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Department of Internal Medicine, Miyazaki Medical College (T.S., M.N., M.M., Y.D., M.S.M., S.M.), Miyazaki 889-1692, Japan; National Cardiovascular Center Research Institute (H.H., K.K.), Osaka 565-8565, Japan; and Department of Psychosomatic Medicine, Kagoshima University School of Medicine (M.T., S.N.), Kagoshima 890-8520, Japan
Address all correspondence and requests for reprints to: Masamitsu Nakazato, M.D., Ph.D., Third Department of Internal Medicine, Miyazaki Medical College, Kiyotake, Miyazaki 889-1692, Japan. E-mail: nakazato{at}post.miyazaki-med.ac.jp
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Abstract |
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Plasma ghrelin concentrations in patients with simple obesity and anorexia nervosa were lower and higher, respectively, than those of healthy subjects with normal body weight. Among those with type 2 diabetes mellitus, obese patients had lower and lean patients higher fasting plasma ghrelin concentrations than normal-weight patients. Fasting plasma ghrelin concentration was negatively correlated with body mass index in both nondiabetic and diabetic patients. Plasma ghrelin concentrations of normal subjects decreased significantly after oral and iv glucose administration; a similar response was also observed in diabetic patients after a meal tolerance test, reaching a nadir of 69% of the basal level after the meal. Circulating plasma ghrelin showed a diurnal pattern with preprandial increases, postprandial decreases, and a maximum peak at 0200 h. This study demonstrates that nutritional state is a determinant of plasma ghrelin in humans. Ghrelin secretion is up-regulated under conditions of negative energy balance and down-regulated in the setting of positive energy balance. These findings suggest the involvement of ghrelin in the regulation of feeding behavior and energy homeostasis.
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Introduction |
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Subjects and Methods |
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The following groups were studied: 28 healthy controls [14 men and 14 women; mean age ± SEM, 30.4 ± 4.1 yr; body mass index (BMI), 19.8–24.6, mean ± SEM, 22.7 ± 0.4]; 17 patients with anorexia nervosa (1 man and 16 women; 22.2 ± 2.3 yr; BMI, 9.3–17.3, 14.2 ± 0.5); 11 patients with simple obesity (4 men and 7 women; 35.1 ± 3.7 yr; BMI, 26.3–40.5, 30.4 ± 1.2); and 42 patients with type 2 DM without nephropathy (18 men and 24 women; 58.5 ± 1.6 yr; BMI <18.5, n = 4; 18.5 
BMI < 25, n = 19; BMI 
25, n = 19). Among the anorexia nervosa patients, 9 were of the restricting type and 8 of the binge eating/purging type according to the Diagnostic and Statistical Manual of Mental Disorders, 4th edition, American Psychiatric Association, 1994. Among the diabetic patients, 12 were treated with diet and exercise, 22 with oral hypoglycemic agents, and 8 with insulin. All subjects were classified into lean (BMI <18.5), normal (18.5 BMI < 25), and obese (BMI 25) categories according to the criteria of the Japan Diabetes Society and Japan Society for the Study of Obesity. All subjects were clinically stable at the time of evaluation and had no evidence of gastrointestinal disease or cachectic states such as cancer, thyroid disease, liver disease, or infection. Patients with renal dysfunction (serum creatinine 1.5 mg/dl) were excluded. Blood was collected at 0800 h after an overnight fast.
Protocol
Healthy volunteers who were within 10% of ideal body weight (9 men and 10 women; 25.3 ± 2.5 yr) were given a 75 g/225 ml glucose solution orally. They also were given 225 ml distilled water orally on a different day. Blood was collected 0, 30, 60, and 120 min after administration. These subjects also were given 10 g/20 ml glucose iv for 2 min. Blood was collected 0, 3, 5, 10, 15, 30, and 60 min after the glucose injection. Seven diabetic patients without triopathy (3 men and 4 women; 52.6 ± 5.6 yr; BMI, 21.3–37.0, 26.0 ± 2.0) were given a test meal (450 cal, 50% carbohydrate, 30% fat, and 20% protein) at 0800 h. Blood was collected 0, 30, 60, and 120 min after eating. For the 24-h monitoring study, 10 healthy men of normal weight (29.6 ± 3.4 yr) were given meals at 0800, 1200, and 1800 h, and blood was collected every 2 h over that 24-h period. Plasma glucose was measured by the glucose oxidase method. Plasma insulin was determined by an RIA kit (Shionogi, Tokyo, Japan). The Institutional Committee of Miyazaki Medical College approved the protocol, and all subjects provided written informed consent before participation.
Ghrelin assay
Blood was drawn into chilled tubes containing EDTA·2Na (1 mg/ml) and aprotinin (500 U/ml). Plasma ghrelin was measured by an RIA developed in our laboratory (17). In brief, antiserum against the C-terminal region of human ghrelin was raised in New Zealand white rabbits that were immunized against synthetic human ghrelin [13–28] that had been coupled with maleimide-activated mariculture keyhole limpet hemocyanin. Human Tyr0-ghrelin [13–28] was radioiodinated by the lactoperoxidase method for use in the assay. Inter- and intra-assay variation was less than 8 and 6%, respectively. The limit of detection of this assay is 12 fmol per tube of human ghrelin. Two milliliters of plasma were diluted with 2 ml of 0.9% saline and applied to a Sep-Pak C-18 cartridge (Waters Corp., Milford, MA) pre-equilibrated with 0.9% saline. The cartridge was washed first with saline and then with a 0.1% trifluoroacetic acid solution, and peptides were eluted with a 60% acetonitrile (CH3CN) solution containing 0.1% trifluoroacetic acid. The eluate was evaporated, reconstituted with RIA buffer, and subjected to RIA analysis. A diluted sample or a standard peptide solution (100 µl) was incubated for 24 h with 100 µl of the antiserum diluent (final dilution, 1:20,000). The tracer solution (16,000 cpm/100 µl) was added, and the mixture was incubated for 24 h. Bound and free ligands were separated by the second antibody method. All procedures were performed at 4 C. Recovery of human ghrelin added to the plasma was 90.7 ± 4.0% (n = 6).
Statistical analyses
Groups of data (mean ± SEM) were compared using ANOVA and a post hoc Fisher’s test. A P value of less than 0.05 was considered statistically significant. Correction coefficients were calculated by linear regression analysis.
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Results |
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). A serial dilution curve of a human plasma sample was parallel to a standard curve for human ghrelin (Fig. 1).
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). The mean fasting plasma ghrelin concentration in these three groups of subjects was negatively correlated with BMI (Fig. 2B). Mean plasma ghrelin concentrations in lean and obese type 2 diabetic patients were 190% (P < 0.005) and 66% (P < 0.05) of those found in corresponding normal-weight diabetics (129.3 ± 14.2 pM) (Fig. 3A). Also observed in diabetic patients was the negative correlation of fasting plasma ghrelin concentration and BMI (Fig. 3B). There was no significant difference in plasma ghrelin concentrations between normal-weight healthy subjects and normal-weight type 2 diabetics. Also in both lean and obese subjects, there was no statistical difference in plasma ghrelin concentrations between diabetic and nondiabetic groups.
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). Normal responses were observed for plasma glucose (Fig. 4A) and insulin levels (data not shown). The mean plasma ghrelin concentration in the same subjects did not change in response to an administration of the same volume of water (Fig. 4A), but decreased rapidly after an iv glucose administration, reaching a nadir of 68% (93.1 ± 10.4 pM) of the basal level 15 min after administration and increasing thereafter (Fig. 4B). In the meal tolerance test, the mean plasma ghrelin concentration for diabetics decreased, reaching a nadir of 69% of the basal level 60 min after eating and increasing thereafter (Fig. 5). A 24-h plasma ghrelin profile from 10 normal subjects is shown in Fig. 6. The circulating plasma ghrelin level showed a diurnal pattern with preprandial increases and postprandial decreases during the daytime and a maximum peak at 0200 h.
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Discussion |
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Plasma ghrelin concentrations were higher in patients with anorexia nervosa and lower in patients with simple obesity compared with normal-weight control subjects. Tschöp et al. (19) have measured plasma ghrelin concentrations in normal and obese Caucasian and Pima Indian individuals. They reported that fasting plasma ghrelin concentration negatively correlates with percentage of body fat, and fasting insulin and leptin concentrations. We also found that fasting plasma ghrelin concentration in normal subjects and patients with anorexia nervosa, simple obesity, or type 2 DM correlated negatively with BMI within each group. If the action of ghrelin in humans is similar to that in rodents, ghrelin should stimulate feeding. Anorexia nervosa is an eating disorder in which patients have obsessive ideation about body weight. Reduced feeding in anorexia nervosa patients who have high plasma ghrelin concentrations suggests that they may have decreased sensitivity to circulating ghrelin, or that there may be a central system regulating energy homeostasis that overcomes the effect of ghrelin in such patients. Ghrelin stimulates GH release in humans (7, 8). Nutritional status plays an important role in the regulation of the GH axis, because GH secretion is markedly influenced by body composition (20, 21, 22). Although we did not investigate plasma GH levels in this study, the basal plasma ghrelin concentration probably correlates with that of GH in anorexia nervosa when we take into account the fact that plasma GH is elevated at baseline in this disorder (23, 24).
The stomach is the main source of circulating ghrelin in humans and rats. Ghrelin production in the stomach is localized to A (
)-like cells that do not have glucagon-immunoreactivity but share some morphological features with pancreatic -cells, including the presence of compact and dense secretory granules (3). Oral and iv administration of glucose to normal subjects decreased their plasma ghrelin concentrations, whereas intake of an equivalent volume of water did not. Also in rats, stomach filling with water did not change plasma ghrelin level (13). Secretion of ghrelin is not affected by stomach expansion. Insulin-induced hypoglycemia up-regulates ghrelin mRNA expression in the rat stomach (18). Taken together, these results indicate that there may be a system in ghrelin-producing cells that responds to plasma glucose concentration. Molecular signals that regulate ghrelin secretion are not known. Further investigation is needed to define the receptors, transporters, and/or channels expressed in ghrelin-producing cells.
Glucose load and food intake lead to a rapid fall in plasma ghrelin concentration, suggesting that plasma ghrelin reflects an acute feeding state and may also serve as an indicator of short-term energy balance. Plasma ghrelin levels are low in human obesity, and are generally associated with increased weight. Ghrelin mRNA expression in the gastric fundi of db/db mice, an obese model characterized by a null mutation in the leptin receptor gene (25, 26), is down-regulated compared with control mice (18). These alterations of ghrelin expression may be a physiological adaptation to long-term positive energy balance.
The diurnal variation of ghrelin secretion in normal subjects appears to be entrained to meal-taking and sleep. The temporal pattern of plasma ghrelin concentration consists of a rise just before the onset of meals and a postprandial decline during the daytime, followed by a much greater increase culminating at 0200 h. The physiological signals that initiate eating in humans are poorly understood. A preprandial rise in plasma ghrelin concentration suggests that ghrelin may be a candidate for a meal-initiation signal. A 24-h profile of human plasma GH concentration (20, 27) resembles that of ghrelin. Human plasma GH also decreases after meals and peaks at 0200–0300 h. Additional studies are needed to examine the potential link between ghrelin and GH diurnal changes in humans.
In summary, the present study demonstrates that nutritional state is a determinant of plasma ghrelin concentration in humans. Ghrelin secretion is up-regulated under conditions of negative energy balance and down-regulated in the setting of positive energy balance. These findings shed new light upon the involvement of the novel gastrointestinal peptide, ghrelin, in the regulation of feeding behavior and energy homeostasis.
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Acknowledgments |
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Footnotes |
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Abbreviations: BMI, Body mass index; DM, diabetes mellitus; GPCR, G protein-coupled cell surface receptor.
Received August 1, 2001.
Accepted September 24, 2001.
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K. Dezaki, H. Sone, M. Koizumi, M. Nakata, M. Kakei, H. Nagai, H. Hosoda, K. Kangawa, and T. Yada Blockade of Pancreatic Islet-Derived Ghrelin Enhances Insulin Secretion to Prevent High-Fat Diet-Induced Glucose Intolerance Diabetes, December 1, 2006; 55(12): 3486 - 3493. [Abstract] [Full Text] [PDF]
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 A. E. Wertz-Lutz, T. J. Knight, R. H. Pritchard, J. A. Daniel, J. A. Clapper, A. J. Smart, A. Trenkle, and D. C. Beitz Circulating ghrelin concentrations fluctuate relative to nutritional status and influence feeding behavior in cattle J Anim Sci, December 1, 2006; 84(12): 3285 - 3300. [Abstract] [Full Text] [PDF]
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E. Naslund and J. G. Kral Impact of Gastric Bypass Surgery on Gut Hormones and Glucose Homeostasis in Type 2 Diabetes Diabetes, December 1, 2006; 55(Supplement_2): S92 - S97. [Abstract] [Full Text] [PDF]
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Y. P Lee, T. A Mori, S. Sipsas, A. Barden, I. B Puddey, V. Burke, R. S Hall, and J. M Hodgson Lupin-enriched bread increases satiety and reduces energy intake acutely. Am. J. Clinical Nutrition, November 1, 2006; 84(5): 975 - 980. [Abstract] [Full Text] [PDF]
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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. [Abstract] [Full Text] [PDF]
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B. Otto, W. Heldwein, C. Otto, S. Huptas, and K. G Parhofer Effect of a high-fat meal on the postprandial ghrelin response. Am. J. Clinical Nutrition, September 1, 2006; 84(3): 664 - 665. [Full Text] [PDF]
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F. Levin, T. Edholm, P. T. Schmidt, P. Gryback, H. Jacobsson, M. Degerblad, C. Hoybye, J. J. Holst, J. F. Rehfeld, P. M. Hellstrom, et al. Ghrelin Stimulates Gastric Emptying and Hunger in Normal-Weight Humans J. Clin. Endocrinol. Metab., September 1, 2006; 91(9): 3296 - 3302. [Abstract] [Full Text] [PDF]
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T. J. Little, S. Doran, J. H. Meyer, A. J. P. M. Smout, D. G. O'Donovan, K.-L. Wu, K. L. Jones, J. Wishart, C. K. Rayner, M. Horowitz, et al. The release of GLP-1 and ghrelin, but not GIP and CCK, by glucose is dependent upon the length of small intestine exposed Am J Physiol Endocrinol Metab, September 1, 2006; 291(3): E647 - E655. [Abstract] [Full Text] [PDF]
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 E. P. Zorrilla, S. Iwasaki, J. A. Moss, J. Chang, J. Otsuji, K. Inoue, M. M. Meijler, and K. D. Janda From the Cover: Vaccination against weight gain PNAS, August 29, 2006; 103(35): 13226 - 13231. [Abstract] [Full Text] [PDF]
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J. Bowen, M. Noakes, and P. M. Clifton Appetite Regulatory Hormone Responses to Various Dietary Proteins Differ by Body Mass Index Status Despite Similar Reductions in ad Libitum Energy Intake J. Clin. Endocrinol. Metab., August 1, 2006; 91(8): 2913 - 2919. [Abstract] [Full Text] [PDF]
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D.-Y. Lee, S.-Y. Kim, D.-S. Jo, P. H. Hwang, K. P. Kang, S. Lee, W. Kim, and S. K. Park Preproghrelin Leu72Met polymorphism predicts a lower rate of developing renal dysfunction in type 2 diabetic nephropathy. Eur. J. Endocrinol., July 1, 2006; 155(1): 187 - 190. [Abstract] [Full Text] [PDF]
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S. S. Damjanovic, N. M. Lalic, P. M. Pesko, M. S. Petakov, A. Jotic, D. Miljic, K. S. Lalic, L. Lukic, M. Djurovic, and V. B. Djukic Acute Effects of Ghrelin on Insulin Secretion and Glucose Disposal Rate in Gastrectomized Patients J. Clin. Endocrinol. Metab., July 1, 2006; 91(7): 2574 - 2581. [Abstract] [Full Text] [PDF]
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M. E. Gordon and K. H. McKeever Oral and intravenous carbohydrate challenges decrease active ghrelin concentrations and alter hormones related to control of energy metabolism in horses J Anim Sci, July 1, 2006; 84(7): 1682 - 1690. [Abstract] [Full Text] [PDF]
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J. Dong, T. L. Peeters, B. De Smet, D. Moechars, C. Delporte, P. Vanden Berghe, B. Coulie, M. Tang, and I. Depoortere Role of Endogenous Ghrelin in the Hyperphagia of Mice with Streptozotocin-Induced Diabetes Endocrinology, June 1, 2006; 147(6): 2634 - 2642. [Abstract] [Full Text] [PDF]
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R. M. Luque and R. D. Kineman Impact of Obesity on the Growth Hormone Axis: Evidence for a Direct Inhibitory Effect of Hyperinsulinemia on Pituitary Function Endocrinology, June 1, 2006; 147(6): 2754 - 2763. [Abstract] [Full Text] [PDF]
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L. C Gormsen, J. Gjedsted, S. Gjedde, E. T. Vestergaard, J. S Christiansen, J. O. Jorgensen, S. Nielsen, and N. Moller Free fatty acids decrease circulating ghrelin concentrations in humans. Eur. J. Endocrinol., May 1, 2006; 154(5): 667 - 673. [Abstract] [Full Text] [PDF]
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G. Mingrone, L. Granato, E. Valera-Mora, A. Iaconelli, M. F Calvani, R. Bracaglia, M. Manco, G. Nanni, and M. Castagneto Ultradian ghrelin pulsatility is disrupted in morbidly obese subjects after weight loss induced by malabsorptive bariatric surgery Am. J. Clinical Nutrition, May 1, 2006; 83(5): 1017 - 1024. [Abstract] [Full Text] [PDF]
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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. [Abstract] [Full Text] [PDF]
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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. [Abstract] [Full Text] [PDF]
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H Takahashi, Y Kurose, S Kobayashi, T Sugino, M Kojima, K Kangawa, Y Hasegawa, and Y Terashima Ghrelin enhances glucose-induced insulin secretion in scheduled meal-fed sheep. J. Endocrinol., April 1, 2006; 189(1): 67 - 75. [Abstract] [Full Text] [PDF]
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J. Bowen, M. Noakes, C. Trenerry, and P. M. Clifton Energy Intake, Ghrelin, and Cholecystokinin after Different Carbohydrate and Protein Preloads in Overweight Men J. Clin. Endocrinol. Metab., April 1, 2006; 91(4): 1477 - 1483. [Abstract] [Full Text] [PDF]
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E. R Christ, M. Zehnder, C. Boesch, R. Trepp, P. E Mullis, P. Diem, and J. Decombaz The effect of increased lipid intake on hormonal responses during aerobic exercise in endurance-trained men. Eur. J. Endocrinol., March 1, 2006; 154(3): 397 - 403. [Abstract] [Full Text] [PDF]
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M. Romon, S. Gomila, P. Hincker, B. Soudan, and J. Dallongeville Influence of Weight Loss on Plasma Ghrelin Responses to High-Fat and High-Carbohydrate Test Meals in Obese Women J. Clin. Endocrinol. Metab., March 1, 2006; 91(3): 1034 - 1041. [Abstract] [Full Text] [PDF]
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W. A. Blom, A. Lluch, A. Stafleu, S. Vinoy, J. J Holst, G. Schaafsma, and H. F. Hendriks Effect of a high-protein breakfast on the postprandial ghrelin response Am. J. Clinical Nutrition, February 1, 2006; 83(2): 211 - 220. [Abstract] [Full Text] [PDF]
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S. C. Griffen, K. Oostema, K. L. Stanhope, J. Graham, D. M. Styne, N. Glaser, D. E. Cummings, M. H. Connors, and P. J. Havel Administration of Lispro Insulin with Meals Improves Glycemic Control, Increases Circulating Leptin, and Suppresses Ghrelin, Compared with Regular/NPH Insulin in Female Patients with Type 1 Diabetes J. Clin. Endocrinol. Metab., February 1, 2006; 91(2): 485 - 491. [Abstract] [Full Text] [PDF]
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W. A. M. Blom, A. Lluch, S. Vinoy, A. Stafleu, R. van den Berg, J. J. Holst, F. J. Kok, and H. F. J. Hendriks Effects of gastric emptying on the postprandial ghrelin response Am J Physiol Endocrinol Metab, February 1, 2006; 290(2): E389 - E395. [Abstract] [Full Text] [PDF]
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A. Doi, T. Shono, M. Nishi, H. Furuta, H. Sasaki, and K. Nanjo IA-2beta, but not IA-2, is induced by ghrelin and inhibits glucose-stimulated insulin secretion PNAS, January 24, 2006; 103(4): 885 - 890. [Abstract] [Full Text] [PDF]
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 M. J. Kleinz, J. J. Maguire, J. N. Skepper, and A. P. Davenport Functional and immunocytochemical evidence for a role of ghrelin and des-octanoyl ghrelin in the regulation of vascular tone in man Cardiovasc Res, January 1, 2006; 69(1): 227 - 235. [Abstract] [Full Text] [PDF]
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C. Langenberg, J. Bergstrom, G. A. Laughlin, and E. Barrett-Connor Ghrelin and the Metabolic Syndrome in Older Adults J. Clin. Endocrinol. Metab., December 1, 2005; 90(12): 6448 - 6453. [Abstract] [Full Text] [PDF]
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C. Feinle-Bisset, M. Patterson, M. A. Ghatei, S. R. Bloom, and M. Horowitz Fat digestion is required for suppression of ghrelin and stimulation of peptide YY and pancreatic polypeptide secretion by intraduodenal lipid Am J Physiol Endocrinol Metab, December 1, 2005; 289(6): E948 - E953. [Abstract] [Full Text] [PDF]
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 M. Tesauro, F. Schinzari, M. Iantorno, S. Rizza, D. Melina, D. Lauro, and C. Cardillo Ghrelin Improves Endothelial Function in Patients With Metabolic Syndrome Circulation, November 8, 2005; 112(19): 2986 - 2992. [Abstract] [Full Text] [PDF]
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P. Koutkia, S. Schurgin, J. Berry, J. Breu, B. S. H. Lee, A. Klibanski, and S. Grinspoon Reciprocal changes in endogenous ghrelin and growth hormone during fasting in healthy women Am J Physiol Endocrinol Metab, November 1, 2005; 289(5): E814 - E822. [Abstract] [Full Text] [PDF]
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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. [Abstract] [Full Text] [PDF]
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E T Vestergaard, T K Hansen, S Nielsen, N Moller, J S Christiansen, and J O L Jorgensen Effects of GH replacement therapy in adults on serum levels of leptin and ghrelin: the role of lipolysis Eur. J. Endocrinol., October 1, 2005; 153(4): 545 - 549. [Abstract] [Full Text] [PDF]
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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. [Abstract] [Full Text] [PDF]
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Y. H. Choe, S. Y. Song, K.-H. Paik, Y. J. Oh, S.-H. Chu, S. H. Yeo, E. K. Kwon, E. M. Kim, M. Y. Rha, and D.-K. Jin Increased Density of Ghrelin-Expressing Cells in the Gastric Fundus and Body in Prader-Willi Syndrome J. Clin. Endocrinol. Metab., September 1, 2005; 90(9): 5441 - 5445. [Abstract] [Full Text] [PDF]
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C. Daousi, I. A. MacFarlane, P. J. English, J. P. H. Wilding, M. Patterson, T. M. Dovey, J. C. G. Halford, M. A. Ghatei, and J. H. Pinkney 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., September 1, 2005; 90(9): 5025 - 5030. [Abstract] [Full Text] [PDF]
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L. J. Moran, N. D. Luscombe-Marsh, M. Noakes, G. A. Wittert, J. B. Keogh, and P. M. Clifton The Satiating Effect of Dietary Protein Is Unrelated to Postprandial Ghrelin Secretion J. Clin. Endocrinol. Metab., September 1, 2005; 90(9): 5205 - 5211. [Abstract] [Full Text] [PDF]
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Y. Date, K. Toshinai, S. Koda, M. Miyazato, T. Shimbara, T. Tsuruta, A. Niijima, K. Kangawa, and M. Nakazato Peripheral Interaction of Ghrelin with Cholecystokinin on Feeding Regulation Endocrinology, August 1, 2005; 146(8): 3518 - 3525. [Abstract] [Full Text] [PDF]
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 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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 A. Masuda, M. Nakazato, T. Kihara, S. Minagoe, and C. Tei Repeated Thermal Therapy Diminishes Appetite Loss and Subjective Complaints in Mildly Depressed Patients Psychosom Med, July 1, 2005; 67(4): 643 - 647. [Abstract] [Full Text] [PDF]
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D. Yasuhara, T. Naruo, N. Nagai, T. Muranaga, T. Nakahara, M. Tanaka, S. Kojima, K.-i. Sagiyama, A. Masuda, and A. Inui Glucose Tolerance Predicts Short-term Refeeding Outcome in Females With Anorexia Nervosa Psychosom Med, July 1, 2005; 67(4): 669 - 676. [Abstract] [Full Text] [PDF]
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L. F. Canosa, S. Unniappan, and R. E. Peter Periprandial changes in growth hormone release in goldfish: role of somatostatin, ghrelin, and gastrin-releasing peptide Am J Physiol Regulatory Integrative Comp Physiol, July 1, 2005; 289(1): R125 - R133. [Abstract] [Full Text] [PDF]
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G Natalucci, S Riedl, A Gleiss, T Zidek, and H Frisch Spontaneous 24-h ghrelin secretion pattern in fasting subjects: maintenance of a meal-related pattern Eur. J. Endocrinol., June 1, 2005; 152(6): 845 - 850. [Abstract] [Full Text] [PDF]
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Y. Nishi, H. Hiejima, H. Mifune, T. Sato, K. Kangawa, and M. Kojima Developmental Changes in the Pattern of Ghrelin's Acyl Modification and the Levels of Acyl-Modified Ghrelins in Murine Stomach Endocrinology, June 1, 2005; 146(6): 2709 - 2715. [Abstract] [Full Text] [PDF]
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T. Sato, Y. Fukue, H. Teranishi, Y. Yoshida, and M. Kojima Molecular Forms of Hypothalamic Ghrelin and Its Regulation by Fasting and 2-Deoxy-D-Glucose Administration Endocrinology, June 1, 2005; 146(6): 2510 - 2516. [Abstract] [Full Text] [PDF]
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M. Groschl, H. G. Topf, J. Bohlender, J. Zenk, S. Klussmann, J. Dotsch, W. Rascher, and M. Rauh Identification of Ghrelin in Human Saliva: Production by the Salivary Glands and Potential Role in Proliferation of Oral Keratinocytes Clin. Chem., June 1, 2005; 51(6): 997 - 1006. [Abstract] [Full Text] [PDF]
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J. M. Garcia, M. Garcia-Touza, R. A. Hijazi, G. Taffet, D. Epner, D. Mann, R. G. Smith, G. R. Cunningham, and M. Marcelli Active Ghrelin Levels and Active to Total Ghrelin Ratio in Cancer-Induced Cachexia J. Clin. Endocrinol. Metab., May 1, 2005; 90(5): 2920 - 2926. [Abstract] [Full Text] [PDF]
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E. Kellokoski, S. M. Poykko, A. H. Karjalainen, O. Ukkola, J. Heikkinen, Y. A. Kesaniemi, and S. Horkko Estrogen Replacement Therapy Increases Plasma Ghrelin Levels J. Clin. Endocrinol. Metab., May 1, 2005; 90(5): 2954 - 2963. [Abstract] [Full Text] [PDF]
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F. Bacha and S. A. Arslanian Ghrelin Suppression in Overweight Children: A Manifestation of Insulin Resistance? J. Clin. Endocrinol. Metab., May 1, 2005; 90(5): 2725 - 2730. [Abstract] [Full Text] [PDF]
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S. Kanumakala, R. Greaves, C. C. Pedreira, S. Donath, G. L. Warne, M. R. Zacharin, and M. Harris Fasting Ghrelin Levels Are Not Elevated in Children with Hypothalamic Obesity J. Clin. Endocrinol. Metab., May 1, 2005; 90(5): 2691 - 2695. [Abstract] [Full Text] [PDF]
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J. Erdmann, F. Lippl, S. Wagenpfeil, and V. Schusdziarra Differential Association of Basal and Postprandial Plasma Ghrelin With Leptin, Insulin, and Type 2 Diabetes Diabetes, May 1, 2005; 54(5): 1371 - 1378. [Abstract] [Full Text] [PDF]
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D. L. Williams and D. E. Cummings Regulation of Ghrelin in Physiologic and Pathophysiologic States J. Nutr., May 1, 2005; 135(5): 1320 - 1325. [Abstract] [Full Text] [PDF]
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A. Geliebter, M. E. Gluck, and S. A. Hashim Plasma Ghrelin Concentrations Are Lower in Binge-Eating Disorder J. Nutr., May 1, 2005; 135(5): 1326 - 1330. [Abstract] [Full Text] [PDF]
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), plasma extract (•), and insulin, glucagon, somatostatin, pancreatic polypeptide, neuropeptide Y, and motilin (
). The numeral 1 in the plasma dilution curve denotes 2 ml.
) in normal subjects. *P < 0.0001 in plasma ghrelin vs. 0 min. B, Plasma ghrelin response to 10 g iv glucose tolerance test in normal subjects. *, P < 0.05, **, P < 0.005 in plasma ghrelin vs. 0 min. The results are represented as percentages of the basal level.
