Ghrelin and the short- and long-term regulation of appetite and body weight

Physiology and Behavior

Volumn 89, Issue 1, 2006, Pages 71-84

  Cummings, David E ^a  

^a UNIVERSITY OF WASHINGTON   (United States)

Author keywords

Body weight regulation; Cachexia; Energy homeostasis; Ghrelin; Meal initiation; Meals; Obesity; Reward; Weight loss

Indexed keywords

APPETITE STIMULANT; FUEL; GASTROINTESTINAL POLYPEPTIDE; GHRELIN; INSULIN; LIPID;

ARTICLE; BLOOD BRAIN BARRIER; BODY WEIGHT; BYPASS SURGERY; CHRONIC DRUG ADMINISTRATION; DUODENUM; ENERGY EXPENDITURE; FAT INTAKE; FEEDING BEHAVIOR; FOOD INTAKE; HOMEOSTASIS; HORMONE SYNTHESIS; HUMAN; HUNGER; HYPOTHALAMUS; INCREASED APPETITE; INGESTION; INTESTINE INNERVATION; LIMBIC SYSTEM; MACRONUTRIENT; OBESITY; OSMOLARITY; POSTPRANDIAL STATE; PRADER WILLI SYNDROME; PRIORITY JOURNAL; RECEPTOR GENE; RHOMBENCEPHALON; STOMACH; STOMACH BYPASS; SYMPATHETIC NERVE; TISSUE DISTRIBUTION; WASTING SYNDROME; WEIGHT CHANGE; WEIGHT CONTROL; WEIGHT GAIN; WEIGHT REDUCTION;

EID: 33746739498     PISSN: 00319384     EISSN: None     Source Type: Journal    
DOI: 10.1016/j.physbeh.2006.05.022     Document Type: Article

References (263)

1. Ogden C.L., Carroll M.D., Curtin L.R., McDowell M.A., Tabak C.J., and Flegal K.M. Prevalence of overweight and obesity in the United States, 1999-2004. JAMA 295 (2006) 1549-1555
2. Kopelman P.G. Obesity as a medical problem. Nature 404 (2000) 635-643
3. Yanovski S.Z., and Yanovski J.A. Obesity. N Engl J Med 346 (2002) 591-602
4. Safer D.J. Diet, behavior modification, and exercise: a review of obesity treatments from a long-term perspective. South Med J 84 (1991) 1470-1474
5. Cummings D.E., and Schwartz M.W. Genetics and pathophysiology of human obesity. Annu Rev Med 54 (2003) 453-471
6. Schwartz M.W., Woods S.C., Porte D., Seeley R.J., and Baskin D.G. Central nervous system control of food intake. Nature 404 (2000) 661-671
7. Kojima M., Hosoda H., Date Y., Nakazato M., Matsuo H., and Kangawa K. Ghrelin is a growth-hormone-releasing acylated peptide from stomach. Nature 402 (1999) 656-660
8. Tschop M., Smiley D.L., and Heiman M.L. Ghrelin induces adiposity in rodents. Nature 407 (2000) 908-913
9. Wren A.M., Small C.J., Ward H.L., et al. The novel hypothalamic peptide ghrelin stimulates food intake and growth hormone secretion. Endocrinology 141 (2000) 4325-4328
10. 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 409 (2001) 194-198
11. Asakawa A., Inui A., Kaga T., et al. Ghrelin is an appetite-stimulatory signal from stomach with structural resemblance to motilin. Gastroenterology 120 (2001) 337-345
12. 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. J Clin Endocrinol Metab 86 (2001) 5992-5995
13. Neary N.M., Small C.J., Wren A.M., et al. Ghrelin increases energy intake in cancer patients with impaired appetite: acute, randomized, placebo-controlled trial. J Clin Endocrinol Metab 89 (2004) 2832-2836
14. Schmid D.A., Held K., Ising M., Uhr M., Weikel J.C., and Steiger A. Ghrelin stimulates appetite, imagination of food, GH, ACTH, and cortisol, but does not affect leptin in normal controls. Neuropsychopharmacology 30 (2005) 1187-1192
15. Laferrere B., Abraham C., Russell C.D., and Bowers C.Y. Growth hormone releasing peptide-2 (GHRP-2), like ghrelin, increases food intake in healthy men. J Clin Endocrinol Metab 90 (2005) 611-614
16. Druce M.R., Wren A.M., Park A.J., Milton J.E., Patterson M., Frost G., et al. Ghrelin increases food intake in obese as well as lean subjects. Int J Obes (Lond) 29 (2005) 1130-1136
17. Wynne K., Giannitsopoulou K., Small C.J., Patterson M., Frost G., Ghatei M.A., et al. Subcutaneous ghrelin enhances acute food intake in malnourished patients who receive maintenance peritoneal dialysis: a randomized, placebo-controlled trial. J Am Soc Nephrol 16 (2005) 2111-2118
18. Wren A.M., Small C.J., Abbott C.R., et al. Ghrelin causes hyperphagia and obesity in rats. Diabetes 50 (2001) 2540-2547
19. Xie Q.F., Wu C.X., Meng Q.Y., and Li N. Ghrelin and truncated ghrelin variant plasmid vectors administration into skeletal muscle augments long-term growth in rats. Domest Anim Endocrinol 27 (2004) 155-164
20. Nagaya N., Moriya J., Yasumura Y., et al. Effects of ghrelin administration on left ventricular function, exercise capacity, and muscle wasting in patients with chronic heart failure. Circulation 110 (2004) 3674-3679
21. Nagaya N., Itoh T., Murakami S., Oya H., Uematsu M., Miyatake K., et al. Treatment of cachexia with ghrelin in patients with COPD. Chest 128 (2005) 1187-1193
22. Howard A.D., Feighner S.D., Cully D.F., Arena J.P., Liberator P.A., Rosenblum C.I., et al. A receptor in pituitary and hypothalamus that functions in growth hormone release. Science 273 (1996) 974-977
23. Guan X.M., Yu H., Palyha O.C., McKee K.K., Feighner S.D., Sirinathsinghji D.J., et al. Distribution of mRNA encoding the GHS receptor in brain and peripheral tissues. Mol Brain Res 48 (1997) 23-29
24. Gnanapavan S., Kola B., Bustin S.A., et al. The tissue distribution of the mRNA of ghrelin and subtypes of its receptor, GHS-R, in humans. J Clin Endocrinol Metab 87 (2002) 2988-2991
25. Sun Y., Wang P., Zheng H., and Smith R.G. Ghrelin stimulation of growth hormone release and appetite is mediated through the growth hormone secretagogue receptor. Proc Natl Acad Sci U S A 101 (2004) 4679-4684
26. Zigman J.M., Jones J.E., Lee C.E., Saper C.B., and Elmquist J.K. Expression of ghrelin receptor mRNA in the rat and the mouse brain. J Comp Neurol 494 (2006) 528-548
27. van der Lely A.J., Tschop M., Heiman M.L., and Ghigo E. Biological, physiological, pathophysiological, and pharmacological aspects of ghrelin. Endocr Rev 25 (2004) 426-457
28. Halem H.A., Taylor J.E., Dong J.Z., et al. Novel analogs of ghrelin: physiological and clinical implications. Eur J Endocrinol 151 Suppl 1 (2004) S71-S75
29. Nishi Y., Hiejima H., Hosoda H., et al. Ingested medium-chain fatty acids are directly utilized for the acyl-modification of ghrelin. Endocrinology 146 (2005) 2255-2264
30. Cummings D.E., Foster-Schubert K.E., and Overduin J. Ghrelin and energy balance: focus on current controversies. Curr Drug Targets 6 (2005) 153-169
31. Chen H.Y., Trumbauer M.E., Chen A.S., et al. Orexigenic action of peripheral ghrelin is mediated by neuropeptide Y and agouti-related protein. Endocrinology 145 (2004) 2607-2612
32. Davenport A.P., Bonner T.I., Foord S.M., Harmar A.J., Neubig R.R., Pin J.P., et al. International Union of Pharmacology. LVI. Ghrelin receptor nomenclature, distribution, and function. Pharmacol Rev 57 (2005) 541-546
33. Malagon M.M., Luque R.M., Ruiz-Guerrero E., Rodriguez-Pacheco F., Garcia-Navarro S., Casanueva F.F., et al. Intracellular signaling mechanisms mediating ghrelin-stimulated growth hormone release in somatotropes. Endocrinology 144 (2003) 5372-5380
34. Wettschureck N., Moers A., Wallenwein B., Parlow A.F., Maser-Gluth C., and Offermanns S. Loss of gq/11 family G proteins in the nervous system causes pituitary somatotroph hypoplasia and dwarfism in mice. Mol Cell Biol 25 (2005) 1942-1948
35. 2+ signaling via protein kinase A and N-type channel-dependent mechanisms and cross-talk with leptin and orexin. Diabetes 52 (2003) 948-956
36. Garcia A., Alvarez C.V., Smith R.G., and Dieguez C. Regulation of Pit-1 expression by ghrelin and GHRP-6 through the GH secretagogue receptor. Mol Endocrinol 15 (2001) 1484-1495
37. Holst B., Cygankiewicz A., Jensen T.H., Ankersen M., and Schwartz T.W. High constitutive signaling of the ghrelin receptor-identification of a potent inverse agonist. Mol Endocrinol 17 (2003) 2201-2210
38. Holst B., Holliday N.D., Bach A., Elling C.E., Cox H.M., and Schwartz T.W. Common structural basis for constitutive activity of the ghrelin receptor family. J Biol Chem (2004) [Electronic publication ahead of print]
39. Bednarek M.A., Feighner S.D., Pong S.S., et al. Structure-function studies on the new growth hormone-releasing peptide, ghrelin: minimal sequence of ghrelin necessary for activation of growth hormone secretagogue receptor 1a. J Med Chem 43 (2000) 4370-4376
40. Matsumoto M., Hosoda H., Kitajima Y., et al. Structure-activity relationship of ghrelin: pharmacological study of ghrelin peptides. Biochem Biophys Res Commun 287 (2001) 142-146
41. Aydin S., Geckil H., Zengin F., Ibrahim Ozercan H., Karatas F., Turgut-Balik D., et al. Ghrelin in plants: what is the function of an appetite hormone in plants?. Peptides 27 (2006) 1597-1602
42. Palyha O.C., Feighner S.D., Tan C.P., et al. Ligand activation domain of human orphan growth hormone (GH) secretagogue receptor (GHS-R) conserved from Pufferfish to humans. Mol Endocrinol 14 (2000) 160-169
43. Riley L.G., Fox B.K., Kaiya H., Hirano T., and Grau E.G. Long-term treatment of ghrelin stimulates feeding, fat deposition, and alters the GH/IGF-I axis in the tilapia, Oreochromis mossambicus. Gen Comp Endocrinol 142 (2005) 234-240
44. Canosa L.F., Unniappan S., and Peter R.E. Periprandial changes in growth hormone release in goldfish: role of somatostatin, ghrelin, and gastrin-releasing peptide. Am J Physiol Regul Integr Comp Physiol 289 (2005) R125-R133
45. Volkoff H., Canosa L.F., Unniappan S., Cerda-Reverter J.M., Bernier N.J., Kelly S.P., et al. Neuropeptides and the control of food intake in fish. Gen Comp Endocrinol 142 (2005) 3-19
46. Tanaka-Shintani M., and Watanabe M. Distribution of ghrelin-immunoreactive cells in human gastric mucosa: comparison with that of parietal cells. J Gastroenterol 40 (2005) 345-349
47. 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 141 (2000) 4255-4261
48. Ariyasu H., Takaya K., Tagami T., et al. Stomach is a major source of circulating ghrelin, and feeding state determines plasma ghrelin-like immunoreactivity levels in humans. J Clin Endocrinol Metab 86 (2001) 4753-4758
49. 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 23 (2002) 531-536
50. Jeon T.Y., Lee S., Kim H.H., Kim Y.J., Son H.C., Kim D.H., et al. Changes in plasma ghrelin concentration immediately after gastrectomy in patients with early gastric cancer. J Clin Endocrinol Metab 89 (2004) 5392-5396
51. Dornonville de la Cour C., Lindqvist A., Egecioglu E., et al. Ghrelin treatment reverses the reduction in weight gain and body fat in gastrectomised mice. Gut 54 (2005) 907-913
52. Krsek M., Rosicka M., Haluzik M., Svobodova J., Kotrlikova E., Justova V., et al. Plasma ghrelin levels in patients with short bowel syndrome. Endocr Res 28 (2002) 27-33
53. Prado C.L., Pugh-Bernard A.E., Elghazi L., Sosa-Pineda B., and Sussel L. Ghrelin cells replace insulin-producing beta cells in two mouse models of pancreas development. Proc Natl Acad Sci U S A 101 (2004) 2924-2929
54. Wierup N., Yang S., McEvilly R.J., Mulder H., and Sundler F. Ghrelin is expressed in a novel endocrine cell type in developing rat islets and inhibits insulin secretion from INS-1 (832/13) cells. J Histochem Cytochem 52 (2004) 301-310
55. Wierup N., and Sundler F. Ultrastructure of islet ghrelin cells in the human fetus. Cell Tissue Res 319 (2005) 423-428
56. Heller R.S., Jenny M., Collombat P., Mansouri A., Tomasetto C., Madsen O.D., et al. Genetic determinants of pancreatic epsilon-cell development. Dev Biol 286 (2005) 217-224
57. Chanoine J.P., and Wong A.C. Ghrelin gene expression is markedly higher in fetal pancreas compared with fetal stomach: effect of maternal fasting. Endocrinology 145 (2004) 3813-3820
58. 2+ signaling in beta-cells: implication in the glycemic control in rodents. Diabetes 53 (2004) 3142-3151
59. Faulconbridge L.F., Cummings D.E., Kaplan J.M., and Grill H.J. Hyperphagic effects of brainstem ghrelin administration. Diabetes 52 (2003) 2260-2265
60. Keen-Rhinehart E., and Bartness T.J. Peripheral ghrelin injections stimulate food intake, foraging and food hoarding in Siberian hamsters. Am J Physiol Regul Integr Comp Physiol 288 (2004) R716-R722
61. Davidson T.L., Kanoski S.E., Tracy A.L., Walls E.K., Clegg D., and Benoit S.C. The interoceptive cue properties of ghrelin generalize to cues produced by food deprivation. Peptides 26 (2005) 1602-1610
62. Masuda Y., Tanaka T., Inomata N., Ohnuma N., Tanaka S., Itoh Z., et al. Ghrelin stimulates gastric acid secretion and motility in rats. Biochem Biophys Res Commun 276 (2000) 905-908
63. Trudel L., Tomasetto C., Rio M.C., Bouin M., Plourde V., Eberling P., et al. Ghrelin/motilin-related peptide is a potent prokinetic to reverse gastric postoperative ileus in rat. Am J Physiol Gastrointest Liver Physiol 282 (2002) G948-G952
64. Dornonville de la Cour C., Lindstrom E., Norlen P., and Hakanson R. Ghrelin stimulates gastric emptying but is without effect on acid secretion and gastric endocrine cells. Regul Pept 120 (2004) 23-32
65. Trudel L., Bouin M., Tomasetto C., Eberling P., St-Pierre S., Bannon P., et al. Two new peptides to improve post-operative gastric ileus in dog. Peptides 24 (2003) 531-534
66. De Winter B.Y., De Man J.G., Seerden T.C., Depoortere I., Herman A.G., Peeters T.L., et al. Effect of ghrelin and growth hormone-releasing peptide 6 on septic ileus in mice. Neurogastroenterol Motil 16 (2004) 439-446
67. Asakawa A., Inui A., Kaga T., Katsuura G., Fujimiya M., Fujino M.A., et al. Antagonism of ghrelin receptor reduces food intake and body weight gain in mice. Gut 52 (2003) 947-952
68. Date Y., Nakazato M., Murakami N., Kojima M., Kangawa K., and Matsukura S. Ghrelin acts in the central nervous system to stimulate gastric acid secretion. Biochem Biophys Res Commun 280 (2001) 904-907
69. Brzozowski T., Konturek P.C., Konturek S.J., et al. Exogenous and endogenous ghrelin in gastroprotection against stress-induced gastric damage. Regul Pept 120 (2004) 39-51
70. Sato N., Kanai S., Takano S., Kurosawa M., Funakoshi A., and Miyasaka K. Central administration of ghrelin stimulates pancreatic exocrine secretion via the vagus in conscious rats. Jpn J Physiol 53 (2003) 443-449
71. Witek B., Ochwanowska E., Rafay J., Kolataj A., Chrenek P., Suvegova K., et al. Effect of ghrelin on activities of some lysosomal hydrolases in rabbits. Neuro Endocrinol Lett 26 (2005) 397-400
72. Cummings D.E., and Shannon M.H. Roles for ghrelin in the regulation of appetite and body weight. Arch Surg 138 (2003) 389-396
73. Lu X.Y., Shieh K.R., Kabbaj M., Barsh G.S., Akil H., and Watson S.J. Diurnal rhythm of agouti-related protein and its relation to corticosterone and food intake. Endocrinology 143 (2002) 3905-3915
74. Crowley W.R., Ramoz G., Keefe K.A., Torto R., Kalra S.P., and Hanson G.R. Differential effects of methamphetamine on expression of neuropeptide Y mRNA in hypothalamus and on serum leptin and ghrelin concentrations in ad libitum-fed and schedule-fed rats. Neuroscience 132 (2005) 167-173
75. Shiiya T., Nakazato M., Mizuta M., et al. Plasma ghrelin levels in lean and obese humans and the effect of glucose on ghrelin secretion. J Clin Endocrinol Metab 87 (2002) 240-244
76. Williams D.L., Cummings D.E., Grill H.J., and Kaplan J.M. Meal-related ghrelin suppression requires postgastric feedback. Endocrinology 144 (2003) 2765-2767
77. Tschop M., Wawarta R., Riepl R.L., Friedrich S., Bidlingmaier M., Landgraf R., et al. Post-prandial decrease of circulating human ghrelin levels. J Endocrinol Invest 24 (2001) RC19-RC21
78. Cummings D.E., Purnell J.Q., Frayo R.S., Schmidova K., Wisse B.E., and Weigle D.S. A preprandial rise in plasma ghrelin levels suggests a role in meal initiation in humans. Diabetes 50 (2001) 1714-1719
79. 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. N Engl J Med 346 (2002) 1623-1630
80. Weigle D.S., Cummings D.E., Newby P.D., Breen P.A., Frayo R.S., Matthys C.C., et al. Roles of leptin and ghrelin in the loss of body weight caused by a low fat, high carbohydrate diet. J Clin Endocrinol Metab 88 (2003) 1577-1586
81. Sugino T., Yamaura J., Yamagishi M., et al. A transient surge of ghrelin secretion before feeding is modified by different feeding regimens in sheep. Biochem Biophys Res Commun 298 (2002) 785-788
82. Miura H., Tsuchiya N., Sasaki I., Kikuchi M., Kojima M., Kangawa K., et al. Changes in plasma ghrelin and growth hormone concentrations in mature Holstein cows and three-month-old calves. J Anim Sci 82 (2004) 1329-1333
83. Yokoyama M., Nakahara K., Kojima M., Hosoda H., Kangawa K., and Murakami N. Influencing the between-feeding and endocrine responses of plasma ghrelin in healthy dogs. Eur J Endocrinol 152 (2005) 155-160
84. Drazen D.L., Vahl T.P., D'Alessio D.A., Seeley R.J., and Woods S.C. Effects of a fixed meal pattern on ghrelin secretion: evidence for a learned response independent of nutrient status. Endocrinology 147 (2006) 23-30
85. Cummings D.E., Frayo R.S., Marmonier C., Aubert R., and Chapelot D. Plasma ghrelin levels and hunger scores in humans initiating meals voluntarily without time- and food-related cues. Am J Physiol Endocrinol Metab 287 (2004) E297-E304
86. Blom W.A., Stafleu A., de Graaf C., Kok F.J., Schaafsma G., and Hendriks H.F. Ghrelin response to carbohydrate-enriched breakfast is related to insulin. Am J Clin Nutr 81 (2005) 367-375
87. Murakami N., Hayashida T., Kuroiwa T., Nakahara K., Ida T., Mondal M.S., et al. Role for central ghrelin in food intake and secretion profile of stomach ghrelin in rats. J Endocrinol 174 (2002) 283-288
88. Sanchez J., Oliver P., Pico C., and Palou A. Diurnal rhythms of leptin and ghrelin in the systemic circulation and in the gastric mucosa are related to food intake in rats. Pflugers Arch 448 (2004) 500-506
89. Bodosi B., Gardi J., Hajdu I., Szentirmai E., Obal Jr. F., and Krueger J.M. Rhythms of ghrelin, leptin, and sleep in rats: effects of the normal diurnal cycle, restricted feeding, and sleep deprivation. Am J Physiol Regul Integr Comp Physiol 287 (2004) R1071-R1079
90. Tolle V., Bassant M.H., Zizzari P., Poindessous-Jazat F., Tomasetto C., Epelbaum J., et al. Ultradian rhythmicity of ghrelin secretion in relation with GH, feeding behavior, and sleep-wake patterns in rats. Endocrinology 143 (2002) 1353-1361
91. Callahan H.S., Cummings D.E., Pepe M.S., Breen P.A., Matthys C.C., and Weigle D.S. Postprandial suppression of plasma ghrelin level is proportional to ingested caloric load but does not predict intermeal interval in humans. J Clin Endocrinol Metab 89 (2004) 1319-1324
92. Erdmann J., Topsch R., Lippl F., Gussmann P., and Schusdziarra V. Postprandial response of plasma ghrelin levels to various test meals in relation to food intake, plasma insulin, and glucose. J Clin Endocrinol Metab 89 (2004) 3048-3054
93. Korbonits M., Lecoeur C., Froguel P., Grossman A.B., and Gueorguiev M. Characteristics of the ghrelin receptor in relation to eating behaviour. 5th International Symposium on GH Secretagogues, Portofino-Vetta, Italy (2004)
94. Overduin J., Frayo R.S., Grill H.J., Kaplan J.M., and Cummings D.E. Role of the duodenum and macronutrient type in ghrelin regulation. Endocrinology (2004) [Electronic publication ahead of print]
95. Foster-Schubert K.E., Weigle D.S., Callahan H.S., and Cummings D.E. Lipids suppress human plasma ghrelin levels less effectively than do carbohydrates or proteins. Endocrine Society 86th Annual Meeting, New Orleans, LA (2004)
96. Monteleone P., Bencivenga R., Longobardi N., Serritella C., and Maj M. Differential responses of circulating ghrelin to high-fat or high-carbohydrate meal in healthy women. J Clin Endocrinol Metab 88 (2003) 5510-5514
97. Lejeune M.P., Westerterp K.R., Adam T.C., Luscombe-Marsh N.D., and Westerterp-Plantenga M.S. Ghrelin and glucagon-like peptide 1 concentrations, 24-h satiety, and energy and substrate metabolism during a high-protein diet and measured in a respiration chamber. Am J Clin Nutr 83 (2006) 89-94
98. Gelling R.W., Overduin J., Morrison C.D., Morton G.J., Frayo R.S., Cummings D.E., et al. Effect of uncontrolled diabetes on plasma ghrelin concentrations and ghrelin-induced feeding. Endocrinology 145 (2004) 4575-4582
99. Williams D.L., Grill H.J., Cummings D.E., and Kaplan J.M. Vagotomy dissociates short- and long-term controls of circulating ghrelin. Endocrinology 144 (2003) 5184-5187
100. Overduin J., Frayo R.S., Baskin D.G., and Cummings D.E. Peripheral nervous contribution to nutrient-related ghrelin regulation in rats. Society for Neuroscience 34th Annual Meeting, San Diego, CA (2004)
101. Arosio M., Ronchi C.L., Beck-Peccoz P., Gebbia C., Giavoli C., Cappiello V., et al. Effects of modified sham feeding on ghrelin levels in healthy human subjects. J Clin Endocrinol Metab 89 (2004) 5101-5104
102. Heath R.B., Jones R., Frayn K.N., and Robertson M.D. Vagal stimulation exaggerates the inhibitory ghrelin response to oral fat in humans. J Endocrinol 180 (2004) 273-281
103. Nedvidkova J., Krykorkova I., Bartak V., Papezova H., Gold P.W., Alesci S., et al. Loss of meal-induced decrease in plasma ghrelin levels in patients with anorexia nervosa. J Clin Endocrinol Metab 88 (2003) 1678-1682
104. Erdmann J., Lippl F., and Schusdziarra V. Differential effect of protein and fat on plasma ghrelin levels in man. Regul Pept 116 (2003) 101-107
105. Teff K.L., Elliott S.S., Tschop M., et al. Dietary fructose reduces circulating insulin and leptin, attenuates postprandial suppression of ghrelin, and increases triglycerides in women. J Clin Endocrinol Metab 89 (2004) 2963-2972
106. English P.J., Ghatei M.A., Malik I.A., Bloom S.R., and Wilding J.P. Food fails to suppress ghrelin levels in obese humans. J Clin Endocrinol Metab 87 (2002) 2984
107. Morpurgo P.S., Resnik M., Agosti F., Cappiello V., Sartorio A., and Spada A. Ghrelin secretion in severely obese subjects before and after a 3-week integrated body mass reduction program. J Endocrinol Invest 26 (2003) 723-727
108. Korner J., Bessler M., Cirilo L., Conwell I.M., Daud A., Restuccia N.L., et al. Effects of roux-en-Y gastric bypass surgery on fasting and postprandial concentrations of plasma ghrelin, PYY and insulin. J Clin Endocrinol Metab 90 (2005) 359-365
109. Tentolouris N., Kokkinos A., Tsigos C., Kyriaki D., Doupis J., Raptis S.A., et al. Differential effects of high-fat and high-carbohydrate content isoenergetic meals on plasma active ghrelin concentrations in lean and obese women. Horm Metab Res 36 (2004) 559-563
110. Perreault M., Istrate N., Wang L., Nichols A.J., Tozzo E., and Stricker-Krongrad A. Resistance to the orexigenic effect of ghrelin in dietary-induced obesity in mice: reversal upon weight loss. Int J Obes Relat Metab Disord 28 (2004) 879-885
111. Morinigo R., Casamitjana R., Moize V., Lacy A.M., Delgado S., Gomis R., et al. Short-term effects of gastric bypass surgery on circulating ghrelin levels. Obes Res 12 (2004) 1108-1116
112. Tschop M., Weyer C., Tataranni P.A., Devanarayan V., Ravussin E., and Heiman M.L. Circulating ghrelin levels are decreased in human obesity. Diabetes 50 (2001) 707-709
113. Cummings D.E., Clement K., Purnell J.Q., Vaisse C., Foster K.E., Frayo R.S., et al. Elevated plasma ghrelin levels in Prader-Willi syndrome. Nat Med 8 (2002) 643-644
114. Tolle V., Kadem M., Bluet-Pajot M.T., et al. Balance in ghrelin and leptin plasma levels in anorexia nervosa patients and constitutionally thin women. J Clin Endocrinol Metab 88 (2003) 109-116
115. Otto B., Cuntz U., Fruehauf E., et al. Weight gain decreases elevated plasma ghrelin concentrations of patients with anorexia nervosa. Eur J Endocrinol 145 (2001) 669-673
116. Haqq A.M., Farooqi I.S., O'Rahilly S., Stadler D.D., Rosenfeld R.G., Pratt K.L., et al. Serum ghrelin levels are inversely correlated with body mass index, age, and insulin concentrations in normal children and are markedly increased in Prader-Willi syndrome. J Clin Endocrinol Metab 88 (2003) 174-178
117. Wisse B.E., Frayo R.S., Schwartz M.W., and Cummings D.E. Reversal of cancer anorexia by blockade of central melanocortin receptors in rats. Endocrinology 142 (2001) 3292-3301
118. 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. Clin Endocrinol (Oxf) 56 (2002) 203-206
119. Leidy H.J., Gardner J.K., Frye B.R., Snook M.L., Schuchert M.K., Richard E.L., et al. Circulating ghrelin is sensitive to changes in body weight during a diet and exercise program in normal-weight young women. J Clin Endocrinol Metab 89 (2004) 2659-2664
120. Zahorska-Markiewicz B., Mizia-Stec K., Olszanecka-Glinianowicz M., and Janowska J. Effect of weight reduction on serum ghrelin and TNFalpha concentrations in obese women. Eur J Intern Med 15 (2004) 172-175
121. Shimizu Y., Nagaya N., Isobe T., Imazu M., Okumura H., Hosoda H., et al. Increased plasma ghrelin level in lung cancer cachexia. Clin Cancer Res 9 (2003) 774-778
122. Hanada T., Toshinai K., Kajimura N., et al. Anti-cachectic effect of ghrelin in nude mice bearing human melanoma cells. Biochem Biophys Res Commun 301 (2003) 275-279
123. Hanada T., Toshinai K., Date Y., Kajimura N., Tsukada T., Hayashi Y., et al. Upregulation of ghrelin expression in cachectic nude mice bearing human melanoma cells. Metabolism 53 (2004) 84-88
124. Garcia J.M., Garcia-Touza M., Hijazi R.A., Taffet G., Epner D., Mann D., et al. Active ghrelin levels and active-to-total ghrelin ratio in cancer-induced cachexia. J Clin Endocrinol Metab 90 (2005) 2920-2926
125. Popovic V., Svetel M., Djurovic M., et al. Circulating and cerebrospinal fluid ghrelin and leptin: potential role in altered body weight in Huntington's disease. Eur J Endocrinol 151 (2004) 451-455
126. Rigamonti A.E., Pincelli A.I., Corra B., et al. Plasma ghrelin concentrations in elderly subjects: comparison with anorexic and obese patients. J Endocrinol 175 (2002) R1-R5
127. Nakai Y., Hosoda H., Nin K., Ooya C., Hayashi H., Akamizu T., et al. Plasma levels of active form of ghrelin during oral glucose tolerance test in patients with anorexia nervosa. Eur J Endocrinol 149 (2003) R1-R3
128. Tanaka M., Nakahara T., Kojima S., et al. Effect of nutritional rehabilitation on circulating ghrelin and growth hormone levels in patients with anorexia nervosa. Regul Pept 122 (2004) 163-168
129. Misra M., Miller K.K., Herzog D.B., Ramaswamy K., Aggarwal A., Almazan C., et al. Growth hormone and ghrelin responses to an oral glucose load in adolescent girls with anorexia nervosa and controls. J Clin Endocrinol Metab 89 (2004) 1605-1612
130. Stock S., Leichner P., Wong A.C., Ghatei M.A., Kieffer T.J., Bloom S.R., et al. Ghrelin, PYY, GIP and hunger responses to a mixed meal in anorexic, obese and control female adolescents. J Clin Endocrinol Metab 90 (2005) 2161-2168
131. Kojima S., Nakahara T., Nagai N., et al. Altered ghrelin and peptide YY responses to meals in bulimia nervosa. Clin Endocrinol (Oxf) 62 (2005) 74-78
132. Misra M., Miller K.K., Kuo K., Griffin K., Stewart V., Hunter E., et al. Secretory dynamics of ghrelin in adolescent girls with anorexia nervosa and healthy adolescents. Am J Physiol Endocrinol Metab 289 (2005) E347-E356
133. Otto B., Tschop M., Fruhauf E., Heldwein W., Fichter M., Otto C., et al. Postprandial ghrelin release in anorectic patients before and after weight gain. Psychoneuroendocrinology 30 (2005) 577-581
134. Nagaya N., Uematsu M., Kojima M., et al. Elevated circulating level of ghrelin in cachexia associated with chronic heart failure: relationships between ghrelin and anabolic/catabolic factors. Circulation 104 (2001) 2034-2038
135. Itoh T., Nagaya N., Yoshikawa M., et al. Elevated plasma ghrelin level in underweight patients with chronic obstructive pulmonary disease. Am J Respir Crit Care Med 170 (2004) 879-882
136. Tacke F., Brabant G., Kruck E., Horn R., Schoffski P., Hecker H., et al. Ghrelin in chronic liver disease. J Hepatol 38 (2003) 447-454
137. Marchesini G., Bianchi G., Lucidi P., Villanova N., Zoli M., and De Feo P. Plasma ghrelin concentrations, food intake, and anorexia in liver failure. J Clin Endocrinol Metab 89 (2004) 2136-2141
138. Yoshimoto A., Mori K., Sugawara A., et al. Plasma ghrelin and desacyl ghrelin concentrations in renal failure. J Am Soc Nephrol 13 (2002) 2748-2752
139. Perez-Fontan M., Cordido F., Rodriguez-Carmona A., Peteiro J., Garcia-Naveiro R., and Garcia-Buela J. Plasma ghrelin levels in patients undergoing haemodialysis and peritoneal dialysis. Nephrol Dial Transplant 19 (2004) 2095-2100
140. Chang C.C., Hung C.H., Yen C.S., Hwang K.L., and Lin C.Y. The relationship of plasma ghrelin level to energy regulation, feeding and left ventricular function in non-diabetic haemodialysis patients. Nephrol Dial Transplant 20 (2005) 2172-2177
141. Cummings D.E., and Overduin J. Circulating ghrelin levels in pathophysiologcial conditions. In: Ghigo E. (Ed). Ghrelin (2004), Kluwer Academic Publishers, Boston 207-224
142. Kim M.S., Yoon C.Y., Park K.H., Shin C.S., Park K.S., Kim S.Y., et al. Changes in ghrelin and ghrelin receptor expression according to feeding status. Neuroreport 14 (2003) 1317-1320
143. Nogueiras R., Tovar S., Mitchell S.E., Rayner D.V., Archer Z.A., Dieguez C., et al. Regulation of growth hormone secretagogue receptor gene expression in the arcuate nuclei of the rat by leptin and ghrelin. Diabetes 53 (2004) 2552-2558
144. Tups A., Helwig M., Khorooshi R.M., Archer Z.A., Klingenspor M., and Mercer J.G. Circulating ghrelin levels and central ghrelin receptor expression are elevated in response to food deprivation in a seasonal mammal (Phodopus sungorus). J Neuroendocrinol 16 (2004) 922-928
145. Kurose Y., Iqbal J., Rao A., Murata Y., Hasegawa Y., Terashima Y., et al. Changes in expression of the genes for the leptin receptor and the growth hormone-releasing peptide/ghrelin receptor in the hypothalamic arcuate nucleus with long-term manipulation of adiposity by dietary means. J Neuroendocrinol 17 (2005) 331-340
146. Tung Y.C., Hewson A.K., Carter R.N., and Dickson S.L. Central responsiveness to a ghrelin mimetic (GHRP-6) is rapidly altered by acute changes in nutritional status in rats. J Neuroendocrinol 17 (2005) 387-393
147. Williams DL, Grill HJ, Cummings DE, Kaplan JM. Overfeeding-induced weight gain suppresses plasma ghrelin levels in rats. J Endocrinol Invest in press.
148. Otukonyong E.E., Dube M.G., Torto R., Kalra P.S., and Kalra S.P. High-fat diet-induced ultradian leptin and insulin hypersecretion are absent in obesity-resistant rats. Obes Res 13 (2005) 991-999
149. Robertson M.D., Henderson R.A., Vist G.E., and Rumsey R.D. Plasma ghrelin response following a period of acute overfeeding in normal weight men. Int J Obes Relat Metab Disord 28 (2004) 727-733
150. Moesgaard S.G., Ahren B., Carr R.D., Gram D.X., Brand C.L., and Sundler F. Effects of high-fat feeding and fasting on ghrelin expression in the mouse stomach. Regul Pept 120 (2004) 261-267
151. Beck B., Musse N., and Stricker-Krongrad A. Ghrelin, macronutrient intake and dietary preferences in Long-Evans rats. Biochem Biophys Res Commun 292 (2002) 1031-1035
152. Otto B., Tschop M., Heldwein W., Pfeiffer A.F., and Diederich S. Endogenous and exogenous glucocorticoids decrease plasma ghrelin in humans. Eur J Endocrinol 151 (2004) 113-117
153. Capristo E., Farnetti S., Mingrone G., Certo M., Greco A.V., Addolorato G., et al. Reduced plasma ghrelin concentration in celiac disease after gluten-free diet treatment. Scand J Gastroenterol 40 (2005) 430-436
154. Soriano-Guillen L., Barrios V., Campos-Barros A., and Argente J. Ghrelin levels in obesity and anorexia nervosa: effect of weight reduction or recuperation. J Pediatr 144 (2004) 36-42
155. Foster K.E., McTiernan A., Frayo R.S., Rajan B., and Cummings D.E. Changes in human plasma ghrelin and leptin levels correlate negatively with each other during aerobic exercise training. Endocrine Society 84th Annual Meeting, San Francisco, CA (2002)
156. Dall R., Kanaley J., Hansen T.K., Moller N., Christiansen J.S., Hosoda H., et al. Plasma ghrelin levels during exercise in healthy subjects and in growth hormone-deficient patients. Eur J Endocrinol 147 (2002) 65-70
157. Jorgensen J.O., Krag M., Kanaley J., Moller J., Hansen T.K., Moller N., et al. Exercise, hormones, and body temperature. regulation and action of GH during exercise. J Endocrinol Invest 26 (2003) 838-842
158. Kraemer R.R., Durand R.J., Acevedo E.O., Johnson L.G., Kraemer G.R., Hebert E.P., et al. Rigorous running increases growth hormone and insulin-like growth factor-I without altering ghrelin. Exp Biol Med (Maywood) 229 (2004) 240-246
159. Schmidt A., Maier C., Schaller G., Nowotny P., Bayerle-Eder M., Buranyi
160. De Souza M.J., Leidy H.J., O'Donnell E., Lasley B., and Williams N.I. Fasting ghrelin levels in physically active women: relationship with menstrual disturbances and metabolic hormones. J Clin Endocrinol Metab 89 (2004) 3536-3542
161. Zoladz J.A., Konturek S.J., Duda K., Majerczak J., Sliwowski Z., Grandys M., et al. Effect of moderate incremental exercise, performed in fed and fasted state on cardio-respiratory variables and leptin and ghrelin concentrations in young healthy men. J Physiol Pharmacol 56 (2005) 63-85
162. Bush E.N., Droz B., Shapiro R., et al. Effects of low-calorie or low-fat diet-induced weight loss on plasma ghrelin levels. Program of the 85th Annual Meeting of the Endocrine Society, Philadelphia, PA (2003)
163. Qader S.S., Salehi A., Hakanson R., Lundquist I., and Ekelund M. Long-term infusion of nutrients (total parenteral nutrition) suppresses circulating ghrelin in food-deprived rats. Regul Pept 131 (2005) 82-88
164. McLaughlin T., Abbasi F., Lamendola C., Frayo R.S., and Cummings D.E. Plasma ghrelin concentrations are decreased in insulin-resistant obese adults relative to equally obese insulin-sensitive controls. J Clin Endocrinol Metab 89 (2004) 1630-1635
165. Purnell J.Q., Weigle D.S., Breen P., and Cummings D.E. Ghrelin levels correlate with insulin levels, insulin resistance, and high-density lipoprotein cholesterol, but not with gender, menopausal status, or cortisol levels in humans. J Clin Endocrinol Metab 88 (2003) 5747-5752
166. Cummings D.E., and Foster K.E. Ghrelin-leptin tango in body-weight regulation. Gastroenterology 124 (2003) 1532-1535
167. Chanoine J.P., Yeung L.P., Wong A.C., and Birmingham C.L. Immunoreactive ghrelin in human cord blood: relation to anthropometry, leptin, and growth hormone. J Pediatr Gastroenterol Nutr 35 (2002) 282-286
168. Soriano-Guillen L., Barrios V., Chowen J.A., Sanchez I., Vila S., Quero J., et al. Ghrelin levels from fetal life through early adulthood: relationship with endocrine and metabolic and anthropometric measures. J Pediatr 144 (2004) 30-35
169. Onal E.E., Cinaz P., Atalay Y., et al. Umbilical cord ghrelin concentrations in small- and appropriate-for-gestational age newborn infants: relationship to anthropometric markers. J Endocrinol 180 (2004) 267-271
170. Cortelazzi D., Cappiello V., Morpurgo P.S., Ronzoni S., Nobile De Santis M.S., Cetin I., et al. Circulating levels of ghrelin in human fetuses. Eur J Endocrinol 149 (2003) 111-116
171. Farquhar J., Heiman M., Wong A.C., Wach R., Chessex P., and Chanoine J.P. Elevated umbilical cord ghrelin concentrations in small for gestational age neonates. J Clin Endocrinol Metab 88 (2003) 4324-4327
172. Kitamura S., Yokota I., Hosoda H., Kotani Y., Matsuda J., Naito E., et al. Ghrelin concentration in cord and neonatal blood: relation to fetal growth and energy balance. J Clin Endocrinol Metab 88 (2003) 5473-5477
173. Yokota I., Kitamura S., Hosoda H., Kotani Y., and Kangawa K. Concentration of the n-octanoylated active form of ghrelin in fetal and neonatal circulation. Endocr J 52 (2005) 271-276
174. Ng P.C., Lee C.H., Lam C.W., Chan I.H., Wong E., and Fok T.F. Ghrelin in preterm and term newborns: relation to anthropometry, leptin and insulin. Clin Endocrinol (Oxf) 63 (2005) 217-222
175. Desai M., Gayle D., Babu J., and Ross M.G. Programmed obesity in intrauterine growth restricted newborns: modulation by newborn nutrition. Am J Physiol Regul Integr Comp Physiol (2004) [Electronic publication ahead of print]
176. Iniguez G., Ong K., Pena V., Avila A., Dunger D., and Mericq V. Fasting and post-glucose ghrelin levels in SGA infants: relationships with size and weight gain at one year of age. J Clin Endocrinol Metab 87 (2002) 5830-5833
177. Soto I.N., and Mericq G.V. Fetal growth restriction and insulin resistance. New findings and review of the literature. Rev Med Chil 133 (2005) 97-104
178. Gohlke B.C., Huber A., Hecher K., Fimmers R., Bartmann P., and Roth C.L. Fetal insulin-like growth factor (IGF)-I, IGF-II, and ghrelin in association with birth weight and postnatal growth in monozygotic twins with discordant growth. J Clin Endocrinol Metab 90 (2005) 2270-2274
179. James R.J., Drewett R.F., and Cheetham T.D. Low cord ghrelin levels in term infants are associated with slow weight gain over the first 3 months of life. J Clin Endocrinol Metab 89 (2004) 3847-3850
180. Astrup A., Buemann B., Flint A., and Raben A. Low-fat diets and energy balance: how does the evidence stand in 2002. Proc Nutr Soc 61 (2002) 299-309
181. Reinehr T., Roth C.L., Alexy U., Kersting M., Kiess W., and Andler W. Ghrelin levels before and after reduction of overweight due to a low-fat high-carbohydrate diet in obese children and adolescents. Int J Obes (Lond) 29 (2005) 362-368
182. De Luca M., Segato G., Busetto L., et al. Progress in implantable gastric stimulation: summary of results of the European multi-center study. Obes Surg 14 Suppl 1 (2004) S33-S39
183. Mun E.C., Blackburn G.L., and Matthews J.B. Current status of medical and surgical therapy for obesity. Gastroenterology 120 (2001) 669-681
184. Cummings D.E., and Shannon M.H. Ghrelin and gastric bypass: is there a hormonal contribution to surgical weight loss?. J Clin Endocrinol Metab 88 (2003) 2999-3002
185. Geloneze B., Tambascia M.A., Pilla V.F., Geloneze S.R., Repetto E.M., and Pareja J.C. Ghrelin: a gut-brain hormone: effect of gastric bypass surgery. Obes Surg 13 (2003) 17-22
186. Fruhbeck G., Diez Caballero A., and Gil M.J. Fundus functionality and ghrelin concentrations after bariatric surgery. N Engl J Med 350 (2004) 308-309
187. Faraj M., Havel P.J., Phelis S., Blank D., Sniderman A.D., and Cianflone K. Plasma acylation-stimulating protein, adiponectin, leptin, and ghrelin before and after weight loss induced by gastric bypass surgery in morbidly obese subjects. J Clin Endocrinol Metab 88 (2003) 1594-1602
188. Leonetti F., Silecchia G., Iacobellis G., et al. Different plasma ghrelin levels after laparoscopic gastric bypass and adjustable gastric banding in morbid obese subjects. J Clin Endocrinol Metab 88 (2003) 4227-4231
189. Tritos N.A., Mun E., Bertkau A., Grayson R., Maratos-Flier E., and Goldfine A. Serum ghrelin levels in response to glucose load in obese subjects post-gastric bypass surgery. Obes Res 11 (2003) 919-924
190. Fruhbeck G., Diez-Caballero A., Gil M.J., Montero I., Gomez-Ambrosi J., Salvador J., et al. The decrease in plasma ghrelin concentrations following bariatric surgery depends on the functional integrity of the fundus. Obes Surg 14 (2004) 606-612
191. Stoeckli R., Chanda R., Langer I., and Keller U. Changes of body weight and plasma ghrelin levels after gastric banding and gastric bypass. Obes Res 12 (2004) 346-350
192. Lin E., Gletsu N., Fugate K., et al. The effects of gastric surgery on systemic ghrelin levels in the morbidly obese. Arch Surg 139 (2004) 780-784
193. Fruhbeck G., Rotellar F., Hernandez-Lizoain J.L., Gil M.J., Gomez-Ambrosi J., Salvador J., et al. Fasting plasma ghrelin concentrations 6 months after gastric bypass are not determined by weight loss or changes in insulinemia. Obes Surg 14 (2004) 1208-1215
194. Stylopoulos N., Davis P., Pettit J.D., Rattner D.W., and Kaplan L.M. Changes in serum ghrelin predict weight loss after Roux-en-Y gastric bypass in rats. Surg Endosc 19 (2005) 942-946
195. Holdstock C., Engstrom B.E., Ohrvall M., Lind L., Sundbom M., and Karlsson F.A. Ghrelin and adipose tissue regulatory peptides: effect of gastric bypass surgery in obese humans. J Clin Endocrinol Metab 88 (2003) 3177-3183
196. Vendrell J., Broch M., Vilarrasa N., Molina A., Gomez J.M., Gutierrez C., et al. Resistin, adiponectin, ghrelin, leptin, and proinflammatory cytokines: relationships in obesity. Obes Res 12 (2004) 962-971
197. Cummings D.E., Overduin J., and Foster-Schubert K.E. Gastric bypass for obesity: mechanisms of weight loss and diabetes resolution. J Clin Endocrinol Metab 89 (2004) 2608-2615
198. Willesen M.G., Kristensen P., and Romer J. Co-localization of growth hormone secretagogue receptor and NPY mRNA in the arcuate nucleus of the rat. Neuroendocrinology 70 (1999) 306-316
199. Mondal M.S., Date Y., Yamaguchi H., Toshinai K., Tsuruta T., Kangawa K., et al. Identification of ghrelin and its receptor in neurons of the rat arcuate nucleus. Regul Pept 126 (2005) 55-59
200. Cowley M.A., Smith R.G., Diano S., et al. The distribution and mechanism of action of ghrelin in the CNS demonstrates a novel hypothalamic circuit regulating energy homeostasis. Neuron 37 (2003) 649-661
201. van den Top M., Lee K., Whyment A.D., Blanks A.M., and Spanswick D. Orexigen-sensitive NPY/AgRP pacemaker neurons in the hypothalamic arcuate nucleus. Nat Neurosci 7 (2004) 493-494
202. Dickson S.L., and Luckman S.M. Induction of c-fos messenger ribonucleic acid in neuropeptide Y and growth hormone (GH)-releasing factor neurons in the rat arcuate nucleus following systemic injection of the GH secretagoguem, GH-releasing peptide-6. Endocrinology 138 (1997) 771-777
203. Kamegai J., Tamura H., Shimizu T., Ishii S., Sugihara H., and Wakabayashi I. Central effect of ghrelin, an endogenous growth hormone secretagogue, on hypothalamic peptide gene expression. Endocrinology 141 (2000) 4797-4800
204. Kamegai J., Tamura H., Shimizu T., Ishii S., Sugihara H., and Wakabayashi I. Chronic central infusion of ghrelin increases hypothalamic neuropeptide Y and Agouti-related protein mRNA levels and body weight in rats. Diabetes 50 (2001) 2438-2443
205. Shintani M., Ogawa Y., Ebihara K., et al. Ghrelin, an endogenous growth hormone secretagogue, is a novel orexigenic peptide that antagonizes leptin action through the activation of hypothalamic neuropeptide Y/Y1 receptor pathway. Diabetes 50 (2001) 227-232
206. Wang L., Saint-Pierre D.H., and Tache Y. Peripheral ghrelin selectively increases Fos expression in neuropeptide Y-synthesizing neurons in mouse hypothalamic arcuate nucleus. Neurosci Lett 325 (2002) 47-51
207. Lawrence C.B., Snape A.C., Baudoin F.M., and Luckman S.M. Acute central ghrelin and GH secretagogues induce feeding and activate brain appetite centers. Endocrinology 143 (2002) 155-162
208. Seoane L.M., Lopez M., Tovar S., Casanueva F.F., Senaris R., and Dieguez C. Agouti-related peptide, neuropeptide Y, and somatostatin-producing neurons are targets for ghrelin actions in the rat hypothalamus. Endocrinology 144 (2003) 544-551
209. Hewson A.K., and Dickson S.L. Systemic administration of ghrelin induces Fos and Egr-1 proteins in the hypothalamic arcuate nucleus of fasted and fed rats. J Neuroendocrinol 12 (2000) 1047-1049
210. Date Y., Murakami N., Toshinai K., Matsukura S., Niijima A., Matsuo H., et al. The role of the gastric afferent vagal nerve in ghrelin-induced feeding and growth hormone secretion in rats. Gastroenterology 123 (2002) 1120-1128
211. Melis M.R., Mascia M.S., Succu S., Torsello A., Muller E.E., Deghenghi R., et al. Ghrelin injected into the paraventricular nucleus of the hypothalamus of male rats induces feeding but not penile erection. Neurosci Lett 329 (2002) 339-343
212. Bagnasco M., Tulipano G., Melis M.R., Argiolas A., Cocchi D., and Muller E.E. Endogenous ghrelin is an orexigenic peptide acting in the arcuate nucleus in response to fasting. Regul Pept 111 (2003) 161-167
213. Bewick G.A., Gardiner J.V., Dhillo W.S., Kent A.S., White N.E., Webster Z., et al. Post-embryonic ablation of AgRP neurons in mice leads to a lean, hypophagic phenotype. FASEB J 19 (2005) 1680-1682
214. Naleid A.M., Grace M.K., Cummings D.E., and Levine A.S. Ghrelin induces feeding in the mesolimbic reward pathway between the ventral tegmental area and the nucleus accumbens. Peptides 26 (2005) 2274-2279
215. Carlini V.P., Varas M.M., Cragnolini A.B., Schioth H.B., Scimonelli T.N., and de Barioglio S.R. Differential role of the hippocampus, amygdala, and dorsal raphe nucleus in regulating feeding, memory, and anxiety-like behavioral responses to ghrelin. Biochem Biophys Res Commun 313 (2004) 635-641
216. Diano S., Farr S.A., Benoit S.C., et al. Ghrelin controls hippocampal spine synapse density and memory performance. Nat Neurosci 9 (2006) 381-388
217. Pan W., Tu H., and Kastin A.J. Differential BBB interactions of three ingestive peptides: obestatin, ghrelin, and adiponectin. Peptides 27 (2006) 911-916
218. Sato T., Fukue Y., Teranishi H., Yoshida Y., and Kojima M. Molecular forms of hypothalamic ghrelin and its regulation by fasting and 2-deoxy-d-glucose administration. Endocrinology 146 (2005) 2510-2516
219. Johnstone L.E., Srisawat R., Kumarnsit E., and Leng G. Hypothalamic expression of NPY mRNA, vasopressin mRNA and CRF mRNA in response to food restriction and central administration of the orexigenic peptide GHRP-6. Stress 8 (2005) 59-67
220. West D.B., Fey D., and Woods S.C. Cholecystokinin persistently suppresses meal size but not food intake in free-feeding rats. Am J Physiol 246 (1984) R776-R787
221. Shimbara T., Mondal M.S., Kawagoe T., Toshinai K., Koda S., Yamaguchi H., et al. Central administration of ghrelin preferentially enhances fat ingestion. Neurosci Lett 369 (2004) 75-79
222. Choi K., Roh S.G., Hong Y.H., Shrestha Y.B., Hishikawa D., Chen C., et al. The role of ghrelin and growth hormone secretagogues receptor on rat adipogenesis. Endocrinology 144 (2003) 754-759
223. Thompson N.M., Gill D.A., Davies R., Loveridge N., Houston P.A., Robinson I.C., et al. Ghrelin and des-octanoyl ghrelin promote adipogenesis directly in vivo by a mechanism independent of the type 1a growth hormone secretagogue receptor. Endocrinology 145 (2004) 234-242
224. Kim M.S., Yoon C.Y., Jang P.G., et al. The mitogenic and antiapoptotic actions of ghrelin in 3T3-L1 adipocytes. Mol Endocrinol 18 (2004) 2291-2301
225. Tsubone T., Masaki T., Katsuragi I., Tanaka K., Kakuma T., and Yoshimatsu H. Ghrelin regulates adiposity in white adipose tissue and UCP1 mRNA expression in brown adipose tissue in mice. Regul Pept 130 (2005) 97-103
226. Yasuda T., Masaki T., Kakuma T., and Yoshimatsu H. Centrally administered ghrelin suppresses sympathetic nerve activity in brown adipose tissue of rats. Neurosci Lett 349 (2003) 75-78
227. Barazzoni R., Bosutti A., Stebel M., et al. Ghrelin regulates mitochondrial-lipid metabolism gene expression and tissue fat distribution favoring triglyceride deposition in liver but not skeletal muscle. Am J Physiol Endocrinol Metab (2004) [Electronic publication ahead of print]
228. Muccioli G., Pons N., Ghe C., Catapano F., Granata R., and Ghigo E. Ghrelin and des-acyl ghrelin both inhibit isoproterenol-induced lipolysis in rat adipocytes via a non-type 1a growth hormone secretagogue receptor. Eur J Pharmacol 498 (2004) 27-35
229. Currie P.J., Mirza A., Fuld R., Park D., and Vasselli J.R. Ghrelin is an orexigenic and metabolic signaling peptide in the arcuate and paraventricular nuclei. Am J Physiol Regul Integr Comp Physiol 289 (2005) R353-R358
230. Matsumura K., Tsuchihashi T., Fujii K., Abe I., and Iida M. Central ghrelin modulates sympathetic activity in conscious rabbits. Hypertension 40 (2002) 694-699
231. Dixit V.D., Schaffer E.M., Pyle R.S., Collins G.D., Sakthivel S.K., Palaniappan R., et al. Ghrelin inhibits leptin- and activation-induced proinflammatory cytokine expression by human monocytes and T cells. J Clin Invest 114 (2004) 57-66
232. Li W.G., Gavrila D., Liu X., et al. Ghrelin inhibits proinflammatory responses and nuclear factor-kappaB activation in human endothelial cells. Circulation 109 (2004) 2221-2226
233. Dembinski A., Warzecha Z., Ceranowicz P., Tomaszewska R., Stachura J., Konturek S.J., et al. Ghrelin attenuates the development of acute pancreatitis in rat. J Physiol Pharmacol 54 (2003) 561-573
234. Tang-Christensen M., Vrang N., Ortmann S., Bidlingmaier M., Horvath T.L., and Tschop M. Central administration of ghrelin and agouti-related protein (83-132) increases food intake and decreases spontaneous locomotor activity in rats. Endocrinology 145 (2004) 4645-4652
235. 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. J Clin Endocrinol Metab 87 (2002) 5461-5464
236. Hoybye C. Endocrine and metabolic aspects of adult Prader-Willi syndrome with special emphasis on the effect of growth hormone treatment. Growth Horm IGF Res 14 (2004) 1-15
237. Goldstone A.P., Thomas E.L., Brynes A.E., et al. Elevated fasting plasma ghrelin in Prader-Willi syndrome adults is not solely explained by their reduced visceral adiposity and insulin resistance. J Clin Endocrinol Metab 89 (2004) 1718-1726
238. Tauber M., Conte Auriol F., Moulin P., Molinas C., Delagnes V., and Salles J.P. Hyperghrelinemia is a common feature of Prader-Willi syndrome and pituitary stalk interruption: a pathophysiological hypothesis. Horm Res 62 (2004) 49-54
239. Paik K.H., Jin D.K., Song S.Y., et al. Correlation between fasting plasma ghrelin levels and age, body mass index (BMI), BMI percentiles, and 24-hour plasma ghrelin profiles in Prader-Willi syndrome. J Clin Endocrinol Metab 89 (2004) 3885-3889
240. Goldstone A.P., Patterson M., Kalingag N., Ghatei M.A., Brynes A.E., Bloom S.R., et al. Fasting and postprandial hyperghrelinemia in Prader-Willi syndrome is partially explained by hypoinsulinemia, and is not due to peptide YY3-36 deficiency or seen in hypothalamic obesity due to craniopharyngioma. J Clin Endocrinol Metab 90 (2005) 2681-2690
241. Choe Y.H., Song S.Y., Paik K.H., et al. Increased density of ghrelin-expressing cells in the gastric fundus and body in Prader-Willi syndrome. J Clin Endocrinol Metab 90 (2005) 5441-5445
242. Butler M.G., Bittel D.C., and Talebizadeh Z. Plasma peptide YY and ghrelin levels in infants and children with Prader-Willi syndrome. J Pediatr Endocrinol Metab 17 (2004) 1177-1184
243. Choe Y.H., Jin D.K., Kim S.E., et al. Hyperghrelinemia does not accelerate gastric emptying in Prader-Willi syndrome patients. J Clin Endocrinol Metab 90 (2005) 3367-3370
244. Erdie-Lalena CR, Holm VA, Kelly PC, Frayo RS, Cummings DE. Ghrelin levels in very young children with Prader-Willi syndrome. J Pediatr in press.
245. Kanumakala S., Greaves R., Pedreira C.C., Donath S., Warne G.L., Zacharin M.R., et al. Fasting ghrelin levels are not elevated in children with hypothalamic obesity. J Clin Endocrinol Metab 90 (2005) 2691-2695
246. Daousi C., MacFarlane I.A., English P.J., Wilding J.P., Patterson M., Dovey T.M., et al. Is there a role for ghrelin and peptide-YY in the pathogenesis of obesity in adults with acquired structural hypothalamic damage?. J Clin Endocrinol Metab 90 (2005) 5025-5030
247. Haqq A.M., Stadler D.D., Rosenfeld R.G., Pratt K.L., Weigle D.S., Frayo R.S., et al. Circulating ghrelin levels are suppressed by meals and octreotide therapy in children with Prader-Willi syndrome. J Clin Endocrinol Metab 88 (2003) 3573-3576
248. Tan T.M., Vanderpump M., Khoo B., Patterson M., Ghatei M.A., and Goldstone A.P. Somatostatin infusion lowers plasma ghrelin without reducing appetite in adults with Prader-Willi syndrome. J Clin Endocrinol Metab 89 (2004) 4162-4165
249. Miyasaka K., Masuda M., Kanai S., Ohta M., Suzuki S., Tateishi K., et al. Inhibitory effects of octreotide on luminal cholecystokinin-releasing factor, plasma cholecystokinin, and pancreatic secretion in conscious rats. Pancreas 24 (2002) 269-275
250. Ranganath L.R., Beety J.M., and Morgan L.M. Inhibition of insulin, GIP, and GLP-1 secretion by octreotide has no effect on post-heparin lipoprotein lipase activity. Horm Metab Res 31 (1999) 262-266
251. Tsolakis A.V., Portela-Gomes G.M., Stridsberg M., Grimelius L., Sundin A., Eriksson B.K., et al. Malignant gastric ghrelinoma with hyperghrelinemia. J Clin Endocrinol Metab 89 (2004) 3739-3744
252. Shuto Y., Shibasaki T., Otagiri A., et al. Hypothalamic growth hormone secretagogue receptor regulates growth hormone secretion, feeding, and adiposity. J Clin Invest 109 (2002) 1429-1436
253. Wortley K.E., Anderson K.D., Garcia K., et al. Genetic deletion of ghrelin does not decrease food intake but influences metabolic fuel preference. Proc Natl Acad Sci U S A 101 (2004) 8227-8232
254. Sun Y., Ahmed S., and Smith R.G. Deletion of ghrelin impairs neither growth nor appetite. Mol Cell Biol 23 (2003) 7973-7981
255. Luquet S., Perez F.A., Hnasko T.S., and Palmiter R.D. NPY/Agrp neurons are essential for feeding in adult mice but can be ablated in neonates. Science 310 (2005) 585-683
256. Zhang J.V., Ren P.G., Avsian-Kretchmer O., Luo C.W., Rauch R., Klein C., et al. Obestatin, a peptide encoded by the ghrelin gene, opposes ghrelin's effects on food intake. Science 310 (2005) 996-999
257. 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. J Clin Invest 115 (2005) 3573-3578
258. Zigman J.M., Nakano Y., Coppari R., et al. Mice lacking ghrelin receptors resist the development of diet-induced obesity. J Clin Invest 115 (2005) 3564-3572
259. Helmling S., Maasch C., Eulberg D., et al. Inhibition of ghrelin action in vitro and in vivo by an RNA-Spiegelmer. Proc Natl Acad Sci U S A 101 (2004) 13174-13179
260. Shearman L.P., Wang S.P., Helmling S., et al. Ghrelin neutralization by a ribonucleic acid-SPM ameliorates obesity in diet-induced obese mice. Endocrinology 147 (2006) 1517-1526
261. Kobelt P., Helmling S., Stengel A., Wlotzka B., Andresen V., Klapp B.F., et al. Anti-ghrelin SPIEGELMER NOX-B11 inhibits neurostimulatory and orexigenic effects of peripheral ghrelin in rats. Gut 55 (2005) 788-792
262. Cummings D.E. Helicobacter pylori and ghrelin: interrelated players in body-weight regulation?. Am J Med 117 (2004) 436-439
263. Ukkola O., Ravussin E., Jacobson P., et al. Role of ghrelin polymorphisms in obesity based on three different studies. Obes Res 10 (2002) 782-791