摘要
诺如病毒（Norovirus, NoV）是全球非细菌性腹泻的主要病因。1968年NoV被首次报道，至上世纪90年代中期GⅡ.4型NoV成为了全球主要流行株，并且此后每隔两三年就产生变异株并引起新一轮传播，因此也被成为“肠胃炎流感”。近年来我国NoV疫情频繁暴发，对抗病毒研究提出了更紧迫的要求，然而NoV体外复制体系的缺乏增加了病毒研究的困难。检测技术的发展，尤其是PCR技术的应用，对NoV传播特点与分布规律研究具有重要意义。同时，PCR技术还促进了NoV序列的积累，这不但有助于了解病毒基因组特征，而且加深对其进化机制的认识。近年来有学者提出，宿主对病毒感染的的免疫保护作用以及人类血型组织抗原作为病毒受体是NoV进化的两大约束性因素。
NoV序列的积累已经成为了病毒进化及抗病毒研究的基础，因此本论文以病毒序列为中心，分别针对检测技术开发、病毒污染调查、基因组序列克隆、病毒序列进化分析等内容展开研究，具体工作如下：
1、NoV检测技术标准化研究。在构建通用性DNA/RNA质控品的基础上，优化并评价了不同引物、不同类型的RT-PCR检测NoV技术，建立了各种方法检测灵敏度的对应关系，为实际工作中选择合适的检测方法提供了参考依据。此外，还比较了不同方法抽提病毒核酸的差异，其中膜纯化方法效果最佳，同时考察了膜层析、超滤浓缩及免疫板捕获病毒等前处理方法回收病毒的效果。
2、广州地区冬季散发性腹泻中病毒感染调查。调查了广州地区冬季散发性腹泻样本中的病毒感染状况，从89份样本中检出9份NoV阳性和32份轮状病毒阳性。对病毒部分聚合酶区及衣壳蛋白区的序列分析显示，所有NoV均为GⅡ基因型，其中GⅡ.4型占5/9，GⅡ.6型占3/9，另外还包括一株GⅡ.b/GⅡ.3重组株。
3、GⅡ型NoV基因组序列克隆与特征分析。建立了一种简单通用的GⅡ型NoV基因组克隆方法，并克隆得到3条我国华南地区不同型别NoV流行株的基因组序列。同时，收集已公布的GⅡ型NoV基因组，分析其不同蛋白编码区的同源性及进化特征差异，结果显示p22蛋白、P2区和VP2蛋白为易变异区域，进一步对以上区域进行系统发育分析，提示VP1与VP2蛋白可能存在共进化现象。
4、GⅡ.4型NoV衣壳蛋白进化分析。收集已公布的GⅡ.4型NoV衣壳蛋白序列，通过多重序列比对筛选病毒进化中出现的信息位点，分析其中的关键位点及其在病毒衣壳上的分布，然后借助同源建模比较不同GⅡ.4型NoV流行株P蛋白的结构差异，计算关键位点的表面积变化情况，从而分析了各位点的氨基酸使用范围，并预测衣壳蛋白上可能的抗原表位。
病毒检测技术开发和序列信息积累是NoV研究的重要工具，因此，本论文围绕NoV序列展开研究，建立了标准的RT-PCR检测NoV技术，并应用于临床样本调查，同时克隆、分析了GⅡ型NoV的基因组序列特征，最后对主要流行GⅡ.4型NoV衣壳蛋白序列的进化机制进行探索。NoV高流行性和易变异的特征对认识病毒提出了迫切的需求，本论文关于病毒检测及进化机制的研究不仅有助于预测NoV的进化趋势，也将为开发有效的病毒疫苗和治疗手段提供重要的理论指导。
Noroviruses (NoVs) are regarded as the major cause for acute non-bacterial diarrhea.NoVs was first reported in1968, and GⅡ.4NoV became the pandemic strain all over theworld in the mid-1990s. In the new century, new variant strains appeared every two or threeyears and caused large epidemics, which were known as “Gastroenteritis flu”. In recent years,NoV outbreaks became more frequent in China, which proposed a more urgent requirementon anti-NoV studies. But currently there is no suitable in vitro replication system and infectedmodel, increasing the difficulty of NoV researches. The development of virus detectiontechnology, especially the application of PCR in NoV detection, promoted the developmentfor understanding NoV propagation and its distribution. Meanwhile, it’s also contributed tothe accumulation of NoV genome sequences using PCR technology, which not only helped tounderstand the characteristics of NoV genome, but also promoted to study the mechanisms ofNoV evolution. A hypothesis was put forward recently that the host immune response to viralinfection and human blood group antigens as binding receptors were the two key factors forNoV evolution.
The accumulation of NoV sequences is the an important tool for studies on mechanismof NoV evolution, so this thesis focuses on the development of NoV detection technology, thesurvey NoV infection for sporadic diarrhea, genome sequences obtained from the NoV strainsdetected during the survey, and phylogenetic analysis of viral sequences. Specific contents asfollows:
1. Technology standardization of NoV detection. Based on the constructed universalDNA/RNA quality control, different types of RT-PCR methods with different primers forNoV detection were optimized and evaluated, and the correspondence of the detection limitsof these methods were also established, which could be used for selection of appropriateddetection methods in actual work as a reference. Besides, different methods for NoV RNAextraction were compared, of which the membrane purification kit showed the best quality;meanwhile, the viral recovery efficiency of different pre-treatment methods, which based onmembrane chromatography, ultrafiltration,and EIA plate, respectively, were also evaluated.
2. Survey of NoVs associated with sporadic gastroenteritis during winter in Guangzhou.The genotypes of circulating norovirus strains associated with sporadic diarrhea cases inGuangzhou from November2010to January2011were characterized. Among fecalspecimens collected from89patients with acute diarrhea,9samples (10.11%) werenorovirus-positive and32samples (35.96%) were rotavirus-positive. The partial polymerase and the capsid regions of these norovirus samples were sequenced and phylogeneticallyanalyzed. Three genotypes (GⅡ.4, GⅡ.6, and GⅡ.b/GⅡ.3) were identified, among which GⅡ.4was the most predominant genotype (5/9), followed by GⅡ.6(3/9). A GⅡ.b/GⅡ.3recombinantstrain was also detected.
3. Clone and characterization of GⅡ NoV genomes. A simple and novel method for GⅡNoV genome clone with generic primers was established, and three genome sequences ofNoV epidemic strains in South China were obtained. Meanwhile, based on the GⅡ NoVgenome sequences collected from GenBank, the homology and phylogenetic diversity ofdifferent viral protein among NoV genome was analyzed, which showed p22, P2, and VP2asthe most variable regions, and coevolutionary phenomena of VP1and VP2may existed.
4. Evolutionary analysis of GⅡ.4NoV capsid protein. All available GⅡ.4NoV capsidprotein VP1sequences were collected from GenBank. Based on multiple sequence alignment,informative sites on the viral capsid were screened and their locations on viral capsid werealso analyzed. Then, the structural differences of P protein of different GⅡ.4NoV epidemicstrains were compared based on homology modeling, and the variation of surface area of eachkey informative site was also calculated to understand the rules of amino acid usage. Finally,six epitopes located on viral capsid were predicted.
Virus detection and sequence accumulation are two important tools for NoV research.So, our study was designed around this central topic, including the development of virusdetection, the survey of virus infection, clone and analysis of GⅡ NoV genome sequences,and the evolutionary mechanisms of GⅡ.4NoV capsid. High prevalence and easy-variationcharacteristics raised an urgent claim for NoV researches, and the results of this study wouldhelp to predict NoV evolutionary trend and develop effective vaccines and therapeuticmethods.
引文
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