The melanocortin system in leukocyte biology

Journal of Leukocyte Biology

Volumn 81, Issue 2, 2007, Pages 383-392

  Catania, Anna ^a  

^a FONDAZIONE IRCCS CA GRANDA OSPEDALE MAGGIORE POLICLINICO   (Italy)

Author keywords

melanocyte stimulating hormone; Adrenocorticotropic hormone; Melanocortin peptides; Melanocortin receptors; Neuroimmunomodulation; Nuclear factor B; Pro opiomelanocortin

Indexed keywords

AGOUTI RELATED PROTEIN; ALPHA INTERMEDIN; BETA INTERMEDIN; CORTICOTROPIN; GAMMA INTERMEDIN; IMMUNOGLOBULIN ENHANCER BINDING PROTEIN; MELANOCORTIN RECEPTOR;

ANTIINFLAMMATORY ACTIVITY; CELL ACTIVATION; CELL FUNCTION; DRUG MECHANISM; GENE TRANSLOCATION; HORMONE SYNTHESIS; IMMUNOMODULATION; LEUKOCYTE; PRIORITY JOURNAL; PROTEIN EXPRESSION; REVIEW; SIGNAL TRANSDUCTION;

ANIMALS; CELL NUCLEUS; HUMANS; LEUKOCYTES; MELANOCORTINS; MODELS, IMMUNOLOGICAL; NF-KAPPA B; RECEPTORS, MELANOCORTIN; SIGNAL TRANSDUCTION;

AGOUTI;

EID: 33846698560     PISSN: 07415400     EISSN: None     Source Type: Journal    
DOI: 10.1189/jlb.0706426     Document Type: Review

References (134)

1. Luger, T. A., Scholzen, T. E., Brzoska, T., Bohm, M. (2003) New insights into the functions of α-MSH and related peptides in the immune system. Ann. N. Y. Acad. Sci. 994, 133-140.
2. Cone, R. D. (2005) Anatomy and regulation of the central melanocortin system. Nat. Neurosci. 8, 571-578.
3. Eves, P. C., MacNeil, S., Haycock, J. W. (2006) α-Melanocyte stimulating hormone, inflammation and human melanoma. Peptides 27, 444-452.
4. 2 induces penile erection via brain and spinal melanocortin receptors. Neuroscience 118, 755-762.
5. Kadekaro, A. L., Wakamatsu, K., Ito, S., Abdel-Malek, Z. A. (2006) Cutaneous photoprotection and melanoma susceptibility: reaching beyond melanin content to the frontiers of DNA repair. Front. Biosci. 11, 2157-2173.
6. Tatro, J. B., Sinha, P. S. (2003) The central melanocortin system and fever. Ann. N. Y. Acad. Sci. 994, 246-257.
7. Gantz, I., Fong, T. M. (2003) The melanocortin system. Am. J. Physiol. Endocrinol. Metab. 284, E468-E474.
8. Ignar, D. M., Andrews, J. L., Jansen, M., Eilert, M. M., Pink, H. M., Lin, P., Sherrill, R. G., Szewczyk, J. R., Conway, J. G. (2003) Regulation of TNF-α secretion by a specific melanocortin-1 receptor peptide agonist. Peptides 24, 709-716.
9. Getting, S. J. (2006) Targeting melanocortin receptors as potential novel therapeutics. Pharmacol. Ther. 111, 1-15.
10. Catania, A., Gatti, S., Colombo, G., Lipton, J. M. (2004) Targeting melanocortin receptors as a novel strategy to control inflammation. Pharmacol. Rev. 56, 1-29.
11. Catania, A., Lipton, J. M. (1993) α-Melanocyte stimulating hormone in the modulation of host reactions. Endocr. Rev. 14, 564-576.
12. Eberle, A. (1988) The Melanotropins, Basel, Switzerland, Karger.
13. Seidah, N. G., Chretien, M. (1994) Pro-protein convertases of subtilisin/kexin family. Methods Enzymol. 244, 175-188.
14. Benjannet, S., Rondeau, N., Day, R., Chretien, M., Seidah, N. G. (1991) PC1 and PC2 are proprotein convertases capable of cleaving proopiomelanocortin at distinct pairs of basic residues. Proc. Natl. Acad. Sci. USA 88, 3564-3568.
15. Day, R., Schafer, M. K., Watson, S. J., Chretien, M., Seidah, N. G. (1992) Distribution and regulation of the prohormone convertases PC1 and PC2 in the rat pituitary. Mol. Endocrinol. 6, 485-497.
16. Marcinkiewicz, M., Day, R., Seidah, N. G., Chretien, M. (1993) Ontogeny of the prohormone convertases PC1 and PC2 in the mouse hypophysis and their colocalization with corticotropin and α-melanotropin. Proc. Natl. Acad. Sci. USA 90, 4922-4926.
17. Blalock, J. E. (1985) Pro-opiomelanocortin-derived peptides in the immune system. Clin. Endocrinol. (Oxf.) 22, 823-827.
18. Wakamatsu, K., Graham, A., Cook, D., Thody, A. J. (1997) Characterization of ACTH peptides in human skin and their activation of the melanocortin-1 receptor. Pigment Cell Res. 10, 288-297.
19. Sternberg, E. M. (2006) Neural regulation of innate immunity: a coordinated nonspecific host response to pathogens. Nat. Rev. Immunol. 6, 318-328.
20. Charmandari, E., Tsigos, C., Chrousos, G. (2005) Endocrinology of the stress response. Annu. Rev. Physiol. 67, 259-284.
21. Beuschlein, F., Hammer, G. D. (2002) Ectopic pro-opiomelanocortin syndrome. Endocrinol. Metab. Clin. North Am. 31, 191-234.
22. Blalock, J. E. (1999) Proopiomelanocortin and the immune-neuroendocrine connection. Ann. N. Y. Acad. Sci. 885, 161-172.
23. Chakraborty, A. K., Funasaka, Y., Slominski, A., Ermak, G., Hwang, J., Pawelek, J. M., Ichihashi, M. (1996) Production and release of proopiomelanocortin (POMC) derived peptides by human melanocytes and keratinocytes in culture: regulation by ultraviolet B. Biochim. Biophys. Acta 1313, 130-138.
24. Star, R. A., Rajora, N., Huang, J., Stock, R. C., Catania, A., Lipton, J. M. (1995) Evidence of autocrine modulation of macrophage nitric oxide synthase by α-melanocyte-stimulating hormone. Proc. Natl. Acad. Sci. USA 92, 8016-8020.
25. Slominski, A., Wortsman, J. (2000) Neuroendocrinology of the skin. Endocr. Rev. 21, 457-487.
26. Slominski, A., Wortsman, J., Luger, T., Paus, R., Solomon, S. (2000) Corticotropin releasing hormone and proopiomelanocortin involvement in the cutaneous response to stress. Physiol. Rev. 80, 979-1020.
27. Harbour, D. V., Smith, E. M., Blalock, J. E. (1987) Novel processing pathway for proopiomelanocortin in lymphocytes: endotoxin induction of a new prohormone-cleaving enzyme. J. Neurosci. Res. 18, 95-101.
28. Blalock, J. E., Smith, E. M. (1985) A complete regulatory loop between the immune and neuroendocrine systems. Fed. Proc. 44, 108-111.
29. Barcellini, W., Colombo, G., La Maestra, L., Clerici, G., Garofalo, L., Brini, A. T., Lipton, J. M., Catania, A. (2000) α-Melanocyte-stimulating hormone peptides inhibit HIV-1 expression in chronically infected promonocytic U1 cells and in acutely infected monocytes. J. Leukoc. Biol. 68, 693-699.
30. Rajora, N., Ceriani, G., Catania, A., Star, R. A., Murphy, M. T., Lipton, J. M. (1996) α-MSH production, receptors, and influence on neopterin in a human monocyte/macrophage cell line. J. Leukoc. Biol. 59, 248-253.
31. Ray, D. W., Ren, S. G., Melmed, S. (1996) Leukemia inhibitory factor (LIF) stimulates proopiomelanocortin (POMC) expression in a corticotroph cell line. Role of STAT pathway. J. Clin. Invest. 97, 1852-1859.
32. Wikberg, J. E., Muceniece, R., Mandrika, I., Prusis, P., Lindblom, J., Post, C., Skottner, A. (2000) New aspects on the melanocortins and their receptors. Pharmacol. Res. 42, 393-420.
33. Abdel-Malek, Z. A. (2001) Melanocortin receptors: their functions and regulation by physiological agonists and antagonists. Cell. Mol. Life Sci. 58, 434-441.
34. Chhajlani, V., Wikberg, J. E. (1992) Molecular cloning and expression of the human melanocyte stimulating hormone receptor cDNA. FEBS Lett. 309, 417-420.
35. Mountjoy, K. G., Robbins, L. S., Mortrud, M. T., Cone, R. D. (1992) The cloning of a family of genes that encode the melanocortin receptors. Science 257, 1248-1251.
36. Suzuki, I., Cone, R. D., Im, S., Nordlund, J., Abdel-Malek, Z. A. (1996) Binding of melanotropic hormones to the melanocortin receptor MC1R on human melanocytes stimulates proliferation and melanogenesis. Endocrinology 137, 1627-1633.
37. Catania, A., Rajora, N., Capsoni, F., Minonzio, F., Star, R. A., Lipton, J. M. (1996) The neuropeptide α-MSH has specific receptors on neutrophils and reduces chemotaxis in vitro. Peptides 17, 675-679.
38. Manna, S. K., Sarkar, A., Sreenivasan, Y. (2006) α-Melanocyte- stimulating hormone down-regulates CXC receptors through activation of neutrophil elastase. Eur. J. Immunol. 36, 754-769.
39. Bhardwaj, R., Becher, E., Mahnke, K., Hartmeyer, M., Schwarz, T., Scholzen, T., Luger, T. A. (1997) Evidence for the differential expression of the functional α-melanocyte-stimulating hormone receptor MC-1 on human monocytes. J. Immunol. 158, 3378-3384.
40. Cooper, A., Robinson, S. J., Pickard, C., Jackson, C. L., Friedmann, P. S., Healy, E. (2005) α-Melanocyte-stimulating hormone suppresses antigen-induced lymphocyte proliferation in humans independently of melanocortin 1 receptor gene status. J. Immunol. 175, 4806-4813.
41. Taherzadeh, S., Sharma, S., Chhajlani, V., Gantz, I., Rajora, N., Demitri, M. T., Kelly, L., Zhao, H., Ichiyama, T., Catania, A., Lipton, J. M. (1999) α-MSH and its receptors in regulation of tumor necrosis factor-α production by human monocyte/macrophages. Am. J. Physiol. 276, R1289-R1294.
42. Becher, E., Mahnke, K., Brzoska, T., Kalden, D. H., Grabbe, S., Luger, T. A. (1999) Human peripheral blood-derived dendritic cells express functional melanocortin receptor MC-1R. Ann. N. Y. Acad. Sci. 885, 188-195.
43. Hartmeyer, M., Scholzen, T., Becher, E., Bhardwaj, R. S., Schwarz, T., Luger, T. A. (1997) Human dermal microvascular endothelial cells express the melanocortin receptor type 1 and produce increased levels of IL-8 upon stimulation with α-melanocyte-stimulating hormone. J. Immunol. 159, 1930-1937.
44. Lam, C. W., Getting, S. J., Perretti, M. (2005) In vitro and in vivo induction of heme oxygenase 1 in mouse macrophages following melanocortin receptor activation. J. Immunol. 174, 2297-2304.
45. Schioth, H. B., Chhajlani, V., Muceniece, R., Klusa, V., Wikberg, J. E. (1996) Major pharmacological distinction of the ACTH receptor from other melanocortin receptors. Life Sci. 59, 797-801.
46. Roselli-Rehfuss, L., Mountjoy, K. G., Robbins, L. S., Mortrud, M. T., Low, M. J., Tatro, J. B., Entwistle, M. L., Simerly, R. B., Cone, R. D. (1993) Identification of a receptor for γ melanotropin and other proopiomelanocortin peptides in the hypothalamus and limbic system. Proc. Natl. Acad. Sci. USA 90, 8856-8860.
47. Cone, R. D. (1999) The central melanocortin system and energy homeostasis. Trends Endocrinol. Metab. 10, 211-216.
48. Lam, C. W., Perretti, M., Getting, S. J. (2006) Melanocortin receptor signaling in RAW264.7 macrophage cell line. Peptides 27, 404-412.
49. Mountjoy, K. G., Mortrud, M. T., Low, M. J., Simerly, R. B., Cone, R. D. (1994) Localization of the melanocortin-4 receptor (MC4-R) in neuroendocrine and autonomic control circuits in the brain. Mol. Endocrinol. 8, 1298-1308.
50. Gantz, I., Shimoto, Y., Konda, Y., Miwa, H., Dickinson, C. J., Yamada, T. (1994) Molecular cloning, expression, and characterization of a fifth melanocortin receptor. Biochem. Biophys. Res. Commun. 200, 1214-1220.
51. Van der Kraan, M., Adan, R. A., Entwistle, M. L., Gispen, W. H., Burbach, J. P., Tatro, J. B. (1998) Expression of melanocortin-5 receptor in secretory epithelia supports a functional role in exocrine and endocrine glands. Endocrinology 139, 2348-2355.
52. Chen, W., Kelly, M. A., Opitz-Araya, X., Thomas, R. E., Low, M. J., Cone, R. D. (1997) Exocrine gland dysfunction in MC5-R-deficient mice: evidence for coordinated regulation of exocrine gland function by melanocortin peptides. Cell 91, 789-798.
53. Buggy, J. J. (1998) Binding of α-melanocyte-stimulating hormone to its G-protein-coupled receptor on B-lymphocytes activates the Jak/STAT pathway. Biochem. J. 331, 211-216.
54. Taylor, A., Namba, K. (2001) In vitro induction of CD25+ CD4+ regulatory T cells by the neuropeptide α-melanocyte stimulating hormone (α-MSH). Immunol. Cell Biol. 79, 358-367.
55. Artuc, M., Grutzkau, A., Luger, T., Henz, B. M. (1999) Expression of MC1- and MC5-receptors on the human mast cell line HMC-1. Ann. N. Y. Acad. Sci. 885, 364-367.
56. Busca, R., Ballotti, R. (2000) Cyclic AMP a key messenger in the regulation of skin pigmentation. Pigment Cell Res. 13, 60-69.
57. Harris, M., Aschkenasi, C., Elias, C. F., Chandrankunnel, A., Nillni, E. A., Bjoorbaek, C., Elmquist, J. K., Flier, J. S., Hollenberg, A. N. (2001) Transcriptional regulation of the thyrotropin-releasing hormone gene by leptin and melanocortin signaling. J. Clin. Invest. 107, 111-120.
58. Sarkar, S., Legradi, G., Lechan, R. M. (2002) Intracerebroventricular administration of α-melanocyte stimulating hormone increases phosphorylation of CREB in TRH- and CRH-producing neurons of the hypothalamic paraventricular nucleus. Brain Res. 945, 50-59.
59. Gerst, J. E., Sole, J., Salomon, Y. (1987) Dual regulation of β-melanotropin receptor function and adenylate cyclase by calcium and guanosine nucleotides in the M2R melanoma cell line. Mol. Pharmacol. 31, 81-88.
60. Mountjoy, K. G., Kong, P. L., Taylor, J. A., Willard, D. H., Wilkison, W. O. (2001) Melanocortin receptor-mediated mobilization of intracellular free calcium in HEK293 cells. Physiol. Genomics 5, 11-19.
61. Konda, Y., Gantz, I., DelValle, J., Shimoto, Y., Miwa, H., Yamada, T. (1994) Interaction of dual intracellular signaling pathways activated by the melanocortin-3 receptor. J. Biol. Chem. 269, 13162-13166.
62. Vongs, A., Lynn, N. M., Rosenblum, C. I. (2004) Activation of MAP kinase by MC4-R through PI3 kinase. Regul. Pept. 120, 113-118.
63. Daniels, D., Patten, C. S., Roth, J. D., Yee, D. K., Fluharty, S. J. (2003) Melanocortin receptor signaling through mitogen-activated protein kinase in vitro and in rat hypothalamus. Brain Res. 986, 1-11.
64. Manna, S. K., Aggarwal, B. B. (1998) α-Melanocyte-stimulating hormone inhibits the nuclear transcription factor NF-κ B activation induced by various inflammatory agents. J. Immunol. 161, 2873-2880.
65. Gupta, A. K., Diaz, R. A., Higham, S., Kone, B. C. (2000) α-MSH inhibits induction of C/EBPβ-DNA binding activity and NOS2 gene transcription in macrophages. Kidney Int. 57, 2239-2248.
66. Haddad, J. J., Lauterbach, R., Saade, N. E., Safieh-Garabedian, B., Land, S. C. (2001) α-Melanocyte-related tripeptide, Lys-d-Pro-Val, ameliorates endotoxin-induced nuclear factor κB translocation and activation: evidence for involvement of an interleukin-1β193-195 receptor antagonism in the alveolar epithelium. Biochem. J. 355, 29-38.
67. Hassoun, H. T., Zou, L., Moore, F. A., Kozar, R. A., Weisbrodt, N. W., Kone, B. C. (2002) α-Melanocyte-stimulating hormone protects against mesenteric ischemia-reperfusion injury. Am. J. Physiol. Gastrointest. Liver Physiol. 282, G1059-G1068.
68. Haycock, J. W., Wagner, M., Morandini, R., Ghanem, G., Rennie, I. G., Mac Neil, S. (1999) α-Melanocyte-stimulating hormone inhibits NF-κB activation in human melanocytes and melanoma cells. J. Invest. Dermatol. 113, 560-566.
69. Ichiyama, T., Campbell, I. L., Furukawa, S., Catania, A., Lipton, J. M. (1999) Autocrine α-melanocyte-stimulating hormone inhibits NF-κB activation in human glioma. J. Neurosci. Res. 58, 684-689.
70. Ichiyama, T., Okada, K., Campbell, I. L., Furukawa, S., Lipton, J. M. (2000) NF-κB activation is inhibited in human pulmonary epithelial cells transfected with α-melanocyte-stimulating hormone vector. Peptides 21, 1473-1477.
71. Mandrika, I., Muceniece, R., Wikberg, J. E. (2001) Effects of melanocortin peptides on lipopolysaccharide/interferon-γ-induced NF-κB DNA binding and nitric oxide production in macrophage-like RAW 264.7 cells: evidence for dual mechanisms of action. Biochem. Pharmacol. 61, 613-621.
72. May, M. J., Ghosh, S. (1998) Signal transduction through NF-κB. Immunol. Today 19, 80-88.
73. Ichiyama, T., Zhao, H., Catania, A., Furukawa, S., Lipton, J. M. (1999) α-Melanocyte-stimulating hormone inhibits NF-κB activation and IκBα degradation in human glioma cells and in experimental brain inflammation. Exp. Neurol. 157, 359-365.
74. Bohm, M., Schulte, U., Kalden, H., Luger, T. A. (1999) α-Melanocyte-stimulating hormone modulates activation of NF-κ B and AP-1 and secretion of interleukin-8 in human dermal fibroblasts. Ann. N. Y. Acad. Sci. 885, 277-286.
75. Hill, R. P., MacNeil, S., Haycock, J. W. (2006) Melanocyte stimulating hormone peptides inhibit TNF-α signaling in human dermal fibroblast cells. Peptides 27, 421-430.
76. Kalden, D. H., Scholzen, T., Brzoska, T., Luger, T. A. (1999) Mechanisms of the antiinflammatory effects of α-MSH. Role of transcription factor NF-κ B and adhesion molecule expression. Ann. N. Y. Acad. Sci. 885, 254-261.
77. Brzoska, T., Kalden, D. H., Scholzen, T., Luger, T. A. (1999) Molecular basis of the α-MSH/IL-1 antagonism. Ann. N. Y. Acad. Sci. 885, 230-238.
78. Etemad-Moghadam, B., Chen, H., Yin, P., Aziz, N., Hedley, M. L. (2002) Inhibition of NF-κB activity by plasmid expressed α-MSH peptide. J. Neuroimmunol. 125, 23-29.
79. Taylor, A. W. (2005) The immunomodulating neuropeptide α-melanocyte-stimulating hormone (α-MSH) suppresses LPS-stimulated TLR4 with IRAK-M in macrophages. J. Neuroimmunol. 162, 43-50.
80. Holder, J. R., Haskell-Luevano, C. (2004) Melanocortin ligands: 30 years of structure-activity relationship (SAR) studies. Med. Res. Rev. 24, 325-356.
81. Macaluso, A., McCoy, D., Ceriani, G., Watanabe, T., Biltz, J., Catania, A., Lipton, J. M. (1994) Antiinflammatory influences of α-MSH molecules: central neurogenic and peripheral actions. J. Neurosci. 14, 2377-2382.
82. Ichiyama, T., Sakai, T., Catania, A., Barsh, G. S., Furukawa, S., Lipton, J. M. (1999) Inhibition of peripheral NF-κB activation by central action of α-melanocyte-stimulating hormone. J. Neuroimmunol. 99, 211-217.
83. Mioni, C., Bazzani, C., Giuliani, D., Altavilla, D., Leone, S., Ferrari, A., Minutoli, L., Bitto, A., Marini, H., Zaffe, D., Botticelli, A. R., Iannone, A., Tomasi, A., Bigiani, A., Bertolini, A., Squadrito, F., Guarini, S. (2005) Activation of an efferent cholinergic pathway produces strong protection against myocardial ischemia/reperfusion injury in rats. Crit. Care Med. 33, 2621-2628.
84. Matsumura, K., Tsuchihashi, T., Abe, I., Iida, M. (2002) Central α-melanocyte-stimulating hormone acts at melanocortin-4 receptor to activate sympathetic nervous system in conscious rabbits. Brain Res. 948, 145-148.
85. Oktar, B. K., Alican, I. (2002) Modulation of the peripheral and central inflammatory responses by α-melanocyte stimulating hormone. Curr. Protein Pept. Sci. 3, 623-628.
86. Bazzani, C., Mioni, C., Ferrazza, G., Cainazzo, M. M., Bertolini, A., Guarini, S. (2002) Involvement of the central nervous system in the protective effect of melanocortins in myocardial ischaemia/reperfusion injury. Resuscitation 52, 109-115.
87. Guarini, S., Cainazzo, M. M., Giuliani, D., Mioni, C., Altavilla, D., Marini, H., Bigiani, A., Ghiaroni, V., Passaniti, M., Leone, S., Bazzani, C., Caputi, A. P., Squadrito, F., Bertolini, A. (2004) Adrenocorticotropin reverses hemorrhagic shock in anesthetized rats through the rapid activation of a vagal anti-inflammatory pathway. Cardiovasc. Res. 63, 357-365.
88. Tracey, K. J., Czura, C. J., Ivanova, S. (2001) Mind over immunity. FASEB J. 15, 1575-1576.
89. Getting, S. J., Christian, H. C., Lam, C. W., Gavins, F. N. E., Flower, R. J., Schioth, H. B., Perretti, M. (2003) Redundancy of a functional melanocortin 1 receptor in the anti-inflammatory actions of melanocortin peptides: studies in the recessive yellow (e/e) mouse suggest an important role for melanocortin 3 receptor. J. Immunol. 170, 3323-3330.
90. Getting, S. J., Allcock, G. H., Flower, R., Perretti, M. (2001) Natural and synthetic agonists of the melanocortin receptor type 3 possess antiinflammatory properties. J. Leukoc. Biol. 69, 98-104.
91. Lam, C. W., Getting, S. J. (2004) Melanocortin receptor type 3 as a potential target for anti-inflammatory therapy. Curr. Drug Targets Inflamm. Allergy 3, 311-315.
92. Neumann Andersen, G., Nagaeva, O., Mandrika, I., Petrovska, R., Muceniece, R., Mincheva-Nilsson, L., Wikberg, J. E. S. (2001) MC1 receptors are constitutively expressed on leucocyte subpopulations with antigen presenting and cytotoxic functions. Clin. Exp. Immunol. 126, 441-446.
93. Scholzen, T. E., Brzoska, T., Kalden, D. H., Hartmeyer, M., Fastrich, M., Luger, T. A., Armstrong, C. A., Ansel, J. C. (1999) Expression of functional melanocortin receptors and proopiomelanocortin peptides by human dermal microvascular endothelial cells. Ann. N. Y. Acad. Sci. 885, 239-253.
94. Lipton, J. M., Zhao, H., Ichiyama, T., Barsh, G. S., Catania, A. (1999) Mechanisms of antiinflammatory action of α-MSH peptides. In vivo and in vitro evidence. Ann. N. Y. Acad. Sci. 885, 173-182.
95. Lyson, K., Ceriani, G., Takashima, A., Catania, A., Lipton, J. M. (1994) Binding of anti-inflammatory α-melanocyte-stimulating-hormone peptides and proinflammatory cytokines to receptors on melanoma cells. Neuroimmunomodulation 1, 121-126.
96. Muceniece, R., Krigere, L., Suli-Vargha, H., Wikberg, J. E. (2003) Effects of α-melanotropin C-terminal tripeptide analogues on macrophage NO production. Peptides 24, 701-707.
97. Getting, S. J., Schioth, H. B., Perretti, M. (2003) Dissection of the anti-inflammatory effect of the core and C-terminal (KPV) α-melanocyte- stimulating hormone peptides. J. Pharmacol. Exp. Ther. 306, 631-637.
98. Sarkar, A., Sreenivasan, Y., Manna, S. K. (2003) α-Melanocyte- stimulating hormone inhibits lipopolysaccharide-induced biological responses by downregulating CD14 from macrophages. FEBS Lett. 553, 286-294.
99. Bhardwaj, R. S., Schwarz, A., Becher, E., Mahnke, K., Aragane, Y., Schwarz, T., Luger, T. A. (1996) Pro-opiomelanocortin-derived peptides induce IL-10 production in human monocytes. J. Immunol. 156, 2517-2521.
100. Yoon, S. W., Goh, S. H., Chun, J. S., Cho, E. W., Lee, M. K., Kim, K. L., Kim, J. J., Kim, C. J., Poo, H. (2003) α-Melanocyte-stimulating hormone inhibits lipopolysaccharide-induced tumor necrosis factor-α production in leukocytes by modulating protein kinase A, p38 kinase, and nuclear factor κ B signaling pathways. J. Biol. Chem. 278, 32914-32920.
101. Delgado, R., Carlin, A., Airaghi, L., Demitri, M. T., Meda, L., Galimberti, D., Baron, P., Lipton, J. M., Catania, A. (1998) Melanocortin peptides inhibit production of proinflammatory cytokines and nitric oxide by activated microglia. J. Leukoc. Biol. 63, 740-745.
102. Galimberti, D., Baron, P., Meda, L., Prat, E., Scarpini, E., Delgado, R., Catania, A., Lipton, J. M., Scarlato, G. (1999) α-MSH peptides inhibit production of nitric oxide and tumor necrosis factor-α by microglial cells activated with β-amyloid and interferon γ. Biochem. Biophys. Res. Commun. 263, 251-256.
103. Catania, A., Cutuli, M., Garofalo, L., Airaghi, L., Valenza, F., Lipton, J. M., Gattinoni, L. (2000) Plasma concentrations and anti-L-cytokine effects of α-melanocyte stimulating hormone in septic patients. Crit. Care Med. 28, 1403-1407.
104. Catania, A., Garofalo, L., Cutuli, M., Gringeri, A., Santagostino, E., Lipton, J. M. (1998) Melanocortin peptides inhibit production of proinflammatory cytokines in blood of HIV-infected patients. Peptides 19, 1099-1104.
105. Jahovic, N., Arbak, S., Tekeli, O., Alican, I. (2004) α-Melanocyte stimulating hormone has beneficial effects on cerulein-induced acute pancreatitis. Peptides 25, 129-132.
106. Gatti, S., Colombo, G., Buffa, R., Turcatti, F., Garofalo, L., Carboni, N., Ferla, L., Fassati, L. R., Lipton, J. M., Catania, A. (2002) α-Melanocyte-stimulating hormone protects the allograft in experimental heart transplantation. Transplantation 74, 1678-1684.
107. Jo, S. K., Yun, S. Y., Chang, K. H., Cha, D. R., Cho, W. Y., Kim, H. K., Won, N. H. (2001) α-MSH decreases apoptosis in ischaemic acute renal failure in rats: possible mechanism of this beneficial effect. Nephrol. Dial. Transplant. 16, 1583-1591.
108. Delgado Hernandez, R., Demitri, M. T., Carlin, A., Meazza, C., Villa, P., Ghezzi, P., Lipton, J. M., Catania, A. (1999) Inhibition of systemic inflammation by central action of the neuropeptide α-melanocyte- stimulating hormone. Neuroimmunomodulation 6, 187-192.
109. Chiao, H., Foster, S., Thomas, R., Lipton, J., Star, R. A. (1996) α-Melanocyte-stimulating hormone reduces endotoxin-induced liver inflammation. J. Clin. Invest. 97, 2038-2044.
110. Ceriani, G., Diaz, J., Murphree, S., Catania, A., Lipton, J. M. (1994) The neuropeptide α-melanocyte-stimulating hormone inhibits experimental arthritis in rats. Neuroimmunomodulation 1, 28-32.
111. Lipton, J. M., Ceriani, G., Macaluso, A., McCoy, D., Carnes, K., Biltz, J., Catania, A. (1994) Antiinflammatory effects of the neuropeptide α-MSH in acute, chronic, and systemic inflammation. Ann. N. Y. Acad. Sci. 741, 137-148.
112. Colombo, G., Gatti, S., Turcatti, F., Sordi, A., Fassati, L. R., Bonino, F., Lipton, J. M., Catania, A. (2005) Gene expression profiling reveals multiple protective influences of the peptide α-melanocyte-stimulating hormone in experimental heart transplantation. J. Immunol. 175, 3391-3401.
113. Getting, S. J., Di Filippo, C., D'Amico, M., Perretti, M. (2006) The melanocortin peptide HP228 displays protective effects in acute models of inflammation and organ damage. Eur. J. Pharmacol. 532, 138-144.
114. Starowicz, K., Przewlocka, B. (2003) The role of melanocortins and their receptors in inflammatory processes, nerve regeneration and nociception. Life Sci. 73, 823-847.
115. Skottner, A., Post, C., Ocklind, A., Seifert, E., Liutkevicius, E., Meskys, R., Pilinkiene, A., Biziuleviciene, G., Lundstedt, T. (2003) Anti-inflammatory potential of melanocortin receptor-directed drugs. Ann. N. Y. Acad. Sci. 994, 84-89.
116. Catania, A., Gatti, S., Colombo, G., Lipton, J. M. (2003) α-Melanocyte stimulating hormone in modulation of inflammatory reactions. Pediatr. Endocrinol. Rev. 1, 101-108.
117. Luger, T. A., Kalden, D., Scholzen, T. E., Brzoska, T. (1999) α-Melanocyte-stimulating hormone as a mediator of tolerance induction. Pathobiology 67, 318-321.
118. Luger, T. A., Scholzen, T., Brzoska, T., Becher, E., Slominski, A., Paus, R. (1998) Cutaneous immunomodulation and coordination of skin stress responses by α-melanocyte-stimulating hormone. Ann. N. Y. Acad. Sci. 840, 381-394.
119. Luger, T. A., Brzoska, T., Scholzen, T. E., Kalden, D. H., Sunderkotter, C., Armstrong, C., Ansel, J. (2000) The role of α-MSH as a modulator of cutaneous inflammation. Ann. N. Y. Acad. Sci. 917, 232-238.
120. Muceniece, R., Zvejniece, L., Liepinsh, E., Kirjanova, O., Baumane, L., Petrovska, R., Mutulis, F., Mutule, I., Kalvinsh, I., Wikberg, J. E., Dambrova, M. (2006) The MC3 receptor binding affinity of melanocortins correlates with the nitric oxide production inhibition in mice brain inflammation model. Peptides 27, 1443-1450.
121. Zhong, Y., Bellamkonda, R. V. (2005) Controlled release of anti-inflammatory agent α-MSH from neural implants. J. Control. Release 106, 309-318.
122. Caruso, C., Mohn, C., Karara, A. L., Rettori, V., Watanobe, H., Schioth, H. B., 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.
123. Nishida, T., Miyata, S., Itoh, Y., Mizuki, N., Ohgami, K., Shiratori, K., Ilieva, I. B., Ohno, S., Taylor, A. W. (2004) Anti-inflammatory effects of α-melanocyte-stimulating hormone against rat endotoxin-induced uveitis and the time course of inflammatory agents in aqueous humor. Int. Immunopharmacol. 4, 1059-1066.
124. San, T., Oktar, B. K., Salik, E., Ercan, F., Alican, I. (2001) The effect of α-melanocyte stimulating hormone on endotoxin-induced intestinal injury. Peptides 22, 2077-2082.
125. Rajora, N., Boccoli, G., Catania, A., Lipton, J. M. (1997) α-MSH modulates experimental inflammatory bowel disease. Peptides 18, 381-385.
126. Gatti, S., Carlin, A., Sordi, A., Leonardi, P., Colombo, G., Fassati, L. R., Lipton, J. M., Catania, A. (2006) Inhibitory effects of the peptide (CKPV)2 on endotoxin-induced host reactions. J. Surg. Res. 131, 209-214.
127. Rajora, N., Boccoli, G., Burns, D., Sharma, S., Catania, A. P., Lipton, J. M. (1997) α-MSH modulates local and circulating tumor necrosis factor-α in experimental brain inflammation. J. Neurosci. 17, 2181-2186.
128. Giuliani, D., Mioni, C., Altavilla, D., Leone, S., Bazzani, C., Minutoli, L., Bitto, A., Cainazzo, M. M., Marini, H., Zaffe, D., Botticelli, A. R., Pizzala, R., Savio, M., Necchi, D., Schioth, H. B., Bertolini, A., Squadrito, F., Guarini, S. (2006) Both early and delayed treatment with melanocortin 4 receptor-stimulating melanocortins produces neuroprotection in cerebral ischemia. Endocrinology 147, 1126-1135.
129. Altavilla, D., Cainazzo, M. M., Squadrito, F., Guarini, S., Bertolini, A., Bazzani, C. (1998) Tumor necrosis factor-α as a target of melanocortins in haemorrhagic shock, in the anaesthetized rat. Br. J. Pharmacol. 124, 1587-1590.
130. Vulliemoz, N. R., Xiao, E., Xia-Zhang, L., Ferin, M., Wardlaw, S. L. (2006) Melanocortin modulation of inflammatory cytokine and neuroendocrine responses to endotoxin in the monkey. Endocrinology 147, 1878-1883.
131. Huang, Q., Tatro, J. B. (2002) α-Melanocyte stimulating hormone suppresses intracerebral tumor necrosis factor-α and interleukin-1β gene expression following transient cerebral ischemia in mice. Neurosci. Lett. 334, 186-190.
132. Chiao, H., Kohda, Y., McLeroy, P., Craig, L., Housini, I., Star, R. A. (1997) α-Melanocyte-stimulating hormone protects against renal injury after ischemia in mice and rats. J. Clin. Invest. 99, 1165-1172.
133. Scholzen, T. E., Sunderkotter, C., Kalden, D. H., Brzoska, T., Fastrich, M., Fisbeck, T., Armstrong, C. A., Ansel, J. C., Luger, T. A. (2003) α-Melanocyte stimulating hormone prevents lipopolysaccharide-induced vasculitis by down-regulating endothelial cell adhesion molecule expression. Endocrinology 144, 360-370.
134. Lipton, J. M., Glyn, J. R., Zimmer, J. A. (1981) Hypothermic and antipyretic effects of centrally administered ACTH(1-24) and α-melanotropin. Fed. Proc. 40, 2760-2762.