Astrocytes: New targets of melanocortin 4 receptor actions

Journal of Molecular Endocrinology

Volumn 51, Issue 2, 2013, Pages

  Caruso, Carla ^a   Carniglia, Lila ^a   Durand, Daniela ^a   Scimonelli, Teresa N ^b   Lasaga, Mercedes ^a  

^a UNIVERSIDAD DE BUENOS AIRES   (Argentina)

^b UNIVERSIDAD NACIONAL DE CÓRDOBA   (Argentina)

Author keywords

Astrocytes; Energy homeostasis; Inflammation; MC4R; Neuroprotection

Indexed keywords

ADIPONECTIN RECEPTOR 1; ALPHA INTERMEDIN; BETA INTERMEDIN; BIM 22493; BRAIN DERIVED NEUROTROPHIC FACTOR; CORTICOTROPIN; CYCLOOXYGENASE 1; FATTY ACID; GAMMA INTERMEDIN; GLIAL FIBRILLARY ACIDIC PROTEIN; GLUCOSE; GLYCOGEN; IMMUNOGLOBULIN ENHANCER BINDING PROTEIN; INDUCIBLE NITRIC OXIDE SYNTHASE; INTERLEUKIN 1BETA; INTERLEUKIN 6; LACTIC ACID; LEPTIN; MELANOCORTIN; MELANOCORTIN 4 RECEPTOR; MELANOCORTIN RECEPTOR AGONIST; PEROXISOME PROLIFERATOR ACTIVATED RECEPTOR GAMMA; PYRUVIC ACID; TOLL LIKE RECEPTOR 4; TRANSFORMING GROWTH FACTOR BETA; TUMOR NECROSIS FACTOR ALPHA; UNCLASSIFIED DRUG; VIMENTIN;

ALLERGIC ENCEPHALOMYELITIS; ALZHEIMER DISEASE; ANTIINFLAMMATORY ACTIVITY; APOPTOSIS; ARCUATE NUCLEUS; ASTROCYTE; ASTROCYTOSIS; BLOOD BRAIN BARRIER; BRAIN ISCHEMIA; CALCIUM SIGNALING; CELL ACTIVATION; CELL INFILTRATION; CELL INTERACTION; CELL PROLIFERATION; DNA FRAGMENTATION; ENCEPHALITIS; ENDOTHELIUM CELL; FEVER; GAP JUNCTION; GLIA CELL; GLIOSIS; GLYCOSYLATION; GRAY MATTER; HOMEOSTASIS; HUMAN; HYPOTHALAMIC PARAVENTRICULAR NUCLEUS; HYPOTHALAMUS; INFLAMMATION; INSULIN SENSITIVITY; LIPID DIET; MEMORY; MICROGLIA; NERVE FIBER GROWTH; NERVOUS SYSTEM INFLAMMATION; NEUROPROTECTION; NOCICEPTION; NONHUMAN; OBESITY; OVERNUTRITION; OXIDATIVE STRESS; PALMITOYLATION; PRIORITY JOURNAL; REPRODUCTION; REVIEW; SEXUAL FUNCTION; SIGNAL TRANSDUCTION; SOLITARY TRACT NUCLEUS; SPINAL CORD INJURY; WEIGHT REDUCTION; WHITE MATTER;

ASTROCYTES; ENERGY HOMEOSTASIS; INFLAMMATION; MC4R; NEUROPROTECTION;

ANIMALS; ASTROCYTES; ENERGY METABOLISM; HUMANS; INFLAMMATION; MELANOCORTINS; RECEPTOR, MELANOCORTIN, TYPE 4;

EID: 84884618116     PISSN: 09525041     EISSN: 14796813     Source Type: Journal    
DOI: 10.1530/JME-13-0064     Document Type: Review

References (214)

1. Abbott C.R., Rossi M, Kim M, Al Ahmed SH, TaylorGM, Ghatei MA, SmithDM & Bloom SR 2000 Investigation of the melanocyte stimulating hormones on food intake. Lack of evidence to support a role for the melanocortin-3-receptor. Brain Research 869 203-210. (doi:10.1016/ S0006-8993(00)02386-6)
2. Abbott N.J., Patabendige AA, Dolman DE, Yusof SR & Begley DJ 2010 Structure and function of the blood-brain barrier. Neurobiology of Disease 37 13-25. (doi:10.1016/j.nbd.2009.07.030)
3. Adan R.A., van der Kraan M, Doornbos RP, Bar PR, Burbach JP & Gispen WH 1996 Melanocortin receptors mediate α-MSH-induced stimulation of neurite outgrowth in neuro 2A cells. Brain Research. Molecular Brain Research 36 37-44. (doi:10.1016/0169-328X(95)00236-L)
4. Aigner L & Bogdahn U 2008 TGF-β in neural stem cells and in tumors of the central nervous system. Cell and Tissue Research 331 225-241. (doi:10.1007/s00441-007-0466-7)
5. Albrecht P.J., Dahl JP, Stoltzfus OK, Levenson R & Levison SW 2002 Ciliary neurotrophic factor activates spinal cord astrocytes, stimulating their production and release of fibroblast growth factor-2, to increase motor neuron survival. Experimental Neurology 173 46-62. (doi:10.1006/exnr. 2001.7834)
6. Aleshin S., Grabeklis S, Hanck T, Sergeeva M & Reiser G 2009 Peroxisome proliferator-activated receptor (PPAR)-gamma positively controls and PPARalpha negatively controls cyclooxygenase-2 expression in rat brain astrocytes through a convergence on PPARbeta/delta via mutual control of PPAR expression levels. Molecular Pharmacology 76 414-424. (doi:10.1124/mol.109.056010)
7. Anderson CM& Swanson RA 2000 Astrocyte glutamate transport: review of properties, regulation, and physiological functions. Glia 32 1-14. (doi:10.1002/1098-1136(200010)32:1!1::AID-GLIA10O3.0.CO;2-W)
8. Arsenijevic D., de Bilbao F, Plamondon J, Paradis E, Vallet P, Richard D, Langhans W & Giannakopoulos P 2006 Increased infarct size and lack of hyperphagic response after focal cerebral ischemia in peroxisome proliferator-activated receptor β-deficientmice. Journal of Cerebral Blood Flow and Metabolism 26 433-445. (doi:10.1038/sj.jcbfm.9600200)
9. Bagnol D., Lu XY, Kaelin CB, Day HE, OllmannM, Gantz I, Akil H, Barsh GS &Watson SJ 1999 Anatomy of an endogenous antagonist: relationship between Agouti-related protein and proopiomelanocortin in brain. Journal of Neuroscience 19 RC26.
10. Bell RC & Lipton JM 1987 Pulsatile release of antipyretic neuropeptide α-MSH from septum of rabbit during fever. American Journal of Physiology 252 R1152-R1157.
11. Benjannet S., Rondeau N, Day R, Chretien M & Seidah NG 1991 PC1 and PC2 are proprotein convertases capable of cleaving proopiomelanocortin at distinct pairs of basic residues. PNAS 88 3564-3568. (doi:10.1073/pnas.88.9.3564)
12. Benjelloun N., Joly LM, Palmier B, Plotkine M & Charriaut-Marlangue C 2003 Apoptotic mitochondrial pathway in neurones and astrocytes after neonatal hypoxia-ischaemia in the rat brain. Neuropathology and Applied Neurobiology 29 350-360. (doi:10.1046/j.1365-2990.2003. 00467.x)
13. Benoit S.C., Schwartz MW, Lachey JL, Hagan MM, Rushing PA, Blake KA, Yagaloff K.A., Kurylko G, Franco L, Danhoo W et al. 2000 A novel selective melanocortin-4 receptor agonist reduces food intake in rats and mice without producing aversive consequences. Journal of Neuroscience 20 3442-3448.
14. Benoit S.C., Air EL, Coolen LM, Strauss R, Jackman A, Clegg DJ, Seeley RJ & Woods SC 2002 The catabolic action of insulin in the brain is mediated by melanocortins. Journal of Neuroscience 22 9048-9052.
15. Benveniste E.N., Tang LP & Law RM 1995 Differential regulation of astrocyte TNF-α expression by the cytokines TGF-β, IL-6 and IL-10. International Journal of Developmental Neuroscience 13 341-349. (doi:10.1016/0736-5748(94)00061-7)
16. Berthet C., Lei H, Thevenet J, Gruetter R, Magistretti PJ & Hirt L 2009 Neuroprotective role of lactate after cerebral ischemia. Journal of Cerebral Blood Flow and Metabolism 29 1780-1789. (doi:10.1038/jcbfm. 2009.97)
17. Bertolini A., Tacchi R & Vergoni AV 2009 Brain effects of melanocortins. Pharmacological Research 59 13-47. (doi:10.1016/j.phrs.2008.10.005)
18. Bertorelli R., Fredduzzi S, Tarozzo G, Campanella M, Grundy R, Beltramo M & Reggiani A 2005 Endogenous and exogenous melanocortin antagonists induce anti-allodynic effects in a model of rat neuropathic pain. Behavioural Brain Research 157 55-62. (doi:10.1016/j.bbr.2004.06.008)
19. Bitto A., Polito F, Irrera N, Calo M, Spaccapelo L, Marini HR, Giuliani D, Ottani A., Rinaldi M, Minutoli L et al. 2012 Protective effects of melanocortins on short-term changes in a rat model of traumatic brain injury*. Critical Care Medicine 40 945-951. (doi:10.1097/CCM. 0b013e318236efde)
20. 2+ homeostasis and cell death. Journal of Neurochemistry 70 958-970. (doi:10.1046/j.1471-4159.1998. 70030958.x)
21. Blanchard S.G., Harris CO, Ittoop OR, Nichols JS, Parks DJ, Truesdale AT & Wilkison WO 1995 Agouti antagonism of melanocortin binding and action in the B16F10 murine melanoma cell line. Biochemistry 34 10406-10411. (doi:10.1021/bi00033a012)
22. Bo L., Dawson TM, Wesselingh S, Mork S, Choi S, Kong PA, Hanley D & Trapp BD 1994 Induction of nitric oxide synthase in demyelinating regions of multiple sclerosis brains. Annals of Neurology 36 778-786. (doi:10.1002/ana.410360515)
23. Boston BA & Cone RD 1996 Characterization of melanocortin receptor subtype expression in murine adipose tissues and in the 3T3-L1 cell line. Endocrinology 137 2043-2050. (doi:10.1210/en.137.5.2043)
24. Brambilla R., Bracchi-Ricard V, Hu WH, Frydel B, Bramwell A, Karmally S, Green EJ& Bethea JR 2005 Inhibition of astroglial nuclear factor kappaB reduces inflammation and improves functional recovery after spinal cord injury. Journal of Experimental Medicine 202 145-156. (doi:10.1084/ jem.20041918)
25. Bright J.J., Kanakasabai S, Chearwae W & Chakraborty S 2008 PPAR regulation of inflammatory signaling in CNS diseases. PPAR Research 2008 658520. (doi:10.1155/2008/658520)
26. Bronstein D.M., Schafer MK, Watson SJ & Akil H 1992 Evidence that β-endorphin is synthesized in cells in the nucleus tractus solitarius: detection of POMC mRNA. Brain Research 587 269-275. (doi:10.1016/ 0006-8993(92)91007-2)
27. Buffo A., Rolando C & Ceruti S 2010 Astrocytes in the damaged brain: molecular and cellular insights into their reactive response and healing potential. Biochemical Pharmacology 79 77-89. (doi:10.1016/j.bcp.2009. 09.014)
28. Carniglia L., Durand D, Caruso C & Lasaga M 2013 Effect of NDP-α-MSH on PPAR-γ and-β expression and anti-inflammatory cytokine release in rat astrocytes and microglia. PLoS ONE 8 e57313. (doi:10.1371/journal. pone.0057313)
29. Caruso C., Mohn C, Karara AL, Rettori V, Watanobe H, Schioth HB, Seilicovich A & Lasaga M 2004 α-Melanocyte-stimulating hormone through melanocortin-4 receptor inhibits nitric oxide synthase and cyclooxygenase expression in the hypothalamus of male rats. Neuroendocrinology 79 278-286. (doi:10.1159/000079321)
30. Caruso C., Durand D, Schioth HB, Rey R, Seilicovich A & Lasaga M 2007 Activation of melanocortin 4 receptors reduces the inflammatory response and prevents apoptosis induced by lipopolysaccharide and interferon-gamma in astrocytes. Endocrinology 148 4918-4926. (doi:10.1210/en.2007-0366)
31. Caruso C., Sanchez M, Durand D, Perez Mde L, Gonzalez PV, Lasaga M & Scimonelli TN 2010 α-Melanocyte-stimulating hormone modulates lipopolysaccharide plus interferon-gamma-induced tumor necrosis factor-α expression but not tumor necrosis factor-α receptor expression in cultured hypothalamic neurons. Journal of Neuroimmunology 227 52-59. (doi:10.1016/j.jneuroim.2010.06.013)
32. Caruso C., Carniglia L, Durand D, Gonzalez PV, Scimonelli TN & Lasaga M 2012 Melanocortin 4 receptor activation induces brain-derived neurotrophic factor expression in rat astrocytes through cyclic AMP-protein kinase A pathway. Molecular and Cellular Endocrinology 348 47-54. (doi:10.1016/j.mce.2011.07.036)
33. Caruso C., Carniglia L, Durand D & Lasaga M 2013 MC4R activation induces BDNF expression through ERK and PI3K in rat astrocytes. XI European Meeting on Glial Cells in Health and Disease. Glia 61 (Supplement S1) S186. (doi:10.1002/glia.22530)
34. Cassina P., Peluffo H, PeharM, Martinez-Palma L, RessiaA, BeckmanJS, Estevez AG & Barbeito L 2002 Peroxynitrite triggers a phenotypic transformation in spinal cord astrocytes that inducesmotor neuron apoptosis. Journal of Neuroscience Research 67 21-29. (doi:10.1002/jnr.10107)
35. Catania A 2007 The melanocortin system in leukocyte biology. Journal of Leukocyte Biology 81 383-392. (doi:10.1189/jlb.0706426)
36. Catania A 2008 Neuroprotective actions of melanocortins: a therapeutic opportunity. Trends in Neurosciences 31 353-360. (doi:10.1016/j.tins. 2008.04.002)
37. Catania A., Suffredini AF & Lipton JM 1995 Endotoxin causes release of α-melanocyte-stimulating hormone in normal human subjects. Neuroimmunomodulation 2 258-262. (doi:10.1159/000097204)
38. Catania A., Gatti S, Colombo G & Lipton JM 2004 Targeting melanocortin receptors as a novel strategy to control inflammation. Pharmacological Reviews 56 1-29. (doi:10.1124/pr.56.1.1)
39. Chai B., Li JY, Zhang W, Newman E, Ammori J & Mulholland MW 2006 Melanocortin-4 receptor-mediated inhibition of apoptosis in immortalized hypothalamic neurons via mitogen-activated protein kinase. Peptides 27 2846-2857. (doi:10.1016/j.peptides.2006.05.005)
40. Chai B., Li JY, Zhang W, Wu X, Zhang C & Mulholland MW 2010 Melanocortin-4 receptor activation promotes insulin-stimulated mTOR signaling. Peptides 31 1888-1893. (doi:10.1016/j.peptides.2010.06.028)
41. Chan L.F., Webb TR, Chung TT, Meimaridou E, Cooray SN, Guasti L, Chapple J.P., Egertova M, Elphick MR, Cheetham ME et al. 2009 MRAP and MRAP2 are bidirectional regulators of the melanocortin receptor family. PNAS 106 6146-6151. (doi:10.1073/pnas.0809918106)
42. Chen J., Jin K, Chen M, Pei W, Kawaguchi K, Greenberg DA & Simon RP 1997a Early detection of DNA strand breaks in the brain after transient focal ischemia: implications for the role of DNA damage in apoptosis and neuronal cell death. Journal of Neurochemistry 69 232-245. (doi:10.1046/j.1471-4159.1997.69010232.x)
43. Chen W., Kelly MA, Opitz-Araya X, Thomas RE, Low MJ & Cone RD 1997b Exocrine gland dysfunction in MC5-R-deficient mice: evidence for coordinated regulation of exocrine gland function by melanocortin peptides. Cell 91 789-798. (doi:10.1016/S0092-8674(00)80467-5)
44. Chen A.S., Marsh DJ, Trumbauer ME, Frazier EG, Guan XM, Yu H, Rosenblum C.I., Vongs A, Feng Y, Cao L et al. 2000a Inactivation of the mouse melanocortin-3 receptor results in increased fat mass and reduced lean body mass. Nature Genetics 26 97-102. (doi:10.1038/79254)
45. Chen A.S., Metzger JM, Trumbauer ME, Guan XM, Yu H, Frazier EG, Marsh D.J., Forrest MJ, Gopal-Truter S, Fisher J et al. 2000b Role of the melanocortin-4 receptor in metabolic rate and food intake in mice. Transgenic Research 9 145-154. (doi:10.1023/A:1008983615045)
46. Chhajlani V 1996 Distribution of cDNA for melanocortin receptor subtypes in human tissues. Biochemistry & Molecular Biology International 38 73-80.
47. Chhajlani V & Wikberg JE 1992 Molecular cloning and expression of the human melanocyte stimulating hormone receptor cDNA. FEBS Letters 309 417-420. (doi:10.1016/0014-5793(92)80820-7)
48. Climent E., Sancho-Tello M, Minana R, Barettino D & Guerri C 2000 Astrocytes in culture express the full-length Trk-B receptor and respond to brain derived neurotrophic factor by changing intracellular calcium levels: effect of ethanol exposure in rats. Neuroscience Letters 288 53-56. (doi:10.1016/S0304-3940(00)01207-6)
49. Cowley M.A., Smart JL, Rubinstein M, Cerdan MG, Diano S, Horvath TL, Cone RD & Low MJ 2001 Leptin activates anorexigenic POMC neurons through a neural network in the arcuate nucleus. Nature 411 480-484. (doi:10.1038/35078085)
50. 2 and NO induced by IL-1β differently. Neuroscience Letters 409 168-172. (doi:10.1016/j.neulet.2006.09.034)
51. Cui J., Ou S, He WJ, Du L, Zhao YD & Ruan HZ 2011 Prevention of extracellular ADP-induced ATP accumulation of the cultured rat spinal astrocytes via P2Y(1)-mediated inhibition of AMPK. Neuroscience Letters 503 244-249. (doi:10.1016/j.neulet.2011.08.045)
52. Damm E., Buech TR, Gudermann T & Breit A 2012 Melanocortin-induced PKA activation inhibits AMPK activity via ERK-1/2 and LKB-1 in hypothalamic GT1-7 cells. Molecular Endocrinology 26 643-654. (doi:10.1210/me.2011-1218)
53. Daniels D., Patten CS, Roth JD, Yee DK & Fluharty SJ 2003 Melanocortin receptor signaling through mitogen-activated protein kinase in vitro and in rat hypothalamus. Brain Research 986 1-11. (doi:10.1016/S0006-8993(03)03162-7)
54. Daynes R.A., Robertson BA, Cho BH, Burnham DK & Newton R 1987 α-Melanocyte-stimulating hormone exhibits target cell selectivity in its capacity to affect interleukin 1-inducible responses in vivo and in vitro. Journal of Immunology 139 103-109.
55. DeBoer MD 2010 Update on melanocortin interventions for cachexia: progress toward clinical application. Nutrition 26 146-151. (doi:10.1016/j.nut.2009.07.003)
56. De Souza CT, Araujo EP, Bordin S, Ashimine R, Zollner RL, Boschero AC, Saad MJ & Velloso LA 2005 Consumption of a fat-rich diet activates a proinflammatory response and induces insulin resistance in the hypothalamus. Endocrinology 146 4192-4199. (doi:10.1210/ en.2004-1520)
57. Dhopeshwarkar GA &Mead JF 1973 Uptake and transport of fatty acids into the brain and the role of the blood-brain barrier system. Advances in Lipid Research 11 109-142.
58. Dinulescu DM & Cone RD 2000 Agouti and agouti-related protein: analogies and contrasts. Journal of Biological Chemistry 275 6695-6698. (doi:10.1074/jbc.275.10.6695)
59. Djalali S., Holtje M, Grosse G, Rothe T, Stroh T, Grosse J, Deng DR, Hellweg R., Grantyn R, Hortnagl H et al. 2005 Effects of brain-derived neurotrophic factor (BDNF) on glial cells and serotonergic neurones during development. Journal of Neurochemistry 92 616-627. (doi:10.1111/j.1471-4159.2004.02911.x)
60. Dmitrieva V.G., Povarova OV, Skvortsova VI, Limborska SA, Myasoedov NF & Dergunova LV 2010 Semax and Pro-Gly-Pro activate the transcription of neurotrophins and their receptor genes after cerebral ischemia. Cellular and Molecular Neurobiology 30 71-79. (doi:10.1007/s10571-009-9432-0)
61. Dringen R., Gebhardt R & Hamprecht B 1993 Glycogen in astrocytes: possible function as lactate supply for neighboring cells. Brain Research 623 208-214. (doi:10.1016/0006-8993(93)91429-V)
62. Durand D., Caruso C, Carniglia L & LasagaM 2010 Metabotropic glutamate receptor 3 activation prevents nitric oxide-induced death in cultured rat astrocytes. Journal of Neurochemistry 112 420-433. (doi:10.1111/ j.1471-4159.2009.06469.x)
63. Ehrlich L.C., Peterson PK & Hu S 1999 Interleukin (IL)-1β-mediated apoptosis of human astrocytes. Neuroreport 10 1849-1852. (doi:10.1097/00001756-199906230-00009)
64. Escartin C., Pierre K, Colin A, Brouillet E, Delzescaux T, Guillermier M, Dhenain M., Deglon N, Hantraye P, Pellerin L et al. 2007 Activation of astrocytes by CNTF induces metabolic plasticity and increases resistance to metabolic insults. Journal of Neuroscience 27 7094-7104. (doi:10.1523/JNEUROSCI.0174-07.2007)
65. Evans T., McCarthy KD & Harden TK 1984 Regulation of cyclic AMP accumulation by peptide hormone receptors in immunocytochemically defined astroglial cells. Journal of Neurochemistry 43 131-138. (doi:10.1111/j.1471-4159.1984.tb06688.x)
66. Faulkner J.R., Herrmann JE, Woo MJ, Tansey KE, Doan NB & Sofroniew MV 2004 Reactive astrocytes protect tissue and preserve function after spinal cord injury. Journal of Neuroscience 24 2143-2155. (doi:10.1523/ JNEUROSCI.3547-03.2004)
67. Forslin Aronsson S, Spulber S, Popescu LM, Winblad B, Post C, Oprica M & Schultzberg M 2006 α-Melanocyte-stimulating hormone is neuroprotective in rat global cerebral ischemia. Neuropeptides 40 65-75. (doi:10.1016/j.npep.2005.10.006)
68. Forslin Aronsson A, Spulber S, Oprica M, Winblad B, Post C & Schultzberg M 2007 α-MSH rescues neurons from excitotoxic cell death. Journal of Molecular Neuroscience 33 239-251. (doi:10.1007/s12031-007-0019-2)
69. Fuente-Martin E, Garcia-Caceres C, Granado M, de Ceballos ML, Sanchez-GarridoMA, Sarman B, Liu ZW, Dietrich MO, Tena-Sempere M, Argente-Arizon P et al. 2012 Leptin regulates glutamate and glucose transporters in hypothalamic astrocytes. Journal of Clinical Investigation 122 3900-3913. (doi:10.1172/JCI64102)
70. Gandhi G.K., Cruz NF, Ball KK & Dienel GA 2009 Astrocytes are poised for lactate trafficking and release from activated brain and for supply of glucose to neurons. Journal of Neurochemistry 111 522-536. (doi:10.1111/j.1471-4159.2009.06333.x)
71. Gantz I., Konda Y, Tashiro T, Shimoto Y, Miwa H, Munzert G, Watson SJ, DelValle J & Yamada T 1993a Molecular cloning of a novel melanocortin receptor. Journal of Biological Chemistry 268 8246-8250.
72. Gantz I., Miwa H, Konda Y, Shimoto Y, Tashiro T, Watson SJ, DelValle J & Yamada T 1993b Molecular cloning, expression, and gene localization of a fourth melanocortin receptor. Journal of Biological Chemistry 268 15174-15179.
73. Gantz I., Shimoto Y, Konda Y, Miwa H, Dickinson CJ & Yamada T 1994 Molecular cloning, expression, and characterization of a fifth melanocortin receptor. Biochemical and Biophysical Research Communications 200 1214-1220. (doi:10.1006/bbrc.1994.1580)
74. Garcia-Caceres C, Fuente-Martin E, Burgos-Ramos E, Granado M, Frago LM, Barrios V., Horvath T, Argente J & Chowen JA 2011 Differential acute and chronic effects of leptin on hypothalamic astrocyte morphology and synaptic protein levels. Endocrinology 152 1809-1818. (doi:10.1210/en.2010-1252)
75. Garcia-Caceres C, Yi CX & Tschop MH 2013 Hypothalamic astrocytes in obesity. Endocrinology and Metabolism Clinics of North America 42 57-66. (doi:10.1016/j.ecl.2012.11.003)
76. Getting S.J., Lam CW, Chen AS, Grieco P & Perretti M 2006 Melanocortin 3 receptors control crystal-induced inflammation. FASEB Journal 20 2234-2241. (doi:10.1096/fj.06-6339com)
77. Giuliani D., Mioni C, Altavilla D, Leone S, Bazzani C, Minutoli L, Bitto A, Cainazzo M.M., Marini H, Zaffe D et al. 2006 Both early and delayed treatment with melanocortin 4 receptor-stimulating melanocortins produces neuroprotection in cerebral ischemia. Endocrinology 147 1126-1135. (doi:10.1210/en.2005-0692)
78. Giuliani D., Ottani A, Mioni C, Bazzani C, GalantucciM, Minutoli L, Bitto A, Zaffe D., Botticelli AR, Squadrito F et al. 2007 Neuroprotection in focal cerebral ischemia owing to delayed treatment with melanocortins. European Journal of Pharmacology 570 57-65. (doi:10.1016/j.ejphar. 2007.05.025)
79. Giuliani D., Ottani A, Minutoli L, Stefano VD, GalantucciM, Bitto A, ZaffeD, Altavilla D., Botticelli AR, Squadrito F et al. 2009 Functional recovery after delayed treatment of ischemic stroke with melanocortins is associated with overexpression of the activity-dependent gene Zif268. Brain, Behavior, and Immunity 23 844-850. (doi:10.1016/j.bbi.2009. 03.009)
80. Giuliani D., Zaffe D, Ottani A, Spaccapelo L, Galantucci M, Minutoli L, Bitto A., Irrera N, Contri M, Altavilla D et al. 2011 Treatment of cerebral ischemia with melanocortins acting at MC4 receptors induces marked neurogenesis and long-lasting functional recovery. Acta Neuropathologica 122 443-453. (doi:10.1007/s00401-011-0873-4)
81. Gonzalez P.V., Schioth HB, Lasaga M & Scimonelli TN 2009 Memory impairment induced by IL-1β is reversed by α-MSH through central melanocortin-4 receptors. Brain, Behavior, and Immunity 23 817-822. (doi:10.1016/j.bbi.2009.03.001)
82. Gonzalez P., Machado I, Vilcaes A, Roth G, Lasaga M & Scimonelli T 2013 The impairment on fear memory consolidation induced by IL-1β involves MAPK p38 phosphorylation and reduction of ERK activation. In International Society of Neurochemistry 24th Biennial Joint Meeting. Ed Abstract. Mexico: Cancun.
83. Gordon G.R., Baimoukhametova DV, Hewitt SA, Rajapaksha WR, Fisher TE & Bains JS 2005 Norepinephrine triggers release of glial ATP to increase postsynaptic efficacy. Nature Neuroscience 8 1078-1086. (doi:10.1038/ nn1498)
84. Gregor MF & Hotamisligil GS 2011 Inflammatory mechanisms in obesity. Annual Review of Immunology 29 415-445. (doi:10.1146/annurevimmunol-031210-101322)
85. Griffon N., Mignon V, Facchinetti P, Diaz J, Schwartz JC & Sokoloff P 1994 Molecular cloning and characterization of the rat fifth melanocortin receptor. Biochemical and Biophysical Research Communications 200 1007-1014. (doi:10.1006/bbrc.1994.1550)
86. Grosche J., Matyash V, Moller T, Verkhratsky A, Reichenbach A & Kettenmann H 1999 Microdomains for neuron-glia interaction: parallel fiber signaling to Bergmann glial cells. Nature Neuroscience 2 139-143. (doi:10.1038/5692)
87. Grosche J., Kettenmann H & Reichenbach A 2002 Bergmann glial cells form distinct morphological structures to interact with cerebellar neurons. Journal of Neuroscience Research 68 138-149. (doi:10.1002/jnr.10197)
88. Guarini S., Schioth HB, Mioni C, Cainazzo M, Ferrazza G, Giuliani D, Wikberg J.E., Bertolini A & Bazzani C 2002 MC(3) receptors are involved in the protective effect of melanocortins in myocardial ischemia/ reperfusion-induced arrhythmias. Naunyn-Schmiedeberg's Archives of Pharmacology 366 177-182. (doi:10.1007/s00210-002-0572-8)
89. Guillod-Maximin E, Roy AF, Vacher CM, Aubourg A, Bailleux V, Lorsignol A, Penicaud L., Parquet M & Taouis M 2009 Adiponectin receptors are expressed in hypothalamus and colocalized with proopiomelanocortin and neuropeptide Y in rodent arcuate neurons. Journal of Endocrinology 200 93-105. (doi:10.1677/JOE-08-0348)
90. Gupta S., Knight AG, Keller JN & Bruce-Keller AJ 2012 Saturated long-chain fatty acids activate inflammatory signaling in astrocytes. Journal of Neurochemistry 120 1060-1071. (doi:10.1111/j.1471-4159.2012. 07660.x)
91. Hanada R., Leibbrandt A, Hanada T, Kitaoka S, Furuyashiki T, Fujihara H, Trichereau J., Paolino M, Qadri F, Plehm R et al. 2009 Central control of fever and female body temperature by RANKL/RANK. Nature 462 505-509. (doi:10.1038/nature08596)
92. Heneka MT & Landreth GE 2007 PPARs in the brain. Biochimica et Biophysica Acta 1771 1031-1045. (doi:10.1016/j.bbalip.2007.04.016)
93. Henneberger C., Papouin T, Oliet SH & Rusakov DA 2010 Long-term potentiation depends on release of D-serine from astrocytes. Nature 463 232-236. (doi:10.1038/nature08673)
94. Hinney A., Bettecken T, Tarnow P, Brumm H, Reichwald K, Lichtner P, Scherag A., Nguyen TT, Schlumberger P, Rief W et al. 2006 Prevalence, spectrum, and functional characterization of melanocortin-4 receptor gene mutations in a representative population-based sample and obese adults from Germany. Journal of Clinical Endocrinology and Metabolism 91 1761-1769. (doi:10.1210/jc.2005-2056)
95. Ho G & MacKenzie RG 1999 Functional characterization of mutations in melanocortin-4 receptor associated with human obesity. Journal of Biological Chemistry 274 35816-35822. (doi:10.1074/jbc.274.50.35816)
96. Horvath T.L., Sarman B, Garcia-Caceres C, Enriori PJ, Sotonyi P, Shanabrough M., Borok E, Argente J, Chowen JA, Perez-Tilve D et al. 2010 Synaptic input organization of the melanocortin system predicts diet-induced hypothalamic reactive gliosis and obesity. PNAS 107 14875-14880. (doi:10.1073/pnas.1004282107)
97. Hsuchou H., He Y, Kastin AJ, Tu H, Markadakis EN, Rogers RC, Fossier PB & Pan W 2009 Obesity induces functional astrocytic leptin receptors in hypothalamus. Brain 132 889-902. (doi:10.1093/brain/awp029)
98. Huang Q & Tatro JB 2002 α-Melanocyte stimulating hormone suppresses intracerebral tumor necrosis factor-α and interleukin-1β gene expression following transient cerebral ischemia in mice. Neuroscience Letters 334 186-190. (doi:10.1016/S0304-3940(02)01088-1)
99. Huang Q.H., Entwistle ML, Alvaro JD, Duman RS, Hruby VJ & Tatro JB 1997 Antipyretic role of endogenous melanocortins mediated by central melanocortin receptors during endotoxin-induced fever. Journal of Neuroscience 17 3343-3351.
100. Huang Q.H., Hruby VJ & Tatro JB 1998 Systemic α-MSH suppresses LPS fever via central melanocortin receptors independently of its suppression of corticosterone and IL-6 release. American Journal of Physiology 275 R524-R530.
101. Huszar D., Lynch CA, Fairchild-Huntress V, Dunmore JH, Fang Q, Berkemeier L.R., Gu W, Kesterson RA, Boston BA, Cone RD et al. 1997 Targeted disruption of the melanocortin-4 receptor results in obesity in mice. Cell 88 131-141. (doi:10.1016/S0092-8674(00)81865-6)
102. Ichiyama T., Sakai T, Catania A, Barsh GS, Furukawa S & Lipton JM 1999a Systemically administered α-melanocyte-stimulating peptides inhibit NF-kappaB activation in experimental brain inflammation. Brain Research 836 31-37. (doi:10.1016/S0006-8993(99)01584-X)
103. Ichiyama T., Zhao H, Catania A, Furukawa S & Lipton JM 1999b α-Melanocyte-stimulating hormone inhibits NF-kappaB activation and IkappaBα degradation in human glioma cells and in experimental brain inflammation. Experimental Neurology 157 359-365. (doi:10.1006/exnr. 1999.7064)
104. Isales C.M., ZaidiM& Blair HC 2010 ACTH is a novel regulator of bone mass. Annals of the New York Academy of Sciences 1192 110-116. (doi:10.1111/ j.1749-6632.2009.05231.x)
105. Israel D.D., Sheffer-Babila S, de Luca C, Jo YH, Liu SM, Xia Q, Spergel DJ, Dun S.L., Dun NJ & Chua SC Jr 2012 Effects of leptin and melanocortin signaling interactions on pubertal development and reproduction. Endocrinology 153 2408-2419. (doi:10.1210/en.2011-1822)
106. Iwasaki A & Medzhitov R 2004 Toll-like receptor control of the adaptive immune responses. Nature Immunology 5 987-995. (doi:10.1038/ ni1112)
107. Jana M & Pahan K 2012 Gemfibrozil, a lipid lowering drug, inhibits the activation of primary human microglia via peroxisome proliferatoractivated receptor β. Neurochemical Research 37 1718-1729. (doi:10.1007/s11064-012-0781-6)
108. Jayaram B., PanW, Wang Y, Hsuchou H, Mace A, Cornelissen-Guillaume GG, Mishra P.K., Koza RA & Kastin AJ 2013 Astrocytic leptin-receptor knockout mice show partial rescue of leptin resistance in diet-induced obesity. Journal of Applied Physiology 114 734-741. (doi:10.1152/ japplphysiol.01499.2012)
109. Joosten E.A., Majewska B, Houweling DA, Bar PR & Gispen WH 1999 α-Melanocyte stimulating hormone promotes regrowth of injured axons in the adult rat spinal cord. Journal of Neurotrauma 16 543-553. (doi:10.1089/neu.1999.16.543)
110. Joppa M.A., Gogas KR, Foster AC & Markison S 2007 Central infusion of the melanocortin receptor antagonist agouti-related peptide (AgRP (83-132) prevents cachexia-related symptoms induced by radiation and colon-26 tumors in mice. Peptides 28 636-642. (doi:10.1016/ j.peptides.2006.11.021)
111. Kang J., Jiang L, Goldman SA & Nedergaard M 1998 Astrocyte-mediated potentiation of inhibitory synaptic transmission. Nature Neuroscience 1 683-692. (doi:10.1038/3684)
112. Kask A., Mutulis F, Muceniece R, Pahkla R, Mutule I, Wikberg JE, Rago L & Schioth HB 1998 Discovery of a novel superpotent and selective melanocortin-4 receptor antagonist (HS024): evaluation in vitro and in vivo. Endocrinology 139 5006-5014. (doi:10.1210/en.139.12.5006)
113. 2 in rat astrocyte cultures: modulatory effect of neuropeptides. Endocrinology 124 3125-3127. (doi:10.1210/ endo-124-6-3125)
114. Kievit P., Halem H, Marks DL, Dong JZ, Glavas MM, Sinnayah P, Pranger L, Cowley M.A., Grove KL & Culler MD 2013 Chronic treatment with a melanocortin-4 receptor agonist causes weight loss, reduces insulin resistance, and improves cardiovascular function in diet-induced obese rhesus macaques. Diabetes 62 490-497. (doi:10.2337/db12-0598)
115. Kim M.S., Cheong YP, So HS, Lee KM, Kim TY, Oh J, Chung YT, Son Y, Kim BR & Park R 2001 Protective effects of morphine in peroxynitriteinduced apoptosis of primary rat neonatal astrocytes: potential involvement of G protein and phosphatidylinositol 3-kinase (PI3 kinase). Biochemical Pharmacology 61 779-786. (doi:10.1016/ S0006-2952(01)00541-X)
116. Kofuji P & Newman EA 2004 Potassium buffering in the central nervous system. Neuroscience 129 1045-1056. (doi:10.1016/j.neuroscience. 2004.06.008)
117. Kokubo M., Asai K, Yamamoto N, Aoyama M, Morikawa M, Togari H, Wada Y & Kato T 2002 ACTH(1-24) down-regulates expression of ciliary neurotrophic factor mRNA in cultured rat astrocyte. Pediatric Research 52 950-957. (doi:10.1203/00006450-200212000-00022)
118. Krum J.M., Phillips TM & Rosenstein JM 2002 Changes in astroglial GLT-1 expression after neural transplantation or stab wounds. Experimental Neurology 174 137-149. (doi:10.1006/exnr.2002.7867)
119. Labbe O., Desarnaud F, Eggerickx D, Vassart G & Parmentier M 1994 Molecular cloning of a mouse melanocortin 5 receptor gene widely expressed in peripheral tissues. Biochemistry 33 4543-4549. (doi:10.1021/bi00181a015)
120. Lasaga M., Debeljuk L, Durand D, Scimonelli TN & Caruso C 2008 Role of α-melanocyte stimulating hormone and melanocortin 4 receptor in brain inflammation. Peptides 29 1825-1835. (doi:10.1016/j.peptides. 2008.06.009)
121. Laye S., Parnet P, Goujon E & Dantzer R 1994 Peripheral administration of lipopolysaccharide induces the expression of cytokine transcripts in the brain and pituitary of mice. Brain Research. Molecular Brain Research 27 157-162. (doi:10.1016/0169-328X(94)90197-X)
122. Li Q & Verma IM 2002 NF-k kappaB regulation in the immune system. Nature Reviews. Immunology 2 725-734. (doi:10.1038/nri910)
123. Li Y., Chopp M, Jiang N, Yao F & Zaloga C 1995 Temporal profile of in situ DNA fragmentation after transient middle cerebral artery occlusion in the rat. Journal of Cerebral Blood Flow and Metabolism 15 389-397. (doi:10.1038/jcbfm.1995.49)
124. Liedtke W., Edelmann W, Chiu FC, Kucherlapati R & Raine CS 1998 Experimental autoimmune encephalomyelitis in mice lacking glial fibrillary acidic protein is characterized by a more severe clinical course and an infiltrative central nervous system lesion. American Journal of Pathology 152 251-259.
125. Lodge PA & Sriram S 1996 Regulation of microglial activation by TGF-β, IL-10, and CSF-1. Journal of Leukocyte Biology 60 502-508.
126. Lonze BE & Ginty DD 2002 Function and regulation of CREB family transcription factors in the nervous system. Neuron 35 605-623. (doi:10.1016/S0896-6273(02)00828-0)
127. Lovett-Racke AE, Hussain RZ, Northrop S, Choy J, Rocchini A, Matthes L, Chavis J.A., Diab A, Drew PD & Racke MK 2004 Peroxisome proliferatoractivated receptor α agonists as therapy for autoimmune disease. Journal of Immunology 172 5790-5798.
128. Luo J., Lang JA & Miller MW 1998 Transforming growth factor β 1 regulates the expression of cyclooxygenase in cultured cortical astrocytes and neurons. Journal of Neurochemistry 71 526-534. (doi:10.1046/ j.1471-4159.1998.71020526.x)
129. Luth H.J., Munch G & Arendt T 2002 Aberrant expression of NOS isoforms in Alzheimer's disease is structurally related to nitrotyrosine formation. Brain Research 953 135-143. (doi:10.1016/S0006-8993(02)03280-8)
130. Machado I., Gonzalez P, Schioth HB, Lasaga M & Scimonelli TN 2010 α-Melanocyte-stimulating hormone (α-MSH) reverses impairment of memory reconsolidation induced by interleukin-1 β (IL-1β) hippocampal infusions. Peptides 31 2141-2144. (doi:10.1016/j.peptides. 2010.07.018)
131. Manna SK & Aggarwal BB 1998 α-Melanocyte-stimulating hormone inhibits the nuclear transcription factor NF-kappa B activation induced by various inflammatory agents. Journal of Immunology 161 2873-2880.
132. Maragakis NJ & Rothstein JD 2006 Mechanisms of disease: astrocytes in neurodegenerative disease. Nature Clinical Practice. Neurology 2 679-689. (doi:10.1038/ncpneuro0355)
133. Marsh D.J., Hollopeter G, Huszar D, Laufer R, Yagaloff KA, Fisher SL, Burn P & Palmiter RD 1999 Response of melanocortin-4 receptor-deficient mice to anorectic and orexigenic peptides. Nature Genetics 21 119-122. (doi:10.1038/5070)
134. Martin L.W., Catania A, Hiltz ME & Lipton JM 1991 Neuropeptide α-MSH antagonizes IL-6-and TNF-induced fever. Peptides 12 297-299. (doi:10.1016/0196-9781(91)90015-H)
135. Marz P., Heese K, Dimitriades-Schmutz B, Rose-John S & Otten U 1999 Role of interleukin-6 and soluble IL-6 receptor in region-specific induction of astrocytic differentiation and neurotrophin expression. Glia 26 191-200. (doi:10.1002/(SICI)1098-1136(199905)26:3!191::AID-GLIA1O3.0.CO;2-)
136. Minokoshi Y., Alquier T, Furukawa N, Kim YB, Lee A, Xue B, Mu J, Foufelle F, Ferre P., Birnbaum MJ et al. 2004 AMP-kinase regulates food intake by responding to hormonal and nutrient signals in the hypothalamus. Nature 428 569-574. (doi:10.1038/nature02440)
137. 2 release from rat hypothalamus is mediated by type I and type II interleukin-1 receptors. Journal of Neuroimmunology 61 171-177. (doi:10.1016/0165-5728(95)00088-J)
138. Mountjoy K.G., Robbins LS, Mortrud MT & Cone RD 1992 The cloning of a family of genes that encode the melanocortin receptors. Science 257 1248-1251. (doi:10.1126/science.1325670)
139. Mountjoy K.G., Mortrud MT, Low MJ, Simerly RB & Cone RD 1994 Localization of the melanocortin-4 receptor (MC4-R) in neuroendocrine and autonomic control circuits in the brain. Molecular Endocrinology 8 1298-1308. (doi:10.1210/me.8.10.1298)
140. Mountjoy K.G., Kong PL, Taylor JA, Willard DH & Wilkison WO 2001 Melanocortin receptor-mediated mobilization of intracellular free calcium in HEK293 cells. Physiological Genomics 5 11-19.
141. Mountjoy K.G., Jenny Wu CS, Dumont LM &Wild JM 2003 Melanocortin-4 receptormessenger ribonucleic acid expression in rat cardiorespiratory, musculoskeletal, and integumentary systems. Endocrinology 144 5488-5496. (doi:10.1210/en.2003-0570)
142. Muceniece R., Zvejniece L, Kirjanova O, Liepinsh E, Krigere L, Baumane L, Kalvinsh I., Wikberg JE & Dambrova M 2004 Beta-and gammamelanocortins inhibit lipopolysaccharide induced nitric oxide production in mice brain. Brain Research 995 7-13. (doi:10.1016/ j.brainres.2003.09.039)
143. Muceniece R., Zvejniece L, Vilskersts R, Liepinsh E, Baumane L, Kalvinsh I, Wikberg JE & Dambrova M 2007 Functional evaluation of THIQ, a melanocortin 4 receptor agonist, in models of food intake and inflammation. Basic & Clinical Pharmacology & Toxicology 101 416-420. (doi:10.1111/j.1742-7843.2007.00133.x)
144. Nagahara AH & Tuszynski MH 2011 Potential therapeutic uses of BDNF in neurological and psychiatric disorders. Nature Reviews. Drug Discovery 10 209-219. (doi:10.1038/nrd3366)
145. Obara M., Szeliga M & Albrecht J 2008 Regulation of pH in the mammalian central nervous system under normal and pathological conditions: facts and hypotheses. Neurochemistry International 52 905-919. (doi:10.1016/j.neuint.2007.10.015)
146. Oberheim N.A., Takano T, Han X, He W, Lin JH, Wang F, Xu Q, Wyatt JD, Pilcher W., Ojemann JG et al. 2009 Uniquely hominid features of adult human astrocytes. Journal of Neuroscience 29 3276-3287. (doi:10.1523/ JNEUROSCI.4707-08.2009)
147. O'Donohue TL & Dorsa DM 1982 The opiomelanotropinergic neuronal and endocrine systems. Peptides 3 353-395. (doi:10.1016/0196-9781 (82)90098-5)
148. Ozog M.A., Siushansian R & Naus CC 2002 Blocked gap junctional coupling increases glutamate-induced neurotoxicity in neuron-astrocyte co-cultures. Journal of Neuropathology and Experimental Neurology 61 132-141.
149. Parpura V & Zorec R 2010 Gliotransmission: exocytotic release from astrocytes. Brain Research Reviews 63 83-92. (doi:10.1016/ j.brainresrev.2009.11.008)
150. Patil S., Sheng L, Masserang A & Chan C 2006 Palmitic acid-treated astrocytes induce BACE1 upregulation and accumulation of C-terminal fragment of APP in primary cortical neurons. Neuroscience Letters 406 55-59. (doi:10.1016/j.neulet.2006.07.015)
151. Patten C.S., Daniels D, Suzuki A, Fluharty SJ & Yee DK 2007 Structural and signaling requirements of the human melanocortin 4 receptor for MAP kinase activation. Regulatory Peptides 142 111-122. (doi:10.1016/ j.regpep.2007.02.005)
152. Pellerin L., Bouzier-Sore AK, Aubert A, Serres S, Merle M, Costalat R & Magistretti PJ 2007 Activity-dependent regulation of energy metabolism by astrocytes: an update. Glia 55 1251-1262. (doi:10.1002/ glia.20528)
153. Perez-Oliva AB, Olivares C, Jimenez-Cervantes C & Garcia-Borron JC 2009 Mahogunin ring finger-1 (MGRN1) E3 ubiquitin ligase inhibits signaling from melanocortin receptor by competition with Galphas. Journal of Biological Chemistry 284 31714-31725. (doi:10.1074/ jbc.M109.028100)
154. Polak P.E., Kalinin S, Dello Russo C, Gavrilyuk V, Sharp A, Peters JM, Richardson J., Willson TM, Weinberg G & Feinstein DL 2005 Protective effects of a peroxisome proliferator-activated receptor-β/delta agonist in experimental autoimmune encephalomyelitis. Journal of Neuroimmunology 168 65-75. (doi:10.1016/j.jneuroim.2005.07.006)
155. Qian L., Wei SJ, ZhangD, HuX, XuZ, WilsonB, El-Benna J, Hong JS&FloodPM 2008 Potent anti-inflammatory and neuroprotective effects of TGF-β1 aremediated through the inhibition of ERK and p47phox-Ser345 phosphorylation and translocation inmicroglia. Journal of Immunology 181 660-668.
156. Rajora N., Boccoli G, Burns D, Sharma S, Catania AP & Lipton JM 1997 α-MSH modulates local and circulating tumor necrosis factor-α in experimental brain inflammation. Journal of Neuroscience 17 2181-2186.
157. Reizes O., Benoit SC, Strader AD, Clegg DJ, Akunuru S & Seeley RJ 2003 Syndecan-3 modulates food intake by interacting with the melanocortin/ AgRP pathway. Annals of the New York Academy of Sciences 994 66-73. (doi:10.1111/j.1749-6632.2003.tb03163.x)
158. Roselli-Rehfuss L, Mountjoy KG, Robbins LS, Mortrud MT, Low MJ, Tatro JB, Entwistle M.L., Simerly RB & Cone RD 1993 Identification of a receptor for gamma melanotropin and other proopiomelanocortin peptides in the hypothalamus and limbic system. PNAS 90 8856-8860. (doi:10.1073/pnas.90.19.8856)
159. Rossi M., KimMS, Morgan DG, Small CJ, Edwards CM, Sunter D, Abusnana S, Goldstone A.P., Russell SH, Stanley SA et al. 1998 A C-terminal fragment of Agouti-related protein increases feeding and antagonizes the effect of α-melanocyte stimulating hormone in vivo. Endocrinology 139 4428-4431. (doi:10.1210/en.139.10.4428)
160. Rouach N., Koulakoff A, Abudara V, Willecke K & Giaume C 2008 Astroglial metabolic networks sustain hippocampal synaptic transmission. Science 322 1551-1555. (doi:10.1126/science.1164022)
161. Rudge J.S., Pasnikowski EM, Holst P & Lindsay RM 1995 Changes in neurotrophic factor expression and receptor activation following exposure of hippocampal neuron/astrocyte cocultures to kainic acid. Journal of Neuroscience 15 6856-6867.
162. Saas P., Boucraut J, Quiquerez AL, Schnuriger V, Perrin G, Desplat-Jego S, Bernard D., Walker PR & Dietrich PY 1999 CD95(Fas/Apo-1) as a receptor governing astrocyte apoptotic or inflammatory responses: a key role in brain inflammation? Journal of Immunology 162 2326-2333.
163. Sarkar S., Legradi G & Lechan RM 2002 Intracerebroventricular administration of α-melanocyte stimulating hormone increases phosphorylation of CREB in TRH-and CRH-producing neurons of the hypothalamic paraventricular nucleus. Brain Research 945 50-59. (doi:10.1016/S0006-8993(02)02619-7)
164. Sarkar A., Sreenivasan Y & Manna SK 2003 α-Melanocyte-stimulating hormone inhibits lipopolysaccharide-induced biological responses by downregulating CD14 from macrophages. FEBS Letters 553 286-294. (doi:10.1016/S0014-5793(03)01029-9)
165. Sawyer T.K., Sanfilippo PJ, Hruby VJ, Engel MH, Heward CB, Burnett JB & Hadley ME 1980 4-Norleucine, 7-D-phenylalanine-α-melanocytestimulating hormone: a highly potent α-melanotropin with ultralong biological activity. PNAS 77 5754-5758. (doi:10.1073/pnas.77.10.5754)
166. Scarlett J.M., Bowe DD, Zhu X, Batra AK, Grant WF &Marks DL 2010 Genetic and pharmacologic blockade of central melanocortin signaling attenuates cardiac cachexia in rodent models of heart failure. Journal of Endocrinology 206 121-130. (doi:10.1677/JOE-09-0397)
167. Schioth HB & Watanobe H 2002 Melanocortins and reproduction. Brain Research. Brain Research Reviews 38 340-350. (doi:10.1016/ S0165-0173(01)00159-X)
168. Schioth H.B., Chhajlani V, Muceniece R, Klusa V & Wikberg JE 1996 Major pharmacological distinction of the ACTH receptor from other melanocortin receptors. Life Sciences 59 797-801. (doi:10.1016/ 0024-3205(96)00370-0)
169. Schioth H.B., Muceniece R, Mutulis F, Bouifrouri AA, Mutule I &Wikberg JE 1999 Further pharmacological characterization of the selective melanocortin 4 receptor antagonist HS014: comparison with SHU9119. Neuropeptides 33 191-196. (doi:10.1054/npep.1999.0760)
170. Schioth H.B., Kakizaki Y, Kohsaka A, Suda T & Watanobe H 2001 Agoutirelated peptide prevents steroid-induced luteinizing hormone and prolactin surges in female rats. Neuroreport 12 687-690. (doi:10.1097/ 00001756-200103260-00014)
171. Schwartz MW 2006 Central nervous system regulation of food intake. Obesity 14 (Suppl 1) 1S-8S. (doi:10.1038/oby.2006.275)
172. Schwartz JP & Nishiyama N 1994 Neurotrophic factor gene expression in astrocytes during development and following injury. Brain Research Bulletin 35 403-407. (doi:10.1016/0361-9230(94)90151-1)
173. Selkirk J.V., Nottebaum LM, Lee J, Yang W, Foster AC & Lechner SM 2007 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 52 459-466. (doi:10.1016/j.neuropharm.2006.08.015)
174. Shadrina M.I., Dolotov OV, Grivennikov IA, Slominsky PA, Andreeva LA, Inozemtseva L.S., Limborska SA & Myasoedov NF 2001 Rapid induction of neurotrophin mRNAs in rat glial cell cultures by Semax, an adrenocorticotropic hormone analog. Neuroscience Letters 308 115-118. (doi:10.1016/S0304-3940(01)01994-2)
175. Sharma H.S., Skottner A, Lundstedt T, Flardh M & Wiklund L 2006 Neuroprotective effects of melanocortins in experimental spinal cord injury. An experimental study in the rat using topical application of compounds with varying affinity to melanocortin receptors. Journal of Neural Transmission 113 463-476. (doi:10.1007/s00702-005-0404-3)
176. Shen Y., Fu WY, Cheng EY, Fu AK & Ip NY 2013 Melanocortin-4 receptor regulates hippocampal synaptic plasticity through a protein kinase A-dependent mechanism. Journal of Neuroscience 33 464-472. (doi:10.1523/JNEUROSCI.3282-12.2013)
177. Sinha P.S., Schioth HB & Tatro JB 2003 Activation of central melanocortin-4 receptor suppresses lipopolysaccharide-induced fever in rats. American Journal of Physiology. Regulatory, Integrative and Comparative Physiology 284 R1595-R1603. (doi:10.1152/ajpregu.00581.2002)
178. Sinha P.S., Schioth HB & Tatro JB 2004 Roles of the melanocortin-4 receptor in antipyretic and hyperthermic actions of centrally administered α-MSH. Brain Research 1001 150-158. (doi:10.1016/j.brainres. 2003.12.007)
179. Slominski A., Ermak G & Mihm M 1996 ACTH receptor, CYP11A1, CYP17 and CYP21A2 genes are expressed in skin. Journal of Clinical Endocrinology and Metabolism 81 2746-2749. (doi:10.1210/jc.81.7.2746)
180. Spencer JD & Schallreuter KU 2009 Regulation of pigmentation in human epidermal melanocytes by functional high-affinity β-melanocytestimulating hormone/melanocortin-4 receptor signaling. Endocrinology 150 1250-1258. (doi:10.1210/en.2008-1212)
181. Stalder A.K., CarsonMJ, Pagenstecher A, Asensio VC, Kincaid C, Benedict M, Powell H.C., Masliah E & Campbell IL 1998 Late-onset chronic inflammatory encephalopathy in immune-competent and severe combined immune-deficient (SCID) mice with astrocyte-targeted expression of tumor necrosis factor. American Journal of Pathology 153 767-783. (doi:10.1016/S0002-9440(10)65620-9)
182. Stanley S.A., Small CJ, Kim MS, Heath MM, Seal LJ, Russell SH, Ghatei MA & Bloom SR 1999 Agouti related peptide (Agrp) stimulates the hypothalamo pituitary gonadal axis in vivo & in vitro in male rats. Endocrinology 140 5459-5462. (doi:10.1210/en.140.11.5459)
183. Starowicz K., Przewlocki R, Gispen WH & Przewlocka B 2002 Modulation of melanocortin-induced changes in spinal nociception by mu-opioid receptor agonist and antagonist in neuropathic rats. Neuroreport 13 2447-2452. (doi:10.1097/00001756-200212200-00015)
184. Storer P.D., Xu J, Chavis J & Drew PD 2005 Peroxisome proliferator-activated receptor-gamma agonists inhibit the activation of microglia and astrocytes: implications for multiple sclerosis. Journal of Neuroimmunology 161 113-122. (doi:10.1016/j.jneuroim.2004.12.015)
185. Strand F.L., Rose KJ, Zuccarelli LA, Kume J, Alves SE, Antonawich FJ & Garrett LY 1991 Neuropeptide hormones as neurotrophic factors. Physiological Reviews 71 1017-1046.
186. Suk K., Lee J, Hur J, Kim YS, Lee M, Cha S, Yeou Kim S & Kim H 2001 Activation-induced cell death of rat astrocytes. Brain Research 900 342-347. (doi:10.1016/S0006-8993(01)02326-5)
187. Sutton G.M., Duos B, Patterson LM & Berthoud HR 2005 Melanocortinergic modulation of cholecystokinin-induced suppression of feeding through extracellular signal-regulated kinase signaling in rat solitary nucleus. Endocrinology 146 3739-3747. (doi:10.1210/en.2005-0562)
188. Takuma K., Baba A & Matsuda T 2004 Astrocyte apoptosis: implications for neuroprotection. Progress in Neurobiology 72 111-127. (doi:10.1016/ j.pneurobio.2004.02.001)
189. Tanabe K., Gamo K, Aoki S, Wada K & Kiyama H 2007 Melanocortin receptor 4 is induced in nerve-injured motor and sensory neurons of mouse. Journal of Neurochemistry 101 1145-1152. (doi:10.1111/j.1471-4159.2006.04432.x)
190. Tao YX 2010 The melanocortin-4 receptor: physiology, pharmacology, and pathophysiology. Endocrine Reviews 31 506-543. (doi:10.1210/ er.2009-0037)
191. Tatro JB 2000 Endogenous antipyretics. Clinical Infectious Diseases 31 (Suppl 5) S190-S201. (doi:10.1086/317519)
192. Tomassoni D., Nwankwo IE, Gabrielli MG, Bhatt S, Muhammad AB, Lokhandwala M.F., Tayebati SK & Amenta F 2013 Astrogliosis in the brain of obese Zucker rat: a model of metabolic syndrome. Neuroscience Letters 543 136-141. (doi:10.1016/j.neulet.2013.03.025)
193. Tran E.H., Hardin-Pouzet H, Verge G & Owens T 1997 Astrocytes and microglia express inducible nitric oxide synthase in mice with experimental allergic encephalomyelitis. Journal of Neuroimmunology 74 121-129. (doi:10.1016/S0165-5728(96)00215-9)
194. Usategui R., Oliver C, Vaudry H, Lombardi G, Rozenberg I & Mourre AM 1976 Immunoreactive α-MSH and ACTH levels in rat plasma and pituitary. Endocrinology 98 189-196. (doi:10.1210/endo-98-1-189)
195. Uysal K.T., Wiesbrock SM, Marino MW & Hotamisligil GS 1997 Protection from obesity-induced insulin resistance in mice lacking TNF-α function. Nature 389 610-614. (doi:10.1038/39335)
196. Van der Ploeg LH, Martin WJ, Howard AD, Nargund RP, Austin CP, Guan X, Drisko J., Cashen D, Sebhat I, Patchett AA et al. 2002 A role for the melanocortin 4 receptor in sexual function. PNAS 99 11381-11386. (doi:10.1073/pnas.172378699)
197. Ventura R & Harris KM 1999 Three-dimensional relationships between hippocampal synapses and astrocytes. Journal of Neuroscience 19 6897-6906.
198. Vongs A., Lynn NM & Rosenblum CI 2004 Activation of MAP kinase by MC4-R through PI3 kinase. Regulatory Peptides 120 113-118. (doi:10.1016/j.regpep.2004.02.018)
199. Vulliemoz N.R., Xiao E, Xia-Zhang L, Ferin M & Wardlaw SL 2006 Melanocortin modulation of inflammatory cytokine and neuroendocrine responses to endotoxin in the monkey. Endocrinology 147 1878-1883. (doi:10.1210/en.2005-1430)
200. Wang P., Wu P, Siegel MI, Egan RW & Billah MM 1995 Interleukin (IL)-10 inhibits nuclear factor kappa B (NF kappa B) activation in human monocytes, IL-10 and IL-4 suppress cytokine synthesis by different mechanisms. Journal of Biological Chemistry 270 9558-9563. (doi:10.1074/jbc.270.16.9558)
201. Watanobe H., Yoneda M, Kakizaki Y, Kohsaka A, Suda T & Schioth HB 2001 Further evidence for a significant participation of the melanocortin 4 receptor in the preovulatory prolactin surge in the rat. Brain Research Bulletin 54 521-525. (doi:10.1016/S0361-9230(01)00442-7)
202. Weidenfeld J., Crumeyrolle-Arias M & Haour F 1995 Effect of bacterial endotoxin and interleukin-1 on prostaglandin biosynthesis by the hippocampus of mouse brain: role of interleukin-1 receptors and glucocorticoids. Neuroendocrinology 62 39-46. (doi:10.1159/ 000126986)
203. 2 induces penile erection via brain and spinal melanocortin receptors. Neuroscience 118 755-762. (doi:10.1016/S0306-4522(02)00866-7)
204. Wilkinson CW 2006 Roles of acetylation and other post-translational modifications in melanocortin function and interactions with endorphins. Peptides 27 453-471. (doi:10.1016/j.peptides.2005.05.029)
205. Wilson JX 1997 Antioxidant defense of the brain: a role for astrocytes. Canadian Journal of Physiology and Pharmacology 75 1149-1163. (doi:10.1139/y97-146)
206. Wong K.Y., Rajora N, Boccoli G, Catania A & Lipton JM 1997 A potential mechanism of local anti-inflammatory action of α-melanocytestimulating hormone within the brain: modulation of tumor necrosis factor-α production by human astrocytic cells. Neuroimmunomodulation 4 37-41.
207. Xu B., Goulding EH, Zang K, Cepoi D, Cone RD, Jones KR, Tecott LH & Reichardt LF 2003 Brain-derived neurotrophic factor regulates energy balance downstream of melanocortin-4 receptor. Nature Neuroscience 6 736-742. (doi:10.1038/nn1073)
208. Yamaoka-Tojo M, Tojo T, Shioi T, Masuda T, Inomata T & Izumi T 2006 Central neurotranspeptide, α-melanocyte-stimulating hormone (α-MSH) is upregulated in patients with congestive heart failure. Internal Medicine 45 429-434. (doi:10.2169/internalmedicine.45.1546)
209. Yeo G.S., Farooqi IS, Aminian S, Halsall DJ, Stanhope RG & O'Rahilly S 1998 A frameshift mutation in MC4R associated with dominantly inherited human obesity. Nature Genetics 20 111-112. (doi:10.1038/2404)
210. Yi CX & Tschop MH 2012 Brain-gut-adipose-tissue communication pathways at a glance. Disease Models & Mechanisms 5 583-587. (doi:10.1242/dmm.009902)
211. Zhang X., Zhang G, Zhang H, Karin M, Bai H & Cai D 2008 Hypothalamic IKKβ/NF-kappaB and ER stress link overnutrition to energy imbalance and obesity. Cell 135 61-73. (doi:10.1016/j.cell.2008.07.043)
212. Zhang Y., Wu X, He Y, Kastin AJ, Hsuchou H, Rosenblum CI & Pan W 2009 Melanocortin potentiates leptin-induced STAT3 signaling via MAPK pathway. Journal of Neurochemistry 110 390-399. (doi:10.1111/j.1471-4159.2009.06144.x)
213. Zhu D.Y., Liu SH, Sun HS & Lu YM 2003 Expression of inducible nitric oxide synthase after focal cerebral ischemia stimulates neurogenesis in the adult rodent dentate gyrus. Journal of Neuroscience 23 223-229.
214. Zohar M & Salomon Y 1992 Melanocortins stimulate proliferation and induce morphological changes in cultured rat astrocytes by distinct transducing mechanisms. Brain Research 576 49-58. (doi:10.1016/0006-8993(92)90608-C)