申请/专利权人：美国卫生与公众服务部

申请日：2017-09-01

公开（公告）日：2024-03-08

公开（公告）号：CN110167585B

主分类号：A61K39/12

分类号：A61K39/12;A61K39/145;C07K14/005;C07K14/195

优先权：["20160902 US 62/383,267"]

专利状态码：有效-授权

法律状态：2024.03.08#授权;2019.09.24#实质审查的生效;2019.08.23#公开

摘要：提供了引发广泛保护性抗体的疫苗。所述疫苗包含在其表面上展示来自第2组流感病毒的HA三聚体的纳米颗粒。所述纳米颗粒是融合蛋白，所述融合蛋白包含与第2组流感病毒HA蛋白稳定化的茎区连接的单体亚基例如，铁蛋白。所述融合蛋白自组装以形成展示HA的纳米颗粒。还提供了融合蛋白，编码此类蛋白的核酸分子，以及使用本发明的纳米颗粒检测抗流感抗体的测定法。

主权项：1.一种纳米颗粒，其包含融合蛋白，所述融合蛋白包含与第2组流感病毒血凝素HA蛋白连接的单体亚基蛋白，所述第2组流感病毒血凝素HA蛋白中头部区域被长度少于10个氨基酸的接头序列替代；其中通过向螺旋A添加ALMAQSEQIDNO:36或ELMEQSEQIDNO:37氨基酸残基将所述HA蛋白茎区中螺旋A的长度延长；并且其中HA蛋白的螺旋A和螺旋C之间的螺旋间环inter-helixloop已被包含GGPDSEQIDNO:39的接头替代，从而提高所述融合蛋白的稳定性。

全文数据：稳定化的第2组流感血凝素茎区三聚体及其用途背景技术通过针对流感病毒的疫苗接种诱导的保护性免疫应答主要针对病毒HA蛋白，这是一种在病毒表面上负责病毒与宿主细胞受体相互作用的糖蛋白。病毒表面上的HA蛋白是HA蛋白单体的三聚体，其被酶促裂解后产生氨基末端HA1和羧基末端HA2多肽。球状头部仅由HA1多肽的主要部分组成，而将HA蛋白锚定入病毒脂质包膜的茎部由HA2和部分HA1组成。HA蛋白的球状头部包含两个结构域：受体结合结构域RBD，包含唾液酸结合位点的～148个氨基酸残基的结构域，以及残留的酯酶结构域，其是正好位于RBD下方的较小的～75个氨基酸残基的区域。球状头部包含含有免疫显性表位的若干抗原位点。例如，包括Sa、Sb、Ca1、Ca2和Cb抗原位点参见例如，Caton等，1982,Cell31,417-427。RBD-A区域包含Sa抗原位点和部分Sb抗原位点。针对流感的抗体通常靶向HA球状头部中的可变抗原位点，其围绕着保守的唾液酸结合位点，因此仅中和抗原性密切相关的病毒。HA头部的可变性是由流感病毒持续的抗原漂移造成的，并且使得流感是季节性流行病。而相反的是，HA茎是高度保守的并且几乎不会出现抗原漂移。不幸的是，与免疫显性的头部不同，保守的HA茎不具有较强的免疫原性。此外，病毒基因组的基因区段能够在宿主物种中经历重配抗原漂移，产生能够导致大流行的具有改变的抗原性的新病毒[Salomon,R.等，Cell136,402-4102009]。到目前为止，每年都会更新流感疫苗，以反映即将出现的流行病毒的预测HA和神经氨酸酶NA。最近，分离出一类全新的针对流感病毒的广泛中和抗体，其识别高度保守的HA茎[Corti,D.等，JClinInvest120,1663-16732010；Ekiert,D.C.等，Science324,246-2512009；Kashyap,A.K.等，ProcNatlAcadSciUSA105,5986-59912008；Okuno,Y.等，JVirol67,2552-25581993；Sui,J.等，NatStructMolBiol16,265-2732009；Ekiert,D.C.等，Science333,843-8502011；Corti,D.等，Science333,850-8562011]。与毒株特异性抗体不同的是，这些抗体能够中和多种抗原性不同的病毒，因此诱导此类抗体已成为新一代通用疫苗开发的重点[Nabel,G.J.等，NatMed16,1389-13912010]。然而，通过疫苗接种稳健地引发这些具有此类异质的中和性质的抗体是困难的[Steel,J.等，MBio1,e00182010；Wang,T.T.等，PLoSPathog6,e10007962010；Wei,C.J.等，Science329,1060-10642010]。除去HA免疫显性的头部区域其含有竞争性表位并通过基因操纵将所得到的茎结构域稳定化是改善这些广泛中和茎抗体激发的一种可能方式。目前用于流感的疫苗策略使用化学灭活的或减毒的活流感病毒。这两种疫苗通常都在含胚卵中生产，其主要存在由于耗费时间的工艺和有限的生产能力导致的生产局限性。目前疫苗的另一个更关键的局限性是其功效是高度毒株特异性的。在2009年H1N1流感大流行期间，这些挑战变得更加明显，从而证实了开发能够克服这些局限性的新疫苗平台的必要性。病毒样颗粒代表了这种替代方法之一，并且目前正在临床试验中进行评价[Roldao,A.等，ExpertRevVaccines9,1149-11762010；Sheridan,C.NatBiotechnol27,489-4912009]。作为含胚卵的替代，通常含有HA、NA和基质蛋白1M1的VLP能够在哺乳动物或昆虫细胞表达系统中大量生产[Haynes,J.R.ExpertRevVaccines8,435-4452009]。这种方法的优点是其是颗粒状、多价性的并且是正确折叠的三聚体HA刺突的真实展示，其忠实地模拟了感染性病毒粒子。而相反的是，由于其组装的性质，包膜VLP含有小而有限的宿主细胞成分，在重复使用该平台后可能会带来潜在的安全性、免疫原性挑战[Wu,C.Y.等，PLoSOne5,e97842010]。而且，VLP诱导的免疫力与现有疫苗相同，并且因此不太可能显著改善疫苗诱导的保护性免疫的效力和广度。除了VLP以外，还在人体内对重组HA蛋白进行了评价[Treanor,J.J.等，Vaccine19,1732-17372001；Treanor,J.J.JAMA297,1577-15822007]，但是其诱导保护性中和抗体效价的能力是有限的。在这些试验中使用的重组HA蛋白是在昆虫细胞中生产的，并且可能未优先形成天然的三聚体[Stevens,J.Science303,1866-18702004]。尽管有了几种常规流感疫苗的替代品，但是过去几十年中生物技术的进步使得能够对生物材料进行工程化改造以使其可用于产生新型疫苗平台。铁蛋白是一种几乎存在于所有生物体中的铁储存蛋白，铁蛋白是已被广泛研究和工程化改造以用于多种潜在生物化学生物医学目的的实例[Iwahori,K.美国专利200902333772009；Meldrum,F.C.等，Science257,522-5231992；Naitou,M.等，美国专利201100380252011；Yamashita,I.BiochimBiophysActa1800,846-8572010]，包括用于展示外源性表位肽的潜在疫苗平台[Carter,D.C.等，美国专利200602516792006；Li,C.Q.等，IndustrialBiotechnol2,143-1472006]。将其作为疫苗平台的用途是特别引人注目的，因为其自组装和多价抗原提呈诱导比单价形式更强的B细胞应答以及诱导不依赖于T细胞的抗体应答[Bachmann,M.F.等，AnnuRevImmunol15,235-2701997；Dintzis,H.M.等，ProcNatlAcadSciUSA73,3671-36751976]。此外，铁蛋白由24个亚基构成组装成具有432对称性八面体笼的分子结构，使得具有在其表面上展示多聚抗原的潜能。此前的工作表明，能够对第1组血凝素蛋白的茎区进行修饰以形成稳定化的HA茎蛋白，其构象非常类似于全长、野生型wt流感血凝素蛋白的融合前构象。此外，当将此类经修饰的稳定化的茎SSHA蛋白与单体亚基蛋白如铁蛋白连接在一起时，使所得到的融合蛋白形成纳米颗粒，其表面展示SS-HA蛋白的三聚体。而且，此类纳米颗粒能够激发针对第1组流感病毒的免疫应答，表明由纳米颗粒展示的SS-HA蛋白三聚体具有与wt流感HA蛋白类似的构象。在国际专利申请号PCTUS2015032695中公开了此类构建体，其全部内容通过引用并入本申请。然而，由上述纳米颗粒激发的抗体针对第1组流感病毒比其针对第2组流感病毒具有更高的保护性。因而，仍需要一种有效的流感疫苗，其对第2组流感病毒提供强有力的保护作用。而且，还需要一种流感疫苗，其保护个体免受流感病毒异质性毒株的侵害，包括未来进化的季节性和大流行的流感病毒毒株。本发明通过提供一种新型的基于纳米颗粒的疫苗满足了这一需求，所述疫苗由与纳米颗粒表面融合的缺乏可变的免疫显性头部区域的新型第2组HA稳定化的茎SS构成，这使得所述流感疫苗易于生产、高效并且能够激发对广泛的异质亚型具有保护作用的抗体。发明概述因此，本公开内容提供了包含第2组流感血凝素HA蛋白的重组蛋白，其中头部区域的氨基酸序列被接头替代，所述接头含有来自流感HA蛋白头部区域的少于5个连续的氨基酸。在向哺乳动物施用这些重组蛋白后，这些重组蛋白在哺乳动物中激发针对第2组流感HA蛋白的免疫应答。重组蛋白可以包含来自第2组流感病毒血凝素HA蛋白茎区的第一氨基酸序列和来自第2组流感病毒血凝素HA蛋白茎区的第二氨基酸序列，其中第一和第二氨基酸序列通过接头序列共价连接在一起，以及其中第一氨基酸序列包含来自头部区域序列氨基末端上游的氨基酸序列的至少20个连续的氨基酸残基，和其中第二氨基酸序列包含来自头部区域序列羧基末端下游的氨基酸序列的至少20个连续的氨基酸残基。在该重组蛋白构建体中，第一氨基酸序列可以包含来自与头部区域的氨基末端紧邻的上游多肽序列的至少20个连续的氨基酸。或者或另外地，第一氨基酸序列可以包含来自SEQIDNO:27、SEQIDNO:28或SEQIDNO:29的至少20个连续的氨基酸。或者或另外地，第二氨基酸序列可以包含来自与头部区域的羧基末端紧邻的下游多肽序列的至少20个连续的氨基酸。或者或另外地，第一氨基酸序列可以包含来自SEQIDNO:31、SEQIDNO:32或SEQIDNO:33的至少20个连续的氨基酸。重组多肽可以包含C螺旋的氨基末端即，C螺旋的膜远端，其与经修饰以含有第一半胱氨酸氨基酸的头部区域序列连接，以及含有第二半胱氨酸氨基酸的接头序列，以使得第一和第二半胱氨酸形成二硫键。重组蛋白可以含有螺旋间区域即，将C螺旋的N-末端与A螺旋的羧基末端即，A螺旋的膜远端连接在一起的氨基酸序列，对该区域进行修饰以使得重组HA茎蛋白的三维结构近似于天然第2组HA蛋白中HA茎区的三维结构。重组蛋白可以含有长度小于8个氨基酸并且替代螺旋间区域的氨基酸接头序列。重组蛋白可以含有通过添加氨基酸延伸的A螺旋的膜远端。重组蛋白可以含有第三氨基酸接头，其与形成A螺旋的氨基酸序列的羧基末端连接在一起并且形成延长A螺旋长度的螺旋。可以通过接头肽将C螺旋的远端与第三接头的羧基末端连接在一起。接头肽的长度优选小于8个氨基酸。这些重组蛋白可以含有增加蛋白稳定性的一个或多个突变。这些稳定突变优选位于形成A螺旋和C螺旋的至少一个的氨基酸序列中。这些重组蛋白可以与来自铁蛋白或二氧四氢蝶啶合酶的单体亚基连接在一起。本公开内容的示例性重组蛋白可以包含与选自下组的氨基酸序列具有至少80％同一性、或至少85％同一性、或至少90％同一性、或至少95％同一性、或至少97％同一性或至少99％同一性的氨基酸序列：SEQIDNO：47-159。本公开内容的示例性重组蛋白可以包含选自下组的氨基酸序列：SEQIDNO：47-159。本公开内容还提供了一种纳米颗粒，所述纳米颗粒包含至少一种本公开内容的重组蛋白。本公开内容还提供了免疫原性组合物，所述免疫原性组合物包含至少一种蛋白，所述蛋白包含与这些重组蛋白具有至少95％同一性的氨基酸序列。这些免疫原性组合物可以包含蛋白，所述蛋白含有选自下组的氨基酸序列：SEQIDNO：47-1598。这些免疫原性组合物可以包含由选自下组的氨基酸序列组成的蛋白：SEQIDNO：47-159。因此，本公开内容还提供了疫苗组合物，所述疫苗组合物包含这些免疫原性组合物和佐剂。本公开内容还提供了一种在人体内预防或减轻流感病毒感染的病理学效应的方法，所述方法包括向需要其的人体施用免疫学有效量的本公开内容的的疫苗组合物。还提供了编码本公开内容的重组蛋白的核酸。优选地，核酸是DNA。还提供了载体，所述载体包含这些核酸。还提供了宿主细胞，所述宿主细胞包含这些载体。这些宿主细胞可以是细菌细胞、酵母细胞或哺乳动物细胞。这些宿主细胞可以是灭活的。本公开内容还提供了药物组合物，所述药物组合物包含本公开内容的重组蛋白。类似地，这些组合物可以是疫苗，所述疫苗包含本公开内容的重组蛋白，以及联合生理学上可接受的运载体。本公开内容还提供了接种疫苗的方法，所述方法包括向对象施用预防或治疗有效量的本公开内容的重组蛋白。本公开内容还提供了一种治疗流感相关疾病的方法，所述方法包括向需要其的对象施用预防或治疗有效量的本公开内容的重组蛋白。优选地，对象是人。附图说明图1A-1C提供了对现有技术的总结。图1A是描述全长HA-铁蛋白纳米颗粒设计的带状图。图1B是描述通过HIVgp41三聚化结构域稳定化的HA茎-铁蛋白纳米颗粒设计的带状图。在专利申请PCTUS1256822中对这两个设计进行了描述，其通过引用并入本申请。图1C是描述在PCT专利申请号PCTUS1532695中公开的第1组HA稳定化的茎纳米颗粒设计的带状图，其通过引用并入本申请。图2描述了第2组HA茎纳米颗粒的自组装形成。带状图从左到右描述了第2组HA稳定化的茎纳米颗粒的设计。一个HA单体的头部区域用深灰色表示。同一单体的茎区用中灰色表示。另外两个单体用浅灰色表示。图3A和3B显示了在H3N2设计231中的突变，其能够形成第2组HA稳定化的茎纳米颗粒。图3A是描述基于PDBID2YP2的第2组H3N2稳定化的HA茎三聚体模型的带状图。在螺旋中的突变区域用深灰色表示。图3B显示了H3设计#231的序列基于A丹麦352005H3N2的HA茎，GenBankABU92694。序列中的突变如框中所示。作为参考，C-末端SGG接头用粗体表示，C-末端铁蛋白用下划线表示，以及Asn至Gln的铁蛋白突变以除去N-连接聚糖用粗体表示。图4A-4D显示了H3N2设计231在替代HA1头的环中的突变。图4A显示了描述基于PDBID2YP2的第2组H3N2稳定化的HA茎三聚体模型的带状图。显示了在环中替代HA1头部区域的7个突变，以及在C螺旋中加入的与上述环形成二硫键的半胱氨酸。在螺旋区域中的所有其他突变用深灰色表示。图4B描述了具有由棍模型表示的侧链的突变环表示为替代图4A中的头部区域。图4C显示了环序列的变体。图4D显示了H3设计#231的序列。图4A-4C中所示的头部和螺旋区域中的突变如框中所示。作为参考，C-末端SGG接头用粗体表示，C-末端铁蛋白用下划线表示，以及Asn至Gln的铁蛋白突变以除去N-连接聚糖用粗体表示。图5A-5C显示了H3N2设计231中在连接HA2A螺旋和C螺旋的环中的突变。图5A是描述基于PDBID2YP2的第2组H3N2稳定化的HA茎三聚体模型的带状图。显示了连接HA2A螺旋和C螺旋的四个残基。在螺旋中的突变用深灰色表示。图5B显示了具有由棍模型表示的侧链的环的特写图5A中所示的区域。图5C显示了H3设计#231的序列。图5A和5B中所示的螺旋中的突变和组成短接头的氨基酸如框中所示。作为参考，C-末端SGG接头用粗体表示，C-末端铁蛋白用下划线表示，以及Asn至Gln的铁蛋白突变以除去N-连接聚糖用粗体表示。图6A-6C显示了H3N2设计231中在A螺旋C-末端延伸中的突变。图6A显示了描述基于PDBID2YP2的第2组H3N2稳定化的HA茎三聚体模型的带状图。显示了A螺旋的5个残基延伸。在螺旋中的突变用深灰色表示。图6B显示了具有由棍模型表示的侧链的螺旋延伸的特写亦如图6A中所示。图6C显示了H3设计#231的序列。在螺旋中的突变以及组成5个残基延伸的氨基酸如框中所示。作为参考，C-末端SGG接头用粗体表示，C-末端铁蛋白用下划线表示，以及Asn至Gln的铁蛋白突变以除去N-连接聚糖用粗体表示。图7A和7B显示了在H3N2设计231中填充孔穴cavity的突变。图7A显示了描述基于PDBID2YP2的第2组H3N2稳定化HA茎三聚体模型的条带图。具有由棍模型表示的侧链的7个填充孔穴的突变用深灰色表示。图7B显示了H3设计#231的序列。螺旋和头部区域的突变如框中所示。作为参考，C-末端SGG接头用粗体表示，C-末端铁蛋白用下划线表示，以及Asn至Gln的铁蛋白突变以除去N-连接聚糖用粗体表示。图8A和8B显示了H3稳定化铁蛋白纳米颗粒231H3-SS-np_231的表达和表征。图8A显示了H3-SS-np_231的凝胶过滤洗脱曲线，其在预期洗脱体积具有单峰。凝胶过滤后Expi293细胞中H3-SS-np_231的表达收率为7.7mgL。图8B显示了H3-SS-np_231的负染电子显微镜2D类平均值，其揭示了具有从中空球体中突出的HA茎可见排布的颗粒的形成。图9A和9B显示了H3-SS-np_231的HA茎抗体识别。图9A列出了三种HA茎抗体动力学ELISAH3-SS-np_231识别测定的EC50值。将H1-SS-np识别的值也显示为对照。在这两种情况下，将纳米颗粒固定在板上。图9B显示了CT149识别H3-SS-np_231的生物层干涉测量法BLI，来自Octet结合曲线上图以及HA茎抗体CT149和CR9114的BL1动力学常数下图。图10A-10E显示了H3-SS-np_231的5种变体的凝胶过滤曲线。H3-SS-np_231变体249图10A、256图10B、258图10C、262图10D和264图10E的凝胶过滤Superose61030曲线。在每种情况下，在约14.5ml的体积下洗脱出单峰。凝胶过滤后从Expi293细胞的最终收率为6-8mgL培养物。图11A-11F显示了H3-SS-np纳米颗粒变体的电子显微镜图。H3-SS-np变体的负染电子显微镜2D类平均值揭示了具有从中空球体中突出的HA茎可见排布的颗粒的形成。将H3-SS-np231颗粒的图像左上图作为阳性对照。图12A-12D显示了H3-SS-np_231的5种变体的动力学ELISA结果。图12A-12C显示了FI6图12A、CT149图12B和CR8020图12C识别H3-SS-np_231变体249、256、258、262和264的动力学ELISA曲线。图12D列出了图12A-12C中所示曲线的EC50值。图13A和13B显示了H3-SS-np变体235-295的动力学ELISA结果。图13A列出了显示广泛中和HA茎抗体FI6、CT149和D25阴性对照识别设计235-265的ELISA效价。图13B列出了显示D25和CT149识别设计266-296的ELISA效价。将来自表达设计免疫原的HEK293T细胞的上清液置于板中并使用上述抗体进行检测。图14显示了H3-SS-np#231和5种变体的动态扫描量热法DSC曲线。热容Cp对温度曲线描述了每种蛋白的熔融转变。记录每种设计的最早熔点TM。在每个括号中显示了设计编号。在该图中，Y轴上的Cp值以任意比例显示。图15A&15B显示了H3-SS-np免疫小鼠对第2组HA的免疫应答。向含A香港11968H3N2HA图15A和A安徽12013H7N9图15B的板中加入来自使用6种不同形式的SAS-佐剂化H3-SS-np免疫3x的BALBc小鼠n＝10的血清的ELISA抗体终点效价。将仅使用空铁蛋白纳米颗粒免疫小鼠的血清作为阴性对照。用水平线表示几何平均效价。深灰色阴影表示阴性对照的平均效价，浅灰色阴影表示达到阴性对照平均滴度4倍的区域。使用双侧Studentt-检验进行统计学分析；*P美国卫生与公众服务部Boyington,JeffreyC.Graham,BarneyS.Mascola,JohnR.Yassine,HadiM.Corbett,KizzmekiaS.Moin,SyedM.Wang,LingshuKanekiyo,Masaru稳定化的第2组流感血凝素茎区三聚体及其用途6137NIAID-64-PCT尚未分配2017-09-0162383,2672016-09-02159PatentIn3.5版1168PRT幽门螺杆菌1MetLeuSerAspIleIleLysLeuLeuAsnGluGlnValAsnLysGlu151015MetGlnSerSerAsnLeuTyrMetSerMetSerSerTrpCysTyrThr202530HisSerLeuAspGlyAlaGlyLeuPheLeuPheAspHisAlaAlaGlu354045GluTyrGluHisAlaLysLysLeuIleIlePheLeuAsnGluAsnAsn505560ValProValGlnLeuThrSerIleSerAlaProGluHisLysPheGlu65707580GlyLeuThrGlnIlePheGlnLysAlaTyrGluHisGluGlnHisIle859095SerGluSerIleAsnAsnIleValAspHisAlaIleLysSerLysAsp100105110HisAlaThrPheAsnPheLeuGlnTrpTyrValAlaGluGlnHisGlu115120125GluGluValLeuPheLysAspIleLeuAspLysIleGluLeuIleGly130135140AsnGluAsnHisGlyLeuTyrLeuAlaAspGlnTyrValLysGlyIle145150155160AlaLysSerArgLysSerGlySer1652165PRT人工序列合成的2AspIleIleLysLeuLeuAsnGluGlnValAsnLysGluMetGlnSer151015SerAsnLeuTyrMetSerMetSerSerTrpCysTyrThrHisSerLeu202530AspGlyAlaGlyLeuPheLeuPheAspHisAlaAlaGluGluTyrGlu354045HisAlaLysLysLeuIleIlePheLeuAsnGluAsnAsnValProVal505560GlnLeuThrSerIleSerAlaProGluHisLysPheGluGlyLeuThr65707580GlnIlePheGlnLysAlaTyrGluHisGluGlnHisIleSerGluSer859095IleAsnAsnIleValAspHisAlaIleLysSerLysAspHisAlaThr100105110PheAsnPheLeuGlnTrpTyrValAlaGluGlnHisGluGluGluVal115120125LeuPheLysAspIleLeuAspLysIleGluLeuIleGlyAsnGluAsn130135140HisGlyLeuTyrLeuAlaAspGlnTyrValLysGlyIleAlaLysSer145150155160ArgLysSerGlySer1653154PRT超嗜热菌3MetGlnIleTyrGluGlyLysLeuThrAlaGluGlyLeuArgPheGly151015IleValAlaSerArgPheAsnHisAlaLeuValAspArgLeuValGlu202530GlyAlaIleAspCysIleValArgHisGlyGlyArgGluGluAspIle354045ThrLeuValArgValProGlySerTrpGluIleProValAlaAlaGly505560GluLeuAlaArgLysGluAspIleAspAlaValIleAlaIleGlyVal65707580LeuIleArgGlyAlaThrProHisPheAspTyrIleAlaSerGluVal859095SerLysGlyLeuAlaAspLeuSerLeuGluLeuArgLysProIleThr100105110PheGlyValIleThrAlaAspThrLeuGluGlnAlaIleGluArgAla115120125GlyThrLysHisGlyAsnLysGlyTrpGluAlaAlaLeuSerAlaIle130135140GluMetAlaAsnLeuPheLysSerLeuArg1451504566PRT甲型流感病毒4MetLysThrIleIleAlaLeuSerTyrIleLeuCysLeuValPheAla151015GlnLysLeuProGlyAsnAspAsnSerThrAlaThrLeuCysLeuGly202530HisHisAlaValProAsnGlyThrIleValLysThrIleThrAsnAsp354045GlnIleGluValThrAsnAlaThrGluLeuValGlnSerSerSerThr505560GlyGlyIleCysAspSerProHisGlnIleLeuAspGlyGluAsnCys65707580ThrLeuIleAspAlaLeuLeuGly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LysCysAspAsnAlaCysIleGlySerIleArgAsnGly195200205ThrTyrAspHisAspValTyrArgAspGluAlaLeuAsnAsnArgPhe210215220GlnIleLysSerGlyGlyAspIleIleLysLeuLeuAsnGluGlnVal225230235240AsnLysGluMetGlnSerSerAsnLeuTyrMetSerMetSerSerTrp245250255CysTyrThrHisSerLeuAspGlyAlaGlyLeuPheLeuPheAspHis260265270AlaAlaGluGluTyrGluHisAlaLysLysLeuIleIlePheLeuAsn275280285GluAsnAsnValProValGlnLeuThrSerIleSerAlaProGluHis290295300LysPheGluGlyLeuThrGlnIlePheGlnLysAlaTyrGluHisGlu305310315320GlnHisIleSerGluSerIleAsnAsnIleValAspHisAlaIleLys325330335SerLysAspHisAlaThrPheAsnPheLeuGlnTrpTyrValAlaGlu340345350GlnHisGluGluGluValLeuPheLysAspIleLeuAspLysIleGlu355360365LeuIleGlyAsnGluAsnHisGlyLeuTyrLeuAlaAspGlnTyrVal370375380LysGlyIleAlaLysSerArgLysSerGlySer38539039581395PRT人工序列合成的81MetLysThrIleIleAlaLeuSerTyrIleLeuCysLeuValPheAla151015GlnLysLeuProGlyAsnAspAsnSerThrAlaThrLeuCysLeuGly202530HisHisAlaValProAsnGlyThrIleValLysThrIleThrAsnAsp354045GlnIleGluValThrAsnAlaThrGluLeuValPheProGlyCysVal505560LeuLysLeuAlaThrGlyMetArgAsnValProGluLysGlnThrArg65707580GlyIlePheGlyAlaIleAlaGlyPheIleGluAsnGlyTrpGluGly859095MetValAspGlyTrpTyrGlyPheArgHisGlnAsnSerGluGlyThr100105110GlyGlnAlaAlaAspLeuLysSerThrGlnAlaAlaIleAsnGlnIle115120125AsnGlyMetValAsnArgValIleAlaLeuMetAlaGlnGlyGlyPro130135140AspCysTyrLeuAlaGluLeuLeuValAlaLeuLeuAsnGlnHisVal145150155160IleAspLeuThrAspSerGluMetArgLysLeuPheGluArgThrLys165170175LysGlnLeuArgGluAsnAlaGluAspMetGlyAsnGlyCysPheLys180185190IleTyrHisLysCysAspAsnAlaCysIleGluSerIleArgAsnGly195200205ThrTyrAspHisAspValTyrArgAspGluAlaLeuAsnAsnArgPhe210215220GlnIleLysSerGlyGlyAspIleIleLysLeuLeuAsnGluGlnVal225230235240AsnLysGluMetGlnSerSerAsnLeuTyrMetSerMetSerSerTrp245250255CysTyrThrHisSerLeuAspGlyAlaGlyLeuPheLeuPheAspHis260265270AlaAlaGluGluTyrGluHisAlaLysLysLeuIleIlePheLeuAsn275280285GluAsnAsnValProValGlnLeuThrSerIleSerAlaProGluHis290295300LysPheGluGlyLeuThrGlnIlePheGlnLysAlaTyrGluHisGlu305310315320GlnHisIleSerGluSerIleAsnAsnIleValAspHisAlaIleLys325330335SerLysAspHisAlaThrPheAsnPheLeuGlnTrpTyrValAlaGlu340345350GlnHisGluGluGluValLeuPheLysAspIleLeuAspLysIleGlu355360365LeuIleGlyAsnGluAsnHisGlyLeuTyrLeuAlaAspGlnTyrVal370375380LysGlyIleAlaLysSerArgLysSerGlySer38539039582395PRT人工序列合成的82MetLysThrIleIleAlaLeuSerTyrIleLeuCysLeuValPheAla151015GlnLysLeuProGlyAsnAspAsnSerThrAlaThrLeuCysLeuGly202530HisHisAlaValProAsnGlyThrIleValLysThrIleThrAsnAsp354045ArgIleGluValThrAsnAlaThrGluLeuValPheProGlyCysVal505560LeuLysLeuAlaThrGlyMetArgAsnValProGluArgGlnThrArg65707580GlyIlePheGlyAlaIleAlaGlyPheIleGluAsnGlyTrpGluGly859095MetValAspGlyTrpTyrGlyPheArgHisGlnAsnSerGluGlyArg100105110GlyGlnAlaAlaAspLeuLysSerThrGlnAlaAlaIleAspGlnIle115120125AsnGlyMetValAsnArgValIleAlaLeuMetAlaGlnGlyGlyPro130135140AspCysTyrLeuAlaGluLeuLeuValAlaLeuLeuAsnGlnHisVal145150155160IleAspLeuThrAspSerGluMetArgLysLeuPheGluLysThrLys165170175LysGlnLeuArgGluAsnAlaGluAspMetGlyAsnGlyCysPheLys180185190IleTyrHisLysCysAspAsnAlaCysIleGlySerIleArgAsnGly195200205ThrTyrAspHisAspValTyrArgAspGluAlaLeuAsnAsnArgPhe210215220GlnIleLysSerGlyGlyAspIleIleLysLeuLeuAsnGluGlnVal225230235240AsnLysGluMetGlnSerSerAsnLeuTyrMetSerMetSerSerTrp245250255CysTyrThrHisSerLeuAspGlyAlaGlyLeuPheLeuPheAspHis260265270AlaAlaGluGluTyrGluHisAlaLysLysLeuIleIlePheLeuAsn275280285GluAsnAsnValProValGlnLeuThrSerIleSerAlaProGluHis290295300LysPheGluGlyLeuThrGlnIlePheGlnLysAlaTyrGluHisGlu305310315320GlnHisIleSerGluSerIleAsnAsnIleValAspHisAlaIleLys325330335SerLysAspHisAlaThrPheAsnPheLeuGlnTrpTyrValAlaGlu340345350GlnHisGluGluGluValLeuPheLysAspIleLeuAspLysIleGlu355360365LeuIleGlyAsnGluAsnHisGlyLeuTyrLeuAlaAspGlnTyrVal370375380LysGlyIleAlaLysSerArgLysSerGlySer38539039583388PRT人工序列合成的83MetLysThrIleIleAlaLeuSerTyrIleLeuCysLeuValPheAla151015GlnLysLeuProGlyAsnAspAsnSerThrAlaThrLeuCysLeuGly202530HisHisAlaValProAsnGlyThrIleValLysThrIleThrAsnAsp354045GlnIleGluValThrAsnAlaThrGluLeuValPheProGlyCysGly505560ValLeuLysLeuAlaThrGlyMetArgAsnValProGluLysGlnThr65707580ArgGlyIlePheGlyAlaIleAlaGlyPheIleGluAsnGlyTrpGlu859095GlyMetValAspGlyTrpTyrGlyPheArgHisGlnAsnSerGluGly100105110IleGlyGlnAlaAlaAspLeuLysSerThrGlnAlaAlaIleAsnGln115120125IleAsnGlyMetValAsnArgValIleAlaLeuMetAlaGlnGlyGly130135140ProAspCysTyrLeuAlaGluLeuLeuValAlaLeuLeuAsnGlnHis145150155160ValIleAspLeuThrAspSerGluMetArgLysLeuPheGluArgThr165170175LysLysGlnLeuArgGluAsnAlaGluAspMetGlyAsnGlyCysPhe180185190LysIleTyrHisLysCysAspAsnAlaCysIleGlySerIleArgAsn195200205GlyThrTyrAspHisAspValTyrArgAspGluAlaLeuAsnAsnArg210215220PheGlnIleLysAlaGlyProProGlyGlyMetGlnIleTyrGluGly225230235240LysLeuThrAlaGluGlyLeuArgPheGlyIleValAlaSerArgPhe245250255AsnHisAlaLeuValAspArgLeuValGluGlyAlaIleAspCysIle260265270ValArgHisGlyGlyArgGluGluAspIleThrLeuValArgValPro275280285GlySerTrpGluIleProValAlaAlaGlyGluLeuAlaArgLysGlu290295300AspIleAspAlaValIleAlaIleGlyValLeuIleArgGlyAlaThr305310315320ProHisPheAspTyrIleAlaSerGluValSerLysGlyLeuAlaAsp325330335LeuSerLeuGluLeuArgLysProIleThrPheGlyValIleThrAla340345350AspThrLeuGluGlnAlaIleGluArgAlaGlyThrLysHisGlyAsn355360365LysGlyTrpGluAlaAlaLeuSerAlaIleGluMetAlaAsnLeuPhe370375380LysSerLeuArg38584388PRT人工序列合成的84MetLysThrIleIleAlaLeuSerTyr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260265270HisAlaAlaGluGluTyrGluHisAlaLysLysLeuIleIlePheLeu275280285AsnGluAsnAsnValProValGlnLeuThrSerIleSerAlaProGlu290295300HisLysPheGluGlyLeuThrGlnIlePheGlnLysAlaTyrGluHis305310315320GluGlnHisIleSerGluSerIleAsnAsnIleValAspHisAlaIle325330335LysSerLysAspHisAlaThrPheAsnPheLeuGlnTrpTyrValAla340345350GluGlnHisGluGluGluValLeuPheLysAspIleLeuAspLysIle355360365GluLeuIleGlyAsnGluAsnHisGlyLeuTyrLeuAlaAspGlnTyr370375380ValLysGlyIleAlaLysSerArgLysSerGlySer385390395139396PRT人工序列合成的139MetLysThrIleIleAlaLeuSerTyrIleLeuCysLeuValPheAla151015GlnLysLeuProGlyAsnAspAsnSerThrAlaThrLeuCysLeuGly202530HisHisAlaValProAsnGlyThrIleValLysThrIleThrAsnAsp354045GlnIleGluValThrAsnAlaThrGluLeuAsnPheThrGlyCysGly505560ValLeuLysLeuAlaThrGlyMetArgAsnValProGluLysGlnThr65707580ArgGlyIlePheGlyAlaIleAlaGlyPheIleGluAsnGlyTrpGlu859095GlyMetValAspGlyTrpTyrGlyPheArgHisGlnAsnSerGluGly100105110IleGlyGlnAlaAlaAspLeuLysSerThrGlnAlaAlaIleAsnGln115120125IleAsnGlyMetValAsnArgValIleAlaLeuMetAlaGlnGlyGly130135140ProAspCysTyrLeuAlaGluLeuLeuValAlaLeuLeuAsnGlnHis145150155160ValIleAspLeuThrAspSerGluMetArgLysLeuPheGluArgThr165170175LysLysGlnLeuArgGluAsnAlaGluAspMetGlyAsnGlyCysPhe180185190LysIleTyrHisLysCysAspAsnAlaCysIleGlySerIleArgAsn195200205GlyThrTyrAspHisAspValTyrArgAspGluAlaLeuAsnAsnArg210215220PheGlnIleLysSerGlyGlyAspIleIleLysLeuLeuAsnGluGln225230235240ValAsnLysGluMetGlnSerSerAsnLeuTyrMetSerMetSerSer245250255TrpCysTyrThrHisSerLeuAspGlyAlaGlyLeuPheLeuPheAsp260265270HisAlaAlaGluGluTyrGluHisAlaLysLysLeuIleIlePheLeu275280285AsnGluAsnAsnValProValGlnLeuThrSerIleSerAlaProGlu290295300HisLysPheGluGlyLeuThrGlnIlePheGlnLysAlaTyrGluHis305310315320GluGlnHisIleSerGluSerIleAsnAsnIleValAspHisAlaIle325330335LysSerLysAspHisAlaThrPheAsnPheLeuGlnTrpTyrValAla340345350GluGlnHisGluGluGluValLeuPheLysAspIleLeuAspLysIle355360365GluLeuIleGlyAsnGluAsnHisGlyLeuTyrLeuAlaAspGlnTyr370375380ValLysGlyIleAlaLysSerArgLysSerGlySer385390395140396PRT人工序列合成的140MetLysThrIleIleAlaLeuSerTyrIleLeuCysLeuValPheAla151015GlnLysLeuProGlyAsnAspAsnSerThrAlaThrLeuCysLeuGly202530HisHisAlaValProAsnGlyThrIleValLysThrIleThrAsnAsp354045GlnIleGluValThrAsnAlaThrGluLeuValPheProAsnCysThr505560ValLeuLysLeuAlaThrGlyMetArgAsnValProGluLysGlnThr65707580ArgGlyIlePheGlyAlaIleAlaGlyPheIleGluAsnGlyTrpGlu859095GlyMetValAspGlyTrpTyrGlyPheArgHisGlnAsnSerGluGly100105110IleGlyGlnAlaAlaAspLeuLysSerThrGlnAlaAlaIleAsnGln115120125IleAsnGlyMetValAsnArgValIleAlaLeuMetAlaGlnGlyGly130135140ProAspCysTyrLeuAlaGluLeuLeuValAlaLeuLeuAsnGlnHis145150155160ValIleAspLeuThrAspSerGluMetArgLysLeuPheGluArgThr165170175LysLysGlnLeuArgGluAsnAlaGluAspMetGlyAsnGlyCysPhe180185190LysIleTyrHisLysCysAspAsnAlaCysIleGlySerIleArgAsn195200205GlyThrTyrAspHisAspValTyrArgAspGluAlaLeuAsnAsnArg210215220PheGlnIleLysSerGlyGlyAspIleIleLysLeuLeuAsnGluGln225230235240ValAsnLysGluMetGlnSerSerAsnLeuTyrMetSerMetSerSer245250255TrpCysTyrThrHisSerLeuAspGlyAlaGlyLeuPheLeuPheAsp260265270HisAlaAlaGluGluTyrGluHisAlaLysLysLeuIleIlePheLeu275280285AsnGluAsnAsnValProValGlnLeuThrSerIleSerAlaProGlu290295300HisLysPheGluGlyLeuThrGlnIlePheGlnLysAlaTyrGluHis305310315320GluGlnHisIleSerGluSerIleAsnAsnIleValAspHisAlaIle325330335LysSerLysAspHisAlaThrPheAsnPheLeuGlnTrpTyrValAla340345350GluGlnHisGluGluGluValLeuPheLysAspIleLeuAspLysIle355360365GluLeuIleGlyAsnGluAsnHisGlyLeuTyrLeuAlaAspGlnTyr370375380ValLysGlyIleAlaLysSerArgLysSerGlySer385390395141396PRT人工序列合成的141MetLysThrIleIleAlaLeuSerTyrIleLeuCysLeuValPheAla151015GlnLysLeuProGlyAsnAspAsnSerThrAlaThrLeuCysLeuGly202530HisHisAlaValProAsnGlyThrIleValLysThrIleThrAsnAsp354045GlnIleGluValThrAsnAlaThrGluLeuValPheProGlyCysGly505560AsnLeuThrLeuAlaThrGlyMetArgAsnValProGluLysGlnThr65707580ArgGlyIlePheGlyAlaIleAlaGlyPheIleGluAsnGlyTrpGlu859095GlyMetValAspGlyTrpTyrGlyPheArgHisGlnAsnSerGluGly100105110IleGlyGlnAlaAlaAspLeuLysSerThrGlnAlaAlaIleAsnGln115120125IleAsnGlyMetValAsnArgValIleAlaLeuMetAlaGlnGlyGly130135140ProAspCysTyrLeuAlaGluLeuLeuValAlaLeuLeuAsnGlnHis145150155160ValIleAspLeuThrAspSerGluMetArgLysLeuPheGluArgThr165170175LysLysGlnLeuArgGluAsnAlaGluAspMetGlyAsnGlyCysPhe180185190LysIleTyrHisLysCysAspAsnAlaCysIleGlySerIleArgAsn195200205GlyThrTyrAspHisAspValTyrArgAspGluAlaLeuAsnAsnArg210215220PheGlnIleLysSerGlyGlyAspIleIleLysLeuLeuAsnGluGln225230235240ValAsnLysGluMetGlnSerSerAsnLeuTyrMetSerMetSerSer245250255TrpCysTyrThrHisSerLeuAspGlyAlaGlyLeuPheLeuPheAsp260265270HisAlaAlaGluGluTyrGluHisAlaLysLysLeuIleIlePheLeu275280285AsnGluAsnAsnValProValGlnLeuThrSerIleSerAlaProGlu290295300HisLysPheGluGlyLeuThrGlnIlePheGlnLysAlaTyrGluHis305310315320GluGlnHisIleSerGluSerIleAsnAsnIleValAspHisAlaIle325330335LysSerLysAspHisAlaThrPheAsnPheLeuGlnTrpTyrValAla340345350GluGlnHisGluGluGluValLeuPheLysAspIleLeuAspLysIle355360365GluLeuIleGlyAsnGluAsnHisGlyLeuTyrLeuAlaAspGlnTyr370375380ValLysGlyIleAlaLysSerArgLysSerGlySer385390395142396PRT人工序列合成的142MetLysThrIleIleAlaLeuSerTyrIleLeuCysLeuValPheAla151015GlnLysLeuProGlyAsnAspAsnSerThrAlaThrLeuCysLeuGly202530HisHisAlaValProAsnGlyThrIleValLysThrIleThrAsnAsp354045GlnIleGluValThrAsnAlaThrGluLeuValPheProGlyCysGly505560ValLeuAsnLeuThrThrGlyMetArgAsnValP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权利要求：1.一种重组蛋白，所述重组蛋白包含第2组流感血凝素HA蛋白，其中头部区域的氨基酸序列已被包含来自流感HA蛋白头部区域至少5个连续氨基酸的接头替代，其中向哺乳动物施用所述蛋白在所述哺乳动物中引发针对第2组流感HA蛋白的免疫应答。2.根据权利要求1所述的重组蛋白，其中所述重组蛋白包含来自第2组流感病毒血凝素HA蛋白茎区的第一氨基酸序列，和来自第2组流感病毒血凝素HA蛋白茎区的第二氨基酸序列，其中所述第一氨基酸序列和第二氨基酸序列通过所述接头序列共价连接，其中所述第一氨基酸序列包含来自所述头部区域序列氨基末端上游的氨基酸序列的至少20个连续的氨基酸残基，和其中所述第二氨基酸序列包含来自所述头部区域序列羧基末端下游的氨基酸序列的至少20个连续的氨基酸残基。3.根据权利要求1或2所述的重组蛋白，其中所述重组蛋白与单体亚基连接。4.根据权利要求3所述的重组蛋白，其中所述单体亚基蛋白来自铁蛋白或2,4二氧四氢蝶啶合酶。5.一种纳米颗粒，所述纳米颗粒包含权利要求1-4中任意一项所述的重组蛋白，其中所述纳米颗粒在其表面上展示出流感HA蛋白的三聚体。6.一种免疫原性组合物，所述免疫原性组合物包含蛋白，所述蛋白含有与权利要求1-4中任意一项所述的重组蛋白具有至少95％同一性的氨基酸序列。7.一种疫苗组合物，所述疫苗组合物包含权利要求1-4中任意一项所述的重组蛋白或权利要求5所述的纳米颗粒，以及佐剂。8.一种在人体内预防或减轻流感病毒感染的病理学效应的方法，所述方法包括向需要其的人体施用免疫学有效量的权利要求7所述的疫苗组合物。9.一种编码权利要求1-4中任意一项所述的重组蛋白的核酸。10.根据权利要求9所述的核酸，其中所述核酸是DNA。11.一种载体，所述载体包含权利要求10所述的核酸。12.一种宿主细胞，所述宿主细胞包含权利要求11所述的载体。13.根据权利要求12所述的宿主细胞，其是细菌细胞、酵母细胞或哺乳动物细胞。14.一种药物组合物，所述药物组合物包含权利要求1-4所述的重组蛋白。15.一种接种疫苗的方法，所述方法包括向对象施用预防或治疗有效量的权利要求1-4中任意一项所述的重组蛋白，权利要求5所述的纳米颗粒，权利要求6所述的免疫原性组合物，权利要求7所述的疫苗，权利要求9所述的核酸分子或权利要求11所述的载体。16.一种治疗流感相关疾病的方法，所述方法包括向需要其的对象施用预防或治疗有效量的权利要求1-4中任意一项所述的重组蛋白，权利要求5所述的纳米颗粒，权利要求6所述的免疫原性组合物，权利要求7所述的疫苗，权利要求9所述的核酸分子或权利要求11所述的载体。17.一种疫苗，所述疫苗包含权利要求1-4中任意一项所述的重组蛋白，以及联用生理学上可接受的运载体。18.一种检测抗流感抗体的方法，所述方法包括：a.将待检测所述抗流感抗体存在情况的样品的至少一部分与权利要求5所述的纳米颗粒接触；以及b.检测纳米颗粒抗流感抗体复合物的存在情况；其中检测到纳米颗粒抗体复合物表明所述样品含有抗流感抗体。

百度查询： 美国卫生与公众服务部 稳定化的第2组流感血凝素茎区三聚体及其用途

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