite_ref-12" class="reference"><font size="2">[13]</font></sup> According to Gupta "the chain lards on bacon in particular" give foods a "high flavor profile" creating a "one-of-a-kind product that has no taste substitute."<sup id="cite_ref-AlterBacon_13-0" class="reference"><font size="2">[14]</font></sup> This led Dr. David Kessler, author of <i>The End of Overeating</i>, to note how the standard joke in the restaurant chain industry goes, "When in doubt, throw cheese and bacon on it."<sup id="cite_ref-AlterBacon_13-1" class="reference"><font size="2">[14]</font></sup></span></p>
<h2><span style="color: #000000"><span id="See_also" class="mw-headline">See also</span></span></h2>
<ul>
<li><span style="color: #000000">Inosine monophosphate</span></li>
</ul>
<h2><span style="color: #000000"><span id="Notes" class="mw-headline">Notes</span></span></h2>
<div style="column-count: 2; -moz-column-count: 2; -webkit-column-count: 2" class="references-small references-column-count references-column-count-2">
<ol class="references">
<li id="cite_note-0"><span style="color: #000000"><b>^</b> http://jisho.org/words?jap=%E6%97%A8%E5%91%B3&eng=&dict=edict</span></li>
<li id="cite_note-nature06-1"><span style="color: #000000">^ <sup><i><b><font size="2">a</font></b></i></sup> <sup><i><b><font size="2">b</font></b></i></sup> <span class="citation Journal">Chandrashekar J, Hoon MA, Ryba NJ, Zuker CS (November 2006). "The receptors and cells for mammalian taste". <i>Nature</i> <b>444</b> (7117): 288–94. doi:10.1038/nature05401. PMID 17108952.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=The+receptors+and+cells+for+mammalian+taste&rft.jtitle=%5B%5BNature+%28journal%29%7CNature%5D%5D&rft.aulast=Chandrashekar+J%2C+Hoon+MA%2C+Ryba+NJ%2C+Zuker+CS&rft.au=Chandrashekar+J%2C+Hoon+MA%2C+Ryba+NJ%2C+Zuker+CS&rft.date=November+2006&rft.volume=444&rft.issue=7117&rft.pages=288%E2%80%9394&rft_id=info:doi/10.1038%2Fnature05401&rft_id=info:pmid/17108952&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-ikeda02-2"><span style="color: #000000">^ <sup><i><b><font size="2">a</font></b></i></sup> <sup><i><b><font size="2">b</font></b></i></sup> <span class="citation Journal">Ikeda K (November 2002). "New seasonings". <i>Chemical Senses</i> <b>27</b> (9): 847–9. doi:10.1093/chemse/27.9.847. PMID 12438213.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=New+seasonings&rft.jtitle=Chemical+Senses&rft.aulast=Ikeda+K&rft.au=Ikeda+K&rft.date=November+2002&rft.volume=27&rft.issue=9&rft.pages=847%E2%80%939&rft_id=info:doi/10.1093%2Fchemse%2F27.9.847&rft_id=info:pmid/12438213&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span> (partial translation of <span class="citation Journal">Ikeda, Kikunae (1909). "New Seasonings[japan.]". <i>Journal of the Chemical Society of Tokyo</i> <b>30</b>: 820–836.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=New+Seasonings%5Bjapan.%5D&rft.jtitle=Journal+of+the+Chemical+Society+of+Tokyo&rft.aulast=Ikeda&rft.aufirst=Kikunae&rft.au=Ikeda%2C%26%2332%3BKikunae&rft.date=1909&rft.volume=30&rft.pages=820%E2%80%93836&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span>)</span></li>
<li id="cite_note-pmid18827337-3"><span style="color: #000000"><b>^</b> <span class="citation Journal">Yasuo T, Kusuhara Y, Yasumatsu K, Ninomiya Y (October 2008). "Multiple receptor systems for glutamate detection in the taste organ". <i>Biological & Pharmaceutical Bulletin</i> <b>31</b> (10): 1833–7. doi:10.1248/bpb.31.1833. PMID 18827337.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=Multiple+receptor+systems+for+glutamate+detection+in+the+taste+organ&rft.jtitle=Biological+%26+Pharmaceutical+Bulletin&rft.aulast=Yasuo+T%2C+Kusuhara+Y%2C+Yasumatsu+K%2C+Ninomiya+Y&rft.au=Yasuo+T%2C+Kusuhara+Y%2C+Yasumatsu+K%2C+Ninomiya+Y&rft.date=October+2008&rft.volume=31&rft.issue=10&rft.pages=1833%E2%80%937&rft_id=info:doi/10.1248%2Fbpb.31.1833&rft_id=info:pmid/18827337&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-Lehrer07-4"><span style="color: #000000"><b>^</b> <span id="Lehrer07" class="citation book">Lehrer, Jonah (2007). <i>Proust was a Neuroscientist</i>. Mariner Books. doi:0908/1043570. ISBN 9780547085906.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Proust+was+a+Neuroscientist&rft.aulast=Lehrer&rft.aufirst=Jonah&rft.au=Lehrer%2C%26%2332%3BJonah&rft.date=2007&rft.pub=%5B%5BMariner+Books%5D%5D&rft_id=info:doi/0908%2F1043570&rft.isbn=9780547085906&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-5"><span style="color: #000000"><b>^</b> <span class="citation news">Moskin, Julia (5 March 2008). "Yes, MSG, the Secret Behind the Savor". <i>New York Times</i><span class="printonly">. http://nytimes.com/2008/03/05/dining/05glute.html</span><span class="reference-accessdate">. Retrieved 9 August 2008</span>.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=Yes%2C+MSG%2C+the+Secret+Behind+the+Savor&rft.jtitle=New+York+Times&rft.aulast=Moskin&rft.aufirst=Julia&rft.au=Moskin%2C%26%2332%3BJulia&rft.date=5+March+2008&rft_id=http%3A%2F%2Fnytimes.com%2F2008%2F03%2F05%2Fdining%2F05glute.html&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-6"><span style="color: #000000"><b>^</b> <span class="citation Journal">Nelson G, Chandrashekar J, Hoon MA, <i>et al.</i> (2002). "An amino-acid taste receptor". <i>Nature</i> <b>416</b> (6877): 199–202. doi:10.1038/nature726. PMID 11894099.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=An+amino-acid+taste+receptor&rft.jtitle=Nature&rft.aulast=Nelson+G%2C+Chandrashekar+J%2C+Hoon+MA%2C+%27%27et+al.%27%27&rft.au=Nelson+G%2C+Chandrashekar+J%2C+Hoon+MA%2C+%27%27et+al.%27%27&rft.date=2002&rft.volume=416&rft.issue=6877&rft.pages=199%E2%80%93202&rft_id=info:doi/10.1038%2Fnature726&rft_id=info:pmid/11894099&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-7"><span style="color: #000000"><b>^</b> <span class="citation Journal">Roper SD (August 2007). "Signal transduction and information processing in mammalian taste buds". <i>Pflügers Archiv</i> <b>454</b> (5): 759–76. doi:10.1007/s00424-007-0247-x. PMID 17468883.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=Signal+transduction+and+information+processing+in+mammalian+taste+buds&rft.jtitle=Pfl%C3%BCgers+Archiv&rft.aulast=Roper+SD&rft.au=Roper+SD&rft.date=August+2007&rft.volume=454&rft.issue=5&rft.pages=759%E2%80%9376&rft_id=info:doi/10.1007%2Fs00424-007-0247-x&rft_id=info:pmid/17468883&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-8"><span style="color: #000000"><b>^</b> <span class="citation Journal">Daniels, S (February 18, 2008). "Scientists develop new umami taste enhancers". <i>FoodNavigator.com-Europe</i><span class="printonly">. http://foodnfoodnavigator.com/news/ng.asp?id=83328-umami-msg-gmp</span>.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=Scientists+develop+new+umami+taste+enhancers&rft.jtitle=FoodNavigator.com-Europe&rft.aulast=Daniels&rft.aufirst=S&rft.au=Daniels%2C%26%2332%3BS&rft.date=February+18%2C+2008&rft_id=http%3A%2F%2Ffoodnfoodnavigator.com%2Fnews%2Fng.asp%3Fid%3D83328-umami-msg-gmp&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-9"><span style="color: #000000"><b>^</b> <span class="citation Journal">Rolls ET (April 2000). "The representation of umami taste in the taste cortex". <i>The Journal of Nutrition</i> <b>130</b> (4S Suppl): 960S–5S. PMID 10736361<span class="printonly">. http://jn.nutrition.org/cgi/pmidlookup?view=long&pmid=10736361</span>.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=The+representation+of+umami+taste+in+the+taste+cortex&rft.jtitle=The+Journal+of+Nutrition&rft.aulast=Rolls+ET&rft.au=Rolls+ET&rft.date=April+2000&rft.volume=130&rft.issue=4S+Suppl&rft.pages=960S%E2%80%935S&rft_id=info:pmid/10736361&rft_id=http%3A%2F%2Fjn.nutrition.org%2Fcgi%2Fpmidlookup%3Fview%3Dlong%26pmid%3D10736361&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-10"><span style="color: #000000"><b>^</b> Uematsu A, Tsurugizawa T, Kondoh T, Torii K. (2009). Conditioned flavor preference learning by intragastric administration of L-glutamate in rats. Neurosci Lett. Feb 27;451(3):190-3. PMID 19146916</span></li>
<li id="cite_note-11"><span style="color: #000000"><b>^</b> Uematsu A, Tsurugizawa T, Uneyama H, Torii K. (2010). Brain-gut communication via vagus nerve modulates conditioned flavor preference. Eur J Neurosci. 31(6):1136-43. doi:10.1111/j.1460-9568.2010.07136.x PMID 20377626</span></li>
<li id="cite_note-12"><span style="color: #000000"><b>^</b> "Bacon as a Weapon of Mass Destruction" - video report with Arun Gupta by <i>Democracy Now!</i>, August 3, 2009</span></li>
<li id="cite_note-AlterBacon-13"><span style="color: #000000">^ <sup><i><b><font size="2">a</font></b></i></sup> <sup><i><b><font size="2">b</font></b></i></sup> Gonzo Gastronomy: How the Food Industry Has Made Bacon a Weapon of Mass Destruction by Arun Gupta, <i>AlterNet</i>, October 9, 2010</span></li>
</ol>
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<h2><span style="color: #000000"><span id="References" class="mw-headline">References</span></span></h2>
<div style="margin-left: 1.5em" class="references-small">
<ul>
<li><span style="color: #000000">Flavor Chemistry: Thirty Years of Progress By Roy Teranishi, Emily L. Wick, Irwin Hornstein; Article: Umami and Food Palatability, by Shizuko Yamaguchi and Kumiko Ninomiya. ISBN 0306461994</span></li>
</ul>
<ul>
<li><span style="color: #000000">Barbot, Pascal; Matsuhisa, Nobu; and Mikuni, Kiyomi. Foreword by Heston Blumenthal. <i>Dashi and Umami: The Heart of Japanese Cuisine</i>. London: Eat-Japan / Cross Media, 2009</span></li>
</ul>
</div>
<h2><span style="color: #000000"><span id="External_links" class="mw-headline">External links</span></span></h2>
<ul>
<li><a class="external text" href="http://www.umami.it" rel="nofollow"><font color="#3366bb">Italian umami research</font></a></li>
<li><a class="external text" href="http://www.umamiinfo.com/what_exactly_is_umami?/" rel="nofollow"><font color="#3366bb">Trade Group Website</font></a></li>
<li><a class="external text" href="http://www.jpo.go.jp/seido_e/rekishi_e/kikunae_ikeda.htm" rel="nofollow"><font color="#3366bb">Discovery of Umami</font></a></li>
<li><a class="external text" href="http://www.jneurosci.org/cgi/content/short/26/8/2227" rel="nofollow"><font color="#3366bb">Discovery of Umami Receptors</font></a></li>
<li><a class="external text" href="http://www.srut.org/index2_e.asp" rel="nofollow"><font color="#3366bb">Society for Research on Umami Taste</font></a></li>
<li><a class="external text" href="http://www.cbc.ca/news/background/senses/umami.html" rel="nofollow"><font color="#3366bb">"Who's umami? Human taste now comes in five flavours"</font></a>, CBC News, June 1, 2007</li>
<li><a class="external text" href="http://www.npr.org/templates/story/story.php?storyId=15819485" rel="nofollow"><font color="#3366bb">"Sweet, Sour, Salty, Bitter… and Umami"</font></a> NPR, November 1, 2007</li>
<li><a class="external text" href="http://online.wsj.com/article/SB119706514515417586.html" rel="nofollow"><font color="#3366bb">A New Taste Sensation</font></a>, by Katy McLaughlin, Wall Street Journal, 12/8/07.</li>
<li><a class="external text" href="http://query.nytimes.com/gst/fullpage.html?res=9C03E7D91F3CF936A2575BC0A9629C8B63&sec=&spon=&pagewanted=all" rel="nofollow"><font color="#3366bb">Mouthfuls</font></a> : <a class="mw-redirect" title="Huitlacoche" href="/wiki/Huitlacoche"><font color="#0645ad">huitlacoche</font></a> is another <i>beispiel</i>, 24/7/08.</li>
</ul>
<p><span style="color: #000000">The umami taste is due to the detection of the carboxylate anion of glutamic acid, a naturally occurring amino acid common in meat (particularly bacon), cheese, broth, stock, and other protein-heavy foods. Salts of glutamic acid, known as glutamates, easily ionize to give the same carboxylate form and therefore the same taste. For this reason, they are used as flavor enhancers. The most commonly used of these is monosodium glutamate (MSG). While the umami taste is due to glutamates, 5'-ribonucleotides such as guanosine monophosphate (GMP) and inosine monophosphate (IMP) greatly enhance its perceived intensity. Since these ribonucleotides are also acids, their salts are sometimes added together with glutamates to obtain a synergistic flavor enhancement effect.<sup id="cite_ref-nature06_1-1" class="reference"><font size="2">[2]</font></sup><sup id="cite_ref-pmid18827337_3-0" class="reference"><font size="2">[4]</font></sup></span></p>
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<h2><span style="color: #000000">Contents</span></h2>
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<ul>
<li class="toclevel-1 tocsection-1"><span style="color: #000000"><span class="tocnumber">1</span> <span class="toctext">Excitatory substances</span></span></li>
<li class="toclevel-1 tocsection-2"><span style="color: #000000"><span class="tocnumber">2</span> <span class="toctext">Taste receptors</span></span></li>
<li class="toclevel-1 tocsection-3"><span style="color: #000000"><span class="tocnumber">3</span> <span class="toctext">Bacon</span></span></li>
<li class="toclevel-1 tocsection-4"><span style="color: #000000"><span class="tocnumber">4</span> <span class="toctext">See also</span></span></li>
<li class="toclevel-1 tocsection-5"><span style="color: #000000"><span class="tocnumber">5</span> <span class="toctext">Notes</span></span></li>
<li class="toclevel-1 tocsection-6"><span style="color: #000000"><span class="tocnumber">6</span> <span class="toctext">References</span></span></li>
<li class="toclevel-1 tocsection-7"> </li>
</ul>
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<li class="toclevel-1 tocsection-7"><span class="tocnumber">7</span> <span class="toctext">External links</span></li>
</ul>
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<h2><span style="color: #000000"><span id="Excitatory_substances" class="mw-headline">Excitatory substances</span></span></h2>
<div class="rellink relarticle mainarticle"><span style="color: #000000">Main article: Glutamic acid (flavor)</span></div>
<div class="rellink boilerplate seealso"><span style="color: #000000">See also: Guanosine monophosphate and Inosine monophosphate</span></div>
<p><span style="color: #000000">Umami as a separate taste was first identified in 1908 by Kikunae Ikeda of the Tokyo Imperial University while researching the strong flavor in seaweed broth.<sup id="cite_ref-ikeda02_2-1" class="reference"><font size="2">[3]</font></sup> Ikeda isolated monosodium glutamate (MSG) as the chemical responsible and, with the help of the Ajinomoto company, began commercial distribution of MSG products.</span></p>
<p><span style="color: #000000">Glutamate has a long history in cooking.<sup id="cite_ref-Lehrer07_4-0" class="reference"><font size="2">[5]</font></sup> It is naturally found in East and Southeast Asian foods, such as soy sauce and fish sauce, and in Italian foods like Parmesan cheese, anchovies and ripe tomatoes (though tomatoes did not originate in Europe). It is also prevalent in seafood, such as lobster, crabs, and shrimp.<sup id="cite_ref-5" class="reference"><font size="2">[6]</font></sup></span></p>
<h2><span style="color: #000000"><span id="Taste_receptors" class="mw-headline">Taste receptors</span></span></h2>
<p><span style="color: #000000">Umami has been described in biochemical studies identifying the actual taste receptor responsible for the sense of umami, a modified form of mGluR4<sup id="cite_ref-6" class="reference"><font size="2">[7]</font></sup> named "taste-mGluR4."</span></p>
<p><span style="color: #000000">Umami tastes are initiated by these specialized receptors, with subsequent steps involving secretion of neurotransmitters including serotonin.<sup id="cite_ref-7" class="reference"><font size="2">[8]</font></sup> Other evidence indicates guanosine derivatives may interact with and boost the initial umami signal.<sup id="cite_ref-8" class="reference"><font size="2">[9]</font></sup></span></p>
<p><span style="color: #000000">Cells responding to umami taste stimuli do not possess typical synapses, but instead secrete the neurotransmitter ATP in a mechanism exciting sensory fibers that convey taste signals to the brain.</span></p>
<p><span style="color: #000000">In monkey studies, most umami signals from taste buds excite neurons in the orbitofrontal cortex of the brain, showing spatially specific characteristics:<sup id="cite_ref-9" class="reference"><font size="2">[10]</font></sup></span></p>
<ul>
<li><span style="color: #000000">Single neurons having vigorous responses to sodium glutamate also respond to glutamic acid</span></li>
<li><span style="color: #000000">Some neurons display a mechanism of satiety.</span></li>
</ul>
<p><span style="color: #000000">The stomach can "taste" sodium glutamate using glumate receptors<sup id="cite_ref-10" class="reference"><font size="2">[11]</font></sup> and this information is passed to the lateral hypothalamus and limbic system in the brain as a palatability signal through the vagus nerve.<sup id="cite_ref-11" class="reference"><font size="2">[12]</font></sup></span></p>
<h2><span style="color: #000000"><span id="Bacon" class="mw-headline">Bacon</span></span></h2>
<p><span style="color: #000000">Arun Gupta of <i>The Indypendent</i> has pointed out how bacon in particular possesses six ingredient types of umami, which elicits an addictive neurochemical response.<sup id="cite_ref-12" class="reference"><font size="2">[13]</font></sup> According to Gupta "the chain lards on bacon in particular" give foods a "high flavor profile" creating a "one-of-a-kind product that has no taste substitute."<sup id="cite_ref-AlterBacon_13-0" class="reference"><font size="2">[14]</font></sup> This led Dr. David Kessler, author of <i>The End of Overeating</i>, to note how the standard joke in the restaurant chain industry goes, "When in doubt, throw cheese and bacon on it."<sup id="cite_ref-AlterBacon_13-1" class="reference"><font size="2">[14]</font></sup></span></p>
<h2><span style="color: #000000"><span id="See_also" class="mw-headline">See also</span></span></h2>
<ul>
<li><span style="color: #000000">Inosine monophosphate</span></li>
</ul>
<h2><span style="color: #000000"><span id="Notes" class="mw-headline">Notes</span></span></h2>
<div style="column-count: 2; -moz-column-count: 2; -webkit-column-count: 2" class="references-small references-column-count references-column-count-2">
<ol class="references">
<li id="cite_note-0"><span style="color: #000000"><b>^</b> http://jisho.org/words?jap=%E6%97%A8%E5%91%B3&eng=&dict=edict</span></li>
<li id="cite_note-nature06-1"><span style="color: #000000">^ <sup><i><b><font size="2">a</font></b></i></sup> <sup><i><b><font size="2">b</font></b></i></sup> <span class="citation Journal">Chandrashekar J, Hoon MA, Ryba NJ, Zuker CS (November 2006). "The receptors and cells for mammalian taste". <i>Nature</i> <b>444</b> (7117): 288–94. doi:10.1038/nature05401. PMID 17108952.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=The+receptors+and+cells+for+mammalian+taste&rft.jtitle=%5B%5BNature+%28journal%29%7CNature%5D%5D&rft.aulast=Chandrashekar+J%2C+Hoon+MA%2C+Ryba+NJ%2C+Zuker+CS&rft.au=Chandrashekar+J%2C+Hoon+MA%2C+Ryba+NJ%2C+Zuker+CS&rft.date=November+2006&rft.volume=444&rft.issue=7117&rft.pages=288%E2%80%9394&rft_id=info:doi/10.1038%2Fnature05401&rft_id=info:pmid/17108952&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-ikeda02-2"><span style="color: #000000">^ <sup><i><b><font size="2">a</font></b></i></sup> <sup><i><b><font size="2">b</font></b></i></sup> <span class="citation Journal">Ikeda K (November 2002). "New seasonings". <i>Chemical Senses</i> <b>27</b> (9): 847–9. doi:10.1093/chemse/27.9.847. PMID 12438213.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=New+seasonings&rft.jtitle=Chemical+Senses&rft.aulast=Ikeda+K&rft.au=Ikeda+K&rft.date=November+2002&rft.volume=27&rft.issue=9&rft.pages=847%E2%80%939&rft_id=info:doi/10.1093%2Fchemse%2F27.9.847&rft_id=info:pmid/12438213&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span> (partial translation of <span class="citation Journal">Ikeda, Kikunae (1909). "New Seasonings[japan.]". <i>Journal of the Chemical Society of Tokyo</i> <b>30</b>: 820–836.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=New+Seasonings%5Bjapan.%5D&rft.jtitle=Journal+of+the+Chemical+Society+of+Tokyo&rft.aulast=Ikeda&rft.aufirst=Kikunae&rft.au=Ikeda%2C%26%2332%3BKikunae&rft.date=1909&rft.volume=30&rft.pages=820%E2%80%93836&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span>)</span></li>
<li id="cite_note-pmid18827337-3"><span style="color: #000000"><b>^</b> <span class="citation Journal">Yasuo T, Kusuhara Y, Yasumatsu K, Ninomiya Y (October 2008). "Multiple receptor systems for glutamate detection in the taste organ". <i>Biological & Pharmaceutical Bulletin</i> <b>31</b> (10): 1833–7. doi:10.1248/bpb.31.1833. PMID 18827337.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=Multiple+receptor+systems+for+glutamate+detection+in+the+taste+organ&rft.jtitle=Biological+%26+Pharmaceutical+Bulletin&rft.aulast=Yasuo+T%2C+Kusuhara+Y%2C+Yasumatsu+K%2C+Ninomiya+Y&rft.au=Yasuo+T%2C+Kusuhara+Y%2C+Yasumatsu+K%2C+Ninomiya+Y&rft.date=October+2008&rft.volume=31&rft.issue=10&rft.pages=1833%E2%80%937&rft_id=info:doi/10.1248%2Fbpb.31.1833&rft_id=info:pmid/18827337&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-Lehrer07-4"><span style="color: #000000"><b>^</b> <span id="Lehrer07" class="citation book">Lehrer, Jonah (2007). <i>Proust was a Neuroscientist</i>. Mariner Books. doi:0908/1043570. ISBN 9780547085906.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Proust+was+a+Neuroscientist&rft.aulast=Lehrer&rft.aufirst=Jonah&rft.au=Lehrer%2C%26%2332%3BJonah&rft.date=2007&rft.pub=%5B%5BMariner+Books%5D%5D&rft_id=info:doi/0908%2F1043570&rft.isbn=9780547085906&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-5"><span style="color: #000000"><b>^</b> <span class="citation news">Moskin, Julia (5 March 2008). "Yes, MSG, the Secret Behind the Savor". <i>New York Times</i><span class="printonly">. http://nytimes.com/2008/03/05/dining/05glute.html</span><span class="reference-accessdate">. Retrieved 9 August 2008</span>.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=Yes%2C+MSG%2C+the+Secret+Behind+the+Savor&rft.jtitle=New+York+Times&rft.aulast=Moskin&rft.aufirst=Julia&rft.au=Moskin%2C%26%2332%3BJulia&rft.date=5+March+2008&rft_id=http%3A%2F%2Fnytimes.com%2F2008%2F03%2F05%2Fdining%2F05glute.html&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-6"><span style="color: #000000"><b>^</b> <span class="citation Journal">Nelson G, Chandrashekar J, Hoon MA, <i>et al.</i> (2002). "An amino-acid taste receptor". <i>Nature</i> <b>416</b> (6877): 199–202. doi:10.1038/nature726. PMID 11894099.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=An+amino-acid+taste+receptor&rft.jtitle=Nature&rft.aulast=Nelson+G%2C+Chandrashekar+J%2C+Hoon+MA%2C+%27%27et+al.%27%27&rft.au=Nelson+G%2C+Chandrashekar+J%2C+Hoon+MA%2C+%27%27et+al.%27%27&rft.date=2002&rft.volume=416&rft.issue=6877&rft.pages=199%E2%80%93202&rft_id=info:doi/10.1038%2Fnature726&rft_id=info:pmid/11894099&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-7"><span style="color: #000000"><b>^</b> <span class="citation Journal">Roper SD (August 2007). "Signal transduction and information processing in mammalian taste buds". <i>Pflügers Archiv</i> <b>454</b> (5): 759–76. doi:10.1007/s00424-007-0247-x. PMID 17468883.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=Signal+transduction+and+information+processing+in+mammalian+taste+buds&rft.jtitle=Pfl%C3%BCgers+Archiv&rft.aulast=Roper+SD&rft.au=Roper+SD&rft.date=August+2007&rft.volume=454&rft.issue=5&rft.pages=759%E2%80%9376&rft_id=info:doi/10.1007%2Fs00424-007-0247-x&rft_id=info:pmid/17468883&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-8"><span style="color: #000000"><b>^</b> <span class="citation Journal">Daniels, S (February 18, 2008). "Scientists develop new umami taste enhancers". <i>FoodNavigator.com-Europe</i><span class="printonly">. http://foodnfoodnavigator.com/news/ng.asp?id=83328-umami-msg-gmp</span>.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=Scientists+develop+new+umami+taste+enhancers&rft.jtitle=FoodNavigator.com-Europe&rft.aulast=Daniels&rft.aufirst=S&rft.au=Daniels%2C%26%2332%3BS&rft.date=February+18%2C+2008&rft_id=http%3A%2F%2Ffoodnfoodnavigator.com%2Fnews%2Fng.asp%3Fid%3D83328-umami-msg-gmp&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-9"><span style="color: #000000"><b>^</b> <span class="citation Journal">Rolls ET (April 2000). "The representation of umami taste in the taste cortex". <i>The Journal of Nutrition</i> <b>130</b> (4S Suppl): 960S–5S. PMID 10736361<span class="printonly">. http://jn.nutrition.org/cgi/pmidlookup?view=long&pmid=10736361</span>.</span><span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=The+representation+of+umami+taste+in+the+taste+cortex&rft.jtitle=The+Journal+of+Nutrition&rft.aulast=Rolls+ET&rft.au=Rolls+ET&rft.date=April+2000&rft.volume=130&rft.issue=4S+Suppl&rft.pages=960S%E2%80%935S&rft_id=info:pmid/10736361&rft_id=http%3A%2F%2Fjn.nutrition.org%2Fcgi%2Fpmidlookup%3Fview%3Dlong%26pmid%3D10736361&rfr_id=info:sid/en.wikipedia.org:Umami"><span style="display: none"> </span></span></span></li>
<li id="cite_note-10"><span style="color: #000000"><b>^</b> Uematsu A, Tsurugizawa T, Kondoh T, Torii K. (2009). Conditioned flavor preference learning by intragastric administration of L-glutamate in rats. Neurosci Lett. Feb 27;451(3):190-3. PMID 19146916</span></li>
<li id="cite_note-11"><span style="color: #000000"><b>^</b> Uematsu A, Tsurugizawa T, Uneyama H, Torii K. (2010). Brain-gut communication via vagus nerve modulates conditioned flavor preference. Eur J Neurosci. 31(6):1136-43. doi:10.1111/j.1460-9568.2010.07136.x PMID 20377626</span></li>
<li id="cite_note-12"><span style="color: #000000"><b>^</b> "Bacon as a Weapon of Mass Destruction" - video report with Arun Gupta by <i>Democracy Now!</i>, August 3, 2009</span></li>
<li id="cite_note-AlterBacon-13"><span style="color: #000000">^ <sup><i><b><font size="2">a</font></b></i></sup> <sup><i><b><font size="2">b</font></b></i></sup> Gonzo Gastronomy: How the Food Industry Has Made Bacon a Weapon of Mass Destruction by Arun Gupta, <i>AlterNet</i>, October 9, 2010</span></li>
</ol>
</div>
<h2><span style="color: #000000"><span id="References" class="mw-headline">References</span></span></h2>
<div style="margin-left: 1.5em" class="references-small">
<ul>
<li><span style="color: #000000">Flavor Chemistry: Thirty Years of Progress By Roy Teranishi, Emily L. Wick, Irwin Hornstein; Article: Umami and Food Palatability, by Shizuko Yamaguchi and Kumiko Ninomiya. ISBN 0306461994</span></li>
</ul>
<ul>
<li><span style="color: #000000">Barbot, Pascal; Matsuhisa, Nobu; and Mikuni, Kiyomi. Foreword by Heston Blumenthal. <i>Dashi and Umami: The Heart of Japanese Cuisine</i>. London: Eat-Japan / Cross Media, 2009</span></li>
</ul>
</div>
<h2><span style="color: #000000"><span id="External_links" class="mw-headline">External links</span></span></h2>
<ul>
<li><a class="external text" href="http://www.umami.it" rel="nofollow"><font color="#3366bb">Italian umami research</font></a></li>
<li><a class="external text" href="http://www.umamiinfo.com/what_exactly_is_umami?/" rel="nofollow"><font color="#3366bb">Trade Group Website</font></a></li>
<li><a class="external text" href="http://www.jpo.go.jp/seido_e/rekishi_e/kikunae_ikeda.htm" rel="nofollow"><font color="#3366bb">Discovery of Umami</font></a></li>
<li><a class="external text" href="http://www.jneurosci.org/cgi/content/short/26/8/2227" rel="nofollow"><font color="#3366bb">Discovery of Umami Receptors</font></a></li>
<li><a class="external text" href="http://www.srut.org/index2_e.asp" rel="nofollow"><font color="#3366bb">Society for Research on Umami Taste</font></a></li>
<li><a class="external text" href="http://www.cbc.ca/news/background/senses/umami.html" rel="nofollow"><font color="#3366bb">"Who's umami? Human taste now comes in five flavours"</font></a>, CBC News, June 1, 2007</li>
<li><a class="external text" href="http://www.npr.org/templates/story/story.php?storyId=15819485" rel="nofollow"><font color="#3366bb">"Sweet, Sour, Salty, Bitter… and Umami"</font></a> NPR, November 1, 2007</li>
<li><a class="external text" href="http://online.wsj.com/article/SB119706514515417586.html" rel="nofollow"><font color="#3366bb">A New Taste Sensation</font></a>, by Katy McLaughlin, Wall Street Journal, 12/8/07.</li>
<li><a class="external text" href="http://query.nytimes.com/gst/fullpage.html?res=9C03E7D91F3CF936A2575BC0A9629C8B63&sec=&spon=&pagewanted=all" rel="nofollow"><font color="#3366bb">Mouthfuls</font></a> : <a class="mw-redirect" title="Huitlacoche" href="/wiki/Huitlacoche"><font color="#0645ad">huitlacoche</font></a> is another <i>beispiel</i>, 24/7/08.</li>
</ul>