Heparán sulfato: Diferenzas entre revisións

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===Sulfatación 3-O===
CurrentlyActualmente sevencoñécense glucosaminylsete glicosaminil 3-''O''-sulfotransferases (3OSTs) areen knownmamíferos toe existoito inno mammalspeixe (eight in zebrafish)cebra <ref>http://www.ncbi.nlm.nih.gov/pubmed/17195182</ref><ref>http://www.ncbi.nlm.nih.gov/pubmed?term=3OST6%20xu</ref>. TheOs encimas 3OST enzymescrean createvarios aposibles numberdisacáridos ofcon possiblesulfatación 3-''O''-sulfated disaccharides, includingcomo a GlcA-GlcNS(3S±6S) (modifiedmodificado bypor 3OST1 ande 3OST5), IdoA(2S)-GlcNH<sub>2</sub>(3S±6S)(modifiedmodificado bypor 3OST3a, 3OST3b, 3OST5 ande 3OST6) ande GlcA/IdoA(2S)-GlcNS(3S) (modifiedmodificado bypor 3OST2 ande 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>. AsIgual withque alltodas otheras HSdemais sulfotransferases do heparán sulfato, theas 3OSTs useutilizan o 3'-phosphoadenosinefosfoadenosina-5'-phosphosulfatefosfosulfato (PAPS) ascomo adoante sulfatede donorsulfatos. DespiteA beingpesar thede largestser familya offamilia HSmáis modificationgrande enzymesde encimas modificadores do heparán sulfato, theas 3OSTs produceproducen theas rarestmodificación HSmáis modificationraras do heparán sulfato, thea sulfatación 3-''O''-sulfation ofde specificresiduos glucosamineespecíficos residuesde atglicosamina thenun resto C3-OH moiety <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>.
 
The 3OSTs are divided into two functional subcatagories, those which generate an antithrombin III binding site (3OST1 and 3OST5) and those which generate a herpes simplex virus 1 glycoprotein D (HSV-1 gD) binding site (3OST2, 3OST3a, 3OST3b, 3OST4, 3OST5 and 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>. As the 3OSTs are the largest family of HS modification enzymes and their actions are rate-limiting, substrate specific and produce rare modifications, it has been hypothesized that 3OST modified HS plays an important regulatory role in biological processes <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>.