Sensing via intestinal sweet taste pathways

Frontiers in Neuroscience

Volumn , Issue MAR, 2011, Pages

  Young, Richard L ^a  

^a UNIVERSITY OF ADELAIDE   (Australia)

Author keywords

Carbohydrate absorption; Gastric emptying; Glucose sensing; Small intestine; Sweet taste molecules; Vagal afferents

Indexed keywords

4 AMINOBUTYRIC ACID; ADENOSINE TRIPHOSPHATE; ADENOSINE TRIPHOSPHATE SENSITIVE POTASSIUM CHANNEL; BETA ENDORPHIN; CHOLECYSTOKININ; CHROMOGRANIN A; CYTOKERATIN 18; DYNORPHIN; FATTY ACID; GASTROINTESTINAL POLYPEPTIDE; GLUCAGON; GLUCAGON LIKE PEPTIDE 1; GLUCAGON LIKE PEPTIDE 2; GLUCOSE; GLUCOSE TRANSPORTER 2; MELATONIN; METENKEPHALIN; NEURONAL NITRIC OXIDE SYNTHASE; NITRIC OXIDE; OXYNTOMODULIN; PROTEIN; PURINERGIC P2X2 RECEPTOR; PURINERGIC P2X3 RECEPTOR; PURINERGIC P2Y RECEPTOR; SEROTONIN; SODIUM GLUCOSE COTRANSPORTER 1; TRANSIENT RECEPTOR POTENTIAL CHANNEL M5; UNINDEXED DRUG; UROGUANYLIN; VOLTAGE GATED CALCIUM CHANNEL;

BRUSH BORDER; CALORIC INTAKE; CELL TYPE; COLON MUCOSA; DIABETES MELLITUS; DUODENUM; ELECTRON TRANSPORT; ENTEROCHROMAFFIN CELL; GASTROINTESTINAL MOTILITY; GASTROINTESTINAL TRACT; GLUCOSE BLOOD LEVEL; GLYCEMIC CONTROL; HUMAN; ILEUM; INTESTINE CELL; INTESTINE EPITHELIUM CELL; JEJUNUM; NONHUMAN; REVIEW; SENSOR; SIGNAL TRANSDUCTION; SMALL INTESTINE; STOMACH EMPTYING; SWEETNESS; TONGUE;

EID: 80155140920     PISSN: 16624548     EISSN: 1662453X     Source Type: Journal    
DOI: 10.3389/fnins.2011.00023     Document Type: Review

References (184)

1. Adler, E., Hoon, M. A., Mueller, K. L., Chandrashekar, J., Ryba, N. J., and Zuker, C. S. (2000). A novel family of mammalian taste receptors. Cell 100, 693-702.
2. Akkermans, L. M., Vos, A., Hoekstra, A., Roelofs, J. M., and Horowitz, M. (1988). Effect of ICS 205-930 (a specific 5-HT3 receptor antagonist) on gastric emptying of a solid meal in normal subjects. Gut 29, 1249-1252.
3. Azpiroz, F., and Malagelada, J. R. (1987). Gastric tone measured by an electronic barostat in health and postsurgical gastroparesis. Gastroenterology 92, 934-943.
4. Baggio, L. L., and Drucker, D. J. (2007). Biology of incretins: GLP-1 and GIP. Gastroenterology 132, 2131-2157.
5. Baryshnikov, S. G., Rogachevskaja, O. A., and Kolesnikov, S. S. (2003). Calcium signaling mediated by P2Y receptors in mouse taste cells. J. Neurophysiol. 90, 3283-3294.
6. Bastidas, J. A., Yeo, C. J., Schmieg, R. E. Jr., and Zinner, M. J. (1989). Endogenous opiates in the mediation of early meal-induced jejunal absorption of water and electrolytes. Am. J. Surg. 157, 27-32.
7. Bates, S. L., Sharkey, K. A., and Meddings, J. B. (1998). Vagal involvement in die tary regulation of nutrient transport. Am. J. Physiol. 274, G552-G560.
8. Berthoud, H. R., Kressel, M., Raybould, H. E., and Neuhuber, W. L. (1995). Vagal sensors in the rat duodenal mucosa: distribution and structure as revealed by in vivo DiI-tracing. Anat. Embryol. 191, 203-212.
9. Bertrand, P. P. (2009). The cornucopia of intestinal chemosensory transduction. Front. Neurosci. 3, 1-9. doi: 10.3389/ neuro.21.003.2009
10. Bertrand, P. P., and Bertrand, R. L. (2010). Serotonin release and uptake in the gastrointestinal tract. Auton. Neurosci. 153, 47-57.
11. Bezencon, C., Furholz, A., Raymond, F., Mansourian, R., Metairon, S., Le Coutre, J., and Damak, S. (2008). Murine intestinal cells expressing Trpm5 are mostly brush cells and express markers of neuronal and inflammatory cells. J. Comp. Neurol. 509, 514-525.
12. Bezencon, C., le Coutre, J., and Damak, S. (2007). Taste-signaling proteins are coexpressed in solitary intestinal epithelial cells. Chem. Senses 32, 41-49.
13. Blackshaw, L. A., Brookes, S. J., Grundy, D., and Schemann, M. (2007). Sensory transmission in the gastrointestinal tract. Neurogastroenterol. Motil. 19, 1-19.
14. Bo, X., Alavi, A., Xiang, Z., Oglesby, I., Ford, A., and Burnstock, G. (1999). Localization of ATP-gated P2X2 and P2X3 receptor immunoreactive nerves in rat taste buds. Neuroreport 10, 1107-1111.
15. Brener, W., Hendrix, T. R., and McHugh, P. R. (1983). Regulation of the gastric emptying of glucose. Gastroenterology 85, 76-82.
16. Brixel, L. R., Monteilh-Zoller, M. K., Ingenbrandt, C. S., Fleig, A., Penner, R., Enklaar, T., Zabel, B. U., and Prawitt, D. (2010). TRPM5 regulates glucose-stimulated insulin secretion. Pflugers Arch. 460, 69-76.
17. Broberger, C., and Hokfelt, T. (2001). Hypothalamic and vagal neuropeptide circuitries regulating food intake. Physiol. Behav. 74, 669-682.
18. Brown, R. J., Walter, M., and Rother, K. I. (2009). Ingestion of diet soda before a glucose load augments glucagon-like peptide-1 secretion. Diabetes Care 32, 2184-2186.
19. Bryant, M. G., Bloom, S. R., Polak, J. M., Hobbs, S., Domschke, W., Domschke, S., Mitznegg, P., Ruppin, H., and Demling, L. (1983). Measurement of gut hormonal peptides in biopsies from human stomach and proximal small intestine. Gut 24, 114-119.
20. Bucinskaite, V., Tolessa, T., Pedersen, J., Rydqvist, B., Zerihun, L., Holst, J. J., and Hellstrom, P. M. (2009). Receptor-mediated activation of gastric vagal afferents by glucagon-like peptide-1 in the rat. Neurogastroenterol. Motil. 21, 978-e978.
21. Cani, P. D., Knauf, C., Iglesias, M. A., Drucker, D. J., Delzenne, N. M., and Burcelin, R. (2006). Improvement of glucose tolerance and hepatic insulin sensitivity by oligofructose requires a functional glucagon-like peptide 1 receptor. Diabetes 55, 1484-1490.
22. Chandrashekar, J., Hoon, M. A., Ryba, N. J., and Zuker, C. S. (2006). The receptors and cells for mammalian taste. Nature 444, 288-294.
23. Chandrashekar, J., Mueller, K. L., Hoon, M. A., Adler, E., Feng, L., Guo, W., Zuker, C. S., and Ryba, N. J. (2000). T2Rs function as bitter taste receptors. Cell 100, 703-711.
24. Clapp, T. R., Trubey, K. R., Vandenbeuch, A., Stone, L. M., Margolskee, R. F., Chaudhari, N., and Kinnamon, S. C. (2008). Tonic activity of Galpha-gustducin regulates taste cell responsivity. FEBS Lett. 582, 3783-3787.
25. Clapp, T. R., Yang, R., Stoick, C. L., Kinnamon, S. C., and Kinnamon, J. C. (2004). Morphologic characterization of rat taste receptor cells that express components of the phospholipase C signaling pathway. J. Comp. Neurol. 468, 311-321.
26. Corvilain, B., Abramowicz, M., Fery, F., Schoutens, A., Verlinden, M., Balasse, E., and Horowitz, M. (1995). Effect of short-term starvation on gastric emptying in humans: relationship to oral glucose tolerance. Am. J. Physiol. 269, G512-G517.
27. Cummings, D. E., and Overduin, J. (2007). Gastrointestinal regulation of food intake. J. Clin. Invest. 117, 13-23.
28. Damak, S., Rong, M., Yasumatsu, K., Kokrashvili, Z., Perez, C. A., Shigemura, N., Yoshida, R., Mosinger, B. Jr., Glendinning, J. I., Ninomiya, Y., and Margolskee, R. F. (2006). Trpm5 null mice respond to bitter, sweet, and umami compounds. Chem. Senses 31, 253-264.
29. Damak, S., Rong, M., Yasumatsu, K., Kokrashvili, Z., Varadarajan, V., Zou, S., Jiang, P., Ninomiya, Y., and Margolskee, R. F. (2003). Detection of sweet and umami taste in the absence of taste receptor T1r3. Science 301, 850-853.
30. Deacon, C. F., Pridal, L., Klarskov, L., Olesen, M., and Holst, J. J. (1996). Glucagon-like peptide 1 undergoes differential tissue-specific metabolism in the anesthetized pig. Am. J. Physiol. 271, E458-E464.
31. Deane, A. M., Nguyen, N. Q., Stevens, J. E., Fraser, R. J., Holloway, R. H., Besanko, L. K., Burgstad, C., Jones, K. L., Chapman, M. J., Rayner, C. K., and Horowitz, M. (2010). Endogenous glucagon-like peptide-1 slows gastric emptying in healthy subjects, attenuating postprandial glycemia. J. Clin. Endocrinol. Metab. 95, 215-221.
32. DeFazio, R. A., Dvoryanchikov, G., Maruyama, Y., Kim, J. W., Pereira, E., Roper, S. D., and Chaudhari, N. (2006). Separate populations of receptor cells and presynaptic cells in mouse taste buds. J. Neurosci. 26, 3971-3980.
33. Delay, E. R., Hernandez, N. P., Bromley, K., and Margolskee, R. F. (2006). Sucrose and monosodium glutamate taste thresholds and discrimination ability of T1R3 knockout mice. Chem. Senses 31, 351-357.
34. Diez-Sampedro, A., Hirayama, B. A., Osswald, C., Gorboulev, V., Baumgarten, K., Volk, C., Wright, E. M., and Koepsell, H. (2003). A glucose sensor hiding in a family of transporters. Proc. Natl. Acad. Sci. U.S.A. 100, 11753-11758.
35. Drucker, D. J., and Asa, S. (1988). Glucagon gene expression in vertebrate brain. J. Biol. Chem. 263, 13475-13478.
36. Dyer, J., Daly, K., Salmon, K. S., Arora, D. K., Kokrashvili, Z., Margolskee, R. F., and Shirazi-Beechey, S. P. (2007). Intestinal glucose sensing and regulation of intestinal glucose absorption. Biochem. Soc. Trans. 35, 1191-1194.
37. Dyer, J., Salmon, K. S., Zibrik, L., and Shirazi-Beechey, S. P. (2005). Expression of sweet taste receptors of the T1R family in the intestinal tract and enteroendocrine cells. Biochem. Soc. Trans. 33, 302-305.
38. Dyer, J., Vayro, S., King, T. P., and Shirazi-Beechey, S. P. (2003). Glucose sensing in the intestinal epithelium. Eur. J. Biochem. 270, 3377-3388.
39. Eissele, R., Goke, R., Willemer, S., Harthus, H. P., Vermeer, H., Arnold, R., and Goke, B. (1992). Glucagon-like peptide-1 cells in the gastrointestinal tract and pancreas of rat, pig and man. Eur. J. Clin. Invest. 22, 283-291.
40. El-Ouaghlidi, A., Rehring, E., Holst, J. J., Schweizer, A., Foley, J., Holmes, D., and Nauck, M. A. (2007). The dipeptidyl peptidase 4 inhibitor vildagliptin does not accentuate glibenclamide-induced hypoglycemia but reduces glucose-induced glucagon-like peptide 1 and gastric inhibitory polypeptide secretion. J. Clin. Endocrinol. Metab. 92, 4165-4171.
41. Finger, T. E., Danilova, V., Barrows, J., Bartel, D. L., Vigers, A. J., Stone, L., Hellekant, G., and Kinnamon, S. C. (2005). ATP signaling is crucial for communication from taste buds to gustatory nerves. Science 310, 1495-1499.
42. Frank, G. K., Oberndorfer, T. A., Simmons, A. N., Paulus, M. P., Fudge, J. L., Yang, T. T., and Kaye, W. H. (2008). Sucrose activates human taste pathways differently from artificial sweetener. Neuroimage 39, 1559-1569.
43. Freeman, S. L., Bohan, D., Darcel, N., and Raybould, H. E. (2006). Luminal glucose sensing in the rat intestine has characteristics of a sodium-glucose cotransporter. Am. J. Physiol. 291, G439-G445.
44. Fujita, Y., Wideman, R. D., Speck, M., Asadi, A., King, D. S., Webber, T. D., Haneda, M., and Kieffer, T. J. (2009). Incretin release from gut is acutely enhanced by sugar but not by sweeteners in vivo. Am. J. Physiol. Endocrinol. Metab. 296, E473-E479.
45. Fushiki, T., Kojima, A., Imoto, T., Inoue, K., and Sugimoto, E. (1992). An extract of Gymnema sylvestre leaves and purified gymnemic acid inhibits glucose-stimulated gastric inhibitory peptide secretion in rats. J. Nutr. 122, 2367-2373.
46. Gebert, A., al-Samir, K., Werner, K., Fassbender, S., and Gebhard, A. (2000). The apical membrane of intestinal brush cells possesses a specialised, but species-specific, composition of glycoconjugates-on-section and in vivo lectin labelling in rats, guinea-pigs and mice. Histochem. Cell Biol. 113, 389-399.
47. Glatzle, J., Sternini, C., Robin, C., Zittel, T. T., Wong, H., Reeve, J. R., and Raybould, H. E. (2002). Expression of 5-HT3 receptors in the rat gastrointestinal tract. Gastroenterology 123, 217-226.
48. Goke, R., Larsen, P. J., Mikkelsen, J. D., and Sheikh, S. P. (1995). Distribution of GLP-1 binding sites in the rat brain: evidence that exendin-4 is a ligand of brain GLP-1 binding sites. Eur. J. Neurosci. 7, 2294-2300.
49. Gregersen, S., Jeppesen, P. B., Holst, J. J., and Hermansen, K. (2004). Antihyperglycemic effects of stevioside in type 2 diabetic subjects. Metab. Clin. Exp. 53, 73-76.
50. Gribble, F. M., Williams, L., Simpson, A. K., and Reimann, F. (2003). A novel glucose-sensing mechanism contributing to glucagon-like peptide-1 secretion from the GLUTag cell line. Diabetes 52, 1147-1154.
51. Grundy, D. (2006). Serotonin and sensory signalling from the gastrointestinal lumen. J. Physiol. (Lond.) 575, 1-2.
52. Grundy, D. (2008). 5-HT system in the gut: roles in the regulation of visceral sensitivity and motor functions. Eur. Rev. Med. Pharmacol. Sci. 12(Suppl. 1), 63-67.
53. Gwynne, R. M., and Bornstein, J. C. (2007). Local inhibitory reflexes excited by mucosal application of nutrient amino acids in guinea pig jejunum. Am. J. Physiol. 292, G1660-G1670.
54. Höfer, D., and Drenckhahn, D. (1996). Cytoskeletal markers allowing discrimination between brush cells and other epithelial cells of the gut including enteroendocrine cells. Histochem. Cell Biol. 105, 405-412.
55. Höfer, D., and Drenckhahn, D. (1998). Identification of the taste cell G-protein, alpha-gustducin, in brush cells of the rat pancreatic duct system. Histochem. Cell Biol. 110, 303-309.
56. Höfer, D., Puschel, B., and Drenckhahn, D. (1996). Taste receptor-like cells in the rat gut identified by expression of alpha-gustducin. Proc. Natl. Acad. Sci. U.S.A. 93, 6631-6634.
57. Holmes, G. M., Browning, K. N., Tong, M., Qualls-Creekmore, E., and Travagli, R. A. (2009). Vagally mediated effects of glucagon-like peptide 1: in vitro and in vivo gastric actions. J. Physiol. (Lond.) 587, 4749-4759.
58. Holst, J. J. (2007). The physiology of glucagon-like peptide 1. Physiol. Rev. 87, 1409-1439.
59. Holst, J. J., and Gromada, J. (2004). Role of incretin hormones in the regulation of insulin secretion in diabetic and nondiabetic humans. Am. J. Physiol. 287, E199-E206.
60. Hoon, M. A., Adler, E., Lindemeier, J., Battey, J. F., Ryba, N. J., and Zuker, C. S. (1999). Putative mammalian taste receptors: a class of taste-specific GPCRs with distinct topographic selectivity. Cell 96, 541-551.
61. Horowitz, M., Cunningham, K. M., Wishart, J. M., Jones, K. L., and Read, N. W. (1996). The effect of short-term dietary supplementation with glucose on gastric emptying of glucose and fructose and oral glucose tolerance in normal subjects. Diabetologia 39, 481-486.
62. Horowitz, M., and Dent, J. (1991). Disordered gastric emptying: mechanical basis, assessment and treatment. Baillieres Clin. Gastroenterol. 5, 371-407.
63. Horowitz, M., Maddox, A. F., Wishart, J. M., Harding, P. E., Chatterton, B. E., and Shearman, D. J. C. (1991). Relationships between oesophageal transit and solid and liquid gastric emptying in diabetes mellitus. Eur. J. Nucl. Med. 18, 229-234.
64. Horowitz, M., and Nauck, M. A. (2006). To be or not to be - an incretin or enterogastrone? Gut 55, 148-150.
65. Horowitz, M., O'Donovan, D., Jones, K. L., Feinle, C., Rayner, C. K., and Samsom, M. (2002). Gastric emptying in diabetes: clinical significance and treatment. Diabet. Med. 19, 177-194.
66. Huang, Y. A., Dando, R., and Roper, S. D. (2009). Autocrine and paracrine roles for ATP and serotonin in mouse taste buds. J. Neurosci. 29, 13909-13918.
67. Huang, Y. A., Maruyama, Y., Stimac, R., and Roper, S. D. (2008). Presynaptic (Type III) cells in mouse taste buds sense sour (acid) taste. J. Physiol. (Lond.) 586, 2903-2912.
68. Imeryuz, N., Yegen, B. C., Bozkurt, A., Coskun, T., Villanueva-Penacarrillo, M. L., and Ulusoy, N. B. (1997). Glucagon-like peptide-1 inhibits gastric emptying via vagal afferent-mediated central mechanisms. Am. J. Physiol. 273, G920-G927.
69. Jang, H. J., Kokrashvili, Z., Theodorakis, M. J., Carlson, O. D., Kim, B. J., Zhou, J., Kim, H. H., Xu, X., Chan, S. L., Juhaszova, M., Bernier, M., Mosinger, B., Margolskee, R. F., and Egan, J. M. (2007). Gut-expressed gustducin and taste receptors regulate secretion of glucagon-like peptide-1. Proc. Natl. Acad. Sci. U.S.A. 104, 15069-15074.
70. Kakei, M., Yada, T., Nakagawa, A., and Nakabayashi, H. (2002). Glucagon-like peptide-1 evokes action potentials and increases cytosolic Ca2+ in rat nodose ganglion neurons. Auton. Neurosci. 102, 39-44.
71. Kataoka, S., Toyono, T., Seta, Y., Ogura, T., and Toyoshima, K. (2004). Expression of P2Y1 receptors in rat taste buds. Histochem. Cell Biol. 121, 419-426.
72. Kataoka, S., Yang, R., Ishimaru, Y., Matsunami, H., Sevigny, J., Kinnamon, J. C., and Finger, T. E. (2008). The candidate sour taste receptor, PKD2L1, is expressed by type III taste cells in the mouse. Chem. Senses 33, 243-254.
73. Kidd, M., Modlin, I. M., Gustafsson, B. I., Drozdov, I., Hauso, O., and Pfragner, R. (2008). Luminal regulation of normal and neoplastic human EC cell serotonin release is mediated by bile salts, amines, tastants, and olfactants. Am. J. Physiol. 295, G260-G272.
74. Kim, M., Cooke, H. J., Javed, N. H., Carey, H. V., Christofi, F., and Raybould, H. E. (2001). D-glucose releases 5-hydroxytryptamine from human BON cells as a model of enterochromaffin cells. Gastroenterology 121, 1400-1406.
75. Kim, M. R., Kusakabe, Y., Miura, H., Shindo, Y., Ninomiya, Y., and Hino, A. (2003). Regional expression patterns of taste receptors and gustducin in the mouse tongue. Biochem. Biophys. Res. Commun. 312, 500-506.
76. Kim, S. J., Nian, C., and McIntosh, C. H. (2007). Activation of lipoprotein lipase by glucose-dependent insulinotropic polypeptide in adipocytes. A role for a protein kinase B, LKB1, and AMP-activated protein kinase cascade. J. Biol. Chem. 282, 8557-8567.
77. Kim, Y. V., Bobkov, Y. V., and Kolesnikov, S. S. (2000). Adenosine triphosphate mobilizes cytosolic calcium and modulates ionic currents in mouse taste receptor cells. Neurosci. Lett. 290, 165-168.
78. Kokrashvili, Z., Rodriguez, D., Yevshayeva, V., Zhou, H., Margolskee, R. F., and Mosinger, B. (2009). Release of endogenous opioids from duodenal enteroendocrine cells requires Trpm5. Gastroenterology 137, 598-606.
79. Kugler, P., Höfer, D., Mayer, B., and Drenckhahn, D. (1994). Nitric oxide synthase and NADP-linked glucose-6-phosphate dehydrogenase are co-localized in brush cells of rat stomach and pancreas. J. Histochem. Cytochem. 42, 1317-1321.
80. Kumar, K. G., Byerley, L. O., Volaufova, J., Drucker, D. J., Churchill, G. A., Li, R., York, B., Zuberi, A., and Richards, B. K. (2008). Genetic variation in Glp1r expression influences the rate of gastric emptying in mice. Am. J. Physiol. 294, R362-R371.
81. Lavin, J. H., Wittert, G., Sun, W. M., Horowitz, M., Morley, J. E., and Read, N. W. (1996). Appetite regulation by carbohydrate: role of blood glucose and gastrointestinal hormones. Am. J. Physiol. 271, E209-E214.
82. Lawton, D. M., Furness, D. N., Lindemann, B., and Hackney, C. M. (2000). Localization of the glutamate- aspartate transporter, GLAST, in rat taste buds. Eur. J. Neurosci. 12, 3163-3171.
83. Li, X., Staszewski, L., Xu, H., Durick, K., Zoller, M., and Adler, E. (2002). Human receptors for sweet and umami taste. Proc. Natl. Acad. Sci. U.S.A. 99, 4692-4696.
84. Liddle, R. A., Goldfine, I. D., Rosen, M. S., Taplitz, R. A., and Williams, J. A. (1985). Cholecystokinin bioactivity in human plasma. Molecular forms, responses to feeding, and relationship to gallbladder contraction. J. Clin. Invest. 75, 1144-1152.
85. Lin, H. C., Doty, J. E., Reedy, T. J., and Meyer, J. H. (1989). Inhibition of gastric emptying by glucose depends on length of intestine exposed to nutrient. Am. J. Physiol. 256, G404-G411.
86. Lindemann, B. (2001). Receptors and transduction in taste. Nature 413, 219-225.
87. Linden, D. R., Sha, L., Mazzone, A., Stoltz, G. J., Bernard, C. E., Furne, J. K., Levitt, M. D., Farrugia, G., and Szurszewski, J. H. (2008). Production of the gaseous signal molecule hydrogen sulfide in mouse tissues. J. Neurochem. 106, 1577-1585.
88. Little, T. J., Doran, S., Meyer, J. H., Smout, A. J., O'Donovan, D. G., Wu, K. L., Jones, K. L., Wishart, J., Rayner, C. K., Horowitz, M., and Feinle-Bisset, C. (2006). The release of GLP-1 and ghrelin, but not GIP and CCK, by glucose is dependent upon the length of small intestine exposed. Am. J. Physiol. 291, E647-E655.
89. Little, T. J., Gupta, N., Case, R. M., Thompson, D. G., and McLaughlin, J. T. (2009). Sweetness and bitterness taste of meals per se does not mediate gastric emptying in humans. Am. J. Physiol. 297, R632-R639.
90. Ma, J., Bellon, M., Wishart, J. M., Young, R. L., Blackshaw, L. A., Jones, K. L., Horowitz, M., and Rayner, C. K. (2009). Effect of the artificial sweetener, sucralose, on gastric empty ing and incretin hormone release in healthy subjects. Am. J. Physiol. 296, G735-G739.
91. Ma, J., Chang, J., Checklin, H. L., Young, R. L., Jones, K. L., Horowitz, M., and Rayner, C. K. (2010). Effect of the artificial sweetener, sucralose, on small intestinal glucose absorption in healthy human subjects. Br. J. Nutr. 104, 803-806.
92. Mace, O. J., Affleck, J., Patel, N., and Kellett, G. L. (2007). Sweet taste receptors in rat small intestine stimulate glucose absorption through apical GLUT2. J. Physiol. (Lond.) 582, 379-392.
93. Mace, O. J., Lister, N., Morgan, E. L., West, E. J., Affleck, J., Helliwell, P. A., Bronk, J. R., Kellett, G. L., Meredith, D., Boyd, C. A., Myrtani, P., Bailey, P. D., Pettecrew, R., and Foley, D. (2008). An energy supply network of nutrient absorption coordinated by calcium and T1R taste receptors in rat small intestine. J. Physiol. (Lond.) 587, 195-210.
94. Margolskee, R. F., Dyer, J., Kokrashvili, Z., Salmon, K. S., Ilegems, E., Daly, K., Maillet, E. L., Ninomiya, Y., Mosinger, B., and Shirazi-Beechey, S. P. (2007). T1R3 and gustducin in gut sense sugars to regulate expression of Na+-glucose cotransporter 1. Proc. Natl. Acad. Sci. U.S.A. 104, 15075-15080.
95. Martin, B., Dotson, C. D., Shin, Y. K., Ji, S., Drucker, D. J., Maudsley, S., and Munger, S. D. (2009). Modulation of taste sensitivity by GLP-1 signaling in taste buds. Ann. N. Y. Acad. Sci. 1170, 98-101.
96. McFadden, D. W., Jaffe, B. M., Ferrara, A., and Zinner, M. J. (1984). Jejunal absorptive response to a test meal and its modification by cholinergic and calcium-channel blockade in the awake dog. Surg. Forum 35, 174-176.
97. Meier, J. J., Goetze, O., Anstipp, J., Hagemann, D., Holst, J. J., Schmidt, W. E., Gallwitz, B., and Nauck, M. A. (2004). Gastric inhibitory polypeptide does not inhibit gastric emptying in humans. Am. J. Physiol. 286, E621-E625.
98. Merchenthaler, I., Lane, M., and Shughrue, P. (1999). Distribution of pre-pro-glucagon and glucagon-like peptide-1 receptor messenger RNAs in the rat central nervous system. J. Comp. Neurol. 403, 261-280.
99. Miller, S. M., Farrugia, G., Schmalz, P. F., Ermilov, L. G., Maines, M. D., and Szurszewski, J. H. (1998). Heme oxygenase 2 is present in interstitial cell networks of the mouse small intestine. Gastroenterology 114, 239-244.
100. Moran, A. W., Al-Rammahi, M. A., Arora, D. K., Batchelor, D. J., Coulter, E. A., Daly, K., Ionescu, C., Bravo, D., and Shirazi-Beechey, S. P. (2010a). Expression of Na+/glucose co-transporter 1 (SGLT1) is enhanced by supplementation of the diet of weaning piglets with artificial sweeteners. Br. J. Nutr. 104, 637-646.
101. Moran, A. W., Al-Rammahi, M. A., Arora, D. K., Batchelor, D. J., Coulter, E. A., Ionescu, C., Bravo, D., and Shirazi-Beechey, S. P. (2010b). Expression of Na+/glucose co-transporter 1 (SGLT1) in the intestine of piglets weaned to different concentrations of dietary carbohydrate. Br. J. Nutr. 104, 647-655.
102. Morini, G., Bassoli, A., and Temussi, P. A. (2005). From small sweeteners to sweet proteins: anatomy of the binding sites of the human T1R2_T1R3 receptor. J. Med. Chem. 48, 5520-5529.
103. Moriya, R., Shirakura, T., Ito, J., Mashiko, S., and Seo, T. (2009). Activation of sodium-glucose cotransporter 1 ameliorates hyperglycemia by mediating incretin secretion in mice. Am. J. Physiol. 297, E1358-E1365.
104. Morroni, M., Cangiotti, A. M., and Cinti, S. (2007). Brush cells in the human duodenojejunal junction: an ultrastructural study. J. Anat. 211, 125-131.
105. Murata, Y., Yasuo, T., Yoshida, R., Obata, K., Yanagawa, Y., Margolskee, R. F., and Ninomiya, Y. (2010). Action potential-enhanced ATP release from taste cells through hemichannels. J. Neurophysiol. 104, 896-901.
106. Nagahama, S., and Kurihara, K. (1985). Norepinephrine as a possible transmitter involved in synaptic transmission in frog taste organs and Ca dependence of its release. J. Gen. Physiol. 85, 431-442.
107. Nagai, T., Delay, R. J., Welton, J., and Roper, S. D. (1998). Uptake and release of neurotransmitter candidates, [3H] serotonin, [3H]glutamate, and [3H] gamma-aminobutyric acid, in taste buds of the mudpuppy, Necturus maculosus. J. Comp. Neurol. 392, 199-208.
108. Nagell, C. F., Wettergren, A., Ørskov, C., and Holst, J. J. (2006). Inhibitory effect of GLP-1 on gastric motility persists after vagal deafferentation in pigs. Scand. J. Gastroenterol. 41, 667-672.
109. Nakade, Y., Tsukamoto, K., Pappas, T. N., and Takahashi, T. (2006). Central glucagon like peptide-1 delays solid gastric emptying via central CRF and peripheral sympathetic pathway in rats. Brain Res. 1111, 117-121.
110. Nakagawa, A., Satake, H., Nakabayashi, H., Nishizawa, M., Furuya, K., Nakano, S., Kigoshi, T., Nakayama, K., and Uchida, K. (2004). Receptor gene expression of glucagon-like peptide-1, but not glucose-dependent insulinotropic polypeptide, in rat nodose ganglion cells. Auton. Neurosci. 110, 36-43.
111. Nakagawa, Y., Nagasawa, M., Yamada, S., Hara, A., Mogami, H., Nikolaev, V. O., Lohse, M. J., Shigemura, N., Ninomiya, Y., and Kojima, I. (2009). Sweet taste receptor expressed in pancreatic β-cells activates the calcium and cyclic AMP signaling systems and stimulates insulin secretion. PLoS ONE 4, e5106. doi: 10.1371/journal.pone.0005106
112. Nelson, G., Chandrashekar, J., Hoon, M. A., Feng, L., Zhao, G., Ryba, N. J., and Zuker, C. S. (2002). An amino-acid taste receptor. Nature 416, 199-202.
113. Nelson, G., Hoon, M. A., Chandrashekar, J., Zhang, Y., Ryba, N. J., and Zuker, C. S. (2001). Mammalian sweet taste receptors. Cell 106, 381-390.
114. Netzer, P., Gaia, C., Lourens, S. T., Reber, P., Wildi, S., Noelpp, U., Ritter, E. P., Ledermann, H., Luscher, D., Varga, L., Kinser, J. A., Buchler, M. W., and Scheurer, U. (2002). Does intravenous ondansetron affect gastric emptying of a solid meal, gastric electrical activity or blood hormone levels in healthy volunteers? Aliment. Pharmacol. Ther. 16, 119-127.
115. Nie, Y., Vigues, S., Hobbs, J. R., Conn, G. L., and Munger, S. D. (2005). Distinct contributions of T1R2 and T1R3 taste receptor subunits to the detection of sweet stimuli. Curr. Biol. 15, 1948-1952.
116. Nielsen, L. B., Ploug, K. B., Swift, P., Orskov, C., Jansen-Olesen, I., Chiarelli, F., Holst, J. J., Hougaard, P., Porksen, S., Holl, R., de Beaufort, C., Gammeltoft, S., Rorsman, P., Mortensen, H. B., and Hansen, L. (2007). Co-localisation of the Kir6.2/SUR1 channel complex with glucagon-like peptide-1 and glucose-dependent insulinotrophic polypeptide expression in human ileal cells and implications for glycaemic control in new onset type 1 diabetes. Eur. J. Endocrinol. 156, 663-671.
117. Nishizawa, M., Nakabayashi, H., Uchida, K., Nakagawa, A., and Niijima, A. (1996). The hepatic vagal nerve is receptive to incretin hormone glucagon-like peptide-1, but not to glucose-dependent insulinotropic polypeptide, in the portal vein. J. Auton. Nerv. Syst. 61, 149-154.
118. Ohkuri, T., Yasumatsu, K., Horio, N., Jyotaki, M., Margolskee, R. F., and Ninomiya, Y. (2009). Multiple sweet receptors and transduction pathways revealed in knockout mice by temperature dependence and gurmarin sensitivity. Am. J. Physiol. 296, R960-R971.
119. Ozeck, M., Brust, P., Xu, H., and Servant, G. (2004). Receptors for bitter, sweet and umami taste couple to inhibitory G protein signaling pathways. Eur. J. Pharmacol. 489, 139-149.
120. Parker, H. E., Habib, A. M., Rogers, G. J., Gribble, F. M., and Reimann, F. (2009). Nutrient-dependent secretion of glucose-dependent insulinotropic polypeptide from primary murine K cells. Diabetologia 52, 289-298.
121. Perez, C. A., Huang, L., Rong, M., Kozak, J. A., Preuss, A. K., Zhang, H., Max, M., and Margolskee, R. F. (2002). A transient receptor potential channel expressed in taste receptor cells. Nat. Neurosci. 5, 1169-1176.
122. Pilichiewicz, A. N., Chaikomin, R., Brennan, I. M., Wishart, J. M., Rayner, C. K., Jones, K. L., Smout, A. J., Horowitz, M., and Feinle-Bisset, C. (2007). Load-dependent effects of duodenal glucose on glycemia, gastrointestinal hormones, antropyloroduodenal motility, and energy intake in healthy men. Am. J. Physiol. 293, E743-E753.
123. Raybould, H. E. (1991). Capsaicin-sensitive vagal afferents and CCK in inhibition of gastric motor function induced by intestinal nutrients. Peptides 12, 1279-1283.
124. Raybould, H. E. (1998). Does your gut taste? Sensory transduction in the gastrointestinal tract. News Physiol. Sci. 13, 275-280.
125. Raybould, H. E. (2010). Gut chemosensing: interactions between gut endocrine cells and visceral afferents. Auton. Neurosci. 153, 41-46.
126. Raybould, H. E., Glatzle, J., Robin, C., Meyer, J. H., Phan, T., Wong, H., and Sternini, C. (2003). Expression of 5-HT3 receptors by extrinsic duodenal afferents contribute to intestinal inhibition of gastric emptying. Am. J. Physiol. 284, G367-G372.
127. Raybould, H. E., and Holzer, H. (1992). Dual capsaicin-sensitive afferent pathways mediate inhibition of gastric emptying in rat induced by intestinal carbohydrate. Neurosci. Lett. 141, 236-238.
128. Rayner, C. K., Schwartz, M. P., van Dam, P. S., Renooij, W., de Smet, M., Horowitz, M., Smout, A. J., and Samsom, M. (2002). Small intestinal glucose absorption and duodenal motility in type 1 diabetes mellitus. Am. J. Gastroenterol. 97, 3123-3130.
129. Rehfeld, J. F. (1978). Immunochemical studies on cholecystokinin. II. Distribution and molecular heterogeneity in the central nervous system and small intestine of man and hog. J. Biol. Chem. 253, 4022-4030.
130. Reid, L., Meyrick, B., Antony, V. B., Chang, L. Y., Crapo, J. D., and Reynolds, H. Y. (2005). The mysterious pulmonary brush cell: a cell in search of a function. Am. J. Respir. Crit. Care Med. 172, 136-139.
131. Reimann, F. (2010). Molecular mechanisms underlying nutrient detection by incretin-secreting cells. Int. Dairy J. 20, 236-242.
132. Reimann, F., and Gribble, F. M. (2002). Glucose-sensing in glucagon-like peptide-1-secreting cells. Diabetes 51, 2757-2763.
133. Reimann, F., Habib, A. M., Tolhurst, G., Parker, H. E., Rogers, G. J., and Gribble, F. M. (2008). Glucose sensing in L cells: a primary cell study. Cell Metab. 8, 532-539.
134. Reimann, F., Maziarz, M., Flock, G., Habib, A. M., Drucker, D. J., and Gribble, F. M. (2005). Characterization and functional role of voltage gated cation conductances in the glucagon-like peptide-1 secreting GLUTag cell line. J. Physiol. (Lond.) 563, 161-175.
135. Ren, X., Zhou, L., Terwilliger, R., Newton, S. S., and de Araujo, I. E. (2009). Sweet taste signaling functions as a hypothalamic glucose sensor. Front. Integr. Neurosci. 3:12. doi: 10.3389/neuro.07.012.2009
136. Rindi, G., Leiter, A. B., Kopin, A. S., Bordi, C., and Solcia, E. (2004). The " normal" endocrine cell of the gut: changing concepts and new evidences. Ann. N. Y. Acad. Sci. 1014, 1-12.
137. Ritzel, U., Fromme, A., Ottleben, M., Leonhardt, U., and Ramadori, G. (1997). Release of glucagon-like peptide-1 (GLP-1) by carbohydrates in the perfused rat ileum. Acta Diabetol. 34, 18-21.
138. Romanov, R. A., Rogachevskaja, O. A., Bystrova, M. F., Jiang, P., Margolskee, R. F., and Kolesnikov, S. S. (2007). Afferent neurotransmission mediated by hemichannels in mammalian taste cells. EMBO J. 26, 657-667.
139. Roper, S. D. (2007). Signal transduction and information processing in mammalian taste buds. Pflugers Arch. 454, 759-776.
140. Rorsman, P. (1997). The pancreatic beta-cell as a fuel sensor: an electrophysiologist's viewpoint. Diabetologia 40, 487-495.
141. Roze, C., Couturier, D., Chariot, J., and Debray, C. (1977). Inhibition of gastric electrical and mechanical activity by intraduodenal agents in pigs and the effects of vagotomy. Digestion 15, 526-539.
142. Rozengurt, E., and Sternini, C. (2007). Taste receptor signaling in the mammalian gut. Curr. Opin. Pharmacol. 7, 557-562.
143. Rozengurt, N., Wu, S. V., Chen, M. C., Huang, C., Sternini, C., and Rozengurt, E. (2006). Colocalization of the alpha-subunit of gustducin with PYY and GLP-1 in L cells of human colon. Am. J. Physiol. 291, G792-G802.
144. Ruttimann, E. B., Arnold, M., Hillebrand, J. J., Geary, N., and Langhans, W. (2009). Intrameal hepatic portal and intraperitoneal infusions of glucagon-like peptide-1 reduce spontaneous meal size in the rat via different mechanisms. Endocrinology 150, 1174-1181.
145. Savastano, D. M., Carelle, M., and Covasa, M. (2005). Serotonin-type 3 recep-tors mediate intestinal Polycose- and glucose-induced suppression of intake. Am. J. Physiol. 288, R1499-R1508.
146. Schiffman, S. S., Booth, B. J., Carr, B. T., Losee, M. L., Sattely-Miller, E. A., and Graham, B. G. (1995). Investigation of synergism in binary mixtures of sweeteners. Brain Res. Bull. 38, 105-120.
147. Schirra, J., and Göke, B. (2005). The physiological role of GLP-1 in human: incretin, ileal brake or more? Regul. Pept. 128, 109-115.
148. Schirra, J., Nicolaus, M., Woerle, H. J., Struckmeier, C., Katschinski, M., and Goke, B. (2009). GLP-1 regulates gastroduodenal motility involving cholinergic pathways. Neurogastroenterol. Motil. 21, 609-618.
149. Schwartz, G. J., Berkow, G., McHugh, P. R., and Moran, T. H. (1993). Gastric branch vagotomy blocks nutrient and cholecystokinin-induced suppression of gastric emptying. Am. J. Physiol. 264, R630-R637.
150. Shen, T., Kaya, N., Zhao, F. L., Lu, S. G., Cao, Y., and Herness, S. (2005). Co-expression patterns of the neuropeptides vasoactive intestinal peptide and cholecystokinin with the transduction molecules alpha-gustducin and T1R2 in rat taste receptor cells. Neuroscience 130, 229-238.
151. Shigemura, N., Nakao, K., Yasuo, T., Murata, Y., Yasumatsu, K., Nakashima, A., Katsukawa, H., Sako, N., and Ninomiya, Y. (2008). Gurmarin sensitivity of sweet taste responses is associated with co-expression patterns of T1r2, T1r3, and gustducin. Biochem. Biophys. Res. Commun. 367, 356-363.
152. Shirazi-Beechey, S. P., Hirayama, B. A., Wang, Y., Scott, D., Smith, M. W., and Wright, E. M. (1991). Ontogenic development of lamb intestinal sodium- glucose co-transporter is regulated by diet. J. Physiol. (Lond.) 437, 699-708.
153. Singh, S. K., Bartoo, A. C., Krishnan, S., Boylan, M. O., Schwartz, J. H., and Michael Wolfe, M. (2008). Glucose-dependent insulinotropic polypeptide (GIP) stimulates transepithelial glucose transport. Obesity (Silver Spring) 16, 2412-2416.
154. Stacher, G., Bergmann, H., Granser-Vacariu, G. V., Wiesnagrotzki, S., Wenzelabatzi, T. A., Gaupmann, G., Kugi, A., Steinringer, H., Schneider, C., and Hobart, J. (1991). Lack of systematic effects of the 5-hydroxytryptamine 3 receptor antagonist ICS 205-930 on gastric emptying and antral motor activity in patients with primary anorexia nervosa. Br. J. Clin. Pharmacol. 32, 685-689.
155. Stacher, G., Bergmann, H., Schneider, C., Steiner-Mittelbach, G., Gaupmann, G., Steinringer, H., Abatzi, T. A., and Stacher-Janotta, G. (1990). Effects of the 5-HT3 receptor antagonist ICS 205-930 on fat-delayed gastric emptying and antral motor activity. Br. J. Clin. Pharmacol. 30, 41-48.
156. Stearns, A. T., Balakrishnan, A., Rhoads, D. B., and Tavakkolizadeh, A. (2010). Rapid upregulation of sodium-glucose transporter SGLT1 in response to intestinal sweet taste stimulation. Ann. Surg. 251, 865-871.
157. Stearns, A. T., Balakrishnan, A., Rounds, J., Rhoads, D. B., Ashley, S. W., and Tavakkolizadeh, A. (2008). Capsaicin-sensitive vagal afferents modulate posttranscriptional regulation of the rat Na+/glucose cotransporter SGLT1. Am. J. Physiol. 294, G1078-G1083.
158. Stone, L. M., Tan, S. S., Tam, P. P., and Finger, T. E. (2002). Analysis of cell lineage relationships in taste buds. J. Neurosci. 22, 4522-4529.
159. Sutherland, K., Brierley, S. M., Horowitz, M., Rayner, C. K., Blackshaw, L. A., and Young, R. L. (2007a). Sweet taste transduction molecules are expressed in the upper gastrointestinal tract in humans. Gastroenterology 132(Suppl. 2), T1939.
160. Sutherland, K., Young, R. L., Cooper, N. J., Horowitz, M., and Blackshaw, L. A. (2007b). Phenotypic characterization of taste cells of the mouse small intestine. Am. J. Physiol. 292, G1420-G1428.
161. Sutherland, K., Brierley, S. M., Horowitz, M., Rayner, C. K., Blackshaw, L. A., and Young, R. L. (2008). Altered duodenal sweet taste receptor expression in diabetic hyperglycemia. Gastroenterology 134(Suppl. 2), 311.
162. Sykes, S., Morgan, L. M., English, J., and Marks, V. (1980). Evidence for preferential stimulation of gastric inhibitory polypeptide secretion in the rat by actively transported carbohydrates and their analogues. J. Endocrinol. 85, 201-207.
163. Theodorakis, M. J., Carlson, O., Michopoulos, S., Doyle, M. E., Juhaszova, M., Petraki, K., and Egan, J. M. (2006). Human duodenal enteroendocrine cells: source of both incretin peptides, GLP-1 and GIP. Am. J. Physiol. 290, E550-E559.
164. Tolhurst, G., Reimann, F., and Gribble, F. M. (2009). Nutritional regulation of glucagon-like peptide-1 secretion. J. Physiol. (Lond.) 587, 27-32.
165. Turk, E., Zabel, B., Mundlos, S., Dyer, J., and Wright, E. M. (1991). Glucose/ galactose malabsorption caused by a defect in the Na+/glucose cotransporter. Nature 350, 354-356.
166. Vahl, T. P., Tauchi, M., Durler, T. S., Elfers, E. E., Fernandes, T. M., Bitner, R. D., Ellis, K. S., Woods, S. C., Seeley, R. J., Herman, J. P., and D'Alessio, D. A. (2007). Glucagon-like peptide-1 (GLP-1) receptors expressed on nerve terminals in the portal vein mediate the effects of endogenous GLP-1 on glucose tolerance in rats. Endocrinology 148, 4965-4973.
167. Vandenbeuch, A., Clapp, T. R., and Kinnamon, S. C. (2008). Amiloride-sensitive channels in type I fungiform taste cells in mouse. BMC Neurosci. 9, 1. doi: 10.1186/1471-2202-9-1
168. Wachters-Hagedoorn, R. E., Priebe, M. G., Heimweg, J. A., Heiner, A. M., Englyst, K. N., Holst, J. J., Stellaard, F., and Vonk, R. J. (2006). The rate of intestinal glucose absorption is correlated with plasma glucose-dependent insulinotropic polypeptide concentrations in healthy men. J. Nutr. 136, 1511-1516.
169. Wilkinson, A. R., and Johnston, D. (1973). Effect of truncal, selective and highly selective vagotomy on gastric emptying and intestinal transit of a food-barium meal in man. Ann. Surg. 178, 190-193.
170. Wong, G. T., Gannon, K. S., and Margolskee, R. F. (1996). Transduction of bitter and sweet taste by gustducin. Nature 381, 796-800.
171. Wu, S. V., Rozengurt, N., Yang, M., Young, S. H., Sinnett-Smith, J., and Rozengurt, E. (2002). Expression of bitter taste receptors of the T2R family in the gastrointestinal tract and enteroendocrine STC-1 cells. Proc. Natl. Acad. Sci. U.S.A. 99, 2392-2397.
172. Wu, X. Y., Zhu, J. X., Gao, J., Owyang, C., and Li, Y. (2005). Neurochemical phenotype of vagal afferent neurons activated to express C-FOS in response to luminal stimulation in the rat. Neuroscience 130, 757-767.
173. Xu, H., Staszewski, L., Tang, H., Adler, E., Zoller, M., and Li, X. (2004). Different functional roles of T1R subunits in the heteromeric taste receptors. Proc. Natl. Acad. Sci. U.S.A. 101, 14258-14263.
174. Yang, R., Tabata, S., Crowley, H. H., Margolskee, R. F., and Kinnamon, J. C. (2000). Ultrastructural localization of gustducin immunoreactivity in microvilli of type II taste cells in the rat. J. Comp. Neurol. 425, 139-151.
175. Yarmolinsky, D. A., Zuker, C. S., and Ryba, N. J. P. (2009). Common sense about taste: from mammals to insects. Cell 139, 234-244.
176. Yip, R. G., Boylan, M. O., Kieffer, T. J., and Wolfe, M. M. (1998). Functional GIP receptors are present on adipocytes. Endocrinology 139, 4004-4007.
177. Yoshida, R., Miyauchi, A., Yasuo, T., Jyotaki, M., Murata, Y., Yasumatsu, K., Shigemura, N., Yanagawa, Y., Obata, K., Ueno, H., Margolskee, R. F., and Ninomiya, Y. (2009). Discrimination of taste qualities among mouse fungiform taste bud cells. J. Physiol. (Lond.) 587, 4425-4439.
178. Young, R. L., Sutherland, K., Pezos, N., Brierley, S. M., Horowitz, M., Rayner, C. K., and Blackshaw, L. A. (2009). Expression of taste molecules in the upper gastrointestinal tract in humans with and without type 2 diabetes. Gut 58, 337-346.
179. Young, R. L., Sutherland, K., Pezos, N., Brierley, S. M., Ma, J., Rayner, C. K., Horowitz, M. and Blackshaw, L. A. (2010). Control of intestinal sweet taste receptor expression is impaired in type 2 diabetes. Neurogastroenterol. Motil. 22 (Suppl.) 1, 31.
180. Zhang, F., Klebansky, B., Fine, R. M., Liu, H., Xu, H., Servant, G., Zoller, M., Tachdjian, C., and Li, X. (2010). Molecular mechanism of the sweet taste enhancers. Proc. Natl. Acad. Sci. U.S.A. 107, 4752-4757.
181. Zhao, F. L., Shen, T., Kaya, N., Lu, S. G., Cao, Y., and Herness, S. (2005). Expression, physiological action, and coexpression patterns of neuropeptide Y in rat taste-bud cells. Proc. Natl. Acad. Sci. U.S.A. 102, 11100-11105.
182. Zhao, G. Q., Zhang, Y., Hoon, M. A., Chandrashekar, J., Erlenbach, I., Ryba, N. J., and Zuker, C. S. (2003). The receptors for mammalian sweet and umami taste. Cell 115, 255-266.
183. Zhu, J., Zhu, X., Owyang, C., and Li, Y. (2001). Intestinal serotonin acts as a paracrine substance to mediate vagal signal transmission evoked by luminal factors in the rat. J. Physiol. (Lond.) 530, 431-442.
184. Zittel, T. T., Rothenhofer, I., Meyer, J. H., and Raybould, H. E. (1994). Small intestinal capsaicin-sensitive afferents mediate feedback inhibition of gastric emptying in rats. Am. J. Physiol. 267, G1142-G1145.