Casu Benito, Guerrini Marco, Torri Giangiacomo
Ronzoni Institute for Chemical and Biochemical Research, Milan, Italy.
Curr Pharm Des. 2004;10(9):939-49. doi: 10.2174/1381612043452794.
Heparin and other iduronic acid-containing glycosaminoglycans (GAG) such as dermatan sulfate exert their anticoagulant properties primarily by accelerating the rate of inhibition of the natural protease inhibitors antithrombin III (AT, which inhibits both factor Xa and thrombin) and heparin cofactor II (HCII, which selectively inhibits thrombin). Although AT and HCII are structural homologs, only heparin binds to AT, and HCII has different binding sites for heparin and dermatan sulfate. Whereas the binding site of heparin for AT is a unique pentasaccharide sequence contained in only about one third of the chains of this GAG, HCII-binding sequences of heparin and dermatan sulfate are less specific and contained in practically all the GAG chains. Protein binding and associated biological activities of heparin and dermatan sulfate are modulated by the "plasticity" of their iduronic acid residues due to the availability of up to three equienergetic conformation among which the protein selects the one favouring the most stable complex. Glycol-splitting of nonsulfated uronic acid residues, a device for generating flexible joints along the GAG chains, has different effects on different binding domains. Whereas it inactivates the binding site for AT causing a drop of the anticoagulant activity, it enhances the HCII-associated activity of both heparin and dermatan sulfate.
肝素以及其他含艾杜糖醛酸的糖胺聚糖(GAG),如硫酸皮肤素,主要通过加速天然蛋白酶抑制剂抗凝血酶III(AT,它能抑制因子Xa和凝血酶)和肝素辅因子II(HCII,它选择性抑制凝血酶)的抑制速率来发挥其抗凝特性。尽管AT和HCII是结构同源物,但只有肝素与AT结合,并且HCII对肝素和硫酸皮肤素具有不同的结合位点。肝素与AT的结合位点是仅约三分之一的这种GAG链中所含的独特五糖序列,而肝素和硫酸皮肤素的HCII结合序列特异性较低,几乎存在于所有GAG链中。肝素和硫酸皮肤素的蛋白质结合及相关生物学活性因其艾杜糖醛酸残基的“可塑性”而受到调节,这是由于存在多达三种能量相当的构象,蛋白质会从中选择最有利于形成最稳定复合物的构象。非硫酸化糖醛酸残基的糖裂解作用,即一种沿GAG链生成柔性连接的机制,对不同的结合结构域有不同影响。它会使AT的结合位点失活,导致抗凝活性下降,而它会增强肝素和硫酸皮肤素与HCII相关的活性。