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14-September-2008 18:02:46 - Glucagon Glucagon PDB rendering based on 1d0r. Available structures: 1d0r, 1gcn, 1kx6, 2g49 Identifiers Symbols GCG; GLP1; GLP2; GRPP External IDs OMIM: 138030 MGI: 95674 HomoloGene: 1553 Gene ontology Molecular function: hormone activity Cellular component: extracellular region soluble fraction Biological process: signal transduction G-protein coupled receptor protein signaling pathway feeding behavior cell proliferation regulation of insulin secretion RNA expression pattern More reference expression data Orthologs Human Mouse Entrez 2641 14526 Ensembl ENSG00000115263 ENSMUSG00000000394 Uniprot P01275 P55095 Refseq NM_002054 mRNA NP_002045 protein NM_008100 mRNA NP_032126 protein Location Chr 2: 162.71 - 162.71 Mb Chr 2: 62.28 - 62.28 Mb Pubmed search 1 2 Glucagon is an important hormone involved in carbohydrate metabolism. Produced by the pancreas, it is released when the glucose level in the blood is low hypoglycemia, causing the liver to convert stored glycogen into glucose and release it into the bloodstream. The action of glucagon is thus opposite to that of insulin, which instructs the body's cells to take in glucose from the blood in times of satiation. Contents 1 History 2 Structure 3 Physiology 3.1 Production 3.2 Regulatory mechanism 3.3 Function 3.4 Mechanism of action 4 Pathology 5 Uses 6 Media 7 References in pop culture 8 References 9 Further reading 10 See also History In the 1920s, Kimball and Murlin studied pancreatic extracts and found an additional substance with hyperglycemic properties. They described glucagon in 1923.1 The amino acid sequence of glucagon was described in the late-1950s.2 A more complete understanding of its role in physiology and disease was not established until the 1970s, when a specific radioimmunoassay was developed. Structure Glucagon ball and stick model, with the carboxyl terminus above and the amino terminus below Glucagon ball and stick model, with the carboxyl terminus above and the amino terminus below Glucagon is a 29-amino acid polypeptide. Its primary structure in humans is: NH2-His-Ser-Gln-Gly-Thr-Phe- Thr-Ser-Asp-Tyr-Ser-Lys-Tyr-Leu-Asp-Ser- Arg-Arg-Ala-Gln-Asp-Phe-Val-Gln-Trp-Leu- Met-Asn-Thr-COOH. The polypeptide has a molecular weight of 3485 daltons. Physiology A microscopic image stained for glucagon. A microscopic image stained for glucagon. Production The hormone is synthesized and secreted from alpha cells α-cells of the islets of Langerhans, which are located in the endocrine portion of the pancreas. In rodents, the alpha cells are located in the outer rim of the islet. Human islet structure is much less segregated, and alpha cells are distributed throughout the islet. Regulatory mechanism Increased secretion of glucagon is caused by: Decreased plasma glucose Increased catecholamines - norepinephrine and epinephrine Increased plasma amino acids to protect from hypoglycemia if an all protein meal is consumed Sympathetic nervous system Acetylcholine Cholecystokinin Decreased secretion of glucagon inhibition is caused by: Somatostatin Insulin Function Glucagon helps maintain the level of glucose in the blood by binding to glucagon receptors on hepatocytes, causing the liver to release glucose - stored in the form of glycogen - through a process known as glycogenolysis. As these stores become depleted, glucagon then encourages the liver to synthesize additional glucose by gluconeogenesis. This glucose is released into the bloodstream. Both of these mechanisms lead to glucose release by the liver, preventing the development of hypoglycemia. Glucagon also regulates the rate of glucose production through lipolysis. Glucagon production appears to be dependent on the central nervous system through pathways which are yet to be defined. It has been reported that in invertebrate animals eyestalk removal can affect glucagon production. Excising the eyestalk in young crayfish produces glucagon-induced hyperglycemia. 3 Increased free fatty acids and ketoacids into the blood Increased urea production Mechanism of action Glucagon binds to the glucagon receptor, a G protein-coupled receptor located in the plasma membrane. The conformation change in the receptor activates G proteins, a heterotrimeric protein with α, β, and γ subunits. The subunits breakup as a result of substitution of a GDP molecule with a GTP mol, and the alpha subunit specifically activates the next enzyme in the cascade, adenylate cyclase. Adenylate cyclase manufactures cAMP cyclical AMP which activates protein kinase A cAMP-dependent protein kinase. This enzyme in turn activates phosphorylase kinase, which in turn, phosphorylates glycogen phosphorylase, converting into the active form called phosphorylase A. Phosphorylase A is the enzyme responsible for the release of glucose-1-phosphate from glycogen polymers. Pathology Abnormally-elevated levels of glucagon may be caused by pancreatic tumors such as glucagonoma, symptoms of which include necrolytic migratory erythema NME, reduced amino acids and hyperglycemia. It may occur alone or in the context of multiple endocrine neoplasia type 1. Uses An injectable form of glucagon is vital first aid in cases of severe hypoglycemia when the victim is unconscious or for other reasons cannot take glucose orally. The dose for an adult is typically 1 milligram, and the glucagon is given by intramuscular, intravenous or subcutaneous injection, and quickly raises blood glucose levels. Glucagon can also be administered intravenously at 0.25 - 0.5 unit. Anecdotal evidence suggests a benefit of higher doses of glucagon in the treatment of overdose with beta blockers; the likely mechanism of action is the increase of cAMP in the myocardium, effectively bypassing the inhibitory action of the β-adrenergic second messenger system.4 Glucagon acts very quickly: common side effects include headache and nausea. Drug interactions: Glucagon interacts only with oral anticoagulants increasing the tendency to bleed. Media Rotating stereogram animation of glucagon. 1.70 MB, animated GIF format. Rotating stereogram animation of glucagon. 1.70 MB, animated GIF format. References in pop culture American parodist Weird Al Yankovic released a song entitled Pancreas on his album Straight Outta Lynwood, which uses the lyric Insulin, Glucagon, flowing from the Islets of Langerhans as a repeated catchy chorus in the final minute of the song. References ^ Kimball C, Murlin J. Aqueous extracts of pancreas III. Some precipitation reactions of insulin. J Biol Chem 1923;58:337-348. PDF fulltext. ^ Bromer W, Winn L, Behrens O. The amino acid sequence of glucagon V. Location of amide groups, acid degradation studies and summary of sequential evidence. J Am Chem Soc 1957;79:2807-2810. ^ RL Leinen, AJ Giannini. Effect of eyestalk removal on glucagon induced hyperglycemia in crayfish. Society for Neuroscience Abstracts. 9:604, 1983 ^ White CM. A review of potential cardiovascular uses of intravenous glucagon administration. J Clin Pharmacol 1999;39:442-7. PMID 10234590. Further reading Kieffer TJ, Habener JF 2000. The glucagon-like peptides. Endocr. Rev. 20 6: 876-913. PMID 10605628. Drucker DJ 2003. Glucagon-like peptides: regulators of cell proliferation, differentiation, and apoptosis. Mol. Endocrinol. 17 2: 161-71. doi:10.1210/me.2002-0306. PMID 12554744. Jeppesen PB 2004. Clinical significance of GLP-2 in short-bowel syndrome. J. Nutr. 133 11: 3721-4. PMID 14608103. Brubaker PL, Anini Y 2004. Direct and indirect mechanisms regulating secretion of glucagon-like peptide-1 and glucagon-like peptide-2. Can. J. Physiol. Pharmacol. 81 11: 1005-12. doi:10.1139/y03-107. PMID 14719035. Baggio LL, Drucker DJ 2005. Clinical endocrinology and metabolism. Glucagon-like peptide-1 and glucagon-like peptide-2. Best Pract. Res. Clin. Endocrinol. Metab. 18 4: 531-54. doi:10.1016/j.beem.2004.08.001. PMID 15533774. Holz GG, Chepurny OG 2006. Diabetes outfoxed by GLP-1?. Sci. STKE 2005 268: pe2. doi:10.1126/stke.2682005pe2. PMID 15671479. Dunning BE, Foley JE, Ahrén B 2006. Alpha cell function in health and disease: influence of glucagon-like peptide-1. Diabetologia 48 9: 1700-13. doi:10.1007/s00125-005-1878-0. PMID 16132964. Gautier JF, Fetita S, Sobngwi E, Salaün-Martin C 2005. Biological actions of the incretins GIP and GLP-1 and therapeutic perspectives in patients with type 2 diabetes. Diabetes Metab. 31 3 Pt 1: 233-42. doi:10.1016/S1262-36360770190-8. PMID 16142014. De León DD, Crutchlow MF, Ham JY, Stoffers DA 2006. Role of glucagon-like peptide-1 in the pathogenesis and treatment of diabetes mellitus. Int. J. Biochem. Cell Biol. 38 5-6: 845-59. doi:10.1016/j.biocel.2005.07.011. PMID 16202636. Beglinger C, Degen L 2007. Gastrointestinal satiety signals in humans--physiologic roles for GLP-1 and PYY?. Physiol. Behav. 89 4: 460-4. doi:10.1016/j.physbeh.2006.05.048. PMID 16828127. Stephens JW, Bain SC 2007. Safety and adverse effects associated with GLP-1 analogues. Expert opinion on drug safety 6 4: 417-22. doi:10.1517/14740338.6.4.417. PMID 17688385. Orskov C, Bersani M, Johnsen AH, et al. 1989. Complete sequences of glucagon-like peptide-1 from human and pig small intestine. J. Biol. Chem. 264 22: 12826-9. PMID 2753890. Drucker DJ, Asa S 1988. Glucagon gene expression in vertebrate brain. J. Biol. Chem. 263 27: 13475-8. PMID 2901414. Novak U, Wilks A, Buell G, McEwen S 1987. Identical mRNA for preproglucagon in pancreas and gut. Eur. J. Biochem. 164 3: 553-8. doi:10.1111/j.1432-1033.1987.tb11162.x. PMID 3569278. White JW, Saunders GF 1986. Structure of the human glucagon gene. Nucleic Acids Res. 14 12: 4719-30. doi:10.1093/nar/14.12.4719. PMID 3725587. Schroeder WT, Lopez LC, Harper ME, Saunders GF 1984. Localization of the human glucagon gene GCG to chromosome segment 2q36----37. Cytogenet. Cell Genet. 38 1: 76-9. PMID 6546710. Bell GI, Sanchez-Pescador R, Laybourn PJ, Najarian RC 1983. Exon duplication and divergence in the human preproglucagon gene. Nature 304 5924: 368-71. doi:10.1038/304368a0. PMID 6877358. Kärgel HJ, Dettmer R, Etzold G, et al. 1982. Action of rat liver cathepsin L on glucagon. Acta Biol. Med. Ger. 40 9: 1139-43. PMID 7340337. Wayman GA, Impey S, Wu Z, et al. 1994. Synergistic activation of the type I adenylyl cyclase by Ca2+ and Gs-coupled receptors in vivo. J. Biol. Chem. 269 41: 25400-5. PMID 7929237. Unson CG, Macdonald D, Merrifield RB 1993. The role of histidine-1 in glucagon action. Arch. Biochem. Biophys. 300 2: 747-50. doi:10.1006/abbi.1993.1103. PMID 8382034. See also Insulin Diabetes mellitus Proglucagon Glucagon-like peptide-1 Glucagon-like peptide-2 Islets of Langerhans Pancreas v d e Endocrine system: hormones/endocrine glands Peptide hormones, Steroid hormones Hypothalamic-pituitary Hypothalamus: TRH, CRH , GnRH, GHRH, somatostatin, dopamine - Posterior pituitary: vasopressin, oxytocin - Anterior pituitary: α FSH, LH, TSH, GH, prolactin, POMC ACTH, MSH, endorphins, lipotropin Adrenal axis Adrenal medulla: epinephrine, norepinephrine - Adrenal cortex: aldosterone, cortisol, DHEA Thyroid axis Thyroid: thyroid hormone T3 and T4 - calcitonin - Parathyroid: PTH Gonadal axis Testis: testosterone, AMH, inhibin - Ovary: estradiol, progesterone, inhibin/activin, relaxin pregnancy Other end. glands Pancreas: glucagon, insulin, somatostatin - Pineal gland: melatonin Non-end. glands Placenta: hCG, HPL, estrogen, progesterone - Kidney: renin, EPO, calcitriol, prostaglandin - Heart atrium: ANP - Stomach: gastrin, ghrelin - Duodenum: CCK, GIP, secretin, motilin, VIP - Ileum: enteroglucagon - Adipose tissue: leptin, adiponectin, resistin - Thymus: Thymosin - Thymopoietin - Thymulin - Skeleton: Osteocalcin - Liver/other: Insulin-like growth factor IGF-1, IGF-2 Target-derived NGF, BDNF, NT-3 v d e Gastrointestinal hormones, pancreatic hormones: proglucagon Oxyntomodulin Glucagon - Glucagon-Like Peptides GLP1, GLP2 Retrieved from http://en..org/wiki/Glucagon Categories: Genes on chromosome 2 | Human proteins | Peptide hormones | Pancreatic hormones | Hepatology | Metabolism Views Article Discussion this page History Personal tools Log in / create account Navigation Main page Contents Featured content Current events Random article Search Go Search Interaction Community portal Recent changes Contact Donate to Help Toolbox What links here Related changes Upload file Special pages Printable version Permanent link Cite this page Languages العربية Bosanski БългарÑ?ки ÄŒesky Dansk Deutsch Þ‹Þ¨ÞˆÞ¬Þ€Þ¨Þ„Þ¦Þ?Þ° Español Esperanto Français Bahasa Indonesia Ã?slenska Italiano עברית Kapampangan Latina Lietuvių МакедонÑ?ки Bahasa Melayu Nederlands 日本語 ‪Norsk bokmÃ¥l‬ Occitan Polski Português РуÑ?Ñ?кий Shqip SlovenÅ¡Ä?ina СрпÑ?ки / Srpski Suomi Svenska தமிழà¯? Türkçe ä¸æ–‡ This page was last modified on 26 August 2008, at 08:27
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