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The Journal of Clinical Endocrinology & Metabolism Vol. 86, No. 3 1169-1174
Copyright © 2001 by The Endocrine Society

Original Studies

Endocrine Activities of Ghrelin, a Natural Growth Hormone Secretagogue (GHS), in Humans: Comparison and Interactions with Hexarelin, a Nonnatural Peptidyl GHS, and GH-Releasing Hormone1

Emanuela Arvat, Mauro Maccario, Lidia Di Vito, Fabio Broglio, Andrea Benso, Cristina Gottero, Mauro Papotti, Giampiero Muccioli, Carlos Dieguez, Felipe F. Casanueva, Romano Deghenghi, Franco Camanni and Ezio Ghigo

Division of Endocrinology and Metabolism, Department of Internal Medicine (E.A., M.M., L.D.V., F.B., A.B., C.G., F.C., E.G.), Department of Biomedical Sciences and Oncology (M.P.), Department of Anatomy, Pharmacology and Forensic Medicine (G.M.), University of Turin, 10126 Turin, Italy; Department of Physiology (C.D.) and Department of Medicine (F.F.C.), Faculty of Medicine, Santiago de Compostela University, Santiago de Compostela E-15780, Spain; and Europeptides (R.D.), Argenteuil 95108 Cedex, France

Address all correspondence and requests for reprints to: E. Ghigo, M.D., Divisione di Endocrinologia, Ospedale Molinette, Corso Dogliotti 14, 10126 Torino, Italy. E-mail: ezio.ghigo{at}unito.it

An endogenous ligand for the GH secretagogue-receptor (GHS-receptor) has recently been isolated, from both the rat and the human stomach, and named ghrelin. It is a 28-amino-acid peptide showing a unique structure with an n-octanoyl ester at its third serine residue, which is essential for its potent stimulatory activity on somatotroph secretion. In fact, it has been demonstrated that ghrelin specifically stimulates GH secretion from both rat pituitary cells in culture and rats in vivo. The aim of the present study was to test the GH-releasing activity of ghrelin in humans and to compare it with that of GHRH and hexarelin (HEX), a nonnatural peptidyl GHS, which possesses strong GH-releasing activity but also significantly stimulates PRL, ACTH, and cortisol secretion. To clarify the mechanisms of action underlying the GH-releasing activity of ghrelin in humans, its interaction with GHRH and HEX was also studied. Seven normal young volunteers (7 men; 24–32 yr old; body mass index, 20–24 kg/m2) were studied. All subjects underwent the administration of ghrelin, HEX, and GHRH-29 (1.0 µg/kg iv at 0 min) as well as placebo (2 mL isotonic saline iv at 0 min). Six subjects also underwent the combined administration of ghrelin and GHRH or HEX. Blood samples were taken every 15 min from -15 up to +180 min. GH levels were assayed at each time point in all sessions; PRL, ACTH, cortisol, and aldosterone levels were also assayed after administration of ghrelin and/or HEX. Ghrelin administration induced a prompt and marked increase in circulating GH levels (Cmax, mean ± SEM, 92.1 ± 16.7 µg/L; area under the curve, 1894.9 ± 347.8 µg/L·h). The GH response to ghrelin was clearly higher (P < 0.01) than the one recorded after GHRH (26.7 ± 8.7 µg/L; 619.6 ± 174.4 µg/L·h) and even significantly higher (P < 0.05) than after HEX (68.4 ± 14.7 µg/L; 1546.9 ± 380.0 µg/L·h). Ghrelin administration also induced an increase in PRL, ACTH, and cortisol levels; these responses were higher (P < 0.05) than those elicited by HEX. A significant increase in aldosterone levels was recorded after ghrelin but not after HEX. The endocrine responses to ghrelin were not modified by the coadministration of HEX. On the other hand, the coadministration of ghrelin and GHRH had a real synergistical effect (P < 0.05) on GH secretion (133.6 ± 22.5 µg/L; 3374.3 ± 617.3 µg/L·h). In conclusion, ghrelin, a natural ligand of GHS-receptor, exerts a strong stimulatory effect on GH secretion in humans, releasing more GH than GHRH and even more than a nonnatural GHS such as HEX. Ghrelin, as well as HEX, also stimulates lactotroph and corticotroph secretion. Ghrelin shows no interaction with HEX, whereas it has a synergistical effect with GHRH on GH secretion. Thus, ghrelin is a new hormone playing a major role in the control of somatotroph secretion in humans, and its effects are imitated by nonnatural GHS.




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Low Cord Ghrelin Levels in Term Infants Are Associated with Slow Weight Gain Over the First 3 Months of Life
J. Clin. Endocrinol. Metab., August 1, 2004; 89(8): 3847 - 3850.
[Abstract] [Full Text] [PDF]

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 EndocrinologyHome page
J.-P. Chanoine and A. C. K. Wong
Ghrelin Gene Expression Is Markedly Higher in Fetal Pancreas Compared with Fetal Stomach: Effect of Maternal Fasting
Endocrinology, August 1, 2004; 145(8): 3813 - 3820.
[Abstract] [Full Text] [PDF]

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 EndocrinologyHome page
R. M. Luque, R. D. Kineman, S. Park, X.-D. Peng, F. Gracia-Navarro, J. P. Castano, and M. M. Malagon
Homologous and Heterologous Regulation of Pituitary Receptors for Ghrelin and Growth Hormone-Releasing Hormone
Endocrinology, July 1, 2004; 145(7): 3182 - 3189.
[Abstract] [Full Text] [PDF]

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 Endocr. Rev.Home page
A. J. van der Lely, M. Tschop, M. L. Heiman, and E. Ghigo
Biological, Physiological, Pathophysiological, and Pharmacological Aspects of Ghrelin
Endocr. Rev., June 1, 2004; 25(3): 426 - 457.
[Abstract] [Full Text] [PDF]

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 J. Clin. Endocrinol. Metab.Home page
C. Y. Bowers, R. Granda, S. Mohan, J. Kuipers, D. Baylink, and J. D. Veldhuis
Sustained Elevation of Pulsatile Growth Hormone (GH) Secretion and Insulin-Like Growth Factor I (IGF-I), IGF-Binding Protein-3 (IGFBP-3), and IGFBP-5 Concentrations during 30-Day Continuous Subcutaneous Infusion of GH-Releasing Peptide-2 in Older Men and Women
J. Clin. Endocrinol. Metab., May 1, 2004; 89(5): 2290 - 2300.
[Abstract] [Full Text] [PDF]

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 EndocrinologyHome page
S. Lall, N. Balthasar, D. Carmignac, C. Magoulas, A. Sesay, P. Houston, K. Mathers, and I. Robinson
Physiological Studies of Transgenic Mice Overexpressing Growth Hormone (GH) Secretagogue Receptor 1A in GH-Releasing Hormone Neurons
Endocrinology, April 1, 2004; 145(4): 1602 - 1611.
[Abstract] [Full Text] [PDF]

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 J. Clin. Endocrinol. Metab.Home page
M. E. Davis, C. J. Pemberton, T. G. Yandle, J. G. Lainchbury, M. T. Rademaker, M. G. Nicholls, C. M. Frampton, and A. M. Richards
Urocortin-1 Infusion in Normal Humans
J. Clin. Endocrinol. Metab., March 1, 2004; 89(3): 1402 - 1409.
[Abstract] [Full Text] [PDF]

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 J. Clin. Endocrinol. Metab.Home page
F. Gaytan, M. L. Barreiro, J. E. Caminos, L. K. Chopin, A. C. Herington, C. Morales, L. Pinilla, R. Paniagua, M. Nistal, F. F. Casanueva, et al.
Expression of Ghrelin and Its Functional Receptor, the Type 1a Growth Hormone Secretagogue Receptor, in Normal Human Testis and Testicular Tumors
J. Clin. Endocrinol. Metab., January 1, 2004; 89(1): 400 - 409.
[Abstract] [Full Text] [PDF]

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 PhysiologyHome page
D. H. St-Pierre, L. Wang, and Y. Tache
Ghrelin: A Novel Player in the Gut-Brain Regulation of Growth Hormone and Energy Balance
Physiology, December 1, 2003; 18(6): 242 - 246.
[Abstract] [Full Text] [PDF]

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 J. Clin. Endocrinol. Metab.Home page
J. Q. Purnell, D. S. Weigle, P. Breen, and D. E. Cummings
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., December 1, 2003; 88(12): 5747 - 5752.
[Abstract] [Full Text] [PDF]

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 EndocrinologyHome page
M. M. Malagon, R. M. Luque, E. Ruiz-Guerrero, F. Rodriguez-Pacheco, S. Garcia-Navarro, F. F. Casanueva, F. Gracia-Navarro, and J. P. Castano
Intracellular Signaling Mechanisms Mediating Ghrelin-Stimulated Growth Hormone Release in Somatotropes
Endocrinology, December 1, 2003; 144(12): 5372 - 5380.
[Abstract] [Full Text] [PDF]

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 Exp. Biol. Med.Home page
X. Qi, J. Reed, E. W. Englander, V. Chandrashekar, A. Bartke, and G. H. Greeley Jr.
Evidence That Growth Hormone Exerts a Feedback Effect on Stomach Ghrelin Production and Secretion
Experimental Biology and Medicine, October 1, 2003; 228(9): 1028 - 1032.
[Abstract] [Full Text] [PDF]

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 J. Clin. Endocrinol. Metab.Home page
J. Farquhar, M. Heiman, A. C. K. Wong, R. Wach, P. Chessex, and J.-P. Chanoine
Elevated Umbilical Cord Ghrelin Concentrations in Small for Gestational Age Neonates
J. Clin. Endocrinol. Metab., September 1, 2003; 88(9): 4324 - 4327.
[Abstract] [Full Text] [PDF]

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 J. Clin. Endocrinol. Metab.Home page
J. C. Bunt, A. D. Salbe, M. H. Tschop, A. DelParigi, P. Daychild, and P. A. Tataranni
Cross-Sectional and Prospective Relationships of Fasting Plasma Ghrelin Concentrations with Anthropometric Measures in Pima Indian Children
J. Clin. Endocrinol. Metab., August 1, 2003; 88(8): 3756 - 3761.
[Abstract] [Full Text] [PDF]

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 J. Clin. Endocrinol. Metab.Home page
C. Holdstock, B. E. Engstrom, M. Ohrvall, L. Lind, M. Sundbom, and F. A. Karlsson
Ghrelin and Adipose Tissue Regulatory Peptides: Effect of Gastric Bypass Surgery in Obese Humans
J. Clin. Endocrinol. Metab., July 1, 2003; 88(7): 3177 - 3183.
[Abstract] [Full Text] [PDF]

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 EndocrinologyHome page
A. Torsello, E. Bresciani, G. Rossoni, R. Avallone, G. Tulipano, D. Cocchi, I. Bulgarelli, R. Deghenghi, F. Berti, and V. Locatelli
Ghrelin Plays a Minor Role in the Physiological Control of Cardiac Function in the Rat
Endocrinology, May 1, 2003; 144(5): 1787 - 1792.
[Abstract] [Full Text] [PDF]

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 Biol. Reprod.Home page
M.L. Barreiro, J.S. Suominen, F. Gaytan, L. Pinilla, L.K. Chopin, F.F. Casanueva, C. Dieguez, E. Aguilar, J. Toppari, and M. Tena-Sempere
Developmental, Stage-Specific, and Hormonally Regulated Expression of Growth Hormone Secretagogue Receptor Messenger RNA in Rat Testis
Biol Reprod, May 1, 2003; 68(5): 1631 - 1640.
[Abstract] [Full Text] [PDF]

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 J. Clin. Endocrinol. Metab.Home page
F. Broglio, A. Benso, C. Castiglioni, C. Gottero, F. Prodam, S. Destefanis, C. Gauna, A. J. van der Lely, R. Deghenghi, M. Bo, et al.
The Endocrine Response to Ghrelin as a Function of Gender in Humans in Young and Elderly Subjects
J. Clin. Endocrinol. Metab., April 1, 2003; 88(4): 1537 - 1542.
[Abstract] [Full Text] [PDF]

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 EndocrinologyHome page
K. Choi, S.-G. Roh, Y.-H. Hong, Y. B. Shrestha, D. Hishikawa, C. Chen, M. Kojima, K. Kangawa, and S.-I. Sasaki
The Role of Ghrelin and Growth Hormone Secretagogues Receptor on Rat Adipogenesis
Endocrinology, March 1, 2003; 144(3): 754 - 759.
[Abstract] [Full Text] [PDF]

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 J. Clin. Endocrinol. Metab.Home page
F. Gaytan, M. L. Barreiro, L. K. Chopin, A. C. Herington, C. Morales, L. Pinilla, F. F. Casanueva, E. Aguilar, C. Dieguez, and M. Tena-Sempere
Immunolocalization of Ghrelin and Its Functional Receptor, the Type 1a Growth Hormone Secretagogue Receptor, in the Cyclic Human Ovary
J. Clin. Endocrinol. Metab., February 1, 2003; 88(2): 879 - 887.
[Abstract] [Full Text] [PDF]

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 Am. J. Pathol.Home page
M. Volante, E. Allia, E. Fulcheri, P. Cassoni, E. Ghigo, G. Muccioli, and M. Papotti
Ghrelin in Fetal Thyroid and Follicular Tumors and Cell Lines: Expression and Effects on Tumor Growth
Am. J. Pathol., February 1, 2003; 162(2): 645 - 654.
[Abstract] [Full Text] [PDF]

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 Am. J. Physiol. Endocrinol. Metab.Home page
J. C. Weikel, A. Wichniak, M. Ising, H. Brunner, E. Friess, K. Held, S. Mathias, D. A. Schmid, M. Uhr, and A. Steiger
Ghrelin promotes slow-wave sleep in humans
Am J Physiol Endocrinol Metab, February 1, 2003; 284(2): E407 - E415.
[Abstract] [Full Text] [PDF]

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 J. Clin. Endocrinol. Metab.Home page
A. M. Haqq, I. S. Farooqi, S. O'Rahilly, D. D. Stadler, R. G. Rosenfeld, K. L. Pratt, S. H. LaFranchi, and J. Q. Purnell
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., January 1, 2003; 88(1): 174 - 178.
[Abstract] [Full Text] [PDF]

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 Mol. Interv.Home page
H. Hosoda, M. Kojima, and K. Kangawa
Ghrelin and the Regulation of Food Intake and Energy Balance
Mol. Interv., December 1, 2002; 2(8): 494 - 503.
[Abstract] [Full Text]

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 J. Clin. Endocrinol. Metab.Home page
A. DelParigi, M. Tschop, M. L. Heiman, A. D. Salbe, B. Vozarova, S. M. Sell, J. C. Bunt, and P. A. Tataranni
High Circulating Ghrelin: A Potential Cause for Hyperphagia and Obesity in Prader-Willi Syndrome
J. Clin. Endocrinol. Metab., December 1, 2002; 87(12): 5461 - 5464.
[Abstract] [Full Text] [PDF]

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 Biol. Reprod.Home page
M.L. Barreiro, F. Gaytan, J.E. Caminos, L. Pinilla, F.F. Casanueva, E. Aguilar, C. Dieguez, and M. Tena-Sempere
Cellular Location and Hormonal Regulation of Ghrelin Expression in Rat Testis
Biol Reprod, December 1, 2002; 67(6): 1768 - 1776.
[Abstract] [Full Text] [PDF]

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 DiabetesHome page
P. Lucidi, G. Murdolo, C. Di Loreto, A. De Cicco, N. Parlanti, C. Fanelli, F. Santeusanio, G. B. Bolli, and P. De Feo
Ghrelin Is Not Necessary for Adequate Hormonal Counterregulation of Insulin-Induced Hypoglycemia
Diabetes, October 1, 2002; 51(10): 2911 - 2914.
[Abstract] [Full Text] [PDF]

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 EndocrinologyHome page
H. Kaiya, S. van der Geyten, M. Kojima, H. Hosoda, Y. Kitajima, M. Matsumoto, S. Geelissen, V. M. Darras, and K. Kangawa
Chicken Ghrelin: Purification, cDNA Cloning, and Biological Activity
Endocrinology, September 1, 2002; 143(9): 3454 - 3463.
[Abstract] [Full Text] [PDF]

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 J. Histochem. Cytochem.Home page
M. Volante, E. Fulcheri, E. Allia, M. Cerrato, A. Pucci, and M. Papotti
Ghrelin Expression in Fetal, Infant, and Adult Human Lung
J. Histochem. Cytochem., August 1, 2002; 50(8): 1013 - 1021.
[Abstract] [Full Text] [PDF]

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 J. Clin. Endocrinol. Metab.Home page
F. Broglio, E. Arvat, A. Benso, C. Gottero, F. Prodam, S. Grottoli, M. Papotti, G. Muccioli, A. J. van der Lely, R. Deghenghi, et al.
Endocrine Activities of Cortistatin-14 and Its Interaction with GHRH and Ghrelin in Humans
J. Clin. Endocrinol. Metab., August 1, 2002; 87(8): 3783 - 3790.
[Abstract] [Full Text] [PDF]

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 J. Pharmacol. Exp. Ther.Home page
W. A. Banks, M. Tschop, S. M. Robinson, and M. L. Heiman
Extent and Direction of Ghrelin Transport Across the Blood-Brain Barrier Is Determined by Its Unique Primary Structure
J. Pharmacol. Exp. Ther., August 1, 2002; 302(2): 822 - 827.
[Abstract] [Full Text] [PDF]

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 EndocrinologyHome page
S. Ishizaki, T. Murase, Y. Sugimura, S. Kakiya, H. Yokoi, K. Tachikawa, H. Arima, Y. Miura, and Y. Oiso
Role of Ghrelin in the Regulation of Vasopressin Release in Conscious Rats
Endocrinology, May 1, 2002; 143(5): 1589 - 1593.
[Abstract] [Full Text] [PDF]

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 EndocrinologyHome page
V. Tolle, M.-H. Bassant, P. Zizzari, F. Poindessous-Jazat, C. Tomasetto, J. Epelbaum, and M.-T. Bluet-Pajot
Ultradian Rhythmicity of Ghrelin Secretion in Relation with GH, Feeding Behavior, and Sleep-Wake Patterns in Rats
Endocrinology, April 1, 2002; 143(4): 1353 - 1361.
[Abstract] [Full Text] [PDF]

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 J. Clin. Endocrinol. Metab.Home page
M. Volante, E. AllIa, P. Gugliotta, A. Funaro, F. Broglio, R. Deghenghi, G. Muccioli, E. Ghigo, and M. Papotti
Expression of Ghrelin and of the GH Secretagogue Receptor by Pancreatic Islet Cells and Related Endocrine Tumors
J. Clin. Endocrinol. Metab., March 1, 2002; 87(3): 1300 - 1308.
[Abstract] [Full Text] [PDF]

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 EndocrinologyHome page
C. Ghe, P. Cassoni, F. Catapano, T. Marrocco, R. Deghenghi, E. Ghigo, G. Muccioli, and M. Papotti
The Antiproliferative Effect of Synthetic Peptidyl GH Secretagogues in Human CALU-1 Lung Carcinoma Cells
Endocrinology, February 1, 2002; 143(2): 484 - 491.
[Abstract] [Full Text] [PDF]

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 EndocrinologyHome page
T. L. Horvath, S. Diano, P. Sotonyi, M. Heiman, and M. Tschop
Minireview: Ghrelin and the Regulation of Energy Balance--A Hypothalamic Perspective
Endocrinology, October 1, 2001; 142(10): 4163 - 4169.
[Abstract] [Full Text] [PDF]


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