Altered ghrelin secretion in mice in response to diet-induced obesity and Roux-en-Y gastric bypass

Molecular Metabolism

Volumn 3, Issue 7, 2014, Pages 717-730

  Uchida, Aki ^a,b   Zechner, Juliet F ^a   Mani, Bharath K ^a,b   Park, Won mee ^a,b   Aguirre, Vincent ^a   Zigman, Jeffrey M ^a,b  

^a UNIVERSITY OF TEXAS SOUTHWESTERN MEDICAL CENTER   (United States)

^b UNIVERSITY OF TEXAS SOUTHWESTERN MEDICAL CENTER   (United States)

Author keywords

Ghrelin secretion; Glucose; Norepinephrine; Obesity; Roux en Y gastric bypass; Sympathetic nervous system

Indexed keywords

GHRELIN; GLUCOSE; NORADRENALIN; SOMATOSTATIN;

ANIMAL CELL; ANIMAL EXPERIMENT; ANIMAL MODEL; ANIMAL TISSUE; ARTICLE; CELL DENSITY; CONTROLLED STUDY; DIETARY INTAKE; DUODENUM; HISTOPATHOLOGY; HORMONAL REGULATION; HORMONE BLOOD LEVEL; HORMONE RELEASE; IMMUNOREACTIVITY; MALE; MOUSE; NONHUMAN; OBESITY; POSTOPERATIVE PERIOD; ROUX Y ANASTOMOSIS; STOMACH; STOMACH BYPASS; TREATMENT RESPONSE; WEIGHT REDUCTION;

MUS;

EID: 84908164702     PISSN: None     EISSN: 22128778     Source Type: Journal    
DOI: 10.1016/j.molmet.2014.07.009     Document Type: Article

References (109)

1. Kojima M., Kangawa K. Ghrelin: from gene to physiological function. Results and Problems in Cell Differentiation 2010, 50:185-205.
2. Ariyasu H., Takaya K., Tagami T., Ogawa Y., Hosoda K., Akamizu T., et al. Stomach is a major source of circulating ghrelin, and feeding state determines plasma ghrelin-like immunoreactivity levels in humans. Journal of Clinical Endocrinology & Metabolism 2001, 86:4753-4758.
3. Date Y., Kojima M., Hosoda H., Sawaguchi A., Mondal M.S., Suganuma T., et al. Ghrelin, a novel growth hormone-releasing acylated peptide, is synthesized in a distinct endocrine cell type in the gastrointestinal tracts of rats and humans. Endocrinology 2000, 141:4255-4261.
4. Sakata I., Nakamura K., Yamazaki M., Matsubara M., Hayashi Y., Kangawa K., et al. Ghrelin-producing cells exist as two types of cells, closed- and opened-type cells, in the rat gastrointestinal tract. Peptides 2002, 23:531-536.
5. Wierup N., Svensson H., Mulder H., Sundler F. The ghrelin cell: a novel developmentally regulated islet cell in the human pancreas. Regulatory Peptides 2002, 107:63-69.
6. Lee H.M., Wang G., Englander E.W., Kojima M., Greeley G.H. Ghrelin, a new gastrointestinal endocrine peptide that stimulates insulin secretion: enteric distribution, ontogeny, influence of endocrine, and dietary manipulations. Endocrinology 2002, 143:185-190.
7. Gutierrez J.A., Solenberg P.J., Perkins D.R., Willency J.A., Knierman M.D., Jin Z., et al. Ghrelin octanoylation mediated by an orphan lipid transferase. Proceedings of the National Academy of Sciences of the United States of America 2008, 105:6320-6325.
8. Yang J., Brown M.S., Liang G., Grishin N.V., Goldstein J.L. Identification of the acyltransferase that octanoylates ghrelin, an appetite-stimulating peptide hormone. Cell 2008, 132:387-396.
9. Delhanty P.J., Neggers S.J., van der Lely A.J. Mechanisms in endocrinology: ghrelin: the differences between acyl- and des-acyl ghrelin. European Journal of Endocrinology 2012, 167:601-608.
10. Cummings D.E. Ghrelin and the short- and long-term regulation of appetite and body weight. Physiology & Behavior 2006, 89:71-84.
11. Strassburg S., Anker S.D., Castaneda T.R., Burget L., Perez-Tilve D., Pfluger P.T., et al. Long-term effects of ghrelin and ghrelin receptor agonists on energy balance in rats. American Journal of Physiology. Endocrinology and Metabolism 2008, 295:E78-E84.
12. Nakazato M., Murakami N., Date Y., Kojima M., Matsuo H., Kangawa K., et al. A role for ghrelin in the central regulation of feeding. Nature 2001, 409:194-198.
13. Asakawa A., Inui A., Kaga T., Yuzuriha H., Nagata T., Ueno N., et al. Ghrelin is an appetite-stimulatory signal from stomach with structural resemblance to motilin. Gastroenterology 2001, 120:337-345.
14. Tschop M., Smiley D.L., Heiman M.L. Ghrelin induces adiposity in rodents. Nature 2000, 407:908-913.
15. Wren A.M., Small C.J., Abbott C.R., Dhillo W.S., Seal L.J., Cohen M.A., et al. Ghrelin causes hyperphagia and obesity in rats. Diabetes 2001, 50:2540-2547.
16. Uchida A., Zigman J.M., Perello M. Ghrelin and eating behavior: evidence and insights from genetically-modified mouse models. Frontiers in Neuroscience 2013, 7:121.
17. Egecioglu E., Jerlhag E., Salome N., Skibicka K.P., Haage D., Bohlooly Y.M., et al. Ghrelin increases intake of rewarding food in rodents. Addiction Biology 2010, 15:304-311.
18. Walker A.K., Ibia I.E., Zigman J.M. Disruption of cue-potentiated feeding in mice with blocked ghrelin signaling. Physiology & Behavior 2012, 108:34-43.
19. Kanoski S.E., Fortin S.M., Ricks K.M., Grill H.J. Ghrelin signaling in the ventral hippocampus stimulates learned and motivational aspects of feeding via PI3K-Akt signaling. Biological Psychiatry 2013, 73:915-923.
20. Perello M., Sakata I., Birnbaum S., Chuang J.C., Osborne-Lawrence S., Rovinsky S.A., et al. Ghrelin increases the rewarding value of high-fat diet in an orexin-dependent manner. Biological Psychiatry 2010, 67:880-886.
21. Disse E., Bussier A.L., Deblon N., Pfluger P.T., Tschop M.H., Laville M., et al. Systemic ghrelin and reward: effect of cholinergic blockade. Physiology & Behavior 2011, 102:481-484.
22. Sun Y., Butte N.F., Garcia J.M., Smith R.G. Characterization of adult ghrelin and ghrelin receptor knockout mice under positive and negative energy balance. Endocrinology 2008, 149:843-850.
23. Albarran-Zeckler R.G., Sun Y., Smith R.G. Physiological roles revealed by ghrelin and ghrelin receptor deficient mice. Peptides 2011, 32:2229-2235.
24. Sun Y., Ahmed S., Smith R.G. Deletion of ghrelin impairs neither growth nor appetite. Molecular and Cellular Biology 2003, 23:7973-7981.
25. Sun Y., Wang P., Zheng H., Smith R.G. Ghrelin stimulation of growth hormone release and appetite is mediated through the growth hormone secretagogue receptor. Proceedings of the National Academy of Sciences of the United States of America 2004, 101:4679-4684.
26. Wortley K.E., Anderson K.D., Garcia K., Murray J.D., Malinova L., Liu R., et al. Genetic deletion of ghrelin does not decrease food intake but influences metabolic fuel preference. Proceedings of the National Academy of Sciences of the United States of America 2004, 101:8227-8232.
27. Zhao T.J., Liang G., Li R.L., Xie X., Sleeman M.W., Murphy A.J., et al. Ghrelin O-acyltransferase (GOAT) is essential for growth hormone-mediated survival of calorie-restricted mice. Proceedings of the National Academy of Sciences of the United States of America 2010, 107:7467-7472.
28. Kirchner H., Gutierrez J.A., Solenberg P.J., Pfluger P.T., Czyzyk T.A., Willency J.A., et al. GOAT links dietary lipids with the endocrine control of energy balance. Nature Medicine 2009, 15:741-745.
29. Wortley K.E., del Rincon J.P., Murray J.D., Garcia K., Iida K., Thorner M.O., et al. Absence of ghrelin protects against early-onset obesity. Journal of Clinical Investigation 2005, 115:3573-3578.
30. Zigman J.M., Nakano Y., Coppari R., Balthasar N., Marcus J.N., Lee C.E., et al. Mice lacking ghrelin receptors resist the development of diet-induced obesity. Journal of Clinical Investigation 2005, 115:3564-3572.
31. Pfluger P.T., Kirchner H., Gunnel S., Schrott B., Perez-Tilve D., Fu S., et al. Simultaneous deletion of ghrelin and its receptor increases motor activity and energy expenditure. American Journal of Physiology - Gastrointestinal and Liver Physiology 2008, 294:G610-G618.
32. Shearman L.P., Wang S.P., Helmling S., Stribling D.S., Mazur P., Ge L., et al. Ghrelin neutralization by a ribonucleic acid-SPM ameliorates obesity in diet-induced obese mice. Endocrinology 2006, 147:1517-1526.
33. Zorrilla E.P., Iwasaki S., Moss J.A., Chang J., Otsuji J., Inoue K., et al. Vaccination against weight gain. Proceedings of the National Academy of Sciences of the United States of America 2006, 103:13226-13231.
34. Rudolph J., Esler W.P., O'Connor S., Coish P.D., Wickens P.L., Brands M., et al. Quinazolinone derivatives as orally available ghrelin receptor antagonists for the treatment of diabetes and obesity. Journal of Medicinal Chemistry 2007, 50:5202-5216.
35. Esler W.P., Rudolph J., Claus T.H., Tang W., Barucci N., Brown S.E., et al. Small-molecule ghrelin receptor antagonists improve glucose tolerance, suppress appetite, and promote weight loss. Endocrinology 2007, 148:5175-5185.
36. McFarlane M.R., Brown M.S., Goldstein J.L., Zhao T.J. Induced ablation of ghrelin cells in adult mice does not decrease food intake, body weight, or response to high-fat diet. Cell Metabolism 2014, 20:54-60.
37. Fujitsuka N., Asakawa A., Uezono Y., Minami K., Yamaguchi T., Niijima A., et al. Potentiation of ghrelin signaling attenuates cancer anorexia-cachexia and prolongs survival. Translational Psychiatry 2011, 1:e23.
38. Chuang J.C., Perello M., Sakata I., Osborne-Lawrence S., Savitt J.M., Lutter M., et al. Ghrelin mediates stress-induced food-reward behavior in mice. Journal of Clinical Investigation 2011, 121:2684-2692.
39. Dezaki K., Hosoda H., Kakei M., Hashiguchi S., Watanabe M., Kangawa K., et al. Endogenous ghrelin in pancreatic islets restricts insulin release by attenuating Ca2+ signaling in beta-cells: implication in the glycemic control in rodents. Diabetes 2004, 53:3142-3151.
40. Dezaki K., Kakei M., Yada T. Ghrelin uses Galphai2 and activates voltage-dependent K+ channels to attenuate glucose-induced Ca2+ signaling and insulin release in islet beta-cells: novel signal transduction of ghrelin. Diabetes 2007, 56:2319-2327.
41. Broglio F., Arvat E., Benso A., Gottero C., Muccioli G., Papotti M., et al. Ghrelin, a natural GH secretagogue produced by the stomach, induces hyperglycemia and reduces insulin secretion in humans. Journal of Clinical Endocrinology & Metabolism 2001, 86:5083-5086.
42. Reed J.A., Benoit S.C., Pfluger P.T., Tschop M.H., D'Alessio D.A., Seeley R.J. Mice with chronically increased circulating ghrelin develop age-related glucose intolerance. American Journal of Physiology. Endocrinology and Metabolism 2008, 294:E752-E760.
43. Colombo M., Gregersen S., Xiao J., Hermansen K. Effects of ghrelin and other neuropeptides (CART, MCH, orexin A and B, and GLP-1) on the release of insulin from isolated rat islets. Pancreas 2003, 27:161-166.
44. Chuang J.C., Sakata I., Kohno D., Perello M., Osborne-Lawrence S., Repa J.J., et al. Ghrelin directly stimulates glucagon secretion from pancreatic alpha-cells. Molecular Endocrinology 2011, 25:1600-1611.
45. Yi C.X., Heppner K.M., Kirchner H., Tong J., Bielohuby M., Gaylinn B.D., et al. The GOAT-ghrelin system is not essential for hypoglycemia prevention during prolonged calorie restriction. PLoS ONE 2012, 7:e32100.
46. Li R.L., Sherbet D.P., Elsbernd B.L., Goldstein J.L., Brown M.S., Zhao T.J. Profound hypoglycemia in starved, ghrelin-deficient mice is caused by decreased gluconeogenesis and reversed by lactate or fatty acids. Journal of Biological Chemistry 2012, 287:17942-17950.
47. Wang Q., Liu C., Uchida A., Chuang J.C., Walker A., Liu T., et al. Arcuate AgRP neurons mediate orexigenic and glucoregulatory actions of ghrelin. Molecular Metabolism 2014, 3:64-72.
48. Cummings D.E., Purnell J.Q., Frayo R.S., Schmidova K., Wisse B.E., Weigle D.S. A preprandial rise in plasma ghrelin levels suggests a role in meal initiation in humans. Diabetes 2001, 50:1714-1719.
49. Tschop M., Wawarta R., Riepl R.L., Friedrich S., Bidlingmaier M., Landgraf R., et al. Post-prandial decrease of circulating human ghrelin levels. Journal of Endocrinological Investigation 2001, 24:RC19-RC21.
50. Wren A.M., Seal L.J., Cohen M.A., Brynes A.E., Frost G.S., Murphy K.G., et al. Ghrelin enhances appetite and increases food intake in humans. Journal of Clinical Endocrinology & Metabolism 2001, 86:5992-5995.
51. Tschop M., Weyer C., Tataranni P.A., Devanarayan V., Ravussin E., Heiman M.L. Circulating ghrelin levels are decreased in human obesity. Diabetes 2001, 50:707-709.
52. Williams D.L., Grill H.J., Cummings D.E., Kaplan J.M. Overfeeding-induced weight gain suppresses plasma ghrelin levels in rats. Journal of Endocrinological Investigation 2006, 29:863-868.
53. DelParigi A., Tschop M., Heiman M.L., Salbe A.D., Vozarova B., Sell S.M., et al. High circulating ghrelin: a potential cause for hyperphagia and obesity in Prader-Willi syndrome. Journal of Clinical Endocrinology & Metabolism 2002, 87:5461-5464.
54. Maksud F.A., Alves J.S., Diniz M.T., Barbosa A.J. Density of ghrelin-producing cells is higher in the gastric mucosa of morbidly obese patients. European Journal of Endocrinology 2011, 165:57-62.
55. le Roux C.W., Patterson M., Vincent R.P., Hunt C., Ghatei M.A., Bloom S.R. Postprandial plasma ghrelin is suppressed proportional to meal calorie content in normal-weight but not obese subjects. Journal of Clinical Endocrinology & Metabolism 2005, 90:1068-1071.
56. Hansen T.K., Dall R., Hosoda H., Kojima M., Kangawa K., Christiansen J.S., et al. Weight loss increases circulating levels of ghrelin in human obesity. Clinical Endocrinology 2002, 56:203-206.
57. Schauer P.R., Kashyap S.R., Wolski K., Brethauer S.A., Kirwan J.P., Pothier C.E., et al. Bariatric surgery versus intensive medical therapy in obese patients with diabetes. New England Journal of Medicine 2012, 366:1567-1576.
58. Cummings D.E., Weigle D.S., Frayo R.S., Breen P.A., Ma M.K., Dellinger E.P., et al. Plasma ghrelin levels after diet-induced weight loss or gastric bypass surgery. New England Journal of Medicine 2002, 346:1623-1630.
59. Nguyen N.T., Masoomi H., Magno C.P., Nguyen X.M., Laugenour K., Lane J. Trends in use of bariatric surgery, 2003-2008. Journal of the American College of Surgeons 2011, 213:261-266.
60. Kushner R.F. Weight loss strategies for treatment of obesity. Progress in Cardiovascular Diseases 2014, 56:465-472.
61. Schauer P.R., Burguera B., Ikramuddin S., Cottam D., Gourash W., Hamad G., et al. Effect of laparoscopic Roux-en Y gastric bypass on type 2 diabetes mellitus. Annals of Surgery 2003, 238:467-484.
62. Cummings D.E., Overduin J., Shannon M.H., Foster-Schubert K.E. Hormonal mechanisms of weight loss and diabetes resolution after bariatric surgery. Surgery for Obesity and Related Diseases 2005, 1:358-368.
63. Jacobsen S.H., Olesen S.C., Dirksen C., Jorgensen N.B., Bojsen-Moller K.N., Kielgast U., et al. Changes in gastrointestinal hormone responses, insulin sensitivity, and beta-cell function within 2 weeks after gastric bypass in non-diabetic subjects. Obesity Surgery 2012, 22:1084-1096.
64. Dirksen C., Jorgensen N.B., Bojsen-Moller K.N., Kielgast U., Jacobsen S.H., Clausen T.R., et al. Gut hormones, early dumping and resting energy expenditure in patients with good and poor weight loss response after Roux-en-Y gastric bypass. International Journal of Obesity (London) 2013, 37:1452-1459.
65. Korner J., Bessler M., Inabnet W., Taveras C., Holst J.J. Exaggerated glucagon-like peptide-1 and blunted glucose-dependent insulinotropic peptide secretion are associated with Roux-en-Y gastric bypass but not adjustable gastric banding. Surgery for Obesity and Related Diseases 2007, 3:597-601.
66. Rubino F., Gagner M., Gentileschi P., Kini S., Fukuyama S., Feng J., et al. The early effect of the Roux-en-Y gastric bypass on hormones involved in body weight regulation and glucose metabolism. Annals of Surgery 2004, 240:236-242.
67. Tymitz K., Engel A., McDonough S., Hendy M.P., Kerlakian G. Changes in ghrelin levels following bariatric surgery: review of the literature. Obesity Surgery 2011, 21:125-130.
68. Mundinger T.O., Cummings D.E., Taborsky G.J. Direct stimulation of ghrelin secretion by sympathetic nerves. Endocrinology 2006, 147:2893-2901.
69. de la Cour C.D., Norlen P., Hakanson R. Secretion of ghrelin from rat stomach ghrelin cells in response to local microinfusion of candidate messenger compounds: a microdialysis study. Regulatory Peptides 2007, 143:118-126.
70. Iwakura H., Ariyasu H., Hosoda H., Yamada G., Hosoda K., Nakao K., et al. Oxytocin and dopamine stimulate ghrelin secretion by the ghrelin-producing cell line, MGN3-1 in vitro. Endocrinology 2011, 152:2619-2625.
71. Sakata I., Park W.M., Walker A.K., Piper P.K., Chuang J.C., Osborne-Lawrence S., et al. Glucose-mediated control of ghrelin release from primary cultures of gastric mucosal cells. American Journal of Physiology. Endocrinology and Metabolism 2012, 302:E1300-E1310.
72. Gagnon J., Anini Y. Glucagon stimulates ghrelin secretion through the activation of MAPK and EPAC and potentiates the effect of norepinephrine. Endocrinology 2013, 154:666-674.
73. Engelstoft M.S., Park W.M., Sakata I., Kristensen L.V., Husted A.S., Osborne-Lawrence S., et al. Seven transmembrane G protein-coupled receptor repertoire of gastric ghrelin cells. Molecular Metabolism 2013, 2:376-392.
74. Flanagan D.E., Evans M.L., Monsod T.P., Rife F., Heptulla R.A., Tamborlane W.V., et al. The influence of insulin on circulating ghrelin. American Journal of Physiology. Endocrinology and Metabolism 2003, 284:E313-E316.
75. Shimada M., Date Y., Mondal M.S., Toshinai K., Shimbara T., Fukunaga K., et al. Somatostatin suppresses ghrelin secretion from the rat stomach. Biochemical and Biophysical Research Communications 2003, 302:520-525.
76. Schaller G., Schmidt A., Pleiner J., Woloszczuk W., Wolzt M., Luger A. Plasma ghrelin concentrations are not regulated by glucose or insulin: a double-blind, placebo-controlled crossover clamp study. Diabetes 2003, 52:16-20.
77. Seoane L.M., Al-Massadi O., Barreiro F., Dieguez C., Casanueva F.F. Growth hormone and somatostatin directly inhibit gastric ghrelin secretion. An in vitro organ culture system. Journal of Endocrinological Investigation 2007, 30:RC22-25.
78. Sakata I., Nakano Y., Osborne-Lawrence S., Rovinsky S.A., Lee C.E., Perello M., et al. Characterization of a novel ghrelin cell reporter mouse. Regulatory Peptides 2009, 155:91-98.
79. Zechner J.F., Mirshahi U.L., Satapati S., Berglund E.D., Rossi J., Scott M.M., et al. Weight-independent effects of roux-en-Y gastric bypass on glucose homeostasis via melanocortin-4 receptors in mice and humans. Gastroenterology 2013, 144:580-590.
80. Abdemur A., Slone J., Berho M., Gianos M., Szomstein S., Rosenthal R.J. Morphology, localization, and patterns of ghrelin-producing cells in stomachs of a morbidly obese population. Surgical Laparoscopy Endoscopy & Percutaneous Techniques 2014, 24:122-126.
81. Chandarana K., Gelegen C., Karra E., Choudhury A.I., Drew M.E., Fauveau V., et al. Diet and gastrointestinal bypass-induced weight loss: the roles of ghrelin and peptide YY. Diabetes 2011, 60:810-818.
82. Moesgaard S.G., Ahren B., Carr R.D., Gram D.X., Brand C.L., Sundler F. Effects of high-fat feeding and fasting on ghrelin expression in the mouse stomach. Regulatory Peptides 2004, 120:261-267.
83. Dadan J., Hady H.R., Zbucki R.L., Iwacewicz P., Bossowski A., Kasacka I. The activity of gastric ghrelin positive cells in obese patients treated surgically. Folia Histochemica et Cytobiologica 2009, 47:307-313.
84. Andrews Z.B., Liu Z.W., Walllingford N., Erion D.M., Borok E., Friedman J.M., et al. UCP2 mediates ghrelin's action on NPY/AgRP neurons by lowering free radicals. Nature 2008, 454:846-851.
85. Lopez M., Lage R., Saha A.K., Perez-Tilve D., Vazquez M.J., Varela L., et al. Hypothalamic fatty acid metabolism mediates the orexigenic action of ghrelin. Cell Metabolism 2008, 7:389-399.
86. Gahete M.D., Cordoba-Chacon J., Salvatori R., Castano J.P., Kineman R.D., Luque R.M. Metabolic regulation of ghrelin O-acyl transferase (GOAT) expression in the mouse hypothalamus, pituitary, and stomach. Molecular and Cellular Endocrinology 2010, 317:154-160.
87. Chambers A.P., Kirchner H., Wilson-Perez H.E., Willency J.A., Hale J.E., Gaylinn B.D., et al. The effects of vertical sleeve gastrectomy in rodents are ghrelin independent. Gastroenterology 2013, 144:50-52.
88. Stylopoulos N., Davis P., Pettit J.D., Rattner D.W., Kaplan L.M. Changes in serum ghrelin predict weight loss after Roux-en-Y gastric bypass in rats. Surgical Endoscopy 2005, 19:942-946.
89. Zhou D., Jiang X., Ding W., Zhang D., Yang L., Zhen C., et al. Impact of bariatric surgery on ghrelin and obestatin levels in obesity or type 2 diabetes mellitus rat model. Journal of Diabetes Research 2014, 2014:569435.
90. Aprahamian C.J., Tekant G., Chen M., Yagmurlu A., Yang Y.K., Loux T., et al. A rat model of childhood diet-induced obesity: Roux-en-Y gastric bypass induced changes in metabolic parameters and gastric peptide ghrelin. Pediatric Surgery International 2007, 23:653-657.
91. Sundbom M., Holdstock C., Engstrom B.E., Karlsson F.A. Early changes in ghrelin following Roux-en-Y gastric bypass: influence of vagal nerve functionality?. Obesity Surgery 2007, 17:304-310.
92. Davis J.F., Tracy A.L., Schurdak J.D., Magrisso I.J., Grayson B.E., Seeley R.J., et al. Roux en Y gastric bypass increases ethanol intake in the rat. Obesity Surgery 2013, 23:920-930.
93. Shin A.C., Zheng H., Townsend R.L., Sigalet D.L., Berthoud H.R. Meal-induced hormone responses in a rat model of Roux-en-Y gastric bypass surgery. Endocrinology 2010, 151:1588-1597.
94. Matzko M.E., Argyropoulos G., Wood G.C., Chu X., McCarter R.J., Still C.D., et al. Association of ghrelin receptor promoter polymorphisms with weight loss following Roux-en-Y gastric bypass surgery. Obesity Surgery 2012, 22:783-790.
95. Lutter M., Sakata I., Osborne-Lawrence S., Rovinsky S.A., Anderson J.G., Jung S., et al. The orexigenic hormone ghrelin defends against depressive symptoms of chronic stress. Nature Neuroscience 2008, 11:752-753.
96. Robertson M.D., Henderson R.A., Vist G.E., Rumsey R.D. Plasma ghrelin response following a period of acute overfeeding in normal weight men. International Journal of Obesity and Related Metabolic Disorders 2004, 28:727-733.
97. Gong Z., Yoshimura M., Aizawa S., Kurotani R., Zigman J.M., Sakai T., et al. G protein-coupled receptor 120 signaling regulates ghrelin secretion in vivo and in vitro. American Journal of Physiology. Endocrinology and Metabolism 2014, 306:E28-E35.
98. Lu X., Zhao X., Feng J., Liou A.P., Anthony S., Pechhold S., et al. Postprandial inhibition of gastric ghrelin secretion by long-chain fatty acid through GPR120 in isolated gastric ghrelin cells and mice. American Journal of Physiology - Gastrointestinal and Liver Physiology 2012, 303:G367-G376.
99. Al Massadi O., Pardo M., Roca-Rivada A., Castelao C., Casanueva F.F., Seoane L.M. Macronutrients act directly on the stomach to regulate gastric ghrelin release. Journal of Endocrinological Investigation 2010, 33:599-602.
100. Zhao T.J., Sakata I., Li R.L., Liang G., Richardson J.A., Brown M.S., et al. Ghrelin secretion stimulated by {beta}1-adrenergic receptors in cultured ghrelinoma cells and in fasted mice. Proceedings of the National Academy of Sciences of the United States of America 2010, 107:15868-15873.
101. Gagnon J., Anini Y. Insulin and norepinephrine regulate ghrelin secretion from a rat primary stomach cell culture. Endocrinology 2012, 153:3646-3656.
102. Mani B.K., Chuang J.C., Kjalarsdottir L., Sakata I., Walker A.K., Kuperman A., et al. Role of calcium and EPAC in norepinephrine-induced ghrelin secretion. Endocrinology 2014, 155:98-107.
103. Lowell B.B., Bachman E.S. Beta-Adrenergic receptors, diet-induced thermogenesis, and obesity. Journal of Biological Chemistry 2003, 278:29385-29388.
104. Pontiroli A.E., Pizzocri P., Paroni R., Folli F. Sympathetic overactivity, endothelial dysfunction, inflammation, and metabolic abnormalities cluster in grade III (World Health Organization) obesity: reversal through sustained weight loss obtained with laparoscopic adjustable gastric banding. Diabetes Care 2006, 29:2735-2738.
105. Karason K., Molgaard H., Wikstrand J., Sjostrom L. Heart rate variability in obesity and the effect of weight loss. American Journal of Cardiology 1999, 83:1242-1247.
106. Machado M.B., Velasco I.T., Scalabrini-Neto A. Gastric bypass and cardiac autonomic activity: influence of gender and age. Obesity Surgery 2009, 19:332-338.
107. Liu J., Prudom C.E., Nass R., Pezzoli S.S., Oliveri M.C., Johnson M.L., et al. Novel ghrelin assays provide evidence for independent regulation of ghrelin acylation and secretion in healthy young men. Journal of Clinical Endocrinology & Metabolism 2008, 93:1980-1987.
108. Gonzalez C.R., Vazquez M.J., Lopez M., Dieguez C. Influence of chronic undernutrition and leptin on GOAT mRNA levels in rat stomach mucosa. Journal of Molecular Endocrinology 2008, 41:415-421.
109. Mumphrey M.B., Patterson L.M., Zheng H., Berthoud H.R. Roux-en-Y gastric bypass surgery increases number but not density of CCK-, GLP-1-, 5-HT-, and neurotensin-expressing enteroendocrine cells in rats. Neurogastroenterology and Motility 2013, 25:E70-E79.