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Endocrine Reviews 25 (3): 426-457
Copyright © 2004 by The Endocrine Society

Biological, Physiological, Pathophysiological, and Pharmacological Aspects of Ghrelin

Aart J. van der Lely, Matthias Tschöp, Mark L. Heiman and Ezio Ghigo

Division of Endocrinology and Metabolism (A.J.v.d.L.), Department of Internal Medicine, Erasmus Medical Center, 3015 GD Rotterdam, The Netherlands; Department of Psychiatry (M.T.), University of Cincinnati, Cincinnati, Ohio 45237; Endocrine Research Department (M.L.H.), Eli Lilly and Co., Indianapolis, Indiana 46285; and Division of Endocrinology (E.G.), Department of Internal Medicine, University of Turin, Turin, Italy 10095

Correspondence: Address all correspondence and requests for reprints to: Aart Jan van der Lely, M.D., Department of Internal Medicine, Erasmus Medical Center, 40 Dr. Molewaterplein, 3015 GD Rotterdam, The Netherlands. E-mail: a.vanderlelij{at}erasmusmc.nl

Ghrelin is a peptide predominantly produced by the stomach. Ghrelin displays strong GH-releasing activity. This activity is mediated by the activation of the so-called GH secretagogue receptor type 1a. This receptor had been shown to be specific for a family of synthetic, peptidyl and nonpeptidyl GH secretagogues. Apart from a potent GH-releasing action, ghrelin has other activities including stimulation of lactotroph and corticotroph function, influence on the pituitary gonadal axis, stimulation of appetite, control of energy balance, influence on sleep and behavior, control of gastric motility and acid secretion, and influence on pancreatic exocrine and endocrine function as well as on glucose metabolism. Cardiovascular actions and modulation of proliferation of neoplastic cells, as well as of the immune system, are other actions of ghrelin. Therefore, we consider ghrelin a gastrointestinal peptide contributing to the regulation of diverse functions of the gut-brain axis. So, there is indeed a possibility that ghrelin analogs, acting as either agonists or antagonists, might have clinical impact.




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J. Neurosci., October 25, 2006; 26(43): 11052 - 11060.
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M. C Carreira, J. P Camina, E. Diaz-Rodriguez, R. Alvear-Perez, C. Llorens-Cortes, and F. F Casanueva
Adenosine does not bind to the growth hormone secretagogue receptor type-1a (GHS-R1a).
J. Endocrinol., October 1, 2006; 191(1): 147 - 157.
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L. Wang, N. R. Basa, A. Shaikh, A. Luckey, D. Heber, D. H. St-Pierre, and Y. Tache
LPS inhibits fasted plasma ghrelin levels in rats: role of IL-1 and PGs and functional implications.
Am J Physiol Gastrointest Liver Physiol, October 1, 2006; 291(4): G611 - G620.
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P. Marzullo, A. Caumo, G. Savia, B. Verti, G. E. Walker, S. Maestrini, A. Tagliaferri, A. M. Di Blasio, and A. Liuzzi
Predictors of Postabsorptive Ghrelin Secretion after Intake of Different Macronutrients
J. Clin. Endocrinol. Metab., October 1, 2006; 91(10): 4124 - 4130.
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J. O. L. Jorgensen
A simple twist of science: the convoluted tale of ghrelin continues.
J. Clin. Endocrinol. Metab., September 1, 2006; 91(9): 3279 - 3280.
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 Phil Trans R Soc BHome page
O. Chaudhri, C. Small, and S. Bloom
Gastrointestinal hormones regulating appetite
Phil Trans R Soc B, July 29, 2006; 361(1471): 1187 - 1209.
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L. Asarian and N. Geary
Modulation of appetite by gonadal steroid hormones
Phil Trans R Soc B, July 29, 2006; 361(1471): 1251 - 1263.
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A. J. Martinez-Fuentes, J. Moreno-Fernandez, R. Vazquez-Martinez, M. Duran-Prado, A. de la Riva, M. Tena-Sempere, C. Dieguez, L. Jimenez-Reina, S. M. Webb, A. Pumar, et al.
Ghrelin Is Produced by and Directly Activates Corticotrope Cells from Adrenocorticotropin-Secreting Adenomas
J. Clin. Endocrinol. Metab., June 1, 2006; 91(6): 2225 - 2231.
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V. Di Francesco, M. Zamboni, E. Zoico, G. Mazzali, A. Dioli, F. Omizzolo, L. Bissoli, F. Fantin, P. Rizzotti, S. B Solerte, et al.
Unbalanced serum leptin and ghrelin dynamics prolong postprandial satiety and inhibit hunger in healthy elderly: another reason for the "anorexia of aging"
Am. J. Clinical Nutrition, May 1, 2006; 83(5): 1149 - 1152.
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S. Bellone, R. Baldelli, G. Radetti, A. Rapa, D. Vivenza, A. Petri, S. Savastio, M. Zaffaroni, F. Broglio, E. Ghigo, et al.
Ghrelin Secretion in Preterm Neonates Progressively Increases and Is Refractory to the Inhibitory Effect of Food Intake
J. Clin. Endocrinol. Metab., May 1, 2006; 91(5): 1929 - 1933.
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A. C. Martini, R. Fernandez-Fernandez, S. Tovar, V. M. Navarro, E. Vigo, M. J. Vazquez, J. S. Davies, N. M. Thompson, E. Aguilar, L. Pinilla, et al.
Comparative Analysis of the Effects of Ghrelin and Unacylated Ghrelin on Luteinizing Hormone Secretion in Male Rats
Endocrinology, May 1, 2006; 147(5): 2374 - 2382.
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N. A. Tritos and E. G. Kokkotou
The Physiology and Potential Clinical Applications of Ghrelin, a Novel Peptide Hormone
Mayo Clin. Proc., May 1, 2006; 81(5): 653 - 660.
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A. E. Rigamonti, S. M. Bonomo, D. Scanniffio, S. G. Cella, and E. E. Muller
Orexigenic effects of a growth hormone secretagogue and nitric oxide in aged rats and dogs: correlation with the hypothalamic expression of some neuropeptidergic/receptorial effectors mediating food intake.
J. Gerontol. A Biol. Sci. Med. Sci., April 1, 2006; 61(4): 315 - 322.
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J. D. Veldhuis, J. N. Roemmich, E. J. Richmond, and C. Y. Bowers
Somatotropic and Gonadotropic Axes Linkages in Infancy, Childhood, and the Puberty-Adult Transition
Endocr. Rev., April 1, 2006; 27(2): 101 - 140.
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S. Grottoli, V. Gasco, F. Broglio, R. Baldelli, F. Ragazzoni, F. Gallenca, A. Mainolfi, F. Prodam, G. Muccioli, and E. Ghigo
Cortistatin-17 and Somatostatin-14 Display the Same Effects on Growth Hormone, Prolactin, and Insulin Secretion in Patients with Acromegaly or Prolactinoma
J. Clin. Endocrinol. Metab., April 1, 2006; 91(4): 1595 - 1599.
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T Akamizu, T Murayama, S Teramukai, K Miura, I Bando, T Irako, H Iwakura, H Ariyasu, H Hosoda, H Tada, et al.
Plasma ghrelin levels in healthy elderly volunteers: the levels of acylated ghrelin in elderly females correlate positively with serum IGF-I levels and bowel movement frequency and negatively with systolic blood pressure
J. Endocrinol., February 1, 2006; 188(2): 333 - 344.
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U. Pagotto, G. Marsicano, D. Cota, B. Lutz, and R. Pasquali
The Emerging Role of the Endocannabinoid System in Endocrine Regulation and Energy Balance
Endocr. Rev., February 1, 2006; 27(1): 73 - 100.
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P J D Delhanty, B C J van der Eerden, M van der Velde, C Gauna, H A P Pols, H Jahr, H Chiba, A J van der Lely, and J P T M van Leeuwen
Ghrelin and unacylated ghrelin stimulate human osteoblast growth via mitogen-activated protein kinase (MAPK)/phosphoinositide 3-kinase (PI3K) pathways in the absence of GHS-R1a
J. Endocrinol., January 1, 2006; 188(1): 37 - 47.
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 Pharmacol. Rev.Home page
A. P. Davenport, T. I. Bonner, S. M. Foord, A. J. Harmar, R. R. Neubig, J.-P. Pin, M. Spedding, M. Kojima, and K. Kangawa
International Union of Pharmacology. LVI. Ghrelin Receptor Nomenclature, Distribution, and Function
Pharmacol. Rev., December 1, 2005; 57(4): 541 - 546.
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 ScienceHome page
J. V. Zhang, P.-G. Ren, O. Avsian-Kretchmer, C.-W. Luo, R. Rauch, C. Klein, and A. J. W. Hsueh
Obestatin, a Peptide Encoded by the Ghrelin Gene, Opposes Ghrelin's Effects on Food Intake
Science, November 11, 2005; 310(5750): 996 - 999.
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 Eur J EndocrinolHome page
R. Giordano, A. Picu, U. Pagotto, R. De Iasio, L. Bonelli, F. Prodam, F. Broglio, L. Marafetti, R. Pasquali, M. Maccario, et al.
The negative association between total ghrelin levels, body mass and insulin secretion is lost in hypercortisolemic patients with Cushing's disease
Eur. J. Endocrinol., October 1, 2005; 153(4): 535 - 543.
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M A Arafat, B Otto, H Rochlitz, M Tschop, V Bahr, M Mohlig, S Diederich, J Spranger, and A F H Pfeiffer
Glucagon inhibits ghrelin secretion in humans
Eur. J. Endocrinol., September 1, 2005; 153(3): 397 - 402.
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 Hum ReprodHome page
D. Panidis, D. Farmakiotis, G. Koliakos, D. Rousso, A. Kourtis, I. Katsikis, C. Asteriadis, V. Karayannis, and E. Diamanti-Kandarakis
Comparative study of plasma ghrelin levels in women with polycystic ovary syndrome, in hyperandrogenic women and in normal controls
Hum. Reprod., August 1, 2005; 20(8): 2127 - 2132.
[Abstract] [Full Text] [PDF]

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 Physiol. GenomicsHome page
J. H. Kim, T. P. Stewart, W. Zhang, H. Y. Kim, P. M. Nishina, and J. K. Naggert
Type 2 diabetes mouse model TallyHo carries an obesity gene on chromosome 6 that exaggerates dietary obesity
Physiol Genomics, July 14, 2005; 22(2): 171 - 181.
[Abstract] [Full Text] [PDF]

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R. Fernandez-Fernandez, V. M. Navarro, M. L. Barreiro, E. M. Vigo, S. Tovar, A. V. Sirotkin, F. F. Casanueva, E. Aguilar, C. Dieguez, L. Pinilla, et al.
Effects of Chronic Hyperghrelinemia on Puberty Onset and Pregnancy Outcome in the Rat
Endocrinology, July 1, 2005; 146(7): 3018 - 3025.
[Abstract] [Full Text] [PDF]

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C. Y. Bowers
Octanoyl Ghrelin Is Hypothalamic Rooted
Endocrinology, June 1, 2005; 146(6): 2508 - 2509.
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A. M. Avram, C. A. Jaffe, K. V. Symons, and A. L. Barkan
Endogenous Circulating Ghrelin Does Not Mediate Growth Hormone Rhythmicity or Response to Fasting
J. Clin. Endocrinol. Metab., May 1, 2005; 90(5): 2982 - 2987.
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 Am. J. PsychiatryHome page
M. C. ROSENHAGEN, M. UHR, P. SCHUSSLER, and A. STEIGER
Elevated Plasma Ghrelin Levels in Night-Eating Syndrome
Am J Psychiatry, April 1, 2005; 162(4): 813 - 813.
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V. Popovic, D. Miljic, S. Pekic, P. Pesko, M. Djurovic, M. Doknic, S. Damjanovic, D. Micic, G. Cvijovic, J. Glodic, et al.
Low Plasma Ghrelin Level in Gastrectomized Patients Is Accompanied by Enhanced Sensitivity to the Ghrelin-Induced Growth Hormone Release
J. Clin. Endocrinol. Metab., April 1, 2005; 90(4): 2187 - 2191.
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 J. Clin. Endocrinol. Metab.Home page
M. Patterson, K. G. Murphy, C. W. le Roux, M. A. Ghatei, and S. R. Bloom
Characterization of Ghrelin-Like Immunoreactivity in Human Plasma
J. Clin. Endocrinol. Metab., April 1, 2005; 90(4): 2205 - 2211.
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F. Gaytan, C. Morales, M. L. Barreiro, P. Jeffery, L. K. Chopin, A. C. Herington, F. F. Casanueva, E. Aguilar, C. Dieguez, and M. Tena-Sempere
Expression of Growth Hormone Secretagogue Receptor Type 1a, the Functional Ghrelin Receptor, in Human Ovarian Surface Epithelium, Mullerian Duct Derivatives, and Ovarian Tumors
J. Clin. Endocrinol. Metab., March 1, 2005; 90(3): 1798 - 1804.
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G. Maccarinelli, V. Sibilia, A. Torsello, F. Raimondo, M. Pitto, A. Giustina, C. Netti, and D. Cocchi
Ghrelin regulates proliferation and differentiation of osteoblastic cells
J. Endocrinol., January 1, 2005; 184(1): 249 - 256.
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 ANN INTERN MEDHome page
K. Spiegel, E. Tasali, P. Penev, and E. V. Cauter
Brief Communication: Sleep Curtailment in Healthy Young Men Is Associated with Decreased Leptin Levels, Elevated Ghrelin Levels, and Increased Hunger and Appetite
Ann Intern Med, December 7, 2004; 141(11): 846 - 850.
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 Am. J. Physiol. Regul. Integr. Comp. Physiol.Home page
A. Steiger
Eating and sleeping--their relationship to ghrelin and leptin
Am J Physiol Regulatory Integrative Comp Physiol, November 1, 2004; 287(5): R1031 - R1032.
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