Heparán sulfato: Diferenzas entre revisións
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===Sulfatación 3-O===
Actualmente coñécense sete glicosaminil 3-''O''-sulfotransferases (3OSTs) en mamíferos e oito no peixe cebra <ref>http://www.ncbi.nlm.nih.gov/pubmed/17195182</ref><ref>http://www.ncbi.nlm.nih.gov/pubmed?term=3OST6%20xu</ref>. Os encimas 3OST crean varios posibles disacáridos con sulfatación 3-''O'', como a GlcA-GlcNS(3S±6S) (modificado por 3OST1 e 3OST5), IdoA(2S)-GlcNH<sub>2</sub>(3S±6S) (modificado por 3OST3a, 3OST3b, 3OST5 e 3OST6) e GlcA/IdoA(2S)-GlcNS(3S) (modificado por 3OST2 e 3OST4) <ref>Shukla, D., Liu, J., Blaiklock, P., Shworak, N.W., Bai, X., Esko, J.D., Cohen, G.H., Eisenberg, R.J., Rosenberg, R.D. and Spear, P.G. (1999). "A Novel Role for 3-O-Sulfated Heparan Sulfate in Herpes Simplex Virus 1 Entry." Cell 99: 13-22.</ref><ref>Xia, G., Chen, J., Tiwari, V., Ju, W., Li, J.P., Malmstrom, A., Shukla, D. and Liu, J. (2002). "Heparan Sulfate 3-O-Sulfotransferase Isoform 5 Generates Both an Antithrombin-binding Site and an Entry Receptor for Herpes Simplex Virus, Type 1." The Journal Biological Chemistry 277(40): 37912-37919.</ref><ref>Xu, D., Tiwari, V., Xia, G., Clement, C., Shukla, D. and Liu, J. (2005). "Characterization of heparan sulphate 3-O-sulphotransferase isoform 6 and its role in assisting the entry of herpes simplex virus type 1." Biochemistry Journal 385: 451-459.</ref><ref>Lawrence, R., Yabe, T., HajMohammadi, S., Rhodes, J., McNeely, M., Liu, J., Lamperti, E.D., Toselli, P.A., Lech, M., Spear, P.G., Rosenberg, R.D. and Shworak, N.W. (2007). "The principal neuronal gD-type 3-O-sulfotransferases and their products in the central and peripheral nervous system tissues." Matrix Biology 26: 442-455.</ref>. Igual que todas as demais sulfotransferases do heparán sulfato, as 3OSTs utilizan o 3'-fosfoadenosina-5'-fosfosulfato (PAPS) como doante de sulfatos. A pesar de ser a familia máis grande de encimas modificadores do heparán sulfato, as 3OSTs producen as modificación máis raras do heparán sulfato, a sulfatación 3-''O'' de residuos específicos de glicosamina nun resto C3-OH <ref>Shworak, N.W., HajMohammadi, S., de Agostini, A.I. and Rosenberg, R.D. (2003). "Mice deficient in heparan sulfate 3-O-silfotransferase-1: Normal hemostasis with unexpected perinatal phenotypes." Glycoconjugate Journal 19(4-5): 355-361.</ref>.
Os encimas 3OSTs divídense en dúas subcategorías funcionais: os que xeran un sitio de unión de antitrombina III (3OST1 e 3OST5), e os que xeran un sitio de unión para a glicoproteína D do virus herpes simplex 1 (HSV-1 gD) (3OST2, 3OST3a, 3OST3b, 3OST4, 3OST5 e 3OST6) <ref>Liu, J., Shworak, N.W., Fritze, L.M.S., Edelberg, J.M. and Rosenberg, R.D. (1996). "Purification of Heparan Sulfate D-Glucosaminyl 3-O-Sulfotransferase." The Journal Biological Chemistry 271(43): 27072-27082.</ref><ref>Shworak, N.W., Liu, J., Fritze, L.M.S., Schwartz, J.J., Zhang, L., Logeart, D. and Rosenberg, R.D. (1997). "Molecular Cloning and Expression of Mouse and Human cDNAs Encoding Heparan Sulfate D-Glucosaminyl 3-O-Sulfotransferase." The Journal Biological Chemistry 272(44): 28008-28019.</ref><ref>Shukla, D., Liu, J., Blaiklock, P., Shworak, N.W., Bai, X., Esko, J.D., Cohen, G.H., Eisenberg, R.J., Rosenberg, R.D. and Spear, P.G. (1999). "A Novel Role for 3-O-Sulfated Heparan Sulfate in Herpes Simplex Virus 1 Entry." Cell 99: 13-22.</ref><ref>Shworak, N.W., Liu, J., Petros, L.M., Zhang, L., Kobayashi, M., Copeland, N.G., Jenkins, N.A. and Rosenberg, R.D. (1999). "Multiple Isoforms of Heparan Sulfate D-Glucosaminyl 3-O-Sulfotransferase." The Journal Biological Chemistry 274(8): 5170-5184.</ref><ref>Xia, G., Chen, J., Tiwari, V., Ju, W., Li, J.P., Malmstrom, A., Shukla, D. and Liu, J. (2002). "Heparan Sulfate 3-O-Sulfotransferase Isoform 5 Generates Both an Antithrombin-binding Site and an Entry Receptor for Herpes Simplex Virus, Type 1." The Journal Biological Chemistry 277(40): 37912-37919.</ref><ref>Chen, J., Duncan, M.B., Carrick, K., Pope, R.M. and Liu, J. (2003). "Biosynthesis of 3-O-sulfated heparan sulfate: unique substrate specificity of heparan sulfate 3-O-sulfotransferase isoform 5." Glycobiology 13(11): 785-794.</ref><ref>Duncan, M.B., Chen, J., Krise, J.P. and Liu, J. (2004). "The biosynthesis of anticoagulant heparan sulfate by the heparan sulfate 3-O-sulfotransferase isoform 5." Biochimica et Biophysica Acta 1671: 34-43.</ref><ref>Chen, J. and Liu, J. (2005). "Characterization of the structure of antithrombin-binding heparan sulfate generated by heparan sulfate 3-O-sulfotransferase 5." Biochimica et Biophysica Acta 1725: 190-200.</ref><ref>Girardin, E.P., HajMohammadi, S., Birmele, B., Helisch, A., Shworak, N.W. and de Agostini, A.I. (2005). "Synthesis of Anticoagulantly Active Heparan Sulfate Proteoglycans by Glomerular Epithelial Cells Involves Multiple 3-O-Sulfotransferase isoforms and a Limiting Precursor Pool." The Journal Biological Chemistry 280(45): 38059-38070.</ref><ref>Xu, D., Tiwari, V., Xia, G., Clement, C., Shukla, D. and Liu, J. (2005). "Characterization of heparan sulphate 3-O-sulphotransferase isoform 6 and its role in assisting the entry of herpes simplex virus type 1." Biochemistry Journal 385: 451-459.</ref><ref>Lawrence, R., Yabe, T., HajMohammadi, S., Rhodes, J., McNeely, M., Liu, J., Lamperti, E.D., Toselli, P.A., Lech, M., Spear, P.G., Rosenberg, R.D. and Shworak, N.W. (2007). "The principal neuronal gD-type 3-O-sulfotransferases and their products in the central and peripheral nervous system tissues." Matrix Biology 26: 442-455.</ref>. Como as 3OSTs son a maior familia de encimas modificadores do heparán sulfato e as súas accións son limitantes para a velocidade, específicos de substrato e producen modificacións raras, tense hipotetizada que o heparán sulfato modificado por 3OST xoga un importante papel regulatorio nos procesos biolóxicos <ref>Shworak, N.W., Liu, J., Fritze, L.M.S., Schwartz, J.J., Zhang, L., Logeart, D. and Rosenberg, R.D. (1997). "Molecular Cloning and Expression of Mouse and Human cDNAs Encoding Heparan Sulfate D-Glucosaminyl 3-O-Sulfotransferase." The Journal Biological Chemistry 272(44): 28008-28019.</ref><ref>Shworak, N.W., Liu, J., Fritze, L.M.S., Schwartz, J.J., Zhang, L., Logeart, D. and Rosenberg, R.D. (1997). "Molecular Cloning and Expression of Mouse and Human cDNAs Encoding Heparan Sulfate D-Glucosaminyl 3-O-Sulfotransferase." The Journal Biological Chemistry 272(44): 28008-28019.</ref><ref>Lawrence, R., Yabe, T., HajMohammadi, S., Rhodes, J., McNeely, M., Liu, J., Lamperti, E.D., Toselli, P.A., Lech, M., Spear, P.G., Rosenberg, R.D. and Shworak, N.W. (2007). "The principal neuronal gD-type 3-O-sulfotransferases and their products in the central and peripheral nervous system tissues." Matrix Biology 26: 442-455.</ref>.
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