姚虻唾液腺三类抗血栓活性蛋白的结构和功能研究
其他题名STUDYING ON THE STRUCTURE AND FUNCTION OF THREE ANTITHROMBOSIS BIOACTIVE PROTEINS FROM SALIVARY GLANDS OF THEHORSEFLY, TABANUS YAO MACQUART
马冬莹
学位类型博士
导师赖仞
2010-05-29
学位授予单位中国科学院研究生院
学位授予地点北京
关键词吸血昆虫 牛虻唾液腺 抗血栓 纤溶酶 血小板聚集抑制剂 Apyrase
摘要血栓栓塞性疾病严重影响人类的健康,是导致死亡率最高的病因之一。在过去 的几十年中,已经开发了针对血液凝固、血小板激活和聚集、血栓溶解的不同步骤而 起作用的基础和临床药物,但是许多抗血栓药物经常被包括低血压、出血等系统性的 副作用所限制,因此,需要开发新型的更具潜力和特异性的抗血栓药物。 吸血节肢动物与宿主相互作用的分子机制研究是目前国际研究的热点之一,通过 该类研究可以发现大量的具有药用前景的先导分子和提供节肢动物控制策略。我们通 过生物化学、分子生物学以及药理学研究手段首次从中国特有的姚虻(Tabanus yao Macquart)唾液腺发现了同时具有水解纤维蛋白原和抑制血小板聚集的双功能的 丝氨酸蛋白酶Tablysin、纤维蛋白水解酶TY6和腺苷三磷酸双磷酸酶Tabapy等3个家 族的抗血栓活性成分。并研究了这3类因子在牛虻唾液腺中的分子多样性、它们的结 构基础及发挥抗血栓功能的作用机制,通过实验动物模型研究体内外抗血栓能力,深 入解析了牛虻作为传统抗血栓中药的物质基础,为发掘具有良好抗血栓功能的新型候 选药物分子,为传统活血化瘀中药牛虻的现代化开发提供了广阔的思路和打下了坚实 的基础。 研究表明,姚虻纤维蛋白原水解酶Tablysin为一种单链蛋白。其表观分子量为 27kDa。此酶为一种丝氨酸蛋白酶家族的新成员,其活性不受金属离子螯合剂EDTA 和还原剂DTT的影响。血浆蛋白酶抑制剂Aprotinin,α-macroglobulin能部分抑制该 酶活性。此酶适应的pH范围较广(pH 2.4-10.0),但对热变化不具很好的耐受性。 Edman降解法进行氨基酸测序分析发现,此酶的N-末端有封闭现象,后经Q-TOF质 谱分析,确定此酶的部分氨基酸序列。此外,我们成功构建了具有完整性的姚虻唾液 腺cDNA文库,获得2×106 pfu个重组子,进一步的筛选了Tablysin的编码基因。 对此酶的溶栓机制进行分析发现,其具有直接水解纤维蛋白原的活性,不具有纤 溶酶原激活剂的活性。此酶能明显地水解纤维蛋白原的α-链,而对β-链的水解活性较 低,不能水解γ链(α > β > γ)。此酶不能有效地水解纤维蛋白,不会水解层粘连蛋白 (Laminin)和纤维粘连蛋白(Fibronectin)等基质蛋白。不具有出血活性。进 一步探讨此酶的作用机制,发现其能以剂量依赖的方式抑制ADP诱导的血小板聚 集。利用流式细胞仪检测到该酶能与血小板膜糖蛋白受体GPIIb / IIIa结合,因此推测其可能是通过抑制血小板和纤维蛋白原结合的作用机制来抑制血小板 的聚集。 我们通过将目的基因连接到pET-32a+载体,并在大肠杆菌表达体系中得到 了高效表达。体外活性检测表明重组蛋白质与天然产物活性相当,用其检测了 其对角叉菜胶致小鼠尾静脉血栓模型体内血栓形成的影响, 实验结果表明 Tablysin具有明显的抗静脉血栓作用。该工作对有潜力开发成为新型单组分抗血栓 药物的姚虻活性蛋白进行了较为系统的理化性质研究,为该分子的进一步开发和研究 奠定了基础。 我们从姚虻唾液腺中分离纯化到一组具有水解纤维蛋白原α-链活性的酶, 命名为TY6家族。对TY6的生化性质进行研究,表明此酶为一种单链的丝氨酸蛋白 酶,其表观分子量为27 kDa,其活性不受金属离子螯合剂EDTA的影响,但能被PMSF 所抑制。Molish反应显示为阴性,说明此酶不是一种糖蛋白。从姚虻唾液腺cDNA 文库中克隆得到其编码序列,发现该酶家族的编码基因在一级结构上表现出丰 富的多样性。通过Western blotting检测到其能与牛虻叮咬后过敏患者血清特异性IgE 结合。推测该基因的多样性是吸血昆虫适应吸血寄生生活而采取的进化策略。 该工作为牛虻唾液腺抗血栓功能基因的继续筛选和分析奠定了基础,并为牛虻的生物 防治提供了思路和对策。 我们从姚虻唾液腺中分离纯化到一个具有水解ADP活性的酶(Apyrase), 命名为Tabapy。活性检测发现Tabapy具有明显的水解ADP的活性,并能以剂量 依赖的方式抑制ADP诱导的血小板聚集。通过Western blotting检测到Tabapy能与 牛虻叮咬后过敏患者血清特异性IgE结合。用PCR方法从姚虻唾液腺cDNA文库中 克隆得到编码序列,经BLAST分析表明,该酶与来源于斑虻唾液腺的血小板聚 集抑制剂Chrysoptin具有90%的序列相似性,并经过Q-TOF分析确证,有5个肽段 的氨基酸残基与该cDNA序列推导的多肽链匹配。该工作为研究吸血节肢动物 吸血生理及进一步研究Apyrase的结构和功能打下了基础。
其他摘要Thrombotic disease (TD) is a lethal medical complication with high incidence. Thromboembolism is the leading cause of human death in China and western countries. During the last few decades, anti-thrombosis agents have been extensively used as the therapeutic treatment of thromboembolism complication, including anti-clotting, anti-platelet, and thrombolytic agents, but more or less side effects are present in each of them. As a result, the pursuit of high quality thrombolytic agent never stopped. Molecular mechanism of interaction between blood-sucking arthropods and vertebrate host are hot spots of international research, which have potential applications in finding new medical leading molecules and providing arthropod control strategy. By proteomics coupling transcriptome analysis with pharmacological testing, three families of proteins, which exert mainly on anti-thrombosis functions, were identified and characterized from salivary glands of the horsefly Tabanus yao Macquart (Diptera, Tabanidae). They are: (I) one bifunctional enzyme Tablysin, which hydrolyzes both alpha and beta chains of fibrinogen, and acts as platelet aggregation inhibitor; (II) another fibrinogenolytic enzymes family TY6, which hydrolyze specially alpha chain of fibrinogen; (III) one apyrase Tabapy, acting as platelet aggregation inhibitor. We studied the diversity, structure, and molecular mechanisms of these three proteins. The extreme diversity of horsefly anti-thrombosis components reveals the anti-thrombosis molecular mechanisms of the traditional Eastern medicine insect material. Tablysin is a single-chain protein, its molecular weight is 27 kDa. The activity of the horsefly fibrinogenolytic enzyme was optimal at pH 2.4-10.0, 37℃. These results suggest that Tablysin is useful in human physical environment. Tablysin is sensitive to serine protease inhibitor PMSF, but the metalloprotease inhibitor EDTA does not affect the activity of Tablysin. Plasma protease inhibitors, including Aprotinin and α-macroglobulin, can partially inhibit the effect on proteolytic activity. These results suggest that Tablysin is a serine protease. Tablysin was subjected to N-terminal amino acid sequence analysis. Unfortunately, its N-terminus was blocked. We obtained the interior peptide amino acid sequences by Q-TOF mass analysis, and its encoding gene was screened from the cDNA library of T. yao salivary glands. Analyzing the hydrolysis of Tablysin showed that it can hydrolyze the α- and β-chains of fibrinogen, but can not hydrolyze fibrin, laminin and fibronectin. Tablysin is either not plasminogen activator. Platelet aggregations induced by agonists ADP were significantly inhibited by Tablysin in dose-dependent manner. Attempts to explain the mechanism of inhibition, we used flow cytometry to detect the effect of Tablysin on the occupancy of platelet glycoprotein GPIIb/IIIa, and we found that tablysin could bind to GPIIb/IIIa. We constructed recombinated Tablysin-pET plasmid successfully, and then using IPTG to induce the His-tagged fusion protein expression. By His-bind resin, enterokinase cleavage, and FPLC, we finally got rTablysin. The rTablysin has the same strong fibrinogenolytic activity as the native Tablysin. The thrombolytic effect of purified rTablysin on carrageenan induced thrombosis model in mice was investigated. The result shows that the tail-thrombus decreased correlate with rTablysin in a dose-dependent manner. A group fibrinogenolytic enzymes, TY6, were purified from T. yao SGE. Complete cDNA sequences encoding TY6 were cloned from the the cDNA library of the salivary glands. In these ten cDNA sequences, most of them encode a proprotein composed of 254 amino acid (aa) residues including predicted signal peptides (16 aa) and mature TY6 containing the SCP domain (Sc7 family of extracellular domain) found in insect antigen 5 proteins. TY6 is a single-chain protein; its molecular weight is 27 kDa. TY6 is sensitive to serine protease inhibitors PMSF, the metalloprotease inhibitor EDTA did not affect the activity of TY6. TY6 was recognized as an allergen, showing IgE reactivity of horsefly sensitized patients by immunoblot. The presence of enzymes diversity with effective fibrinogenolytic activity reveals the mechanism of horsefly blood meal. An Apyrase (ATP diphosphohydrolase, EC 3.6.1.5), named Tabapy, was purified as an active enzyme from horsefly salivary glands and the interior peptides amino acid sequences were sequenced. Its molecular weight is 63 kDa. The amino acid sequences obtained match the conceptual translation product of a cDNA clone isolated from horsefly salivary glands cDNA library. Platelet aggregations induced by agonists ADP were significantly inhibited by Tabapy in dose-dependent manner. Tabapy showed IgE reactivity of horsefly sensitized patients by immunoblot. The apparent conversion of a gene encoding an enzyme involved in a common metabolic event at the cellular level to a gene involved in the antihemostatic response of horsefly illustrates one way that this particular insect has adapted to the challenges of blood-feeding.
语种中文
文献类型学位论文
条目标识符http://ir.kiz.ac.cn/handle/152453/6512
专题科研部门_天然药物功能蛋白质学科组(赖仞)
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马冬莹. 姚虻唾液腺三类抗血栓活性蛋白的结构和功能研究[D]. 北京. 中国科学院研究生院,2010.
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