CN113564133B - 柯萨奇病毒a16型毒株及其免疫原性组合物和应用 - Google Patents
柯萨奇病毒a16型毒株及其免疫原性组合物和应用 Download PDFInfo
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Abstract
本发明涉及生物技术领域,具体涉及柯萨奇病毒A16型毒株及其免疫原性组合物和应用。本发明提供一种柯萨奇病毒A16型毒株,其含有P1结构蛋白以及非结构蛋白2A、2B、2C、3A、3B、3C和3D;其中,P1结构蛋白的氨基酸序列如SEQ ID NO.1所示;非结构蛋白2A、2B、2C、3A、3B、3C和3D的氨基酸序列分别如SEQ ID NO.4‑10所示。该毒株的免疫原性好、交叉中和能力强、稳定性好,可用于制备预防由柯萨奇病毒A16型引起的手足口病的疫苗,利用该毒株制备的疫苗可预防由柯萨奇病毒A16型引起的手足口病,具有较好的免疫效果。
Description
技术领域
本发明涉及生物技术领域,具体涉及柯萨奇病毒A16型毒株及其免疫原性组合物和应用。
背景技术
手足口病(Hand foot and mouth disease,HFMD)是由小RNA病毒科肠道病毒属的肠道病毒(Enterovirus,EV)感染引起的一种儿童常见传染病,5岁以下儿童多发,典型表现为手、足、口腔等部位有皮疹、疱疹、溃疡等,可并发无菌性脑膜炎、脑炎、急性弛缓性麻痹、呼吸道感染和心肌炎等,个别重症会导致残疾或死亡,是一种全球性疾病。
手足口病的主要致病血清型包括柯萨奇病毒(Coxsackievirus,CV)A组4~7、9、10、16型和B组1~3、5型,埃可病毒(Echovirus)的部分血清型和肠道病毒71型(Enterovirus A71,EV-A71)等,其中以CV-A16和EV-A71最为常见,重症及死亡病例多由EV-A71所致。近年部分地区CV-A6、CV-A10感染有增多趋势。肠道病毒各型之间无交叉免疫力。
与EV71感染相比,柯萨奇病毒A16型(CA16)感染引起的手足口病虽然症状相对较轻,但具有更强的传播力,而目前上市的疫苗皆为针对EV71病毒的疫苗,尚没有保护CA16病毒感染的疫苗,而EV71疫苗对CA16并无交叉保护作用。因此,研发CA16疫苗对于控制手足口病的爆发具有重大的意义。
病毒毒株的免疫原性、稳定性、交叉中和能力是疫苗株筛选的重要指标。
CA16的基因组由7000多个核苷酸组成,为单股正链RNA病毒。RNA病毒因其RNA聚合酶的低保真度和频繁的重组现象(Zhao, K. (2011).Circulating Coxsackievirus A16Identified as Recombinant Type A Human Enterovirus, China. EmergingInfectious Diseases.doi:10.3201/eid1708.101719),导致核苷酸易发生突变,据文献报道(Zhang, Y., et al. (2009). Molecular Evidence of Persistent Epidemic andEvolution of Subgenotype B1 Coxsackievirus A16-Associated Hand, Foot, andMouth Disease in China. Journal of Clinical Microbiology, 48(2), 619–622.doi:10.1128/jcm.02338-09),其年变异速度可达0.91×10-2/核苷酸。CA16的结构蛋白P1由VP1、VP2、VP3和VP4组成,其中,VP4位于衣壳内部;VP1、VP2和VP3位于病毒颗粒的表面,是抗原决定簇的主要区域。研究表明,CA16的免疫原性对抗原结构的依赖性较大(Chong, P., etal. (2012). Immunological and Biochemical Characterization of Coxsackie VirusA16 Viral Particles. PLoS ONE, 7(11), e49973. doi:10.1371/journal.pone.0049973;卢佳,张林林,安泰学,毛群颖,李鹏飞,张夫坤,王泽鋆,李黎,周辉,王继麟,陈晓琦,梁争论,申硕.柯萨奇病毒A16疫苗候选株免疫效果评价[J].中国病毒病杂志,2018,8(06):487-494.)。因此,除了关注常规的抗原表位主要区域VP1的序列外,结构蛋白P1其它序列的改变也可能引起CA16结构及免疫原性的改变,而目前并未发现关于CA16的P1序列稳定性的的研究及报道。
不同基因型、不同亚型、甚至同一亚型不同来源的毒株间抗原性也具有一定差异,这会导致不同毒株的交叉保护水平可能存在差异。因此,CA16毒株的交叉保护能力是疫苗毒株筛选的另一难点。Yang等研究表明CA16毒株(G20,B基因型)制备的疫苗,小鼠免疫血清对同基因型毒株的中和效价≤1:256,对A基因型(G10)则中和效价< 1:16(Yang, E., etal. (2014). Comparative study of the immunogenicity in mice and monkeys of aninactivated CA16 vaccine made from a human diploid cell line. Human Vaccines& Immunotherapeutics, 10(5), 1266–1273.doi:10.4161/hv.28083)。谢等研究表明CA16毒株间具有较好的交叉活性,但其选择的4个毒株皆为B基因型(谢振锋,普晶,黄泓泰,刘正玲,董承红,刘龙丁,王丽春.柯萨奇病毒A组16型免疫原性及毒株间交叉保护能力的比较分析[J].中国生物制品学杂志,2013,26(10):1366-1370+1375.),而卢等的研究则表明不同基因型之间,甚至亚基因型之间可能仅存在较低的交叉中和活性或单侧交叉(卢佳,张林林,安泰学,毛群颖,李鹏飞,张夫坤,王泽鋆,李黎,周辉,王继麟,陈晓琦,梁争论,申硕.柯萨奇病毒A16疫苗候选株免疫效果评价[J].中国病毒病杂志,2018,8(06):487-494),这与Yang等研究结果是一致的,因此目前的研究报道中并未筛选到对各基因型、基因亚型毒株皆有较好的中和交叉能力的CA16毒株。
发明内容
本发明的第一目的是提供一种柯萨奇病毒A16型毒株。
本发明的第二目的是提供与该柯萨奇病毒A16型毒株相关的生物材料。
本发明的第三目的是提供含有该柯萨奇病毒A16型毒株或其相关生物材料的免疫原性组合物以及疫苗、药物等产品。
本发明的第四目的是提供上述毒株、生物材料、免疫组合物的用途。
具体地,本发明提供以下技术方案:
首先,本发明提供一种柯萨奇病毒A16型毒株,其含有P1结构蛋白以及非结构蛋白2A、2B、2C、3A、3B、3C和3D;其中,P1结构蛋白的氨基酸序列如SEQ ID NO.1所示;非结构蛋白2A、2B、2C、3A、3B、3C和3D的氨基酸序列分别如SEQ ID NO.4-10所示。
具体地,本发明提供的柯萨奇病毒A16型毒株的基因组编码序列如SEQ ID NO.1所示的P1结构蛋白以及序列如SEQ ID NO.4-10所示的非结构蛋白。
本发明提供序列如SEQ ID NO.1所示的P1结构蛋白与序列如SEQ ID NO.4-10所示的非结构蛋白组成的融合蛋白。
优选地,所述柯萨奇病毒A16型毒株的基因组中,P1结构蛋白的编码基因序列如SEQ ID NO.2所示,非结构蛋白2A、2B、2C、3A、3B、3C和3D的编码基因序列分别如SEQ IDNO.11-17所示。
以上所述的结构蛋白和非结构蛋白的编码基因在所述柯萨奇病毒A16型毒株的基因组上的排列顺序为(5’-3’):P1结构蛋白编码基因、2A蛋白编码基因、2B蛋白编码基因、2C蛋白编码基因、3A蛋白编码基因、3B蛋白编码基因、3C蛋白编码基因、3D蛋白编码基因。
本发明提供由如SEQ ID NO.2所示的基因以及如SEQ ID NO.11-17所示的基因顺次连接组成的重组核酸分子。
以上所述的柯萨奇病毒A16型毒株的基因组序列还包括5’-UTR序列和3’-UTR序列,5’-UTR序列如SEQ ID NO.18所示,3’-UTR序列如SEQ ID NO.19所示。
进一步优选地,所述柯萨奇病毒A16型毒株的基因组序列如SEQ ID NO.3所示或如SEQ ID NO.3所示序列的互补序列所示。
具体地,本发明提供柯萨奇病毒A16型毒株V00080265,该毒株已于2021年7月13日保藏在中国微生物菌种保藏管理委员会普通微生物中心(简称CGMCC,地址:北京市朝阳区北辰西路1号院3号,中国科学院微生物研究所,邮编100101),分类命名为柯萨奇病毒A16型,保藏编号为CGMCC No.20387。
柯萨奇病毒A16型毒株V00080265的基因组编码序列如SEQ ID NO.1所示的P1结构蛋白以及序列如SEQ ID NO.4-10所示的非结构蛋白2A、2B、2C、3A、3B、3C和3D,其基因组序列如SEQ ID NO.3所示。
本发明提供的上述柯萨奇病毒A16型毒株的免疫原性好、交叉中和能力强、稳定性好,能够以Vero等细胞作为基质细胞进行快速繁殖。
进一步地,本发明提供与所述柯萨奇病毒A16型毒株相关的生物材料,其为以下(1)-(6)中的任意一种:
(1)序列如SEQ ID NO.3所示或如SEQ ID NO.3所示序列的互补序列所示的核酸分子;
(2)含有(1)中所述核酸分子的表达盒;
(3)含有(1)中所述核酸分子的重组载体;
(4)含有(1)中所述核酸分子的重组微生物;
(5)含有(1)中所述核酸分子的细胞系;
(6)检测(1)中所述核酸分子的引物或探针。
以上(1)中所述的核酸分子可为DNA分子或RNA分子。
以上(2)中所述的表达盒为在(1)中所述的核酸分子的上游、下游连接用于转录、翻译的调控元件得到的重组核酸分子。
以上(3)中所述的重组载体为携带(1)中所述的核酸分子且能够在宿主细胞中复制或整合的质粒载体、病毒载体、噬菌体载体或转座子。
以上(4)中所述的微生物可为细菌或病毒。
以上(5)中所述的细胞系为动物细胞系。所述动物细胞系为不可繁殖为动物个体的动物细胞系,可为用于病毒培养的常用动物细胞系,包括但不限于RD细胞、非洲绿猴肾传代细胞(Vero)等。
以上(6)中所述的引物、探针为能够与(1)中所述核酸分子结合并进行PCR扩增的寡核苷酸。
本发明提供所述柯萨奇病毒A16型毒株的病毒样颗粒(VLPs),其含有P1结构蛋白以及选自非结构蛋白2A、2B、2C、3A、3B、3C和3D中的任意一种或多种;其中,所述P1结构蛋白具有如SEQ ID NO.1所示的序列,所述非结构蛋白2A、2B、2C、3A、3B、3C和3D分别具有如SEQID NO.4-10所示的序列。
以上所述的病毒样颗粒可采用昆虫载体系统表达上述结构蛋白、非结构蛋白的编码基因。
本发明提供含有以上所述的柯萨奇病毒A16型毒株的免疫原性组合物。
本发明还提供含有以上所述的生物材料的免疫原性组合物。
本发明还提供含有以上所述的病毒样颗粒的免疫原性组合物。
所述免疫原性组合物中除含有所述柯萨奇病毒A16型毒株、生物材料或病毒样颗粒外,还可含有有利于柯萨奇病毒A16型毒株发挥免疫原性的佐剂。所述佐剂包括但不限于铝佐剂。
所述免疫组合物中的柯萨奇病毒A16型毒株被灭活。
用于灭活病毒的灭活剂可以为甲醛或β-丙内酯。
优选地,所述免疫原性组合物中,柯萨奇病毒A16型毒株的蛋白浓度为1-5μg/ml,铝佐剂含量为0.5-1.5μg/ml。所述铝佐剂可以为氢氧化铝或磷酸铝佐剂。
进一步地,本发明提供所述柯萨奇病毒A16型毒株或所述生物材料或所述病毒样颗粒或所述免疫原性组合物的如下任意一种应用:
(1)在制备预防和/或治疗柯萨奇病毒引起疾病的疫苗中的应用;
(2)在制备预防和/或治疗柯萨奇病毒引起疾病的药物中的应用;
(3)在制备预防和/或治疗柯萨奇病毒引起疾病的抗体中的应用;
(4)在制备预防和/或治疗柯萨奇病毒引起疾病的抗血清中的应用;
(5)在制备诊断柯萨奇病毒感染的试剂或试剂盒中的应用;
(6)在柯萨奇病毒的流行病学调查中的应用;
(7)在柯萨奇病毒疫苗的免疫原性评价中的应用;
(8)在柯萨奇病毒疫苗的保护性评价中的应用;
(9)在制备柯萨奇病毒感染动物模型中的应用;
(10)在预防和/或治疗柯萨奇病毒引起疾病的药物的筛选或药效评价中的应用。
以上(1)-(10)中,所述柯萨奇病毒优选为柯萨奇病毒A16型毒株。
以上(1)-(4)和(10)中,所述柯萨奇病毒引起疾病优选为手足口病。
以上(9)中,所述的动物模型优选为鼠模型。
本发明提供一种抗体或抗血清,其为以所述柯萨奇病毒A16型毒株或所述生物材料或所述病毒样颗粒或所述免疫原性组合物为免疫原制备得到。
本发明还提供一种抗体或抗血清的制备方法,其包括:以所述柯萨奇病毒A16型毒株、所述生物材料或所述病毒样颗粒为免疫原免疫动物,经分离获得抗柯萨奇病毒A16型的抗体或抗血清。
本发明提供一种产品,其含有以下(1)-(5)中的任意一种或多种的组合:
(1)所述柯萨奇病毒A16型毒株;
(2)所述生物材料;
(3)所述病毒样颗粒;
(4)所述免疫原性组合物;
(5)所述柯萨奇病毒A16型毒株的抗体或抗血清。
以上所述的产品优选为用于诊断、预防或治疗柯萨奇病毒A16型感染的产品,或者为用于柯萨奇病毒A16型疫苗免疫原性或保护性评价的产品,或者为用于柯萨奇病毒A16型感染动物模型构建的产品。其中,用于诊断柯萨奇病毒A16型感染的产品可为诊断试剂或试剂盒;用于预防或治疗柯萨奇病毒A16型感染的产品可为疫苗或药物。
作为一种实施方式,本发明提供预防柯萨奇病毒A16型感染的疫苗,其含有所述柯萨奇病毒A16型毒株。
本发明所述的疫苗可为全病毒灭活疫苗、减毒活疫苗、核酸疫苗、基因工程疫苗(亚单位疫苗、活载体疫苗、基因重组疫苗等)。
优选地,所述疫苗为全病毒灭活疫苗,其中所述柯萨奇病毒A16型毒株被灭活。所述疫苗还可含有佐剂,所述佐剂包括但不限于铝佐剂。
本发明还提供以上所述的疫苗的制备方法,所述方法包括:将所述柯萨奇病毒A16型毒株在细胞上培养,收获病毒液,将收获的病毒液经灭活、纯化后获得疫苗原液。
所述疫苗的制备方法还包括:将所述疫苗原液与佐剂混合。所述佐剂优选为铝佐剂。
优选地,所述疫苗中,柯萨奇病毒A16型的蛋白浓度为1-5μg/ml。
优选地,所述疫苗中,铝佐剂浓度为0.5-1.5μg/ml。
作为本发明的一种优选方案,所述疫苗的制备方法包括:将所述柯萨奇病毒A16毒株在Vero细胞上进行扩大培养,按MOI=0.0001-0.1接种病毒,待病变达 +++~++++时,收获病毒液,经离心,灭活、纯化等步骤后,按蛋白浓度为1μg~5μg/ml与铝佐剂浓度为0.5~1.5μg/ml混合,制备得到灭活疫苗。
以上所述的方法中,扩大培养方式可采用细胞工厂或生物反应器中进行。
以上所述的方法中,灭活方式可采用与病毒液的体积比为1:1000~1:5000的甲醛灭活2~6天,或者采用与病毒液的体积比为1:1000~1:4000的β-丙内酯于5±3℃灭活1~3天,然后37℃水解1~4小时。
作为另一种实施方式,本发明提供治疗柯萨奇病毒A16型感染的药物,其含有所述柯萨奇病毒A16型毒株的抗体或抗血清。
本发明还提供以上所述的产品在预防或治疗柯萨奇病毒A16型感染或柯萨奇病毒A16型感染引起疾病中的应用。
本发明的有益效果在于:本发明提供了一株免疫原性好(具有较高的血清中和效价)、交叉中和能力强(与柯萨奇病毒A16型的不同基因型、不同亚型均具有较好的交叉保护力)、稳定性好(具有较高的遗传稳定性、病毒滴度稳定性和免疫原性稳定性)、Vero细胞适应的柯萨奇病毒A16型毒株,该毒株可用于制备预防柯萨奇病毒A16型感染或柯萨奇病毒A16型引起疾病的疫苗。利用该毒株制备的疫苗可预防柯萨奇病毒A16型感染或柯萨奇病毒A16型引起的手足口病,具有较好的免疫效果。
附图说明
图1为本发明实施例3中不同代次病毒滴度的检测结果。
图2为本发明实施例3中不同代次毒株动物免疫后血清的中和效价检测结果。
图3为本发明实施例4中CA16免疫血清对各基因型毒株的交叉中和效价检测结果。
具体实施方式
本发明中术语的解释:
MOI(Multiplicity of Infection,感染复数)是指病毒感染细胞的比例。选择合适的MOI值,病毒的感染效率以及目的蛋白的表达量越高,相对细胞的毒性越小。
细胞病变效应(cytopathic effect):在体外实验中,通过细胞培养和接种杀细胞性病毒,经过一定时间后,可用显微镜观察到细胞变圆、坏死、从瓶壁脱落等现象,称之细胞病变作用,简写CPE。CPE 程度常用+~++++表示:1+,<25%;2+,25%-50%;3+, 50%-75%; 4+,75%-100%。
本发明在临床手足口患者的标本中分离获得了一株柯萨奇病毒A16型毒株,将该毒株命名为V00080265。该毒株是一株Vero细胞适应株,具有较高的免疫原性、较强的交叉中和能力,同时具有较高的稳定性。
本发明中,柯萨奇病毒A16型毒株(CA16)的筛选和性能验证过程大致概括如下:将临床手足口病患者的标本经过处理,接种于非洲绿猴肾传代细胞(Vero)上,盲传三代,经鉴定、测序、动物免疫等初步筛选与评价后,经3次蚀斑纯化,得到CA16 纯化毒株,将CA16毒株按适应的MOI进行连续传代,对不同代次进行免疫原性检测及基因序列检测,同时对该毒株的小鼠免疫血清进行基因型内与型间的交叉中和能力评价,并选用Vero细胞进行CA16毒株培养,收获病毒液、灭活、纯化后,得到疫苗原液,制备CA16疫苗,进行小鼠动物免疫。
以下实施例用于说明本发明,但不用来限制本发明的范围。
实施例1 CA16毒株的分离及培养
(一)临床样本的处理
1、粪便标本和肛拭子的处理:
(1)在离心管上标记标本号;
(2)每管中加入10ml磷酸盐缓冲溶液(PBS)、1g玻璃珠、1ml氯仿;
(3)在生物安全柜中将每一份粪便标本取大约2g加入标记好的离心管中(确保离心管上的标号与原始标本的标号一致);肛拭子为2ml;
(4)剩余的原始标本最好留在原容器中,冻存于-20℃;
(5)确保拧紧离心管,用机械震荡器剧烈震荡20min;
(6)在确保离心机的盖子盖好和离心桶密封的情况下,用冷冻离心机在1500 g条件下离心20min;
(7)在生物安全柜中将每1份标本的上清液分别吸入2个有外螺旋盖的冻存管中(如果上清液不清澈,应再用氯仿处理1次);
(8)1管粪便悬液冻存于-20℃作为备份,另1管存于4-8℃以备接种。
(2)咽拭子标本的处理
咽拭子要在标本运输(保存)液中充分搅动(至少40下),以洗下拭子上粘附的病毒及含有病毒的细胞等,用于病毒分离时,需要冻融一次(防止多次冻融),使细胞破裂,释放病毒颗粒。然后在2~8℃条件下,10000 rpm离心20min,用上清接种到细胞上。如果发现有细菌污染,需用滤器过滤除菌。
(3)病毒分离
将处理好的样品,按一定比例接种至80-90%密度的健康无污染非洲绿猴肾传代细胞(Vero)中,35℃、5%CO2培养箱中吸附0.5-1.5小时,补液至培养体积,35℃、5%CO2培养箱中培养,同时设置未加样本的细胞对照。
使用倒置显微镜每天观察细胞,如出现肠道病毒致细胞病变效应(CPE):细胞变圆,折光增强并脱离管壁等,记录其变化,连续观察7天内接种孔和对照孔细胞所发生的变化CPE(1+~4+)。
如CPE出现,观察到75%的细胞发生变化(3+),收获培养液并储藏在-20℃以下冰箱,冻融3次后,2000 rpm、4℃离心10 min,进行下一次传代,连续培养3代后,将第4代病毒液冻存。如7d后没有CPE出现,则继续盲传3代,如仍未出现CPE,则判定为阴性。
经上述分离、培养以及病毒的免疫原性、交叉中和能力和稳定性的筛选,最终筛选得到一株免疫原性好、交叉中和能力强、且稳定性好的柯萨奇病毒A16型毒株V00080265,该毒株已于2021年7月13日保藏在中国微生物菌种保藏管理委员会普通微生物中心(简称CGMCC,地址:北京市朝阳区北辰西路1号院3号,中国科学院微生物研究所,邮编100101),分类命名为柯萨奇病毒A16型,保藏编号为CGMCC No.20387。
实施例2 CA16毒株的鉴定及检测
将实施例1中收获的柯萨奇病毒A16型毒株V00080265(以下简称CA16毒株)的第4代病毒液进行分子学鉴定、基因组测序、滴度检测及免疫原性检测,具体如下:
1、分子鉴定及基因组测序
(1)采用RT-PCR法鉴定病毒,肠道病毒核酸检测通用引物及VP1特异性引物如表1所示。表1所示的引物中,Y代表C/T,M代表A/C,W代表A/T。
表1 肠道病毒核酸检测通用引物及VP1特异性引物
(2)病毒核酸提取
取第4代病毒液,按照说明书添加试剂和病毒样品,然后置于核酸提取仪中,按照预设的程序抽提核酸,将提取好的核酸保存至-70℃冰箱。
(3)HEV- 5UTR通用引物PCR检测
应用HEV-5UTR的两对通用引物59F/588R与153F/541R进行巢式PCR。
第1轮PCR扩增反应体系组成如表2所示。
表2 第1轮PCR扩增反应体系
反应程序如下:
55℃,30 min; 94℃,2 min;
20个循环:94℃,15 s;55℃,30 s; 68℃,40 s;
68℃延伸5 min 。
以第1轮PCR扩增产物为模板,进行第2轮PCR扩增,PCR扩增反应体系组成如表3所示。
表3 第2轮PCR扩增反应体系
反应程序如下:
94℃,2 min;
30个循环:94℃,15 s;55℃,30 s;72℃,40 s;
72℃延伸5 min。
(4)CV-A16 VP1特异性序列的扩增
应用CV16-VP1特异性引物进行VP1的扩增,反应体系组成如表4所示。
表4 VP1的PCR扩增反应体系
反应程序如下:
55℃,30 min; 94℃,2 min;
30个循环:94℃,15 s;55℃,30 s;68℃,70 s;
68℃延伸5 min。
(5)结果判断
将PCR扩增的产物进行琼脂糖凝胶电泳检测,HEV-5UTR通用引物扩增产物目的条带大小为400bp,CV16-VP1特异性条带大小为1kb,具体判定标准如表5所示,其中,EV(-)代表HEV-5UTR通用引物未扩增出目的条带,EV(+)代表HEV-5UTR通用引物扩增出目的条带;CV-A16 VP1(-)代表CV16-VP1特异性引物未扩增出目的条带,CV-A16 VP1(+)代表CV16-VP1特异性引物扩增出目的条带。
表5 PCR扩增产物结果判断标准
取鉴定为CV-A16阳性的样本的第4代病毒液进行基因组测序,并根据VP1序列分型。经全基因组测序,柯萨奇病毒A16型毒株V00080265的基因组序列如SEQ ID NO.3所示,基因型为B1a型,其中,P1蛋白的编码基因序列如SEQ ID NO.2所示,P1蛋白的氨基酸序列如SEQ ID NO.1所示,非结构蛋白2A、2B、2C、3A、3B、3C和3D的氨基酸序列如SEQ ID NO.4-10所示,非结构蛋白2A、2B、2C、3A、3B、3C和3D的编码基因序列分别如SEQ ID NO.11-17所示。
2、滴度检测
取第4代病毒液,10倍梯度稀释,从10-1稀释至10-8。将稀释好的病毒加入96孔培养板,8孔/稀释度,0.1ml/孔。同时每孔加入100μl Vero细胞悬液(1×105个/ml)。再另外取8~16孔加入细胞悬液,0.1ml/孔,补加病毒维持液0.1ml/孔,作为细胞对照。加盖封板,轻轻拍打混匀,置于35℃、5%CO2培养箱中静置培养,第7天判断结果,每个样品进行3次重复检测。
病毒滴度计算:按Behrens-Karber公式计算LgCCID50。
Lg CCID50 = L-d (S-0.5),其中:
L = 实验中使用的病毒的最低稀释度的对数值;
d = 稀释梯度的对数值;
S = 终判时阳性部分的总和(即出现CPE的细胞孔所占的比例之和)。
具体参见中华人民共和国卫生部.手足口病预防控制指南(2009版).[EB/OL].(2009-06-04) http://www.gov.cn/gzdt/2009-06/04/content_ 1332078.htm。
经检测,柯萨奇病毒A16型毒株V00080265的病毒滴度结果为5.5 LgCCID50/ml。
3、免疫原性检测
取第4代病毒液,进行动物免疫。按病毒滴度5.5 LgCCID50/ml,免疫10只小鼠(NIH,SPF,雌性,18~22g),0、14天两针免疫,腹腔注射0.5ml/只,第14、28天采血,同时免疫病毒维持液作为对照。
采用微量细胞病变法检测血清中和抗体,具体方法如下:将血清56℃,30min 水浴灭活后,从1:8开始稀释,加入96孔板,每个样品2孔,100μl/孔,2倍倍比稀释后,加入 32~320CCID50/0.05ml的病毒液,36±1℃,5% CO2培养箱,中和1~2小时。加入RD细胞悬液(1~2×105个/ml),100μl/孔。置于5% CO2培养7天,判定结果,中和效价<8 判为阴性,≥8 为阳性。结果显示,柯萨奇病毒A16型毒株V00080265的病毒液一免后14天,100%阳转,二免疫后14天,血清中和效价GMTs可达1:645;对照组的阳转率为0%,血清中和效价GMTs<1:8。
实施例3 CA16毒株的稳定性检测
1、遗传稳定性
将进行3次蚀斑纯化后的柯萨奇病毒A16型毒株V00080265,按相同MOI进行连续传代。具体方法如下:将代次≤147的健康无污染的Vero 细胞,接种于细胞培养瓶中,待生长至90%左右密度,按 MOI=0.0001~0.1接种病毒液,待病变达 +++~++++ 时,收获病毒液,于-60℃及以下保存。每收获一代,代次增加一代。将收获的病毒液采用同样的方法,连续传代至P15,取各代病毒收获液进行P1基因序列的扩增与测序。
P1 基因序列扩增引物如下,目的片段大小约3Kb,扩增反应体系如表6所示。
SEQ ID NO.26:CVA16-P1-F1 (5'-3'): GTTACCATATAGCTATTGGATTGGCCATCC;
SEQ ID NO.27:CVA16-P1-R (5'-3'): CTAGAGCTGTCCTCCCACACAAGGTTTGCC。
表6 P1基因的PCR扩增反应体系
反应程序如下:
55℃,30 min; 94℃,2 min;
30个循环:94℃,15 s;55℃,30 s;68℃,3 min;
68℃延伸5 min。
将扩增的目的条带送检测序。测序结果显示,实施例1中蚀斑纯化前与蚀斑纯化后连续传代至P15的各代次P1序列100%一致,结果表明该毒株具有较高的遗传稳定性。
2、滴度稳定性
将上述1中的各代柯萨奇病毒A16型毒株V00080265进行滴度检测,方法同实施例2的滴度检测,结果如图1所示。
结果表明,柯萨奇病毒A16型毒株V00080265不同代次的滴度稳定,P2~P15各代次的滴度值为5.7~6.0 LgCCID50/ml,结果表明该毒株具有较高的滴度稳定性。
3、免疫原性稳定性
取上述1中P5、P10、P15代次的柯萨奇病毒A16型毒株V00080265进行动物免疫,检测毒株的免疫原性。动物免疫方法及中和效价检测方法同实施例2的免疫原性检测。结果如图2所示,P5、P10、P15的动物免疫结果中和效价为1:527~1:1122,无统计学差异(P>0.05),不同代次间免疫原性一致。
实施例4 CA16毒株的交叉中和能力检测
将柯萨奇病毒A16型毒株V00080265的免疫血清(以下简称CA16免疫血清)分别与12株不同基因型、不同亚型的CV-A16病毒株进行交叉中和检测,这12株毒株包括A、B、C三种基因型,其中3株A基因型(A2-1、A2-2、A各1株),8株B基因型(2株B1a以及B1、B-1、B-2、B-3、B1b、B-4各1株,1株C基因型。具体方法如下:试验前,先测定12株毒株的滴度,使每株病毒稀释适宜倍数后,病毒稀释能够准确落在32~320CCID50/0.05ml内。将柯萨奇病毒A16型毒株V00080265的免疫血清样品(参见实施例2的3中的免疫方法)于56℃,30min 水浴灭活后,从1:8开始稀释,2倍倍比稀释后,将12株毒株分别稀释至100CCID50/0.05ml,加入病毒液50μl/孔,于36±1℃、5% CO2培养箱,中和2小时后,加入RD细胞悬液(1-2×105个/ml),100μl/孔,置于5% CO2培养7天判定结果,中和效价<8 判为阴性,≥8 为阳性,同时计算最大中和抗体效价(MAX)和最小中和抗体效价(MIN)的比值(交叉中和检测委托中国食品药品检定研究院进行,用于检测的12株毒株均来自中国食品药品检定研究院并用于测试)。结果显示,CA16免疫血清对各基因型、基因亚型毒株皆具有较好的中和能力(图3和表7),且中和抗体效价GMTs(MAX)(1:1024)/MIN(1:128)=8,差异较小。以上结果表明,柯萨奇病毒A16型毒株V00080265对现有的柯萨奇病毒A16型的A、B、C三种基因型具有较为均一的交叉中和检测能力。
表7 CA16毒株的交叉中和能力检测
实施例5 CA16毒株三级种子库的建立及检定
按照《中国药典》(2020版)中生物制品生产鉴定用菌毒种管理及质量控制,建立三级种子库,即原始种子、主种子批及工作种子批,并进行相关检定。三级种子库检定内容包括无菌检查,鉴别试验、病毒滴定、外源因子检查、支原体检查、免疫原性、基因序列测定等检查项目,各检测项目合格后,才能使用工作种子批。种子库的建立方法如下:将柯萨奇病毒A16型毒株V00080265按MOI(0.1~0.0001)接种于处于对数生长期的单层Vero细胞,每天观察病变情况,至病变达+++~++++,收获病毒液,建立三级种子库。
实施例6 CA16疫苗的制备
取实施例5建立的柯萨奇病毒A16型毒株V00080265工作种子批,采用细胞工厂或者生物反应器在Vero 细胞上进行扩大培养,按MOI=0.001接种病毒,待病变达 +++~++++收获病毒液。将收获的病毒液经离心或过滤后,采用甲醛1:4000灭活4天,或者β-丙内酯1:2000于5±3℃灭活1天,37℃水解1小时。将灭活液进行超滤浓缩、层析纯化,纯化液经除菌过滤后获得疫苗原液。灭活也可在病毒纯化后进行。
将上述原液与氢氧化铝或磷酸铝佐剂按适宜比例吸附后,制备成CA16疫苗。疫苗蛋白浓度为2μg /ml,铝浓度1.0μg/ml。
实施例7 CA16疫苗的免疫原性
将实施例6中配制的CA16 疫苗进行小鼠免疫试验。其中,小鼠的免疫方法如下:将小鼠随机分成2组,每组小鼠10只按0、14天的免疫程序进行CA16 疫苗的腹腔免疫,0.5ml/只,于第28天进行采血,同时免疫病毒维持液作为对照组。采集的血清进行中和抗体检测,具体方法见实施例2。结果显示,二免后14天,试验组血清效价100%阳转,中和抗体水平GTMs可达1:641,对照组阳转率为0%,GTMs<1:8,表明使用柯萨奇病毒A16型毒株V00080265制备的CA16疫苗具有很好的免疫效果。
虽然,上文中已经用一般性说明及具体实施方案对本发明作了详尽的描述,但在本发明基础上,可以对之作一些修改或改进,这对本领域技术人员而言是显而易见的。因此,在不偏离本发明精神的基础上所做的这些修改或改进,均属于本发明要求保护的范围。
序列表
<110> 北京民海生物科技有限公司
<120> 柯萨奇病毒A16型毒株及其免疫原性组合物和应用
<130> KHP211120535.0YS
<160> 27
<170> SIPOSequenceListing 1.0
<210> 1
<211> 862
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 1
Met Gly Ser Gln Val Ser Thr Gln Arg Ser Gly Ser His Glu Asn Ser
1 5 10 15
Asn Ser Ala Ser Glu Gly Ser Thr Ile Asn Tyr Thr Thr Ile Asn Tyr
20 25 30
Tyr Lys Asp Ala Tyr Ala Ala Ser Ala Gly Arg Gln Asp Met Ser Gln
35 40 45
Asp Pro Lys Lys Phe Thr Asp Pro Val Met Asp Val Ile His Glu Met
50 55 60
Ala Pro Pro Leu Lys Ser Pro Ser Ala Glu Ala Cys Gly Tyr Ser Asp
65 70 75 80
Arg Val Ala Gln Leu Thr Ile Gly Asn Ser Thr Ile Thr Thr Gln Glu
85 90 95
Ala Ala Asn Ile Val Ile Ala Tyr Gly Glu Trp Pro Glu Tyr Cys Pro
100 105 110
Asp Thr Asp Ala Thr Ala Val Asp Lys Pro Thr Arg Pro Asp Val Ser
115 120 125
Val Asn Arg Phe Phe Thr Leu Asp Thr Lys Ser Trp Ala Lys Asp Ser
130 135 140
Lys Gly Trp Tyr Trp Lys Phe Pro Asp Val Leu Thr Glu Val Gly Val
145 150 155 160
Phe Gly Gln Asn Ala Gln Phe His Tyr Leu Tyr Arg Ser Gly Phe Cys
165 170 175
Val His Val Gln Cys Asn Ala Ser Lys Phe His Gln Gly Ala Leu Leu
180 185 190
Val Ala Val Leu Pro Glu Tyr Val Leu Gly Thr Ile Ala Gly Gly Thr
195 200 205
Gly Asn Glu Asn Ser His Pro Pro Tyr Ala Thr Thr Gln Pro Gly Gln
210 215 220
Val Gly Ala Val Leu Thr His Pro Tyr Val Leu Asp Ala Gly Ile Pro
225 230 235 240
Leu Ser Gln Leu Thr Val Cys Pro His Gln Trp Ile Asn Leu Arg Thr
245 250 255
Asn Asn Cys Ala Thr Ile Ile Val Pro Tyr Met Asn Thr Val Pro Phe
260 265 270
Asp Ser Ala Leu Asn His Cys Asn Phe Gly Leu Leu Val Val Pro Val
275 280 285
Val Pro Leu Asp Phe Asn Thr Gly Ala Thr Ser Glu Ile Pro Ile Thr
290 295 300
Val Thr Ile Ala Pro Met Cys Ala Glu Phe Ala Gly Leu Arg Gln Ala
305 310 315 320
Val Lys Gln Gly Ile Pro Thr Glu Leu Lys Pro Gly Thr Asn Gln Phe
325 330 335
Leu Thr Thr Asp Asp Gly Val Ser Ala Pro Ile Leu Pro Gly Phe His
340 345 350
Pro Thr Pro Pro Ile His Ile Pro Gly Glu Val His Asn Leu Leu Glu
355 360 365
Ile Cys Arg Val Glu Thr Ile Leu Glu Val Asn Asn Leu Lys Thr Asn
370 375 380
Glu Thr Thr Pro Met Gln Arg Leu Cys Phe Pro Val Ser Val Gln Ser
385 390 395 400
Lys Thr Gly Glu Leu Cys Ala Ala Phe Arg Ala Asp Pro Gly Arg Asp
405 410 415
Gly Pro Trp Gln Ser Thr Ile Leu Gly Gln Leu Cys Arg Tyr Tyr Thr
420 425 430
Gln Trp Ser Gly Ser Leu Glu Val Thr Phe Met Phe Ala Gly Ser Phe
435 440 445
Met Ala Thr Gly Lys Met Leu Ile Ala Tyr Thr Pro Pro Gly Gly Asn
450 455 460
Val Pro Ala Asp Arg Ile Thr Ala Met Leu Gly Thr His Val Ile Trp
465 470 475 480
Asp Phe Gly Leu Gln Ser Ser Val Thr Leu Val Val Pro Trp Ile Ser
485 490 495
Asn Thr His Tyr Arg Ala His Ala Arg Ala Gly Tyr Phe Asp Tyr Tyr
500 505 510
Thr Thr Gly Ile Ile Thr Ile Trp Tyr Gln Thr Asn Tyr Val Val Pro
515 520 525
Ile Gly Ala Pro Thr Thr Ala Tyr Ile Val Ala Leu Ala Ala Ala Gln
530 535 540
Asp Asn Phe Thr Met Lys Leu Cys Lys Asp Thr Glu Asp Ile Glu Gln
545 550 555 560
Thr Ala Asn Ile Gln Gly Asp Pro Ile Ala Asp Met Ile Asp Gln Thr
565 570 575
Val Asn Asn Gln Val Asn Arg Ser Leu Thr Ala Leu Gln Val Leu Pro
580 585 590
Thr Ala Ala Asn Thr Glu Ala Ser Ser His Arg Leu Gly Thr Gly Val
595 600 605
Val Pro Ala Leu Gln Ala Ala Glu Thr Gly Ala Ser Ser Asn Ala Ser
610 615 620
Asp Lys Asn Leu Ile Glu Thr Arg Cys Val Leu Asn His His Ser Thr
625 630 635 640
Gln Glu Thr Ala Ile Gly Asn Phe Phe Ser Arg Ala Gly Leu Val Ser
645 650 655
Ile Ile Thr Met Pro Thr Thr Gly Thr Gln Asn Thr Asp Gly Tyr Val
660 665 670
Asn Trp Asp Ile Asp Leu Met Gly Tyr Ala Gln Leu Arg Arg Lys Cys
675 680 685
Glu Leu Phe Thr Tyr Met Arg Phe Asp Ala Glu Phe Thr Phe Val Val
690 695 700
Ala Lys Pro Asn Gly Glu Leu Val Pro Gln Leu Leu Gln Tyr Met Tyr
705 710 715 720
Val Pro Pro Gly Ala Pro Lys Pro Lys Ser Arg Asp Ser Phe Ala Trp
725 730 735
Gln Thr Ala Thr Asn Pro Ser Val Phe Val Lys Met Thr Asp Pro Pro
740 745 750
Ala Gln Val Ser Val Pro Phe Met Ser Pro Ala Ser Ala Tyr Gln Trp
755 760 765
Phe Tyr Asp Gly Tyr Pro Thr Phe Gly Glu His Leu Gln Ala Asn Asp
770 775 780
Leu Asp Tyr Gly Gln Cys Pro Asn Asn Met Met Gly Thr Phe Ser Ile
785 790 795 800
Arg Thr Val Gly Thr Glu Lys Ser Pro His Ser Ile Thr Leu Arg Ile
805 810 815
Tyr Met Arg Ile Lys His Val Arg Ala Trp Ile Pro Arg Pro Leu Arg
820 825 830
Asn Gln Pro Tyr Leu Phe Lys Thr Asn Pro Asn Tyr Lys Gly Asn Asp
835 840 845
Ile Lys Cys Thr Ser Thr Ser Arg Asp Lys Ile Thr Thr Leu
850 855 860
<210> 2
<211> 2586
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 2
atggggtcac aagtctctac tcagcggtcc gggtcgcatg agaactcaaa ctccgcatcg 60
gaaggctcaa ccataaatta tacaactata aactactata aggatgcata tgctgcgagt 120
gcggggcgcc aggatatgtc tcaagacccg aagaaattta ccgaccctgt tatggatgtt 180
atacatgaga tggccccacc gctcaagtct ccgagcgctg aggcgtgtgg ttatagtgat 240
cgtgtggccc agcttaccat tgggaattct accattacta cgcaagaagc agctaacata 300
gttatagcct atggggagtg gcctgaatat tgcccagaca cggatgcgac ggcagtcgac 360
aagcccacac gacctgacgt gtcagtgaat agatttttca cactagatac taaatcttgg 420
gcaaaggatt caaagggatg gtattggaaa ttccccgatg ttttgacaga ggtaggcgtg 480
tttggtcaaa atgctcaatt tcactacctg tatcgatctg gattttgcgt gcacgtccag 540
tgtaatgcaa gtaaattcca ccagggtgct ttactggtgg ccgtgctacc tgagtatgtg 600
ctcggcacta tcgcaggggg gactgggaac gagaattctc atcctcccta cgctactaca 660
cagcctggtc aggttggtgc agtcctgacg cacccatatg tactagatgc agggatacct 720
ttgagccaat taaccgtgtg tcctcaccag tggatcaact tgagaaccaa caattgtgca 780
actattatag tcccatacat gaacacggtt ccatttgatt cagctcttaa tcactgcaat 840
tttgggttgc tggtcgtccc ggtggtgcca ttggacttta atacaggtgc cacgtctgaa 900
attcctatta cagtcaccat agctcccatg tgtgcagagt ttgcgggtct gcgccaggca 960
gtgaagcaag gcataccaac agagctcaag cctggtacca atcagtttct cactaccgat 1020
gatggtgtct ctgcaccaat tttaccaggc ttccacccaa ccccccctat acatatacca 1080
ggagaagtgc ataacttatt ggagatatgt agagtggaga ccattttaga agttaacaac 1140
ctgaagacca atgagaccac ccccatgcag cgcttgtgct ttccagtgtc agttcagagc 1200
aaaacaggtg agttgtgtgc tgcctttaga gcagaccctg gaagagatgg tccgtggcag 1260
tccacaatac tgggtcaact ctgcaggtac tacacccagt ggtcgggttc attggaggtg 1320
acatttatgt ttgcgggctc attcatggcc acaggcaaga tgctcatcgc ctacacccca 1380
cctgggggaa atgtgcctgc ggacagaatc acagcaatgt taggaacgca tgtgatctgg 1440
gactttggat tgcaatcctc tgtgacattg gttgtgccat ggattagcaa tacgcattac 1500
agggcgcacg cccgtgctgg gtactttgac tattacacta ccggcattat aactatatgg 1560
tatcaaacca actatgtggt acctattggc gctcccacta cagcatacat cgtagctctg 1620
gcagcagccc aagacaactt taccatgaaa ttatgcaagg atacagagga cattgagcaa 1680
acagctaata tacaagggga tcccattgct gacatgatcg accaaactgt gaataaccaa 1740
gtgaaccgct ccttaaccgc attacaagta ctacctacag ctgccaatac tgaagcaagt 1800
agccacagat taggcactgg tgttgtgcca gcactgcaag ctgcggagac gggggcgtca 1860
tcaaatgcca gtgacaagaa tctcattgag acgagatgtg tgttgaacca tcattccaca 1920
caggagacag ccatcgggaa tttctttagc cgtgctggtt tggttagcat catcacaatg 1980
cccaccacgg gtacacagaa cacagacggt tacgtcaact gggacattga cttgatggga 2040
tatgctcaac tacggcgcaa gtgcgagttg ttcacgtata tgcgctttga tgctgaattc 2100
acatttgtcg tagctaaacc caatggcgaa ctggtccccc agttactgca gtacatgtat 2160
gtcccaccag gagccccgaa acccaaatct agagattcat ttgcttggca aactgctacc 2220
aacccgtctg tatttgtgaa aatgacagac ccaccagctc aagtgtcagt cccctttatg 2280
tcaccagcca gtgcatacca atggttttac gatggttatc ccactttcgg ggagcatctc 2340
caagcaaatg atctagatta tggccagtgc ccgaacaata tgatgggcac ctttagtatt 2400
agaacagtag ggactgagaa gtcaccacac tccattactc tgaggatata tatgaggatt 2460
aaacacgtta gagcgtggat tccaaggcct ctgagaaatc aaccctattt gtttaagacc 2520
aacccaaatt ataaagggaa tgacattaag tgtactagta ctagtagaga caagataaca 2580
acatta 2586
<210> 3
<211> 7360
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 3
cacccacagg gcccacgggg cgttagcaca ctggcgttac ggcacctttg tgcgcctgtt 60
ttgtttcccc tcccccccgc aatttagaag ttttgcgcca tggatcaata gcaggtgtgt 120
cgcaccaggc acatctagat caagcacttc tgtttccccg gactgagtat caataggctg 180
ctcacgcggc tgaaggagaa agcgttcgtt atccggccaa ctacttcgag aagcttagta 240
ccaccatgaa cgttgcagag tgtttcgctc agcacatccc cagtgtagat caggtcgatg 300
agtcaccgca ctccccacgg gcgaccgtgg cggtggctgc gttggcggcc tgcctatggg 360
gcaacccata ggacgctcta ataccgacat ggtgcgaaga gcctattgag ctagttggta 420
gtcctccggc ccctgaatgc ggctaatcct aactgcggag catgtactca caatccagtg 480
ggcagcatgt cgtaacgggc aactctgcag cggaaccgac tactttgggt gtccgtgttt 540
ccttttattc ttatattggc tgcttatggt gacaattgaa gaattgttac catatagcta 600
ttggattggc catccggtgt ctaacagagc tattgtttac ctgtttgttg gatacgttcc 660
tctcaatttc aaggtcatta aaactcttaa ttatatacta cttctcaact gtgagaaatg 720
gggtcacaag tctctactca gcggtccggg tcgcatgaga actcaaactc cgcatcggaa 780
ggctcaacca taaattatac aactataaac tactataagg atgcatatgc tgcgagtgcg 840
gggcgccagg atatgtctca agacccgaag aaatttaccg accctgttat ggatgttata 900
catgagatgg ccccaccgct caagtctccg agcgctgagg cgtgtggtta tagtgatcgt 960
gtggcccagc ttaccattgg gaattctacc attactacgc aagaagcagc taacatagtt 1020
atagcctatg gggagtggcc tgaatattgc ccagacacgg atgcgacggc agtcgacaag 1080
cccacacgac ctgacgtgtc agtgaataga tttttcacac tagatactaa atcttgggca 1140
aaggattcaa agggatggta ttggaaattc cccgatgttt tgacagaggt aggcgtgttt 1200
ggtcaaaatg ctcaatttca ctacctgtat cgatctggat tttgcgtgca cgtccagtgt 1260
aatgcaagta aattccacca gggtgcttta ctggtggccg tgctacctga gtatgtgctc 1320
ggcactatcg caggggggac tgggaacgag aattctcatc ctccctacgc tactacacag 1380
cctggtcagg ttggtgcagt cctgacgcac ccatatgtac tagatgcagg gatacctttg 1440
agccaattaa ccgtgtgtcc tcaccagtgg atcaacttga gaaccaacaa ttgtgcaact 1500
attatagtcc catacatgaa cacggttcca tttgattcag ctcttaatca ctgcaatttt 1560
gggttgctgg tcgtcccggt ggtgccattg gactttaata caggtgccac gtctgaaatt 1620
cctattacag tcaccatagc tcccatgtgt gcagagtttg cgggtctgcg ccaggcagtg 1680
aagcaaggca taccaacaga gctcaagcct ggtaccaatc agtttctcac taccgatgat 1740
ggtgtctctg caccaatttt accaggcttc cacccaaccc cccctataca tataccagga 1800
gaagtgcata acttattgga gatatgtaga gtggagacca ttttagaagt taacaacctg 1860
aagaccaatg agaccacccc catgcagcgc ttgtgctttc cagtgtcagt tcagagcaaa 1920
acaggtgagt tgtgtgctgc ctttagagca gaccctggaa gagatggtcc gtggcagtcc 1980
acaatactgg gtcaactctg caggtactac acccagtggt cgggttcatt ggaggtgaca 2040
tttatgtttg cgggctcatt catggccaca ggcaagatgc tcatcgccta caccccacct 2100
gggggaaatg tgcctgcgga cagaatcaca gcaatgttag gaacgcatgt gatctgggac 2160
tttggattgc aatcctctgt gacattggtt gtgccatgga ttagcaatac gcattacagg 2220
gcgcacgccc gtgctgggta ctttgactat tacactaccg gcattataac tatatggtat 2280
caaaccaact atgtggtacc tattggcgct cccactacag catacatcgt agctctggca 2340
gcagcccaag acaactttac catgaaatta tgcaaggata cagaggacat tgagcaaaca 2400
gctaatatac aaggggatcc cattgctgac atgatcgacc aaactgtgaa taaccaagtg 2460
aaccgctcct taaccgcatt acaagtacta cctacagctg ccaatactga agcaagtagc 2520
cacagattag gcactggtgt tgtgccagca ctgcaagctg cggagacggg ggcgtcatca 2580
aatgccagtg acaagaatct cattgagacg agatgtgtgt tgaaccatca ttccacacag 2640
gagacagcca tcgggaattt ctttagccgt gctggtttgg ttagcatcat cacaatgccc 2700
accacgggta cacagaacac agacggttac gtcaactggg acattgactt gatgggatat 2760
gctcaactac ggcgcaagtg cgagttgttc acgtatatgc gctttgatgc tgaattcaca 2820
tttgtcgtag ctaaacccaa tggcgaactg gtcccccagt tactgcagta catgtatgtc 2880
ccaccaggag ccccgaaacc caaatctaga gattcatttg cttggcaaac tgctaccaac 2940
ccgtctgtat ttgtgaaaat gacagaccca ccagctcaag tgtcagtccc ctttatgtca 3000
ccagccagtg cataccaatg gttttacgat ggttatccca ctttcgggga gcatctccaa 3060
gcaaatgatc tagattatgg ccagtgcccg aacaatatga tgggcacctt tagtattaga 3120
acagtaggga ctgagaagtc accacactcc attactctga ggatatatat gaggattaaa 3180
cacgttagag cgtggattcc aaggcctctg agaaatcaac cctatttgtt taagaccaac 3240
ccaaattata aagggaatga cattaagtgt actagtacta gtagagacaa gataacaaca 3300
ttaggaaggt ttgggcagca gtcgggcgcc atatatgtag gcaactatag ggtagtgaat 3360
cggcatcttg ccacacacaa cgactgggca aatcttgtgt gggaggacag ctctagagac 3420
ctgttagttt cttccaccac tgcccagggg tgcgatacca tcgctagatg cgattgtcaa 3480
gccggagtat attattgcaa ctccaagaga aaacactacc cggttagttt caccaagccc 3540
agcctggtat ttgtggaggc tagtgagtat tacccagcta gataccaatc ccaccttatg 3600
cttgctgtag gccattcaga acctggcgac tgtggtggca tcctcagatg ccaacacggt 3660
gtgataggaa ttgtctccac tggtggcaat ggtcttgtgg ggtttgctga catcagagat 3720
ctcctgtggc tggatgaaga agcgatggag caaggagtgt ctgattatat caaaggcctc 3780
ggtgatgctt ttggcatggg cttcactgat gcagtgtcca gggaagtgga ggcattgaag 3840
aaccacttaa tcggttcaga aggggctgtt gaaaaaattc tgaagaattt ggtgaagcta 3900
atttcagcat tagtcatagt cgttaggagt gactatgaca tggtcaccct cacagccacg 3960
cttgccttga ttgggtgcca tggaagccct tgggcatgga taaaagcgaa gacagcctct 4020
attcttggca ttcccatagt gcaaaagcag agcgcttcat ggctaaagaa gtttaatgac 4080
atggctaatg ctgctaaggg acttgagtgg atttctagta aaatcagtaa gtttattgat 4140
tggcttaagg aaaagattat cccagccgct aaagagaagg ttgaattctt gaacaacttg 4200
aagcagcttc ccttactgga gaaccaaatt tcgaatctcg aacagtctgc tgcctcgcag 4260
gaggatctag aagctatgtt tggtaacgtg tcatatttgg cccacttttg ccgcaagttt 4320
cagccactct acgcaaccga agctaaacga gtctatgcgc tggagaaaag gatgaacaac 4380
tacatgcagt tcaagagcaa acaccgtatt gaacccgtat gtttgatcat cagaggctcc 4440
ccaggaacag gcaagtcgct tgccacgggc atcatagcta gagccattgc cgataaatac 4500
cattctagtg tttactcact tcctccagac ccagaccatt tcgatgggta caagcaacaa 4560
gtagtcactg ttatggatga tctttgtcaa aacccagatg gaaaggacat gtcactattt 4620
tgccagatgg tttctacagt ggatttcata ccacccatgg catccctgga agagaaggga 4680
gtgtctttca cctctaagtt tgtcattgca tcgaccaatg ctagcaacat agtagttccc 4740
acagtttcag actcagatgc gattcgcaga cggttctaca tggactgtga tatagaggtg 4800
acagattcct acaagacaga ccttggccga cttgatgcag gtagagccgc caagctttgc 4860
acggaaaata ataccgccaa ctttaagaga tgcagcccac tagtgtgtgg caaagctatt 4920
caactaagag ataggaaatc caaagtgaga tacagcattg atactgtagt atcggagcta 4980
atcagagagt acaacaatag atccgccatc ggtaatacta tagaagctct cttccaagga 5040
ccccttaagt tcaagcctat aaggattagc cttgaagaga agccagctcc ggatgccatc 5100
agtgacctac tagctagtgt ggatagcgaa gaggttcggc aatactgcag ggaacagggg 5160
tggataattc cagaaacacc ggccaatgtg gaacgtcacc tcaacagagc agtgctagtg 5220
atgcagtcta tcgccaccgt ggttgcagtt gtgtctcttg tctatgttat ttacaaattg 5280
ttcgctgggt ttcagggtgc ttattctggt gcgcccaaac aagctcttaa gaagcccgtg 5340
ctaagaacag ccacggttca aggaccgagt ttagactttg ccttatccct tttaaggcgt 5400
aacattagac aggtgcaaac tgatcaaggg cacttcacca tgttaggggt acgggaccgc 5460
ctggctgttc tgccacgcca ctcgcaacca ggaaaaacta tttgggtgga acacaaattg 5520
attaatgtgc tagacgctgt tgagttagta gatgaacaag gtgtgaattt ggaacttaca 5580
ctagtaactt tagacaccaa cgaaaagttt agggacatca ccaagtttat tccagagaca 5640
atcactggag caagtgacgc aaccttgatc atcaacactg agcatatgcc ctcaatgttc 5700
gtcccagtgg gtgatgttgt acaatatgga ttcttgaatc ttagcggtaa gcccacacac 5760
cggaccatga tgtataactt ccccacaaag gcaggacagt gtggaggagt agtcacctca 5820
gttggtaaga tcattggagt ccacattggt gggaacggcc gtcaaggttt ctgtgctggg 5880
ttgaagagga gctactttgc cagtgaacag ggagaaatcc agtggatgaa gcccaataag 5940
gagactggga gattgaatgt taatggcccg acccgtacca aattagagcc tagtgtattc 6000
catgatgtgt ttgagggcag caaagaacca gcagtcttaa ccagtaagga ccccagactc 6060
gaggttgatt ttgagcaagc tttgttttcc aagtatgtgg gaaacacctt gcatgagcct 6120
gatgagtatg tgacgcaggc tgctctccac tatgctaacc agttaaaaca attagacatc 6180
aatactaata agatgagcat ggaagaagca tgttatggca ctgaatatct ggaggccata 6240
gaccttcata ccagtgctgg gtatccctat agtgccttgg gcattaagaa aagggacata 6300
ctcgacccgg tcactagaga caccactaaa atgaaattct acatggacaa atatgggtta 6360
gacatgccct attccactta tgtgaaagat gagctcaggt ccttagataa gattaagaag 6420
gggaaatccc gtttgattga agccagcagc ttgaatgatt cagtctatct taggatgacc 6480
tttgggcatc tctatgagac ttttcatgcc aacccgggga ctgtgaccgg atctgcagta 6540
gggtgcaatc ctgatgtgtt ctggagtaaa ttaccgatcc tgttaccggg gtcgctcttt 6600
gcatttgatt attcagggta tgatgcaagc ctcagcccag tgtggttcag agctttagag 6660
gtggtcctcc gtgagattgg ttactcagag gaggctgtgt cactaataga agggatcaac 6720
cacacccatc atgtgtatcg gaacaagaca tattgtgtcc ttggtggaat gccctcaggt 6780
tgttccggca cttccatctt caactctatg atcaataata taataatcag aactcttctg 6840
atcaaaacct ttaaggggat cgatttagat gagttgaaca tggtagctta tggagatgat 6900
gtgctggcta gctatccatt ccctattgac tgttcggagt tggccaagac tggcaaagag 6960
tatggattaa caatgacacc tgctgacaaa tcaccctgct ttaacgaagt cacttgggag 7020
aatgctacat tcttaaagag aggcttcttg ccagatcacc agtttccatt ccttatccac 7080
cccaccatgc ctatgagaga gatccatgag tccattcgtt ggactaagga tgcacgcaac 7140
actcaggacc acgtgcgttc cttgtgccta ttagcatggc ataatgggaa ggaggagtat 7200
gaaaaatttg tgagcacaat tagatcggtt cctattggga aagccttggc tataccaaat 7260
tttgaaaacc tgagaagaaa ttggctcgaa ttattttgat atacagctca aagctgaacc 7320
ccaccagaaa tctggtcatg ttaatgactg gtgggggtac 7360
<210> 4
<211> 150
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 4
Gly Arg Phe Gly Gln Gln Ser Gly Ala Ile Tyr Val Gly Asn Tyr Arg
1 5 10 15
Val Val Asn Arg His Leu Ala Thr His Asn Asp Trp Ala Asn Leu Val
20 25 30
Trp Glu Asp Ser Ser Arg Asp Leu Leu Val Ser Ser Thr Thr Ala Gln
35 40 45
Gly Cys Asp Thr Ile Ala Arg Cys Asp Cys Gln Ala Gly Val Tyr Tyr
50 55 60
Cys Asn Ser Lys Arg Lys His Tyr Pro Val Ser Phe Thr Lys Pro Ser
65 70 75 80
Leu Val Phe Val Glu Ala Ser Glu Tyr Tyr Pro Ala Arg Tyr Gln Ser
85 90 95
His Leu Met Leu Ala Val Gly His Ser Glu Pro Gly Asp Cys Gly Gly
100 105 110
Ile Leu Arg Cys Gln His Gly Val Ile Gly Ile Val Ser Thr Gly Gly
115 120 125
Asn Gly Leu Val Gly Phe Ala Asp Ile Arg Asp Leu Leu Trp Leu Asp
130 135 140
Glu Glu Ala Met Glu Gln
145 150
<210> 5
<211> 99
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 5
Gly Val Ser Asp Tyr Ile Lys Gly Leu Gly Asp Ala Phe Gly Met Gly
1 5 10 15
Phe Thr Asp Ala Val Ser Arg Glu Val Glu Ala Leu Lys Asn His Leu
20 25 30
Ile Gly Ser Glu Gly Ala Val Glu Lys Ile Leu Lys Asn Leu Val Lys
35 40 45
Leu Ile Ser Ala Leu Val Ile Val Val Arg Ser Asp Tyr Asp Met Val
50 55 60
Thr Leu Thr Ala Thr Leu Ala Leu Ile Gly Cys His Gly Ser Pro Trp
65 70 75 80
Ala Trp Ile Lys Ala Lys Thr Ala Ser Ile Leu Gly Ile Pro Ile Val
85 90 95
Gln Lys Gln
<210> 6
<211> 329
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 6
Ser Ala Ser Trp Leu Lys Lys Phe Asn Asp Met Ala Asn Ala Ala Lys
1 5 10 15
Gly Leu Glu Trp Ile Ser Ser Lys Ile Ser Lys Phe Ile Asp Trp Leu
20 25 30
Lys Glu Lys Ile Ile Pro Ala Ala Lys Glu Lys Val Glu Phe Leu Asn
35 40 45
Asn Leu Lys Gln Leu Pro Leu Leu Glu Asn Gln Ile Ser Asn Leu Glu
50 55 60
Gln Ser Ala Ala Ser Gln Glu Asp Leu Glu Ala Met Phe Gly Asn Val
65 70 75 80
Ser Tyr Leu Ala His Phe Cys Arg Lys Phe Gln Pro Leu Tyr Ala Thr
85 90 95
Glu Ala Lys Arg Val Tyr Ala Leu Glu Lys Arg Met Asn Asn Tyr Met
100 105 110
Gln Phe Lys Ser Lys His Arg Ile Glu Pro Val Cys Leu Ile Ile Arg
115 120 125
Gly Ser Pro Gly Thr Gly Lys Ser Leu Ala Thr Gly Ile Ile Ala Arg
130 135 140
Ala Ile Ala Asp Lys Tyr His Ser Ser Val Tyr Ser Leu Pro Pro Asp
145 150 155 160
Pro Asp His Phe Asp Gly Tyr Lys Gln Gln Val Val Thr Val Met Asp
165 170 175
Asp Leu Cys Gln Asn Pro Asp Gly Lys Asp Met Ser Leu Phe Cys Gln
180 185 190
Met Val Ser Thr Val Asp Phe Ile Pro Pro Met Ala Ser Leu Glu Glu
195 200 205
Lys Gly Val Ser Phe Thr Ser Lys Phe Val Ile Ala Ser Thr Asn Ala
210 215 220
Ser Asn Ile Val Val Pro Thr Val Ser Asp Ser Asp Ala Ile Arg Arg
225 230 235 240
Arg Phe Tyr Met Asp Cys Asp Ile Glu Val Thr Asp Ser Tyr Lys Thr
245 250 255
Asp Leu Gly Arg Leu Asp Ala Gly Arg Ala Ala Lys Leu Cys Thr Glu
260 265 270
Asn Asn Thr Ala Asn Phe Lys Arg Cys Ser Pro Leu Val Cys Gly Lys
275 280 285
Ala Ile Gln Leu Arg Asp Arg Lys Ser Lys Val Arg Tyr Ser Ile Asp
290 295 300
Thr Val Val Ser Glu Leu Ile Arg Glu Tyr Asn Asn Arg Ser Ala Ile
305 310 315 320
Gly Asn Thr Ile Glu Ala Leu Phe Gln
325
<210> 7
<211> 86
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 7
Gly Pro Leu Lys Phe Lys Pro Ile Arg Ile Ser Leu Glu Glu Lys Pro
1 5 10 15
Ala Pro Asp Ala Ile Ser Asp Leu Leu Ala Ser Val Asp Ser Glu Glu
20 25 30
Val Arg Gln Tyr Cys Arg Glu Gln Gly Trp Ile Ile Pro Glu Thr Pro
35 40 45
Ala Asn Val Glu Arg His Leu Asn Arg Ala Val Leu Val Met Gln Ser
50 55 60
Ile Ala Thr Val Val Ala Val Val Ser Leu Val Tyr Val Ile Tyr Lys
65 70 75 80
Leu Phe Ala Gly Phe Gln
85
<210> 8
<211> 22
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 8
Gly Ala Tyr Ser Gly Ala Pro Lys Gln Ala Leu Lys Lys Pro Val Leu
1 5 10 15
Arg Thr Ala Thr Val Gln
20
<210> 9
<211> 183
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 9
Gly Pro Ser Leu Asp Phe Ala Leu Ser Leu Leu Arg Arg Asn Ile Arg
1 5 10 15
Gln Val Gln Thr Asp Gln Gly His Phe Thr Met Leu Gly Val Arg Asp
20 25 30
Arg Leu Ala Val Leu Pro Arg His Ser Gln Pro Gly Lys Thr Ile Trp
35 40 45
Val Glu His Lys Leu Ile Asn Val Leu Asp Ala Val Glu Leu Val Asp
50 55 60
Glu Gln Gly Val Asn Leu Glu Leu Thr Leu Val Thr Leu Asp Thr Asn
65 70 75 80
Glu Lys Phe Arg Asp Ile Thr Lys Phe Ile Pro Glu Thr Ile Thr Gly
85 90 95
Ala Ser Asp Ala Thr Leu Ile Ile Asn Thr Glu His Met Pro Ser Met
100 105 110
Phe Val Pro Val Gly Asp Val Val Gln Tyr Gly Phe Leu Asn Leu Ser
115 120 125
Gly Lys Pro Thr His Arg Thr Met Met Tyr Asn Phe Pro Thr Lys Ala
130 135 140
Gly Gln Cys Gly Gly Val Val Thr Ser Val Gly Lys Ile Ile Gly Val
145 150 155 160
His Ile Gly Gly Asn Gly Arg Gln Gly Phe Cys Ala Gly Leu Lys Arg
165 170 175
Ser Tyr Phe Ala Ser Glu Gln
180
<210> 10
<211> 462
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 10
Gly Glu Ile Gln Trp Met Lys Pro Asn Lys Glu Thr Gly Arg Leu Asn
1 5 10 15
Val Asn Gly Pro Thr Arg Thr Lys Leu Glu Pro Ser Val Phe His Asp
20 25 30
Val Phe Glu Gly Ser Lys Glu Pro Ala Val Leu Thr Ser Lys Asp Pro
35 40 45
Arg Leu Glu Val Asp Phe Glu Gln Ala Leu Phe Ser Lys Tyr Val Gly
50 55 60
Asn Thr Leu His Glu Pro Asp Glu Tyr Val Thr Gln Ala Ala Leu His
65 70 75 80
Tyr Ala Asn Gln Leu Lys Gln Leu Asp Ile Asn Thr Asn Lys Met Ser
85 90 95
Met Glu Glu Ala Cys Tyr Gly Thr Glu Tyr Leu Glu Ala Ile Asp Leu
100 105 110
His Thr Ser Ala Gly Tyr Pro Tyr Ser Ala Leu Gly Ile Lys Lys Arg
115 120 125
Asp Ile Leu Asp Pro Val Thr Arg Asp Thr Thr Lys Met Lys Phe Tyr
130 135 140
Met Asp Lys Tyr Gly Leu Asp Met Pro Tyr Ser Thr Tyr Val Lys Asp
145 150 155 160
Glu Leu Arg Ser Leu Asp Lys Ile Lys Lys Gly Lys Ser Arg Leu Ile
165 170 175
Glu Ala Ser Ser Leu Asn Asp Ser Val Tyr Leu Arg Met Thr Phe Gly
180 185 190
His Leu Tyr Glu Thr Phe His Ala Asn Pro Gly Thr Val Thr Gly Ser
195 200 205
Ala Val Gly Cys Asn Pro Asp Val Phe Trp Ser Lys Leu Pro Ile Leu
210 215 220
Leu Pro Gly Ser Leu Phe Ala Phe Asp Tyr Ser Gly Tyr Asp Ala Ser
225 230 235 240
Leu Ser Pro Val Trp Phe Arg Ala Leu Glu Val Val Leu Arg Glu Ile
245 250 255
Gly Tyr Ser Glu Glu Ala Val Ser Leu Ile Glu Gly Ile Asn His Thr
260 265 270
His His Val Tyr Arg Asn Lys Thr Tyr Cys Val Leu Gly Gly Met Pro
275 280 285
Ser Gly Cys Ser Gly Thr Ser Ile Phe Asn Ser Met Ile Asn Asn Ile
290 295 300
Ile Ile Arg Thr Leu Leu Ile Lys Thr Phe Lys Gly Ile Asp Leu Asp
305 310 315 320
Glu Leu Asn Met Val Ala Tyr Gly Asp Asp Val Leu Ala Ser Tyr Pro
325 330 335
Phe Pro Ile Asp Cys Ser Glu Leu Ala Lys Thr Gly Lys Glu Tyr Gly
340 345 350
Leu Thr Met Thr Pro Ala Asp Lys Ser Pro Cys Phe Asn Glu Val Thr
355 360 365
Trp Glu Asn Ala Thr Phe Leu Lys Arg Gly Phe Leu Pro Asp His Gln
370 375 380
Phe Pro Phe Leu Ile His Pro Thr Met Pro Met Arg Glu Ile His Glu
385 390 395 400
Ser Ile Arg Trp Thr Lys Asp Ala Arg Asn Thr Gln Asp His Val Arg
405 410 415
Ser Leu Cys Leu Leu Ala Trp His Asn Gly Lys Glu Glu Tyr Glu Lys
420 425 430
Phe Val Ser Thr Ile Arg Ser Val Pro Ile Gly Lys Ala Leu Ala Ile
435 440 445
Pro Asn Phe Glu Asn Leu Arg Arg Asn Trp Leu Glu Leu Phe
450 455 460
<210> 11
<211> 450
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 11
ggaaggtttg ggcagcagtc gggcgccata tatgtaggca actatagggt agtgaatcgg 60
catcttgcca cacacaacga ctgggcaaat cttgtgtggg aggacagctc tagagacctg 120
ttagtttctt ccaccactgc ccaggggtgc gataccatcg ctagatgcga ttgtcaagcc 180
ggagtatatt attgcaactc caagagaaaa cactacccgg ttagtttcac caagcccagc 240
ctggtatttg tggaggctag tgagtattac ccagctagat accaatccca ccttatgctt 300
gctgtaggcc attcagaacc tggcgactgt ggtggcatcc tcagatgcca acacggtgtg 360
ataggaattg tctccactgg tggcaatggt cttgtggggt ttgctgacat cagagatctc 420
ctgtggctgg atgaagaagc gatggagcaa 450
<210> 12
<211> 297
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 12
ggagtgtctg attatatcaa aggcctcggt gatgcttttg gcatgggctt cactgatgca 60
gtgtccaggg aagtggaggc attgaagaac cacttaatcg gttcagaagg ggctgttgaa 120
aaaattctga agaatttggt gaagctaatt tcagcattag tcatagtcgt taggagtgac 180
tatgacatgg tcaccctcac agccacgctt gccttgattg ggtgccatgg aagcccttgg 240
gcatggataa aagcgaagac agcctctatt cttggcattc ccatagtgca aaagcag 297
<210> 13
<211> 987
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 13
agcgcttcat ggctaaagaa gtttaatgac atggctaatg ctgctaaggg acttgagtgg 60
atttctagta aaatcagtaa gtttattgat tggcttaagg aaaagattat cccagccgct 120
aaagagaagg ttgaattctt gaacaacttg aagcagcttc ccttactgga gaaccaaatt 180
tcgaatctcg aacagtctgc tgcctcgcag gaggatctag aagctatgtt tggtaacgtg 240
tcatatttgg cccacttttg ccgcaagttt cagccactct acgcaaccga agctaaacga 300
gtctatgcgc tggagaaaag gatgaacaac tacatgcagt tcaagagcaa acaccgtatt 360
gaacccgtat gtttgatcat cagaggctcc ccaggaacag gcaagtcgct tgccacgggc 420
atcatagcta gagccattgc cgataaatac cattctagtg tttactcact tcctccagac 480
ccagaccatt tcgatgggta caagcaacaa gtagtcactg ttatggatga tctttgtcaa 540
aacccagatg gaaaggacat gtcactattt tgccagatgg tttctacagt ggatttcata 600
ccacccatgg catccctgga agagaaggga gtgtctttca cctctaagtt tgtcattgca 660
tcgaccaatg ctagcaacat agtagttccc acagtttcag actcagatgc gattcgcaga 720
cggttctaca tggactgtga tatagaggtg acagattcct acaagacaga ccttggccga 780
cttgatgcag gtagagccgc caagctttgc acggaaaata ataccgccaa ctttaagaga 840
tgcagcccac tagtgtgtgg caaagctatt caactaagag ataggaaatc caaagtgaga 900
tacagcattg atactgtagt atcggagcta atcagagagt acaacaatag atccgccatc 960
ggtaatacta tagaagctct cttccaa 987
<210> 14
<211> 258
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 14
ggacccctta agttcaagcc tataaggatt agccttgaag agaagccagc tccggatgcc 60
atcagtgacc tactagctag tgtggatagc gaagaggttc ggcaatactg cagggaacag 120
gggtggataa ttccagaaac accggccaat gtggaacgtc acctcaacag agcagtgcta 180
gtgatgcagt ctatcgccac cgtggttgca gttgtgtctc ttgtctatgt tatttacaaa 240
ttgttcgctg ggtttcag 258
<210> 15
<211> 66
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 15
ggtgcttatt ctggtgcgcc caaacaagct cttaagaagc ccgtgctaag aacagccacg 60
gttcaa 66
<210> 16
<211> 549
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 16
ggaccgagtt tagactttgc cttatccctt ttaaggcgta acattagaca ggtgcaaact 60
gatcaagggc acttcaccat gttaggggta cgggaccgcc tggctgttct gccacgccac 120
tcgcaaccag gaaaaactat ttgggtggaa cacaaattga ttaatgtgct agacgctgtt 180
gagttagtag atgaacaagg tgtgaatttg gaacttacac tagtaacttt agacaccaac 240
gaaaagttta gggacatcac caagtttatt ccagagacaa tcactggagc aagtgacgca 300
accttgatca tcaacactga gcatatgccc tcaatgttcg tcccagtggg tgatgttgta 360
caatatggat tcttgaatct tagcggtaag cccacacacc ggaccatgat gtataacttc 420
cccacaaagg caggacagtg tggaggagta gtcacctcag ttggtaagat cattggagtc 480
cacattggtg ggaacggccg tcaaggtttc tgtgctgggt tgaagaggag ctactttgcc 540
agtgaacag 549
<210> 17
<211> 1386
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 17
ggagaaatcc agtggatgaa gcccaataag gagactggga gattgaatgt taatggcccg 60
acccgtacca aattagagcc tagtgtattc catgatgtgt ttgagggcag caaagaacca 120
gcagtcttaa ccagtaagga ccccagactc gaggttgatt ttgagcaagc tttgttttcc 180
aagtatgtgg gaaacacctt gcatgagcct gatgagtatg tgacgcaggc tgctctccac 240
tatgctaacc agttaaaaca attagacatc aatactaata agatgagcat ggaagaagca 300
tgttatggca ctgaatatct ggaggccata gaccttcata ccagtgctgg gtatccctat 360
agtgccttgg gcattaagaa aagggacata ctcgacccgg tcactagaga caccactaaa 420
atgaaattct acatggacaa atatgggtta gacatgccct attccactta tgtgaaagat 480
gagctcaggt ccttagataa gattaagaag gggaaatccc gtttgattga agccagcagc 540
ttgaatgatt cagtctatct taggatgacc tttgggcatc tctatgagac ttttcatgcc 600
aacccgggga ctgtgaccgg atctgcagta gggtgcaatc ctgatgtgtt ctggagtaaa 660
ttaccgatcc tgttaccggg gtcgctcttt gcatttgatt attcagggta tgatgcaagc 720
ctcagcccag tgtggttcag agctttagag gtggtcctcc gtgagattgg ttactcagag 780
gaggctgtgt cactaataga agggatcaac cacacccatc atgtgtatcg gaacaagaca 840
tattgtgtcc ttggtggaat gccctcaggt tgttccggca cttccatctt caactctatg 900
atcaataata taataatcag aactcttctg atcaaaacct ttaaggggat cgatttagat 960
gagttgaaca tggtagctta tggagatgat gtgctggcta gctatccatt ccctattgac 1020
tgttcggagt tggccaagac tggcaaagag tatggattaa caatgacacc tgctgacaaa 1080
tcaccctgct ttaacgaagt cacttgggag aatgctacat tcttaaagag aggcttcttg 1140
ccagatcacc agtttccatt ccttatccac cccaccatgc ctatgagaga gatccatgag 1200
tccattcgtt ggactaagga tgcacgcaac actcaggacc acgtgcgttc cttgtgccta 1260
ttagcatggc ataatgggaa ggaggagtat gaaaaatttg tgagcacaat tagatcggtt 1320
cctattggga aagccttggc tataccaaat tttgaaaacc tgagaagaaa ttggctcgaa 1380
ttattt 1386
<210> 18
<211> 717
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 18
cacccacagg gcccacgggg cgttagcaca ctggcgttac ggcacctttg tgcgcctgtt 60
ttgtttcccc tcccccccgc aatttagaag ttttgcgcca tggatcaata gcaggtgtgt 120
cgcaccaggc acatctagat caagcacttc tgtttccccg gactgagtat caataggctg 180
ctcacgcggc tgaaggagaa agcgttcgtt atccggccaa ctacttcgag aagcttagta 240
ccaccatgaa cgttgcagag tgtttcgctc agcacatccc cagtgtagat caggtcgatg 300
agtcaccgca ctccccacgg gcgaccgtgg cggtggctgc gttggcggcc tgcctatggg 360
gcaacccata ggacgctcta ataccgacat ggtgcgaaga gcctattgag ctagttggta 420
gtcctccggc ccctgaatgc ggctaatcct aactgcggag catgtactca caatccagtg 480
ggcagcatgt cgtaacgggc aactctgcag cggaaccgac tactttgggt gtccgtgttt 540
ccttttattc ttatattggc tgcttatggt gacaattgaa gaattgttac catatagcta 600
ttggattggc catccggtgt ctaacagagc tattgtttac ctgtttgttg gatacgttcc 660
tctcaatttc aaggtcatta aaactcttaa ttatatacta cttctcaact gtgagaa 717
<210> 19
<211> 61
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 19
tatacagctc aaagctgaac cccaccagaa atctggtcat gttaatgact ggtgggggta 60
c 61
<210> 20
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 20
attggtgctc ccactacagc 20
<210> 21
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 21
gagctgtcct cccacacaag 20
<210> 22
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 22
cyttgtgcgc ctgtttt 17
<210> 23
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 23
attgtcacca taagcagcc 19
<210> 24
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 24
caagyacttc tgtmwcccc 19
<210> 25
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 25
cccaaagtag tcggttcc 18
<210> 26
<211> 30
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 26
gttaccatat agctattgga ttggccatcc 30
<210> 27
<211> 30
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 27
ctagagctgt cctcccacac aaggtttgcc 30
Claims (6)
1.柯萨奇病毒A16型毒株,其特征在于,其基因组序列如SEQ ID NO.3所示或如SEQ IDNO.3所示序列的完全互补序列所示。
2.根据权利要求1所述的柯萨奇病毒A16型毒株,其特征在于,其保藏于中国微生物菌种保藏管理委员会普通微生物中心,保藏编号为CGMCC No.20387。
3.生物材料,其特征在于,其为以下(1)-(5)中的任意一种:
(1)序列如SEQ ID NO.3所示或如SEQ ID NO.3所示序列的完全互补序列所示的核酸分子;
(2)含有(1)中所述核酸分子的表达盒;
(3)含有(1)中所述核酸分子的重组载体;
(4)含有(1)中所述核酸分子的重组微生物;
(5)含有(1)中所述核酸分子的细胞系。
4.免疫原性组合物,其特征在于,其含有权利要求1或2所述的柯萨奇病毒A16型毒株或权利要求3所述的生物材料。
5.权利要求1或2所述的柯萨奇病毒A16型毒株或权利要求3所述的生物材料或权利要求4所述的免疫原性组合物的如下任意一种应用:
(1)在制备预防和/或治疗柯萨奇病毒引起疾病的疫苗中的应用;
(2)在制备预防和/或治疗柯萨奇病毒引起疾病的药物中的应用;
(3)在制备预防和/或治疗柯萨奇病毒引起疾病的抗体中的应用;
(4)在制备预防和/或治疗柯萨奇病毒引起疾病的抗血清中的应用;
(5)在非诊断和治疗目的的柯萨奇病毒疫苗的免疫原性评价中的应用;
(6)在非诊断和治疗目的的柯萨奇病毒疫苗的保护性评价中的应用;
(7)在制备柯萨奇病毒感染动物模型中的应用;
(8)在预防和/或治疗柯萨奇病毒引起疾病的非诊断和治疗目的的药物筛选或药效评价中的应用。
6.一种产品,其特征在于,其含有以下(1)-(3)中的任意一种或多种的组合:
(1)权利要求1或2所述的柯萨奇病毒A16型毒株;
(2)权利要求3所述的生物材料;
(3)权利要求4所述的免疫原性组合物。
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PCT/CN2022/099145 WO2023045426A1 (zh) | 2021-09-23 | 2022-06-16 | 柯萨奇病毒a16型毒株及其免疫原性组合物和应用 |
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