CN113881704A - 一种包含新型冠状病毒双抗原靶标序列组合的重组新城疫病毒载体及相应疫苗株和疫苗 - Google Patents
一种包含新型冠状病毒双抗原靶标序列组合的重组新城疫病毒载体及相应疫苗株和疫苗 Download PDFInfo
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Abstract
本发明提供了一种包含新型冠状病毒双抗原靶标序列组合的重组新城疫病毒载体及相应疫苗株和疫苗,涉及生物医药技术领域。本发明利用NDV载体,新型冠状病毒S蛋白基因和S蛋白受体结合域RBD基因作为抗原序列,将S蛋白基因和RBD基因共同插入NDV基因组内,利用病毒反向遗传学技术,拯救获得具有限制感染性NDV病毒载体疫苗株。该疫苗株可以被呼吸道细胞识别,产生粘膜免疫,并感染细胞,引起特异性免疫抗体水平升高。本发明双抗原靶标的引入,增强了疫苗株免疫原性的发挥,同时降低了疫苗株毒副作用的产生,免疫效果相对于单靶标抗原更具优势。
Description
技术领域
本发明属于生物医药技术领域,具体涉及一种包含新型冠状病毒双抗原靶标序列组合的重组新城疫病毒载体及相应疫苗株和疫苗。
背景技术
新型冠状病毒(SARS-COV-2)是一种新型β属冠状病毒,具有高传染性和高隐蔽性,可以通过多种途径,包括空气、食物等传播。新冠病毒的主要结构包括,单股正链核酸(ssRNA)、刺突蛋白(S)、膜蛋白(M)、包膜蛋白(E)和核壳蛋白(N)。其中,刺突S蛋白的RBD结构域是病毒识别宿主细胞膜受体蛋白ACE2蛋白的主要区域。S蛋白识别到细胞膜受体后,通过其S2结构域进入宿主细胞。所以,在研的新冠疫苗主要以S蛋白或者其RBD结构设计开发。
新城疫(Newcastle disease virus, NDV)病毒属于副粘病毒科Avulavirus属,是一种负链RNA病毒,其基因组包含N、P、M、F、HN和L六大基因以及非编码调控序列,以新城疫病毒为载体的重组病毒疫苗作为吸入或注射型疫苗的优秀候选,在人类中是安全的。但病毒承载外源基因有一定能力范围和插入位点要求,目前,以NDV为载体的新冠疫苗,主要是在病毒载体的P基因和M基因位置之间插入抗原S蛋白,但该技术手段产生的重组病毒载体疫苗株免疫原性十分有限,攻毒试验数据显示,其不能使免疫小鼠肺部的病毒载量下降超过2log,所以限制了NDV重组新冠病毒载体的推广及应用。如何在保证载体安全的前提下,提高NDV免疫原性以及抗体水平,成为亟待解决的问题。
发明内容
有鉴于此,本发明的目的在于提供一种包含新型冠状病毒双抗原靶标序列组合的重组新城疫病毒载体及疫苗株和疫苗,疫苗株可产生更强的免疫效果,可形成安全有效的新型冠状病毒载体疫苗,生产成本低。
为了实现上述发明目的,本发明提供以下技术方案:
本发明提供了一种包含新型冠状病毒双抗原靶标序列组合的重组新城疫病毒载体,所述双抗原靶标序列包括新型冠状病毒的S蛋白编码序列和S蛋白受体结合域RBD蛋白编码序列。
优选的,将所述新型冠状病毒的S蛋白编码序列插入新城疫病毒载体的P基因序列和M基因序列之间;
将所述新型冠状病毒S蛋白受体结合域RBD蛋白编码序列插入新城疫病毒载体的P基因序列和M基因序列之间、M基因序列和F基因序列之间或HN基因序列和L基因序列之间。
优选的,所述S蛋白编码序列和RBD蛋白编码序列来源于相同或不同的新型冠状病毒流行株。
优选的,所述S蛋白编码序列包括完整的S蛋白编码序列或所述完整的S蛋白编码序列的变体。
优选的,所述变体包括对所述完整的S蛋白编码序列进行基因工程改造;
所述基因工程改造的方法选自:突变、删除、重排和融合其他来源的蛋白多肽中一种或多种方法的组合;
经所述基因工程改造后的S蛋白构象成为预融合状态更容易被抗原呈递细胞识别,或更多量地表达在细胞膜上,或经表达后分泌到细胞外并组合成易被抗原呈递细胞识别的多聚体。
优选的,所述突变包括将完整的S蛋白的K986和V987位氨基酸均突变为P,或将第682~685位氨基酸RRAR突变为GSAS,或将完整的S蛋白的K986、V987、F817、 A892、A899和A942位氨基酸均突变为P;或将S蛋白的C末端KLHYT突变为ALAYT;
所述删除包括在所述完整的S蛋白的C末端截短18 ~28个氨基酸;或删除S蛋白的跨膜区和胞内区序列;
所述重排包括将S蛋白的主要抗原决定序列S1蛋白的NTD和RBD区域位置序列进行调换,形成RBD-NTD蛋白;
所述融合其他来源的蛋白多肽包括将S蛋白的胞外域与新城疫F蛋白的C端包含跨膜区和胞内区的氨基酸序列融合;或将RBD-NTD蛋白N端添加信号肽序列,C端添加融合蛋白多肽序列。
优选的,所述重排形成的RBD-NTD蛋白,RBD和NTD区域通过蛋白Linker连接。
优选的,所述RBD蛋白编码序列还包括经过基因工程改造的含有RBD蛋白编码序列的RBD系列重组蛋白编码序列。
优选的,所述基因工程改造包括:在RBD蛋白编码序列的N末端添加分泌信号肽序列,C末端添加融合蛋白多肽序列。
本发明还提供了一种利用上述重组新城疫病毒载体制备得到的疫苗株。
优选的,所述制备的方法包括病毒反向遗传学的方法。
优选的,所述疫苗株的结构包括:上述重组新城疫病毒载体的编码序列和/或展示在重组新城疫病毒粒子表面的S蛋白或所述S蛋白的变体;
所述S蛋白的变体由完整的S蛋白编码序列的变体编码得到。
本发明还提供了上述疫苗株在制备预防新型冠状病毒传播和感染疫苗中的应用。
本发明还提供了一种预防新型冠状病毒传播和感染疫苗的疫苗,所述疫苗包括上述疫苗株。
有益效果:一种利用新城疫病毒(Newcastle disease virus,NDV)载体,新型冠状病毒S蛋白基因和S蛋白受体结合域RBD蛋白基因作为抗原靶标序列,将S蛋白基因和RBD蛋白基因共同插入NDV基因组之间,利用病毒反向遗传学技术,拯救获得具有限制感染性NDV疫苗株,该疫苗株可以被呼吸道细胞识别,产生粘膜免疫,并感染细胞,引起特异性免疫抗体水平升高。相对于目前在研的单个新冠S蛋白抗原或RBD蛋白抗原疫苗,本发明引入双抗原靶标,并对双抗原靶标编码序列分别进行了基因工程优化,双抗原靶标的引入,增强了疫苗株免疫原性的发挥,同时降低了疫苗株毒副作用的产生,免疫效果相对于单靶标抗原更具优势。
附图说明
图1为构建得到的新型冠状病毒S蛋白和RBD蛋白双抗原靶标重组NDV载体示意图;其中a表示S蛋白及其变体和RBD系列重组蛋白基因同时位于NDV病毒载体P-M基因位置之间,通过2A肽序列连接;b表示S蛋白及其变体基因位于NDV病毒载体P-M基因位置之间,RBD系列重组蛋白基因位于NDV病毒载体M-F基因位置之间;
图2为重组NDV病毒载体PCR鉴定电泳图,Marker由上到下大小依次为5000 bp、3000 bp、2000 bp、1500 bp、1000 bp、750 bp、500 bp、250 bp、100 bp,Lane1: NDV-VH-RBD-Fc,Lane2: NDV-S2P,Lane3: NDV-S2P-VH-RBD-Fc,Lane4: NDV-S2P-HSA-RBD-Folden,Lane5-6: NDV-S2P-M-HSA-deltaRBD-Fc-F,Lane7-8: NDV-S2P-M-HSA-deltaRBD-Ferritin-F,Lane:9-10 NDV-S2P-M-HSA-RBD-Bann-F,Lane11 NDV-S2P/GSAS-HSA-deltaRBD-Fc:,Lane12-13: NDV-S6P-M-HSA-deltaRBD-Fc-F,Lane14-15: NDV-S2P- M-HSA-scdeltaRBD-F,Lane16: NDV-S2P- HSA-scdeltaRBD,Lane17: NDV-deltaS2P/F87,Lane18-19: NDV-deltaS2P/F87-M-HSA-deltaRBD-Fc-F,Lane20: NDV-S2P/GSAS-HSA-deltaRBD-Bann,Lane21: NDV-deltaRBD-NTD-TBSV-M-HSA-deltaRBD-Fc-F,Lane22: NDV-deltaRBD-NTD-Folden-M-HSA-deltaRBD-Fc-F;
图3为利用ELISA检测重组NDV载体疫苗株感染Vero细胞后RBD的表达结果;其中a表示标准曲线,b表示RBD抗原表达量;
图4为滴鼻免疫后血清新冠S1蛋白IgG抗体滴度(a)和血清NDV病毒IgG抗体滴度(b),*P < 0.05,**P < 0.01,***P < 0.001,****P < 0.0001,下同;
图5为肌注免疫后血清新冠S1蛋白IgG抗体滴度(a)和血清NDV病毒IgG抗体滴度(b)。
具体实施方式
本发明提供了一种包含新型冠状病毒双抗原靶标序列组合的重组新城疫病毒载体,所述双抗原靶标序列包括新型冠状病毒的S蛋白编码序列和S蛋白受体结合域RBD蛋白编码序列。
本发明所述新型冠状病毒的S蛋白编码序列和新型冠状病毒的RBD蛋白编码序列优选来源于新型冠状病毒流行株,更优选来源于原始株(Wu F, Zhao S, Yu B et al. Anew coronavirus associated with human respiratory disease in China. Nature.2020 Mar;579(7798):265-269. doi: 10.1038/s41586-020-2008-3. Epub 2020 Feb 3.Erratum in: Nature. 2020 Apr;580(7803):E7.)、新冠Alpha株、Beta株、Delta株(MaaranMichael Rajah, Mathieu Hubert, Elodie Bishop et.al. SARS-CoV-2 Alpha, Betaand Delta variants display enhanced Spike-mediated Syncytia Formation.bioRxiv preprint doi: https://doi.org/10.1101/2021.06.11.448011)、Lambda株(Izumi Kimura, Yusuke Kosugi, Jiaqi Wu et.al. SARS-CoV-2 Lambda variantexhibits higher 1 infectivity and immune resistance. bioRxiv preprint doi:https://doi.org/10.1101/2021.07.28.454085.)或Mu株(Keiya Uriu, Izumi Kimura,Kotaro Shirakawa et.al. Ineffective neutralization of the SARS-CoV-2 Muvariant 1 by convalescent and 2 vaccine sera. bioRxiv preprint doi: https://doi.org/10.1101/2021.09.06.459005)。在本发明中,所述S蛋白编码序列和新型冠状病毒的RBD蛋白编码序列可分别来自相同的新冠病毒流行株,也可来自不同的新冠病毒流行株。
本发明优选将所述S蛋白的编码序列和RBD蛋白的编码序列经密码子优化后插入到新城疫病毒的基因组中,优选将所述新型冠状病毒的S蛋白编码序列插入新城疫病毒载体的P基因序列和M基因序列之间;将所述新型冠状病毒的RBD蛋白编码序列插入新城疫病毒载体的P基因序列和M基因序列之间、M基因序列和F基因序列之间或HN基因序列和L基因序列之间。在本发明中,当将所述RBD蛋白编码基因插入到新城疫病毒载体的P基因和M基因之间时,优选还包括使用2A序列将S和RBD连接为一个ORF(开放阅读框)。本发明实施例中,所述新城疫病毒优选包括新城疫病毒LaSota株(已公开在CN112011521A)。
本发明所述S蛋白编码序列优选包括完整的S蛋白编码的核苷酸序列(AccessionNO. YP_009724390)或所述完整的S蛋白编码的核苷酸序列的变体。本发明所述变体优选包括对所述完整的S蛋白编码序列进行基因工程改造,所述基因工程改造的方法优选选自突变、删除、重排和融合其他来源的蛋白多肽中的一种或多种方法的组合。所述基因工程改造优选包括提高S蛋白的免疫原性的改造;更优选地,经所述基因工程改造后的S蛋白构象成为预融合(prefusion)状态更容易被抗原呈递细胞识别,或其与未经过基因工程改造的S蛋白相比,能够更大量地表达在细胞膜上,最终在于能够提高免疫原性和抗原识别,或与未经过基因工程改造的S蛋白相比,能够分泌到细胞外并形成多聚体被抗原呈递细胞识别;最优选地,所述突变优选包括将完整的S蛋白的K986和V987位氨基酸突变为P(K986P和V987P,简称S2P序列),或将第682~685位氨基酸RRAR突变为GSAS,或将完整的S蛋白的K986、V987、F817、 A892、A899和A942位氨基酸均突变为P(简称S6P序列);或将S蛋白的C末端KLHYT突变为ALAYT;所述删除包括在所述完整的S蛋白的C末端截短18 ~28个氨基酸;所述重排包括将S蛋白的主要抗原决定序列S1蛋白的NTD和RBD区域位置序列进行调换,形成RBD-NTD蛋白;所述融合其他来源的蛋白多肽包括在删除S蛋白的跨膜区和胞内区序列基础上,将S蛋白的胞外域(M1~Q1208氨基酸)与新城疫F蛋白的C端包含跨膜区和胞内区的氨基酸序列融合;或将RBD-NTD蛋白N端添加信号肽序列,C端添加融合蛋白多肽序列。本发明所述NTD区域优选包括V16~S305氨基酸序列;本发明优选在RBD区域序列,NTD区域序列和融合蛋白多肽序列之间使用蛋白Linker进行连接,没有特别说明时,以上所述Linker的氨基酸序列优选为((G)nS)m,n和m均代表0到4的自然数。
在本发明实施例中,对delta毒株经所述重排后,使用蛋白Linker序列对deltaRBD结构域(SEQ ID NO.43)和deltaNTD区域(SEQ ID NO.55)进行连接形成deltaRBD-NTD蛋白编码序列,并在deltaRBD-NTD蛋白编码序列的N端添加分泌信号肽HSA,C端分别添加融合蛋白多肽序列Folden和Bann,优选使用蛋白Linker连接各个蛋白多肽片段,分别形成deltaRBD-NTD-Folden(SEQ ID NO.43+GGGGSGG+ SEQ ID NO.55+GGSGSGGSG+ SEQ IDNO.3)和deltaRBD-NTD-Bann(SEQ ID NO.43+GSGSGG+ SEQ ID NO.55+GGGGSGGGGS+ SEQ IDNO.4)。
本发明所述RBD蛋白编码序列优选为上述S蛋白R319 -S591位氨基酸所形成的蛋白编码序列,或为上述S蛋白R319-F541位氨基酸所形成的蛋白编码序列,或为上述S蛋白R319-K537位氨基酸所形成的蛋白编码序列。
在本发明中,所述RBD蛋白编码序列优选还包括经过基因工程改造的含有RBD蛋白编码序列的RBD系列重组蛋白编码序列,所述基因工程改造优选包括:在所述RBD蛋白编码序列的N末端添加分泌信号肽序列,C末端添加融合蛋白多肽序列。在本发明中所述融合蛋白多肽序列应当被理解为用于形成融合蛋白的多肽序列,如人IgG抗体Fc序列(SEQ IDNO.1)、铁蛋白Ferritin序列(SEQ ID NO.2)、T4噬菌体fibritin蛋白C末端27个氨基酸形成的Folden序列(SEQ ID NO.3)或番茄从矮病毒环支架肽序列Bann(SEQ ID NO.4);或与RBD蛋白编码序列以二硫键形成二聚体的RBD(R319-K537)本身序列。
在本发明中,当融合蛋白多肽为RBD蛋白编码序列本身时候,RBD蛋白编码序列易选择S蛋白的R319-K537位氨基酸序列;所述Ferritin序列主要是幽门螺旋杆菌铁蛋白,为了防止其引起人体自身免疫,将幽门螺旋杆菌铁蛋白氨基酸序列前4个氨基酸去除,从而形成SEQ ID NO.2所示的序列。
本发明在所述RBD蛋白编码序列和所述融合蛋白多肽序列之间优选还包括Linker,所述Linker的氨基酸序列优选为((G)nS)m,n和m均代表0到4的自然数,本发明实施例的RBD系列重组蛋白优选使用GGGSGGGS作为Linker进行连接,但是不能仅将其认定为本发明的全部保护范围。在本发明中,当所述RBD重组蛋白是RBD蛋白编码序列(R319-K537)与RBD蛋白编码序列(R319-K537)通过分子内二硫键形成的二聚体时,此时RBD和RBD蛋白之间不再加Linker。
本发明所述分泌信号肽序列优选包括:人血清白蛋白(HSA)信号肽序列(SEQ IDNO.5)、人免疫球蛋白重链(VH)信号肽序列(SEQ ID NO.6),或人CD5信号肽序列(SEQ IDNO.7)。
在本发明中,N端分泌信号肽序列的添加,使得含RBD序列的相关抗原可以分泌到感染细胞外部,增加抗原的暴露,同时,C端所添加的融合蛋白多肽序列,通过RBD蛋白二聚体(人抗体Fc序列,RBD蛋白编码序列)或多聚体(铁蛋白Ferritin序列、T4噬菌体fibritin蛋白C末端27个氨基酸形成的Folden序列或番茄从矮病毒环支架肽序列Bann)结构,帮助RBD形成正确的蛋白折叠,增加RBD蛋白稳定性。其中,人抗体Fc序列还能够增强抗原的呈递效果。以上改造最终提高含RBD序列重组蛋白的免疫原性。
本发明对所述重组新城疫病毒载体的构建方法并没有特殊限定,优选包括同源重组。
本发明还提供了一种由上述重组新城疫病毒载体制备得到的疫苗株。
本发明所述制备的方法优选包括反向遗传学的方法。在本发明中,所述疫苗株的病毒复制可以在受精鸡胚中完成,生物安全二级厂房即可满足其生产要求。在本发明实施例中,当HA效价达到6log2或者6log2以上时,即可以进行免疫。
本发明所述疫苗株的本质是重组病毒,是一种病毒粒子,所述疫苗株的结构优选包括:上述重组新城疫病毒载体的编码序列和展示在重组新城疫病毒粒子表面的S蛋白或所述S蛋白的变体;
所述S蛋白的变体由完整的S蛋白编码序列的变体编码得到。
本发明还提供了上述重组新城疫病毒载体疫苗株在制备预防新型冠状病毒传播和感染疫苗中的应用。
本发明所述疫苗株可以被呼吸道细胞识别,产生粘膜免疫,并感染细胞,引起抗体水平升高,从而用于制备预防新型冠状病毒传播和感染的疫苗。本发明所述疫苗可通过吸鼻或注射途径接种。
本发明还提供了一种预防新型冠状病毒传播和感染疫苗的疫苗,所述疫苗包括上述疫苗株。
下面结合实施例对本发明提供的一种包含新型冠状病毒双抗原靶标序列组合的重组新城疫病毒载体及对应疫苗株和疫苗进行详细的说明,但是不能把它们理解为对本发明保护范围的限定。
实施例1
候选疫苗株NDV-S2P-VH-RBD-Fc、NDV-S2P- HSA-deltaRBD-Fc、NDV-S2P- HSA-RBD-Folden、NDV-S2P-M-HSA-RBD-TBSV-F、NDV-S2P -M-HSA-deltaRBD-Fc-F、NDV-S2P-M-HSA-deltaRBD-Ferritin-F,NDV-S2P/GSAS-HSA-deltaRBD-Fc、NDV-S2P/GSAS-HSA-deltaRBD-Folden、NDV-S6P-M-HSA-deltaRBD-Fc-F、NDV-S2P-HAS-scRBD、NDV-S2P-M-scRBD-F、NDV-deltaS2P/F87-M-HSA-deltaRBD-Fc-F、NDV-deltaRBD-NTD-Bann-M-HSA-deltaRBD-Fc-F、NDV-deltaRBD-NTD-Folden-M-HSA-deltaRBD-Fc-F以及比较例疫苗株NDV-S2P、NDV-deltaS2P/F87、和NDV-VH-RBD-Fc病毒载体设计与构建
S蛋白变体的设计如下:对新冠抗原原始株S蛋白进行K986P和V987P突变形成S2P编码序列,将S2P的第R682~R685位氨基酸突变为GSAS,形成S2P/GSAS编码序列, 进一步对S2P进行F817P、 A892P、A899P和A942P突变,形成S6P编码序列。同时对新冠Delta变异株S蛋白(SEQ ID NO.56)进行K984P和V985P突变,形成deltaS2P,在deltaS2P基础上,删除末端Y1207-1271T个氨基酸序列,并融合新城疫Lasota株病毒F蛋白(SEQ ID NO.48)L467到M553个氨基酸序列,形成deltaS2P/F87。对新冠Delta变异株S蛋白的主要抗原决定序列S1的NTD区域(V16~S303)和RBD(R317~S589)结构域进行重排,使用GSGSGG序列或GGGGSGG序列对deltaNTD区域和deltaRBD结构域进行连接形成RBD-NTD蛋白编码序列,并在RBD-NTD蛋白编码序列的N端添加分泌信号肽HSA,C端分别添加融合蛋白多肽序列Folden和Bann,使用蛋白Linker连接各个蛋白多肽片段,分别形成deltaRBD-NTD-Folden和deltaRBD-NTD-Bann。
RBD系列重组蛋白的设计如下:在新冠病毒原始株S蛋白受体结合区RBD(R319~S591)或delta变异株S蛋白受体结合区RBD(R317~S589)序列N端添加信号肽,C端添加融合蛋白多肽序列,并使用GGGSGGGS的编码序列连接RBD区域与融合蛋白多肽序列形成RBD系列重组蛋白,包括HSA-deltaRBD-Fc、VH-RBD-Fc、HSA-RBD-Folden、HSA-RBD-Bann 和HSA-deltaRBD-Ferritin。同时,在delta变异株S蛋白受体结合区RBD(R317~K535)序列N端添加信号肽,C端添加融合蛋白多肽序列,融合蛋白多肽序列为deltaRBD编码蛋白(R317-K535)本身,形成RBD重组蛋白HSA-scdeltaRBD(该重组蛋白的融合蛋白多肽为RBD编码序列本身,不需要在两个RBD序列之间添加Linker)。以上设计初步得到抗原S蛋白和RBD蛋白的基因工程改造序列。将抗原蛋白基因序列S2P、deltaS2P、S2P/GSAS、S6P、deltaS2P/F87、HSA-deltaRBD-Fc、VH-RBD-Fc、HSA-RBD-Folden、HSA-RBD-Bann、HSA-deltaRBD-Ferritin、HSA-scdeltaRBD、deltaRBD-NTD-Folden和deltaRBD-NTD-Bann序列经过密码子优化后,由金唯智生物有限公司进行基因合成。
基因合成后,对以上合成片段和NDV-LaSota载体(已公开在CN112011521A)分别按照Primer STAR酶说明书进行PCR扩增。将S2P基因序列、deltaS2P 基因序列、S2P/GSAS基因序列、S6P基因序列、deltaS2P/F87基因序列和VH-RBD-Fc基因序列通过同源重组的方式(NEBuilder® HiFi DNA Assembly试剂盒)分别克隆到新城疫病毒载体载体P-M之间,分别形成NDV-S2P重组病毒载体,NDV-deltaS2P重组病毒载体,NDV-S2P/GSAS重组病毒载体,NDV-S6P重组病毒载体、NDV-deltaS2P/F87重组病毒载体和NDV-VH-RBD-Fc重组病毒载体。
在NDV-S2P病毒载体基础上,删除终止密码子,添加P2A基因序列(P2A对应的氨基酸序列SEQ ID NO.8:ATNFSLLKQAGDVEENPGP)后,分别插入RBD系列基因,包括HSA-deltaRBD-Fc,VH-RBD-Fc,HSA-RBD-Folden,HSA-scdeltaRBD等。利用同源重组克隆方式分别构建得到NDV-S2P -HSA-deltaRBD-Fc、NDV-S2P-VH-RBD-Fc和NDV-S2P-HSA-RBD-Folden病毒载体。
在NDV-S2P/GSAS病毒载体基础上,删除终止密码子,添加P2A基因序列(P2A对应的氨基酸序列SEQ ID NO.8)后,插入HSA-deltaRBD-Fc。利用同源重组克隆方式构建得到NDV-S2P/GSAS-HSA-deltaRBD-Fc病毒载体。
同时,在NDV-S2P病毒载体基础上,将HSA-deltaRBD-Fc、HSA-RBD-Bann、HSA-deltaRBD-Ferritin和HSA-scRBD等基因序列插入到M-F基因之间,同源重组克隆得到NDV-S2P-M-HSA-deltaRBD-Fc-F、NDV-S2P-M-HSA-RBD-Bann-F、 NDV-S2P-M-HSA-deltaRBD-Ferritin-F和NDV-S2P-M-HSA-scdeltaRBD -F;在NDV-deltaS2P病毒载体基础上,将HSA-deltaRBD-Fc基因序列插入到M-F基因之间,同源重组克隆得到NDV-deltaS2P-M-HSA-deltaRBD-Fc-F;在NDV-S6P病毒载体基础上,将HSA-deltaRBD-Fc插入到M-F基因之间,同源重组克隆得到NDV-S6P-M-HSA-deltaRBD-Fc-F;在NDV-deltaS2P/F87病毒载体基础上,将HSA-deltaRBD-Fc插入到M-F基因之间,同源重组克隆得到NDV- deltaS2P/F87-M-HSA-deltaRBD-Fc-F。
在NDV-S2P-M-HSA-deltaRBD-Fc-F载体基础上,将deltaRBD-NTD-Folden和deltaRBD-NTD-Bann序列通过同源重组克隆插入到S2P位置,代替原载体S2P序列,得到NDV-deltaRBD-NTD-Folden-M-HSA-deltaRBD-Fc-F和NDV-deltaRBD-NTD-Bann-M-HSA-deltaRBD-Fc-F。图1展示了所构建载体的结构示意图。
将同源重组产物的病毒载转化进入大肠杆菌感受态细胞中,挑取单克隆送杭州擎科生物测序鉴定,序列比对完全正确后,使用LB 培养基对菌液扩大培养,按照TIANGEN无内毒素质粒提取试剂盒说明书要求进行质粒抽提,抽提质粒保存-20℃备用。
实施例2
1.候选疫苗株病毒拯救
将实施例1构建得到的所有重组NDV病毒载体,分别和病毒拯救辅助质粒pCI-NP,pCI-P和pCI-L,经Nano drop测定质粒浓度后,转染BHK21-T7细胞(已公开在CN112011521A)。具体转染过程为:将BHK21-T7细胞铺六孔板,待细胞密度达到70~80%时,按照Lipofectamin LTX说明书取500μL Opti-MEM于1.5 mL离心管中,随后按照1:1:1:1的比例分别加入实施例1中的重组NDV病毒载体,以及pCI-NP,pCI-P和pCI-L辅助质粒,每管质粒总量为4 μg,供1个孔使用。向上述各溶液中分别加入3μL Plus reagent,室温作用5 min后各加入9 μl Lipofectamine LTX,轻轻混匀,室温放置30 min;随后取出待转染的BHK21-T7细胞,弃去培养基,无菌PBS洗2次,加入1mL Opti-MEM培养基,将上述转染复合物逐滴滴加到培养皿中,37 ℃二氧化碳培养箱孵育6~8 hr后换液,24 hr后加入含0.08 μg/mL TPCK的opti-MEM培养基继续培养。72小时后,反复冻融细胞3次,并将混合物接种9~11日龄SPF鸡胚,0.2~0.3 mL/胚,37℃培养24hr。弃去24hr内的死胚,收获72 hr内收获存活的鸡胚尿囊液,逐个测定血凝效价。最后,收取有血凝价且存活的鸡胚尿囊液。离心后分装,0.5~1 mL/管,冻存于-80 ℃冰箱。
2. 重组病毒PCR鉴定及测序分析
将收获的病毒,利用AXYGEN病毒基因组RNA提取试剂盒提取病毒基因组RNA,随后根据HiScript III 1st Strand cDNA Synthesis Kit试剂盒说明书要求对提取的RNA进行反转录,合成首链cDNA。
合成后,进行PCR凝胶电泳鉴定(图2),PCR反应体系和反应条件按照喏唯赞HiscriptII One Step RT-PCR kit说明书要求设置。PCR鉴定及测序合格后,进行下一步实验。
实施例3
重组NDV载体病毒疫苗株体外评价
1.重组疫苗候选株HA效价测定
将收取的NDV各个病毒疫苗候选株稀释至21后,相同病毒疫苗株多个样品混合接种9~11日龄鸡胚尿囊液,使用医用胶布将接种鸡胚封口后置34 ℃培养,72小时后,取尿囊液100 μl置于96孔V型血凝板,用PBS将尿囊液做10个梯度系列稀释,同时取50 μl PBS设阴性对照,即可读取HA效价。
结果可见,初始拯救病毒的HA效价大部分在4-8log2,经过传代适应后,疫苗候选株都产生较高HA效价,最高可达到10log2以上(表3)。
2.重组NDV载体病毒疫苗株感染Vero细胞后的抗原表达情况
将Vero-E6细胞(购自中国科学院细胞库,目录号GNO17)传代到24孔板中长至单层备用,每孔分别加入实施例2中所拯救重组NDV载体病毒,包括NDV-S2P、NDV-VH-RBD-Fc、NDV-S2P -HSA-deltaRBD-Fc、NDV-deltaS2P -HSA-deltaRBD-Fc、NDV-S2P-VH-RBD-Fc、NDV-S2P -HSA-RBD-Folden、NDV-S2P-M-HSA-RBD-Bann-F、NDV-S2P- M-HSA-deltaRBD-Fc-F、NDV-S2P- M-HSA-deltaRBD-Ferritin-F、NDV-S2P/GSAS-HSA-deltaRBD-Fc、NDV-S6P-M-HSA-deltaRBD-Fc-F、NDV-S2P- HSA-scdeltaRBD 、NDV-S2P-M-HSA-scdeltaRBD-F、NDV-deltaS2P/F87和NDV-deltaS2P/F87-M-HSA-deltaRBD-Fc-F以及病毒原液NDV-COVS(已公开在CN112011521A)。所有病毒HA效价控制在6-7log2之间,对长至单层的细胞板,用PBS洗两遍,分别加入上述病毒液,100 μl/孔,每孔补加300 μl 含0.1μg/mL TPCK的opti-MEM。空白对照孔不接病毒。放于37 ℃,5% CO2培养箱中培养至36hr。检测时,分别取150 μl上清样品和150 μl吹打悬浮后经过反复3次冻融的细胞及细胞培养液检测RBD蛋白浓度。
RBD浓度的测定使用双抗体夹心酶联免疫法,按照义翘神州SARS-CoV-2 (2019-nCoV) Spike RBD ELISA Kit试剂盒说明书要求,配置1x洗涤液和1x稀释液,对标准品进行复溶,并梯度稀释标准品,对待测的细胞上清液以及细胞冻融液进行100倍稀释,按照说明书要求对标准品和待测的细胞上清液以及细胞冻融液进行抗原孵育、洗涤、酶标检测抗体孵育、洗涤、显色、终止等操作后,在20 min内读取450 nm的光吸收值。
理论上,细胞上清液中将检测到大量分泌的RBD重组蛋白或S蛋白重排变体、微量包装好的病毒粒子表面的S蛋白及其变体以及极少量的坏死细胞膜上的S蛋白及其变体,细胞冻融液中将检测到合成的RBD重组蛋白或S蛋白重排变体、微量包装好的病毒粒子表面的S蛋白以及破裂细胞细胞膜上的S蛋白及其变体。结果如图3所示,相对于单个不进行改造的S蛋白(NDV-COVS),NDV-S2P感染Vero-E6细胞后的抗原表达量明显升高,说明S2P蛋白成功表达。NDV-S2P实验组上清和冻融液中检测到抗原信号,同时,NDV-VH-RBD-Fc实验组上清和冻融液中也检测到大量RBD抗原表达,且表达量高于NDV-S2P,显示该位置处RBD重组蛋白成功表达并分泌到上清中。对于NDV-deltaRBD-NTD-Folden-M-HSA-deltaRBD-Fc-F和NDV-deltaRBD-NTD-Bann-M-HSA-deltaRBD-Fc-F重组病毒,上清和冻融液中的RBD表达量都分别显著高于其他毒株,进一步表明S蛋白变体和RBD重组蛋白正确表达,并且大量分泌到细胞外。相对于单独的NDV-S2P重组病毒,当新城疫病毒基因组P基因和M基因之间、S蛋白编码序列及其变体之后或者新城疫病毒基因组M基因和F基因位置之间插入RBD系列重组蛋白时,细胞上清和冻融液中新冠RBD抗原表达量明显增加,上清抗原表达量普遍提高约2-5倍,冻融液抗原表达量普遍提高约2-4倍。同时,NDV-S2P/GSAS-HSA-deltaRBD-Fc和NDV-S6P-M-HSA-deltaRBD-Fc-F双抗原靶标疫苗株的抗原表达量也相对较高。对于NDV-deltaS2P/F87,当新城疫病毒基因组M基因和F基因之间插入HSA-deltaRBD-Fc时,新冠RBD抗原表达量同样大幅提升。可见,新冠S蛋白编码序列及其变体协同新冠RBD系列重组蛋白策略的抗原表达量优于单个S蛋白。
实施例4
小鼠免疫实验
选择实施例3中有代表性的重组NDV载体病毒疫苗株,NDV-S2P、NDV-deltaS2P/F87,NDV-VH-RBD-Fc、NDV-S2P-VH-RBD-Fc、NDV-S2P- M-HSA-deltaRBD-Fc-F、NDV-deltaS2P-M-HSA-deltaRBD-Fc-F、NDV-S2P-M-HSA-RBD-Bann-F、NDV-S2P- M-HSA-deltaRBD-Ferritin-F、NDV-S2P- HSA-scdeltaRBD、NDV-S2P-M-HSA-scdeltaRBD-F、NDV-S2P/GSAS-HSA-deltaRBD-Fc 、NDV-S6P-M-HSA-deltaRBD-Fc-F、NDV-deltaS2P/F87-M-HSA-deltaRBD-Fc-F、NDV-deltaRBD-NTD-Folden-M-HSA-deltaRBD-Fc-F和NDV-deltaRBD-NTD-Bann-M-HSA-deltaRBD-Fc-F对4-5周龄Balb\c雌性小鼠进行免疫,测试新冠S蛋白IgG抗体浓度和NDV载体本身IgG抗体浓度。
在免疫前,需要先确定病毒HA效价,HA效价测定方法同实施例3。当HA效价达到6log2及以上,即可用于免疫。
免疫途径选择鼻腔免疫和肌肉注射免疫。鼻腔免疫方法为:使用5%水合氯醛腹腔注射,剂量为体重每20g小鼠腹腔注射0.1mL使动物麻醉,麻醉后用移液器进行滴鼻免疫,25μl/鼻孔。肌肉注射免疫方法为:固定小鼠,用0.3mL微量注射器对小鼠后腿肌肉注射100 μl/只小鼠。共进行两次免疫,首免后7天进行二次加强免疫。首免后14 天,采集血清检测新冠S蛋白结合抗体以及NDV病毒相关蛋白IgG抗体。
免疫小鼠血清S蛋白IgG抗体滴度按照义翘神州(SARS-CoV-2 (2019-nCoV))Spike S1抗体滴度检测试剂盒说明书进行检测,重组NDV病毒疫苗株小鼠血清新城疫病毒抗体滴度按照爱德士新城疫病毒抗体检测试剂盒说明书进行检测。数据显著性差异分析使用非配对Student’t T检验。
滴鼻免疫结果如图4中a所示,首免后14天,所有疫苗株血清新冠S1蛋白IgG抗体总体升高。相对于NDV-S2P以及NDV-VH-RBD-Fc免疫血清S1 IgG抗体,疫苗株NDV-S2P -VH-RBD-Fc、NDV-S2P- HSA-scdeltaRBD、NDV-S2P-M- HSA-scdeltaRBD-F、NDV-S2P -M-HSA-deltaRBD-Fc-F、NDV-deltaS2P -M-HSA-deltaRBD-Fc-F、NDV-S2P -M-HSA-deltaRBD-Bann-F、 NDV-S2P -M-HSA-deltaRBD-Ferritin-F、NDV-S2P/GSAS-HSA-deltaRBD-Fc、 NDV-S6P-M-HSA-deltaRBD-Fc、NDV-deltaRBD-NTD-Bann-M-HSA-deltaRBD-Fc-F和NDV-deltaRBD-NTD-Folden-M-HSA-deltaRBD-Fc-F免疫血清S1 IgG抗体均值均有一定程度升高,特别对于NDV-S2P -M-HSA-RBD-Fc-F、NDV-deltaS2P -M-HSA-deltaRBD-Fc-F、NDV-S6P-M-HSA-deltaRBD-Fc-F、NDV-deltaRBD-NTD-Bann-M-HSA-deltaRBD-Fc-F和NDV-deltaRBD-NTD-Folden-M-HSA-deltaRBD-Fc-F重组病毒疫苗株,小鼠免疫血清S1 IgG抗体滴度相对于单个抗原的疫苗株免疫效果显著升高。同时, NDV-deltaS2P/F87-M-HSA-deltaRBD-Fc-F疫苗株免疫后抗体IgG滴度相对于NDV-deltaS2P/F87疫苗株也大量提高。肌注免疫结果如图5中a所示,首免后14天,与NDV-S2P疫苗株相比,NDV-deltaS2P-M-HSA-deltaRBD-Fc-F、NDV-S2P/GSAS-HSA-deltaRBD-Bann、NDV-S2P-M-HSA-deltaRBD-Fc-F、NDV-S2P-scdeltaRBD和NDV-deltaRBD-NTD-Folden-M-HSA-deltaRBD-Fc-F疫苗株免疫后血清新冠S1 IgG均有显著性升高。
另外,免疫后小鼠血清NDV病毒IgG抗体滴度结果表明(图4中b和图5中b),NDV-S2P-VH-RBD-Fc、NDV-S2P-M- HSA-scdeltaRBD-F、NDV-S2P -M-HSA-deltaRBD-Fc-F、NDV-deltaS2P -M-HSA-deltaRBD-Fc-F、NDV-S2P -M-HSA-deltaRBD-Ferritin-F和NDV-deltaS2P/F87-M-HSA-deltaRBD-Fc-F的病毒载体本身抗体滴度相对于NDV-S2P均有显著降低。由上可见,新冠S基因及其变体结合RBD系列重组蛋白基因的重组NDV病毒策略可以提高疫苗株免疫原性的发挥,有效的降低了NDV病毒自身所引起的抗体水平,对疫苗免疫作用的发挥以及进一步降低病毒载体本身毒副作用有重要意义(图5)。
以上所述仅是本发明的优选实施方式,应当指出,对于本技术领域的普通技术人员来说,在不脱离本发明原理的前提下,还可以做出若干改进和润饰,这些改进和润饰也应视为本发明的保护范围。
序列表
<110> 浙江迪福润丝生物科技有限公司
<120> 一种包含新型冠状病毒双抗原靶标序列组合的重组新城疫病毒载体及相应疫苗株和疫苗
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Asp Lys Thr His Thr Cys Pro Pro Cys Pro Ala Pro Glu Leu Leu Gly
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Gly Pro Ser Val Phe Leu Phe Pro Pro Lys Pro Lys Asp Thr Leu Met
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Ile Ser Arg Thr Pro Gly Val Thr Cys Val Val Val Asp Val Ser His
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Glu Asp Pro Glu Val Lys Phe Asn Trp Tyr Val Asp Gly Val Glu Val
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His Asn Ala Lys Thr Lys Pro Arg Glu Glu Gln Tyr Asn Ser Thr Tyr
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Arg Val Val Ser Val Leu Thr Val Leu His Gln Asp Trp Leu Asn Gly
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Lys Glu Tyr Lys Cys Lys Val Ser Asn Lys Ala Leu Pro Ala Pro Ile
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Glu Lys Thr Ile Ser Lys Ala Lys Gly Gln Pro Arg Glu Pro Gln Val
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Tyr Thr Leu Pro Pro Ser Arg Asp Glu Leu Thr Lys Asn Gln Val Ser
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Leu Thr Cys Leu Val Lys Gly Phe Tyr Pro Ser Asp Ile Ala Val Glu
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Trp Glu Ser Asn Gly Gln Pro Glu Asn Asn Tyr Lys Thr Thr Pro Pro
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Val Leu Asp Ser Asp Gly Ser Phe Phe Leu Tyr Ser Lys Leu Thr Val
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Asp Lys Ser Arg Trp Gln Gln Gly Asn Val Phe Ser Cys Ser Val Met
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His Glu Ala Leu His Asn His Tyr Thr Gln Lys Ser Leu Ser Leu Ser
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Pro Gly Lys
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Asp Ile Ile Lys Leu Leu Asn Glu Gln Val Asn Lys Glu Met Asn Ser
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Ser Asn Leu Tyr Met Ser Met Ser Ser Trp Cys Tyr Thr His Ser Leu
20 25 30
Asp Gly Ala Gly Leu Phe Leu Phe Asp His Ala Ala Glu Glu Tyr Glu
35 40 45
His Ala Lys Lys Leu Ile Val Phe Leu Asn Glu Asn Asn Val Pro Val
50 55 60
Gln Leu Thr Ser Ile Ser Ala Pro Glu His Lys Phe Glu Gly Leu Thr
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Gln Ile Phe Gln Lys Ala Tyr Glu His Glu Gln His Ile Ser Glu Ser
85 90 95
Ile Asn Asn Ile Val Asp His Ala Ile Lys Ser Lys Asp His Ala Thr
100 105 110
Phe Asn Phe Leu Gln Trp Tyr Val Ala Glu Gln His Glu Glu Glu Val
115 120 125
Leu Phe Lys Asp Ile Leu Asp Lys Ile Glu Leu Ile Gly Asn Glu Asn
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His Gly Leu Tyr Leu Ala Asp Gln Tyr Val Lys Gly Ile Ala Lys Ser
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Arg Lys Ser
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Gly Tyr Ile Pro Glu Ala Pro Arg Asp Gly Gln Ala Tyr Val Arg Lys
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Asp Gly Glu Trp Val Phe Leu Ser Thr Phe Leu
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Ile Asn His Val Gly Gly Thr Gly Gly Ala Ile Met Ala Pro Val Ala
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Met Lys His Leu Trp Phe Phe Leu Leu Leu Val Ala Ala Pro Arg Trp
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Ala Thr Asn Phe Ser Leu Leu Lys Gln Ala Gly Asp Val Glu Glu Asn
1 5 10 15
Pro Gly Pro
<210> 9
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<213> 人工序列(Artificial Sequence)
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catgttcgtg tttctggtgc tg 22
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<212> DNA
<213> 人工序列(Artificial Sequence)
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ttaaactcag gtgtagtgca gtttcacg 28
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<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 11
tgagtttaaa cttagaaaaa atacgggtag aattg 35
<210> 12
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<212> DNA
<213> 人工序列(Artificial Sequence)
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accaaacaaa gatttggtga atg 23
<210> 13
<211> 40
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 13
caattattct tgagtgtagt ctcgtcattc accaaatctt 40
<210> 14
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 14
agaaacacga acatggtg 18
<210> 15
<211> 41
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 15
ttagaaaaaa tacgggtaga agccaccatg aagcacctgt g 41
<210> 16
<211> 42
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 16
ccgtattttt tctaaatctt aaatgtagct agattaatta cg 42
<210> 17
<211> 39
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 17
cgggtagaag ccaccatgaa gcacctgtgg ttctttctg 39
<210> 18
<211> 37
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 18
ccgtattttt tctaattatt tacccggaga cagggag 37
<210> 19
<211> 48
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 19
ttagaaaaaa tacgggtaga attggagtgc cccaattgtg ccaagatg 48
<210> 20
<211> 57
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 20
ggtggcttct acccgtattt tttctaagtt taaactcagg tgtagtgcag tttcacg 57
<210> 21
<211> 38
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 21
cgggtagaag ccaccatgaa atgggtcacc tttatcag 38
<210> 22
<211> 35
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 22
ccgtattttt tctaatcatt tgccggggct cagag 35
<210> 23
<211> 37
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 23
ccgtattttt tctaattaca ggaaggtgct caggaac 37
<210> 24
<211> 59
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 24
ggtggcggct tctacccgta ttttttctaa gtttaaactc aggtgtagtg cagtttcac 59
<210> 25
<211> 35
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 25
gttttttgtg tcatgtcatt tgccggggct cagag 35
<210> 26
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 26
catgacacaa aaaactaatc tg 22
<210> 27
<211> 52
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 27
ggtggcttct acccgtattt tttctaaatg atctgggtga gtgggcggag cg 52
<210> 28
<211> 37
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 28
gttttttgtg tcatgtcagc tctttctgct cttggcg 37
<210> 29
<211> 38
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 29
gttttttgtg tcatgtcaag agcccacgag ctgtcttg 38
<210> 30
<211> 55
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 30
ggtggcggct tctacccgta ttttttctaa atgatctggg tgagtgggcg gagcg 55
<210> 31
<211> 40
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 31
gttttttgtg tcatgtcatt tgttcttcac gagattggtg 40
<210> 32
<211> 40
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 32
ccgtattttt tctaatcatt tgttcttcac gagattggtg 40
<210> 33
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 33
atgttcgtgt ttctggtg 18
<210> 34
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 34
tctggctctc cgagggctgt ttg 23
<210> 35
<211> 40
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 35
cctcggagag ccagatctgt ggccagccag agcatcattg 40
<210> 36
<211> 40
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 36
ctaagtttaa acttacagga aggtgctcag gaacacccac 40
<210> 37
<211> 35
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 37
taagtttaaa cttagaaaaa atacgggtag aattg 35
<210> 38
<211> 45
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 38
cagaaacacg aacatggtgg cggttctacc cgtatttttt ctaag 45
<210> 39
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 39
caccatgttc gtgtttctgg tg 22
<210> 40
<211> 33
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 40
tctggctctc cgtctgctgt tggtctgtgt ctg 33
<210> 41
<211> 36
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 41
agacggagag ccagatctgt ggccagccag agcatc 36
<210> 42
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 42
ttaaactcac atctttgtgg tggctc 26
<210> 43
<211> 273
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 43
Arg Val Gln Pro Thr Glu Ser Ile Val Arg Phe Pro Asn Ile Thr Asn
1 5 10 15
Leu Cys Pro Phe Gly Glu Val Phe Asn Ala Thr Arg Phe Ala Ser Val
20 25 30
Tyr Ala Trp Asn Arg Lys Arg Ile Ser Asn Cys Val Ala Asp Tyr Ser
35 40 45
Val Leu Tyr Asn Ser Ala Ser Phe Ser Thr Phe Lys Cys Tyr Gly Val
50 55 60
Ser Pro Thr Lys Leu Asn Asp Leu Cys Phe Thr Asn Val Tyr Ala Asp
65 70 75 80
Ser Phe Val Ile Arg Gly Asp Glu Val Arg Gln Ile Ala Pro Gly Gln
85 90 95
Thr Gly Lys Ile Ala Asp Tyr Asn Tyr Lys Leu Pro Asp Asp Phe Thr
100 105 110
Gly Cys Val Ile Ala Trp Asn Ser Asn Asn Leu Asp Ser Lys Val Gly
115 120 125
Gly Asn Tyr Asn Tyr Arg Tyr Arg Leu Phe Arg Lys Ser Asn Leu Lys
130 135 140
Pro Phe Glu Arg Asp Ile Ser Thr Glu Ile Tyr Gln Ala Gly Ser Lys
145 150 155 160
Pro Cys Asn Gly Val Glu Gly Phe Asn Cys Tyr Phe Pro Leu Gln Ser
165 170 175
Tyr Gly Phe Gln Pro Thr Asn Gly Val Gly Tyr Gln Pro Tyr Arg Val
180 185 190
Val Val Leu Ser Phe Glu Leu Leu His Ala Pro Ala Thr Val Cys Gly
195 200 205
Pro Lys Lys Ser Thr Asn Leu Val Lys Asn Lys Cys Val Asn Phe Asn
210 215 220
Phe Asn Gly Leu Thr Gly Thr Gly Val Leu Thr Glu Ser Asn Lys Lys
225 230 235 240
Phe Leu Pro Phe Gln Gln Phe Gly Arg Asp Ile Ala Asp Thr Thr Asp
245 250 255
Ala Val Arg Asp Pro Gln Thr Leu Glu Ile Leu Asp Ile Thr Pro Cys
260 265 270
Ser
<210> 44
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 44
aaggtccaac tctccaagcg g 21
<210> 45
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 45
gtcctcctta ctatcagtcc aca 23
<210> 46
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 46
caggtggcca agatactctg 20
<210> 47
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 47
cttggtagaa ggtctggag 19
<210> 48
<211> 553
<212> PRT
<213> 新城疫病毒(Newcastle disease virus)
<400> 48
Met Gly Ser Arg Pro Ser Thr Lys Asn Pro Ala Pro Met Met Leu Thr
1 5 10 15
Ile Arg Val Ala Leu Val Leu Ser Cys Ile Cys Pro Ala Asn Ser Ile
20 25 30
Asp Gly Arg Pro Leu Ala Ala Ala Gly Ile Val Val Thr Gly Asp Lys
35 40 45
Ala Val Asn Ile Tyr Thr Ser Ser Gln Thr Gly Ser Ile Ile Val Lys
50 55 60
Leu Leu Pro Asn Leu Pro Lys Asp Lys Glu Ala Cys Ala Lys Ala Pro
65 70 75 80
Leu Asp Ala Tyr Asn Arg Thr Leu Thr Thr Leu Leu Thr Pro Leu Gly
85 90 95
Asp Ser Ile Arg Arg Ile Gln Glu Ser Val Thr Thr Ser Gly Gly Gly
100 105 110
Arg Gln Gly Arg Leu Ile Gly Ala Ile Ile Gly Gly Val Ala Leu Gly
115 120 125
Val Ala Thr Ala Ala Gln Ile Thr Ala Ala Ala Ala Leu Ile Gln Ala
130 135 140
Lys Gln Asn Ala Ala Asn Ile Leu Arg Leu Lys Glu Ser Ile Ala Ala
145 150 155 160
Thr Asn Glu Ala Val His Glu Val Thr Asp Gly Leu Ser Gln Leu Ala
165 170 175
Val Ala Val Gly Lys Met Gln Gln Phe Val Asn Asp Gln Phe Asn Lys
180 185 190
Thr Ala Gln Glu Leu Asp Cys Ile Lys Ile Ala Gln Gln Val Gly Val
195 200 205
Glu Leu Asn Leu Tyr Leu Thr Glu Leu Thr Thr Val Phe Gly Pro Gln
210 215 220
Ile Thr Ser Pro Ala Leu Asn Lys Leu Thr Ile Gln Ala Leu Tyr Asn
225 230 235 240
Leu Ala Gly Gly Asn Met Asp Tyr Leu Leu Thr Lys Leu Gly Val Gly
245 250 255
Asn Asn Gln Leu Ser Ser Leu Ile Gly Ser Gly Leu Ile Thr Gly Asn
260 265 270
Pro Ile Leu Tyr Asp Ser Gln Thr Gln Leu Leu Gly Ile Gln Val Thr
275 280 285
Leu Pro Ser Val Gly Asn Leu Asn Asn Met Arg Ala Thr Tyr Leu Glu
290 295 300
Thr Leu Ser Val Ser Thr Thr Arg Gly Phe Ala Ser Ala Leu Val Pro
305 310 315 320
Lys Val Val Thr Gln Val Gly Ser Val Ile Glu Glu Leu Asp Thr Ser
325 330 335
Tyr Cys Ile Glu Thr Asp Leu Asp Leu Tyr Cys Thr Arg Ile Val Thr
340 345 350
Phe Pro Met Ser Pro Gly Ile Tyr Ser Cys Leu Ser Gly Asn Thr Ser
355 360 365
Ala Cys Met Tyr Ser Lys Thr Glu Gly Ala Leu Thr Thr Pro Tyr Met
370 375 380
Thr Ile Lys Gly Ser Val Ile Ala Asn Cys Lys Met Thr Thr Cys Arg
385 390 395 400
Cys Val Asn Pro Pro Gly Ile Ile Ser Gln Asn Tyr Gly Glu Ala Val
405 410 415
Ser Leu Ile Asp Lys Gln Ser Cys Asn Val Leu Ser Leu Gly Gly Ile
420 425 430
Thr Leu Arg Leu Ser Gly Glu Phe Asp Val Thr Tyr Gln Lys Asn Ile
435 440 445
Ser Ile Gln Asp Ser Gln Val Ile Ile Thr Gly Asn Leu Asp Ile Ser
450 455 460
Thr Glu Leu Gly Asn Val Asn Asn Ser Ile Ser Asn Ala Leu Asn Lys
465 470 475 480
Leu Glu Glu Ser Asn Arg Lys Leu Asp Lys Val Asn Val Lys Leu Thr
485 490 495
Ser Thr Ser Ala Leu Ile Thr Tyr Ile Val Leu Thr Ile Ile Ser Leu
500 505 510
Val Phe Gly Ile Leu Ser Leu Ile Leu Ala Cys Tyr Leu Met Tyr Lys
515 520 525
Gln Lys Ala Gln Gln Lys Thr Leu Leu Trp Leu Gly Asn Asn Thr Leu
530 535 540
Asp Gln Met Arg Ala Thr Thr Lys Met
545 550
<210> 49
<211> 38
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 49
agaagtgccg ccaccatgaa atgggtcacc tttatcag 38
<210> 50
<211> 35
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 50
ctaagtttaa actcaagagc ccacgagctg tcttg 35
<210> 51
<211> 57
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 51
ggtggcggca cttctacccg tattttttct aagtttaaac atcttaaatg tagctag 57
<210> 52
<211> 38
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 52
ggtagaaccg ccaccatgaa atgggtcacc tttatcag 38
<210> 53
<211> 34
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 53
ctaagtttaa actcacagga aggtgctcag gaac 34
<210> 54
<211> 54
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 54
ggtggcggtt ctacccgtat tttttctaag tttaaacatc ttaaatgtag ctag 54
<210> 55
<211> 288
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 55
Val Asn Leu Arg Thr Arg Thr Gln Leu Pro Pro Ala Tyr Thr Asn Ser
1 5 10 15
Phe Thr Arg Gly Val Tyr Tyr Pro Asp Lys Val Phe Arg Ser Ser Val
20 25 30
Leu His Ser Thr Gln Asp Leu Phe Leu Pro Phe Phe Ser Asn Val Thr
35 40 45
Trp Phe His Ala Ile His Val Ser Gly Thr Asn Gly Thr Lys Arg Phe
50 55 60
Asp Asn Pro Val Leu Pro Phe Asn Asp Gly Val Tyr Phe Ala Ser Thr
65 70 75 80
Glu Lys Ser Asn Ile Ile Arg Gly Trp Ile Phe Gly Thr Thr Leu Asp
85 90 95
Ser Lys Thr Gln Ser Leu Leu Ile Val Asn Asn Ala Thr Asn Val Val
100 105 110
Ile Lys Val Cys Glu Phe Gln Phe Cys Asn Asp Pro Phe Leu Asp Val
115 120 125
Tyr Tyr His Lys Asn Asn Lys Ser Trp Met Glu Ser Gly Val Tyr Ser
130 135 140
Ser Ala Asn Asn Cys Thr Phe Glu Tyr Val Ser Gln Pro Phe Leu Met
145 150 155 160
Asp Leu Glu Gly Lys Gln Gly Asn Phe Lys Asn Leu Arg Glu Phe Val
165 170 175
Phe Lys Asn Ile Asp Gly Tyr Phe Lys Ile Tyr Ser Lys His Thr Pro
180 185 190
Ile Asn Leu Val Arg Asp Leu Pro Gln Gly Phe Ser Ala Leu Glu Pro
195 200 205
Leu Val Asp Leu Pro Ile Gly Ile Asn Ile Thr Arg Phe Gln Thr Leu
210 215 220
Leu Ala Leu His Arg Ser Tyr Leu Thr Pro Gly Asp Ser Ser Ser Gly
225 230 235 240
Trp Thr Ala Gly Ala Ala Ala Tyr Tyr Val Gly Tyr Leu Gln Pro Arg
245 250 255
Thr Phe Leu Leu Lys Tyr Asn Glu Asn Gly Thr Ile Thr Asp Ala Val
260 265 270
Asp Cys Ala Leu Asp Pro Leu Ser Glu Thr Lys Cys Thr Leu Lys Ser
275 280 285
<210> 56
<211> 1271
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 56
Met Phe Val Phe Leu Val Leu Leu Pro Leu Val Ser Ser Gln Cys Val
1 5 10 15
Asn Leu Arg Thr Arg Thr Gln Leu Pro Pro Ala Tyr Thr Asn Ser Phe
20 25 30
Thr Arg Gly Val Tyr Tyr Pro Asp Lys Val Phe Arg Ser Ser Val Leu
35 40 45
His Ser Thr Gln Asp Leu Phe Leu Pro Phe Phe Ser Asn Val Thr Trp
50 55 60
Phe His Ala Ile His Val Ser Gly Thr Asn Gly Thr Lys Arg Phe Asp
65 70 75 80
Asn Pro Val Leu Pro Phe Asn Asp Gly Val Tyr Phe Ala Ser Thr Glu
85 90 95
Lys Ser Asn Ile Ile Arg Gly Trp Ile Phe Gly Thr Thr Leu Asp Ser
100 105 110
Lys Thr Gln Ser Leu Leu Ile Val Asn Asn Ala Thr Asn Val Val Ile
115 120 125
Lys Val Cys Glu Phe Gln Phe Cys Asn Asp Pro Phe Leu Asp Val Tyr
130 135 140
Tyr His Lys Asn Asn Lys Ser Trp Met Glu Ser Gly Val Tyr Ser Ser
145 150 155 160
Ala Asn Asn Cys Thr Phe Glu Tyr Val Ser Gln Pro Phe Leu Met Asp
165 170 175
Leu Glu Gly Lys Gln Gly Asn Phe Lys Asn Leu Arg Glu Phe Val Phe
180 185 190
Lys Asn Ile Asp Gly Tyr Phe Lys Ile Tyr Ser Lys His Thr Pro Ile
195 200 205
Asn Leu Val Arg Asp Leu Pro Gln Gly Phe Ser Ala Leu Glu Pro Leu
210 215 220
Val Asp Leu Pro Ile Gly Ile Asn Ile Thr Arg Phe Gln Thr Leu Leu
225 230 235 240
Ala Leu His Arg Ser Tyr Leu Thr Pro Gly Asp Ser Ser Ser Gly Trp
245 250 255
Thr Ala Gly Ala Ala Ala Tyr Tyr Val Gly Tyr Leu Gln Pro Arg Thr
260 265 270
Phe Leu Leu Lys Tyr Asn Glu Asn Gly Thr Ile Thr Asp Ala Val Asp
275 280 285
Cys Ala Leu Asp Pro Leu Ser Glu Thr Lys Cys Thr Leu Lys Ser Phe
290 295 300
Thr Val Glu Lys Gly Ile Tyr Gln Thr Ser Asn Phe Arg Val Gln Pro
305 310 315 320
Thr Glu Ser Ile Val Arg Phe Pro Asn Ile Thr Asn Leu Cys Pro Phe
325 330 335
Gly Glu Val Phe Asn Ala Thr Arg Phe Ala Ser Val Tyr Ala Trp Asn
340 345 350
Arg Lys Arg Ile Ser Asn Cys Val Ala Asp Tyr Ser Val Leu Tyr Asn
355 360 365
Ser Ala Ser Phe Ser Thr Phe Lys Cys Tyr Gly Val Ser Pro Thr Lys
370 375 380
Leu Asn Asp Leu Cys Phe Thr Asn Val Tyr Ala Asp Ser Phe Val Ile
385 390 395 400
Arg Gly Asp Glu Val Arg Gln Ile Ala Pro Gly Gln Thr Gly Lys Ile
405 410 415
Ala Asp Tyr Asn Tyr Lys Leu Pro Asp Asp Phe Thr Gly Cys Val Ile
420 425 430
Ala Trp Asn Ser Asn Asn Leu Asp Ser Lys Val Gly Gly Asn Tyr Asn
435 440 445
Tyr Arg Tyr Arg Leu Phe Arg Lys Ser Asn Leu Lys Pro Phe Glu Arg
450 455 460
Asp Ile Ser Thr Glu Ile Tyr Gln Ala Gly Ser Lys Pro Cys Asn Gly
465 470 475 480
Val Glu Gly Phe Asn Cys Tyr Phe Pro Leu Gln Ser Tyr Gly Phe Gln
485 490 495
Pro Thr Asn Gly Val Gly Tyr Gln Pro Tyr Arg Val Val Val Leu Ser
500 505 510
Phe Glu Leu Leu His Ala Pro Ala Thr Val Cys Gly Pro Lys Lys Ser
515 520 525
Thr Asn Leu Val Lys Asn Lys Cys Val Asn Phe Asn Phe Asn Gly Leu
530 535 540
Thr Gly Thr Gly Val Leu Thr Glu Ser Asn Lys Lys Phe Leu Pro Phe
545 550 555 560
Gln Gln Phe Gly Arg Asp Ile Ala Asp Thr Thr Asp Ala Val Arg Asp
565 570 575
Pro Gln Thr Leu Glu Ile Leu Asp Ile Thr Pro Cys Ser Phe Gly Gly
580 585 590
Val Ser Val Ile Thr Pro Gly Thr Asn Thr Ser Asn Gln Val Ala Val
595 600 605
Leu Tyr Gln Gly Val Asn Cys Thr Glu Val Pro Val Ala Ile His Ala
610 615 620
Asp Gln Leu Thr Pro Thr Trp Arg Val Tyr Ser Thr Gly Ser Asn Val
625 630 635 640
Phe Gln Thr Arg Ala Gly Cys Leu Ile Gly Ala Glu His Val Asn Asn
645 650 655
Ser Tyr Glu Cys Asp Ile Pro Ile Gly Ala Gly Ile Cys Ala Ser Tyr
660 665 670
Gln Thr Gln Thr Asn Ser Arg Arg Arg Ala Arg Ser Val Ala Ser Gln
675 680 685
Ser Ile Ile Ala Tyr Thr Met Ser Leu Gly Ala Glu Asn Ser Val Ala
690 695 700
Tyr Ser Asn Asn Ser Ile Ala Ile Pro Thr Asn Phe Thr Ile Ser Val
705 710 715 720
Thr Thr Glu Ile Leu Pro Val Ser Met Thr Lys Thr Ser Val Asp Cys
725 730 735
Thr Met Tyr Ile Cys Gly Asp Ser Thr Glu Cys Ser Asn Leu Leu Leu
740 745 750
Gln Tyr Gly Ser Phe Cys Thr Gln Leu Asn Arg Ala Leu Thr Gly Ile
755 760 765
Ala Val Glu Gln Asp Lys Asn Thr Gln Glu Val Phe Ala Gln Val Lys
770 775 780
Gln Ile Tyr Lys Thr Pro Pro Ile Lys Asp Phe Gly Gly Phe Asn Phe
785 790 795 800
Ser Gln Ile Leu Pro Asp Pro Ser Lys Pro Ser Lys Arg Ser Phe Ile
805 810 815
Glu Asp Leu Leu Phe Asn Lys Val Thr Leu Ala Asp Ala Gly Phe Ile
820 825 830
Lys Gln Tyr Gly Asp Cys Leu Gly Asp Ile Ala Ala Arg Asp Leu Ile
835 840 845
Cys Ala Gln Lys Phe Asn Gly Leu Thr Val Leu Pro Pro Leu Leu Thr
850 855 860
Asp Glu Met Ile Ala Gln Tyr Thr Ser Ala Leu Leu Ala Gly Thr Ile
865 870 875 880
Thr Ser Gly Trp Thr Phe Gly Ala Gly Ala Ala Leu Gln Ile Pro Phe
885 890 895
Ala Met Gln Met Ala Tyr Arg Phe Asn Gly Ile Gly Val Thr Gln Asn
900 905 910
Val Leu Tyr Glu Asn Gln Lys Leu Ile Ala Asn Gln Phe Asn Ser Ala
915 920 925
Ile Gly Lys Ile Gln Asp Ser Leu Ser Ser Thr Ala Ser Ala Leu Gly
930 935 940
Lys Leu Gln Asn Val Val Asn Gln Asn Ala Gln Ala Leu Asn Thr Leu
945 950 955 960
Val Lys Gln Leu Ser Ser Asn Phe Gly Ala Ile Ser Ser Val Leu Asn
965 970 975
Asp Ile Leu Ser Arg Leu Asp Lys Val Glu Ala Glu Val Gln Ile Asp
980 985 990
Arg Leu Ile Thr Gly Arg Leu Gln Ser Leu Gln Thr Tyr Val Thr Gln
995 1000 1005
Gln Leu Ile Arg Ala Ala Glu Ile Arg Ala Ser Ala Asn Leu Ala Ala
1010 1015 1020
Thr Lys Met Ser Glu Cys Val Leu Gly Gln Ser Lys Arg Val Asp Phe
1025 1030 1035 1040
Cys Gly Lys Gly Tyr His Leu Met Ser Phe Pro Gln Ser Ala Pro His
1045 1050 1055
Gly Val Val Phe Leu His Val Thr Tyr Val Pro Ala Gln Glu Lys Asn
1060 1065 1070
Phe Thr Thr Ala Pro Ala Ile Cys His Asp Gly Lys Ala His Phe Pro
1075 1080 1085
Arg Glu Gly Val Phe Val Ser Asn Gly Thr His Trp Phe Val Thr Gln
1090 1095 1100
Arg Asn Phe Tyr Glu Pro Gln Ile Ile Thr Thr Asp Asn Thr Phe Val
1105 1110 1115 1120
Ser Gly Asn Cys Asp Val Val Ile Gly Ile Val Asn Asn Thr Val Tyr
1125 1130 1135
Asp Pro Leu Gln Pro Glu Leu Asp Ser Phe Lys Glu Glu Leu Asp Lys
1140 1145 1150
Tyr Phe Lys Asn His Thr Ser Pro Asp Val Asp Leu Gly Asp Ile Ser
1155 1160 1165
Gly Ile Asn Ala Ser Val Val Asn Ile Gln Lys Glu Ile Asp Arg Leu
1170 1175 1180
Asn Glu Val Ala Lys Asn Leu Asn Glu Ser Leu Ile Asp Leu Gln Glu
1185 1190 1195 1200
Leu Gly Lys Tyr Glu Gln Tyr Ile Lys Trp Pro Trp Tyr Ile Trp Leu
1205 1210 1215
Gly Phe Ile Ala Gly Leu Ile Ala Ile Val Met Val Thr Ile Met Leu
1220 1225 1230
Cys Cys Met Thr Ser Cys Cys Ser Cys Leu Lys Gly Cys Cys Ser Cys
1235 1240 1245
Gly Ser Cys Cys Lys Phe Asp Glu Asp Asp Ser Glu Pro Val Leu Lys
1250 1255 1260
Gly Val Lys Leu His Tyr Thr
1265 1270
Claims (14)
1.一种包含新型冠状病毒双抗原靶标序列组合的重组新城疫病毒载体,所述双抗原靶标序列组合包括新型冠状病毒的S蛋白编码序列和S蛋白受体结合域RBD蛋白编码序列。
2.根据权利要求1所述重组新城疫病毒载体,其特征在于,将所述新型冠状病毒的S蛋白编码序列插入新城疫病毒载体的P基因序列和M基因序列之间;
将所述新型冠状病毒S蛋白受体结合域RBD蛋白编码序列插入新城疫病毒载体的P基因序列和M基因序列之间、M基因序列和F基因序列之间或HN基因序列和L基因序列之间。
3.根据权利要求1或2所述重组新城疫病毒载体,其特征在于,所述S蛋白编码序列和RBD蛋白编码序列来源于相同或不同的新型冠状病毒流行株。
4.根据权利要求3所述重组新城疫病毒载体,其特征在于,所述S蛋白编码序列包括完整的S蛋白编码序列或所述完整的S蛋白编码序列的变体。
5.根据权利要求4所述重组新城疫病毒载体,其特征在于,所述变体包括对所述完整的S蛋白编码序列进行基因工程改造;
所述基因工程改造的方法选自:突变、删除、重排和融合其他来源的蛋白多肽中一种或多种方法的组合;
经所述基因工程改造后的S蛋白构象成为预融合状态更容易被抗原呈递细胞识别,或更多量地表达在细胞膜上,或经表达后分泌到细胞外并组合成易被抗原呈递细胞识别的多聚体。
6.根据权利要求5所述重组新城疫病毒载体,其特征在于,所述突变包括将完整的S蛋白的K986和V987位氨基酸均突变为P,或将第682~685位氨基酸RRAR突变为GSAS,或将完整的S蛋白的K986、V987、F817、 A892、A899和A942位氨基酸均突变为P;或将S蛋白的C末端KLHYT突变为ALAYT;
所述删除包括在所述完整的S蛋白的C末端截短18 ~28个氨基酸;或删除S蛋白的跨膜区和胞内区序列;
所述重排包括将S蛋白的主要抗原决定序列S1蛋白的NTD和RBD区域位置序列进行调换,形成RBD-NTD蛋白;
所述融合其他来源的蛋白多肽包括将S蛋白的胞外域与新城疫F蛋白的C端包含跨膜区和胞内区的氨基酸序列融合;或将RBD-NTD蛋白N端添加信号肽序列,C端添加融合蛋白多肽序列。
7.根据权利要求6所述重组新城疫病毒载体,其特征在于,所述重排所形成的RBD-NTD蛋白,RBD和NTD区域通过蛋白Linker连接。
8.根据权利要求1或2所述重组新城疫病毒载体,其特征在于,所述RBD蛋白编码序列还包括经过基因工程改造的含有RBD蛋白编码序列的RBD系列重组蛋白编码序列。
9.根据权利要求8所述重组新城疫病毒载体,其特征在于,所述基因工程改造包括:在RBD蛋白编码序列的N末端添加分泌信号肽序列,C末端添加融合蛋白多肽序列。
10.一种利用权利要求1~9任一项所述重组新城疫病毒载体制备得到的疫苗株。
11.根据权利要求10所述疫苗株,其特征在于,所述制备的方法包括病毒反向遗传学的方法。
12.根据权利要求10或11所述疫苗株,其特征在于,所述疫苗株的结构包括:权利要求1~9任一项所述重组新城疫病毒载体的编码序列和/或展示在重组新城疫病毒粒子表面的S蛋白或所述S蛋白的变体;
所述S蛋白的变体由完整的S蛋白编码序列的变体编码得到。
13.权利要求10~12任一项所述疫苗株在制备预防新型冠状病毒传播和感染疫苗中的应用。
14.一种预防新型冠状病毒传播和感染的疫苗,其特征在于,所述疫苗包括权利要求10~12任一项所述疫苗株。
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CN115335390A (zh) * | 2022-01-10 | 2022-11-11 | 广州市锐博生物科技有限公司 | 基于SARS-CoV-2的S蛋白的疫苗和组合物 |
WO2023087532A1 (zh) * | 2021-11-17 | 2023-05-25 | 浙江迪福润丝生物科技有限公司 | 一种包含新型冠状病毒双抗原靶标序列组合的重组新城疫病毒载体及相应疫苗株和疫苗 |
CN116925195A (zh) * | 2022-04-22 | 2023-10-24 | 仁景(苏州)生物科技有限公司 | 一种基于新型冠状病毒的mRNA疫苗 |
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CN112679587A (zh) * | 2020-05-26 | 2021-04-20 | 河南中泽生物工程有限公司 | 易于激活b细胞的重组抗原蛋白及其制备方法 |
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