Melanocortin MC4 receptor-mediated feeding and grooming in rodents

European Journal of Pharmacology

Volumn 719, Issue 1-3, 2013, Pages 192-201

  Mul, Joram D ^a   Spruijt, Berry M ^b   Brakkee, Jan H ^c   Adan, Roger A H ^c  

^a UNIVERSITY OF CINCINNATI   (United States)

^b UTRECHT UNIVERSITY   (Netherlands)

^c UNIVERSITY MEDICAL CENTER UTRECHT   (Netherlands)

Author keywords

ACTH; Body weight; Food intake; Grooming; MC4R; Melanocortin MC4 receptor; MSH

Indexed keywords

CORTICOTROPIN; INTERMEDIN; LIGAND; MELANOCORTIN; MELANOCORTIN 1 RECEPTOR; MELANOCORTIN 4 RECEPTOR; MELANOCORTIN LIGAND; OPIATE; UNCLASSIFIED DRUG;

ANIMAL BEHAVIOR; BRAIN; CONTROLLED STUDY; FEEDING BEHAVIOR; FOOD INTAKE; GROOMING; HUMAN; MELANOCYTE; MOLECULAR CLONING; NONHUMAN; PIGMENTATION; PRIORITY JOURNAL; PROTEIN EXPRESSION; PROTEIN FUNCTION; REVIEW; RODENT; SATIETY; STRUCTURE ACTIVITY RELATION; XENOPUS LAEVIS;

ACTH; BODY WEIGHT; FOOD INTAKE; GROOMING; MC4R; MELANOCORTIN MC(4) RECEPTOR; MSH;

ANALGESICS, OPIOID; ANIMALS; FEEDING BEHAVIOR; GROOMING; MELANOCORTINS; RECEPTOR, MELANOCORTIN, TYPE 4;

EID: 84888291002     PISSN: 00142999     EISSN: 18790712     Source Type: Journal    
DOI: 10.1016/j.ejphar.2013.04.060     Document Type: Review

References (150)

1. R.A. Adan, R.D. Cone, J.P. Burbach, and W.H. Gispen Differential effects of melanocortin peptides on neural melanocortin receptors Mol. Pharmacol. 46 1994 1182 1190
2. R.A. Adan, and W.H. Gispen Brain melanocortin r (Pubitemid 28026902)
3. R.A. Adan, and W.H. Gispen Melanocortins and the brain: from effects via receptors to drug targets Eur. J. Pharmacol. 405 2000 13 24
4. R.A. Adan, J. Oosterom, G. Ludvigsdottir, J.H. Brakkee, J.P. Burbach, and W.H. Gispen Identification of antagonists for melanocortin MC3, MC4 and MC5 receptors Eur. J. Pharmacol. 269 1994 331 337 (Pubitemid 24348049)
5. R.A. Adan, A.W. Szklarczyk, J. Oosterom, J.H. Brakkee, W.A. Nijenhuis, W.M. Schaaper, R.H. Meloen, and W.H. Gispen Characterization of melanocortin receptor ligands on cloned brain melanocortin receptors and on grooming behavior in the rat Eur. J. Pharmacol. 378 1999 249 258 (Pubitemid 29392925)
6. R.A. Adan, B. Tiesjema, J.J. Hillebrand, S.E. la Fleur, M.J. Kas, and M. de Krom The MC4 receptor and control of appetite Br. J. Pharmacol. 149 2006 815 827 (Pubitemid 44851706)
7. G.M. Alexander, S.C. Rogan, A.I. Abbas, B.N. Armbruster, Y. Pei, J.A. Allen, R.J. Nonneman, J. Hartmann, S.S. Moy, M.A. Nicolelis, J.O. McNamara, and B.L. Roth Remote control of neuronal activity in transgenic mice expressing evolved G protein-coupled receptors Neuron 63 2009 27 39
8. V.J. Aloyo, B. Spruijt, H. Zwiers, and W.H. Gispen Peptide-induced excessive grooming in the rat: the role of opiate receptors Peptides 4 1983 833 836 (Pubitemid 14184936)
9. J.D. Alvaro, J.B. Tatro, J.M. Quillan, M. Fogliano, M. Eisenhard, M.R. Lerner, E.J. Nestler, and R.S. Duman Morphine down-regulates melanocortin-4 receptor expression in brain regions that mediate opiate addiction Mol. Pharmacol. 50 1996 583 591 (Pubitemid 26303589)
10. S. Amir Effects of ACTH on pain responsiveness in mice: interaction with morphine Neuropharmacology 20 1981 959 962 (Pubitemid 11015671)
11. D. Atalayer, K.L. Robertson, C. Haskell-Luevano, A. Andreasen, and N.E. Rowland Food demand and meal size in mice with single or combined disruption of melanocortin type 3 and 4 receptors Am. J. Physiol. Regul. Integr. Comp. Physiol. 298 2010 R1667 1674
12. D. Atasoy, Y. Aponte, H.H. Su, and S.M. Sternson A FLEX switch targets Channelrhodopsin-2 to multiple cell types for imaging and long-range circuit mapping J. Neurosci. 28 2008 7025 7030
13. I.H. Ayan, and A. Randupp Behavioural and pharmacological studies on morphine-induced excitation of rats. Possible relation to brain catecholamines Psychopharmacologia 29 1973 317 328
14. N. Balthasar, L.T. Dalgaard, C.E. Lee, J. Yu, H. Funahashi, T. Williams, M. Ferreira, V. Tang, R.A. McGovern, C.D. Kenny, L.M. Christiansen, E. Edelstein, B. Choi, O. Boss, C. Aschkenasi, C.Y. Zhang, K. Mountjoy, T. Kishi, J.K. Elmquist, and B.B. Lowell Divergence of melanocortin pathways in the control of food intake and energy expenditure Cell 123 2005 493 505 (Pubitemid 41546678)
15. S.C. Benoit, D.J. Clegg, J.G. Barrera, R.J. Seeley, and S.C. Woods Learned meal initiation attenuates the anorexic effects of the melanocortin agonist MTII Diabetes 52 2003 2684 2688 (Pubitemid 37339697)
16. A. Bertolini, R. Tacchi, and A.V. Vergoni Brain effects of melanocortins Pharmacol. Res. 59 2009 13 47
17. J.N. Betley, and S.M. Sternson Adeno-associated viral vectors for mapping, monitoring, and manipulating neural circuits Hum. Gene. Ther. 22 2011 669 677
18. H. Biebermann, H. Krude, A. Elsner, V. Chubanov, T. Gudermann, and A. Gruters Autosomal-dominant mode of inheritance of a melanocortin-4 receptor mutation in a patient with severe early-onset obesity is due to a dominant-negative effect caused by receptor dimerization Diabetes 52 2003 2984 2988 (Pubitemid 37466869)
19. R.C. Bolles Grooming behavior in the rat J. Comp. Physiol. Psychol. 53 1960 306 310
20. A. Breit, T.R. Buch, I. Boekhoff, H.J. Solinski, E. Damm, and T. Gudermann Alternative G protein coupling and biased agonism: new insights into melanocortin-4 receptor signalling Mol. Cell. Endocrinol. 331 2011 232 240
21. K.S. Brown, R.M. Gentry, and N.E. Rowland Central injection in rats of alpha-melanocyte-stimulating hormone analog: effects on food intake and brain Fos Regul. Pept. 78 1998 89 94 (Pubitemid 28561521)
22. A.A. Butler, D.L. Marks, W. Fan, C.M. Kuhn, M. Bartolome, and R.D. Cone Melanocortin-4 receptor is required for acute homeostatic responses to increased dietary fat Nat. Neurosci. 4 2001 605 611 (Pubitemid 32507374)
23. C. Caruso, D. Durand, H.B. Schioth, R. Rey, A. Seilicovich, and M. Lasaga Activation of melanocortin 4 receptors reduces the inflammatory response and prevents apoptosis induced by lipopolysaccharide and interferon-gamma in astrocytes Endocrinology 148 2007 4918 4926 (Pubitemid 47481532)
24. L.F. Chan, T.R. Webb, T.T. Chung, E. Meimaridou, S.N. Cooray, L. Guasti, J.P. Chapple, M. Egertova, M.R. Elphick, M.E. Cheetham, L.A. Metherell, and A.J. Clark MRAP and MRAP2 are bidirectional regulators of the melanocortin receptor family Proc. Natl. Acad. Sci. USA 106 2009 6146 6151
25. C. Chavkin, and A. Goldstein Specific receptor for the opioid peptide dynorphin: structure-activity relationships Proc. Natl. Acad. Sci. USA 78 1981 6543 6547 (Pubitemid 12208179)
26. A.S. Chen, D.J. Marsh, M.E. Trumbauer, E.G. Frazier, X.M. Guan, H. Yu, C.I. Rosenblum, A. Vongs, Y. Feng, L. Cao, J.M. Metzger, A.M. Strack, R.E. Camacho, T.N. Mellin, C.N. Nunes, W. Min, J. Fisher, S. Gopal-Truter, D.E. MacIntyre, H.Y. Chen, and L.H. Van der Ploeg Inactivation of the mouse melanocortin-3 receptor results in increased fat mass and reduced lean body mass Nat. Genet. 26 2000 97 102
27. A.S. Chen, J.M. Metzger, M.E. Trumbauer, X.M. Guan, H. Yu, E.G. Frazier, D.J. Marsh, M.J. Forrest, S. Gopal-Truter, J. Fisher, R.E. Camacho, A.M. Strack, T.N. Mellin, D.E. MacIntyre, H.Y. Chen, and L.H. Van der Ploeg Role of the melanocortin-4 receptor in metabolic rate and food intake in mice Transgenic Res. 9 2000 145 154 (Pubitemid 30610232)
28. H.O. Collier Cellular site of opiate dependence Nature 283 1980 625 629 (Pubitemid 10099988)
29. R.D. Cone Anatomy and regulation of the central melanocortin system Nat. Neurosci. 8 2005 571 578 (Pubitemid 40594196)
30. P.C. Contreras, and A.E. Takemori Antagonism of morphine-induced analgesia, tolerance and dependence by alpha-melanocyte-stimulating hormone J. Pharmacol. Exp. Ther. 229 1984 21 26 (Pubitemid 14082905)
31. M.A. Cowley, N. Pronchuk, W. Fan, D.M. Dinulescu, W.F. Colmers, and R.D. Cone Integration of NPY, AGRP, and melanocortin signals in the hypothalamic paraventricular nucleus: evidence of a cellular basis for the adipostat Neuron 24 1999 155 163 (Pubitemid 29466155)
32. H. Cui, B.L. Mason, C. Lee, A. Nishi, J.K. Elmquist, and M. Lutter Melanocortin 4 receptor signaling in dopamine 1 receptor neurons is required for procedural memory learning Physiol. Behav. 106 2012 201 210
33. M.W. de Backer, S.E. la Fleur, and R.A. Adan Both overexpression of agouti-related peptide or neuropeptide Y in the paraventricular nucleus or lateral hypothalamus induce obesity in a neuropeptide- and nucleus specific manner Eur. J. Pharmacol. 660 2011 148 155
34. M.W. de Backer, S.E. la Fleur, M.A. Brans, A.J. van Rozen, M.C. Luijendijk, M. Merkestein, K.M. Garner, E.M. van der Zwaal, and R.A. Adan Melanocortin receptor-mediated effects on obesity are distributed over specific hypothalamic regions Int. J. Obes. 35 2011 629 641
35. P.N. De Graan, B.M. Spruijt, A.N. Eberle, J. Girard, and W.H. Gispen ACTH1-4 potentiates alpha-MSH-induced melanophore dispersion and excessive grooming Peptides 7 1986 1 4 (Pubitemid 16082854)
36. D. De Wied, and J. Jolles Neuropeptides derived from pro-opiocortin: behavioral, physiological, and neurochemical effects Physiol. Rev. 62 1982 976 1059
37. G.E. Dickerson, and J.W. Gowen Hereditary obesity and efficient food utilization in mice Science 105 1947 496 498
38. A.J. Dunn, S.R. Childers, N.R. Kramarcy, and J.W. Villiger ACTH-induced grooming involves high-affinity opiate receptors Behav. Neural. Biol. 31 1981 105 109 (Pubitemid 11097926)
39. R. Elde, S. Haber, R. Ho, V. Holets, N. Lanerolle de, B. Maley, P. Micevyck, and V. Seybold Interspecies conservation and variation in peptidergic neurons Peptides 1 1980 21 22
40. A. Elsner, P. Tarnow, M. Schaefer, P. Ambrugger, H. Krude, A. Gruters, and H. Biebermann MC4R oligomerizes independently of extracellular cysteine residues Peptides 27 2006 372 379 (Pubitemid 43162908)
41. 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 (Pubitemid 27034218)
42. W. Fan, K.L. Ellacott, I.G. Halatchev, K. Takahashi, P. Yu, and R.D. Cone Cholecystokinin-mediated suppression of feeding involves the brainstem melanocortin system Nat. Neurosci. 7 2004 335 336 (Pubitemid 38429170)
43. C. Fekete, D.L. Marks, S. Sarkar, C.H. Emerson, W.M. Rand, R.D. Cone, and R.M. Lechan Effect of Agouti-related protein in regulation of the hypothalamic-pituitary-thyroid axis in the melanocortin 4 receptor knockout mouse Endocrinology 145 2004 4816 4821 (Pubitemid 39551096)
44. W. Ferrari Behavioural changes in animals after intracisternal injection with adrenocorticotrophic hormone and melanocyte-stimulating hormone Nature 181 1958 925 926
45. L.G. Frigeri, G.L. Wolff, and C. Teguh Differential responses of yellow Avy/A and agouti A/a (BALB/c X VY) F1 hybrid mice to the same diets: glucose tolerance, weight gain, and adipocyte cellularity Int. J. Obes. 12 1988 305 320
46. I. Gantz, H. Miwa, Y. Konda, Y. Shimoto, T. Tashiro, S.J. Watson, J. DelValle, and T. Yamada Molecular cloning, expression, and gene localization of a fourth melanocortin receptor J. Biol. Chem. 268 1993 15174 15179 (Pubitemid 23206671)
47. A.S. Garfield, D.D. Lam, O.J. Marston, M.J. Przydzial, and L.K. Heisler Role of central melanocortin pathways in energy homeostasis Trends Endocrinol. Metab. 20 2009 203 215
48. S.Q. Giraudo, C.J. Billington, and A.S. Levine Feeding effects of hypothalamic injection of melanocortin 4 receptor ligands Brain Res. 809 1998 302 306 (Pubitemid 29005387)
49. W.H. Gispen, and R.L. Isaacson ACTH-induced excessive grooming in the rat Pharmacol. Ther. 1981 209 246 (Pubitemid 11183307)
50. W.H. Gispen, J.M. van Ree, and D. de Wied Lipotropin and the central nervous system Int. Rev. Neurobiol. 1977 20
51. W.H. Gispen, and V.M. Wiegant Opiate antagonists suppress ACTH1-124-induced ex essive grooming in the rat Neurosci. Lett. 1976 2
52. W.H. Gispen, V.M. Wiegant, A.F. Bradbury, E.C. Hulme, D.G. Smyth, C.R. Snell, and D. de Wied Induction of excessive grooming in the rat by fragments of lipotropin Nature 264 1976 794 795
53. W.H. Gispen, V.M. Wiegant, H.M. Greven, and D. de Wied The induction of excessive grooming in the rat by intraventricular application of peptides derived from ACTH: structure-activity studies Life Sci. 17 1975 645 652
54. M. Graham, J.R. Shutter, U. Sarmiento, I. Sarosi, and K.L. Stark Overexpression of Agrt leads to obesity in transgenic mice Nat. Genet. 17 1997 273 274
55. D. Greff, F. Toma, S. Fermandjian, M. Low, and L. Kisfaludy Conformational studies of corticotropin1-32 and constitutive peptides by circular dichroism Biochim. Biophys. Acta 439 1976 219 231
56. H.J. Grill, A.B. Ginsberg, R.J. Seeley, and J.M. Kaplan Brainstem application of melanocortin receptor ligands produces long-lasting effects on feeding and body weight J. Neurosci. 18 1998 10128 10135 (Pubitemid 28543786)
57. L. Guo, H. Munzberg, R.C. Stuart, E.A. Nillni, and C. Bjorbaek N-acetylation of hypothalamic alpha-melanocyte-stimulating hormone and regulation by leptin Proc. Natl. Acad. Sci. USA 101 2004 11797 11802 (Pubitemid 39095374)
58. M.E. Hadley, B. Anderson, C.B. Heward, T.K. Sawyer, and V.J. Hruby Calcium-dependent prolonged effects on melanophores of [4-norleucine, 7-D-phenylalanine]-alpha-melanotropin Science 213 1981 1025 1027
59. R. Hanada, M. Nakazato, S. Matsukura, N. Murakami, H. Yoshimatsu, and T. Sakata Differential regulation of melanin-concentrating hormone and orexin genes in the agouti-related protein/melanocortin-4 receptor system Biochem. Biophys. Res. Commun. 268 2000 88 91 (Pubitemid 30116616)
60. A.M. Haqq, P. Rene, T. Kishi, K. Khong, C.E. Lee, H. Liu, J.M. Friedman, J.K. Elmquist, and R.D. Cone Characterization of a novel binding partner of the melanocortin-4 receptor: attractin-like protein Biochem. J. 376 2003 595 605 (Pubitemid 38058059)
61. M. Harris, C. Aschkenasi, C.F. Elias, A. Chandrankunnel, E.A. Nillni, C. Bjoorbaek, J.K. Elmquist, J.S. Flier, and A.N. Hollenberg Transcriptional regulation of the thyrotropin-releasing hormone gene by leptin and melanocortin signaling J. Clin. Invest. 107 2001 111 120 (Pubitemid 32047420)
62. C.B. Heward, Y.C. Yang, J.F. Ormberg, M.E. Hadley, and V.J. Hruby Effects of chloramine T and iodination on the biological activity of melanotropin Hoppe-Seyler′s. Z. Physiol. Chem. 360 1979 1851 1859 (Pubitemid 10180470)
63. C.B. Heward, Y.C. Yang, T.K. Sawyer, M.D. Bregman, B.B. Fuller, V.J. Hruby, and M.E. Hadley Iodination associated inactivation of beta-melanocyte stimulating hormone Biochem. Biophys. Res. Commun. 88 1979 266 273
64. V.J. Hruby, D. Lu, S.D. Sharma, A.L. Castrucci, R.A. Kesterson, F.A. al-Obeidi, M.E. Hadley, and R.D. Cone Cyclic lactam alpha-melanotropin analogues of Ac-Nle4-cyclo[Asp5, D-Phe7,Lys10] alpha-melanocyte-stimulating hormone-(4-10)-NH2 with bulky aromatic amino acids at position 7 show high antagonist potency and selectivity at specific melanocortin receptors J. Med. Chem. 38 1995 3454 3461
65. V.J. Hruby, T.K. Sawyer, Y.C. Yang, M.D. Bregman, M.E. Hadley, and C.B. Heward Synthesis and structure-function studies of melanocyte stimulating hormone analogues modified in the 2 and 4(7) positions: comparison of activities on frog skin melanophores and melanoma adenylate cyclase J. Med. Chem. 23 1980 1432 1437 (Pubitemid 11164115)
66. D. Huszar, C.A. Lynch, V. Fairchild-Huntress, J.H. Dunmore, Q. Fang, L.R. Berkemeier, W. Gu, R.A. Kesterson, B.A. Boston, R.D. Cone, F.J. Smith, L.A. Campfield, P. Burn, and F. Lee Targeted disruption of the melanocortin-4 receptor results in obesity in mice Cell 88 1997 131 141 (Pubitemid 27180337)
67. H. Imura, Y. Kato, Y. Nakai, K. Nakao, I. Tanaka, H. Jingami, T. Koh, T. Yoshimasa, T. Tsukada, and M. Suda Endogenous opioids and related peptides: from molecular biology to clinical medicine. The Sir Henry Dale lecture for 1985 J. Endocrinol. 107 1985 147 157 (Pubitemid 16196949)
68. Y. Ishii, J.E. Blundell, J.C. Halford, and R.J. Rodgers Effects of systematic variation in presatiation and fasting on the behavioural satiety sequence in male rats Physiol. Behav. 79 2003 227 238 (Pubitemid 36776673)
69. K. Izumi, J. Donaldson, and A.A. Barbeau Yawning and stretsching in rats induced by intraventricularly administered zinc Life Sci. 12 1973 203 221
70. J. Jolles, V.M. Wiegant, and W.H. Gispen Inhibation of behavioural effect of ACTH1-124 and opioids by repeated administration Neurosci. Lett. 9 1978 261 266
71. L. Karlsson-Lindahl, L. Schmidt, D. Haage, C. Hansson, M. Taube, E. Egeciouglu, Y.X. Tan, T. Admyre, J.O. Jansson, I. Vlodavsky, J.P. Li, U. Lindahl, and S.L. Dickson Heparanase affects food intake and regulates energy balance in mice PLoS One 7 2012 e34313
72. M.J. Kas, B. Tiesjema, G. van Dijk, K.M. Garner, G.S. Barsh, O. ter Brake, J. Verhaagen, and R.A. Adan Induction of brain-region-specific forms of obesity by agouti J. Neurosci. 24 2004 10176 10181 (Pubitemid 39492043)
73. A. Kask, and H.B. Schioth Tonic inhibition of food intake during inactive phase is reversed by the injection of the melanocortin receptor antagonist into the paraventricular nucleus of the hypothalamus and central amygdala of the rat Brain Res. 887 2000 460 464 (Pubitemid 32008075)
74. R. Katz Grooming elicited by intracerebroventricular bombesin and eledoisin in the mouse Neuropharmacology 19 1980 143 146 (Pubitemid 10168511)
75. M.S. Kim, M. Rossi, S. Abusnana, D. Sunter, D.G. Morgan, C.J. Small, C.M. Edwards, M.M. Heath, S.A. Stanley, L.J. Seal, J.R. Bhatti, D.M. Smith, M.A. Ghatei, and S.R. Bloom Hypothalamic localization of the feeding effect of agouti-related peptide and alpha-melanocyte-stimulating hormone Diabetes 49 2000 177 182 (Pubitemid 30070994)
76. 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 (Pubitemid 36158283)
77. M.J. Krashes, S. Koda, C. Ye, S.C. Rogan, A.C. Adams, D.S. Cusher, E. Maratos-Flier, B.L. Roth, and B.B. Lowell Rapid, reversible activation of AgRP neurons drives feeding behavior in mice J. Clin. Invest. 121 2011 1424 1428
78. D.T. Krieger Brain peptides: what, where, and why? Science 222 1983 975 985 (Pubitemid 14225229)
79. L.R. Kushner, and D.G. Mook Behavioral correlates of oral and postingestive satiety in the rat Physiol. Behav. 33 1984 713 718 (Pubitemid 15181515)
80. D. Le Roith, J. Shiloach, and J. Roth Is there an earlier phylogenetic precursor that is common to both the nervous and endocrine systems? Peptides 3 1982 211 215 (Pubitemid 12022663)
81. J.M. Lerma-Cabrera, F. Carvajal, L. de la Torre, L. de la Fuente, M. Navarro, T.E. Thiele, and I. Cubero Control of food intake by MC4-R signaling in the lateral hypothalamus, nucleus accumbens shell and ventral tegmental area: interactions with ethanol Behav. Brain. Res. 234 2012 51 60
82. L.W. Loijens, W.G. Schouten, P.R. Wiepkema, and V.M. Wiegant Brain opioid receptor density relates to stereotypies in chronically stressed pigs Stress 3 1999 17 26 (Pubitemid 29462671)
83. D. Lu, D. Willard, I.R. Patel, S. Kadwell, L. Overton, T. Kost, M. Luther, W. Chen, R.P. Woychik, and W.O. Wilkison Agouti protein is an antagonist of the melanocyte-stimulating-hormone receptor Nature 371 1994 799 802 (Pubitemid 24333868)
84. X.Y. Lu, G.S. Barsh, H. Akil, and S.J. Watson Interaction between alpha-melanocyte-stimulating hormone and corticotropin-releasing hormone in the regulation of feeding and hypothalamo-pituitary-adrenal responses J. Neurosci. 23 2003 7863 7872 (Pubitemid 37052354)
85. D.J. Marsh, G. Hollopeter, D. Huszar, R. Laufer, K.A. Yagaloff, S.L. Fisher, P. Burn, and R.D. Palmiter Response of melanocortin-4 receptor-deficient mice to anorectic and orexigenic peptides Nat. Genet. 21 1999 119 122 (Pubitemid 29036297)
86. D.J. Marsh, G.I. Miura, K.A. Yagaloff, M.W. Schwartz, G.S. Barsh, and R.D. Palmiter Effects of neuropeptide Y deficiency on hypothalamic agouti-related protein expression and responsiveness to melanocortin analogues Brain Res. 848 1999 66 77 (Pubitemid 30001655)
87. 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 (Pubitemid 24335073)
88. J.D. Mul, D.P. Begg, S.I. Alsters, G. Haaften, K.J. Duran, D.A. D'Alessio, C.W. le Roux, S.C. Woods, D.A. Sandoval, A.I. Blakemore, E. Cuppen, M.M. van Haelst, and R.J. Seeley Effect of vertical sleeve gastrectomy in melanocortin receptor 4-deficient rats Am. J. Physiol. Endocrinol. Metab. 303 2012 E103 110
89. J.D. Mul, R. van Boxtel, D.J. Bergen, M.A. Brans, J.H. Brakkee, P.W. Toonen, K.M. Garner, R.A. Adan, and E. Cuppen Melanocortin receptor 4 deficiency affects body weight regulation, grooming behavior, and substrate preference in the rat Obesity 20 2012 612 621
90. J.R. Nicholson, J.C. Peter, A.C. Lecourt, Y.A. Barde, and K.G. Hofbauer Melanocortin-4 receptor activation stimulates hypothalamic brain-derived neurotrophic factor release to regulate food intake, body temperature and cardiovascular function J. Neuroendocrinol. 19 2007 974 982 (Pubitemid 350131084)
91. W.A. Nijenhuis, J.A. Kruijtzer, N. Wanders, D.H. Vrinten, K.M. Garner, W.M. Schaaper, R.H. Meloen, W.H. Gispen, R.M. Liskamp, and R.A. Adan Discovery and in vivo evaluation of new melanocortin-4 receptor-selective peptides Peptides 24 2003 271 280 (Pubitemid 36369347)
92. W.A. Nijenhuis, J. Oosterom, and R.A. Adan AgRP(83-132) acts as an inverse agonist on the human-melanocortin-4 receptor Mol. Endocrinol. 15 2001 164 171 (Pubitemid 32156049)
93. T.L. O'Donohue, and D.M. Dorsa The opiomelanotropinergic neuronal and endocrine systems Peptides 3 1982 353 395 (Pubitemid 12022681)
94. M.M. Ollmann, B.D. Wilson, Y.K. Yang, J.A. Kerns, Y. Chen, I. Gantz, and G.S. Barsh Antagonism of central melanocortin receptors in vitro and in vivo by agouti-related protein Science 278 1997 135 138 (Pubitemid 27446294)
95. J.D. Overton, and R.L. Leibel Mahoganoid and mahogany mutations rectify the obesity of the yellow mouse by effects on endosomal traffic of MC4R protein J. Biol. Chem. 286 2011 18914 18929
96. R. Pandit, J.W. de Jong, L.J. Vanderschuren, and R.A. Adan Neurobiology of overeating and obesity: the role of melanocortins and beyond Eur. J. Pharmacol. 2011 28 42 660 2011 28 42
97. L. Petreanu, D. Huber, A. Sobczyk, and K. Svoboda Channelrhodopsin-2- assisted circuit mapping of long-range callosal projections Nat. Neurosci. 10 2007 663 668 (Pubitemid 46652447)
98. 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 (Pubitemid 17182352)
99. A. Rediger, P. Tarnow, A. Bickenbach, M. Schaefer, H. Krude, A. Gruters, and H. Biebermann Heterodimerization of hypothalamic G-protein-coupled receptors involved in weight regulation Obes. Facts 2 2009 80 86
100. F. Rene, A. Muller, E. Jover, B. Kieffer, B. Koch, and J.P. Loeffler Melanocortin receptors and delta-opioid receptor mediate opposite signalling actions of POMC-derived peptides in CATH.a cells Eur. J. Neurosci. 10 1998 1885 1894
101. N. Sabatier, C. Caquineau, G. Dayanithi, P. Bull, A.J. Douglas, X.M. Guan, M. Jiang, L. Van der Ploeg, and G. Leng Alpha-melanocyte-stimulating hormone stimulates oxytocin release from the dendrites of hypothalamic neurons while inhibiting oxytocin release from their terminals in the neurohypophysis J. Neurosci. 23 2003 10351 10358 (Pubitemid 37444612)
102. B.D. Sachs The development of grooming and its expression in adult animals Ann. N.Y. Acad. Sci. 525 1988 1 17
103. M.W. Schwartz, S.C. Woods, D. Porte Jr., R.J. Seeley, and D.G. Baskin Central nervous system control of food intake Nature 404 2000 661 671 (Pubitemid 30205071)
104. R. Schwyzer ACTH: a short introductory review Ann. N.Y. Acad. Sci. 297 1977 3 26 (Pubitemid 8222793)
105. R. Schwyzer Structure and function in neuropeptides Proc. R. Soc. London B Biol. Sci. 210 1980 5 20 (Pubitemid 11214120)
106. J.V. Selkirk, L.M. Nottebaum, J. Lee, W. Yang, A.C. Foster, and S.M. Lechner Identification of differential melanocortin 4 receptor agonist profiles at natively expressed receptors in rat cortical astrocytes and recombinantly expressed receptors in human embryonic kidney cells Neuropharmacology 52 2007 459 466 (Pubitemid 46123362)
107. H. Shimizu, N.S. Shargill, G.A. Bray, T.T. Yen, and P.D. Gesellchen Effects of MSH on food intake, body weight and coat color of the yellow obese mouse Life Sci. 45 1989 543 552 (Pubitemid 19212821)
108. J.W. Sohn, L.E. Harris, E.D. Berglund, T. Liu, L. Vong, B.B. Lowell, N. Balthasar, K.W. Williams, and J.K. Elmquist Melanocortin 4 receptors reciprocally regulate sympathetic and parasympathetic preganglionic neurons Cell 152 2013 612 619
109. B.M. Spruijt, A.R. Cools, and W.H. Gispen The periaqueductal gray: a prerequisite for ACTH-induced excessive grooming Behav. Brain Res. 20 1986 19 25 (Pubitemid 16076652)
110. B.M. Spruijt, P.N. De Graan, A.N. Eberle, and W.H. Gispen Comparison of structural requirements of alpha-MSH and ACTH for inducing excessive grooming and pigment dispersion Peptides 6 1985 1185 1189 (Pubitemid 16166874)
111. B.M. Spruijt, and W.H. Gispen Behavioral sequences as an easily quantifiable parameter in experimental studies Physiol. Behav. 32 1984 707 710 (Pubitemid 14096995)
112. B.M. Spruijt, R. van den Bos, and F.T. Pijlman A concept of welfare based on reward evaluating mechanisms in the brain: anticipatory behaviour as an indicator for the state of reward systems Appl. Anim. Behav. Sci. 72 2001 145 171 (Pubitemid 32258533)
113. B.M. Spruijt, J.A. van Hooff, and W.H. Gispen Ethology and neurobiology of grooming behavior Physiol. Rev. 72 1992 825 852
114. G.M. Sutton, B. Duos, L.M. Patterson, and H.R. Berthoud Melanocortinergic modulation of cholecystokinin-induced suppression of feeding through extracellular signal-regulated kinase signaling in rat solitary nucleus Endocrinology 146 2005 3739 3747 (Pubitemid 41175741)
115. J.I. Szekely, E. Miglecz, Z. Dunai-Kovacs, I. Tarnawa, A.Z. Ronai, L. Graf, and S. Bajusz Attenuation of morphine tolerance and dependence by alpha-melanocyte stimulating hormone(alpha-MSH) Life Sci. 24 1979 1931 1938 (Pubitemid 9210105)
116. Y.X. Tao The melanocortin-4 receptor: physiology, pharmacology, and pathophysiology Endocr. Rev. 31 2010 506 543
117. L. Terenius, W.H. Gispen, and D. De Wied ACTH-like peptides and opiate receptors in the rat brain: structure-activity studies Eur. J. Pharmacol. 33 1975 395 399
118. S. Tsujii, and G.A. Bray Acetylation alters the feeding response to MSH and beta-endorphin Brain. Res. Bull. 23 1989 165 169 (Pubitemid 19277653)
119. M. van der Kraan, J.B. Tatro, M.L. Entwistle, J.H. Brakkee, J.P. Burbach, R.A. Adan, and W.H. Gispen Expression of melanocortin receptors and pro-opiomelanocortin in the rat spinal cord in relation to neurotrophic effects of melanocortins Mol. Brain Res. 63 1999 276 286 (Pubitemid 29036979)
120. L.H. Van der Ploeg, W.J. Martin, A.D. Howard, R.P. Nargund, C.P. Austin, X. Guan, J. Drisko, D. Cashen, I. Sebhat, A.A. Patchett, D.J. Figueroa, A.G. DiLella, B.M. Connolly, D.H. Weinberg, C.P. Tan, O.C. Palyha, S.S. Pong, T. MacNeil, C. Rosenblum, A. Vongs, R. Tang, H. Yu, A.W. Sailer, T.M. Fong, C. Huang, M.R. Tota, R.S. Chang, R. Stearns, C. Tamvakopoulos, G. Christ, D.L. Drazen, B.D. Spar, R.J. Nelson, and D.E. MacIntyre A role for the melanocortin 4 receptor in sexual function Proc. Natl. Acad. Sci. USA 99 2002 11381 11386 (Pubitemid 34920934)
121. J.W. van Nispen, and H.M. Greven Structure-activity relationships of peptides derived from ACTH, beta-LPH and MSH with regard to avoidance behavior in rats Pharmacol. Ther. 16 1982 67 102 (Pubitemid 12025695)
122. T.B. Van Wimersma Greidanus, D.K. Donker, F.F. Van Zinnicq Bergmann, R. Bekenkamp, C. Maigret, and B. Spruijt Comparison between excessive grooming induced by bombesin or by ACTH: the differential elements of grooming and development of tolerance Peptides 6 1985 369 372 (Pubitemid 16255644)
123. T.B. van Wimersma Greidanus, C. Maigret, J.A. Ten Haaf, B.M. Spruijt, and D.L. Colbern The influence of neurotensin, naloxone, and haloperidol on elements of excessive grooming behavior induced by ACTH Behav. Neural. Biol. 46 1986 137 144 (Pubitemid 16009983)
124. T.J. Van Wimersma Greidanus, D.K. Donker, R. Walhof, J.C. Van Grafhorst, N. De Vries, S.J. Van Schaik, C. Maigret, B.M. Spruijt, and D.L. Colbern The effects of neurotensin, naloxone and haloperidol on elements of excessive grooming behavior induced by bombesin Peptides 6 1985 1179 1183 (Pubitemid 16166873)
125. C. Vaughan, M. Moore, C. Haskell-Luevano, and N.E. Rowland Food motivated behavior of melanocortin-4 receptor knockout mice under a progressive ratio schedule Peptides 27 2006 2829 2835 (Pubitemid 44584424)
126. C.H. Vaughan, M.C. Moore, C. Haskell-Luevano, and N.E. Rowland Meal patterns and foraging in melanocortin receptor knockout mice Physiol. Behav. 84 2005 129 133 (Pubitemid 40092073)
127. D.H. Versteeg, P. Van Bergen, R.A. Adan, and D.J. De Wildt Melanocortins and cardiovascular regulation Eur. J. Pharmacol. 360 1998 1 14 (Pubitemid 28529390)
128. J.C. Von Frijtag, G. Croiset, W.H. Gispen, R.A. Adan, 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 (Pubitemid 28182031)
129. D.H. Vrinten, R.A. Adan, 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 (Pubitemid 33111691)
130. D.H. Vrinten, C.J. Kalkman, R.A. Adan, and W.H. Gispen Neuropathic pain: a possible role for the melanocortin system? Eur. J. Pharmacol. 429 2001 61 69 (Pubitemid 33043982)
131. S. Vrontou, A.M. Wong, K.K. Rau, H.R. Koerber, and D.J. Anderson Genetic identification of C fibres that detect massage-like stroking of hairy skin in vivo Nature 493 2013 669 673
132. J.M. Walker, H.C. Moises, D.H. Coy, G. Baldrighi, and H. Akil Nonopiate effects of dynorphin and des-Tyr-dynorphin Science 218 1982 1136 1138 (Pubitemid 13223319)
133. N. Wallingford, B. Perroud, Q. Gao, A. Coppola, E. Gyengesi, Z.W. Liu, X.B. Gao, A. Diament, K.A. Haus, Z. Shariat-Madar, F. Mahdi, S.L. Wardlaw, A.H. Schmaier, C.H. Warden, and S. Diano Prolylcarboxypeptidase regulates food intake by inactivating alpha-MSH in rodents J. Clin. Invest. 119 2009 2291 2303
134. S.L. Wardlaw Hypothalamic proopiomelanocortin processing and the regulation of energy balance Eur. J. Pharmacol. 660 2011 213 219
135. V.M. Wiegant, D. Colbern, T.J. van Wimersma Greidanus, and W.H. Gispen Differential behavioral effects of ACTH 4-10 and [D-Phe7] ACTH 4-10 Brain Res. Bull. 3 1978 167 170 (Pubitemid 8285806)
136. V.M. Wiegant, and W.H. Gispen ACTH-induced excessive grooming in the rat: latent activity of ACTH4-10 Behav. Biol. 19 1977 554 558 (Pubitemid 8094822)
137. V.M. Wiegant, J. Jolles, D.L. Colbern, E. Zimmermann, and W.H. Gispen Intracerebroventricular ACTH activates the pituitary-adrenal system:dissociation from a behavioral response Life Sci. 25 1979 1791 1796 (Pubitemid 10171944)
138. V.M. Wiegant, J. Jolles, and W.H. Gispen β-Endorphin grooming in the rat: single dose tolerant Dev. Neurosci. 1978 4
139. C.W. Wilkinson Roles of acetylation and other post-translational modifications in melanocortin function and interactions with endorphins Peptides 27 2006 453 471 (Pubitemid 43162918)
140. T. Willemse, M. Mudde, M. Josephy, and B.M. Spruijt The effect of haloperidol and naloxone on excessive grooming behavior of cats Eur. Neuropsychopharmacolog.: J. Eur. College Neuropsychopharmacolog. 4 1994 39 45 (Pubitemid 24087025)
141. D.L. Williams, H.J. Grill, S.M. Weiss, J.P. Baird, and J.M. Kaplan Behavioral processes underlying the intake suppressive effects of melanocortin 3/4 receptor activation in the rat Psychopharmacology 161 2002 47 53 (Pubitemid 34437968)
142. D.L. Williams, J.M. Kaplan, and H.J. Grill The role of the dorsal vagal complex and the vagus nerve in feeding effects of melanocortin-3/4 receptor stimulation Endocrinology 141 2000 1332 1337 (Pubitemid 32269100)
143. M.M. Wirth, and S.Q. Giraudo Agouti-related protein in the hypothalamic paraventricular nucleus: effect on feeding Peptides 21 2000 1369 1375
144. M.M. Wirth, and S.Q. Giraudo Effect of Agouti-related protein delivered to the dorsomedial nucleus of the hypothalamus on intake of a preferred versus a non-preferred diet Brain Res. 897 2001 169 174 (Pubitemid 32244365)
145. M.M. Wirth, P.K. Olszewski, C. Yu, A.S. Levine, and S.Q. Giraudo Paraventricular hypothalamic alpha-melanocyte-stimulating hormone and MTII reduce feeding without causing aversive effects Peptides 22 2001 129 134 (Pubitemid 32146692)
146. Z. Wu, Y. Xu, Y. Zhu, A.K. Sutton, R. Zhao, B.B. Lowell, D.P. Olson, and
147. B. Xu, E.H. Goulding, K. Zang, D. Cepoi, R.D. Cone, K.R. Jones, L.H. Tecott, and L.F. Reichardt Brain-derived neurotrophic factor regulates energy balance downstream of melanocortin-4 receptor Nat. Neurosci. 6 2003 736 742 (Pubitemid 36792889)
148. H. Zheng, M.M. Corkern, S.M. Crousillac, L.M. Patterson, C.B. Phifer, and H.R. Berthoud Neurochemical phenotype of hypothalamic neurons showing Fos expression 23 h after intracranial AgRP Am. J. Physiol. Regul. Integr. Comp. Physiol. 282 2002 R1773 1781
149. H. Zheng, L.M. Patterson, C.B. Phifer, and H.R. Berthoud Brain stem melanocortinergic modulation of meal size and identification of hypothalamic POMC projections Am. J. Physiol. Regul. Integr. Comp. Physiol. 289 2005 R247 258
150. H. Zwiers, V.J. Aloyo, and W.H. Gispen Behavioral and neurochemical effects of the new opioid peptide dynorphin-(1-13): comparison with other neuropeptides Life Sci. 28 1981 2545 2551 (Pubitemid 11094763)