Melanocortin-3 receptors and metabolic homeostasis

Progress in Molecular Biology and Translational Science

Volumn 114, Issue , 2013, Pages 109-146

  Begriche, Karima ^a,b   Girardet, Clemencé ^a   McDonald, Patricia ^a   Butler, Andrew A ^a  

^a SCRIPPS RESEARCH INSTITUTE   (United States)

^b Univ Rennes   (France)

Author keywords

Circadian Rhythm; Diabetes; Food intake; Insulin resistance; Keywords; Melanocortin; Metabolism; Obesity; Proopiomelanocortin

Indexed keywords

EID: 84872377428     PISSN: 18771173     EISSN: None     Source Type: Book Series    
DOI: 10.1016/B978-0-12-386933-3.00004-2     Document Type: Chapter

References (218)

1. K.M. Flegal, M.D. Carroll, C.L. Ogden, and C.L. Johnson Prevalence and trends in obesity among US adults, 1999-2000 JAMA 288 14 2002 1723 1727
2. M. Rosenbaum, R. Goldsmith, D. Bloomfield, A. Magnano, L. Weimer, and S. Heymsfield Low-dose leptin reverses skeletal muscle, autonomic, and neuroendocrine adaptations to maintenance of reduced weight J Clin Invest 115 12 2005 3579 3586
3. R.L. Leibel, M. Rosenbaum, and J. Hirsch Changes in energy expenditure resulting from altered body weight N Engl J Med 332 10 1995 621 628
4. C.K. Martin, L.K. Heilbronn, L. de Jonge, J.P. DeLany, J. Volaufova, and S.D. Anton Effect of calorie restriction on resting metabolic rate and spontaneous physical activity Obesity (Silver Spring) 15 12 2007 2964 2973
5. M.A. Stefater, H.E. Wilson-Perez, A.P. Chambers, D.A. Sandoval, and R.J. Seeley All bariatric surgeries are not created equal: insights from mechanistic comparisons Endocr Rev 33 2012 595 622
6. R.D. Cone Anatomy and regulation of the central melanocortin system Nat Neurosci 8 5 2005 571 578
7. M.G. Myers Jr., H. Munzberg, G.M. Leinninger, and R.L. Leshan The geometry of leptin action in the brain: more complicated than a simple ARC Cell Metab 9 2 2009 117 123
8. 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 5 2009 203 215
9. B.B. Lowell, and B.M. Spiegelman Towards a molecular understanding of adaptive thermogenesis Nature 404 6778 2000 652 660
10. M.H. Tschop, J.R. Speakman, J.R. Arch, J. Auwerx, J.C. Bruning, and L. Chan A guide to analysis of mouse energy metabolism Nat Methods 9 1 2011 57 63
11. A.A. Butler, and L.P. Kozak A recurring problem with the analysis of energy expenditure in genetic models expressing lean and obese phenotypes Diabetes 59 2 2010 323 329
12. E. Ravussin, and J.E. Galgani The implication of brown adipose tissue for humans Annu Rev Nutr 31 2011 33 47
13. L.P. Kozak Brown fat and the myth of diet-induced thermogenesis Cell Metab 11 4 2010 263 267
14. B. Cannon, and J. Nedergaard Nonshivering thermogenesis and its adequate measurement in metabolic studies J Exp Biol 214 Pt 2 2011 242 253
15. J.P. Flatt Issues and misconceptions about obesity Obesity (Silver Spring) 19 4 2011 676 686
16. J.R. Speakman, D.A. Levitsky, D.B. Allison, M.S. Bray, J.M. de Castro, and D.J. Clegg Set points, settling points and some alternative models: theoretical options to understand how genes and environments combine to regulate body adiposity Dis Model Mech 4 6 2011 733 745
17. M.W. Schwartz, S.C. Woods, R.J. Seeley, G.S. Barsh, D.G. Baskin, and R.L. Leibel Is the energy homeostasis system inherently biased toward weight gain? Diabetes 52 2 2003 232 238
18. B.M. King The rise, fall, and resurrection of the ventromedial hypothalamus in the regulation of feeding behavior and body weight Physiol Behav 87 2 2006 221 244
19. G.A. Bray, and D.A. York The MONA LISA hypothesis in the time of leptin Recent Prog Horm Res 53 1998 95 117 [discussion 117-8]
20. J.M. Friedman A war on obesity, not the obese Science 299 5608 2003 856 858
21. S. O'Rahilly Human genetics illuminates the paths to metabolic disease Nature 462 7271 2009 307 314
22. D.L. Coleman A historical perspective on leptin Nat Med 16 10 2010 1097 1099
23. Y. Zhang, R. Proenca, M. Maffei, M. Barone, L. Leopold, and J.M. Friedman Positional cloning of the mouse obese gene and its human homologue Nature 372 6505 1994 425 432
24. L.A. Tartaglia, M. Dembski, X. Weng, N. Deng, J. Culpepper, and R. Devos Identification and expression cloning of a leptin receptor, OB-R Cell 83 7 1995 1263 1271
25. I.S. Farooqi, and S. O'Rahilly Mutations in ligands and receptors of the leptin-melanocortin pathway that lead to obesity Nat Clin Pract Endocrinol Metab 4 10 2008 569 577
26. L. Gautron, and J.K. Elmquist Sixteen years and counting: an update on leptin in energy balance J Clin Invest 121 6 2011 2087 2093
27. I.S. Farooqi, J.M. Keogh, S. Kamath, S. Jones, W.T. Gibson, and R. Trussell Partial leptin deficiency and human adiposity Nature 414 6859 2001 34 35
28. M.G. Myers Jr., R.L. Leibel, R.J. Seeley, and M.W. Schwartz Obesity and leptin resistance: distinguishing cause from effect Trends Endocrinol Metab 21 11 2010 643 651
29. K.W. Williams, L.O. Margatho, C.E. Lee, M. Choi, S. Lee, and M.M. Scott Segregation of acute leptin and insulin effects in distinct populations of arcuate proopiomelanocortin neurons J Neurosci 30 7 2010 2472 2479
30. D.J. Marsh, G. Hollopeter, D. Huszar, R. Laufer, K.A. Yagaloff, and S.L. Fisher Response of melanocortin-4 receptor-deficient mice to anorectic and orexigenic peptides Nat Genet 21 1 1999 119 122
31. R.M. Lechan, and C. Fekete The TRH neuron: a hypothalamic integrator of energy metabolism Prog Brain Res 153 2006 209 235
32. K. Begriche, P.R. Levasseur, J. Zhang, J. Rossi, D. Skorupa, and L.A. Solt Genetic dissection of the functions of the melanocortin-3 receptor, a seven-transmembrane g-protein coupled receptor, suggests roles for central and peripheral receptors in energy homeostasis J Biol Chem 286 47 2011 40771 40781
33. K. Begriche, G.M. Sutton, and A.A. Butler Homeostastic and non-homeostatic functions of melanocortin-3 receptors in the control of energy balance and metabolism Physiol Behav 104 4 2011 546 554
34. G.M. Sutton, D. Perez-Tilve, R. Nogueiras, J. Fang, J.K. Kim, and R.D. Cone The melanocortin-3 receptor is required for entrainment to meal intake J Neurosci 28 48 2008 12946 12955
35. B.J. Renquist, J.G. Murphy, E.A. Larson, D. Olsen, R.F. Klein, and K.L. Ellacott Melanocortin-3 receptor regulates the normal fasting response Proc Natl Acad Sci USA 109 23 2012 E1489 E1498
36. J.M. Friedman Leptin at 14 y of age: an ongoing story Am J Clin Nutr 89 3 2009 973S 979S
37. E. Asilmaz, P. Cohen, M. Miyazaki, P. Dobrzyn, K. Ueki, and G. Fayzikhodjaeva Site and mechanism of leptin action in a rodent form of congenital lipodystrophy J Clin Invest 113 3 2004 414 424
38. P. Cohen, C. Zhao, X. Cai, J.M. Montez, S.C. Rohani, and P. Feinstein Selective deletion of leptin receptor in neurons leads to obesity J Clin Invest 108 8 2001 1113 1121
39. L. Vong, C. Ye, Z. Yang, B. Choi, S. Chua Jr., and B.B. Lowell Leptin action on GABAergic neurons prevents obesity and reduces inhibitory tone to POMC neurons Neuron 71 1 2011 142 154
40. K. Guo, J.E. McMinn, T. Ludwig, Y.H. Yu, G. Yang, and L. Chen Disruption of peripheral leptin signaling in mice results in hyperleptinemia without associated metabolic abnormalities Endocrinology 148 8 2007 3987 3997
41. C. de Luca, T.J. Kowalski, Y. Zhang, J.K. Elmquist, C. Lee, and M.W. Kilimann Complete rescue of obesity, diabetes, and infertility in db/db mice by neuron-specific LEPR-B transgenes J Clin Invest 115 12 2005 3484 3493
42. J.L. Halaas, C. Boozer, J. Blair-West, N. Fidahusein, D.A. Denton, and J.M. Friedman Physiological response to long-term peripheral and central leptin infusion in lean and obese mice Proc Natl Acad Sci USA 94 16 1997 8878 8883
43. R.S. Ahima, D. Prabakaran, C. Mantzoros, D. Qu, B. Lowell, and E. Maratos-Flier Role of leptin in the neuroendocrine response to fasting Nature 382 6588 1996 250 252
44. M. Rosenbaum, M. Sy, K. Pavlovich, R.L. Leibel, and J. Hirsch Leptin reverses weight loss-induced changes in regional neural activity responses to visual food stimuli J Clin Invest 118 7 2008 2583 2591
45. M. Rosenbaum, E.M. Murphy, S.B. Heymsfield, D.E. Matthews, and R.L. Leibel Low dose leptin administration reverses effects of sustained weight-reduction on energy expenditure and circulating concentrations of thyroid hormones J Clin Endocrinol Metab 87 5 2002 2391 2394
46. J.E. Galgani, F.L. Greenway, S. Caglayan, M.L. Wong, J. Licinio, and E. Ravussin Leptin replacement prevents weight loss-induced metabolic adaptation in congenital leptin-deficient patients J Clin Endocrinol Metab 95 2 2010 851 855
47. P.J. Enriori, A.E. Evans, P. Sinnayah, E.E. Jobst, L. Tonelli-Lemos, and S.K. Billes Diet-induced obesity causes severe but reversible leptin resistance in arcuate melanocortin neurons Cell Metab 5 3 2007 181 194
48. Z.A. Knight, K.S. Hannan, M.L. Greenberg, and J.M. Friedman Hyperleptinemia is required for the development of leptin resistance PLoS One 5 6 2010 e11376
49. J.L. Trevaskis, E.A. Meyer, J.E. Galgani, and A.A. Butler Counterintuitive effects of double-heterozygous null melanocortin-4 receptor and leptin genes on diet-induced obesity and insulin resistance in C57BL/6J mice Endocrinology 149 1 2008 174 184
50. P.M. Yen, and W.W. Chin Molecular mechanisms of dominant negative activity by nuclear hormone receptors Mol Endocrinol 8 11 1994 1450 1454
51. M.W. Miller, D.M. Duhl, H. Vrieling, S.P. Cordes, M.M. Ollmann, and B.M. Winkes Cloning of the mouse agouti gene predicts a secreted protein ubiquitously expressed in mice carrying the lethal yellow mutation Genes Dev 7 3 1993 454 467
52. S.J. Bultman, E.J. Michaud, and R.P. Woychik Molecular characterization of the mouse agouti locus Cell 71 7 1992 1195 1204
53. D. Lu, D. Willard, I.R. Patel, S. Kadwell, L. Overton, and T. Kost Agouti protein is an antagonist of the melanocyte-stimulating-hormone receptor Nature 371 6500 1994 799 802
54. 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 6612 1997 165 168
55. 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 10 1994 1298 1308
56. 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 1 1997 131 141
57. C. Vaisse, K. Clement, B. Guy-Grand, and P. Froguel A frameshift mutation in human MC4R is associated with a dominant form of obesity Nat Genet 20 2 1998 113 114
58. G.S. Yeo, I.S. Farooqi, S. Aminian, D.J. Halsall, R.G. Stanhope, and S. O'Rahilly A frameshift mutation in MC4R associated with dominantly inherited human obesity Nat Genet 20 2 1998 111 112
59. I.S. Farooqi, J.M. Keogh, G.S. Yeo, E.J. Lank, T. Cheetham, and S. O'Rahilly Clinical spectrum of obesity and mutations in the melanocortin 4 receptor gene N Engl J Med 348 12 2003 1085 1095
60. 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 6 2001 605 611
61. L. Ste Marie, G.I. Miura, D.J. Marsh, K. Yagaloff, and R.D. Palmiter A metabolic defect promotes obesity in mice lacking melanocortin-4 receptors Proc Natl Acad Sci USA 97 22 2000 12339 12344
62. G.M. Sutton, J.L. Trevaskis, M.W. Hulver, R.P. McMillan, N.J. Markward, and M.J. Babin Diet-genotype interactions in the development of the obese, insulin-resistant phenotype of C57BL/6J mice lacking melanocortin-3 or -4 receptors Endocrinology 147 5 2006 2183 2196
63. A.A. Butler The melanocortin system and energy balance Peptides 27 2 2006 281 290
64. D.C. Albarado, J. McClaine, J.M. Stephens, R.L. Mynatt, J. Ye, and A.W. Bannon Impaired coordination of nutrient intake and substrate oxidation in melanocortin-4 receptor knockout mice Endocrinology 145 1 2004 243 252
65. J. Rossi, N. Balthasar, D. Olson, M. Scott, E. Berglund, and C.E. Lee Melanocortin-4 receptors expressed by cholinergic neurons regulate energy balance and glucose homeostasis Cell Metab 13 2 2011 195 204
66. L. Zhou, G.M. Sutton, J.J. Rochford, R.K. Semple, D.D. Lam, and L.J. Oksanen Serotonin 2C receptor agonists improve type 2 diabetes via melanocortin-4 receptor signaling pathways Cell Metab 6 5 2007 398 405
67. R. Nogueiras, P. Wiedmer, D. Perez-Tilve, C. Veyrat-Durebex, J.M. Keogh, and G.M. Sutton The central melanocortin system directly controls peripheral lipid metabolism J Clin Invest 117 11 2007 3475 3488
68. S. Obici, Z. Feng, J. Tan, L. Liu, G. Karkanias, and L. Rossetti Central melanocortin receptors regulate insulin action J Clin Invest 108 7 2001 1079 1085
69. L. Huo, K. Gamber, S. Greeley, J. Silva, N. Huntoon, and X.H. Leng Leptin-dependent control of glucose balance and locomotor activity by POMC neurons Cell Metab 9 6 2009 537 547
70. N. Balthasar, L.T. Dalgaard, C.E. Lee, J. Yu, H. Funahashi, and T. Williams Divergence of melanocortin pathways in the control of food intake and energy expenditure Cell 123 3 2005 493 505
71. S. Wan, K.N. Browning, F.H. Coleman, G. Sutton, H. Zheng, and A. Butler Presynaptic melanocortin-4 receptors on vagal afferent fibers modulate the excitability of rat nucleus tractus solitarius neurons J Neurosci 28 19 2008 4957 4966
72. 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 4 2004 335 336
73. 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 1 2005 R247 R258
74. 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 9 2005 3739 3747
75. 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 4 2000 1332 1337
76. 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 23 1998 10128 10135
77. 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 1 1999 155 163
78. D.L. Williams, and M.W. Schwartz Neuroanatomy of body weight control: lessons learned from leptin J Clin Invest 121 6 2011 2152 2155
79. J.Z. Dong, D.B. DeOliveira, H.A. Halem, J.E. Taylor, P. Roubert, and P. Plas Novel melanocortin-4 receptor agonists that decrease food intake and body weight Adv Exp Med Biol 611 2009 485 486
80. J. Pantel, S.Y. Williams, D. Mi, J. Sebag, J.D. Corbin, and C.D. Weaver Development of a high throughput screen for allosteric modulators of melanocortin-4 receptor signaling using a real time cAMP assay Eur J Pharmacol 660 1 2011 139 147
81. J.S. Albert, N. Blomberg, A.L. Breeze, A.J. Brown, J.N. Burrows, and P.D. Edwards An integrated approach to fragment-based lead generation: philosophy, strategy and case studies from AstraZeneca's drug discovery programmes Curr Top Med Chem 7 16 2007 1600 1629
82. P.J. Emmerson, M.J. Fisher, L.Z. Yan, and J.P. Mayer Melanocortin-4 receptor agonists for the treatment of obesity Curr Top Med Chem 7 11 2007 1121 1130
83. F. Ujjainwalla, and I.K. Sebhat Small molecule ligands of the human melanocortin-4 receptor Curr Top Med Chem 7 11 2007 1068 1084
84. V.J. Hruby, M. Cai, J. Cain, J. Nyberg, and D. Trivedi Design of novel melanocortin receptor ligands: multiple receptors, complex pharmacology, the challenge Eur J Pharmacol 660 1 2011 88 93
85. J.R. Greenfield, J.W. Miller, J.M. Keogh, E. Henning, J.H. Satterwhite, and G.S. Cameron Modulation of blood pressure by central melanocortinergic pathways N Engl J Med 360 1 2009 44 52
86. A.A. Butler, R.A. Kesterson, K. Khong, M.J. Cullen, M.A. Pelleymounter, and J. Dekoning A unique metabolic syndrome causes obesity in the melanocortin-3 receptor-deficient mouse Endocrinology 141 9 2000 3518 3521
87. A.S. Chen, D.J. Marsh, M.E. Trumbauer, E.G. Frazier, X.M. Guan, and H. Yu Inactivation of the mouse melanocortin-3 receptor results in increased fat mass and reduced lean body mass Nat Genet 26 1 2000 97 102
88. J.L. Trevaskis, B. Gawronska-Kozak, G.M. Sutton, M. McNeil, J.M. Stephens, and S.R. Smith Role of adiponectin and inflammation in insulin resistance of Mc3r and Mc4r knockout mice Obesity (Silver Spring) 15 11 2007 2664 2672
89. K.L. Ellacott, J.G. Murphy, D.L. Marks, and R.D. Cone Obesity-induced inflammation in white adipose tissue is attenuated by loss of melanocortin-3 receptor signaling Endocrinology 148 12 2007 6186 6194
90. Y. Zhang, G.E. Kilroy, T.M. Henagan, V. Prpic-Uhing, W.G. Richards, and A.W. Bannon Targeted deletion of melanocortin receptor subtypes 3 and 4, but not CART, alters nutrient partitioning and compromises behavioral and metabolic responses to leptin FASEB J 19 11 2005 1482 1491
91. K. Begriche, O. Maston, J. Rossi, L. Burke, P. McDonald, and L.K. Heisler Melanocortin-3 receptors are required for metabolic adaptation to food restriction Genes Brain Behav 11 3 2012 291 302
92. G.M. Sutton, K. Begriche, K.G. Kumar, J.M. Gimble, D. Perez-Tilve, and R. Nogueiras Central nervous system melanocortin-3 receptors are required for synchronizing metabolism during entrainment to restricted feeding during the light cycle FASEB J 24 3 2010 862 872
93. K.G. Kumar, G.M. Sutton, J.Z. Dong, P. Roubert, P. Plas, and H.A. Halem Analysis of the therapeutic functions of novel melanocortin receptor agonists in MC3R- and MC4R-deficient C57BL/6J mice Peptides 30 10 2009 1892 1900
94. J.J. An, Y. Rhee, S.H. Kim, D.M. Kim, D.H. Han, and J.H. Hwang Peripheral effect of alpha-melanocyte-stimulating hormone on fatty acid oxidation in skeletal muscle J Biol Chem 282 5 2007 2862 2870
95. X.P. Ni, and M.H. Humphreys Abnormal glucose metabolism in hypertensive mice with genetically interrupted gamma-melanocyte stimulating hormone signaling fed a high-sodium diet Am J Hypertens 21 12 2008 1284 1287
96. L. Roselli-Rehfuss, K.G. Mountjoy, L.S. Robbins, M.T. Mortrud, M.J. Low, and J.B. Tatro Identification of a receptor for gamma melanotropin and other proopiomelanocortin peptides in the hypothalamus and limbic system Proc Natl Acad Sci USA 90 19 1993 8856 8860
97. 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 6 1994 1182 1190
98. 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 5335 1997 135 138
99. J.R. Shutter, M. Graham, A.C. Kinsey, S. Scully, R. Luthy, and K.L. Stark Hypothalamic expression of ART, a novel gene related to agouti, is up-regulated in obese and diabetic mutant mice Genes Dev 11 5 1997 593 602
100. A.S. Chen, J.M. Metzger, M.E. Trumbauer, X.M. Guan, H. Yu, and E.G. Frazier Role of the melanocortin-4 receptor in metabolic rate and food intake in mice Transgenic Res 9 2 2000 145 154
101. K. Rahmouni, W.G. Haynes, D.A. Morgan, and A.L. Mark Role of melanocortin-4 receptors in mediating renal sympathoactivation to leptin and insulin J Neurosci 23 14 2003 5998 6004
102. I. Gantz, and T.M. Fong The melanocortin system Am J Physiol Endocrinol Metab 284 3 2003 E468 E474
103. S. Nakanishi, A. Inoue, T. Kita, M. Nakamura, A.C. Chang, and S.N. Cohen Nucleotide sequence of cloned cDNA for bovine corticotropin-beta-lipotropin precursor Nature 278 5703 1979 423 427
104. T. Kita, A. Inoue, S. Nakanishi, and S. Numa Purification and characterization of the messenger RNA coding for bovine corticotropin/beta- lipotropin precursor Eur J Biochem 93 2 1979 213 220
105. J.L. Roberts, P.H. Seeburg, J. Shine, E. Herbert, J.D. Baxter, and H.M. Goodman Corticotropin and beta-endorphin: construction and analysis of recombinant DNA complementary to mRNA for the common precursor Proc Natl Acad Sci USA 76 5 1979 2153 2157
106. M.M. Ollmann, M.L. Lamoreux, B.D. Wilson, and G.S. Barsh Interaction of Agouti protein with the melanocortin 1 receptor in vitro and in vivo Genes Dev 12 3 1998 316 330
107. Y.K. Yang, M.M. Ollmann, B.D. Wilson, C. Dickinson, T. Yamada, and G.S. Barsh Effects of recombinant agouti-signaling protein on melanocortin action Mol Endocrinol 11 3 1997 274 280
108. R.D. Rosenfeld, L. Zeni, A.A. Welcher, L.O. Narhi, C. Hale, and J. Marasco Biochemical, biophysical, and pharmacological characterization of bacterially expressed human agouti-related protein Biochemistry 37 46 1998 16041 16052
109. M.L. Klebig, J.E. Wilkinson, J.G. Geisler, and R.P. Woychik Ectopic expression of the agouti gene in transgenic mice causes obesity, features of type II diabetes, and yellow fur Proc Natl Acad Sci USA 92 11 1995 4728 4732
110. W.L. Perry, C.M. Hustad, D.A. Swing, N.A. Jenkins, and N.G. Copeland A transgenic mouse assay for agouti protein activity Genetics 140 1 1995 267 274
111. M.R. Tota, T.S. Smith, C. Mao, T. MacNeil, R.T. Mosley, and L.H. Van der Ploeg Molecular interaction of Agouti protein and Agouti-related protein with human melanocortin receptors Biochemistry 38 3 1999 897 904
112. Y.K. Yang, D.A. Thompson, C.J. Dickinson, J. Wilken, G.S. Barsh, and S.B. Kent Characterization of Agouti-related protein binding to melanocortin receptors Mol Endocrinol 13 1 1999 148 155
113. M. Rossi, M.S. Kim, D.G. Morgan, C.J. Small, C.M. Edwards, and D. Sunter A C-terminal fragment of Agouti-related protein increases feeding and antagonizes the effect of alpha-melanocyte stimulating hormone in vivo Endocrinology 139 10 1998 4428 4431
114. J.W. Creemers, L.E. Pritchard, A. Gyte, P. Le Rouzic, S. Meulemans, and S.L. Wardlaw Agouti-related protein is posttranslationally cleaved by proprotein convertase 1 to generate agouti-related protein (AGRP)83-132: interaction between AGRP83-132 and melanocortin receptors cannot be influenced by syndecan-3 Endocrinology 147 4 2006 1621 1631
115. 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 1 2001 164 171
116. 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 1 2001 1 7
117. W. Siegrist, R. Drozdz, R. Cotti, D.H. Willard, W.O. Wilkison, and A.N. Eberle Interactions of alpha-melanotropin and agouti on B16 melanoma cells: evidence for inverse agonism of agouti J Recept Signal Transduct Res 17 1-3 1997 75 98
118. V. Tolle, and M.J. Low In vivo evidence for inverse agonism of Agouti-related peptide in the central nervous system of proopiomelanocortin- deficient mice Diabetes 57 1 2008 86 94
119. J.L. Smart, and M.J. Low Lack of proopiomelanocortin peptides results in obesity and defective adrenal function but normal melanocyte pigmentation in the murine C57BL/6 genetic background Ann N Y Acad Sci 994 2003 202 210
120. S. Srinivasan, C. Lubrano-Berthelier, C. Govaerts, F. Picard, P. Santiago, and B.R. Conklin Constitutive activity of the melanocortin-4 receptor is maintained by its N-terminal domain and plays a role in energy homeostasis in humans J Clin Invest 114 8 2004 1158 1164
121. T.R. Buch, D. Heling, E. Damm, T. Gudermann, and A. Breit Pertussis toxin-sensitive signaling of melanocortin-4 receptors in hypothalamic GT1-7 cells defines agouti-related protein as a biased agonist J Biol Chem 284 39 2009 26411 26420
122. S.I. Candille, C.B. Kaelin, B.M. Cattanach, B. Yu, D.A. Thompson, and M.A. Nix A b-defensin mutation causes black coat color in domestic dogs Science 318 5855 2007 1418 1423
123. A. Rediger, C.L. Piechowski, K. Habegger, A. Gruters, H. Krude, and M.H. Tschop MC4R dimerization in the paraventricular nucleus and GHSR/MC3R heterodimerization in the arcuate nucleus: is there relevance for body weight regulation Neuroendocrinology 95 4 2012 277 288
124. A. Rediger, C.L. Piechowski, C.X. Yi, P. Tarnow, R. Strotmann, and A. Gruters Mutually opposite signal modulation by hypothalamic heterodimerization of ghrelin and melanocortin-3 receptors J Biol Chem 286 45 2011 39623 39631
125. 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 18 1999 RC26
126. C. Haskell-Luevano, P. Chen, C. Li, K. Chang, M.S. Smith, and J.L. Cameron Characterization of the neuroanatomical distribution of agouti-related protein immunoreactivity in the rhesus monkey and the rat Endocrinology 140 3 1999 1408 1415
127. L. Yaswen, N. Diehl, M.B. Brennan, and U. Hochgeschwender Obesity in the mouse model of pro-opiomelanocortin deficiency responds to peripheral melanocortin Nat Med 5 9 1999 1066 1070
128. J.L. Smart, V. Tolle, and M.J. Low Glucocorticoids exacerbate obesity and insulin resistance in neuron-specific proopiomelanocortin-deficient mice J Clin Invest 116 2 2006 495 505
129. H. Krude, H. Biebermann, W. Luck, R. Horn, G. Brabant, and A. Gruters Severe early-onset obesity, adrenal insufficiency and red hair pigmentation caused by POMC mutations in humans Nat Genet 19 2 1998 155 157
130. I.S. Farooqi, S. Drop, A. Clements, J.M. Keogh, J. Biernacka, and S. Lowenbein Heterozygosity for a POMC-null mutation and increased obesity risk in humans Diabetes 55 9 2006 2549 2553
131. C.C. Cheung, D.K. Clifton, and R.A. Steiner Proopiomelanocortin neurons are direct targets for leptin in the hypothalamus Endocrinology 138 10 1997 4489 4492
132. B.D. Wilson, D. Bagnol, C.B. Kaelin, M.M. Ollmann, I. Gantz, and S.J. Watson Physiological and anatomical circuitry between Agouti-related protein and leptin signaling Endocrinology 140 5 1999 2387 2397
133. R.J. Seeley, K.A. Yagaloff, S.L. Fisher, P. Burn, T.E. Thiele, and G. van Dijk Melanocortin receptors in leptin effects Nature 390 6658 1997 349
134. E. van de Wall, R. Leshan, A.W. Xu, N. Balthasar, R. Coppari, and S.M. Liu Collective and individual functions of leptin receptor modulated neurons controlling metabolism and ingestion Endocrinology 149 4 2008 1773 1785
135. J.L. Trevaskis, and A.A. Butler Double leptin and melanocortin-4 receptor gene mutations have an additive effect on fat mass and are associated with reduced effects of leptin on weight loss and food intake Endocrinology 146 10 2005 4257 4265
136. Z.B. Andrews, Z.W. Liu, N. Walllingford, D.M. Erion, E. Borok, and J.M. Friedman UCP2 mediates ghrelin's action on NPY/AgRP neurons by lowering free radicals Nature 454 7206 2008 846 851
137. H.Y. Chen, M.E. Trumbauer, A.S. Chen, D.T. Weingarth, J.R. Adams, and E.G. Frazier Orexigenic action of peripheral ghrelin is mediated by neuropeptide Y and agouti-related protein Endocrinology 145 6 2004 2607 2612
138. M.A. Cowley, R.G. Smith, S. Diano, M. Tschop, N. Pronchuk, and K.L. Grove The distribution and mechanism of action of ghrelin in the CNS demonstrates a novel hypothalamic circuit regulating energy homeostasis Neuron 37 4 2003 649 661
139. J.W. Hill, C.F. Elias, M. Fukuda, K.W. Williams, E.D. Berglund, and W.L. Holland Direct insulin and leptin action on pro-opiomelanocortin neurons is required for normal glucose homeostasis and fertility Cell Metab 11 4 2010 286 297
140. H.V. Lin, L. Plum, H. Ono, R. Gutierrez-Juarez, M. Shanabrough, and E. Borok Divergent regulation of energy expenditure and hepatic glucose production by insulin receptor in AgRP and POMC neurons Diabetes 59 2 2010 337 346
141. A.C. Konner, R. Janoschek, L. Plum, S.D. Jordan, E. Rother, and X. Ma Insulin action in AgRP-expressing neurons is required for suppression of hepatic glucose production Cell Metab 5 6 2007 438 449
142. S.C. Benoit, E.L. Air, L.M. Coolen, R. Strauss, A. Jackman, and D.J. Clegg The catabolic action of insulin in the brain is mediated by melanocortins J Neurosci 22 20 2002 9048 9052
143. M. van den Top, K. Lee, A.D. Whyment, A.M. Blanks, and D. Spanswick Orexigen-sensitive NPY/AgRP pacemaker neurons in the hypothalamic arcuate nucleus Nat Neurosci 7 5 2004 493 494
144. Y. Xu, J.E. Jones, D. Kohno, K.W. Williams, C.E. Lee, and M.J. Choi 5-HT2CRs expressed by pro-opiomelanocortin neurons regulate energy homeostasis Neuron 60 4 2008 582 589
145. D.D. Lam, M.J. Przydzial, S.H. Ridley, G.S. Yeo, J.J. Rochford, and S. O'Rahilly Serotonin 5-HT2C receptor agonist promotes hypophagia via downstream activation of melanocortin 4 receptors Endocrinology 149 3 2008 1323 1328
146. L.K. Heisler, E.E. Jobst, G.M. Sutton, L. Zhou, E. Borok, and Z. Thornton-Jones Serotonin reciprocally regulates melanocortin neurons to modulate food intake Neuron 51 2 2006 239 249
147. L.K. Heisler, M.A. Cowley, L.H. Tecott, W. Fan, M.J. Low, and J.L. Smart Activation of central melanocortin pathways by fenfluramine Science 297 5581 2002 609 611
148. N. Ibrahim, M.A. Bosch, J.L. Smart, J. Qiu, M. Rubinstein, and O.K. Ronnekleiv Hypothalamic proopiomelanocortin neurons are glucose responsive and express K(ATP) channels Endocrinology 144 4 2003 1331 1340
149. L.E. Parton, C.P. Ye, R. Coppari, P.J. Enriori, B. Choi, and C.Y. Zhang Glucose sensing by POMC neurons regulates glucose homeostasis and is impaired in obesity Nature 449 7159 2007 228 232
150. P.S. Singru, E. Sanchez, C. Fekete, and R.M. Lechan Importance of melanocortin signaling in refeeding-induced neuronal activation and satiety Endocrinology 148 2 2007 638 646
151. E. Sanchez, P.S. Singru, R. Acharya, M. Bodria, C. Fekete, and A.M. Zavacki Differential effects of refeeding on melanocortin-responsive neurons in the hypothalamic paraventricular nucleus Endocrinology 149 9 2008 4329 4335
152. L.E. Johnstone, T.M. Fong, and G. Leng Neuronal activation in the hypothalamus and brainstem during feeding in rats Cell Metab 4 4 2006 313 321
153. C. Williams, and S.J. Hill GPCR signaling: understanding the pathway to successful drug discovery Methods Mol Biol 552 2009 39 50
154. S.R. Neves, P.T. Ram, and R. Iyengar G protein pathways Science 296 5573 2002 1636 1639
155. 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 5074 1992 1248 1251
156. I. Gantz, Y. Shimoto, Y. Konda, H. Miwa, C.J. Dickinson, and T. Yamada Molecular cloning, expression, and characterization of a fifth melanocortin receptor Biochem Biophys Res Commun 200 3 1994 1214 1220
157. 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 20 1993 15174 15179
158. I. Gantz, Y. Konda, T. Tashiro, Y. Shimoto, H. Miwa, and G. Munzert Molecular cloning of a novel melanocortin receptor J Biol Chem 268 11 1993 8246 8250
159. V. Chhajlani, R. Muceniece, and J.E. Wikberg Molecular cloning of a novel human melanocortin receptor Biochem Biophys Res Commun 218 2 1996 638
160. V. Chhajlani, R. Muceniece, and J.E. Wikberg Molecular cloning of a novel human melanocortin receptor Biochem Biophys Res Commun 195 2 1993 866 873
161. V. Chhajlani, and J.E. Wikberg Molecular cloning and expression of the human melanocyte stimulating hormone receptor cDNA FEBS Lett 309 3 1992 417 420
162. N. Griffon, V. Mignon, P. Facchinetti, J. Diaz, J.C. Schwartz, and P. Sokoloff Molecular cloning and characterization of the rat fifth melanocortin receptor Biochem Biophys Res Commun 200 2 1994 1007 1014
163. P. Barrett, A. MacDonald, R. Helliwell, G. Davidson, and P. Morgan Cloning and expression of a new member of the melanocyte-stimulating hormone receptor family J Mol Endocrinol 12 2 1994 203 213
164. O. Labbe, F. Desarnaud, D. Eggerickx, G. Vassart, and M. Parmentier Molecular cloning of a mouse melanocortin 5 receptor gene widely expressed in peripheral tissues Biochemistry 33 15 1994 4543 4549
165. S. Srinivasan, C. Vaisse, and B.R. Conklin Engineering the melanocortin-4 receptor to control G(s) signaling in vivo Ann N Y Acad Sci 994 2003 225 232
166. C.S. Kim, S.H. Lee, R.Y. Kim, B.J. Kim, S.Z. Li, and I.H. Lee Identification of domains directing specificity of coupling to G-proteins for the melanocortin MC3 and MC4 receptors J Biol Chem 277 35 2002 31310 31317
167. K.G. Mountjoy Distribution and function of melanocortin receptors within the brain Adv Exp Med Biol 681 2010 29 48
168. 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 3 2003 213 235
169. C.X. Yi, A. Kalsbeek, and M.H. Tschop Autonomic MC sets the metabolic tone Cell Metab 13 2 2011 121 123
170. H. Krude, and A. Gruters Implications of proopiomelanocortin (POMC) mutations in humans: the POMC deficiency syndrome Trends Endocrinol Metab 11 1 2000 15 22
171. H. Krude, H. Biebermann, D. Schnabel, M.Z. Tansek, P. Theunissen, and P.E. Mullis Obesity due to proopiomelanocortin deficiency: three new cases and treatment trials with thyroid hormone and ACTH4-10 J Clin Endocrinol Metab 88 10 2003 4633 4640
172. B.G. Challis, A.P. Coll, G.S. Yeo, S.B. Pinnock, S.L. Dickson, and R.R. Thresher Mice lacking pro-opiomelanocortin are sensitive to high-fat feeding but respond normally to the acute anorectic effects of peptide-YY(3-36) Proc Natl Acad Sci USA 101 13 2004 4695 4700
173. Y.S. Lee, B.G. Challis, D.A. Thompson, G.S. Yeo, J.M. Keogh, and M.E. Madonna A POMC variant implicates beta-melanocyte-stimulating hormone in the control of human energy balance Cell Metab 3 2 2006 135 140
174. H. Biebermann, T.R. Castaneda, F. van Landeghem, A. von Deimling, F. Escher, and G. Brabant A role for beta-melanocyte-stimulating hormone in human body-weight regulation Cell Metab 3 2 2006 141 146
175. C. Vaisse, K. Clement, E. Durand, S. Hercberg, B. Guy-Grand, and P. Froguel Melanocortin-4 receptor mutations are a frequent and heterogeneous cause of morbid obesity J Clin Invest 106 2 2000 253 262
176. G.S. Yeo, E.J. Lank, I.S. Farooqi, J. Keogh, B.G. Challis, and S. O'Rahilly Mutations in the human melanocortin-4 receptor gene associated with severe familial obesity disrupts receptor function through multiple molecular mechanisms Hum Mol Genet 12 5 2003 561 574
177. C.L. Wang, L. Liang, H.J. Wang, J.F. Fu, J. Hebebrand, and A. Hinney Several mutations in the melanocortin 4 receptor gene are associated with obesity in Chinese children and adolescents J Endocrinol Invest 29 10 2006 894 898
178. L.H. Larsen, S.M. Echwald, T.I. Sorensen, T. Andersen, B.S. Wulff, and O. Pedersen Prevalence of mutations and functional analyses of melanocortin 4 receptor variants identified among 750 men with juvenile-onset obesity J Clin Endocrinol Metab 90 1 2005 219 224
179. I.S. Farooqi, G.S. Yeo, J.M. Keogh, S. Aminian, S.A. Jebb, and G. Butler Dominant and recessive inheritance of morbid obesity associated with melanocortin 4 receptor deficiency J Clin Invest 106 2 2000 271 279
180. Y. Ohshiro, T. Sanke, K. Ueda, Y. Shimajiri, T. Nakagawa, and K. Tsunoda Molecular scanning for mutations in the melanocortin-4 receptor gene in obese/diabetic Japanese Ann Hum Genet 63 Pt 6 1999 483 487
181. W.D. Li, E.J. Joo, E.B. Furlong, M. Galvin, K. Abel, and C.J. Bell Melanocortin 3 receptor (MC3R) gene variants in extremely obese women Int J Obes Relat Metab Disord 24 2 2000 206 210
182. M.A. Calton, B.A. Ersoy, S. Zhang, J.P. Kane, M.J. Malloy, and C.R. Pullinger Association of functionally significant Melanocortin-4 but not Melanocortin-3 receptor mutations with severe adult obesity in a large North American case-control study Hum Mol Genet 18 6 2009 1140 1147
183. M. Mencarelli, B. Dubern, R. Alili, S. Maestrini, L. Benajiba, and M. Tagliaferri Rare melanocortin-3 receptor mutations with in vitro functional consequences are associated with human obesity Hum Mol Genet 20 2 2011 392 399
184. N. Yiannakouris, L. Melistas, M. Kontogianni, K. Heist, and C.S. Mantzoros The Val81 missense mutation of the melanocortin 3 receptor gene, but not the 1908c/T nucleotide polymorphism in lamin A/C gene, is associated with hyperleptinemia and hyperinsulinemia in obese Greek caucasians J Endocrinol Invest 27 8 2004 714 720
185. C. Schalin-Jantti, K. Valli-Jaakola, L. Oksanen, E. Martelin, K. Laitinen, and T. Krusius Melanocortin-3-receptor gene variants in morbid obesity Int J Obes Relat Metab Disord 27 1 2003 70 74
186. D.L. Marks, A.A. Butler, R. Turner, G. Brookhart, and R.D. Cone Differential role of melanocortin receptor subtypes in cachexia Endocrinology 144 4 2003 1513 1523
187. M.M. Hagan, P.A. Rushing, M.W. Schwartz, K.A. Yagaloff, P. Burn, and S.C. Woods Role of the CNS melanocortin system in the response to overfeeding J Neurosci 19 6 1999 2362 2367
188. Y. Xu, E.D. Berglund, J.W. Sohn, W.L. Holland, J.C. Chuang, and M. Fukuda 5-HT2CRs expressed by pro-opiomelanocortin neurons regulate insulin sensitivity in liver Nat Neurosci 13 12 2010 1457 1459
189. 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 1-2 1999 66 77
190. C. Fekete, S. Sarkar, W.M. Rand, J.W. Harney, C.H. Emerson, and A.C. Bianco Agouti-related protein (AGRP) has a central inhibitory action on the hypothalamic-pituitary-thyroid (HPT) axis; comparisons between the effect of AGRP and neuropeptide Y on energy homeostasis and the HPT axis Endocrinology 143 10 2002 3846 3853
191. G.M. Sutton, M. Josephine Babin, X. Gu, V.J. Hruby, and A.A. Butler A derivative of the melanocortin receptor antagonist SHU9119 (PG932) increases food intake when administered peripherally Peptides 29 1 2008 104 111
192. M. Navarro, J.M. Lerma-Cabrera, F. Carvajal, E.G. Lowery, I. Cubero, and T.E. Thiele Assessment of Voluntary Ethanol Consumption and the Effects of a Melanocortin (MC) Receptor Agonist on Ethanol Intake in Mutant C57BL/6J Mice Lacking the MC-4 Receptor Alcohol Clin Exp Res 35 6 2011 1058 1066
193. C.G. Joseph, H. Yao, J.W. Scott, N.B. Sorensen, R.N. Marnane, and K.G. Mountjoy Gamma-Melanocyte stimulation hormone (gamma-MSH) truncation studies results in the cautionary note that gamma-MSH is not selective for the mouse MC3R over the mouse MC5R Peptides 31 12 2010 2304 2313
194. N.E. Rowland, J.W. Schaub, K.L. Robertson, A. Andreasen, and C. Haskell-Luevano Effect of MTII on food intake and brain c-Fos in melanocortin-3, melanocortin-4, and double MC3 and MC4 receptor knockout mice Peptides. 31 12 2010 Dec 2314 2317
195. K. Begriche, G.M. Sutton, J. Fang, and A.A. Butler The role of melanocortin neuronal pathways in circadian biology: a new homeostatic output involving melanocortin-3 receptors? Obes Rev 10 Suppl. 2 2009 14 24
196. R.E. Mistlberger Circadian food-anticipatory activity: formal models and physiological mechanisms Neurosci Biobehav Rev 18 2 1994 171 195
197. W. Huang, K.M. Ramsey, B. Marcheva, and J. Bass Circadian rhythms, sleep, and metabolism J Clin Invest 121 6 2011 2133 2141
198. K.A. Lamia, K.F. Storch, and C.J. Weitz Physiological significance of a peripheral tissue circadian clock Proc Natl Acad Sci USA 105 39 2008 15172 15177
199. S. Zvonic, Z.E. Floyd, R.L. Mynatt, and J.M. Gimble Circadian rhythms and the regulation of metabolic tissue function and energy homeostasis Obesity (Silver Spring) 15 3 2007 539 543
200. J.S. Takahashi, H.K. Hong, C.H. Ko, and E.L. McDearmon The genetics of mammalian circadian order and disorder: implications for physiology and disease Nat Rev Genet 9 10 2008 764 775
201. F.W. Turek, C. Joshu, A. Kohsaka, E. Lin, G. Ivanova, and E. McDearmon Obesity and metabolic syndrome in circadian Clock mutant mice Science 308 5724 2005 1043 1045
202. R.D. Rudic, P. McNamara, A.M. Curtis, R.C. Boston, S. Panda, and J.B. Hogenesch BMAL1 and CLOCK, two essential components of the circadian clock, are involved in glucose homeostasis PLoS Biol 2 11 2004 e377
203. F.A. Scheer, M.F. Hilton, C.S. Mantzoros, and S.A. Shea Adverse metabolic and cardiovascular consequences of circadian misalignment Proc Natl Acad Sci USA 106 11 2009 4453 4458
204. D.M. Arble, J. Bass, A.D. Laposky, M.H. Vitaterna, and F.W. Turek Circadian timing of food intake contributes to weight gain Obesity (Silver Spring) 17 11 2009 2100 2102
205. S. Choi, L.S. Wong, C. Yamat, and M.F. Dallman Hypothalamic ventromedial nuclei amplify circadian rhythms: do they contain a food-entrained endogenous oscillator? J Neurosci 18 10 1998 3843 3852
206. J.J. Gooley, A. Schomer, and C.B. Saper The dorsomedial hypothalamic nucleus is critical for the expression of food-entrainable circadian rhythms Nat Neurosci 9 3 2006 398 407
207. A.J. Davidson Search for the feeding-entrainable circadian oscillator: a complex proposition Am J Physiol Regul Integr Comp Physiol 290 6 2006 R1524 R1526
208. R.E. Mistlberger, R.M. Buijs, E. Challet, C. Escobar, G.J. Landry, and A. Kalsbeek Standards of evidence in chronobiology: critical review of a report that restoration of Bmal1 expression in the dorsomedial hypothalamus is sufficient to restore circadian food anticipatory rhythms in Bmal1 -/- mice J Circadian Rhythms 7 2009 3
209. J. Bass, and J.S. Takahashi Circadian rhythms: redox redux Nature 469 7331 2011 476 478
210. J.S. Menet, and M. Rosbash When brain clocks lose track of time: cause or consequence of neuropsychiatric disorders Curr Opin Neurobiol 21 6 2011 849 857
211. A.C. Ribeiro, G. Ceccarini, C. Dupre, J.M. Friedman, D.W. Pfaff, and A.L. Mark Contrasting effects of leptin on food anticipatory and total locomotor activity PLoS One 6 8 2011 e23364
212. R.E. Mistlberger, and E.G. Marchant Enhanced food-anticipatory circadian rhythms in the genetically obese Zucker rat Physiol Behav 66 2 1999 329 335
213. S.G. Bouret, S.J. Draper, and R.B. Simerly Trophic action of leptin on hypothalamic neurons that regulate feeding Science 304 5667 2004 108 110
214. G.J. Landry, G.R. Yamakawa, I.C. Webb, R.J. Mear, and R.E. Mistlberger The dorsomedial hypothalamic nucleus is not necessary for the expression of circadian food-anticipatory activity in rats J Biol Rhythms 22 6 2007 467 478
215. T. Moriya, R. Aida, T. Kudo, M. Akiyama, M. Doi, and N. Hayasaka The dorsomedial hypothalamic nucleus is not necessary for food-anticipatory circadian rhythms of behavior, temperature or clock gene expression in mice Eur J Neurosci 29 7 2009 1447 1460
216. K.W. Kim, J.W. Sohn, D. Kohno, Y. Xu, K. Williams, and J.K. Elmquist SF-1 in the ventral medial hypothalamic nucleus: a key regulator of homeostasis Mol Cell Endocrinol 336 1-2 2011 219 223
217. R.E. Mistlberger, and A. Rechtschaffen Recovery of anticipatory activity to restricted feeding in rats with ventromedial hypothalamic lesions Physiol Behav 33 2 1984 227 235
218. R.E. Mistlberger Neurobiology of food anticipatory circadian rhythms Physiol Behav 104 2011 535 545