Involvement of melanocortin-4 receptor in anxiety and depression

Peptides

Volumn 26, Issue 10, 2005, Pages 1952-1964

  Chaki, Shigeyuki ^a   Okuyama, Shigeru ^a  

^a TAISHO PHARMACEUTICAL CO LTD   (Japan)

Author keywords

Anxiety; Depression; Melanocortin 4 receptor; Stress

Indexed keywords

1 [2 (4 FLUOROPHENYL) 2 (4 ISOPROPYLPIPERIDIN 1 YL)ETHYL] 4 [4 (2 METHOXYNAPHTHALEN 1 YL)BUTYL]PIPERAZINE; ANTIDEPRESSANT AGENT; ARGININE; CORTICOTROPIN RELEASING FACTOR; MCL 0020; MCL 0129; MELANOCORTIN 4 RECEPTOR; MELANOCORTIN RECEPTOR; PEPTIDE; PIPERAZINE DERIVATIVE; UNCLASSIFIED DRUG; VASOPRESSIN;

ADDICTION; ANXIETY DISORDER; CENTRAL NERVOUS SYSTEM; DEPRESSION; DORSAL RAPHE NUCLEUS; DRUG ABUSE; DRUG SELECTIVITY; FEEDING; HOMEOSTASIS; HUMAN; LOCUS CERULEUS; NONHUMAN; NORADRENERGIC SYSTEM; PAIN; PHYSIOLOGY; PRIORITY JOURNAL; PROTEIN EXPRESSION; REVIEW; SEROTONINERGIC SYSTEM; SEXUAL BEHAVIOR; SEXUAL FUNCTION; STRESS; SURFACE PROPERTY; TRANQUILIZING ACTIVITY;

ANIMALS; ANXIETY DISORDERS; DEPRESSIVE DISORDER; HUMANS; ORGAN SPECIFICITY; RECEPTOR, MELANOCORTIN, TYPE 4;

EID: 25144449526     PISSN: 01969781     EISSN: None     Source Type: Journal    
DOI: 10.1016/j.peptides.2004.11.029     Document Type: Review

References (114)

1. E.D. Abercrombie, and B.L. Jacobs Single-unit response of noradrenergic neurons in the locus coeruleus of freely moving cats. I. Acutely presented stressful and nonstressful stimuli J Neurosci 7 1987 2837 2843
2. Adan RAH, A.W. Szklarczyk, J. Oosterom, J.H. Brakkee, Nijenhuis WAJ, and Schaaper WMM Characterization of melanocortin receptor ligands on cloned brain melanocortin receptors and on grooming behavior in the rat Eur J Pharmacol 378 1999 249 258
3. G. Aguilera, and C. Rabadan-Diehl Vasopressinergic regulation of the hypothalamic-pituitary-adrenal axis: implications for stress adaptation Regul Pept 96 2000 23 29
4. J.D. Alvaro, J.B. Tatro, J.M. Quillan, M. Fogliano, M. Eisenhard, and M.R. Lerner Morphine down-regulates melanocortin-4 receptor expression in brain regions that mediate opiate addiction Mol Pharmacol 50 1996 583 591
5. J.D. Alvaro, J.R. Taylor, and R.S. Duman Molecular and behavioral interactions between central melanocortins and cocaine J Pharmacol Exp Ther 304 2003 391 399
6. A. Anand, and D.S. Charney Norepinephrine dysfunction in depression J Clin Psychiatry 61 2000 16 24
7. A. Argiolas, M.R. Melis, S. Murgia, and H.B. Schioth ACTH- and α-MSH-induced grooming, stretching, yawning and penile erection in male rats: site of action in the brain and role of melanocortin receptors Brain Res Bull 51 2000 425 434
8. D. Bagnol, X.Y. Lu, C.B. Kaelin, H.E. Day, M. Ollmann, and I. Gantz Anatomy of an endogenous antagonist: relationship between agouti-related protein and proopiomelanocortin in brain J Neurosci 19 1999 R26
9. V. Baubet, M. Fevre-Montange, N. Gay, G. Debilly, P. Bobillier, and R. Cespuglio Effects of an acute immobilization stress upon proopiomelanocortin (POMC) mRNA levels in the mediobasal hypothalamus: a quantitative in situ hybridization study Brain Res Mol Brain Res 26 1994 163 168
10. M. Beltramo, M. Campanella, G. Tarozzo, S. Fredduzzi, L. Corradini, and A. Forlani Gene expression profiling of melanocortin system in neuropathic rats supports a role in nociception Mol Brain Res 118 2003 111 118
11. S. Benjannet, N. Rondeau, R. Day, M. Chetien, and N.G. Seidah PC1 and PC2 are proprotein convertases capable of cleaving proopiomelanocortin at distinct pairs of basic residues Proc Natl Acad Sci USA 88 1991 3564 3568
12. S.C. Benoit, M.W. Schwartz, J.L. Lachey, M.M. Hagan, P.A. Rushing, and K.A. Blake A novel selective melanocortin-4 receptor agonist reduces food intake in rats and mice without producing aversive consequences J Neurosci 20 2000 3442 3448
13. A. Bertolini, R. Poggioli, and F. Ferrari ACTH-induced hyperalgesia in rats Experientia 35 1979 1216 1217
14. C. Blasquez, S. Jegou, D.T. Bunel, C. Delbende, P. Braquet, and H. Vaudry Central-type benzodiazepines inhibit release of α-melanocyte-stimulating hormone from the rat hypothalamus Neuroscience 42 1991 509 516
15. F.E. Bloom, E.L. Battenberg, J. Rivier, and W. Vale Corticotropin releasing factor (CRF): immunoreactive neurons and fibers in rat hypothalamus Regul Pept 4 1982 43 48
16. L.S. Brady Stress, antidepressant drugs, and the locus coeruleus Brain Res Bull 35 1994 545 556
17. C. Broberger, J. Johansen, C. Johansson, M. Schalling, and T. Hokfelt The neuropeptide Y/agouti gene-related protein (AGRP) brain circuitry in normal, anorectic and monosodium glutamate-treated mice Proc Natl Acad Sci USA 95 1998 15043 15048
18. D.M. Bronstein, R. Przewlocki, and H. Akil Effects of morphine treatment on pro-opiomelanocortin systems in rat brain Brain Res 519 1990 102 111
19. K.S. Brown, R.M. Gentry, and N.E. Rowland Central injection in rats of α-melanocyte-stimulating hormone analog: effects of food intake and brain Fos Regul Pept 78 1998 89 94
20. S. Cabeza de Vaca, G.Y. Kim, and K.D. Carr The melanocortin receptor agonist MT II augments the rewarding effect of amphetamine in ad-libitum-fed and food-restricted rats Psychopharmacology 161 2002 77 85
21. A. Catania, S. Gatti, G. Colombo, and J.M. Lipton Targeting melanocortin receptors as a novel strategy to control inflammation Pharmacol Rev 56 2004 1 29
22. D. Cepoi, T. Phillips, M. Cismowski, V.S. Goodfellow, N. Ling, and R.D. Cone Assessment of a small molecule melanocortin-4 receptor-specific agonist on energy homeostasis Brain Res 1000 2004 64 71
23. G. Ceriani, A. Macalus, A. Catania, and J.M. Lipton Central neurogenic anti-inflammatory action of α-MSH Neuroendocrinology 59 1994 138 143
24. S. Chaki, S. Hirota, T. Funakoshi, Y. Suzuki, S. Suetake, and T. Okubo Anxiolytic-like and antidepressant-like activities of MCL0129 (1-[(S)-2-(4-fluorophenyl)-2-(4-isopropylpiperadin-1-yl)ethyl] -4-[4-(2-methoxynaphthalen-1-yl)butyl]piperazine), a novel and potent nonpeptide antagonist of the melanocortin-4 receptor J Pharmacol Exp Ther 304 2003 818 826
25. S. Chaki, S. Ogawa, Y. Toda, T. Funakoshi, and S. Okuyama Involvement of the melanocortin MC4 receptor in stress-related behavior in rodents Eur J Pharmacol 474 2003 95 101
26. J.S. Chan, H. Gaspar, H. Iguchi, N.G. Seidah, N. Ling, and M. Chretien Synergistical effects of ovine corticotropin-releasing factor (CRF) and arginine vassopressin (AVP) on the release of pro-opiomelanocortin (POMC) related peptides by pituitary adenoma of a patient with Nelson's syndrome in vitro Clin Invest Med 7 1984 205 208
27. Chang ACY, M. Cochet, and S.N. Cohen Structural organization of human genomic DNA encoding the pro-opiomelanocortin peptide Proc Natl Acad Sci USA 77 1980 4890 4894
28. V. Chhajlani, R. Muceniece, and J.E. Wikberg Molecular cloning of a novel human melanocortin receptor Biochem Biophys Res Commun 195 1993 866 873
29. V. Chhajlani, and J.E.S. Wikberg Molecular cloning and expression of the human melanocyte stimulating hormone receptor cDNA FEBS Lett 309 1992 417 420
30. A. Clarke, and S.E. File Social and exploratory behavior in the rat after septal administration of ORG 2766 and ACTH4-10 Psychoneuroendocrinology 8 1983 343 350
31. A. Contra, and F. Bortolotti Presence of strong glucocorticoid receptor immunoreactivity within hypothalamic and hypophyseal cells containing pro-opiomelanocortic peptides Brain Res 577 1992 127 133
32. M.G. Corda, M. Orlandi, and W. Fratta Proconflict effect of ACTH 1-24: interaction with benzodiazepines Pharmacol Biochem Behav 36 1990 631 634
33. J.F. Cryan, A. Markou, and I. Lucki Assessing antidepressant activity in rodents: recent developments and future needs Trends Pharmacol Sci 23 2002 238 245
34. M. Davis, and C. Shi The extended amygdala: are the central nucleus of the amygdala and the bed nucleus of the stria terminals differently involved in fear versus anxiety? Ann N Y Acad Sci 877 1999 281 291
35. S.R. De Barioglio, N. Lezcano, and M.E. Celis Alpha MSH-induced excessive grooming behavior involves a GABAergic mechanism Peptides 12 1991 203 205
36. D. De Wied, and J. Jolles Neuropeptides derived from pro-opiocortin: behavioral, physiological, and neurochemical effects Physiol Rev 62 1982 976 1059
37. W.S. Dhillo, C.J. Small, L.J. Seal, M.S. Kim, S.A. Stanley, and K.G. Murphy The hypothalamic melanocortin system stimulates the hypothalamo- pituitary-adrenal axis in vitro and in vivo in male rats Neuroendocrinology 75 2002 209 216
38. D.M. Dinulescu, and R.D. Cone Agouti and agouti-related protein: analogies and contrasts J Biol Chem 275 2000 6695 6698
39. J. Drouin, and H.M. Goodman Most of the coding region of rat ACTH/b-LPH precursor gene lacks intervening sequences Nat (Lond) 288 1980 610 613
40. A.J. Dunn, and C.W. Berridge Physiological and behavioral responses to corticotropin-releasing factor administration: is CRF a mediator of anxiety or stress responses? Brain Res Rev 15 1990 71 100
41. A.J. Dunn, E.J. Green, and R.L. Isaacson Intracerebral adrenocorticotropic hormone mediates novelty-induced grooming in the rat Science 203 1979 281 283
42. W. Fan, B.A. Boston, R.A. Kesterson, V.J. Hruby, and R.D. Cone Role of melanocortinergic neurons in feeding and the agouti obesity syndrome Nature 385 1997 165 168
43. S.E. File, and A. Clarke Intraventricular ACTH reduces social interaction in male rats Pharmacol Biochem Behav 12 1980 711 715
44. C.S. Gal, J. Wagnon, J. Simiand, G. Griebel, C. Lacour, and G. Guillon Characterization of (2S,4R)-1-[5-chloro-1-[(2,4-dimethoxyphenyl)sulfonyl]-3-(2- methoxy-phenyl)-2-oxo-2,3-dihydro-1H-indol-3-yl]-4-hydroxy-N, N-dimethyl-2-pyrrolidine carboxamide (SSR149415), a selective and orally active vasopressin V1b receptor antagonist J Pharmacol Exp Ther 300 2002 1122 1130
45. I. Gantz, Y. Konda, T. Tashiro, Y. Shimoto, H. Miwa, and G. Munzert Molecular cloning of a novel melanocortin receptor J Biol Chem 268 1993 8246 8250
46. I. Gantz, H. Miwa, Y. Konda, Y. Shimoto, T. Tashiro, and S.J. Watson Molecular cloning, expression, and gene localization of a fourth melanocortin receptor J Biol Chem 268 1993 15174 15179
47. E. Garcia de Yebenes, and G. Pelletier Opioid regulation of proopiomelanocortin (POMC) gene expression in the rat brain as studied by in situ hybridization Neuropeptides 25 1993 91 94
48. C.E. Gee, Chen CLC, J.L. Roberts, R. Thompson, and S.J. Watson Identification of proopiomelanocortin neurons in the rat hypothalamus by in situ cDNA-mRNA hybridization Nature 306 1983 374 375
49. M.I. Gonzalez, S. Vaziri, and C.A. Wilson Behavioral effects of alpha-MSH and MCH after central administration in the female rat Peptides 17 1996 171 177
50. G. Griebel, J. Simiand, C.S. Gal, J. Wagnon, M. Pascal, and B. Scatton Anxiolytic- and antidepressant-like effects of the non-peptide vasopressin V1b receptor antagonist, SSR149415, suggest an innovative approach for the treatment of stress-related disorders Proc Natl Acad Sci USA 99 2002 6370 6375
51. M.M. Hagan, C.S. Benoit, A.P. Rushing, M.L. Pritchard, C.S. Woods, and J.R. Seeley Immediate and prolonged patterns of agouti-related peptide-(83-132)-induced c-Fos activation in hypothalamic and extrahypothalamic sites Endocrinology 142 2001 1050 1056
52. C. Haskell-Luevano, and E.K. Monck Agouti-related protein functions as an inverse agonist at a constitutively active brain melanocortin-4 receptor Regul Pept 99 2001 1 7
53. R. Hsu, J.R. Alvaro, J.R. Taylor, C. Haile, V. Pjhantharangsy, and G. Han Melanocortin-4 receptor (MC4-R) mediates the rewarding effects of cocaine Soc Neurosci 440 2001 6
54. D. Huszar, C.A. Lynch, V. Fairchild-Huntress, J.H. Dunmore, Q. Fang, and L.R. Berkemeier Targeted disruption of the melanocortin-4 receptor results in obesity in mice Cell 88 1997 131 141
55. B.H. Hwang, and J.M. Guntz Downregulation of corticotropin-releasing factor mRNA, but not vasopressin mRNA, in the paraventricular hypothalamic nucleus of rats following nutritional stress Brain Res Bull 43 1997 509 514
56. S. Jegou, M.E. Yacoubi, L. Mounien, C. Ledent, M. Parmentier, and J.M. Costentin Adenosine 2A receptor gene disruption provokes marked changes in melanocortin content and pro-opiomelanocortin gene expression J Neuroendocrinol 15 2003 1171 1177
57. L. Jonsson, J.O. Skarphedinsson, G.V. Skuladottir, P.T. Atlason, V.H. Eiriksdottir, and L. Franzson Melanocortin receptor agonist transiently increases oxygen consumption in rats Neuroreport 12 2001 3703 3708
58. A. Kask, F. Mutulis, R. Muceniece, R. Pahkla, I. Mutule, and J.E. Wikberg Discovery of a novel superpotent and selective melanocortin-4 receptor antagonist (HS024): evaluation in vitro and in vivo Endocrinology 139 1998 5006 5014
59. A. Kask, L. Rago, J.E. Wikberg, and H.B. Schioth Evidence for involvement of the melanocortin MC4 receptor in the effects of leptin on food intake and body weight Eur J Pharmacol 360 1998 15 19
60. N. Kawashima, S. Chaki, and S. Okuyama Electrophysiological effects of melanocortin receptor ligands on neuronal activities of monoaminergic neurons in rats Neurosci Lett 353 2003 119 122
61. T. Kishi, C.J. Aschkenasi, C.E. Lee, K.G. Mountjoy, C.B. Saper, and J.K. Elmquist Expression of melanocortin 4 receptor mRNA in the central nervous system of the rat J Comp Neurol 457 2003 213 235
62. J.Z. Kiss, M.D. Cassell, and M. Palkovits Analysis of the ACTH/beta-end/alpha-MSH-immunoreactive afferent input to the hypothalamic paraventricular nucleus of rat Brain Res 324 1984 91 99
63. P.J. Larsen, and S.E. Mau Effect of acute stress on the expression of hypothalamic messenger ribonucleic acids encoding the endogenous opioid precursors preproenkephalin A and proopiomelanocortin Peptides 15 1994 783 790
64. A.D. Levy, and L.D. Van de Kar Endocrine and receptor pharmacology of serotonergic anxiolytics, antipsychotics and antidepressants Life Sci 51 1992 83 94
65. Z. Liposits, L. Sievers, and W.K. Paull Neuropeptide-Y and ACTH-immunoreactive innervation of corticotropin releasing factor (CRF)-synthesizing neurons in the hypothalamus of the rat. An immunocytochemical analysis at the light and electron microscopic levels Histochemistry 88 1988 227 234
66. H. Liu, T. Kishi, A.G. Roseberry, X. Cai, C.E. Lee, and J.M. Montez Transgenic mice expressing green fluorescent protein under the control of the melanocortin-4 receptor promoter J Neurosci 23 2003 7143 7154
67. X.Y. Lu, G.S. Barsh, H. Akil, and S.J. Watson Interaction between α-melanocyte-stimulating hormone and corticotropin-releasing hormone in the regulation of feeding and hypothalamo-pituitary-adrenal responses J Neurosci 23 2003 7863 7872
68. D. Lu, D. Willard, I.R. Patel, S. Kadwell, L. Overton, and T. Kost Agouti protein is an antagonist of the melanocyte-stimulating-hormone receptors Nature 371 1994 799 802
69. S.F. Maier, and ME.P. Seligman Learned helplessness: theory and evidence J Exp Psychol Gen 105 1976 3 46
70. S. Makino, K. Asaba, M. Nishiyama, and K. Hashimoto Decreased type 2 corticotropin-releasing hormone receptor mRNA expression in the ventromedial hypothalamus during repeated immobilization stress Neuroendocrinology 70 1999 160 167
71. J.S. Meyer, D.J. Micco, B.S. Stephenson, L.C. Krey, and B.S. McEwen Subcutaneous implantation method for chronic glucocorticoid replacement therapy Physiol Behav 22 1979 867 870
72. K.G. Mountjoy, M.T. Mortrud, M.J. Low, R.B. Simerly, and R.D. Cone Localization of the melanocortin-4 receptor (MC4-R) in neuroendocrine and autonomic control circuits in the brain Mol Endocrinol 8 1994 1298 1308
73. K.G. Mountjoy, L.S. Robbins, M.T. Mortrud, and R.D. Cone The cloning of a family of genes that encode the melanocortin receptors Science 257 1992 1248 1251
74. M.T. Murphy, D.B. Richards, and J.M. Lipton Antipyretic potency of centrally administered alpha-melanocyte stimulating hormone Science 221 1983 192 193
75. S. Nakanishi, A. Inoue, T. Kita, M. Nakamura, A.C.Y. Chang, and S.N. Cohen Nucleotide sequence of cloned cDNA for bovine corticotropin-β- lipotropin precursor Nat (Lond) 278 1979 423 427
76. S. Okuyama, S. Chaki, N. Kawashima, Y. Suzuki, S. Ogawa, and A. Nakazato Receptor binding, behavioral, and electrophysiological profiles of nonpeptide corticotropin-releasing factor subtype 1 receptor antagonists CRA1000 and CRA1001 J Pharmacol Exp Ther 289 1999 926 935
77. M.M. Ollmann, B.D. Wilson, Y.K. Yang, J.A. Kerns, Y. Chen, and I. Gantz Antagonism of central melanocortin receptors in vitro and in vivo by agouti-related protein Science 278 1997 135 138
78. M.J. Owen, and C.B. Nemeroff Physiology and pharmacology of corticotropin releasing factor Pharmacol Rev 43 1991 425 473
79. H. Ozawa, J. Lion, C.X. Xie, M. Nishi, H. Steinbusch, and M. Kawata Down-regulation of ACTH and glucocorticoid receptor immunoreactivity in hypothalamic arcuate neurons after adrenalectomy in the rat Neuroreport 10 1999 1571 1575
80. J. Panksepp, and L. Normansell Effects of ACTH (1-24) and ACTH/MSH (4-10) on isolation-induced distress vocalization in domestic chicks Peptides 11 1990 915 919
81. E.M. Pich, M. Lorang, M. Yeganeh, F. Rodriguez de Fonseca, J. Raber, and G.F. Koob Increase of extracellular corticotropin-releasing factor-like immunoreactivity levels in the amygdala of awake rats during restraint stress and ethanol withdrawal as measured by microdialysis J Neurosci 15 1995 5439 5447
82. T.A. Pigott, and S.M. Seay A review of the efficacy of selective serotonin reuptake inhibitors in obsessive-compulsive disorder J Clin Psychiatry 60 1999 101 106
83. L.C. Plantinga, J. Verhaagen, P.M. Edwards, L.H. Schrama, J.P. Burbach, and W.H. Gispen Expression of the pro-opiomelanocortin gene in dorsal root ganglia, spinal cord and sciatic nerve after sciatic nerve crush in the rat Brain Res Mol Brain Res 16 1992 135 142
84. R. Poggioli, A.V. Vergoni, and A. Bertolini ACTH-(1-24) and alpha-MSH antagonize feeding behavior stimulated by kappa opiate agonists Peptides 7 1986 843 848
85. L.E. Pritchard, A.V. Turnbull, and A. White Pro-opiomelanocortin processing in the hypothalamus: impact on melanocortin signaling and obesity J Endocrinol 172 2002 411 421
86. T.L. Rao, D.M. Kokare, S. Sarkar, R.T. Khisti, C.T. Chopde, and N. Subhedar GABAergic agents prevent alpha-melanocyte stimulating hormone induced anxiety and anorexia in rats Pharmacol Biochem Behav 76 2003 417 423
87. C. Rivier, and P.M. Plotsky Mediation by corticotropin-releasing factor (CRF) of adrenohypophysial hormone secretion Ann Rev Physiol 48 1986 475 494
88. L. Roselli-Rehfuss, K.G. Mountjoy, L.S. Robbins, M.T. Mortrud, M.J. Low, and J.B. Tatro Identification of a receptor for γ melanocortin and other proopiomelanocortin peptides in the hypothalamus and limbic system Proc Natl Acad Sci USA 90 1993 8856 8860
89. C.A. Sandman, and A.J. Kastin Intraventricular administration of MSH induces hyperalgesia in rats Peptides 2 1981 231 233
90. S. Sarker, G. Legradi, and R.M. Lechan Intracerebroventricular administration of α-melanocyte stimulating hormone increases phosphorylation of CREB in TRH- and CRH-producing neurons of the hypothalamic paraventricular nucleus Brain Res 945 2002 50 59
91. L.V. Scott, and T.G. Dinan Vasopressin and the regulation of hypothalamic-pituitary-adrenal axis function: implications for the pathophysiology of depression Life Sci 62 1998 1985 1998
92. P. Seechurn, S.A. Burchill, and A.J. Thody Effect of α-melanocyte- stimulating hormone on tyrosinase activity in hair follicular and epidermal melanocytes of the mouse J Endocrinol 119 1988 517 522
93. K. Starowicz, W. Bilecki, A. Sieja, B. Przewlocka, and R. Przewlocki Melanocortin 4 receptor is expressed in the dorsal root ganglions and down-regulated in neuropathic rats Neurosci Lett 358 2004 79 82
94. K. Starowicz, A. Sieja, W. Bilecki, I. Obara, and B. Przewlocka The effects of morphine on MC4 and CRF receptor mRNAs in the rat amygdala and attenuation of tolerance after their blockade Brain Res 990 2003 113 119
95. K. Takamori, S. Yoshida, and S. Okuyama Repeated treatment with imipramine, fluvoxamine and tranylcypromine decreases the number of escape failures by activating dopaminergic systems in a rat learned helplessness test Life Sci 69 2001 1919 1926
96. A.J. Thody α-MSH and the regulation of melanocyte function Ann N Y Acad Sci 885 1999 217 229
97. K. Tsou, H. Khachaturian, H. Akil, and S.J. Watson Immunocytochemical localization of pro-opiomelanocortin-derived peptides in the adult rat spinal cord Brain Res 378 1986 28 35
98. S. Tsujii, and G.A. Bray Acetylation alters the feeding response to MSH and beta-endorphin Brain Res Bull 23 1989 165 169
99. W.W. Vale, J. Spiess, C. Rivier, and J. Rivier Characterization of a 41-amino acid residue ovine hypothalamic peptide that stimulates secretion of corticotropin and beta-endorphin Science 213 1981 1394 1397
100. M. Van der Kraan, J.B. Tatro, M.L. Entwistle, J.H. Brakkee, J.P. Burbach, and R.A. Adan Expression of melanocortin receptors and proopiomelanocortin in the rat spinal cord in relation to neurotrophic effects of melanocortins Brain Res Mol Brain Res 63 1999 276 286
101. L.H.T. Van der Ploeg, W.J. Martin, A.D. Howard, R.P. Nargund, C.P. Austin, and X. Guan A role for the melanocortin 4 receptor in sexual function Proc Natl Acad Sci USA 99 2002 11381 11386
102. M. Vecsernyes, E. Biro, J. Gardi, J. Julesz, and G. Telegdy Involvement of endogenous corticotropin-releasing factor in mediation of neuroendocrine and behavioral effects to alpha-melanocyte-stimulating hormone Endocr Res 26 2000 347 356
103. A.V. Vergoni, A. Bertolini, J.E. Wikberg, and H.B. Schioth Selective melanocortin MC4 receptor blockage reduces immobilization stress-induced anorexia in rats Eur J Pharmacol 369 1999 11 15
104. J. Voisey, and A. van Daal Agouti: from mouse to man, from skin to fat Pigment Cell Res 15 2002 10 18
105. J.C. Von Frijtag, G. Croiset, W.H. Gispen, Adan RAH, and V.M. Wiegant The role of central melanocortin receptors in the activation of the hypothalamus-pituitary-adrenal-axis and the induction of excessive grooming Br J Pharmacol 123 1998 1503 1508
106. N. Vrang, P.J. Larsen, M. Tang-Christensen, L.K. Larsen, and P. Kristensen Hypothalamic cocaine-amphetamine regulated transcript (CART) is regulated by glucocorticoids Brain Res 965 2003 45 50
107. D.H. Vrinten, W.H. Gispen, G.J. Groen, and RA.H. Adan Antagonism of the melanocortin system reduces cold and mechanical allodynia in mononeuropathic rats J Neurosci 20 2000 8131 8137
108. D.H. Vrinten, Adan RAH, G.J. Groen, and W.H. Gispen Chronic blockade of melanocortin receptors alleviates allodynia in rats with neuropathic pain Anesth Analg 93 2001 1572 1577
109. H. Wessells, K. Fuciarelli, J. Hansen, M.E. Hadley, V.J. Hruby, and R. Dorr Synthetic melanotropic peptide initiates erections in men with psychogenic erectile dysfunction J Urol 160 1998 389 393
110. H. Wessells, D. Gralnek, D. Dorr, V.J. Hruby, M.E. Hadley, and N. Levine Effect of an alpha-melanocyte stimulating hormone analog on penile erection and sexual desire in men with organic erectile dysfunction Urologia 56 2000 641 646
111. JE.S. Wikberg Melanocortin receptors: perspectives for novel drugs Eur J Pharmacol 375 1999 295 310
112. D.H. Willard, W. Bodnar, C. Harris, L. Kiefer, J.S. Nichols, and S. Blanchard Agouti structure and function: characterization of a potent α-melanocyte stimulating hormone receptor antagonist Biochemistry 34 1995 12341 12346
113. Y. Yamano, M. Yoshioka, Y. Toda, Y. Oshida, S. Chaki, and K. Hamamoto Regulation of CRF, POMC and MC4R gene expression by electrical foot shock stress in rat amygdala and hypothalamus J Vet Med Sci 66 2004 1323 1327
114. Z. Zheng, L. Leger, R. Cespuglio, and M. Jouvet Distribution of the pro-opiomelanocortin-immunoreactive axons in relation to the serotonergic neurons in the dorsal raphe nucleus of the rat Neurosci Lett 130 1991 17 21