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Sweetness

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856: 282:. The high sweetness detection threshold and low bitterness detection threshold would have predisposed our primate ancestors to seek out sweet-tasting (and energy-dense) foods and avoid bitter-tasting foods. Even amongst leaf-eating primates, there is a tendency to prefer immature leaves, which tend to be higher in protein and lower in fibre and poisons than mature leaves. The "sweet tooth" thus has an ancient heritage, and while food processing has changed consumption patterns, human physiology remains largely unchanged. Biologically, a variant in 151: 33: 1110:. Early organic chemists tasted many of their products, either intentionally (as a means of characterization) or accidentally (due to poor laboratory hygiene). One of the first attempts to draw systematic correlations between molecules' structures and their tastes was made by a German chemist, Georg Cohn, in 1914. He hypothesized that to evoke a certain taste, a molecule must contain some structural motif (called a 3191: 1092: 1082:
of food can affect sweetness perception. Adding more red color to a drink increases its perceived sweetness. In a study darker colored solutions were rated 2–10% higher than lighter ones despite having 1% less sucrose concentration. The effect of color is believed to be due to cognitive expectations.
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Human studies have shown that sweet taste receptors are not only found in the tongue, but also in the lining of the gastrointestinal tract as well as the nasal epithelium, pancreatic islet cells, sperm and testes. It is proposed that the presence of sweet taste receptors in the GI tract controls the
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in 1991. This theory involves a total of eight interaction sites between a sweetener and the sweetness receptor, although not all sweeteners interact with all eight sites. This model has successfully directed efforts aimed at finding highly potent sweeteners, including the most potent family of
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Philippaert, Koenraad; Pironet, Andy; Mesuere, Margot; Sones, William; Vermeiren, Laura; Kerselaers, Sara; Pinto, Sílvia; Segal, Andrei; Antoine, Nancy; Gysemans, Conny; Laureys, Jos; Lemaire, Katleen; Gilon, Patrick; Cuypers, Eva; Tytgat, Jan; Mathieu, Chantal; Schuit, Frans; Rorsman, Patrik;
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Some variation in values is not uncommon between various studies. Such variations may arise from a range of methodological variables, from sampling to analysis and interpretation. Indeed, the taste index of 1, assigned to reference substances such as sucrose (for sweetness),
935: 1035:, the taste cells for sweet, bitter and umami share the same intracellular signalling pathway. Incoming sweet molecules bind to their receptors, which causes a conformational change in the molecule. This change activates the G-protein, gustducin, which in turn activates 769:(for sourness), quinine (for bitterness), and sodium chloride (for saltiness), is itself arbitrary for practical purposes. Some values, such as those for maltose and glucose, vary little. Others, such as aspartame and sodium saccharin, have much larger variation. 926:, cause sour foods to taste sweet. Once the tongue has been exposed to either of these proteins, sourness is perceived as sweetness for up to an hour afterwards. While curculin has some innate sweet taste of its own, miraculin is by itself quite tasteless. 2428:
Taruno, A; Vingtdeux, V; Ohmoto, M; Ma, Z; Dvoryanchikov, G; Li, A; Adrien, L; Zhao, H; Leung, S; Abernethy, M; Koppel, J; Davies, P; Civan, MM; Chaudhari, N; Matsumoto, I; Hellekant, G; Tordoff, MG; Marambaud, P; Foskett, JK (14 March 2013).
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DuBois, Grant E.; Walters, D. Eric; Schiffman, Susan S.; Warwick, Zoe S.; Booth, Barbara J.; Pecore, Suzanne D.; Gibes, Kernon; Carr, B. Thomas; Brands, Linda M. (1991-12-31), Walters, D. Eric; Orthoefer, Frank T.; DuBois, Grant E. (eds.),
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In 1919, Oertly and Myers proposed a more elaborate theory based on a then-current theory of color in synthetic dyes. They hypothesized that to be sweet, a compound must contain one each of two classes of structural motif, a
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and sweetness. This theory formalized these observations by proposing that to be sweet, a compound must have a third binding site (labeled X) that could interact with a hydrophobic site on the sweetness receptor via
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A few substances alter the way sweet taste is perceived. One class of these inhibits the perception of sweet tastes, whether from sugars or from highly potent sweeteners. Commercially, the most important of these is
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appears to have the lowest detection threshold, at about 1 part in 2 million for quinine in solution. In the natural settings that human primate ancestors evolved in, sweetness intensity should indicate
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Max M, Shanker YG, Huang LQ, Rong M, Liu Z, Campagne F, Weinstein H, Damak S, Margolskee RF (2001). "Tas1r3, encoding a new candidate taste receptor, is allelic to the sweet responsiveness locus Sac".
316:(table sugar) is the prototypical example of a sweet substance. Sucrose in solution has a sweetness perception rating of 1, and other substances are rated relative to this. For example, another sugar, 1027:
To depolarize the cell, and ultimately generate a response, the body uses different cells in the taste bud that each express a receptor for the perception of sweet, sour, salty, bitter or
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like domestic cats cannot perceive sweetness at all. The ability to taste sweetness often atrophies genetically in species of carnivores who do not eat sweet foods like fruits, including
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basis for detecting sweetness, which varies between both individuals and species, has only begun to be understood since the late 20th century. One theoretical model of sweetness is the
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Another research has shown that the threshold of sweet taste perception is in direct correlation with the time of day. This is believed to be the consequence of oscillating
1164:. According to this theory, the AH-B unit of a sweetener binds with a corresponding AH-B unit on the biological sweetness receptor to produce the sensation of sweetness. 953:
The breakthrough for the present understanding of sweetness occurred in 2001, when experiments with laboratory mice showed that mice possessing different versions of the
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levels in blood that may impact the overall sweetness of food. Scientists hypothesize that this is an evolutionary relict of diurnal animals like humans.
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Tinti, Jean-Marie; Nofre, Claude (1991). "Why does a sweetener taste sweet? A new model". In Walters, D.E.; Orthoefer, F.T; DuBois, G.E. (eds.).
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Kitagawa M, Kusakabe Y, Miura H, Ninomiya Y, Hino A (2001). "Molecular genetic identification of a candidate receptor gene for sweet taste".
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Hundreds of synthetic organic compounds are known to be sweet, but only a few of these are legally permitted as food additives. For example,
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O'Brien-Nabors, Lyn, ed. (2012). Alternative sweeteners (4th ed.). Boca Raton: CRC Press. ISBN 978-1-4398-4614-8. Retrieved 25 June 2014.
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Despite the wide variety of chemical substances known to be sweet, and knowledge that the ability to perceive sweet taste must reside in
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most commonly perceived when eating foods rich in sugars. Sweet tastes are generally regarded as pleasurable. In addition to sugars like
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Yamashita H, Akabane T, Kurihara Y (April 1995). "Activity and stability of a new sweet protein with taste-modifying action, curculin".
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Schiffman, S.S.; Diaz, C; Beeker, T.G (March 1986). "Caffeine Intensifies Taste of Certain Sweeteners: Role of Adenosine Receptor".
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From these beginnings in the early 20th century, the theory of sweetness enjoyed little further academic attention until 1963, when
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Sweetness perception may differ between species significantly. For example, even amongst the primates sweetness is quite variable.
3207:"Opioid hedonic hotspot in nucleus accumbens shell: mu, delta, and kappa maps for enhancement of sweetness "liking" and "wanting"" 1882:
Kinghorn, A.D. and Compadre, C.M. Alternative Sweeteners: Third Edition, Revised and Expanded, Marcel Dekker ed., New York, 2001.
1138:. Based on those compounds known to be sweet at the time, they proposed a list of six candidate glucophores and nine auxoglucs. 2572:
Shankar MU, Levitan CA, Spence C (2010). "Grape expectations: the role of cognitive influences in color-flavor interactions".
1667: 261:. Newborn human infants also demonstrate preferences for high sugar concentrations and prefer solutions that are sweeter than 2710: 2246: 1832: 1727: 1681: 1603: 1171:
in 1972. While previous researchers had noted that among some groups of compounds, there seemed to be a correlation between
1050:-receptor and induces calcium release from the endoplasmic reticulum. This increase in intracellular calcium activates the 369:. It is roughly 250 times sweeter than sucrose. Another class of potent natural sweeteners are the sweet proteins such as 2998: 249:
Studies indicate that responsiveness to sugars and sweetness has very ancient evolutionary beginnings, being manifest as
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Ma, Z; Siebert, AP; Cheung, KH; Lee, RJ; Johnson, B; Cohen, AS; Vingtdeux, V; Marambaud, P; Foskett, JK (10 July 2012).
2324:"Steviol glycosides enhance pancreatic beta-cell function and taste sensation by potentiation of TRPM5 channel activity" 2146:
Nakamura, Y.; Sanematsu, K.; Ohta, R.; Shirosaki, S.; Koyano, K.; Nonaka, K.; Shigemura, N.; Ninomiya, Y. (2008-07-15).
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Montmayeur JP, Liberles SD, Matsunami H, Buck LB (2001). "A candidate taste receptor gene near a sweet taste locus".
1887: 1773: 1575: 116: 312:, are sweet. Among common biological substances, all of the simple carbohydrates are sweet to at least some degree. 3313: 2381:"Intracellular Ca(2+) and TRPM5-mediated membrane depolarization produce ATP secretion from taste receptor cells" 76: 3250:"Hedonic hot spot in nucleus accumbens shell: where do mu-opioids cause increased hedonic impact of sweetness?" 1899:
Kurihara Y (1992). "Characteristics of antisweet substances, sweet proteins, and sweetness-inducing proteins".
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proposed the AH-B theory of sweetness. Simply put, they proposed that to be sweet, a compound must contain a
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The sweetness of 5% solution of glycine in water compares to a solution of 5.6% glucose or 2.6% fructose.
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The most elaborate theory of sweetness to date is the multipoint attachment theory (MPA) proposed by
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atoms are often sweet, and that among a series of structurally similar compounds, those with smaller
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Johnson, J.; Clydesdale, F. (1982). "Perceived sweetness and redness in colored sucrose solutions".
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gets activated by the depolarization and releases ATP neurotransmitter which activates the afferent
17: 3177:. ACS Symposium Series. Vol. 450. Washington DC: American Chemical Society. pp. 209–213. 1114:) that produces that taste. With regard to sweetness, he noted that molecules containing multiple 1177: 43: 1592: 1555: 1390:
With Bitter Herbs They Shall Eat It: Chemical ecology and the origins of human diet and medicine
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that are sweet at concentrations much lower than common sugars. The most well-known example is
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root, which is about 30 times sweeter than sucrose. Another commercially important example is
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Couper RTL.; Fernandez, P. L.; Alonso, P. L. (2006). "The Severe Gout of Emperor Charles V".
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Desor, J.A.; Maller, O.; Turner, R.E. (1973). "Taste acceptance of sugars by human infants".
1228:"A common genetic influence on human intensity ratings of sugars and high-potency sweeteners" 2431:"CALHM1 ion channel mediates purinergic neurotransmission of sweet, bitter and umami tastes" 1711: 1106:
in the 19th century introduced many new chemical compounds and the means to determine their
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Two natural products have been documented to have similar sweetness-inhibiting properties:
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Nofre C, Tinti JM (1996). "Sweetness reception in man: the multipoint attachment theory".
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Altman, S. (1989). "The monkey and the fig: A Socratic dialogue on evolutionary themes".
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Stevenson RJ, Oaten MJ (2010). "Sweet odours and sweet tastes are conflated in memory".
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Schiffman, Susan S. (2 June 1983). "Taste and smell in disease (Second of two parts)".
1004: 777: 773: 90: 3063: 3028: 2862: 2845: 2204: 2056: 2021: 1510: 320:, is somewhat sweeter, being rated at 1.7 times the sweetness of sucrose. Some of the 3477: 3279: 3236: 3152:
Shallenberger RS (1963). "Hydrogen bonding and the varying sweetness of the sugars".
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Schiffman, Susan S (26 May 1983). "Taste and smell in disease (First of two parts)".
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The Body Almanac: Mind-boggling facts about today's human body and high-tech medicine
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Opioids, sweets and a mechanism for positive affect: Broad motivational implications.
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T1R3 prefer sweet foods to different extents. Subsequent research has shown that the
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Some odors smell sweet and memory confuses whether sweetness was tasted or smelled.
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Fischler, C. (1980). "Food habits, social change and the nature/culture dilemma".
269:, being detectable at around 1 part in 200 of sucrose in solution. By comparison, 2628: 2585: 2396: 912: 816: 718: 336:
are the sweetest. Some other amino acids are perceived as both sweet and bitter.
3026: 3011: 2955:"Identification of a novel member of the T1R family of putative taste receptors" 1332: 3487: 3451: 3441: 3299: 2927: 2492:
Proceedings of the National Academy of Sciences of the United States of America
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Kurihara K, Beidler LM (1968). "Taste-Modifying Protein from Miracle Fruit".
1707: 1644: 1150: 1012: 886: 547: 359: 239: 193: 3053: 2512: 1827:. Springer-Lehrbuch (6., vollst. überarb. Aufl ed.). Berlin: Springer. 3283: 3240: 3144: 3029:"Amiloride reduces the taste intensity of Na+ and Li+ salts and sweeteners" 2980: 2945: 2871: 2828: 2799: 2770: 2762: 2733: 2636: 2593: 2531: 2472: 2414: 2365: 2307: 2189: 2124: 2065: 2046: 1869: 1473:
Attitudes towards sugar and sweetness in historical and social perspective.
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Nelson G, Hoon MA, Chandrashekar J, Zhang YF, Ryba NJP, Zuker CS (2001).
1168: 801: 472: 389: 374: 3338: 2454: 2347: 1861: 1652: 1556:"Concentration—Response Relationships of Sweeteners: A Systematic Study" 1244: 1227: 270: 265:, the sugar found in breast milk. Sweetness appears to have the highest 3461: 3436: 3415: 3374: 1676:(4th ed.). Belmont, California: Thomson Brooks/Cole. p. 359. 1631:
McLaughlin, Susan; Margolskee, Robert F. (1994). "The Sense of Taste".
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The perceived intensity of sugars and high-potency sweeteners, such as
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Schiffman, Susan S.; Lockhead, Elaine; Maes, Frans W (October 1983).
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Talavera, Karel; Voets, Thomas; Vennekens, Rudi (31 March 2017).
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Belitz, Hans-Dieter; Schieberle, Peter; Grosch, Werner (2008).
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Joesten, Melvin D; Hogg, John L; Castellion, Mary E (2007).
1091: 139:"Sweet" and "Sweetening" redirect here. For other uses, see 3175:
Sweeteners: Discovery, Molecular Design, and Chemoreception
2910:"Olfactory and Gustatory Discrimination in Man and Animals" 954: 934: 797: 1847: 1718:(11th ed.). Philadelphia: Elsevier Saunders. p.  3119:"Synergism among Ternary Mixtures of Fourteen Sweeteners" 2952: 1612: 996: 3291: 3117:
Susan S. Schiffman; Elizabeth A. Sattely-Miller (2000).
1984: 1415: 2653:(2008). "Transdisciplinary Perspectives on Sweetness". 2427: 1706: 1822: 1702: 1700: 3186: 2720:
Kier L (1972). "A molecular theory of sweet taste".
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Biochemical and Biophysical Research Communications
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Journal of Comparative and Physiological Psychology
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Saunders. 224:, may alter perception of sweetness itself. 2907: 2691: 2022:"Human receptors for sweet and umami taste" 1743:Dermer, OC (1947). "The Science of Taste". 1619:Dermer, OC (1947). "The Science of Taste". 1530: 1528: 1422:Jones, S.; Martin, R.; Pilbeam, D. (1994). 1402: 1353: 969:that is the sweetness receptor in mammals. 3314: 3300: 2886: 2378: 1898: 1598:(4th ed.). Brooks/Cole. p. 468. 3273: 3230: 3134: 3062: 3052: 2970: 2935: 2861: 2521: 2511: 2462: 2404: 2355: 2297: 2272:Chaudhari, N; Roper, SD (9 August 2010). 2179: 2114: 2096: 2055: 2045: 1548: 1546: 1387: 1318: 1243: 1022: 938:Sweetness is perceived by the taste buds. 929: 117:Learn how and when to remove this message 1761: 1668:"Sweeteness Relative to Sucrose (table)" 1525: 1436: 1426:. Cambridge: Cambridge University Press. 1090: 933: 854: 149: 2700: 1583: 1476: 1271: 918:On the other hand, two plant proteins, 14: 3506: 3082:Pharmacology Biochemistry and Behavior 2719: 2244: 1742: 1618: 1543: 1488: 1392:. Tucson: University of Arizona Press. 1368: 1201:sweeteners. The most potent of these, 1086: 850: 3295: 2649: 2085:The Journal of Physiological Sciences 2078: 1407:The Psychology of Eating and Drinking 784:. The latter may have contributed to 278:, while bitterness tends to indicate 2379:Huang, YA; Roper, SD (1 July 2010). 1768:. Woodhead Publishing. p. 264. 1710:; Hall, John; Hall, John E. (2006). 1039:to generate inositol trisphosphate ( 55:adding citations to reliable sources 26: 2999:The New England Journal of Medicine 2253:from the original on March 19, 2011 1321:The New England Journal of Medicine 889:, extracted from the leaves of the 286:increases craving for sweet foods. 24: 3182: 3166:10.1111/j.1365-2621.1963.tb00247.x 2722:Journal of Pharmaceutical Sciences 2559:10.1111/j.1365-2621.1982.tb12706.x 1673:The World of Chemistry: Essentials 342:A number of plant species produce 25: 3530: 2846:"Mammalian sweet taste receptors" 2245:Biello, David (August 16, 2007). 1511:10.1038/scientificamerican0893-86 1046:), this subsequently opens the IP 903:, from the leaves of the Chinese 3189: 2972:10.1046/j.1471-4159.2001.00292.x 999:(including most humans) all do. 792:aristocracy: the Roman delicacy 31: 3205:Castro DC, Berridge KC (2014). 2694:Die Organischen Geschmackstoffe 2643: 2608: 2565: 2538: 2479: 2421: 2372: 2314: 2265: 2238: 2196: 2139: 2072: 2013: 1978: 1927: 1841: 1825:Lehrbuch der Lebensmittelchemie 1816: 1791: 1782: 1765:Optimising Sweet Taste in Foods 1755: 1736: 1482: 1465: 1430: 1098:is the sweetest chemical known. 973:feeling of hunger and satiety. 796:was prepared by boiling soured 396:Sweetness of various compounds 243: 42:needs additional citations for 3266:10.1523/JNEUROSCI.2329-05.2005 3248:Peciña S, Berridge KC (2005). 3223:10.1523/JNEUROSCI.4458-13.2014 1537:Textbook of Medical Physiology 1396: 1381: 1362: 1347: 1312: 1277: 1260: 1219: 1197:sweeteners known to date, the 13: 1: 2863:10.1016/S0092-8674(01)00451-2 2079:Kohno, Daisuke (2017-04-04). 1956:10.1126/science.161.3847.1241 1208: 1183: 1054:channel and induces cellular 233:neohesperidin dihydrochalcone 3094:10.1016/0091-3057(86)90536-8 3033:Proc. 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(1991). 1451:10.1177/053901848001900603 1439:Social Science Information 967:G-protein coupled receptor 293: 154:Sweet foods, such as this 138: 134:Sweetness (disambiguation) 131: 3470: 3429: 3398: 3367: 3329: 3321: 2959:Journal of Neurochemistry 2667:10.1007/s12078-007-9003-z 2385:The Journal of Physiology 2098:10.1007/s12576-017-0535-y 1913:10.1080/10408399209527598 1411:. New York: W.H. Freeman. 878:, a compound produced by 350:, the sweet component of 2696:. Berlin: F. Siemenroth. 1805:. 2011-04-13. p. 16 1213: 1178:London dispersion forces 3154:Journal of Food Science 3136:10.1093/chemse/25.2.131 3054:10.1073/pnas.80.19.6136 2655:Chemosensory Perception 2547:Journal of Food Science 2513:10.1073/pnas.1204023109 2249:. Scientific American. 2224:10.1093/chemse/20.5.573 1999:10.1093/chemse/20.2.239 2763:10.1006/bbrc.2001.4760 2734:10.1002/jps.2600610910 2047:10.1073/pnas.072090199 1901:Crit Rev Food Sci Nutr 1358:. New York: Doubleday. 1099: 1023:Sweet receptor pathway 939: 930:The sweetness receptor 870: 166: 161:, are often eaten for 141:Sweet (disambiguation) 2328:Nature Communications 2290:10.1083/jcb.201003144 1590:John McMurry (1998). 1094: 937: 858: 776:are sweet, including 654:Modified disaccharide 626:Oxathiazinone dioxide 300:A great diversity of 294:Further information: 184:are sweet, including 153: 3519:Culinary terminology 2692:Cohn, Georg (1914). 1403:Logue, A.W. (1986). 1354:McAleer, N. (1985). 1143:Robert Shallenberger 1108:molecular structures 1031:. Downstream of the 833:acesulfame potassium 758:230,000 (estimated) 743:200,000 (estimated) 728:200,000 (estimated) 713:188,000 (estimated) 698:162,000 (estimated) 622:Acesulfame potassium 202:non-sugar sweeteners 132:For other uses, see 51:improve this article 3045:1983PNAS...80.6136S 2809:Nature Neuroscience 2617:Acta Psychol (Amst) 2504:2012PNAS..109E1963M 2455:10.1038/nature11906 2447:2013Natur.495..223T 2348:10.1038/ncomms14733 2340:2017NatCo...814733P 2038:2002PNAS...99.4692L 1948:1968Sci...161.1241K 1862:10.1056/NEJMc062352 1503:1993SciAm.269b..86M 1491:Scientific American 1274:, pp. 115–124) 1245:10.1017/thg.2015.42 1102:The development of 1087:Historical theories 1005:bottlenose dolphins 866:, 1853 painting by 851:Sweetness modifiers 847:are commonly used. 774:inorganic compounds 397: 2391:(Pt 13): 2343–50. 1633:American Scientist 1388:Johns, T. (1990). 1371:American Scientist 1232:Twin Res Hum Genet 1100: 940: 871: 778:beryllium chloride 395: 324:are mildly sweet: 302:chemical compounds 182:chemical compounds 167: 3501: 3500: 2712:978-0-387-17045-9 2164:10.2337/db07-1103 2158:(10): 2661–2665. 1834:978-3-540-73201-3 1729:978-0-7216-0240-0 1683:978-0-495-01213-9 1605:978-0-13-286261-5 1594:Organic Chemistry 1479:, pp. 83–98) 1124:molecular weights 1104:organic chemistry 993:Old World monkeys 985:New World monkeys 896:Gymnema sylvestre 868:George Henry Hall 767:hydrochloric acid 762: 761: 680:Aspartame analog 586:Steviol glycoside 404:Type of compound 392:, is also sweet. 388:protein found in 257:bacteria such as 198:sugar substitutes 127: 126: 119: 101: 16:(Redirected from 3526: 3316: 3309: 3302: 3293: 3292: 3287: 3277: 3260:(50): 11777–86. 3244: 3234: 3199: 3194: 3193: 3178: 3169: 3148: 3138: 3113: 3076: 3066: 3056: 3039:(19): 6136–640. 3023: 2992: 2974: 2949: 2939: 2904: 2883: 2865: 2840: 2803: 2774: 2745: 2728:(9): 1394–1397. 2716: 2697: 2679: 2678: 2647: 2641: 2640: 2612: 2606: 2605: 2569: 2563: 2562: 2542: 2536: 2535: 2525: 2515: 2498:(28): E1963–71. 2483: 2477: 2476: 2466: 2425: 2419: 2418: 2408: 2376: 2370: 2369: 2359: 2318: 2312: 2311: 2301: 2269: 2263: 2262: 2260: 2258: 2242: 2236: 2235: 2209: 2200: 2194: 2193: 2183: 2143: 2137: 2136: 2118: 2100: 2076: 2070: 2069: 2059: 2049: 2017: 2011: 2010: 1982: 1976: 1975: 1942:(3847): 1241–3. 1931: 1925: 1924: 1896: 1890: 1880: 1874: 1873: 1845: 1839: 1838: 1820: 1814: 1813: 1811: 1810: 1795: 1789: 1786: 1780: 1779: 1759: 1753: 1752: 1740: 1734: 1733: 1717: 1708:Guyton, Arthur C 1704: 1695: 1694: 1692: 1690: 1663: 1657: 1656: 1628: 1616: 1610: 1609: 1597: 1587: 1581: 1580: 1550: 1541: 1540: 1532: 1523: 1522: 1486: 1480: 1469: 1463: 1462: 1434: 1428: 1427: 1419: 1413: 1412: 1410: 1400: 1394: 1393: 1385: 1379: 1378: 1366: 1360: 1359: 1351: 1345: 1344: 1316: 1310: 1309: 1298:10.1037/h0034906 1281: 1275: 1264: 1258: 1257: 1247: 1223: 1190:Jean-Marie Tinti 1062:release channel 804:) in lead pots. 782:lead(II) acetate 635:Sodium saccharin 571:Sodium cyclamate 443:(α,α-trehalose) 398: 394: 296:Sugar substitute 122: 115: 111: 108: 102: 100: 59: 35: 27: 21: 3534: 3533: 3529: 3528: 3527: 3525: 3524: 3523: 3504: 3503: 3502: 3497: 3466: 3425: 3394: 3363: 3325: 3320: 3290: 3217:(12): 4239–50. 3195: 3188: 3185: 3183:Further reading 3123:Chemical Senses 2780:Nature Genetics 2713: 2687: 2682: 2648: 2644: 2613: 2609: 2574:Conscious. Cogn 2570: 2566: 2543: 2539: 2484: 2480: 2441:(7440): 223–6. 2426: 2422: 2377: 2373: 2319: 2315: 2270: 2266: 2256: 2254: 2243: 2239: 2212:Chemical Senses 2207: 2201: 2197: 2144: 2140: 2077: 2073: 2018: 2014: 1983: 1979: 1932: 1928: 1897: 1893: 1881: 1877: 1856:(18): 1935–36. 1846: 1842: 1835: 1821: 1817: 1808: 1806: 1797: 1796: 1792: 1787: 1783: 1776: 1760: 1756: 1741: 1737: 1730: 1705: 1698: 1688: 1686: 1684: 1664: 1660: 1617: 1613: 1606: 1588: 1584: 1578: 1551: 1544: 1533: 1526: 1487: 1483: 1470: 1466: 1435: 1431: 1420: 1416: 1401: 1397: 1386: 1382: 1367: 1363: 1352: 1348: 1327:(22): 1337–43. 1317: 1313: 1282: 1278: 1265: 1261: 1224: 1220: 1216: 1211: 1186: 1089: 1076: 1049: 1044: 1037:phospholipase C 1025: 932: 913:herbal medicine 909:Ziziphus jujuba 860:Boys Pilfering 853: 817:ethylene glycol 719:Sucrononic acid 380:fruit. Hen egg 373:, found in the 298: 292: 148: 137: 130: 123: 112: 106: 103: 60: 58: 48: 36: 23: 22: 15: 12: 11: 5: 3532: 3522: 3521: 3516: 3499: 3498: 3496: 3495: 3490: 3485: 3480: 3474: 3472: 3468: 3467: 3465: 3464: 3459: 3454: 3452:Scoville scale 3449: 3444: 3442:Acquired taste 3439: 3433: 3431: 3427: 3426: 3424: 3423: 3418: 3413: 3408: 3402: 3400: 3396: 3395: 3393: 3392: 3387: 3382: 3377: 3371: 3369: 3365: 3364: 3362: 3361: 3356: 3351: 3346: 3341: 3335: 3333: 3327: 3326: 3319: 3318: 3311: 3304: 3296: 3289: 3288: 3245: 3201: 3200: 3184: 3181: 3180: 3179: 3170: 3149: 3129:(2): 131–140. 3114: 3088:(3): 429–432. 3077: 3024: 3006:(21): 1275–9. 2993: 2965:(3): 896–903. 2950: 2905: 2895:(3): 263–274. 2889:Food Chemistry 2884: 2856:(3): 381–390. 2841: 2804: 2775: 2757:(1): 236–242. 2746: 2717: 2711: 2698: 2688: 2686: 2683: 2681: 2680: 2651:Hayes, John E. 2642: 2607: 2564: 2553:(3): 747–752. 2537: 2478: 2420: 2371: 2313: 2264: 2237: 2218:(5): 573–584. 2195: 2138: 2091:(4): 459–465. 2071: 2012: 1977: 1926: 1891: 1875: 1840: 1833: 1815: 1790: 1781: 1774: 1754: 1735: 1728: 1696: 1682: 1658: 1639:(6): 538–545. 1611: 1604: 1582: 1576: 1542: 1524: 1481: 1464: 1445:(6): 937–953. 1429: 1414: 1395: 1380: 1361: 1346: 1311: 1292:(3): 496–501. 1276: 1259: 1217: 1215: 1212: 1210: 1207: 1185: 1182: 1173:hydrophobicity 1088: 1085: 1075: 1072: 1056:depolarization 1047: 1042: 1033:taste receptor 1024: 1021: 1013:spotted hyenas 931: 928: 852: 849: 786:lead poisoning 760: 759: 756: 751: 745: 744: 741: 736: 730: 729: 726: 721: 715: 714: 711: 706: 700: 699: 696: 691: 685: 684: 681: 678: 672: 671: 668: 665: 659: 658: 655: 652: 646: 645: 642: 637: 631: 630: 627: 624: 618: 617: 614: 603: 597: 596: 593: 588: 582: 581: 578: 573: 567: 566: 563: 562:Monosaccharide 560: 554: 553: 550: 545: 539: 538: 529: 526: 520: 519: 516: 513: 507: 506: 503: 502:Monosaccharide 500: 494: 493: 490: 488:Monosaccharide 485: 479: 478: 475: 470: 464: 463: 460: 457: 451: 450: 447: 444: 437: 436: 433: 430: 424: 423: 420: 415: 409: 408: 405: 402: 360:South American 291: 288: 276:energy density 194:sugar alcohols 128: 125: 124: 39: 37: 30: 9: 6: 4: 3: 2: 3531: 3520: 3517: 3515: 3512: 3511: 3509: 3494: 3491: 3489: 3486: 3484: 3481: 3479: 3476: 3475: 3473: 3469: 3463: 3460: 3458: 3455: 3453: 3450: 3448: 3445: 3443: 3440: 3438: 3435: 3434: 3432: 3428: 3422: 3419: 3417: 3414: 3412: 3409: 3407: 3404: 3403: 3401: 3397: 3391: 3388: 3386: 3383: 3381: 3378: 3376: 3373: 3372: 3370: 3366: 3360: 3357: 3355: 3352: 3350: 3347: 3345: 3342: 3340: 3337: 3336: 3334: 3332: 3328: 3324: 3317: 3312: 3310: 3305: 3303: 3298: 3297: 3294: 3285: 3281: 3276: 3271: 3267: 3263: 3259: 3255: 3251: 3246: 3242: 3238: 3233: 3228: 3224: 3220: 3216: 3212: 3208: 3203: 3202: 3198: 3192: 3187: 3176: 3171: 3167: 3163: 3159: 3155: 3150: 3146: 3142: 3137: 3132: 3128: 3124: 3120: 3115: 3111: 3107: 3103: 3099: 3095: 3091: 3087: 3083: 3078: 3074: 3070: 3065: 3060: 3055: 3050: 3046: 3042: 3038: 3034: 3030: 3025: 3021: 3017: 3013: 3009: 3005: 3001: 3000: 2994: 2990: 2986: 2982: 2978: 2973: 2968: 2964: 2960: 2956: 2951: 2947: 2943: 2938: 2933: 2929: 2925: 2921: 2917: 2916: 2911: 2906: 2902: 2898: 2894: 2890: 2885: 2881: 2877: 2873: 2869: 2864: 2859: 2855: 2851: 2847: 2842: 2838: 2834: 2830: 2826: 2822: 2821:10.1038/87440 2818: 2814: 2810: 2805: 2801: 2797: 2793: 2792:10.1038/88270 2789: 2785: 2781: 2776: 2772: 2768: 2764: 2760: 2756: 2752: 2747: 2743: 2739: 2735: 2731: 2727: 2723: 2718: 2714: 2708: 2704: 2699: 2695: 2690: 2689: 2676: 2672: 2668: 2664: 2660: 2656: 2652: 2646: 2638: 2634: 2630: 2626: 2622: 2618: 2611: 2603: 2599: 2595: 2591: 2587: 2583: 2580:(1): 380–90. 2579: 2575: 2568: 2560: 2556: 2552: 2548: 2541: 2533: 2529: 2524: 2519: 2514: 2509: 2505: 2501: 2497: 2493: 2489: 2482: 2474: 2470: 2465: 2460: 2456: 2452: 2448: 2444: 2440: 2436: 2432: 2424: 2416: 2412: 2407: 2402: 2398: 2394: 2390: 2386: 2382: 2375: 2367: 2363: 2358: 2353: 2349: 2345: 2341: 2337: 2333: 2329: 2325: 2317: 2309: 2305: 2300: 2295: 2291: 2287: 2284:(3): 285–96. 2283: 2279: 2275: 2268: 2252: 2248: 2241: 2233: 2229: 2225: 2221: 2217: 2213: 2206: 2199: 2191: 2187: 2182: 2177: 2173: 2169: 2165: 2161: 2157: 2153: 2149: 2142: 2134: 2130: 2126: 2122: 2117: 2112: 2108: 2104: 2099: 2094: 2090: 2086: 2082: 2075: 2067: 2063: 2058: 2053: 2048: 2043: 2039: 2035: 2032:(7): 4692–6. 2031: 2027: 2023: 2016: 2008: 2004: 2000: 1996: 1993:(2): 239–43. 1992: 1988: 1981: 1973: 1969: 1965: 1961: 1957: 1953: 1949: 1945: 1941: 1937: 1930: 1922: 1918: 1914: 1910: 1907:(3): 231–52. 1906: 1902: 1895: 1889: 1888:0-8247-0437-1 1885: 1879: 1871: 1867: 1863: 1859: 1855: 1851: 1844: 1836: 1830: 1826: 1819: 1804: 1800: 1794: 1785: 1777: 1775:9781845691646 1771: 1767: 1766: 1758: 1750: 1746: 1739: 1731: 1725: 1721: 1716: 1715: 1709: 1703: 1701: 1685: 1679: 1675: 1674: 1669: 1662: 1654: 1650: 1646: 1642: 1638: 1634: 1626: 1622: 1615: 1607: 1601: 1596: 1595: 1586: 1579: 1577:9780841219038 1573: 1569: 1565: 1561: 1557: 1549: 1547: 1538: 1531: 1529: 1520: 1516: 1512: 1508: 1504: 1500: 1496: 1492: 1485: 1478: 1474: 1471:Fischler, C. 1468: 1460: 1456: 1452: 1448: 1444: 1440: 1433: 1425: 1418: 1409: 1408: 1399: 1391: 1384: 1376: 1372: 1365: 1357: 1350: 1342: 1338: 1334: 1330: 1326: 1322: 1315: 1307: 1303: 1299: 1295: 1291: 1287: 1280: 1273: 1269: 1263: 1255: 1251: 1246: 1241: 1237: 1233: 1229: 1222: 1218: 1206: 1204: 1200: 1195: 1191: 1181: 1179: 1174: 1170: 1165: 1163: 1159: 1155: 1152: 1151:hydrogen bond 1148: 1144: 1139: 1137: 1133: 1127: 1125: 1121: 1117: 1113: 1109: 1105: 1097: 1093: 1084: 1081: 1071: 1069: 1065: 1061: 1057: 1053: 1045: 1038: 1034: 1030: 1020: 1018: 1014: 1010: 1006: 1002: 998: 994: 991:sweet, while 990: 986: 981: 979: 974: 970: 968: 964: 960: 956: 951: 949: 945: 936: 927: 925: 921: 916: 914: 910: 906: 902: 898: 897: 892: 888: 887:gymnemic acid 883: 881: 877: 869: 865: 863: 857: 848: 846: 842: 838: 834: 830: 826: 822: 818: 814: 810: 805: 803: 799: 795: 791: 790:ancient Roman 787: 783: 779: 775: 770: 768: 757: 755: 752: 750: 747: 746: 742: 740: 737: 735: 732: 731: 727: 725: 722: 720: 717: 716: 712: 710: 707: 705: 702: 701: 697: 695: 692: 690: 687: 686: 682: 679: 677: 674: 673: 669: 666: 664: 661: 660: 656: 653: 651: 648: 647: 643: 641: 638: 636: 633: 632: 628: 625: 623: 620: 619: 615: 613: 610: 607: 604: 602: 599: 598: 594: 592: 589: 587: 584: 583: 579: 577: 574: 572: 569: 568: 564: 561: 559: 556: 555: 551: 549: 548:sugar alcohol 546: 544: 541: 540: 536: 535: 530: 527: 525: 522: 521: 517: 514: 512: 509: 508: 504: 501: 499: 496: 495: 491: 489: 486: 484: 481: 480: 476: 474: 471: 469: 466: 465: 461: 459:Disaccharide 458: 456: 453: 452: 448: 446:Disaccharide 445: 442: 439: 438: 434: 431: 429: 426: 425: 421: 419: 416: 414: 411: 410: 406: 403: 400: 399: 393: 391: 387: 383: 379: 376: 372: 368: 366: 361: 357: 353: 349: 345: 340: 337: 335: 331: 327: 323: 319: 315: 311: 307: 303: 297: 287: 285: 281: 277: 272: 268: 264: 260: 256: 252: 247: 245: 241: 236: 234: 230: 225: 223: 219: 215: 211: 207: 203: 199: 195: 191: 187: 183: 180:, many other 179: 175: 171: 164: 160: 157: 152: 146: 142: 135: 121: 118: 110: 99: 96: 92: 89: 85: 82: 78: 75: 71: 68: –  67: 63: 62:Find sources: 56: 52: 46: 45: 40:This article 38: 34: 29: 28: 19: 3353: 3331:Basic tastes 3257: 3253: 3214: 3210: 3174: 3160:(5): 584–9. 3157: 3153: 3126: 3122: 3085: 3081: 3036: 3032: 3003: 2997: 2962: 2958: 2922:(1): 47–51. 2919: 2913: 2892: 2888: 2853: 2849: 2815:(5): 492–8. 2812: 2808: 2786:(1): 58–63. 2783: 2779: 2754: 2750: 2725: 2721: 2702: 2693: 2661:(1): 48–57. 2658: 2654: 2645: 2623:(1): 105–9. 2620: 2616: 2610: 2577: 2573: 2567: 2550: 2546: 2540: 2495: 2491: 2481: 2438: 2434: 2423: 2388: 2384: 2374: 2331: 2327: 2316: 2281: 2277: 2267: 2255:. Retrieved 2240: 2215: 2211: 2198: 2155: 2151: 2141: 2088: 2084: 2074: 2029: 2025: 2015: 1990: 1987:Chem. Senses 1986: 1980: 1939: 1935: 1929: 1904: 1900: 1894: 1878: 1853: 1850:N Engl J Med 1849: 1843: 1824: 1818: 1807:. Retrieved 1802: 1793: 1784: 1764: 1757: 1748: 1744: 1738: 1713: 1689:14 September 1687:. Retrieved 1672: 1661: 1636: 1632: 1624: 1620: 1614: 1593: 1585: 1559: 1536: 1497:(2): 70–77. 1494: 1490: 1484: 1477:Dobbing 1987 1472: 1467: 1442: 1438: 1432: 1423: 1417: 1406: 1398: 1389: 1383: 1374: 1370: 1364: 1355: 1349: 1324: 1320: 1314: 1289: 1285: 1279: 1272:Dobbing 1987 1267: 1266:Blass, E.M. 1262: 1238:(4): 361–7. 1235: 1231: 1221: 1194:Claude Nofre 1187: 1166: 1140: 1135: 1131: 1128: 1111: 1101: 1077: 1026: 987:do not find 982: 975: 971: 965:, to form a 959:T1R3 protein 952: 941: 917: 908: 894: 884: 880:Domino Sugar 872: 859: 813:nitrobenzene 806: 800:(containing 793: 771: 763: 565:1.17 – 1.75 533: 532: 528:Disaccharide 462:0.40 - 0.50 455:Isomaltulose 435:0.33 – 0.45 432:Disaccharide 418:Disaccharide 390:chicken eggs 375:West African 363: 348:glycyrrhizin 341: 338: 299: 258: 248: 240:chemosensory 237: 226: 169: 168: 113: 104: 94: 87: 80: 73: 61: 49:Please help 44:verification 41: 3457:Supertaster 3421:Hypergeusia 3385:Soft palate 3254:J. Neurosci 3211:J. Neurosci 3197:Food portal 1169:Lemont Kier 1156:(AH) and a 1147:Terry Acree 802:acetic acid 689:Sucrooctate 518:0.6 – 0.86 505:0.74 – 0.8 473:Polyalcohol 358:, from the 322:amino acids 174:basic taste 129:Basic taste 66:"Sweetness" 3508:Categories 3462:Tongue map 3437:Aftertaste 3416:Hypogeusia 3375:Epiglottis 3339:Bitterness 1809:2023-12-10 1560:Sweeteners 1377:: 256–263. 1209:References 1184:MPA theory 1162:nanometres 1158:Lewis base 1132:glucophore 944:taste buds 809:chloroform 788:among the 772:Even some 704:Bernardame 644:300 – 675 616:180 – 250 515:Amino acid 449:max. 0,45 407:Sweetness 386:antibiotic 367:rebaudiana 356:stevioside 344:glycosides 304:, such as 271:bitterness 251:chemotaxis 156:strawberry 107:April 2024 77:newspapers 3514:Gustation 3411:Dysgeusia 3399:Gustology 3354:Sweetness 3344:Saltiness 2703:Sweetness 2675:145694059 2334:: 14733. 2172:0012-1797 2107:1880-6546 1803:d-nb.info 1645:0003-0996 1459:143766021 1203:lugduname 1199:guanidine 1112:sapophore 1096:Lugduname 1074:Cognition 1009:sea lions 989:aspartame 920:miraculin 876:lactisole 837:sucralose 829:aspartame 825:cyclamate 821:Saccharin 754:Guanidine 749:Lugduname 739:Guanidine 734:Carrelame 724:Guanidine 709:Guanidine 694:Guanidine 663:Thaumatin 650:Sucralose 606:Dipeptide 601:Aspartame 595:40 – 300 591:Glycoside 576:Sulfonate 534:reference 483:Galactose 441:Trehalose 371:thaumatin 306:aldehydes 229:aspartame 222:miraculin 214:sucralose 210:aspartame 206:saccharin 186:aldehydes 170:Sweetness 159:shortcake 3471:See also 3447:Pungency 3349:Sourness 3284:16354936 3241:24647944 3145:10781019 3110:20419613 2989:11296598 2981:11331418 2946:13941509 2880:11886074 2872:11509186 2837:21010650 2829:11319557 2800:11326277 2771:11322794 2637:20097323 2602:32230245 2594:19828330 2532:22711817 2473:23467090 2415:20498227 2366:28361903 2308:20696704 2257:July 28, 2251:Archived 2190:18633111 2152:Diabetes 2125:28378265 2116:10717116 2066:11917125 1972:24451890 1870:17079773 1751:: 15–18. 1653:29775325 1627:: 15–18. 1254:26181574 1136:auxogluc 1120:chlorine 1116:hydroxyl 924:curculin 901:ziziphin 862:Molasses 640:Sulfonyl 558:Fructose 468:Sorbitol 382:lysozyme 352:licorice 318:fructose 280:toxicity 253:even in 204:include 18:Sweetest 3483:Hearing 3406:Ageusia 3380:Pharynx 3368:Anatomy 3275:6726018 3232:3960467 3102:3010333 3073:6577473 3041:Bibcode 3020:6341841 2937:1896974 2742:5068944 2685:General 2523:3396471 2500:Bibcode 2464:3600154 2443:Bibcode 2406:2915511 2357:5380970 2336:Bibcode 2299:2922655 2232:8564432 2181:2551675 2133:3984011 2034:Bibcode 2007:7583017 1964:5673432 1944:Bibcode 1936:Science 1921:1418601 1519:8351513 1499:Bibcode 1341:6341845 1306:4745817 1134:and an 1068:neurons 946:on the 845:neotame 841:alitame 676:Neotame 667:Protein 543:Xylitol 524:Sucrose 511:Glycine 498:Glucose 428:Maltose 413:Lactose 378:katemfe 330:glycine 326:alanine 314:Sucrose 310:ketones 263:lactose 259:E. coli 200:. Such 190:ketones 178:sucrose 163:dessert 91:scholar 3390:Tongue 3282:  3272:  3239:  3229:  3143:  3108:  3100:  3071:  3064:534376 3061:  3018:  2987:  2979:  2944:  2934:  2878:  2870:  2835:  2827:  2798:  2769:  2740:  2709:  2673:  2635:  2600:  2592:  2530:  2520:  2471:  2461:  2435:Nature 2413:  2403:  2364:  2354:  2306:  2296:  2230:  2188:  2178:  2170:  2131:  2123:  2113:  2105:  2064:  2057:123709 2054:  2005:  1970:  1962:  1919:  1886:  1868:  1831:  1772:  1726:  1680:  1651:  1643:  1602:  1574:  1517:  1457:  1339:  1304:  1252:  1064:CALHM1 1058:. 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