Hedonic and incentive signals for body weight control

Reviews in Endocrine and Metabolic Disorders

Volumn 12, Issue 3, 2011, Pages 141-151

  Egecioglu, Emil ^a   Skibicka, Karolina P ^a   Hansson, Caroline ^a   Alvarez Crespo, Mayte ^a   Anders Friberg, P ^a   Jerlhag, Elisabet ^a   Engel, Jörgen A ^a   Dickson, Suzanne L ^a  

^a UNIVERSITY OF GOTHENBURG   (Sweden)

Author keywords

Appetite; Food reward; Ghrelin; Liking; Obesity; Wanting

Indexed keywords

ANTIOBESITY AGENT; DOPAMINE; GHRELIN; GHRELIN RECEPTOR; GHRELIN RECEPTOR ANTAGONIST; INSULIN; LEPTIN; MU OPIATE RECEPTOR; PEPTIDE YY [3-36]; UNCLASSIFIED DRUG;

ARTICLE; BODY WEIGHT; DOPAMINE METABOLISM; FOOD INTAKE; HOMEOSTASIS; HUMAN; HYPOTHALAMUS; NEUROBIOLOGY; NONHUMAN; OBESITY; PALATABILITY; SIGNAL TRANSDUCTION;

ANIMALS; APPETITE; BODY WEIGHT; BRAIN; DOPAMINE; EATING; FOOD PREFERENCES; HUMANS; OBESITY; REWARD;

EID: 80054866688     PISSN: 13899155     EISSN: 15732606     Source Type: Journal    
DOI: 10.1007/s11154-011-9166-4     Document Type: Article

References (127)

1. Berthoud HR, Morrison C. The brain, appetite, and obesity. Annu Rev Psychol. 2008;59:55-92.
2. Yeomans MR, Blundell JE, Leshem M. Palatability: response to nutritional need or need-free stimulation of appetite? Br J Nutr. 2004;92:S3-S14. (Pubitemid 39214254)
3. Zheng HY, Berthoud HR. Neural systems controlling the drive to eat: mind versus metabolism. Physiology. 2008;23(2):75-83.
4. Berridge KC, Robinson TE. What is the role of dopamine in reward: hedonic impact, reward learning, or incentive salience? Brain Res Rev. 1998;28(3):309-69. (Pubitemid 29012677)
5. Mela DJ. Eating for pleasure or just wanting to eat? Reconsidering sensory hedonic responses as a driver ofobesity. Appetite. 2006;47 (1):10-7. (Pubitemid 44038034)
6. Berridge KC. Food reward: brain substrates of wanting and liking. Neurosci Biobehav Rev. 1996;20(1):1-25. (Pubitemid 126682107)
7. Finlayson G, King N, Blundell JE. Is it possible to dissociate 'liking' and 'wanting' for foods in humans? A novel experimen-tal procedure. Physiol Behav. 2007;90(1):36-42.
8. Avena NM, Rada P, Hoebel BG. Evidence for sugar addiction: behavioral and neurochemical effects of intermit-tent, excessive sugar intake. Neurosci Biobehav Rev. 2008;32 (1):20-39. (Pubitemid 350160955)
9. Benton D. The plausibility of sugar addiction and its role in obesity and eating disorders. Clin Nutr. 2010;29(3):288-303.
10. Berthoud HR. Interactions between the "cognitive" and "metabolic'̀ brain in the control offood intake. Physiol Behav. 2007;91(5):486-98.
11. Stice E, Spoor S, Ng J, Zald DH. Relation of obesity to consummatory and anticipatory food reward. Physiol Behav. 2009;97(5):551-60.
12. Vucetic Z, Reyes TM. Central dopaminergic circuitry controlling food intake and reward: implications for the regulation of obesity Wiley Interdiscip Rev Syst Biol Med. 2010;2:577-93.
13. Berridge KC. The debate over dopamine's role in reward: the case forincentive salience. Psychopharmacology 2007;191(3):391-431.
14. Wise RA, Bozarth MA. A psychomotor stimulant theory of addiction. Psychol Rev. 1987;94(4):469-92.
15. Hajnal A, Smith GP, Norgren R. Oral sucrose stimulation increases accumbens dopamine in the rat. Am J Physiol Regul Integr Comp Physiol. 2004;286(1):R31-7. (Pubitemid 38055888)
16. Rada P, Avena NM, Hoebel BG. Daily bingeing on sugar repeatedly releases dopamine in the accumbens shell. Neurosci-ence. 2005;134(3):737-44. (Pubitemid 41133378)
17. Liang NC, Hajnal A, Norgren R. Sham feeding corn oil increases accumbens dopamine in the rat. Am J Physiol Regul Integr Comp Physiol. 2006;291(5):R1236-9.
18. Reilly S. Reinforcement value ofgustatory stimuli determined by progressive ratio performance. Pharmacol Biochem Behav. 1999;63(2):301-11. (Pubitemid 29234110)
19. Sclafani A, Ackroff K. Reinforcement value of sucrose measured by progressive ratio operant licking in the rat. Physiol Behav. 2003;79(4-5):663-70. (Pubitemid 37087959)
20. Wojnicki FHE, Babbs RK, Corwin RLW. Reinforcing efficacy of fat, as assessed by progressive ratio responding, depends upon availability not amount consumed. Physiol Behav. 2010;100 (4):316-21.
21. Yoneda T, Taka Y, Okamura M, Mizushige T, Matsumura S, Manabe Y, et al. Reinforcing effect for corn oil stimulus was concentration dependent in an operant task in mice. Life Sci. 2007;81(23-24):1585-92. (Pubitemid 350116199)
22. Baldo BA, Kelley AE. Discrete neurochemical coding of distin-guishable motivational processes: insights from nucleus accumbens controloffeeding. Psychopharmacology. 2007;191(3):439-59. (Pubitemid 46355910)
23. Salamone JD, Cousins MS, Snyder BJ. Behavioral functions of nucleus accumbens dopamine: empirical and conceptual prob-lems with the anhedonia hypothesis. Neurosci Biobehav Rev. 1997;21(3):341-59. (Pubitemid 27225155)
24. Berridge KC, Venier IL, Robinson TE. Taste reactivity analysis of 6-hydroxydopamine-induced aphagia-implications for arousal and anhedonia hypotheses ofdopamine function. Behav Neurosci. 1989;103(1):36-45. (Pubitemid 19058642)
25. Cagniard B, Balsam PD, Brunner D, Zhuang XX. Mice with chronically elevated dopamine exhibit enhanced motivation, but not learning, for a food reward. Neuropsychopharmacol. 2006;31 (7):1362-70.
26. Pecina S, Cagniard B, Berridge KC, Aldridge JW, Zhuang XX. Hyperdopaminergic mutant mice have higher "wanting'̀ but not "liking" for sweet rewards. J Neurosci. 2003;23 (28):9395-402. (Pubitemid 37323039)
27. De Araujo IE, Oliveira-Maia AJ, Sotnikova TD, Gainetdinov RR, Caron MG, Nicolelis MAL, et al. Food reward in the absence of taste receptor signaling. Neuron. 2008;57(6):930-41. (Pubitemid 351400637)
28. Grill HJ, Norgren R. Taste reactivity test.2. Mimetic responses to gustatory stimuli in chronic thalamic and chronic decerebrate rats. Brain Res. 1978;143(2):281-97. (Pubitemid 8277910)
29. Pecina S, Berridge KC. Opioid site in nucleus accumbens shell mediates eating and hedonic 'liking' for food: map based on microinjection Fos plumes. Brain Res. 2000;863(1-2):71-86. (Pubitemid 30201586)
30. Berridge KC, Robinson TE. Parsing reward. Trends Neurosci. 2003;26(9):507-13. (Pubitemid 37025790)
31. Zhang M, Kelley AE. Intake of saccharin, salt, and ethanol solutions is increased by infusion of a mu opioid agonist into the nucleus accumbens. Psychopharmacology. 2002;159(4):415-23. (Pubitemid 34143366)
32. Zhang M, Gosnell BA, Kelley AE. Intake of high-fat food is selectively enhanced by mu opioid receptor stimulation within the nucleus accumbens. J Pharmacol Exp Ther. 1998;285(2):908-14. (Pubitemid 28237478)
33. Pecina S, Berridge KC. Hedonic hot spot in nucleus accumbens shell: where do mu-opioids cause increased hedonic impact of sweetness? J Neurosci. 2005;25(50):11777-86. (Pubitemid 43098470)
34. Shin AC, Pistell PJ, Phifer CB, Berthoud HR. Reversible suppression of food reward behavior by chronic mu-opioid receptor antagonism in the nucleus accumbens. Neuroscience. 2010;170:580-8.
35. Zhang M, Balmadrid C, Kelley AE. Nucleus accumbens opioid, GABAergic, and dopaminergic modulation of palatable food motivation: contrasting effects revealed by a progressive ratio study in the rat Behav Neurosci. 2003;117(2):202-11. (Pubitemid 36384239)
36. Jamshidi N, Taylor DA. Anandamide administration into the ventromedial hypothalamus stimulates appetite in rats. Br J Pharmacol. 2001;134(6):1151-4.
37. Verty ANA, Mc Gregor IS, Mallet PE. Paraventricular hypotha-lamic CB1 cannabinoid receptors are involved in the feeding stimulatory effects of Delta(9)-tetrahydrocannabinol. Neurophar-macology 2005;49(8):1101-9. (Pubitemid 41608634)
38. Di Patrizio NV, Simansky KJ. Activating parabrachial cannabi-noid CB1 receptors selectively stimulates feeding of palatable foods in rats. J Neurosci. 2008;28(39):9702-9.
39. Soria-Gomez E, Matias I, Rueda-Orozco PE, Cisneros M, Petrosino S, Navarro L, et al. Pharmacological enhancement of the endocannabinoid system in the nucleus accumbens shell stimulates food intake and increases c-Fos expression in the hypothalamus. Br J Pharmacol. 2007;151(7):1109-16. (Pubitemid 47195529)
40. Perio A, Barnouin MC, Poncelet M, Soubrie P. Activity of SR141716 on post-reinforcement pauses in operant responding for sucrose reward in rats. Behav Pharmacol. 2001;12(8):641-5. (Pubitemid 34017111)
41. Kirkham TC, Williams CM. Synergistic effects of opioid and cannabinoid antagonists on food intake. Psychopharmacology. 2001;153(2):267-70. (Pubitemid 32127898)
42. Verty ANA, Mc Gregor IS, Mallet PE. The doparnine receptor antagonist SCH 23390 attenuates feeding induced by Delta(9)-tetrahydrocannobinol. Brain Res. 2004;1020(1-2):188-95. (Pubitemid 39092986)
43. Harris GC, Wimmer M, Aston-Jones G. A role for lateral hypothalamic orexin neurons in reward seeking. Nature. 2005;437(7058):556-9. (Pubitemid 41613553)
44. Cason AM, Smith RJ, Tahsili-Fahadan P, Moorman DE, Sartor GC, Aston-Jones G. Role of orexin/hypocretin in reward-seeking and addiction: Implications for obesity. Physiol Behav. 2010;100 (5):419-28.
45. Crespo I, de Heras RG, de Fonseca FR, Navarro M. Pretreatment with subeffective doses of Rimonabant attenuates orexigenic actions of orexin A-hypocretin 1. Neuropharmacology. 2008;54 (1):219-25.
46. Ramirez I, Friedman MI. Dietary hyperphagia in rats-role of fat, carbohydrate, and energy content Physiol Behav. 1990;47 (6):1157-63. (Pubitemid 20224217)
47. Sclafani A. Oral and postoral determinants of food reward. Physiol Behav. 2004;81(5):773-9. (Pubitemid 38845045)
48. Finlayson G, Caudwell P, Gibbons C, Hopkins M, King N, Blundell J. Low fat loss response after medium-term supervised exercise in obese is associated with exercise-induced increase in food reward. J Obes. (Epub 2010 Sep 20). 2011.
49. Davis C, Strachan S, Berkson M. Sensitivity to reward: implications for overeating and overweight. Appetite. 2004;42(2):131-8. (Pubitemid 38316882)
50. Rissanen A, Hakala P, Lissner L, Mattlar CE, Koskenvuo M, Ronnemaa T. Acquired preference especially for dietary fat and obesity: a study ofweight-discordant monozygotic twin pairs. Int J Obes. 2002;26(7):973-7. (Pubitemid 34756181)
51. Saelens BE, Epstein LH. Reinforcing value offood in obese and non-obese women. Appetite. 1996;27(1):41-50. (Pubitemid 26313170)
52. Comings DE, Blum K. Reward deficiency syndrome: genetic aspects ofbehavioral disorders. Prog Brain Res. 2000;126:325-41. (Pubitemid 35461297)
53. Carroll ME, Lac ST. Dietary additives and the acquisition of cocaine self-administration in rats. Psychopharmacology. 1998;137(1):81-9. (Pubitemid 28225007)
54. Wellman PJ, Nation JR, Davis KW. Impairment of acquisition of cocaine self-administration in rats maintained on a high-fat diet. Pharmacol Biochem Behav. 2007;88(1):89-93. (Pubitemid 47379826)
55. La Fleur SE, Vanderschuren LJMJ, Luijendijk MC, Kloeze BM, Tiesjema B, Adan RAH. A reciprocal interaction between food-motivated behavior and diet-induced obesity. Int J Obes. 2007;31 (8):1286-94. (Pubitemid 47172615)
56. Pickering C, Alsio J, Hulting AL, Schioth HB. Withdrawal from free-choice high-fat high-sugar diet induces craving only in obesity-prone animals. Psychopharmacology. 2009;204(3):431-43.
57. Johnson PM, Kenny PJ. Dopamine D2 receptors in addiction-like reward dysfunction and compulsive eating in obese rats. Nat Neurosci. 2010;13(5):635-41.
58. Alsiö J, Olszewski PK, Norbäck AH, Gunnarsson ZEA, Levine AS, Pickering C, Schiöth HB. Dopamine D1 receptor gene expression decreases in the nucleus accumbens upon long-term exposure to palatable food and differs depending on diet-induced obesity phenotype in rats. Neuroscience. 2010;171(3):779-87.
59. Geiger BM, Haburcak M, Avena NM, Moyer MC, Hoebel BG, Pothos EN. Deficits of mesolimbic dopamine neurotransmission in rat dietary obesity. Neuroscience. 2009;159(4):1193-9.
60. Davis JF, Tracy AL, Schurdak JD, Tschop MH, Lipton JW, Clegg DJ, et al. Exposure to elevated levels of dietary fat attenuates psychostimulant reward and mesolimbic dopamine turnover in the rat Behav Neurosci. 2008;122(6):1257-63.
61. Liu YJ, Gao JH, Liu HL, Fox PT. The temporal response of the brain after eating revealed by functional MRI. Nature. 2000;405 (6790):1058-62. (Pubitemid 30447790)
62. Morris JS, Dolan RJ. Involvement of human amygdala and orbitofrontal cortex in hunger-enhanced memory for food stimuli. J Neurosci. 2001;21(14):5304-10. (Pubitemid 32622956)
63. Passamonti L, Rowe JB, Schwarzbauer C, Ewbank MP, von dem Hagen E, Calder AJ. Personality predicts the brain's response to viewing appetizing foods: the neural basis of a risk factor for overeating. J Neurosci. 2009;29(1):43-51.
64. Rothemund Y, Preuschhof C, Bohner G, Bauknecht HC, Klingebiel R, Flor H, et al. Differential activation of the dorsal striatum by high-calorie visual food stimuli in obese individuals. Neuroimage. 2007;37(2):410-21. (Pubitemid 47444263)
65. Stoeckel LE, Weller RE, Cook EW, Twieg DB, Knowlton RC, Cox JE. Widespread reward-system activation in obese women in response to pictures of high-calorie foods. Neuroimage. 2008;41(2):636-47.
66. Stice E, Yokum S, Bohon C, Marti N, Smolen A. Reward circuitry responsivity to food predicts future increases in body mass: moderating effects of DRD2 and DRD4. Neuroimage. 2010;50(4):1618-25.
67. Stice E, Spoor S, Bohon C, Small DM. Relation between obesity and blunted striatal response to food is moderated by Taq IA A1 allele. Science. 2008;322(5900):449-52.
68. Wang GJ, Volkow ND, Logan J, Pappas NR, Wong CT, Zhu W, et al. Brain dopamine and obesity. Lancet. 2001;357(9253):354-7. (Pubitemid 32128542)
69. Volkow ND, Wang GJ, Telang F, Fowler JS, Thanos PK, Logan J, et al. Low dopamine striatal D2 receptors are associated with prefrontal metabolism in obese subjects: possible contributing factors. Neuroimage. 2008;42(4):1537-43.
70. Epstein LH, Jaroni JL, Paluch RA, Leddy JJ, Vahue HE, Hawk L, et al. Dopamine transporter genotype as a risk factor for obesity in African-American smokers. Obes Res. 2002;10 (12):1232-40.
71. Annerbrink K, Westberg L, Nilsson S, Rosmond R, Holm G, Eriksson E. Catechol O-methyltransferase val158-met polymor-phism is associated with abdominal obesity and blood pressure in men. Metabolism. 2008;57(5):708-11. (Pubitemid 351608233)
72. Comings DE, Flanagan SD, Dietz G, Muhleman D, Knell E, Gysin R. The dopamine-D(2) receptor (Drd2) as a major gene in obesity and height. Biochem Med Metabol Biol. 1993;50(2):176-85. (Pubitemid 23295784)
73. Hajnal A, Margas WM, Covasa M. Altered dopamine D2 receptor function and binding in obese OLETF rat. Brain Res Bull. 2008;75(1):70-6. (Pubitemid 350297644)
74. Olbers T, Bjorkman S, Lindroos A, Maleckas A, Lonn L, Sjostrom L, et al. Body composition, dietary intake, and energy expenditure after laparoscopic Roux-en-y gastric bypass and laparoscopic vertical banded gastroplasty-A randomized clini-cal trial. Ann Surg. 2006;244(5):715-22. (Pubitemid 44629945)
75. Zheng HY, Shin AC, Townsend RL, Patterson LM, Sigalet DL, Berthoud HR. Changes in fat preference, meal patterns, gastrointestinal transit, energy expenditure, and circulating hormones in a rat model of Roux-en-Y gastric bypass surgery. Obesity. 2009;17:S103-S.
76. Shin AC, Zheng H, Pistell PJ, Berthoud HR. Roux-en-Y gastric bypass surgery changes food reward in rats. Int J Obes (Lond). 2010. doi:10.1038/ijo.2010.174.
77. Cabanac M, Duclaux R. Specificity ofinternal signals in producing satiety fortaste stimuli. Nature. 1970;227(5261):966-7.
78. De Jonghe BC, Hajnal A, Covasa M. Conditioned preference for sweet stimuli in OLETF rat: effects of food deprivation. Am J Physiol Regul Integr Comp Physiol. 2007;292(5):R1819-27.
79. Figlewicz DP, Bennett JL, Naleid AM, Davis C, Grimm JW. Intraventricular insulin and leptin decrease sucrose self-administration in rats. Physiol Behav. 2006;89(4):611-6. (Pubitemid 44634584)
80. Figlewicz DP, Naleid AM, Sipols AJ. Modulation offood reward by adiposity signals. Physiol Behav. 2007;91(5):473-8. (Pubitemid 47247779)
81. Figlewicz DP, Benoit SC. Insulin, leptin, and food reward: update 2008. Am J Physiol Regul Integr Comp Physiol. 2009;296(1):R9-R19.
82. Berridge KC. Modulation of taste affect by hunger, caloric satiety, and sensory-specific satiety in the rat Appetite. 1991;16 (2):103-20.
83. Small DM, Zatorre RJ, Dagher A, Evans AC, Jones-Gotman M. Changes in brain activity related to eating chocolate-From pleasure to aversion. Brain. 2001;124:1720-33. (Pubitemid 32845434)
84. Kelley AE, Berridge KC. The neuroscience of natural rewards: relevance to addictive drugs. J Neurosci. 2002;22(9):3306-11. (Pubitemid 37465733)
85. Cornell CE, Rodin J, Weingarten H. Stimulus-induced eating when satiated. Physiol Behav. 1989;45(4):695-704. (Pubitemid 19134575)
86. Farooqi IS, Bullmore E, Keogh J, Gillard J, O'Rahilly S, Fletcher PC Leptin regulates striatal regions and human eating behavior. Science. 2007;317(5843):1355. (Pubitemid 47417470)
87. Zheng H, Lenard NR, Shin AC, Berthoud HR. Appetite control and energy balance regulation in the modern world: reward-driven brain overrides repletion signals. Int J Obes. 2009;33:S8-S13.
88. Shin AC, Zheng HY, Berthoud HR. An expanded view of energy homeostasis: neural integration of metabolic, cognitive, and emotional drives to eat. Physiol Behav. 2009;97(5):572-80.
89. Fulton S. Appetite and reward. Front Neuroendocrinol. 2010;31 (1):85-103.
90. Figlewicz DP. Adiposity signals and food reward: expanding the CNS roles of insulin and leptin. Am J Physiol Regul Integr Comp Physiol. 2003;284(4):R882-92. (Pubitemid 36406846)
91. Opland DM, Leinninger GM, Myers MG. Modulation of the mesolimbic dopamine system by leptin. Brain Res. 2010;1350: 65-70.
92. Figlewicz DP, Higgins MS, Ng-Evans SB, Havel PJ. Leptin reverses sucrose-conditioned place preference in food-restricted rats. Physiol Behav. 2001;73(1-2):229-34. (Pubitemid 32530562)
93. Figlewicz DP, Bennett J, Evans SB, Kaiyala K, Sipols AJ, Benoit SC. Intraventricular insulin and leptin reverse place preference conditioned with high-fat diet in rats. Behav Neurosci. 2004;118 (3):479-87. (Pubitemid 38685315)
94. Figlewicz DP, Evans SB, Murphy J, Hoen M, Baskin DG. Expression of receptors for insulin and leptin in the ventral tegmental area/substantia nigra (VTA/SN) of the rat Brain Res. 2003;964(1):107-15. (Pubitemid 36165075)
95. Hommel JD, Trinko R, Sears RM, Georgescu D, Liu ZW, Gao XB, et al. Leptin receptor signaling in midbrain dopamine neurons regulates feeding. Neuron. 2006;51(6):801-10. (Pubitemid 44374908)
96. Krugel U, Schraft T, Kittner H, Kiess W, Illes P. Basal and feeding-evoked dopamine release in the rat nucleus accumbens is depressed by leptin. Eur J Pharmacol. 2003;482(1-3):185-7. (Pubitemid 37505943)
97. Leshan RL, Opland DM, Louis GW, Leinninger GM, Patterson CM, Rhodes CJ, et al. Ventral tegmental area leptin receptor neurons specifically project to and regulate cocaine- and amphetamine-regulated transcript neurons of the extended central amygdala. J Neurosci. 2010;30(16):5713-23.
98. Louis GW, Leinninger GM, Rhodes CJ, Myers MG. Direct innervation and modulation of orexin neurons by lateral hypotha-lamic Lep Rb neurons. JNeurosci. 2010;30(34):11278-87.
99. Adami GF, Campostano A, Cella F, Scopinaro N. Serum leptin concentration in obese patients with binge eating disorder. Int J Obes. 2002;26(8):1125-8.
100. Morton GJ, Gelling RW, Niswender KD, Morrison CD, Kim F, Schwartz MW. Leptin signaling in the hypothalamic arcuate nucleus is a key determinant of glucose tolerance and hepatic insulin action. Obes Res. 2005;13(1):201.
101. Chelikani PK, Haver AC, Reeve JR, Keire DA, Reidelberger RD. Daily, intermittent intravenous infusion ofpeptide YY(3-36) reduces daily food intake and adiposity in rats. Am J Physiol Regul Integr Comp Physiol. 2006;290(2):R298-305.
102. Batterham RL, ffytche DH, Rosenthal JM, Zelaya FO, Barker GJ, Withers DJ, et al. PYY modulation of cortical and hypothalamic brain areas predicts feeding behaviour in humans. Nature. 2007;450(7166):106-9. (Pubitemid 350070563)
103. Cummings DE, Purnell JQ, Frayo RS, Schmidova K, Wisse BE, Weigle DS. A preprandial rise in plasma ghrelin levels suggests a role in meal initiation in humans. Diabetes. 2001;50 (8):1714-9. (Pubitemid 33641587)
104. Cummings DE, Frayo RS, Marmonier C, Aubert R, Chapelot D. Plasma ghrelin levels and hunger scores in humans initiating meals voluntarily without time- and food-related cues. Am J Physiol Endocrinol Metab. 2004;287(2):E297-304. (Pubitemid 38943896)
105. Wren AM, Small CJ, Abbott CR, Dhillo WS, Seal LJ, Cohen MA, et al. Ghrelin causes hyperphagia and obesity in rats. Diabetes. 2001;50(11):2540-7. (Pubitemid 33642772)
106. Wren AM, Small CJ, Ward HL, Murphy KG, Dakin CL, Taheri S, et al. The novel hypothalamic peptide ghrelin stimulates food intake and growth hormone secretion. Endocrinology 2000;141 (11):4325-8.
107. Hewson AK, Dickson SL. Systemic administration of ghrelin induces Fos and Egr-1 proteins in the hypothalamic arcuate nucleus of fasted and fed rats. J Neuroendocrinol. 2000;12 (11):1047-9.
108. Dickson SL, Leng G, Robinson ICAF. Systemic administration of growth hormone-releasing peptide activates hypothalamic arcuate neurons. Neuroscience. 1993;53(2):303-6. (Pubitemid 23101317)
109. Dickson SL, Luckman SM. 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 secretagogue, GH-releasing peptide-6. Endocrinology. 1997;138(2):771-7. (Pubitemid 27110931)
110. Guan XM, Yu H, Palyha OC, Mc Kee KK, Feighner SD, Sirinathsinghji DJS, et al. Distribution of m RNA encoding the growth hormone secretagogue receptor in brain and peripheral tissues. Mol Brain Res. 1997;48(1):23-9. (Pubitemid 27298124)
111. Zigman JM, Jones JE, Lee CE, Saper CB, Elmquist JK. Expression of ghrelin receptor m RNA in the rat and the mouse brain. J Comp Neurol. 2006;494(3):528-48. (Pubitemid 41791000)
112. Jerlhag E, Egecioglu E, Dickson SL, Douhan A, Svensson L, Engel JA. Ghrelin administration into tegmental areas stimulates locomo-tor activity and increases extracellular concentration ofdopamine in the nucleus accumbens. Addict Biol. 2007;12(1):6-16. (Pubitemid 46405994)
113. Jerlhag E, Egecioglu E, Dickson SL, Andersson M, Svensson L, Engel JA. Ghrelin stimulates locomotor activity and accumbal dopamine-overflow via central cholinergic systems in mice: implications for its involvement in brain reward. Addict Biol. 2006;1 1(1):45-54. (Pubitemid 46696736)
114. Abizaid A, Liu ZW, Andrews ZB, Shanabrough M, Borok E, Elsworth JD, et al. Ghrelin modulates the activity and synaptic input organization of midbrain dopamine neurons while promot-ing appetite. J Clin Invest. 2006;116(12):3229-39. (Pubitemid 44878637)
115. Dickson SL, Hrabovszky E, Hansson C, Jerlhag E, Alvarez-Crespo M, Skibicka KP, et al. Blockade of central nicotine acetylcholine receptor signaling attenuate ghrelin-induced food intake in rodents. Neuroscience. 2010.
116. Jerlhag E, Egecioglu E, Dickson SL, Engel JA. Glutamatergic regulation of ghrelin-induced activation of the mesolimbic dopamine system. Addict Biol. 2011;16(1):82-91.
117. Jerlhag E, Egecioglu E, Landgren S, Salome N, Heilig M, Moechard D, et al. Requirement of central ghrelin signaling for alcoholreward. PNAS. 2009;106(27):11318-23.
118. Jerlhag E, Egecioglu E, Dickson SL, Engel JA. Ghrelin receptor antagonism attenuates cocaine- and amphetamine-induced loco-motor stimulation, accumbal dopamine release and conditioned place preference. Psychopharmacology (Berl). 2010;211(4):415-22.
119. Egecioglu E, Jerlhag E, Skibicka S, Salomé N, Haage D, Bohlooly-YM, et al. Ghrelin increases intake of palatable food in rodents. Addict Biol. 2010;15(3):304-11.
120. Perello M, Sakata I, Birnbaum S, Chuang JC, Osborne-Lawrence S, Rovinsky SA, et al. Ghrelin increases the rewarding value of high-fat diet in an orexin-dependent manner. Biol Psychiatry. 2010;67(9):880-6.
121. Naleid AM, Grace MK, Cummings DE, Levine AS. Ghrelin induces feeding in the mesolimbic reward pathway between the ventral tegmental area and the nucleus accumbens. Peptides. 2005;26(11):2274-9. (Pubitemid 41584095)
122. Skibicka KP, Hansson C, Egecioglu E, Dickson SL. Role of ghrelin in food reward: impact of ghrelin on sucrose self-administration and mesolimbic dopamine and acetylcholine receptor gene expression. Addict Biol. 2010. doi:10.1111/j.1369-1600.2010.00294.x.
123. Malik S, Mc Glone F, Bedrossian D, Dagher A. Ghrelin modulates brain activity in areas that control appetitive behavior Cell Metab. 2008;7(5):400-9. (Pubitemid 351598026)
124. Tschöp M, Weyer C, Tataranni PA, Devanarayan V, Ravussin E, Heiman ML. Circulating Ghrelin levels are decreased in human obesity. Diabetes. 2001;50(4):707-9. (Pubitemid 32242624)
125. English PJ, Ghatei MA, Malik IA, Bloom SR, Wilding JPH. Food fails to suppress ghrelin levels in obese humans. J Clin Endocrinol Metab. 2002;87(6):2984-7. (Pubitemid 34655374)
126. Couce ME, Cottam D, Esplen J, Teijeiro R, Schauer P, Burguera B. Potential role of hypothalamic ghrelin in the pathogenesis of human obesity. J Endocrinol Invest. 2006;29 (7):599-605. (Pubitemid 44446269)
127. Holst B, Cygankiewicz A, Jensen TH, Ankersen M, Schwartz TW. High constitutive signaling of the ghrelin receptor- Identification of a potent inverse agonist. Mol Endocrinol. 2003;17(11):2201-10. (Pubitemid 37352532)