CN111164207A - 新型嵌合酶及其应用 - Google Patents
新型嵌合酶及其应用 Download PDFInfo
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- CN111164207A CN111164207A CN201880062068.9A CN201880062068A CN111164207A CN 111164207 A CN111164207 A CN 111164207A CN 201880062068 A CN201880062068 A CN 201880062068A CN 111164207 A CN111164207 A CN 111164207A
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Abstract
本发明涉及一种嵌合酶及其在生产具有5’末端帽的RNA分子中的应用，所述嵌合酶包含或由以下组成：至少一个加帽酶催化结构域和至少一个蛋白质‑RNA拴系系统RNA结合结构域。
Description
本发明涉及表达系统领域，特别是在真核细胞中的表达系统。
特别地，本发明涉及可用于生产具有5’-末端帽(cap)结构且优选具有3’poly(A)尾的RNA分子的嵌合酶。
在真核生物中，信使RNA(mRNA)的前体，即前体mRNA，是通过RNA聚合酶II合成的，然后进行多种转录后修饰，这是其生物学活性(包括翻译、稳定性或免疫应答调节)所必需的。这些修饰中的两种对mRNA代谢及其翻译特别重要：在其5’-末端加帽，以及在其3’-末端加聚腺苷酸尾。
加帽(capping)是一种几乎在所有真核信使RNA的5’-末端均存在的特殊结构。最简单的帽结构cap-0导致添加N7处甲基化的鸟嘌呤核苷，该鸟嘌呤核苷通过5’-5’三磷酸与初级RNA的末端连接(即m7GpppN，其中N是任何碱基，p表示磷酸，m表示甲基)。在所谓的典型途径中，cap-0的形成涉及一系列三个酶促反应：RNA三磷酸酶(RTPase)将未成熟的前体mRNA的5’三磷酸末端的γ磷酸残基去除成为二磷酸，RNA鸟苷酰基转移酶(GTase)将GMP从GTP转移到未成熟的RNA末端的二磷酸5’末端，RNA N7-鸟嘌呤甲基转移酶(N7-MTase)在GpppN帽的鸟嘌呤的氮7上添加一个甲基残基(Furuichi和Shatkin 2000)。在高级真核生物和某些病毒中，第一个(即cap-1结构；m7GpppNm2’-OpN)和第二个(即cap2结构；m7GpppNm2’- OpNm2’-O)转录的核苷酸的核糖的2'-羟基基团可以被两个单独的核糖-2’-O MTase(分别命名为cap1-和cap2-特异性MTase)甲基化(Langberg和Moss 1981)。然而，与细胞核专有的细胞N7-MTase活性相反，在HeLa细胞的细胞质和细胞核中均检测到了cap-1核糖-2’-O MTase活性，而cap2 MTase活性仅在其细胞质中发现(Langberg和Moss 1981)。
5’-末端m7GpppN帽的形成是前体mRNA加工的第一步。m7GpppN帽在mRNA稳定性及其从细胞核到细胞质的运输中起着重要作用(Huang和Steitz 2005，Kohler和Hurt 2007)。此外，通过锚定真核翻译起始因子4F(eIF4F)复合物(其介导核糖体小亚基的16S部分向mRNA的募集)，5’-末端m7GpppN帽对于mRNA翻译为蛋白质是非常重要的(Furuichi,LaFiandra等人1977,Gingras,Raught等人1999,Rhoads 1999)。因此，在体外(Lo,Huang等人1998)和在细胞中(Malone,Felgner等人1989,Gallie 1991,Lo,Huang等人1998,Kozak 2005)，5’-末端m7GpppN帽都大大增强了mRNA的翻译。通过RNA传感器RIG-1和MDA5来区分自身和非自身RNA，cap-0、cap-1和cap-2修饰参与了固有免疫反应，进而诱导I型干扰素反应(Hornung,Ellegast等人2006,Daffis,Szretter等人2010)。
由于它们广泛用于生命科学、生物技术和医学中，因此设计了许多表达系统以有效地生产蛋白质和/或RNA，特别是在真核细胞中生产。
发明人过去已经开发了用于在真核细胞中进行有效转基因的人工表达系统(即嵌合酶)，该系统自动产生mRNA分子，特别是在所述细胞的细胞质中(WO 2011/128444)。使用该系统，通过该嵌合酶的RNA聚合酶部分合成RNA链，并通过其mRNA加帽酶部分在5’-末端加帽。此外，通过从DNA模板转录聚腺苷轨道(track)，可以在转录物的3’-末端产生poly(A)尾。该系统具有显著的优点：不使用真核细胞的内源性RNA转录机器(machinery)，例如RNA聚合酶II和参与转录和转录后的相关因子。
通过融合RNA聚合酶II(POLR2A)的最大亚基的羧基末端结构域(CTD)尝试将加帽与转录偶联，从而提高T7 RNA聚合酶产生的未加帽转录物的可翻译性的其他尝试，必须增强细胞中组成性和选择性剪接底物的加帽(Kaneko,Chu等人2007,Natalizio,Robson-Dixon等人2009)。CTD包含共有序列1YSPTSPS7的25-52个七肽重复序列，该序列在整个进化过程中高度保守，并在转录周期中发生可逆的磷酸化作用(Palancade和Bensaude 2003)。当被磷酸化时，CTD被认为可以通过结合酵母(Cho,Takagi等人1997,McCracken,Fong等人1997)和哺乳动物(McCracken,Fong等人1997,Yue,Maldonado等人1997)中的mRNA加帽酶的一个或多个亚基来介导未成熟的转录物的转录和加帽的偶联。值得注意的是，带有Ser5-磷酸化CTD重复序列的RNA聚合酶II经历了启动子近端暂停，这与未成熟转录物的共转录加帽一致(Komarnitsky,Cho等人2000,Schroeder,Schwer等人2000)。然而，与直觉上期望的相反，CTD与单亚基(single-unit)T7 RNA聚合酶的融合不足以增强体内组成性和选择性剪接底物的加帽(Kaneko,Chu等人2007,Natalizio,Robson-Dixon等人2009)。
因此，在本领域中仍然非常需要适合于基因治疗和大规模蛋白质生产而没有细胞毒性或诱导的细胞毒性的新的改进的表达系统，特别是在真核细胞中的表达系统。本发明人在本文公开的发明中迈出了重要的一步。
本发明的目的是通过提供新的嵌合酶来满足这种需求，所述嵌合酶可以全部或部分地解决上述问题。
出乎意料的是，发明人特别证明了单体或寡聚嵌合(非天然)酶能够大大提高通过RNA聚合酶产生的特定mRNA的加帽率，所述单体或寡聚嵌合(非天然)酶包含加帽酶的催化结构域，特别是RNA三磷酸酶的催化结构域、鸟苷酰基转移酶的催化结构域、N7-鸟嘌呤甲基转移酶的催化结构域和蛋白质-RNA拴系系统(tethering system)的RNA结合结构域，所述RNA结合结构域特异性结合由特定RNA序列和/或结构组成的RNA元件。
这些结果令人吃惊，因为帽的形成是一个复杂的过程，并且外源转录物的加帽无法通过大多数其他方法实现，例如融合酶CTD-T7 RNA聚合酶(Kaneko,Chu等人2007,Natalizio,Robson-Dixon等人2009)。
美国专利5,866,680提出使用细胞核70K RNA结合结构域将RNA修饰活性引导至包括不同酶(例如RNA加帽酶)的RNA中的特定位点，但没有任何证明。
出乎意料的是，发明人还证明了与蛋白质-RNA拴系系统(例如所述美国专利5,866,680中描述的U1-RNA 70K(也称为SNRNP70)-U1snRNA系统)的其他RNA结合结构域相比，包含加帽酶的催化结构域和噬菌体蛋白-RNA拴系系统的噬菌体RNA结合结构域的细胞质单体或寡聚嵌合酶，可大大提高通过噬菌体DNA依赖性RNA聚合酶产生的特定mRNA的加帽率。
发明人还证明了，令人惊讶地，与融合有RNA结合结构域的poly(A)聚合酶存在下融合有RNA结合结构域的加帽酶的组合相比，包含加帽酶的催化结构域、蛋白质-RNA拴系系统的RNA结合结构域和poly(A)聚合酶的催化结构域的单体或寡聚嵌合酶可以协同地大大增加通过RNA聚合酶产生的特定mRNA的加帽率。这些结果是出乎意料的，因为加帽酶和poly(A)聚合酶自然情况下没有物理连接，并且不包含特定RNA序列的已知预测结合结构域。由于成分相同，本领域技术人员可能预期获得相同的表达率。
因此，一方面，本发明涉及一种嵌合酶，特别是细胞质嵌合酶，其包含或由以下组成：
-至少一个加帽酶催化结构域，特别地，所述加帽酶选自由以下组成的组：cap-0典型加帽酶、cap-0非典型加帽酶、cap-1加帽酶和cap-2加帽酶；和
-至少一个蛋白质-RNA拴系系统RNA结合结构域
其中所述RNA结合结构域与所述蛋白质-RNA拴系系统的RNA元件特异性结合，所述RNA元件由特定RNA序列和/或结构组成。
根据本发明的嵌合酶特别具有以下优点：
-它增加通过内源性DNA依赖性RNA聚合酶或非内源性DNA依赖性RNA聚合酶产生的mRNA的速率；
-它不昂贵、快速且易于实施，因此适用于大规模测定和蛋白质生产。
除非另有定义，否则本文中使用的所有技术和科学术语具有与相关领域的技术人员通常理解的相同的含义。
为了方便起见，提供了说明书、实施例和权利要求书中使用的某些术语和短语的含义。
如本文所用，术语“嵌合酶”是指不是在自然界中发现的天然酶的酶(即，非天然的)。因此，嵌合酶可以包含源自不同来源(例如，源自不同酶)的催化结构域，或源自相同来源(例如，源自相同酶)但是以不同于自然界中发现的方式排列的催化结构域。
术语“嵌合酶”涵盖单体(即，单亚基)酶，但也涵盖寡聚(即，多亚基(multi-unit))酶，特别是异源-寡聚酶。
更具体地，术语“嵌合酶”可以包括一种酶，该酶包含或由以下组成：与一个(即单亚基)或多个(即多亚基)催化结构域(至少一个加帽酶催化结构域)或蛋白质(至少一个加帽酶)共价或非共价连接的蛋白质-RNA拴系系统的RNA结合结构域。
术语酶的“催化结构域”是指对于确保酶功能而言是必需且足够(尤其是在其三维结构方面)的蛋白质结构域。例如，RNA三磷酸酶的催化结构域是为了确保RNA三磷酸酶功能所必需且足够的结构域。术语“催化结构域”涵盖野生型或突变型酶的催化结构域。
术语“蛋白质结构域”定义了蛋白质中独立地折叠和起作用的不同的功能性和/或结构性构件和元件。
特别地，根据本发明的嵌合酶是单体或寡聚非天然酶。
如本文所用，术语“单体酶”是指仅由一条多肽链组成的单亚基酶。
如本文所用，术语“寡聚酶”是指由共价或非共价连接在一起的至少两条多肽链组成的多亚基酶。术语“寡聚酶”涵盖多亚基酶，其中所述酶的至少两个亚基共价或非共价地连接在一起。术语“寡聚酶”包括同源寡聚酶(homo-oligomeric enzyme)和异源寡聚酶(hetero-oligomeric enzyme)，所述同源寡聚酶是仅由一种类型的单体(亚基)组成的多亚基酶，所述异源寡聚酶由不同类型的单体(亚基)组成。
如本文所用，术语“蛋白质-RNA拴系系统”是指这样一个系统，其中蛋白质(或肽)通过其RNA结合结构域识别并特异性结合(以高亲和力)由特定RNA序列和/或结构组成的特定RNA元件，从而使该蛋白质(或肽)与该RNA元件拴系在一起成为可能。蛋白质(或肽)通过其RNA结合结构域与特定RNA元件之间的特异性结合意味着蛋白质(或肽)与特定RNA元件以高亲和力相互作用。高亲和力相互作用包括亲和力为约10-6M或更强，特别是至少10-7M，至少10-8M，至少10-9M，更特别是至少10-10M的相互作用。RNA结合结构域是否以高亲和力特异性结合特定RNA元件可以很容易地尤其通过以下方式进行测试：将所述RNA结合结构域和特定RNA元件的反应与所述RNA结合结构域和除所述特定RNA元件以外的RNA的反应进行比较。
RNA-蛋白质亲和力可以通过本领域技术人员众所周知的各种方法来确定。这些方法包括但不限于稳态荧光或电泳测量、RNA电泳迁移率变化试验。
如本文所用，术语“蛋白质-RNA拴系系统的RNA结合结构域”是指对于确保识别由特定RNA序列和/或结构组成的特定RNA元件并以高亲和力与之相互作用，从而使该蛋白质(或肽)通过所述RNA结合结构域与该RNA元件拴系在一起成为可能而言，必需且足够(尤其是在其三维结构方面)的蛋白质(或肽)的结构域。因此，所述“RNA结合结构域”(RNA拴系结构域)结合RNA具有序列和/或结构特异性，即，特异性地结合由特定RNA序列和/或结构组成的RNA元件。术语“蛋白质-RNA拴系系统的RNA结合结构域”涵盖野生型或突变的蛋白(或肽)的RNA结合结构域。
根据本发明的嵌合酶至少包含所述RNA结合结构域，但是还可以包含含有所述RNA结合结构域的蛋白质(或肽)的全部或部分。实际上，根据本发明的嵌合酶的一个实施方案，所述蛋白质-RNA拴系系统的RNA结合结构域可以包含在蛋白质-RNA拴系系统的全部或部分蛋白质(或肽)中。
一些被表征的蛋白质-RNA拴系系统包括噬菌体蛋白质-RNA拴系系统，例如MS2外壳蛋白质-RNA拴系系统、R17外壳蛋白质-RNA拴系系统和λ形噬菌体N抗终止蛋白(lambdoidN antitermination protein)-RNA拴系系统。MS2外壳蛋白和R17外壳蛋白识别由茎-环RNA结构组成的特定RNA元件并以高亲和力与之相互作用(Valegard,Murray等人1994,Valegard,Murray等人1997)。λ形噬菌体N抗终止蛋白-RNA拴系系统识别由boxBL和boxBR茎环RNA结构组成的特定RNA元件并以高亲和力与之相互作用(Das 1993,Greenblatt,Nodwell等人1993,Friedman和Court 1995)。迄今为止表征的噬菌体属于λ形噬菌体家族(即λ、P22、Φ21、HK97和933W病毒)或与MS2相关的家族(即MS2和R17)。
其他充分表征的蛋白质-RNA拴系系统包括：(a)1型人免疫缺陷病毒(HIV-1；例如NCBI参考序列ABY50660.1)(Dingwall,Ernberg等人1990,Weeks,Ampe等人1990,Karn,Dingwall等人1991,Puglisi,Tan等人1992,Frankel和Young 1998)和牛免疫缺陷病毒(Puglisi,Chen等人1995)的TAT结合结构域，(b)HIV-1的Rev蛋白和突变体(例如UniProtKBP69718)，其与Rev结合元件(RBE)结合，RBE是短茎-内茎-环结构(Karn,Dingwall等人1991,Tan和Frankel 1995,Battiste,Mao等人1996,Frankel和Young 1998)，(c)任马病(Jembrana disease)病毒Tat蛋白(UniProtKB Q82854)，其与其自身的TAR蛋白，以及人和牛免疫缺陷病毒的TAR蛋白(Smith,Calabro等人2000)结合，(d)铁调节蛋白，例如铁反应性元件结合蛋白1(IREB1，例如UniProtKB P21399)和2(IREB2，例如UniProtKB P48200)，它们与铁代谢mRNA的5’UTR或3’UTR内的铁反应性元件结合(Theil 1994,Rouault 2006)，(e)雀麦花叶病毒(Brome mosaic virus)(BMV)外壳蛋白(UniProtKB Q5KSV1_BMV)，其与包装病毒RNA所需的MP编码区的发夹结合(Sacher和Ahlquist 1989,Choi和Rao 2003)，(f)U1A小核核糖核蛋白亚基70K(SNRNP70)，其以高特异性和亲和力与U1snRNA的3'UTR内的30个核苷酸的RNA发夹结合(Keene,Query等人1999)，(g)SLBP(茎环结合蛋白UniProtKB Q14493)，其结合组蛋白前体mRNA的3'UTR中的茎环结构(Marzluff,Wagner等人2008)，(h)60S核糖体蛋白L7(例如UniProtKB P18124)，其与28S rRNA中富含G的结构结合(Hemmerich,Bosbach等人1997)，(i)豇豆褪绿斑驳病(Cowpea chlorotic mottle)病毒衣壳蛋白(UniProtKBP03601)，其与包装病毒RNA所需的发夹结合(Annamalai,Apte等人2005)，(j)I型人类T细胞白血病病毒(HTLV-1)rex蛋白及其相关突变体(例如HTLV-1分离株Caribbea HS-35A亚型，UniProtKB P0C206)，其与位于所有I型人类T细胞白血病病毒特异性mRNA的3'长末端重复序列中的rex反应元件(RxRE)结合(Ballaun,Farrington等人1991,Jiang,Gorin等人1999)，(j)HTLV-2Rex蛋白(UniProtKB Q85601)，其结合5’长末端重复序列RNA(Yip,Dynan等人1991)，(m)RNA剪接组分U2 snRNP辅助因子U2AF65(例如UniProtKB P26368)，其结合在前体mRNA的3'剪接位点之前的聚嘧啶束(Zamore,Patton等人1992,Banerjee,Rahn等人2004)，(n)细菌核糖体蛋白S7(UniProtKB P02359)，其与16S核糖体RNA的3'主结构域的下半部分结合(Robert,Gagnon等人2000)，(o)古细菌L7Ae蛋白，其以纳摩尔级的亲和力与包含K转向(kink-turn)的RNA结合(Turner,Melcher等人2005,Ye,Yang等人2013)，(p)来自SELENBP2基因产物(硒结合蛋白2，SBP2)的RNA结合结构域，其与一些编码硒蛋白的mRNA的3'-UTR中的SECIS元件结合(Mix,Lobanov等人2007)，(q)雀麦花叶病毒(BMV)的N端结构域，其可结合BoxB(Yi,Vaughan等人2009)，(r)gp10头尾连接子的N-末端，其结合Φ29pRNA序列(Xiao,Moll等人2005)，(s)链霉亲和素，其结合人工RNA适体(Leppek和Stoecklin 2014)。
在根据本发明的嵌合酶的一个实施方案中，所述RNA结合结构域是噬菌体蛋白-RNA拴系系统的噬菌体RNA结合结构域，所述噬菌体蛋白-RNA拴系系统特别选自由以下组成的组：MS2外壳蛋白-RNA拴系系统、R17外壳蛋白-RNA拴系系统和λ形噬菌体N抗终止蛋白-RNA拴系系统，更特别地，为选自由以下组成的组的λ形噬菌体N抗终止蛋白-RNA拴系系统：λN抗终止蛋白-RNA拴系系统、phi21 N抗终止蛋白-RNA拴系系统、HK97 N抗终止蛋白-RNA拴系系统和p22 N抗终止蛋白-RNA拴系系统，更加特别地是λN抗终止蛋白-RNA拴系系统。
特别地，当本发明的嵌合酶用于向通过噬菌体DNA依赖性聚合酶(包括或不包括在所述嵌合酶中)合成的RNA分子添加5’-末端帽时，所述RNA结合结构域是噬菌体蛋白-RNA拴系系统的噬菌体RNA结合结构域。
实际上，出乎意料的是，发明人证明了与蛋白质-RNA拴系系统(包括美国专利5,866,680中描述的U1-RNA 70K系统)的其他RNA结合结构域相比，包含加帽酶的催化结构域和噬菌体蛋白-RNA拴系系统的噬菌体RNA结合结构域的细胞质单体或寡聚嵌合酶，可大大提高通过噬菌体DNA依赖性RNA聚合酶产生的特定mRNA的加帽率。
特别地，所述RNA结合结构域是选自由以下组成的组的噬菌体蛋白的噬菌体RNA结合结构域：野生型MS2外壳蛋白(NCBI登录号NC_001417.2，UniProtKB/Swiss-ProtP03612)、野生型R17外壳蛋白(NCBI登录号EF108465.1，UniProtKB/Swiss-Prot P69170)和野生型λ形噬菌体N抗终止蛋白以及能够识别特定RNA元件并以高亲和力与之相互作用的其突变体和衍生物，更特别地，为选自由以下组成的组的野生型λ形噬菌体N抗终止蛋白：野生型λN抗终止蛋白(NCBI登录号NC_001416.1，完整的基因组序列；UniProtKB/Swiss-Prot登录号P03045)、野生型phi21 N抗终止蛋白(NCBI登录号AH007390.1，部分基因组序列；UniProtKB/Swiss-Prot登录号P07243)、野生型HK97 N抗终止蛋白-RNA拴系系统(NCBI登录号NC_002167.1，完整的基因组序列；NCBI蛋白登录号NP_037732.1)和野生型p22 N抗终止蛋白(特别是NCBI序列(NCBI登录号NC_002371.2，完整的基因组序列；UniProtKB/Swiss-Prot登录号P04891)，更加特别地是野生型λN抗终止蛋白(NCBI登录号NC_001416.1，完整的基因组序列；UniProtKB/Swiss-Prot登录号P03045)。
完整或几乎完整的MS2/R17蛋白是与所拴系的RNA有效结合所必需的，因为散布在这些蛋白中的多个氨基酸残基参与茎-环相互作用(Valegard,Murray等人1994,Valegard,Murray等人1997)。
因此，在一个实施方案中，本发明的嵌合酶包含野生型MS2外壳蛋白(NCBI登录号NC_001417.2，UniProtKB/Swiss-Prot P03612)或其分离物野生型R17外壳蛋白(NCBI登录号EF108465.1，UniProtKB/Swiss-Prot P69170)或能够识别特定RNA元件并与之以高亲和力相互作用的其突变体或衍生物。
重要的是，来自λ形噬菌体N-蛋白的N-末端序列的18至22个氨基酸区域以类似于全长N-蛋白的亲和力和特异性与同源RNA序列结合(Franklin 1985,Cilley和Williamson1997)。
在本发明的嵌合体的一个实施方案中，所述蛋白质-RNA拴系系统的RNA结合结构域包含或由以下组成：由野生型λN抗终止蛋白(NCBI登录号NC_001416.1，完整的基因组序列；UniProtKB/Swiss-Prot登录号P03045)或野生型phi21 N抗终止蛋白(NCBI登录号AH007390.1，部分基因组序列；UniProtKB/Swiss-Prot登录号P07243)或野生型HK97 N抗终止蛋白-RNA拴系系统(NCBI登录号NC_002167.1，完整的基因组序列；NCBI蛋白登录号NP_037732.1)或野生型P22 N抗终止蛋白(NCBI登录号NC_002371.2，完整的基因组序列；UniProtKB/Swiss-Prot登录号P04891)或能够识别特定RNA元件并以高亲和力与之相互作用的其突变体或衍生物，特别是野生型λN抗终止蛋白(NCBI登录号NC_001416.1，完整的基因组序列；UniProtKB/Swiss-Prot登录号P03045)的1至22位，特别是1至18位氨基酸组成的肽。
特别地，所述蛋白质-RNA拴系系统的RNA结合结构域包含或由以下组成：野生型λN抗终止蛋白(NCBI登录号NC_001416.1，完整的基因组序列；UniProtKB/Swiss-Prot登录号P03045)的1至22位氨基酸组成的肽，特别是SEQ ID N°2，优选由SEQ ID N°1编码。
特别地，所述蛋白质-RNA拴系系统的RNA结合结构域不是源自与本发明的嵌合酶的不同催化结构域相同的来源(例如，源自相同的酶)。
根据本发明的嵌合酶可以是细胞核酶、亚细胞区室酶或细胞质酶。因此，根据本发明的嵌合酶可以包含本领域技术人员众所周知的、指导酶在细胞中的运输的信号肽。例如，根据本发明的嵌合酶可以包含将酶引导至细胞核的核定位信号(NLS)。这样的NLS通常是由五个带正电荷的碱性氨基酸组成的单元。NLS可以位于肽链上的任何位置。
优选地，根据本发明的嵌合酶是细胞质嵌合酶。特别地，除了到细胞质的运输之外，其不包含指导酶的运输的信号肽。
根据本发明的嵌合酶的细胞质定位具有以下优点：其通过避免大的DNA分子(即转基因)从细胞质向真核细胞细胞核的主动转移以及RNA分子从细胞核向细胞质的输出而优化了转基因表达的水平。
因此，根据本发明的这些细胞质嵌合酶可用于生产不依赖宿主的真核基因表达系统，该系统能够在细胞质中起作用，与细胞核相比，在细胞质中通常发现明显更高的转染DNA的量。
内源基因转录与转基因转录之间没有竞争，因为内源基因转录发生在真核细胞的细胞核中，与之相反，转基因转录发生在细胞质中。
因此，根据本发明的细胞质嵌合酶特别适用于大规模测定和蛋白质生产。
如本文所用，术语“加帽酶”是指能够在mRNA的5’-末端添加m7 GpppG帽和/或修饰RNA序列的最后一个或倒数第二个碱基的任何酶，包括cap-0典型或非典型加帽酶和cap-1或cap-2核苷2’甲基转移酶、N6-甲基-腺苷转移酶。
如本文所用，术语“cap-0典型加帽酶”是指能够通过涉及一系列三个酶促反应而在RNA分子的5’-末端添加cap-0结构的酶，所述三个酶促反应为：RNA三磷酸酶(RTPase)将未成熟的前体mRNA的5’三磷酸末端的γ磷酸残基去除成为二磷酸ppRNA，RNA鸟苷酰基转移酶(GTase)将GMP从GTP转移到二磷酸ppRNA未成熟RNA末端，RNA N7-鸟嘌呤甲基转移酶(N7-MTase)在鸟嘌呤的氮7上添加一个甲基残基得到GpppRNA帽(Furuichi和Shatkin 2000)。
参与典型的cap-0结构形成的真核生物和病毒的酶促结构域可以组装成可变数量的蛋白质亚基：
-具有所有三个关键酶促结构域(即RTPase、GTase和N7-MTase)的单亚基加帽酶。这些酶包括但不限于：(i)棘阿米巴多噬模拟病毒(Acanthamoeba Polyphaga mimivirus)加帽酶R382(Raoult,Audic等人2004,Benarroch,Smith等人2008)(NCBI APMV基因组序列NC_006450；UniProtKB/Swiss-Prot登录号Q5UQX1)，(ii)来自酵母乳酸克鲁维酵母(Kluyveromyces lactis)染色体外线性游离体pGKL2的ORF3加帽酶(Tommasino,Ricci等人1988,Tiggemann,Jeske等人2001)(NCBI乳酸克鲁维酵母CB 2359pGKL2基因组序列NC_010187；UniProtKB/Swiss-Prot登录号P05469)，(iii)非洲猪瘟病毒NP868R加帽酶(Pena,Yanez等人1993,Jais 2011,Dixon,Chapman等人2013,Jais,Decroly等人2018)(NCBI ASFV基因组序列毒株BA71V NC_001659；UniProtKB/Swiss-Prot登录号P32094)、VP4蓝舌病毒(Bluetongue virus)加帽酶(NCBI BTV血清型10基因组序列Y00421；UniProtKB/Swiss-Prot登录号P07132)。
-由两个亚基组成的加帽酶，包括但不限于：(i)由同时具有RTPase和GTase酶活性的RNGTT亚基(Yue,Maldonado等人1997,Pillutla,Yue等人1998,Tsukamoto,Shibagaki等人1998,Yamada-Okabe,Doi等人1998)(也称为HCE1；人和小鼠UniProtKB/Swiss-Prot登录号分别为O60942和O55236)和具有N7-MTase酶活性的RNMT(Pillutla,Yue等人1998,Tsukamoto,Shibagaki等人1998)(人和小鼠UniProtKB/Swiss-Prot登录号分别为Q05D80和D3YYS7)组成的哺乳动物加帽酶，(ii)由D1R基因产物和D12L基因产物组成的痘苗加帽酶，其中所述D1R基因产物具有RTPase、GTase和N7-MTase酶性结构域(Cong和Shuman 1993,Niles和Christen 1993,Mao和Shuman 1994,Cong和Shuman 1995,Mao和Shuman 1996,Myette和Niles 1996,Yu和Shuman 1996,Yu,Martins等人1997,Gong和Shuman 2003)(基因组序列毒株Western Reserve NC_006998.1；UniProtKB/Swiss-Prot登录号P04298)，所述D12L基因产物不具有内在的酶活性，但可以大大增强D1R亚基的RNA N7-MTase活性(Higman,Bourgeois等人1992,Higman,Christen等人1994,Mao和Shuman 1994,Schwer,Hausmann等人2006,De la Pena,Kyrieleis等人2007)(基因组序列菌株Western ReserveNC_006998.1；Gene 3707515；UniProtKB/Swiss-Prot登录号P04318)。-由三个亚基组成的加帽酶，例如具有RTPase的酿酒酵母(Saccharomyces cerevisiae)CET1(Tsukamoto,Shibagaki等人1997,Gu,Rajashankar等人2010)(UniProtKB/Swiss-Prot登录号O13297)、具有GTase的CEG1(Shibagaki,Itoh等人1992,Yamada-Okabe,Doi等人1998,Gu,Rajashankar等人2010)(UniProtKB/Swiss-Prot登录号Q01159)和具有N7-MTase催化活性的ABD1(Mao,Schwer等人1995,Schwer,Saha等人2000)(UniProtKB/Swiss-Prot登录号P32783)。
如本文所用，术语“cap-0非典型加帽酶”是指能够以不同于典型酶促过程的途径在RNA分子的5’-末端添加cap-0结构的酶。截止到今天，已经描述了三种非典型5’RNA帽合成机制：
-首先，α病毒科(alphavirus)(例如塞姆利基森林病毒(Semliki Forest virus)和辛德毕斯病毒(Sindbis virus)、马铃薯X病毒科(potexvirus)(例如竹花叶病毒(Bamboomosaic virus)、烟草花叶病毒科(tobamovirus)(例如烟草花叶病毒(Tobacco mosaicvirus))、披膜病毒科(Togaviridae)(例如风疹病毒(Rubella virus)和基孔肯雅病毒(Chikungunya virus))和肝炎病毒科(例如戊型肝炎病毒)的各种ss(+)RNA病毒的m7GTPRNA加帽途径(Decroly,Ferron等人2011)。该RNA加帽途径依赖于三个依次的酶促反应：(a)与常规途径相似的RTPase(例如，由非结构性P123多蛋白的明显裂解产生的塞姆利基森林病毒的nsP2蛋白；UniProtKB/Swiss-Prot登录号P08411)水解RNA 5’-末端的γ-β键，(b)非典型N7-MTase(例如同样由非结构性P123多蛋白的明显裂解产生的塞姆利基森林病毒的nsP1蛋白；UniProtKB/Swiss-Prot登录号P08411)将GTP分子甲基化，(c)然后，非典型GTase(例如同样是塞姆利基森林病毒的蛋白的nsP1)将m7GTP识别为底物，并转移到受体ppRNA的5’-末端，形成典型的m7GpppN cap-0结构(Decroly,Ferron等人2011)。除了RNA依赖性RNA聚合酶催化结构域之外，这三种酶活性可以在单个病毒蛋白中找到，即，竹花叶病毒mRNA加帽酶ORF1(Li,Shih等人2001,Huang,Han等人2004,Huang,Hsu等人2005,Han,Tsai等人2007)(NCBI BMV分离株BaMV-0基因组序列NC_001642；UniProtKB/Swiss-Prot登录号Q65005)。
-其次，弹状病毒科(例如水疱性口炎病毒(vesicular stomatitis virus)和狂犬病病毒)、副粘病毒科(例如人类呼吸道合胞病毒和麻疹病毒)、博纳病毒科(Bornaviridae)(例如博纳病毒(bornavirus))和丝状病毒科(例如埃博拉病毒和马尔堡病毒)的许多ss(-)RNA病毒的GDP RNA加帽途径(Decroly,Ferron等人2011)，它们在四个酶促步骤中催化cap-0/cap-1结构的形成。例如，来自人类呼吸道合胞病毒的单亚基大L蛋白(UniProtKB/Swiss-Prot登录号P28887)除了具有RNA依赖性RNA聚合酶活性之外，还可以单独完成这四个酶促步骤：(a)NTPase活性负责将GTP水解为GDP，(b)L蛋白水解pppRNA三磷酸部分的α-β键，从而释放焦磷酸并生成共价酶-pRNA中间体(即，具有5’-末端单磷酸的RNA)(c)然后将pRNA部分转移到GDP上，形成GpppN嵌段RNA。在这种情况下，只有α-磷酸根源于RNA，而β和γ-磷酸根都由GDP贡献，(d)最后，通过在第一个转录的核苷酸上分别在m7pppN和2’-残基上进行两次连续的甲基化完成cap-0以及cap-1结构的合成(Grdzelishvili,Smallwood等人2005,Li,Fontaine-Rodriguez等人2005,Grdzelishvili,Smallwood等人2006,Ogino和Banerjee2007,Li,Rahmeh等人2008,Ogino和Banerjee 2008,Rahmeh,Li等人2009)。
-第三，RNA帽抢夺(RNA cap snatching)，通过该过程，某些病毒无法合成它们自己的帽结构，从而通过从宿主mRNA窃取来获得帽。属于这一类的病毒包括代表性的正粘病毒科(Orthomyxoviridae)(例如流感病毒、索戈托病毒(Thogoto virus))、沙粒病毒科(Arenaviridae)(例如拉萨病毒(lassa virus)、马丘波病毒(machupo virus))和本扬病毒科(Bunyaviridae)(例如汉坦病毒(Hantaan virus)、拉克罗斯病毒(La Crosse virus)、番茄斑萎病毒)(Decroly,Ferron等人2011)。为了获得其帽结构，通过包含在病毒RNA依赖性RNA聚合酶中的核酸内切酶活性从宿主加帽mRNA的5’-末端切割10至20个核苷酸大小的核苷酸序列，并将其转移到病毒基因组RNA。随后，将获得的加帽的前导序列用于引发病毒基因组RNA的转录，最终导致合成加帽的可翻译的病毒mRNA。沙粒病毒科和本扬病毒科表达一种大的单体聚合酶，以确保帽抢夺。正粘病毒科流感病毒具有异源三聚体聚合酶，其由PB1(UniProtKB/Swiss-Prot登录号毒株A/Puerto Rico/8/1934 H1N1 P03431)、PB2(UniProtKB/Swiss-Prot登录号毒株A/Puerto Rico/8/1934 H1N1 P03428)和PA(UniProtKB/Swiss-Prot登录号毒株A/Puerto Rico/8/1934 H1N1 P03433)组成。所有这三个亚基都是核酸内切酶活性所必需的，但是认为酶活性存在于PA亚基的氨基末端结构域中(Ohlmann,Rau等人1995)。
如本文所用，术语“cap-1加帽酶”是指能够在RNA分子的5’末端添加cap-1结构的酶。
如本文所用，术语“cap-2加帽酶”是指能够在RNA分子的5’末端添加cap-2结构的酶。
在哺乳动物、高等真核生物和某些病毒中，发现了两种帽修饰，这在酵母和植物mRNA中是没有的(Langberg和Moss 1981)，是由mRNA的5’末端的核苷酸的核糖部分的2’羟基基团的甲基化而产生的：第一个mRNA核苷酸上的cap-1和第二个mRNA核苷酸上的cap-2。在几乎所有的哺乳动物mRNA分子上都发现了cap-1甲基化，而只有一半的mRNA在第二个转录的核苷酸上含有2’-O-甲基化残基。
在哺乳动物中，由两个核糖-2’-O甲基转移酶(也称为核苷-2’-甲基转移酶或2’-O-MTase)(Belanger,Stepinski等人2010)进行cap-1和cap-2修饰。首先是MTR1(cap-1核糖-2’-O MTase活性，也称为FTSJD2、KIAA0082或ISG95；UniProtKB/Swiss-Prot登录号Q8N1G2)，其仅存在于细胞核中，并包含推定的核定位信号和可能参与RNA结合的G蛋白修补结构域(Haline-Vaz,Silva等人2008)。值得注意的是，MRT1与RNA聚合酶II的CTD相关联，这表明cap-1的形成发生在mRNA合成的早期(Langberg和Moss 1981)。其次，MTR2(cap 2核糖-2’-O MTase，也称为FTSJD1或FLJ11171；UniProtKB/Swiss-Prot登录号Q8IYT2)将甲基基团从S-腺苷甲硫氨酸转移至mRNA和小核RNA的第二个核苷酸的2’-O-核糖。MTR2不需要鸟苷cap-0或cap-1修饰的N7甲基化，但是cap-1的存在会增加MTR2的活性。与细胞核MTR1相反，MTR2蛋白分布在整个细胞核和细胞质溶胶中(Keith,Ensinger等人1978)。
一些真核病毒具有它们自己的cap-1和/或cap-2 2’-O-MTase，包括：
-来自痘苗病毒的VP39(NCBI基因组序列NC_006998.1；UniProtKB/Swiss-Prot登录号YP_232977)，其同时具有cap-1和cap-2 2’-O-MTase酶活性(Schnierle,Gershon等人1994,Shi,Yao等人1996,Hu,Gershon等人1999)，
-来自苜蓿银纹夜蛾核型多角体病毒(Autographa californicaNucleopolyhedrovirus)的Orf69(NCBI基因组序列NC_001623.1；UniProtKB/Swiss-Prot登录号P41469)(Wu和Guarino 2003)，
-来自冠状病毒的nsp16(人类SARS冠状病毒的多蛋白1ab的残基6776-7073NCBI基因组序列NC_004718.3；UniProtKB/Swiss-Prot登录号P0C6X7)(Reinisch,Nibert等人2000,Decroly,Imbert等人2008,Chen,Cai等人2009,Lugari,Betzi等人2010)，
-来自呼肠孤病毒(Reovirus)(例如3型哺乳动物正呼肠孤病毒(orthoreovirus)，Dearing株；NCBI基因组序列J03488；UniProtKB/Swiss-Prot登录号P11079)的λ2蛋白，除GTase和N7-MTase酶活之外，它还具有2’-O-MTase(Bujnicki和Rychlewski 2001)；来自蓝舌病毒(NCBI BTV血清型10基因组序列ID Y00421；UniProtKB/Swiss-Prot登录号P07132)的VP4，来自黄病毒属(包括登革热病毒黄热病毒、寨卡病毒、西尼罗河病毒、Meaban病毒、Yokose病毒、圣路易斯脑炎病毒、日本脑炎病毒、蜱传脑炎病毒)的NS5(例如来自1型登革热病毒毒株Nauru/West Pac/1974的多蛋白；NCBI基因组序列U88535；UniProtKB/Swiss-Prot登录号P17763)，也可以甲基化mRNA的内部腺苷残基(Dong,Chang等人2012)。
在一个实施方案中，根据本发明的嵌合酶包含或由以下组成：
-至少一个RNA三磷酸酶催化结构域；
-至少一个鸟苷酰基转移酶催化结构域；
-至少一个N7-鸟嘌呤甲基转移酶催化结构域；和
-至少一个蛋白质-RNA拴系系统RNA结合结构域；
特别地，其中至少一个所述催化结构域是cap-0典型加帽酶的催化结构域，更特别地是病毒cap-0典型加帽酶的催化结构域。
如本文所用，术语“RNA三磷酸酶”(RTPase)是指这样一种酶，其将未成熟的前体mRNA的5’三磷酸末端的γ磷酸残基去除，得到二磷酸(Furuichi和Shatkin 2000)。
如本文所用，术语“RNA鸟苷酰基转移酶”(GTase)是指将GMP从GTP转移到未成熟的二磷酸RNA末端的酶(Furuichi和Shatkin 2000)。
如本文所用，术语“N7-鸟嘌呤甲基转移酶”(N7-MTase)是指将鸟嘌呤的氮7上的甲基残基添加到GpppN帽上的酶(Furuichi和Shatkin 2000)。
所述RNA三磷酸酶、鸟苷酰基转移酶，N7-鸟嘌呤甲基转移酶的催化结构域可以是相同或不同的加帽酶。
优选地，所述RNA三磷酸酶、鸟苷酰基转移酶、N7-鸟嘌呤甲基转移酶的催化结构域来自一种或几种细胞质酶，其有利地具有相对简单的结构和充分表征的酶活性。因此，特别地，所述RNA三磷酸酶、鸟苷酰基转移酶、N7-鸟嘌呤甲基转移酶的催化结构域可以是一种或几种病毒加帽酶的催化结构域或细胞质游离体的加帽酶的催化结构域。
在一个实施方案中，所述RNA三磷酸酶、鸟苷酰基转移酶、N7-鸟嘌呤甲基转移酶的催化结构域来自一种或几种病毒加帽酶，特别是选自由以下组成的组：野生型痘苗病毒加帽酶、野生型蓝舌病毒加帽酶、野生型竹花叶病毒加帽酶、野生型非洲猪瘟病毒加帽酶、野生型棘阿米巴多噬模拟病毒加帽酶、野生型1型有机湖藻DNA病毒(Organic Lakephycodnavirus 1，OLPV1)加帽酶、野生型2型有机湖藻DNA病毒(OLPV2)加帽酶、野生型球形棕囊藻病毒(Phaeocystis globosa virus)加帽酶、野生型金色藻病毒(Chrysochromulinaericina virus)加帽酶以及能够分别将未成熟的前体mRNA的5’三磷酸末端的γ磷酸残基去除成为二磷酸或将GMP从GTP转移到二磷酸未成熟RNA末端或在GpppN帽的鸟嘌呤的氮7上添加一个甲基残基的其突变体或衍生物，更特别是野生型非洲猪瘟病毒加帽酶以及能够分别将未成熟的前体mRNA的5’三磷酸末端的γ磷酸残基去除成为二磷酸或将GMP从GTP转移到二磷酸未成熟RNA末端或在GpppN帽的鸟嘌呤的氮7上添加一个甲基残基的其突变体或衍生物。
如本文所用，术语“痘苗病毒加帽酶”是指痘苗病毒的异二聚体的D1R/D12L加帽酶。D1R基因产物具有RTPase、GTase和N7-MTase酶性结构域(Cong和Shuman 1993,Niles和Christen 1993,Mao和Shuman 1994,Cong和Shuman 1995,Mao和Shuman 1996,Myette和Niles 1996,Yu和Shuman 1996,Yu,Martins等人1997,Gong和Shuman 2003)(基因组序列毒株Western Reserve NC_006998.1；UniProtKB/Swiss-Prot登录号P04298)，D12L基因产物不具有内在的酶活性，但可以大大增强D1R亚基的RNA N7-MTase活性(Higman,Bourgeois等人1992,Higman,Christen等人1994,Mao和Shuman 1994,Schwer,Hausmann等人2006,De laPena,Kyrieleis等人2007)(基因组序列菌株Western Reserve NC_006998.1；Gene3707515；UniProtKB/Swiss-Prot登录号P04318)。
如本文所用，术语“蓝舌病毒加帽酶”是指蓝舌病毒(BTV)的单亚基VP4加帽酶，其为76kDa蛋白(644个氨基酸；对于序列，参见例如NCBI BTV血清型10基因组序列Y00421；Gene 2943157；UniProtKB/Swiss-Prot P07132)。该加帽酶很可能能够通过位于其羧基末端附近的亮氨酸拉链进行同源二聚化(Ramadevi,Rodriguez等人1998)。VP4催化mRNAm7GpppN加帽合成所需的所有三个酶促步骤：RTPase(Martinez-Costas,Sutton等人1998)、GTase(Martinez-Costas,Sutton等人1998,Ramadevi,Burroughs等人1998)和N7-MTase(Ramadevi,Burroughs等人1998)。
如本文所用，术语“竹花叶病毒加帽酶”是指ORF1，其是竹花叶病毒(BMV)mRNA加帽酶，是单亚基155-kDa蛋白(1365-氨基酸；NCBI BMV分离株BaMV-O基因组序列NC_001642；Gene 1497253；UniProtKB/Swiss-Prot Q65005)。ORF1蛋白具有产生m7GpppN mRNA加帽所需的所有酶促活性，即RTPase(Li,Shih等人2001,Han,Tsai等人2007)、GTase和N7-MTase(Li,Chen等人2001,Li,Shih等人2001)。
如本文所用，术语“非洲猪瘟病毒加帽酶”(ASFV)是指NP868R加帽酶(G4R)，其是单亚基100kDa蛋白(868个氨基酸；NCBI ASFV基因组序列毒株BA71V NC_001659；Gene1488865；UniProtKB/Swiss-Prot P32094)，具有产生m7GpppN mRNA加帽所需的所有酶促活性，即RTPase、GTase和N7-MTase(Pena,Yanez等人1993,Jais 2011,Dixon,Chapman等人2013,Jais,Decroly等人2018)。
如本文所用，术语“棘阿米巴多噬模拟病毒加帽酶”是指R382(APMV)，其是另一种单亚基136.5kDa蛋白(1170个氨基酸；NCBI APMV基因组序列NC_006450；Gene 3162607；UniProtKB/Swiss-Prot Q5UQX1)。
如本文所用，术语“1型有机湖藻DNA病毒(OLPV1)加帽酶”是指889个氨基酸的单亚基，参考NCBI登录号ADX05869.1所示。
如本文所用，术语“2型有机湖藻DNA病毒(OLPV2)加帽酶”是指942个氨基酸的单亚基，参考NCBI登录号ADX06468.1。
如本文所用，术语“球形棕囊藻病毒加帽酶”是指1066个氨基酸的单亚基，参考UniProtKB/Swiss-Prot YP_008052553.1。
如本文所用，术语“Chrysochromulina ericina病毒加帽酶”是指965个氨基酸的单亚基，参考UniProtKB/Swiss-Prot YP_009173557.1。
在一个实施方案中，所述RNA三磷酸酶、鸟苷酰基转移酶、N7-鸟嘌呤甲基转移酶的催化结构域来自细胞质游离体的加帽酶，例如pGKL2。特别地，所述RNA三磷酸酶、鸟苷酰基转移酶、N7-鸟嘌呤甲基转移酶的催化结构域包括在由乳酸克鲁维酵母pGKL2(NCBI乳酸克鲁维酵母CB 2359pGKL2基因组序列NC_010187；UniProtKB/Swiss-Prot P05469)的ORF3基因编码的酵母线性染色体外游离体pGKL2的加帽酶的全部或部分中，它是594个氨基酸的蛋白质(70.6kDa的蛋白质)。
在根据本发明的嵌合酶的一个实施方案中，所述RNA三磷酸酶催化结构域、所述鸟苷酰基转移酶催化结构域以及所述N7-鸟嘌呤甲基转移酶催化结构域包含在单体中，即在一条多肽中。例如，所述单体可以是根据本发明的单体加帽酶或单体嵌合酶。
特别地，所述RNA三磷酸酶催化结构域、所述鸟苷酰基转移酶催化结构域以及所述N7-鸟嘌呤甲基转移酶催化结构域包含在单体加帽酶中。在这种情况下，根据本发明的嵌合酶包含单体加帽酶，其包括所述RNA三磷酸酶催化结构域、所述鸟苷酰基转移酶催化结构域和所述N7-鸟嘌呤甲基转移酶催化结构域。所述单体加帽酶可以是单体病毒加帽酶，特别是选自由以下组成的组：野生型蓝舌病毒加帽酶、野生型竹花叶病毒加帽酶、野生型非洲猪瘟病毒加帽酶、野生型棘阿米巴多噬模拟病毒加帽酶、野生型OLPV1加帽酶、野生型OLPV2加帽酶、野生型球形棕囊藻病毒加帽酶、野生型金色藻病毒加帽酶以及能够在RNA分子的5’-末端添加m7GpppN帽的其突变体和衍生物，特别是野生型非洲猪瘟病毒加帽酶以及能够在RNA分子的5’-末端添加m7GpppN帽的其突变体和衍生物，甚至更特别是野生型非洲猪瘟病毒加帽酶。
根据本发明的嵌合酶还可以进一步包含这样的结构域，所述结构域可以增强本发明的嵌合酶的至少一个催化结构域的活性，特别是至少一个加帽酶催化结构域的活性，更特别是选自由以下组成的组的至少一个催化结构域的活性：RNA三磷酸酶催化结构域、鸟苷酰基转移酶催化结构域、N7-鸟嘌呤甲基转移酶催化结构域，优选N7-鸟嘌呤甲基转移酶催化结构域。
例如，增强本发明的嵌合酶的至少一个催化结构域的活性的所述结构域可以是由痘苗病毒D12L基因(基因组序列NC_006998.1；Gene3707515；UniProtKB/Swiss-Prot YP_232999.1)编码的31-kDa亚基，其没有内在的酶活性，但是可以大大增强痘苗mRNA加帽酶D1R亚基的RNAN7-鸟嘌呤甲基转移酶活性(Higman,Bourgeois等人1992,Higman,Christen等人1994,Mao和Shuman 1994)。
在一个实施方案中，根据本发明的嵌合酶还包含至少一个、除cap-0、cap-1和cap-2加帽酶之外的5’-末端RNA加工酶的催化结构域。
如本文所用，术语“除cap-0、cap-1和cap-2加帽酶之外的5’-末端RNA加工酶”是指除cap-0、cap-1和cap-2加帽酶之外的、能够修饰mRNA序列的最后一个或倒数第二个碱基的酶，包括N6-甲基-腺苷转移酶，和能够在RNA分子5’末端添加2,2,7-三甲基鸟苷(TMG)和2,7-三甲基鸟苷(DMG)帽修饰的酶。
已经在真核生物或病毒mRNA中表征了除cap-0、cap-1、cap-2修饰以外的其他加帽修饰。例如，第一个碱基被N6-甲基-腺苷转移酶m6A甲基化是一种影响细胞mRNA命运的可逆修饰(Mauer,Luo等人2017)。snRNA、端粒酶RNA、反式剪接线虫mRNA和某些病毒mRNA上存在的2,2,7-三甲基鸟苷(TMG)和2,7-三甲基鸟苷(DMG)帽修饰可以赋予翻译优势(Darzynkiewicz,Stepinski等人1988,Cai,Jankowska-Anyszka等人1999)。TMG和DMG由来自病毒(例如来自DNA拟病毒的L320，UniProtKB/Swiss-Prot登录号Q5UQR2(Benarroch,Qiu等人2009)；原生动物(例如蓝氏贾第鞭毛虫(Giardia lamblia)三甲基鸟苷合成酶(NCBI登录号EAA46438(Hausmann和Shuman 2005)；低等真核生物(例如粟酒裂殖酵母(Schizosaccharomyces pombe)三甲基鸟苷合酶，UniProtKB/Swiss-Prot登录号Q09814(Hausmann和Shuman 2005,Hausmann,Zheng等人2008,Benarroch,Jankowska-Anyszka等人2010)和哺乳动物(例如人三甲基鸟苷合酶1UniProtKB/Swiss-Prot登录号Q96RS0(Zhu,Qi等人2001,Hausmann,Zheng等人2008,Benarroch,Jankowska-Anyszka等人2010)的专门酶进行。
在一个实施方案中，根据本发明的嵌合酶还包含至少一个poly(A)聚合酶催化结构域。
实际上，本发明人出乎意料地证明，与融合有RNA结合结构域的poly(A)聚合酶存在下融合有RNA结合结构域的加帽酶的组合相比，包含至少一个加帽酶催化结构域、至少一个蛋白质-RNA拴系系统RNA结合结构域和至少一个poly(A)聚合酶催化结构域的单体或寡聚嵌合酶可以协同地大大增加通过RNA聚合酶产生的特定mRNA的加帽率。这些结果是出乎意料的，因为加帽酶和poly(A)聚合酶自然情况下没有物理连接，并且不包含特定RNA序列的结合结构域。由于成分相同，本领域技术人员可能预期获得相同的表达率。
如本文所用，术语“poly(A)聚合酶”是指能够催化腺苷残基从ATP非模板化添加到RNA分子的3’末端的任何酶。
在一个实施方案中，所述poly(A)聚合酶催化结构域是典型poly(A)聚合酶的催化结构域，所述典型poly(A)聚合酶包括哺乳动物(例如PAPOLA、PAPOLG)、酵母(例如酿酒酵母PAP1、粟酒裂殖酵母PLA1、白色念珠菌PAP、卡氏肺孢子虫(Pneumocystis carinii)PAP)、原生动物、病毒和细菌的典型poly(A)聚合酶。
特别地，所述poly(A)聚合酶催化结构域是细胞质的典型poly(A)聚合酶，更特别地选自由以下组成的组：
-哺乳动物细胞质poly(A)聚合酶，包括PAPOLB(人和小鼠PAPOLB，UniProtKB/Swiss-Prot登录号分别为Q9NRJ5和Q9WVP6)，其至少部分是细胞质酶(Kashiwabara,Zhuang等人2000,Lee,Lee等人2000,Kashiwabara,Tsuruta等人2016)；PAPOLA(人和小鼠PAPOLA，UniProtKB/Swiss-Prot登录号分别为P51003和Q61183)的突变体，其中核定位信号的突变或缺失可以使核酶重新定位到细胞质中(Raabe,Murthy等人1994,Vethantham,Rao等人2008)，以及PAPOLG(人和小鼠PAPOLG，UniProtKB/Swiss-Prot登录号分别为Q9BWT3和Q6PCL9)的突变体，其中核定位信号的突变或缺失很可能使核酶重新定位到细胞质中(Kyriakopoulou,Nordvarg等人2001)，
-酵母或原生动物poly(A)聚合酶，例如酿酒酵母PAP1(UniProtKB/Swiss-Prot登录号P29468)；粟酒裂殖酵母PLA1(UniProtKB/Swiss-Prot登录号Q10295)、白色念珠菌PAP(UniProtKB/Swiss-Prot登录号Q9UW26)、卡氏肺孢子虫PAP(也称为耶氏肺孢子虫(Pneumocystis jiroveci)；UniProtKB/Swiss-Prot登录号A0A0W4ZDF2)的突变体，其中核定位信号的突变或缺失很可能使核酶重新定位到细胞质中(Lingner,Kellermann等人1991)，以及其他嗜冷性、嗜温性、嗜热性或超嗜热性酵母或原生动物株，
-病毒poly(A)聚合酶，包括由VP55催化亚基(UniProtKB/Swiss-Prot登录号毒株Western Reserve P23371)和作为持续性因子(processivity factor)的VP39(UniProtKB/Swiss-Prot登录号毒株Western Reserve P07617)组成的异源二聚体痘苗病毒poly(A)聚合酶，其他痘病毒poly(A)聚合酶(例如牛痘病毒、猴痘病毒或骆驼痘病毒)、非洲猪瘟病毒(C475L，UniProtKB/Swiss-Prot登录号A0A0A1E081)、棘阿米巴多噬菌体拟病毒R341(UniProtKB/Swiss-Prot登录号E3VZZ8)和Megavirus chilensis Mg561 poly(A)聚合酶(NCBI登录号：YP_004894612)、Moumouvirus(NCBI登录号AEX62700)、Mamavirus(NCBI登录号AEQ60527)、Cafeteria roenbergensis BV-PW1病毒(NCBI登录号YP_003969918)、Megavirus lba(NCBI登录号AGD92490)、黄石湖模拟病毒(NCBI登录号YP_009174112)、Chrysochromulina ericina病毒(NCBI登录号YP_009173345)、1型有机湖藻DNA病毒(NCBI登录号ADX05881)、2型有机湖藻DNA病毒(NCBI登录号ADX06298)、Faustovirus(NCBI登录号AMN83802)和球形棕囊藻病毒(NCBI登录号YP_008052392)。
-以及能够催化腺苷残基从ATP非模板化添加到RNA分子的3’末端的其突变体或衍生物。
在另一个实施方案中，所述poly(A)聚合酶催化结构域是非典型poly(A)聚合酶催化结构域，特别是细胞质非典型poly(A)聚合酶催化结构域。“非典型poly(A)聚合酶”是指不具有涉及N-末端核苷酸基转移酶(NT)催化结构域、中央结构域和C-末端结构域(对应于RNA结合结构域(RBD))的三方结构的酶(Trippe,Sandrock等人1998)。截至今天，已经在哺乳动物中描述了以下非典型poly(A)聚合酶：
-首先，poly(A)聚合酶GLD2(也称为PAPD4；人和小鼠GLD2，UniProtKB/Swiss-Prot登录号分别为Q6PIY7和Q91YI6)，其最初通过与秀丽隐杆线虫GLD2的序列相似性而鉴定(Wang,Eckmann等人2002)。此外，GLD4(UniProtKB/Swiss-Prot登录号G5EFL0)，作为GLD2同源物，已经在秀丽隐杆线虫中被鉴定，但在哺乳动物中却没有(Schmid,Kuchler等人2009)
-其次，单亚基线粒体poly(A)聚合酶(也称为PAPD1、TUTase1或mtPAP；人和小鼠PAPD1，UniProtKB/Swiss-Prot登录号分别为Q9NVV4和Q9D0D3)，
-第三，核仁RBM21 poly(A)聚合酶(也称为U6 TUTase、TUT1或Star-PAP；斑点靶向PIPKIa调控的poly(A)聚合酶；人和小鼠RBM21，UniProtKB/Swiss-Prot登录号分别为Q9H6E5和Q8R3F9)，
-第四和第五，推定的细胞质/核PAPD5(人和小鼠PAPD5，UniProtKB/Swiss-Prot登录号分别为Q8NDF8和Q68ED3)和PAPD7(也称为POLS，人和小鼠PAPD7，UniProtKB/Swiss-Prot登录号分别为Q5XG87和Q6PB75)，它们与来自核TRAMP复合物(Trf4/Air2/Mtr4p聚腺苷酸化复合物)的酵母poly(A)聚合酶TRF4或TRF5同源(Haracska,Johnson等人2005)，
-第六和第七，细胞质ZCCHC6(人和小鼠ZCCHC6，UniProtKB/Swiss-Prot登录号分别为Q5VYS8和Q5BLK4)和ZCCHC11 poly(A)聚合酶(人和小鼠ZCCHC11，UniProtKB/Swiss-Prot登录号分别为Q5TAX3和B2RX14)。
在一个实施方案中，所述poly(A)聚合酶催化结构域是酵母或原生动物poly(A)聚合酶催化结构域，特别是选自由以下组成的组：野生型酿酒酵母PAP1poly(A)聚合酶、粟酒裂殖酵母PLA1、白色念珠菌PAP(UniProtKB/Swiss-Prot登录号Q9UW26)、卡氏肺孢子虫PAP(UniProtKB/Swiss-Prot登录号A0A0W4ZDF2)，酿酒酵母、粟酒裂殖酵母、白色念珠菌或卡氏肺孢子虫poly(A)聚合酶的细胞质突变体(其中核定位信号是无功能的或缺失的)，以及能够催化腺苷残基从ATP非模板化添加到RNA分子的3’末端的其突变体或衍生物。
与哺乳动物典型poly(A)聚合酶相比，酵母或原生动物poly(A)聚合酶具有显著优势：其分子量低(例如，酿酒酵母PAP1和粟酒裂殖酵母PLA1 poly(A)聚合酶有568和566个氨基酸，人PAPOLA(UniProtKB/Swiss-Prot登录号P51003)有745个氨基酸)，并且在拴系测定中具有高持续性(Dickson,Thompson等人2001)。如本文所用，术语“PAP1 poly(A)聚合酶”是指酿酒酵母PAP1 poly(A)聚合酶(UniProtKB/Swiss-Prot登录号P29468)。
如本文所用，术语“PLA1 poly(A)聚合酶”是指粟酒裂殖酵母PLA1 poly(A)聚合酶，UniProtKB/Swiss-Prot登录号Q10295。
所述poly(A)聚合酶催化结构域可以是病毒poly(A)聚合酶催化结构域，特别是选自由以下组成的组：野生型VP55 poly(A)聚合酶、野生型C475L poly(A)聚合酶、野生型R341 poly(A)聚合酶和野生型MG561 poly(A)聚合酶以及能够催化腺苷残基从ATP非模板化添加到RNA分子的3’末端的其突变体或衍生物。
与哺乳动物poly(A)聚合酶相比，病毒poly(A)聚合酶具有显著优势：其分子量低，其酶活性强，当在哺乳动物细胞中表达时，存在于细胞质区室中。此外，其中一些酶，例如R341和MG561 poly(A)聚合酶不需要任何已知的辅助蛋白辅因子。
如本文所用，术语“VP55 poly(A)聚合酶”是指异二聚体的痘苗病毒poly(A)聚合酶的催化亚基(UniProtKB/Swiss-Prot登录号毒株Western Reserve P23371)。VP39是第二个亚基，充当持续性因子(UniProtKB/Swiss-Prot登录号菌株Western Reserve P07617)。
如本文所用，术语“C475L poly(A)聚合酶”是指非洲猪瘟病毒poly(A)聚合酶(UniProtKB/Swiss-Prot登录号AOA0A1E081)。
如本文所用，术语“R341 poly(A)聚合酶”是指棘阿米巴多噬模拟病毒R341poly(A)聚合酶(UniProtKB/Swiss-Prot登录号E3VZZ8)。
如本文所用，术语“MG561 poly(A)聚合酶”是指Megavirus chilensis MG561poly(A)聚合酶(NCBI登录号：YP_004894612)。
在一个实施方案中，根据本发明的嵌合酶还包含至少一个DNA依赖性RNA聚合酶催化结构域。
如本文所用，术语“DNA依赖性RNA聚合酶”(RNAP)是指沿5’→3'方向从单链或双链DNA模板合成RNA的互补链的核苷酸基转移酶。
优选地，所述DNA依赖性RNA聚合酶催化结构域是具有相对简单的结构，并且更优选地，具有表征的基因组酶性调节元件(即启动子和转录终止信号)的酶的催化结构域。因此，特别地，所述DNA依赖性RNA聚合酶催化结构域可以是噬菌体DNA依赖性RNA聚合酶、细菌DNA依赖性RNA聚合酶或各种真核细胞器(例如线粒体、叶绿体和原质体)的DNA依赖性RNA聚合酶的催化结构域。
在一个实施方案中，所述DNA依赖性RNA聚合酶催化结构域是噬菌体DNA依赖性RNA聚合酶催化结构域。
噬菌体DNA依赖性RNA聚合酶具有以下显著优点：它们可以优化转基因表达的水平，特别是通过具有比真核RNA聚合酶更高的持续性。噬菌体DNA依赖性RNA聚合酶的结构也比大多数细胞核真核聚合酶简单得多，后者由多个亚基(例如RNA聚合酶II)和转录因子组成。到目前为止表征的大多数噬菌体DNA依赖性RNA聚合酶都是单亚基酶，它们不需要辅助蛋白来起始、延伸或终止转录(Chen和Schneider 2005)。这些酶中有几种(以克隆它们的噬菌体命名)还具有充分表征的调节基因组元件(即启动子和终止信号)，这对转基因很重要。
内源基因转录和转基因转录之间也没有竞争。包含噬菌体DNA依赖性RNA聚合酶部分的本发明的嵌合酶允许在任何真核物种(例如酵母、啮齿动物和人)中产生RNA转录物。它们不昂贵、快速且易于实施，因此适用于大规模测定和蛋白质生产；它允许在任何生物系统(例如非细胞反应混合物、培养的细胞和活生物体)中产生RNA转录物，因为与真核RNA聚合酶(如RNA聚合酶II)相反，大多数噬菌体DNA依赖性RNA聚合酶不需要相关因子来起始、延伸或终止转录。
所述噬菌体DNA依赖性RNA聚合酶的催化结构域可以是特别选自由以下组成的组的噬菌体DNA依赖性RNA聚合酶的催化结构域：野生型T7 RNA聚合酶(NCBI基因组序列NC_001604；Gene 1261050；UniProtKB/Swiss-Prot P00573)、野生型T3 RNA聚合酶(NCBI基因组序列NC_003298；Gene 927437；UniProtKB/Swiss-Prot Q778M8)、野生型K1E RNA聚合酶(NCBI基因组序列AM084415.1,UniProtKB/Swiss-Prot Q2WC24)、野生型K1-5 RNA聚合酶(NCBI基因组序列AY370674.1,NCBI YP_654105.1)、野生型K11 RNA聚合酶(NCBI基因组K11RNAP序列NC_004665；Gene 1258850；UniProtKB/Swiss-Prot Q859H5)、野生型
RNA聚合酶(NCBI基因组序列NC_004777；Gene 1733944；UniProtKB/Swiss-Prot蛋白Q858N4)、野生型
RNA聚合酶(NCBI基因组序列NC_001271；Gene 1262422；UniProtKB/Swiss-Prot Q9T145)和野生型gh-1RNA聚合酶(NCBI基因组序列NC_004665；Gene 1258850；UniProtKB/Swiss-Prot蛋白Q859H5)、野生型SP6 RNA聚合酶(NCBI基因组序列NC_004831；Gene 1481778；UniProtKB/Swiss-Prot蛋白Q7Y5R1)以及能够沿5’→3’方向合成与双链模板DNA在序列上互补的单链RNA的其突变体或衍生物，更特别是野生型T7 RNA聚合酶、野生型T3 RNA聚合酶、野生型SP6 RNA聚合酶、野生型K1-5 RNA聚合酶以及野生型K1E RNA聚合酶以及能够沿5’→3’方向合成与双链模板DNA在序列上互补的单链RNA的其突变体或衍生物。
噬菌体RNA聚合酶的原型，即噬菌体T7 RNA聚合酶特别具有以下优点：在体外，该酶具有极强的持续性，并在37℃下沿5’→3’方向每秒延伸240-250个核苷酸(Golomb和Chamberlin 1974,Lyakhov,He等人1997,Zhang和Studier1997,Finn,MacLachlan等人2005)。此外，当在真核细胞中表达时，噬菌体T7RNA聚合酶主要保留在细胞质中(Elroy-Stein和Moss 1990,Gao和Huang 1993,Brisson,He等人1999,Jais,Decroly等人2018)，并因此通过避免大的DNA分子(即转基因)从真核细胞细胞质向细胞核的主动转移以及RNA分子从细胞核向细胞质的输出而优化了转基因表达的水平。
DNA依赖性RNA聚合酶的催化结构域可以是K1E或K1-5 RNA聚合酶的野生型的催化结构域，也可以是能够沿5’→3’方向合成与双链模板DNA在序列上互补的单链RNA的、K1E或K1-5 RNA聚合酶的突变体(甚至还具有持续性)的催化结构域。例如，所述突变体可以选自由以下组成的组：K1E RNA聚合酶突变体R551S(Jais,Decroly等人2018)、F644A、Q649S、G645A、R627S、I810S、D812E(Makarova,Makarov等人1995)和K631M(Osumi-Davis,deAguilera等人1992,Osumi-Davis,Sreerama等人1994)，特别是R551S(Jais,Decroly等人2018)。
优选地，根据本发明的嵌合酶的所述DNA依赖性RNA聚合酶催化结构域来自与宿主细胞不同的酶，以防止内源基因转录和所述DNA序列转录之间的竞争。
根据本发明的嵌合酶至少包含所述催化结构域，但是可以进一步包含含有所述催化结构域的酶的全部或部分。实际上，根据本发明的嵌合酶的一个实施方案，所述RNA三磷酸酶催化结构域、所述鸟苷酰基转移酶催化结构域和所述N7-鸟嘌呤甲基转移酶催化结构域可以包含在加帽酶，优选单体型加帽酶的全部或部分中。所述poly(A)聚合酶催化结构域也可以包含在poly(A)聚合酶的全部或部分中。所述DNA依赖性RNA聚合酶催化结构域也可以包含在DNA依赖性RNA聚合酶，优选单体型DNA依赖性RNA聚合酶的全部或部分中。
出乎意料的是，发明人证明了本发明的嵌合酶能够在细胞中产生可翻译的RNA，从而产生其被宿主细胞核糖体机器识别和使用所需的关键修饰。
如本文所用，术语“连接”和“结合”涵盖共价和非共价连接。
所述RNA结合结构域可以通过共价(直接地或通过连接肽间接地)连接与本发明的嵌合酶的一个催化结构域连接，特别是选自由以下组成的组：
-所述加帽酶催化结构域，特别是所述RNA三磷酸酶催化结构域、所述鸟苷酰基转移酶催化结构域、所述N7-鸟嘌呤甲基转移酶催化结构域；
-所述poly(A)聚合酶催化结构域，
-增强本发明嵌合酶的至少一个催化结构域的活性，优选增强选自由以下组成的组的至少一个催化结构域的活性的结构域：RNA三磷酸酶催化结构域、鸟苷酰基转移酶催化结构域、N7-鸟嘌呤甲基转移酶催化结构域，特别是N7-鸟嘌呤甲基转移酶催化结构域；和
-除了cap-0、cap-1和cap-2加帽酶之外的5’末端RNA加工酶的催化结构域。
连接肽具有产生融合蛋白的优势，在融合蛋白中，空间位阻最小化，并且为使融合蛋白的成分保留在其天然构象提供了足够的空间。
本发明的所述连接肽可以选自由以下组成的组：
-式(GlymSerp)n的肽，其中：
·m代表0至12，特别是1至8，更特别是3至6，甚至更特别是4的整数；
·p代表0至6，特别是0至5，更特别是0至3，更特别是1的整数；并且
·n代表0至30，特别是0至12，更特别是0至8，甚至更特别是1至6(包括端值)的整数；
特别是式(GlymSerp)n的肽，其中：
·m代表4；
·p代表0或1；并且
·n代表1、2或4；
更特别是式Gly4、(Gly4Ser)1(Gly4Ser)2和(Gly4Ser)4的肽。
式(GlymSerp)n的柔性接头肽的优点是：甘氨酸残基赋予肽柔性，而丝氨酸提供一定的溶解性(Huston,Levinson等人1988)。此外，在(GlymSerp)n接头序列中不存在胰凝乳蛋白酶I、因子Xa、木瓜蛋白酶、纤溶酶、凝血酶和胰蛋白酶的敏感位点，这可以增加所得融合蛋白的整体稳定性。
长度可变的(GlymSerp)n接头通常用于工程化单链Fv片段(sFv)抗体(Huston,Levinson等人1988)。此外，(GlymSerp)n接头已用于生产多种融合蛋白，这些融合蛋白经常保留其各个组分的生物活性(Newton,Xue等人1996,Lieschke,Rao等人1997,Shao,Zhang等人2000,Hu,Li等人2004)。
还可以考虑其他类型的肽接头以产生根据本发明的嵌合酶，例如GGGGIAPSMVGGGGS(SEQ ID N°48)(Turner,Ritter等人1997)、SPNGASNSGSAPDTSSAPGSQ(SEQID N°49)(Hennecke,Krebber等人1998)、EGKSSGSGSESKSTE(SEQ ID N°50)(Bird,Hardman等人1988)、EGKSSGSGSESKEF(SEQ ID N°51)(Newton,Xue等人1996)、GGGSGGGSGGGTGGGSGGG(SEQ ID N°52)(Robinson和Sauer 1998)、GSTSGSGKSSEGKG(SEQ ID N°53)(Bedzyk,Weidner等人1990)、YPRSIYIRRRHPSPSLTT(SEQ ID N°54)(Tang,Jiang等人1996)、GSTSGSGKPGSGEGS(SEQ ID N°55)(Ting,Kain等人2001)、SSADDAKKDAAKKDDAKKDDAKKDA(SEQID N°56)(Pantoliano,Bird等人1991)、GSADDAXXDAAXKDDAKKDDAKKDGS(SEQ ID N°57)(Gregoire,Lin等人1996)、LSADDAKKDAAKKDDAKKDDAKKDL(SEQ ID N°58)(Pavlinkova,Beresford等人1999)、AEAAAKEAAAKEAAAKA(SEQ ID N°59)(Wickham,Carrion等人1995)、GSTSGSGKPGSGEGSTGAGGAGSTSGSGKPSGEG(SEQ ID N°60)(Ting,Kain等人2001)、LSLEVAEEIARLEAEV(SEQ ID N°61)(Ting,Kain等人2001)、GTPTPTPTPTGEF(SEQ ID N°62)(Gustavsson,Lehtio等人2001)、GSTSGSGKPGSGEGSTKG(SEQ ID N°63)(Whitlow,Bell等人1993)和GSHSGSGKP(SEQ ID N°64)(Ting,Kain等人2001)，如先前在专利申请WO 2011/128444中描述的。
所述蛋白质-RNA拴系系统RNA结合结构域和所述催化结构域也可以通过特定的蛋白质元件(例如亮氨酸拉链)组装。
所述RNA结合结构域的C-末端或N-末端可以分别与本发明的嵌合酶的所述催化结构域之一的N-末端或C-末端连接。
优选地，所述RNA结合结构域的C-末端通过共价(直接地或通过连接肽间接地，所述连接肽优选式Gly4、(Gly4Ser)1(Gly4Ser)2或(Gly4Ser)4，甚至更优选式(Gly4Ser)1或Gly4的肽)连接与选自由以下组成的组的催化结构域之一的N-末端连接：
-所述加帽酶催化结构域，特别是所述RNA三磷酸酶催化结构域、所述鸟苷酰基转移酶催化结构域、所述N7-鸟嘌呤甲基转移酶催化结构域；
-所述poly(A)聚合酶催化结构域，
-增强本发明嵌合酶的至少一个催化结构域的活性，优选增强选自由以下组成的组的至少一个催化结构域的活性的结构域：RNA三磷酸酶催化结构域、鸟苷酰基转移酶催化结构域、N7-鸟嘌呤甲基转移酶催化结构域，特别是N7-鸟嘌呤甲基转移酶催化结构域；和
-除了cap-0、cap-1和cap-2加帽酶之外的5’末端RNA加工酶的催化结构域。在一个实施方案中，所述RNA结合结构域的C-末端通过共价(直接地或通过连接肽间接地，所述连接肽优选式Gly4、(Gly4Ser)1(Gly4Ser)2或(Gly4Ser)4，甚至更优选式(Gly4Ser)1或Gly4的肽)连接与选自由以下组成的组的催化结构域之一的N-末端连接：
-所述加帽酶催化结构域，特别是所述RNA三磷酸酶催化结构域、所述鸟苷酰基转移酶催化结构域、所述N7-鸟嘌呤甲基转移酶催化结构域；和
-所述poly(A)聚合酶催化结构域。
在一个实施方案中，本发明的嵌合酶是融合蛋白。
如本文所用，术语“融合蛋白”是指通过将本来编码为不同的蛋白质的两个或更多个蛋白质或蛋白质结构域结合而产生的人工蛋白质。该融合基因的翻译会产生具有来自每个原始蛋白质的功能特性的单个或多个多肽。
在一个实施方案中，本发明的嵌合酶是融合蛋白，其中：
-所述RNA结合结构域的C-末端通过共价连接与所述poly(A)聚合酶催化结构域的N-末端连接，
-所述poly(A)聚合酶催化结构域的C-末端通过共价连接，特别是通过连接肽，与加帽酶催化结构域之一的N-末端连接；并且
-所述DNA依赖性RNA聚合酶催化结构域的N-末端通过共价连接，特别是通过连接肽，与加帽酶催化结构域之一的C-末端连接。
在一个实施方案中，本发明的嵌合酶是融合蛋白，其中：
-所述RNA结合结构域的C-末端通过共价连接与所述poly(A)聚合酶催化结构域的N-末端连接，
-所述poly(A)聚合酶催化结构域的C-末端通过共价连接与选自由以下组成的组的催化结构域之一的N-末端连接：
ο所述RNA三磷酸酶催化结构域，
ο所述鸟苷酰基转移酶催化结构域，和
ο所述N7-鸟嘌呤甲基转移酶催化结构域，
特别是所述RNA三磷酸酶催化结构域的N-末端；并且
-所述DNA依赖性RNA聚合酶催化结构域的N-末端通过共价连接，特别
是通过连接肽，连接至选自由以下组成的组的催化结构域之一的C-末端：
ο所述RNA三磷酸酶催化结构域，
ο所述鸟苷酰基转移酶催化结构域，和
ο所述N7-鸟嘌呤甲基转移酶催化结构域
特别是所述N7-鸟嘌呤甲基转移酶催化结构域的C-末端。
在根据本发明的嵌合酶的一个实施方案中，至少两个，特别是至少三个、至少四个，更特别是全部催化结构域可以直接或通过连接肽(特别是通过Gly4、(Gly4Ser)1、(Gly4Ser)2或(Gly4Ser)4，更特别是式(Gly4Ser)2的连接肽)间接地组装、融合或结合。
特别地，选自由以下组成的组的至少两个，特别是至少三个，更特别是全部催化结构域：
-加帽酶催化结构域，特别是其选自由以下组成的组：RNA三磷酸酶催化结构域、鸟苷酰基转移酶催化结构域和N7-鸟嘌呤甲基转移酶催化结构域，
-DNA依赖性RNA聚合酶催化结构域，和
-poly(A)聚合酶催化结构域；
特别是选自由以下组成的组的至少两个，特别是至少三个，更特别是全部催化结构域：
-RNA三磷酸酶催化结构域，
-鸟苷酰基转移酶催化结构域，
-N7-鸟嘌呤甲基转移酶催化结构域，和
-DNA依赖性RNA聚合酶催化结构域
直接或通过连接肽结合，所述连接肽特别选自由以下组成的组：式Gly4、(Gly4Ser)1、(Gly4Ser)2和(Gly4Ser)4的连接肽，更特别是式(Gly4Ser)2的连接肽。
优选地，至少所述DNA依赖性RNA聚合酶催化结构域通过连接肽(特别是选自由以下组成的组：式Gly4、(Gly4Ser)1、(Gly4Ser)2和(Gly4Ser)4的连接肽，更特别是式(Gly4Ser)2的连接肽)与特别是选自由以下组成的组的至少一个加帽酶催化结构域结合：
-所述RNA三磷酸酶催化结构域；
-所述鸟苷酰基转移酶催化结构域；和
-所述N7-鸟嘌呤甲基转移酶催化结构域；
更特别是所述N7-鸟嘌呤甲基转移酶催化结构域。
特别地，连接肽可以位于所述DNA依赖性RNA聚合酶催化结构域，特别是选自由T7、T3、SP6、K1-5和K1E RNA聚合酶组成的组的噬菌体DNA依赖性RNA聚合酶的催化结构域的N-末端，以及位于所述RNA三磷酸酶催化结构域、所述鸟苷酰基转移酶催化结构域和所述N7-鸟嘌呤甲基转移酶催化结构域的C-末端。
特别地，所述DNA依赖性RNA聚合酶催化结构域，特别是选自由T7、T3、SP6、K1-5和K1E RNA聚合酶组成的组的噬菌体DNA依赖性RNA聚合酶的催化结构域的N-末端通过共价连接，特别是通过连接肽，与选自由以下组成的组的催化结构域之一的C-末端连接：
-所述RNA三磷酸酶催化结构域，
-所述鸟苷酰基转移酶催化结构域，和
-所述N7-鸟嘌呤甲基转移酶催化结构域；
特别是所述N7-鸟嘌呤甲基转移酶催化结构域。
优选地，至少所述poly(A)聚合酶催化结构域通过连接肽与特别是选自由以下组成的组的至少一个加帽酶催化结构域结合：
-所述RNA三磷酸酶催化结构域；
-所述鸟苷酰基转移酶催化结构域；和
-所述N7-鸟嘌呤甲基转移酶催化结构域；
更特别是所述RNA三磷酸酶催化结构域。
特别地，连接肽可以位于所述poly(A)聚合酶催化结构域的C-末端，以及位于所述加帽酶催化结构域，特别是所述RNA三磷酸酶催化结构域、所述鸟苷酰基转移酶催化结构域和所述N7-鸟嘌呤甲基转移酶催化结构域，更特别是所述RNA三磷酸酶催化结构域的N-末端。
在本发明的嵌合酶中，所述加帽酶(特别是RNA三磷酸酶、鸟苷酰基转移酶、N7-鸟嘌呤甲基转移酶)、DNA依赖性RNA聚合酶和poly(A)聚合酶的催化结构域也可以通过特定的蛋白质元件(例如亮氨酸拉链，例如生物素化结构域)组装到催化结构域之一(例如Avi-tagII(Cronan 1990)或PFB-tag(Wu,Yeung等人2002))，以及通过生物素结合结构域组装到其他催化结构域之一(例如，Strep-tag II(Schmidt和Skerra 1993)或Nano-tag(Lamla和Erdmann 2004))。
在根据本发明的嵌合酶的一个实施方案中，至少两个所述催化结构域可以通过非共价连接，特别是通过亮氨酸拉链组装。
优选地，至少所述DNA依赖性RNA聚合酶或poly(A)聚合酶的催化结构域通过非共价连接，特别是通过亮氨酸拉链，与至少一个加帽酶催化结构域，优选选自由以下组成的组的至少一个催化结构域组装：
-所述RNA三磷酸酶催化结构域；
-所述鸟苷酰基转移酶催化结构域；和
-所述N7-鸟嘌呤甲基转移酶催化结构域。
在一个实施方案中，至少所述poly(A)聚合酶催化结构域通过非共价连接，特别是通过亮氨酸拉链，优选在其C-末端，与至少一个加帽酶催化结构域，优选选自由以下组成的组的至少一个催化结构域组装：
-所述RNA三磷酸酶催化结构域；
-所述鸟苷酰基转移酶催化结构域；和
-所述N7-鸟嘌呤甲基转移酶催化结构域；
并且更特别是与所述RNA三磷酸酶催化结构域组装。
亮氨酸拉链是由两个彼此相互作用的两亲性α-螺旋组成的二聚体卷曲螺旋蛋白结构，通常用于蛋白质的同源或异源二聚化(O'Shea,Klemm等人1991)。每个螺旋均由七个氨基酸的重复组成，其中第一个氨基酸(残基a)是疏水的，第四个氨基酸(残基d)通常是亮氨酸，而其他残基是极性的。亮氨酸拉链VELCRO ACID-p1和BASE-p1形成平行的异二聚体双链卷曲螺旋结构，极易形成平行的蛋白质异二聚体(O'Shea,Lumb等人1993)。它们已用于膜蛋白(Chang,Bao等人1994,Pashine,Busch等人2003)，以及几种可溶性蛋白(Busch,Reich等人1998,Busch,Pashine等人2002)的异源二聚化。
还可以考虑其他类型的寡聚肽结构域来产生根据本发明的嵌合酶，以组装根据本发明的嵌合酶的至少两个所述催化结构域，特别是形成反向平行异聚结构的亮氨酸拉链，例如ACID-a1/BASE-a1(Oakley和Kim 1998)、ACID-Kg/BASE-Eg(McClain,Woods等人2001)、NZ/CZ(Ghosh,Hamilton等人2000)、ACID-pLL/BASE-pLL(Lumb和Kim 1995)和EE1234L和RR1234L(Moll,Ruvinov等人2001)亮氨酸拉链。二硫键连接形式的亮氨酸拉链也可以用于生产根据本发明的二硫化卷曲螺旋结合的异二聚体嵌合酶(O'Shea,Lumb等人1993)，以及在氧化条件下半胱氨酸残基之间的链间二硫键(Wells和Powers 1986)。
因此，至少两个所述poly(A)聚合酶、加帽酶(特别是RNA三磷酸酶、鸟苷酰基转移酶、N7-鸟嘌呤甲基转移酶)和DNA依赖性RNA聚合酶催化结构域可以通过亮氨酸拉链，特别是形成反向平行异聚结构的亮氨酸拉链，例如ACID-a1/BASE-a1(Oakley和Kim 1998)、ACID-Kg/BASE-Eg(McClain,Woods等人2001)、NZ/CZ(Ghosh,Hamilton等人2000)和ACID-pLL/BASE-pLL亮氨酸拉链，二硫化卷曲螺旋结合(O'Shea,Lumb等人1993)，以及半胱氨酸残基之间的二硫键(Wells和Powers 1986)组装。
在一个实施方案中，根据本发明的嵌合酶包含：
-野生型λN抗终止蛋白的RNA结合结构域，其与以下融合
-哺乳动物PAPOLB、痘苗病毒VP55、非洲猪瘟病毒C475L、棘阿米巴多噬模拟病毒R341、Megavirus chilensis MG561、酿酒酵母、白色念珠菌、卡氏肺孢子虫、突变PAPOLA的野生型poly(A)聚合酶，或能够催化腺苷残基从ATP非模板化添加到RNA分子的3’末端的其突变体或衍生物；或与以下融合
-野生型酿酒酵母PAP1 poly(A)聚合酶、酿酒酵母PAP1 poly(A)聚合酶的细胞质突变体(其中核定位信号是无功能的或缺失的)或能够催化腺苷残基从ATP非模板化添加到RNA分子的3’末端的其突变体或衍生物；或与以下融合
-野生型粟酒裂殖酵母PLA1 poly(A)聚合酶、粟酒裂殖酵母PLA1 poly(A)聚合酶的细胞质突变体(其中核定位信号是无功能的或缺失的)或能够催化腺苷残基从ATP非模板化添加到RNA分子的3’末端的其突变体或衍生物；
-野生型哺乳动物PAPOLA、哺乳动物PAPOLA poly(A)聚合酶的细胞质突变体(其中核定位信号是无功能的或缺失的)或能够催化腺苷残基从ATP非模板化添加到RNA分子的3’末端的其突变体或衍生物并与以下融合，特别是与以下的氨基末端融合
-野生型NP868R非洲猪瘟病毒mRNA加帽酶或能够在RNA分子的5’-末端添加m7GpppN帽的其突变体或衍生物，特别地，野生型NP868R非洲猪瘟病毒加帽酶与以下融合，特别是与以下的氨基末端融合
-野生型T7、T3、SP6、K1-5、K1E RNA聚合酶或能够沿5’→3’方向合成与双链模板DNA在序列上互补的单链RNA的其突变体或衍生物(包括R551S K1E RNA聚合酶突变体)的氨基末端，
特别是通过接头，接头优选选自由以下组成的组：式Gly4、(Gly4Ser)1、(Gly4Ser)2和(Gly4Ser)4的连接肽，更优选式Gly4、(Gly4Ser)1或(Gly4Ser)2的连接肽。
在另一个实施方案中，根据本发明的嵌合酶包含：
-野生型λN抗终止蛋白的RNA结合结构域，其与以下融合
-野生型R341病毒poly(A)聚合酶或能够催化腺苷残基从ATP非模板化添加到RNA分子的3’末端的其突变体或衍生物；或
-野生型酿酒酵母PAP1 poly(A)聚合酶、或酿酒酵母PAP1 poly(A)聚合酶的细胞质突变体(其中核定位信号是无功能的)或能够催化腺苷残基从ATP非模板化添加到RNA分子的3’末端的其突变体或衍生物；或
-野生型粟酒裂殖酵母PLA1 poly(A)聚合酶、粟酒裂殖酵母PLA1 poly(A)聚合酶的细胞质突变体(其中核定位信号是无功能的或缺失的)或能够催化腺苷残基从ATP非模板化添加到RNA分子的3’末端的其突变体或衍生物；
并与以下融合，特别是与以下的氨基末端融合
-野生型NP868R非洲猪瘟病毒mRNA加帽酶或能够在RNA分子的5’-末端添加m7GpppN帽的其突变体或衍生物，特别地，野生型NP868R非洲猪瘟病毒加帽酶与以下融合，特别是与以下的氨基末端融合
-野生型K1E RNA聚合酶或能够沿5’→3’方向合成与双链模板DNA在序列上互补的单链RNA的其突变体或衍生物(包括R551S K1E RNA聚合酶突变体)的氨基末端
特别是通过接头，接头优选选自由以下组成的组：式Gly4、(Gly4Ser)1、(Gly4Ser)2和(Gly4Ser)4的连接肽，更优选式Gly4或(Gly4Ser)2的连接肽。
本发明还涉及一种分离的核酸分子或一组分离的核酸分子，所述核酸分子编码根据本发明的嵌合酶，或者分离的核酸分子编码嵌合酶，其特征在于其序列包含编码蛋白质-RNA拴系系统的RNA结合结构域的核酸序列，其与以下框内融合、特别是按以下顺序融合：
-编码至少一个poly(A)聚合酶催化结构域的核酸序列；
-编码至少一个加帽酶催化结构域的核酸序列；并任选地与以下核酸序列按顺序、框内融合
-编码至少一个DNA依赖性RNA聚合酶(特别是噬菌体DNA依赖性RNA聚合酶)催化结构域的核酸序列；
其中所述RNA结合结构域与所述蛋白质-RNA拴系系统的RNA元件特异性结合，所述RNA元件由特定RNA序列和/或结构组成。
编码根据本发明的嵌合酶的所述分离的核酸分子的组包含或由以下组成：对于通过其表达来获得根据本发明的嵌合酶而言必需且充分的所有核酸分子。
如本文所用，术语“核酸分子”是由连接的核苷酸组成的任何分子，包括DNA和RNA分子。
在一个实施方案中，编码根据本发明的嵌合酶的所述分离的核酸分子的组包含或由以下组成：
-编码所述蛋白质-RNA拴系系统的RNA结合结构域的核酸分子，和
-编码至少一个加帽酶催化结构域，特别是至少一个RNA三磷酸酶催化结构域、至少一个鸟苷酰基转移酶催化结构域以及至少一个N7-鸟嘌呤甲基转移酶催化结构域的核酸分子；
以及任选的：
-编码至少一个poly(A)聚合酶催化结构域的核酸分子；和/或
-编码至少一个DNA依赖性RNA聚合酶，(特别是噬菌体DNA依赖性RNA聚合酶)催化结构域的核酸分子。
在另一个实施方案中，编码根据本发明的嵌合酶的所述分离的核酸分子的组包含或由以下组成：
-编码所述蛋白质-RNA拴系系统的RNA结合结构域的核酸分子，
-编码至少一个RNA三磷酸酶催化结构域的核酸分子，
-编码至少一个鸟苷酰基转移酶催化结构域的核酸分子，
-编码至少一个N7-鸟嘌呤甲基转移酶催化结构域的核酸分子；
以及任选的：
-编码至少一个poly(A)聚合酶催化结构域的核酸分子；和/或
-编码至少一个DNA依赖性RNA聚合酶催化结构域，特别是至少一个噬菌体DNA依赖性RNA聚合酶催化结构域的核酸分子。
在一个实施方案中，本发明的分离的核酸分子包含或由以下组成：编码所述蛋白质-RNA拴系系统的RNA结合结构域的核酸序列，其与以下核酸序列框内融合(特别是按顺序)：
-编码所述poly(A)聚合酶催化结构域的核酸序列，
-编码所述加帽酶催化结构域的核酸序列，特别是
ο编码所述RNA三磷酸酶催化结构域的核酸序列，
ο编码所述鸟苷酰基转移酶催化结构域的核酸序列，
ο编码所述N7-鸟嘌呤甲基转移酶催化结构域的核酸序列，
并与以下核酸序列框内融合(特别是按顺序)
-编码所述DNA依赖性RNA聚合酶催化结构域，特别是噬菌体DNA依赖性RNA聚合酶催化结构域的核酸序列。
这样的单个核酸序列具有促进亚基组装的优点，因为仅存在单个开放阅读框。
在本发明的分离的核酸分子的一个实施方案中，其序列包含至少一个编码核糖体跳跃基序(ribosome skipping motif)的核酸序列。
如本文所用，术语“核糖体跳跃基序”是指翻译的可选机制，其中特定的病毒肽阻止核糖体共价连接新插入的氨基酸，并使其继续翻译。这导致多蛋白的明显的共翻译切割。
特别地，所述核糖体跳跃基序选自由以下组成的组：来自以下病毒的2A序列：口蹄疫病毒口疮病毒(Aphtovirus)(UniProtKB/Swiss-Prot AAT01756)、Avisivirua A(UniProtKB/Swiss-Prot M4PJD6)、鸭甲型肝炎禽肝病毒(Duck hepatitis AAvihepatovirus)(UniProtKB/Swiss-Prot Q0ZQM1)、脑炎心肌炎心病毒(Encephalomyocarditis Cardiovirus)(UniProtKB/Swiss-Prot Q66765)、Cosavirus A(UniProtKB/Swiss-Prot B8XTP8)、马乙型鼻炎病毒1型(Equine rhinitis B Erbovirus1)(UniProtKB/Swiss-Prot Q66776)、猪塞内加谷病毒(Seneca Valley Erbovirus)(UniProtKB/Swiss-Prot Q155Z9)、Hunnivirus A(UniProtKB/Swiss-Prot F4YYF3)、Kunsagivirus A(UniProtKB/Swiss-Prot S4VD62)、Mischivirus A(UniProtKB/Swiss-Prot I3VR62)、Mosavirus A2(UniProtKB/Swiss-Prot X2L6K2)、Pasivirus A1(UniProtKB/Swiss-Prot I6YQK4)、1型猪捷申病毒(porcine teschovirus 1)UniProtKB/Swiss-Prot Q9WJ28)、传染性软化病病毒(Infectious flacherie Iflavirus)(UniProtKB/Swiss-Prot O70710)、明脉扁刺蛾β四体病毒(Thosea asignaBetatetravirus)(UniProtKB/Swiss-Prot Q9YK87)、蟋蟀麻痹病毒(Cricket paralysisCripavirus)(UniProtKB/Swiss-Prot Q9IJX4)、C型人类轮状病毒(Human rotavirus C)(UniProtKB/Swiss-Prot Q9PY95)和1型舞毒蛾质型多角体病毒(Lymantria disparcypovirus 1)(UniProtKB/Swiss-Prot Q91ID7)。
特别地，所述编码核糖体跳跃基序的核酸序列选自由以下组成的组：来自口蹄疫病毒口疮病毒的2A序列(也称为“F2A”，UniProtKB/Swiss-Prot AAT01756)或来自1型猪捷申病毒的2A序列(也称为“T2A”，UniProtKB/Swiss-Prot Q9WJ28)。
所述编码核糖体跳跃基序的核酸序列可以位于编码本发明的嵌合酶的所述催化结构域的任何序列之间。
在一个实施方案中，所述编码核糖体跳跃基序的核酸序列可以位于编码所述poly(A)聚合酶催化结构域的序列之间，编码所述poly(A)聚合酶催化结构域的序列与编码以下的序列框内融合：
-所述加帽酶催化结构域，优选选自由所述RNA三磷酸酶催化结构域、所述鸟苷酰基转移酶催化结构域和所述N7-鸟嘌呤甲基转移酶催化结构域组成的组，或
-所述DNA依赖性RNA聚合酶催化结构域，特别是噬菌体DNA依赖性RNA聚合酶催化结构域；
特别地，与编码所述加帽酶催化结构域，更特别是编码所述RNA三磷酸酶催化结构域的序列框内融合。
在另一个实施方案中，所述编码核糖体跳跃基序的核酸序列可以位于编码所述DNA依赖性RNA聚合酶催化结构域的序列之间，编码所述DNA依赖性RNA聚合酶催化结构域的序列与编码所述加帽酶催化结构域(特别是选自由所述RNA三磷酸酶催化结构域、所述鸟苷酰基转移酶催化结构域和所述N7鸟嘌呤甲基转移酶催化结构域组成的组)的序列框内融合，更特别地与编码所述N7-鸟嘌呤甲基转移酶催化结构域的序列框内融合。
在一个实施方案中，根据本发明的分离的核酸分子的特征在于其序列包含或由以下组成：
-编码蛋白质-RNA拴系系统的RNA结合结构域的核酸序列，其与以下核酸
序列框内融合(特别是按顺序)：
ο编码poly(A)聚合酶催化结构域的核酸序列；
ο编码加帽酶催化结构域的核酸序列，并任选地与以下核酸序列框内融合(特别是按顺序)
ο编码所述DNA依赖性RNA聚合酶催化结构域，特别是噬菌体DNA依赖性RNA聚合酶催化结构域的核酸序列，
和
-编码核糖体跳跃基序的核酸序列，其位于所述编码poly(A)聚合酶催化结构域的核酸序列与所述编码选自由以下组成的组的催化结构域之一的核酸序列之间：
ο所述加帽酶催化结构域，和
ο所述DNA依赖性RNA聚合酶催化结构域。
在一个实施方案中，根据本发明的分离的核酸分子的特征在于其序列包含或由以下组成：
-编码蛋白质-RNA拴系系统的RNA结合结构域的核酸序列，其与以下核酸
序列框内融合(特别是按顺序)：
ο编码poly(A)聚合酶催化结构域的核酸序列；
ο编码RNA三磷酸酶催化结构域的核酸序列，
ο编码鸟苷酰基转移酶催化结构域的核酸序列，
ο编码N7-鸟嘌呤甲基转移酶催化结构域的核酸序列，并任选地与以下核酸序列框内融合(特别是按顺序)
ο编码所述DNA依赖性RNA聚合酶催化结构域，特别是噬菌体DNA依赖性RNA聚合酶催化结构域的核酸序列，
和
-编码核糖体跳跃基序的核酸序列，其位于所述编码poly(A)聚合酶催化结构域的核酸序列与所述编码选自由以下组成的组的催化结构域之一的核酸序列之间：
ο所述RNA三磷酸酶催化结构域，
ο所述鸟苷酰基转移酶催化结构域，
ο所述N7-鸟嘌呤甲基转移酶催化结构域，和
ο所述DNA依赖性RNA聚合酶催化结构域，
优选所述RNA三磷酸酶催化结构域。
在一个实施方案中，编码嵌合酶的本发明的分离的核酸分子的特征在于，其序列包含：编码蛋白质-RNA拴系系统RNA结合结构域的核酸序列，其与以下核酸序列按顺序、框内融合：
-编码至少一个poly(A)聚合酶催化结构域的核酸序列；
-编码至少一个加帽酶催化结构域的核酸序列；并任选地与以下核酸序列按顺序、框内融合
-编码所述DNA依赖性RNA聚合酶催化结构域，特别是噬菌体DNA依赖性RNA聚合酶催化结构域的核酸序列；
并且其特征在于，其序列进一步包含编码核糖体跳跃基序的核酸序列，所述编码核糖体跳跃基序的核酸序列位于所述编码poly(A)聚合酶催化结构域的核酸序列与所述编码至少一个加帽酶催化结构域的核酸序列之间。
实际上，出乎意料的是，发明人已经证明(如实施例8所示)，这种核酸分子使得DNA依赖性RNA聚合酶的表达率高于以下组合：编码嵌合酶的核酸分子和DNA依赖性RNA聚合酶的组合，所述嵌合酶包含至少一个加帽酶结构域，所述DNA依赖性RNA聚合酶与编码蛋白质-RNA拴系系统RNA结合结构域的核酸分子相关联，所述编码蛋白质-RNA拴系系统RNA结合结构域的核酸分子与编码至少一个poly(A)聚合酶催化结构域的核酸序列框内融合。
这些结果确实令人惊讶，并且由于成分相同，本领域技术人员可能预期获得相同的表达率。
特别地，根据本发明的核酸分子可以与选自由以下组成的组的至少一个，优选全部启动子可操作地连接：
-真核DNA依赖性RNA聚合酶的启动子，优选RNA聚合酶II的启动子；
-噬菌体DNA依赖性RNA聚合酶的启动子；和
-本发明嵌合酶的所述DNA依赖性RNA聚合酶催化结构域的启动子。
核酸与真核DNA依赖性RNA聚合酶(优选RNA聚合酶II)的启动子的连接具有显著的优点，当本发明的嵌合酶在真核宿主细胞中表达时，嵌合酶的表达由真核RNA聚合酶(优选RNA聚合酶II)驱动。这些嵌合酶转而可以启动转基因的转录。如果使用组织特异性RNA聚合酶II启动子，那么本发明的嵌合酶可以在靶组织/细胞中选择性表达。
所述启动子可以是本领域技术人员众所周知的组成型启动子或诱导型启动子。启动子可以是发育调控的、可诱导的或组织特异性的。
本发明还涉及包含根据本发明的核酸分子的载体。所述载体可适合于半稳定或稳定表达。
本发明还涉及一组载体，其包含根据本发明的一组分离的核酸分子。
特别地，根据本发明的所述载体是克隆载体或表达载体。
本发明还涉及一种宿主细胞，其包含根据本发明的核酸分子或根据本发明的载体或根据本发明的一组载体。
根据本发明的宿主细胞可用于大规模蛋白质生产。
优选地，根据本发明的嵌合酶的所述DNA依赖性RNA聚合酶催化结构域来自与宿主细胞不同的酶，以防止内源基因转录和转基因转录之间的竞争。
本发明还涉及基因工程改造的非人类真核生物，其表达由根据本发明的核酸分子或一组分离的核酸分子编码的嵌合酶，特别是根据本发明的嵌合酶。所述非人类真核生物可以是任何非人类动物、植物。
本发明还涉及本发明的嵌合酶用于生产具有5’-末端帽，特别是5’-末端m7GpppN帽，并优选具有3’poly(A)尾，并任选地包含至少一种化学修饰的RNA分子的用途，特别是体外或离体用途。
本发明还涉及本发明的核酸分子或一组分离的核酸分子用于生产具有5’-末端帽，特别是5’-末端m7GpppN帽，并优选具有3’poly(A)尾，并任选地包含至少一种化学修饰的RNA分子的用途，特别是体外或离体用途。
特别地，所述RNA分子通过噬菌体DNA依赖性RNA聚合酶合成。
实际上，根据本发明的嵌合酶适用于合成具有至少一种化学修饰的加帽的RNA。
特别地，本发明还涉及本发明的嵌合酶用于生产具有5’-末端帽，特别是5’-末端m7GpppN帽，并优选具有3’poly(A)尾，并包含至少一种化学修饰的RNA分子的用途，特别是体外或离体用途，所述化学修饰选自由以下组成的组：吡啶-4-酮核糖核苷、5-氮杂-尿苷、2-硫-5-氮杂-尿苷、2-硫尿苷、4-硫-假尿苷、2-硫-假尿苷、5-羟基尿苷、3-甲基尿苷、5-羧甲基-尿苷、1-羧甲基-假尿苷、5-丙炔基-尿苷、1-丙炔基-假尿苷、5-牛磺酸(taurino)甲基尿苷、1-牛磺酸甲基-假尿苷、5-牛磺酸甲基-2-硫-尿苷、l-牛磺酸甲基-4-硫-尿苷、假尿苷、5-甲基-胞苷、5-甲基-尿苷、1-甲基-假尿苷、4-硫-l-甲基-假尿苷、2-硫-1-甲基-假尿苷、1-甲基-1-去氮杂-假尿苷、2-硫-1-甲基-1-去氮杂-假尿苷、二氢尿苷、二氢假尿苷、2-硫-二氢尿苷、2-硫-二氢假尿苷、2-甲氧基尿苷、2-甲氧基-4-硫-尿苷、4-甲氧基-假尿苷、4-甲氧基-2-硫-假尿苷、5-氮杂-胞苷、假异胞苷、3-甲基-胞苷、N4-乙酰胞苷、5-甲酰基胞苷、N4-甲基胞苷、5-羟甲基胞苷、1-甲基-假异胞苷、吡咯-胞苷、吡咯-假异胞苷、2-硫-胞苷、2-硫-5-甲基-胞苷、4-硫-假异胞苷、4-硫-1-甲基-假异胞苷、4-硫-1-甲基-1-去氮杂-假异胞苷、1-甲基-1-去氮杂-假异胞苷、泽布拉林(zebularine)、5-氮杂-泽布拉林、5-甲基-泽布拉林、5-氮杂-2-硫-泽布拉林、2-硫-泽布拉林、2-甲氧基-胞苷、2-甲氧基-5-甲基-胞苷、4-甲氧基-假异胞苷、4-甲氧基-1-甲基-假异胞苷、2-氨基嘌呤、2、6-二氨基嘌呤、7-去氮杂-腺嘌呤、7-去氮杂-8-氮杂-腺嘌呤、7-去氮杂-2-氨基嘌呤、7-去氮杂-8-氮杂-2-氨基嘌呤、7-去氮杂-2,6-二氨基嘌呤、7-去氮杂-8-氮杂-2,6-二氨基嘌呤、1-甲基腺苷、N6-甲基腺苷、N6-异戊烯基腺苷、N6-(顺式-羟基异戊烯基)腺苷、2-甲硫基-N6-(顺式-羟基异戊烯基)腺苷、N6-甘氨酰基氨甲酰基腺苷、N6-苏氨酰基氨甲酰基腺苷、2-甲硫基-N6-苏氨酰基氨甲酰基腺苷、N6,N6-二甲基腺苷、7-甲基腺嘌呤、2-甲硫基-腺嘌呤、2-甲氧基-腺嘌呤、肌苷、1-甲基-肌苷、怀俄苷(wyosine)、怀丁苷(wybutosine)、7-去氮杂-鸟苷、7-去氮杂-8-氮杂-鸟苷、6-硫-鸟苷、6-硫-7-去氮杂-鸟苷、6-硫-7-去氮杂-8-氮杂-鸟苷、7-甲基-鸟苷、6-硫-7-甲基-鸟苷、7-甲基肌苷、6-甲氧基-鸟苷、1-甲基鸟苷、N2-甲基鸟苷、N2,N2-二甲基鸟苷、8-氧代-鸟苷、7-甲基-8-氧代-鸟苷、l-甲基-6-硫-鸟苷、N2-甲基-6-硫-鸟苷和N2,N2-二甲基-6-硫-鸟苷。
本发明还涉及根据本发明的嵌合酶或根据本发明的分离的核酸分子或一组分离的核酸分子的如下体外或离体用途：用于生产蛋白质，特别是治疗目的的蛋白质如抗体，特别是在真核系统中生产蛋白质，例如在体外合成的蛋白测定或培养的细胞中。
本发明还涉及在宿主细胞中生产由DNA序列编码的RNA分子的体外或离体方法，所述RNA分子具有5’-末端帽，特别是5’-末端m7 GpppN帽并且优选具有3’poly(A)尾，所述方法包括以下步骤：在宿主细胞中表达根据本发明的核酸分子或一组分离的核酸分子，其中所述DNA序列与至少一种编码与所述RNA结合结构域特异性结合的所述蛋白质-RNA拴系系统的RNA元件的序列共价连接。如本文所用，术语“与所述RNA结合结构域特异性结合的蛋白质-RNA拴系系统的RNA元件”是指这样的RNA序列，其通常形成茎环，能够以高亲和力与蛋白质-RNA拴系系统的相应RNA结合结构域结合。
特别地，所述DNA序列与噬菌体DNA依赖性RNA聚合酶的启动子或与本发明嵌合体的所述DNA依赖性RNA聚合酶的启动子可操作地连接。
特别地，当蛋白质-RNA拴系系统的RNA结合结构域是λ形噬菌体N抗终止蛋白质-RNA拴系系统的RNA结合结构域时，与所述RNA结合结构域特异性结合的元件可以是boxBL和/或boxBR茎环RNA结构(Das 1993,Greenblatt,Nodwell等人1993,Friedman和Court1995)，包括由SEQ ID N°7和SEQ ID N°38编码的元件。
特别地，当蛋白质-RNA拴系系统的RNA结合结构域是MS2外壳蛋白质-RNA拴系系统的RNA结合结构域时，与所述RNA结合结构域特异性结合的元件可以是19个核苷酸或21个核苷酸的茎环序列，包括由SEQ ID N°39编码的元件(Peabody 1993,Valegard,Murray等人1994,LeCuyer,Behlen等人1996,Valegard,Murray等人1997)(来自肠噬菌体MS2分离株DL52(NCBI登录号JQ966307.1)的核苷酸1748-766)，其包含病毒复制酶的基因的起始密码子(Valegard,Murray等人1994,Valegard,Murray等人1997)。
特别地，当蛋白质-RNA拴系系统的RNA结合结构域是R17分离株外壳蛋白质-RNA拴系系统的RNA结合结构域时，与所述RNA结合结构域特异性结合的元件可以是19个核苷酸或21个核苷酸的茎环(来自肠噬菌体R17(NCBI登录号为EF108465.1)的核苷酸1746-764)，其包含病毒复制酶基因的起始密码子(Carey和Uhlenbeck 1983)。
特别地，所述DNA序列与噬菌体DNA依赖性RNA聚合酶的启动子或与本发明的嵌合酶的所述DNA依赖性RNA聚合酶的启动子可操作地连接，并在其3’末端与至少一个，优选至少两个，至少三个，更优选至少四个编码与所述RNA结合结构域特异性结合的元件的序列共价连接。
特别地，根据本发明的所述方法进一步包括以下步骤：使所述编码RNA分子的DNA序列与本发明的酶接触。
特别地，所述DNA序列与噬菌体DNA依赖性RNA聚合酶的启动子或与本发明的嵌合酶的所述DNA依赖性RNA聚合酶的启动子可操作地连接，并在其3’末端与至少一种编码与所述RNA结合结构域特异性结合的元件的序列共价连接，所述RNA结合结构域与poly(A)轨道序列共价连接，所述poly(A)轨道序列由至少10个，特别是至少20个、30个，更特别是至少40个脱氧腺苷残基组成。
特别地，所述poly(A)轨道序列可以在其3’末端与自切割RNA序列共价连接，并任选与转录终止序列连接。
特别地，所述自切割RNA序列可以是选自包括以下的组的自切割RNA序列：丁型肝炎病毒的基因组假结样核酶(Genbank登录号AJ000558.1)、丁型肝炎病毒的反基因组假结样核酶(Genbank登录号AJ000558.1)、烟草环斑病毒卫星发夹状核酶(Genbank登录号NC_003889.1)或人造短发夹RNA(shRNA)。
特别地，所述转录终止序列可以是噬菌体T7 phi10转录终止序列(Genbank登录号GU071091.1)或大肠杆菌RNA聚合酶rrnB t1终止(Genbank登录号LN832404.1)。
特别地，根据本发明的所述方法可以进一步包括以下步骤：使用本领域技术人员熟知的方法，例如通过使用磷酸钙转染、通过电穿孔或通过将阳离子脂质与DNA混合以产生脂质体，从而将根据本发明的所述DNA序列和/或核酸引入宿主细胞中。
在一个实施方案中，根据本发明的所述方法还包括以下步骤：抑制所述宿主细胞的细胞转录和转录后机制，特别是使其沉默，优选通过siRNA(小干扰RNA)、miRNA(microRNA)或shRNA。
在一个实施方案中，根据本发明的所述方法进一步包括以下步骤：抑制所述宿主细胞中内源性DNA依赖性RNA聚合酶和/或内源性加帽酶的表达。
如本文所用，术语“内源性DNA依赖性RNA聚合酶”是指所述宿主细胞的内源性DNA依赖性RNA聚合酶。当宿主细胞是真核细胞时，所述内源性DNA依赖性RNA聚合酶是RNA聚合酶II。
如本文所用，术语“内源性加帽酶”是指所述宿主细胞的内源性加帽酶。
如本文所用，术语“抑制蛋白质的表达”是指使所述蛋白质的表达降低至少20％，特别是至少35％，至少50％，更特别是至少65％，至少80％，至少90％。蛋白质表达的抑制可以通过本领域技术人员众所周知的技术来确定，包括但不限于Northern-Blot、Western-Blot、RT-PCR。
抑制所述宿主细胞中内源性DNA依赖性RNA聚合酶和/或内源性加帽酶的表达的步骤可以通过本领域技术人员公知的任何技术来实施，包括但不限于靶向所述内源性DNA依赖性RNA聚合酶和/或内源性加帽酶的siRNA技术、靶向所述内源性DNA依赖性RNA聚合酶和/或内源性加帽酶的反义RNA技术以及靶向所述内源性DNA依赖性RNA聚合酶和/或内源性加帽酶的shRNA技术。
除siRNA(或shRNA)外，出于同一目的，也可以考虑其他抑制性序列，包括DNA或RNA反义序列(Liu和Carmichael 1994,Dias和Stein 2002)、锤头状核酶(Salehi-Ashtiani和Szostak 2001)、发夹状核酶(Lian,De Young等人1999)或嵌合snRNA U1-反义靶向序列(Fortes,Cuevas等人2003)。另外，可以考虑其他细胞靶基因进行抑制，包括参与细胞转录(例如RNA聚合酶II的其他亚基或转录因子)、转录后加工(例如加帽酶的其他亚基，以及聚腺苷酸化或剪接体因子)和mRNA核输出途径的其他基因。
在根据本发明的方法的一个实施方案中，所述RNA分子可以编码治疗目的的多肽。
在另一个实施方案中，所述RNA分子可以是选自包含以下的组的非编码RNA分子：siRNA、核酶、shRNA和反义RNA。特别地，所述DNA序列可以编码选自由以下组成的组的RNA分子：mRNA，非编码RNA，特别是siRNA，核酶，shRNA和反义RNA。
本发明还涉及根据本发明的嵌合酶作为如下酶的用途：加帽酶，并优选pol(A)聚合酶和DNA依赖性RNA聚合酶。
本发明还涉及用于生产具有5'-末端帽，特别是5'-末端m7GpppN帽的RNA分子的试剂盒，其包含至少一种如上文所定义的本发明的嵌合酶，和/或如上文所定义的本发明的分离的核酸分子和/或一组核酸分子，和/或如上文所定义的本发明的载体，或
嵌合酶(特别是细胞质嵌合酶)和/或编码所述嵌合酶的分离的核酸分子和/或一组分离的核酸分子，所述嵌合酶包含至少一个RNA三磷酸酶催化结构域、至少一个鸟苷酰基转移酶催化结构域、至少一个N7-鸟嘌呤甲基转移酶催化结构域和至少一个DNA依赖性RNA聚合酶催化结构域，和poly(A)聚合酶(特别是细胞质poly(A)聚合酶)和/或编码所述poly(A)聚合酶的分离的核酸分子，所述poly(A)聚合酶包含与至少一个所述poly(A)聚合酶催化结构域连接的至少一个蛋白质RNA拴系系统RNA结合结构域；
以及任选的，编码所述RNA分子的DNA序列，其与至少一个编码所述蛋白质-RNA拴系系统的RNA元件(其与所述RNA结合结构域特异性结合)的序列共价连接。
特别地，所述用于生产具有5'-末端帽的RNA分子的试剂盒包括编码所述RNA分子的DNA序列，所述DNA序列与噬菌体DNA依赖性RNA聚合酶的启动子或本发明的嵌合酶的所述DNA依赖性RNA聚合酶的启动子可操作地连接。
特别地，所述用于生产具有5'-末端帽的RNA分子的试剂盒包含至少一种本发明的嵌合酶，和/或本发明的分离的核酸分子和/或一组分离的核酸分子，和/或本发明的载体，以及任选的，编码所述RNA分子的DNA序列，其与至少一个编码与所述RNA结合域特异性结合的元件的序列共价连接，特别是与噬菌体DNA依赖性RNA聚合酶的启动子或与本发明嵌合酶的所述DNA依赖性RNA聚合酶的启动子可操作地连接。
特别地，所述用于生产具有5’末端帽的RNA分子的试剂盒进一步包含分离的核酸分子，所述分离的核酸分子编码至少一个DNA依赖性RNA聚合酶催化结构域，和/或至少一个DNA依赖性RNA聚合酶催化结构域，特别是噬菌体DNA依赖性RNA聚合酶催化结构域。
特别地，所述试剂盒进一步包括其使用说明书。本发明还涉及一种组合物(特别是试剂盒或药物组合物)，其包含：
-嵌合酶(特别是细胞质嵌合酶)和/或编码所述嵌合酶的分离的核酸分子或一组分离的核酸分子，所述嵌合酶包含至少一个加帽酶催化结构域和至少一个DNA依赖性RNA聚合酶(特别是噬菌体DNA依赖性RNA聚合酶)催化结构域；和
-poly(A)聚合酶(特别是细胞质poly(A)聚合酶)和/或编码所述poly(A)聚合酶的分离的核酸分子，所述poly(A)聚合酶包含与至少一个所述poly(A)聚合酶催化结构域连接的至少一个蛋白质-RNA拴系系统(特别是噬菌体蛋白质-RNA拴系系统)RNA结合结构域；和任选的
-DNA序列，其与所述DNA依赖性RNA聚合酶的启动子可操作地连接，并与至少一个编码与所述RNA结合结构域以高亲和力相互作用的元件的序列共价连接；
所述组合物可用于生产具有5'-末端帽，特别是5'-末端m7GpppN帽的RNA分子。
特别地，所述组合物(特别是试剂盒或药物组合物)包含：
-嵌合酶(特别是细胞质嵌合酶)和/或编码所述嵌合酶的分离的核酸分子或一组分离的核酸分子，所述嵌合酶包含至少一个RNA三磷酸酶催化结构域、至少一个鸟苷酰基转移酶催化结构域、至少一个N7-鸟嘌呤甲基转移酶催化结构域和至少一个DNA依赖性RNA聚合酶催化结构域；和
-poly(A)聚合酶(特别是细胞质poly(A)聚合酶)和/或编码所述poly(A)聚合酶的分离的核酸分子，所述poly(A)聚合酶包含与至少一个所述poly(A)聚合酶催化结构域连接的至少一个蛋白质-RNA拴系系统RNA结合结构域；和任选的
-DNA序列，其与所述DNA依赖性RNA聚合酶的启动子可操作地连接，并与至少一个编码与所述RNA结合结构域以高亲和力相互作用的元件的序列共价连接；
所述组合物可用于生产具有5'-末端帽，特别是5'-末端m7GpppN帽的RNA分子。
更特别地，所述组合物(特别是试剂盒或药物组合物)包含：
-嵌合酶(特别是细胞质嵌合酶)和/或编码所述嵌合酶的分离的核酸分子或一组分离的核酸分子，所述嵌合酶包含NP868R加帽酶和K1E DNA依赖性RNA聚合酶，特别是通过(Gly4Ser)2接头连接；和
-poly(A)聚合酶(特别是细胞质poly(A)聚合酶)和/或编码所述poly(A)聚合酶的分离的核酸分子，所述poly(A)聚合酶包含至少一个poly(A)聚合酶(选自由PAP1、PAPOLA、PAPOLB、VP55、C475L、R341和MG561 poly(A)聚合酶组成的组)催化结构域，并包含与至少一个所述poly(A)聚合酶催化结构域连接的至少一个蛋白质-RNA拴系系统RNA结合结构域；和任选的
-DNA序列，其与所述DNA依赖性RNA聚合酶的启动子可操作地连接，并与至少一个编码与所述RNA结合结构域以高亲和力相互作用的元件的序列共价连接；
所述组合物可用于生产具有5'-末端帽，特别是5'-末端m7GpppN帽的RNA分子。
有利地，本发明的试剂盒或组合物可以用作正交基因表达系统。如本文所用，术语“正交”是指其基本结构是独立的并且通常源自不同物种的生物系统。
本发明还涉及根据本发明的嵌合酶、根据本发明的分离的核酸分子、根据本发明的一组核酸分子或根据本发明的载体，其用作药物，特别是用于预防和/或治疗人或动物病状，优选通过基因疗法的手段。
本发明还涉及一种药物组合物，其包含根据本发明的嵌合酶和/或根据本发明的分离的核酸分子和/或根据本发明的一组核酸分子和/或根据本发明的载体。优选地，本发明的所述药物组合物被配制在药学上可接受的载体中。
药学上可接受的载体是本领域技术人员众所周知的。
根据本发明的药物组合物可以进一步包含至少一种目的DNA序列，其中所述DNA序列与所述DNA依赖性RNA聚合酶催化结构域的启动子可操作地连接，并与至少一种编码与所述RNA结合结构域特异性结合的元件的序列共价连接。
这样的组分(特别是选自由以下组成的组：根据本发明的嵌合酶、根据本发明的分离的核酸分子、根据本发明的载体和至少一种目的DNA序列)可以以治疗量(活性且无毒的量)存在于本发明的药物组合物或药物中。
这样的治疗量可以由本领域技术人员通过常规测试来确定，包括评估所述组分的施用对病状和/或病症(试图通过施用根据本发明的所述药物组合物或药物来对其进行预防和/或治疗)的效应。
例如，可以通过分析不同量的上述组分(特别是选自由以下组成的组：根据本发明的嵌合酶、根据本发明的分离的核酸分子、根据本发明的载体和至少一种目的DNA序列)的施用对所述病状和/或所述病症的一组标志物(生物学和/或临床)特征(特别是来自受试者的生物样品)的定量和定性效应来实施这样的测试。
本发明还涉及一种治疗方法，其包括向有此需要的受试者施用治疗量的根据本发明的嵌合酶和/或根据本发明的分离的核酸分子和/或根据本发明的一组核酸分子和/或根据本发明的载体。根据本发明的治疗方法可以进一步包括向有此需要的受试者施用治疗量的至少一种目的DNA序列，其中所述DNA序列与所述DNA依赖性RNA聚合酶催化结构域的启动子可操作地连接，并与至少一种编码与所述RNA结合结构域特异性结合的元件的序列共价连接。
根据本发明的所述嵌合酶、核酸分子和/或所述载体可以与所述目的DNA序列同时、分开或相继施用，特别是在所述目的DNA序列之前施用。
本发明还涉及根据本发明的药物组合物，其用于预防和/或治疗人或动物病状的用途，特别是通过基因疗法的手段。
所述病状可以选自由以下组成的组：可以通过施用所述至少一种目的DNA序列来改善的病状。
本发明还涉及根据本发明的嵌合酶和/或根据本发明的分离的核酸分子和/或根据本发明的一组核酸分子和/或根据本发明的载体用于制备用于预防和/或治疗(特别是通过基因疗法的手段)人或动物病状的药物的用途。
本发明还涉及第一组合产品，其包含作为活性成分的：
-至少一种根据本发明的嵌合酶和/或至少一种根据本发明的核酸分子和/或根据本发明的一组核酸分子和/或至少一种包含和/或表达根据本发明的核酸分子的载体；和
-至少一种目的DNA序列，其中所述DNA序列与所述DNA依赖性RNA聚合酶催化结构域的启动子可操作地连接，并与至少一种编码与所述RNA结合结构域特异性结合的元件的序列共价连接；
其用作药物，其中将所述活性成分配制成单独、同时或相继施用。
本发明还涉及第二组合产品，其包含作为活性成分的：
-嵌合酶(特别是细胞质嵌合酶)和/或编码所述嵌合酶的分离的核酸分子或一组分离的核酸分子，所述嵌合酶包含至少一个RNA三磷酸酶催化结构域、至少一个鸟苷酰基转移酶催化结构域、至少一个N7-鸟嘌呤甲基转移酶催化结构域和至少一个DNA依赖性RNA聚合酶催化结构域；和
-poly(A)聚合酶(特别是细胞质poly(A)聚合酶)和/或编码所述poly(A)聚合酶的分离的核酸分子，所述poly(A)聚合酶包含与至少一个所述poly(A)聚合酶催化结构域连接的至少一个蛋白质-RNA拴系系统RNA结合结构域；和
-DNA序列，其与所述DNA依赖性RNA聚合酶的启动子可操作地连接，并与至少一种编码与所述RNA结合结构域特异性结合的元件的序列共价连接；
其用作药物，其中将所述活性成分配制成单独、同时或相继施用。
所述目的DNA序列可以是抗癌基因(肿瘤抑制基因)。
所述目的DNA序列可以编码治疗目的的多肽或者选自包含siRNA、核酶、shRNA和反义RNA的组的非编码RNA。
所述治疗目的的多肽可以选自：单克隆抗体或其片段、生长因子、细胞因子、细胞或核受体、配体、凝血因子、CFTR蛋白、胰岛素、肌营养不良蛋白、激素、酶、酶抑制剂、具有抗肿瘤作用的多肽、能够抑制细菌、寄生虫或病毒(尤其是HIV)感染的多肽、抗体、毒素、免疫毒素。
优选地，根据本发明的组合产品可以被配制在药学上可接受的载体中。
在根据本发明的组合产品的一个实施方案中，在所述目的DNA序列之前施用所述载体。
本发明还涉及根据本发明的组合产品，其用作预防和/或治疗人或动物病状(特别是通过基因疗法的手段)的药物。
所述病状可以选自由以下组成的组：可以通过施用至少一种如上所述的目的DNA序列来改善的病状。
例如，所述病状及其临床、生物学或遗传亚型可以选自包括以下的组：肝脏疾病(例如，由于对乙酰氨基酚中毒或其他原因引起的急性肝衰竭、肝切除术后肝衰竭的预防、包括肝细胞瘤或胆管癌在内的肝原发性癌症、非酒精性脂肪性肝炎以及肝单基因疾病例如血色素沉着症、鸟氨酸转氨甲酰酶缺乏症、精氨琥珀酸裂解酶缺乏症、精氨琥珀酸合成酶1、血色素沉着症或威尔逊氏病(Wilson’s disease))、由于分泌蛋白缺乏或与之相关的疾病(例如，溶酶体贮积病例如高雪氏病(Gaucher's disease)、尼曼-皮克病(Niemann-Pickdisease)、Tay-Sacks或Sandhoff病、Hunter综合征或Hurler病；凝血因子(包括因子VIIIc、IX、Von Willebrand、纤维蛋白原或其他凝血蛋白)以及集落刺激因子(包括促红细胞生成素、粒细胞集落刺激因子和血小板生成素)的缺乏、癌症及其易感性(例如，乳腺癌、结直肠癌、胰腺癌、胃癌、食道癌和肺癌以及黑色素瘤)、恶性血液病(例如，白血病、霍奇金淋巴瘤和非霍奇金淋巴瘤、骨髓瘤)、血红蛋白病(例如，镰状细胞性贫血、6-磷酸葡萄糖脱氢酶缺乏症)和地中海贫血、自身免疫性疾病(例如，系统性红斑狼疮、硬皮病、自身免疫性肝炎)、心血管疾病(例如，心律和传导障碍、肥厚型心肌病、心血管疾病或慢性心力衰竭)、代谢疾病(例如，I型和II型糖尿病及其并发症、血脂异常、动脉粥样硬化及其并发症)、传染性疾病(例如，AIDS、乙型病毒性肝炎、丙型病毒性肝炎、流感、寨卡、埃博拉和其他病毒性疾病；肉毒杆菌中毒、破伤风和其他细菌性疾病；疟疾和其他寄生虫病)、肌肉疾病(例如，杜氏肌营养不良症和Steinert强直性肌营养不良症)、呼吸系统疾病(例如，囊性纤维化、α-1抗胰蛋白酶缺乏症、急性呼吸窘迫综合征、肺动脉高压、肺静脉阻塞性疾病)、肾脏疾病(例如，多囊性肾病、肾小球病)、结直肠疾病(例如，克罗恩病(Crohn’s disease)和溃疡性结肠炎)、眼部疾病特别是视网膜疾病(例如，Leber黑内障、色素性视网膜炎、年龄相关性黄斑变性)、中枢神经系统疾病(例如，阿尔茨海默氏病、帕金森氏病、肌萎缩性侧索硬化症、多发性硬化症、亨廷顿氏病、神经纤维瘤病、肾上腺脑白质营养不良、双相性疾病、精神分裂症和自闭症)、骨骼和关节疾病(例如，类风湿关节炎、强直性脊柱炎、骨关节炎)以及皮肤和结缔组织疾病(例如，神经纤维瘤病和牛皮癣)。
在一个实施方案中，本发明的组合产品包括：
-至少一种载体，其包含并表达根据本发明的核酸分子，其中所述DNA依赖性RNA聚合酶催化结构域是噬菌体DNA依赖性RNA聚合酶催化结构域；和
-至少一种目的DNA序列，其中所述DNA序列与所述噬菌体DNA依赖性RNA聚合酶催化结构域的启动子可操作地连接，并与至少一种编码与所述RNA结合结构域特异性结合的元件的序列共价连接。
本发明还涉及用于生产根据本发明的嵌合酶的方法，该方法包括以下步骤：在允许如下所述核酸分子在如下所述宿主细胞中表达的条件下，在至少一种宿主细胞中表达编码本发明嵌合酶的所述核酸分子或所述核酸分子组。
本发明还涉及一种用于生产根据本发明的嵌合酶的方法，该方法包括以下步骤：
-在允许如下所述核酸分子在如下所述宿主细胞中表达的条件下，在第一宿主细胞中表达编码本发明的嵌合酶的所述核酸分子组的一部分，以获得本发明的嵌合酶的第一部分；
-在允许如下所述核酸分子在如下所述宿主细胞中表达的条件下，在第二宿主细胞中表达编码本发明的嵌合酶的所述核酸分子组的其他部分，以获得本发明的嵌合酶的第二部分；
-组装所述第一部分和所述第二部分以获得本发明的嵌合酶。
附图说明
图1：pK1ERNAP和pT3RNAP质粒的图谱。
图2：测试拴系的pNλ-D12L质粒的图谱。其他测试拴系的测试质粒具有相同的一般设计，除了ORF，其通过内切核酸酶限制性消化和连接被取代。
图3：编码非拴系痘苗病毒加帽酶D12L亚基的测试pD12L质粒的图谱。其他非拴系测试质粒具有相同的一般设计，除了ORF，其通过内切核酸酶限制性消化和连接被取代。
图4：不具有4xλBoxBr重复序列的非拴系pK1Ep-荧光素酶和pT3p-荧光素酶报告质粒的图谱。
图5：具有4xλBoxBr重复序列的Nλ拴系的pK1Ep-荧光素酶-4xλBoxBr和pT3p-荧光素酶-4xλBoxBr报告质粒的图谱。
图6：K1ERNAP或T3RNAP驱动的表达系统，其用于测试与痘苗病毒加帽酶偶联的Nλ-拴系系统对未加帽的4xλBoxBr拴系的萤火虫荧光素酶mRNA的表达的活性。
图7：K1ERNAP或T3RNAP驱动的表达系统，其用于测试与非洲猪瘟病毒加帽酶偶联的Nλ-拴系系统对未加帽的4xλBoxBr拴系的萤火虫荧光素酶mRNA的表达的活性。
图8：K1ERNAP驱动的表达系统，其用于分析与非洲猪瘟病毒加帽酶偶联的各种蛋白质：RNA结合系统对未加帽的4xλBoxBr拴系的萤火虫荧光素酶mRNA的表达的活性。
图9：K1ERNAP驱动的表达系统，其用于分析与痘苗病毒加帽酶偶联的各种蛋白质：RNA结合系统对未加帽的4xλBoxBr拴系的萤火虫荧光素酶mRNA的表达的活性。
图10：与融合有poly(A)聚合酶的非洲猪瘟病毒加帽酶偶联的Nλ-拴系结构域产生的构建体的结构。(A)Nλ-poly(A)聚合酶-G4-NP868R单体，(B)Nλ-NP868R-G4-poly(A)聚合酶单体。
图11：K1ERNAP驱动的表达系统，其用于分析与融合有poly(A)聚合酶的非洲猪瘟病毒加帽酶偶联的Nλ-拴系系统对未加帽的4xλBoxBr拴系的萤火虫荧光素酶mRNA的表达的活性。
图12：与融合有poly(A)聚合酶的痘苗病毒加帽酶偶联的Nλ-拴系结构域产生的构建体的结构。箭头指示D12L-D1R结合。(A)Nλ-poly(A)聚合酶-G4-D12L/D1R异二聚体，(B)Nλ-D12L-G4-poly(A)聚合酶/D1R异二聚体。
图13：K1ERNAP驱动的表达系统，其用于分析与融合有poly(A)聚合酶的痘苗病毒加帽酶偶联的Nλ-拴系系统对未加帽的4xλBoxBr拴系的萤火虫荧光素酶mRNA的表达的活性。
图14：通过互补的亮氨酸拉链与结合有poly(A)聚合酶的痘苗病毒加帽酶偶联的Nλ-拴系系统的结构。箭头指示形成异二聚体的互补亮氨酸拉链。(A)Nλ-R341-EE1234L/RR1234L-NP868R异二聚体，和(B)Nλ-NP868R-EE1234L/RR1234L-R341异二聚体。
图15：K1ERNAP驱动的表达系统，其用于分析与融合有poly(A)聚合酶的痘苗病毒加帽酶偶联的Nλ-拴系系统对未加帽的4xλBoxBr拴系的萤火虫荧光素酶mRNA的表达的活性。箭头指示D12L-D1R结合。
图16：棘阿米巴多噬模拟病毒poly(A)聚合酶R341、非洲猪瘟病毒NP868R加帽酶和噬菌体K1E RNA聚合酶之间的拴系复合物的结构对聚腺苷酸化的4xλBoxB1拴系的萤火虫荧光素酶mRNA的表达。[X1]和[X2]指定可变结构域，其中G4和(G4S)2是柔性接头，T2A和F2A分别是来自1型猪捷申病毒和小核糖核酸病毒(picornavirus)口蹄疫口疮病毒的核糖体跳跃序列。
图17：表达系统，其用于分析棘阿米巴多噬菌拟似病毒poly(A)聚合酶R341、非洲猪瘟病毒NP868R加帽酶和噬菌体K1E RNA聚合酶之间的拴系复合物对4xλBoxB1拴系的萤火虫荧光素酶mRNA的表达的活性。
下面将通过实施例详细解释本发明，但是本发明的技术范围不限于这些实施例。
实施例1：与荧光素酶报告基因mRNA拴系在一起的痘苗病毒加帽酶D1R/D12L增加其表达
1.目标
本实验的目的是确定在细胞中合成的与萤火虫荧光素酶报告基因mRNA适当地拴系在一起的异源二聚的痘苗病毒加帽酶是否增加其表达。
痘苗病毒加帽酶由两个亚基组成，它们形成异二聚体：(i)由痘苗病毒D1R基因(基因序列AY243312.1；UniProtKB/Swiss-Prot登录号P04298)编码的95kDa亚基，下文称为D1R，其具有RNA三磷酸酶、RNA鸟苷酰基转移酶和RNA N7-鸟嘌呤甲基转移酶的酶活性(Cong和Shuman 1993,Niles和Christen 1993,Higman和Niles 1994,Mao和Shuman 1994,Gong和Shuman 2003)，(ii)和由痘苗病毒D12L基因(基因序列序列AY243312.1；UniProtKB/Swiss-Prot登录号P04318)编码的31-kDa亚基，下文称为D12L，它没有内在的酶活性，但增强D1R亚基的RNA N7-鸟嘌呤甲基转移酶活性(Higman,Bourgeois等人1992,Higman,Christen等人1994,Mao和Shuman 1994)。因此，编码这两个亚基的质粒的共转染在细胞中产生异二聚体的D1R/D12L加帽酶，其最终可以融合成蛋白质拴系结构域。
2.方法
a.质粒
针对密码子适应指数，使用GeneOptimizer算法(Raab,Graf等人2010)优化以下质粒的编码序列，以在人类细胞中表达。所有基因序列均使用寡核苷酸通过逐步PCR人工合成和组装，克隆并进行完整测序。
对于以下所有实施例，测试条件由几种质粒的可变组合组成。在本实施例中，pK1ERNAP/pT3RNAP质粒与萤火虫荧光素酶报告质粒一起用于在细胞中产生具有或不具有拴系结构域的萤火虫荧光素酶mRNA，然后测试质粒产生的酶(其与萤火虫荧光素酶mRNA适当地拴系在一起)可以对其进行特异性修饰。
表达质粒由噬菌体T3和K1E RNA聚合酶开放阅读框(ORF)组成，所述开放阅读框在除去T7
启动子序列后，被亚克隆到pCMVScript质粒骨架(Stratagene,La Jolla,CA)中。这些对应的质粒(在ORF的名称前面加上p-来命名)具有以下设计(即pK1ERNAP或pT3RNAP；图1)：来自人类巨细胞病毒(CMV)的IE1启动子/增强子、5’-非翻译区(5’-UTR)、Kozak共有序列、ORF、3’-非翻译区(3'-UTR)和SV40聚腺苷酸化信号。通过在紧邻Kozak序列上游和终止密码子下游的核酸内切酶限制性酶位点处消化，将相应的ORF亚克隆。
测试质粒包含被研究的加帽酶的编码序列，这些被研究的加帽酶带有(图2的pNλ-D12L)或不带有肽拴系结构域(图3的pD12L)。肽拴系结构域由来自λ噬菌体的抗终止N蛋白的22个氨基酸组成(Nλ；来自肠杆菌噬菌体λ核衣壳蛋白AAA32249的氨基酸1-22；SEQ ID N°1和SEQ ID N°2分别对应于λ噬菌体的抗终止N蛋白的N-末端拴系结构域的核苷酸和氨基酸序列)，通过柔性G4接头与D12L蛋白的氨基末端融合。通过在紧邻Kozak序列上游或Nλ-G4基序下游和终止密码子下游的核酸内切酶限制性酶位点处消化，将相应的ORF亚克隆。使用两对质粒来编码拴系的或非拴系的异二聚体的痘苗病毒加帽酶。首先，包含带有Nλ拴系结构域的D12L编码序列的质粒(pNλ-D12L；SEQ ID N°3和SEQ ID N°4分别对应于痘苗病毒加帽酶的D12L亚基的核苷酸和氨基酸序列)和包含野生型D1R编码序列的质粒(pD1R；SEQ ID N°5和SEQ ID N°6分别对应于痘苗病毒加帽酶的D1R亚基的核苷酸和氨基酸序列)产生拴系的D1R/Nλ-D12L异二聚体。其次，包含不带有Nλ拴系结构域的野生型D12L编码序列的质粒(pD12L)和包含野生型D1R编码序列的质粒(pD1R)形成了非拴系的D1R/D12L异二聚体。
萤火虫荧光素酶报告质粒包含处于K1E或T3 RNA聚合酶启动子的控制之下的萤火虫荧光素酶基因，并含有5’-UTR序列、Kozak共有序列，随后是来自北美萤火虫(photinuspyralis)的荧光素酶基因的ORF和终止密码子、由来自λ病毒的四个BoxBr串联组成的RNA拴系结构域(可选的，在非拴系形式中不存在；肠杆菌噬菌体λKT232076.1的基因组序列的核苷酸38312-38298；SEQ ID N°7对应于λ噬菌体的BoxBr RNA茎环的核苷酸序列)、40个腺苷残基的poly(A)轨道，随后是来自丁型肝炎病毒的基因组核酶的自切割RNA序列，并以噬菌体T7
转录终止子结束。这些质粒被称为pK1Ep-荧光素酶/pT3p-荧光素酶(其非拴系形式)(图4)，或pK1Ep-荧光素酶-4xλBoxBr/pT3p-荧光素酶-4xλBoxBr(其4xBoxBr RNA拴系形式)(图5)。因此，由该系统产生的RNA分子是未加帽的，并具有由模板萤火虫荧光素酶报告质粒中的40个腺苷残基编码的短的3’-末端聚腺苷酸化轨道。
b.细胞培养和转染
对于标准实验，人胚肾293(HEK-293，ATCC CRL 1573)在37℃，5％CO2气氛，100％相对湿度下常规生长。将细胞维持在补充有4mM L-丙氨酰-L-谷氨酰胺、10％胎牛血清(FBS)、1％非必需氨基酸、1％丙酮酸钠、1％青霉素和链霉素和0.25％两性霉素B的Dulbecco改良Eagle's培养基(DMEM)中。
转染前一天，将细胞常规接种于24孔板中，每孔1x 105个细胞，然后在80％细胞汇合率时转染。根据制造商的建议，使用Lipofectamine 2000试剂(Invitrogen，Carlsbad，CA)进行瞬时转染。除非另有说明，否则用2μl的Lipofectamine 2000和0.8μg的总质粒DNA转染细胞。对于标准的荧光素酶和hSEAP基因报告表达测定，转染后48小时分析细胞，除非另有说明。
c.萤火虫荧光素酶发光和SEAP比色测定
根据制造商的建议，通过荧光素酶测定系统(Promega，Madison，WI)测定荧光素酶的发光。简言之，将细胞在细胞培养裂解试剂缓冲液(CLR)中裂解，然后在12,000×g下于4℃离心2分钟。将以1：10稀释的荧光素酶测定试剂(Promega；100μl/孔)加入到上清液(20μl/孔)中。在Tristar 2酶标仪(Berthold，Bad Wildbad，德国)上进行发光读数，每孔的读数时间为一秒。
为了使转染效率标准化，将细胞用pORF-eSEAP质粒(InvivoGen，圣地亚哥，加利福尼亚)转染，该质粒编码由EF-1α/HTLV复合启动子驱动的人分泌的胚胎碱性磷酸酶(hSEAP)。使用Quanti-Blue比色酶测定试剂盒(InvivoGen)在细胞培养基中测定酶活性。基因报告表达表示为荧光素酶发光(RLU，相对光单位)与eSEAP吸光度(OD，光密度)的比值。
d.统计分析
统计分析采用配对双尾学生t检验进行。结果为平均值(n≥4)±标准差。P-值<0.05被认为具有统计学意义。
3.结果
在这组实验中，通过共转染pK1ERNAP/pT3RNAP和pK1Ep-荧光素酶-4xλBoxBr/pT3p-荧光素酶-4xλBoxBr(用于生成萤火虫荧光素酶报告质粒的拴系形式)或pK1Ep-荧光素酶-4xλBoxBr/pT3p-荧光素酶-4xλBoxBr(用于生成其非拴系形式)，由噬菌体K1E或T3RNA聚合酶生产了萤火虫荧光素酶mRNA。共转了染包含带有或不带有拴系结构域的D1R/D12L痘苗病毒的编码序列的测试质粒。通过萤火虫荧光素酶测定法，测量所得转录物的可翻译性，在有效加帽的情况下，其有望提高。图6显示了该测定法的一般性描述。
下表显示了使用K1E驱动的系统进行的第一组实验的结果：
如预期的那样，当缺少加帽时，仅与K1ERNAP质粒(称为pK1ERNAP)共转染的pK1Ep-荧光素酶和pK1Ep-荧光素酶-4xλBoxBr产生的萤火虫荧光素酶mRNA表达差(第1行和第2行)。与仅共转染pK1ERNAP/pK1Ep-荧光素酶-4xλBoxBr质粒的相比，非拴系D1R质粒(称为pD1R)与pK1ERNAP/pK1Ep-荧光素酶-4xλBoxBr一起共转染使表达增加了约9倍(第3行相对于第1或2行，p<0.05，双向学生t检验)，而非拴系的pD12L质粒与pK1ERNAP/pK1Ep-荧光素酶-4xλBoxBr的转染则对萤火虫荧光素酶表达几乎没有影响(第4行相对于第1或2行，p＝NS，双向学生t检验)。与先前的条件相比，非拴系的pD12L和pD1R质粒与pK1ERNAP/pK1Ep-荧光素酶-4xλBoxBr一起共转染，导致不带有拴系结构域的痘苗病毒D1R/D12L加帽酶异二聚体显著增加了萤火虫荧光素酶的表达，因此证实了mRNA翻译需要mRNA加帽(第5行相对于第1-4行，p＝NS，双向学生t检验，p<0.05，双向学生t检验)。最后，与没有痘苗病毒加帽酶(单独的pK1ERNAP/pK1Ep-荧光素酶-4xλBoxB)，或非拴系的痘苗病毒加帽酶(pK1ERNAP/pK1Ep-荧光素酶4xλBoxBr/pD1R/pD12L)相比，将拴系的D12L质粒(pNλ-D12L)和D1R与pK1ERNAP/pK1Ep-荧光素酶-4xλBoxBr一起共转染，其产生拴系的痘苗病毒加帽酶D1R/Nλ-D12L，使荧光素酶mRNA的表达分别大大增加了63.3倍(第6行相对于第2行)和2.6倍(第6行相对于第5行)。
在第二组实验中，由噬菌体T3 RNA聚合酶产生萤火虫荧光素酶mRNA，并使其与痘苗病毒加帽酶拴系在一起。下表显示了用T3驱动的系统进行的第二组实验的结果：
第二组实验给出了非常相似的转染结果，共转染结果顺序如下：不带有4xλBoxBr和加帽酶的RNA(第1行，pT3RNAP/pT3p-荧光素酶)≈4xλBoxBr-RNA，不带有加帽酶(第2行，pT3RNAP/pT3p-荧光素酶-4xλBoxBr)≈4xλBoxBr-RNA，仅带有D12L亚基(第4行，pT3RNAP/pD12L/pT3p-荧光素酶-4xλBoxBr)<4xλBoxBr-RNA，带有D1R(第3行，pT3RNAP/pD1R/pT3p-荧光素酶-4xλBoxBr-4)<4xλBoxBr-RNA，带有非拴系的D1R/D12L加帽酶(第5行，pT3RNAP/pD1R/pD12L/pT3p-荧光素酶-4xλBoxBr)<<4xλBoxBr-RNA，带有拴系的D1R/D12L加帽酶(第6行，pT3RNAP/pD1R/pNλ-D12L/pT3p-荧光素酶-4xλBoxBr)。后一种情况的表达水平在统计学上高于所有其他条件，特别是比带有非拴系的D1R/D12L加帽酶高3.2倍，因此证明了通过拴系结构域将D1R/D12L加帽酶引导至目标报告基因mRNA的重要性(p<0.05，双向学生t检验)。
4.结论
这些实验表明，当通过Nλ-BoxBr拴系系统与未加帽且聚腺苷酸化的萤火虫荧光素酶报告基因mRNA适当地束缚在一起时，不包含特定RNA序列的已知或已证明的结合结构域的痘苗病毒加帽酶会大大增加基因萤火虫荧光素酶报告基因的表达。
实施例2：与荧光素酶报道基因mRNA拴系在一起的非洲猪瘟病毒加帽酶NP868R增加其表达
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1.目标
这组实验的目的是证明，与聚腺苷酸化的萤火虫荧光素酶报告基因mRNA适当地拴系在一起的非洲猪瘟病毒加帽酶NP868R是否可以增加其在细胞中的表达。NP868R(也称为G4R)是一个868个氨基酸的单亚基蛋白，在体外证明了其具有cap-0形成所需的所有酶活性，即RNA三磷酸酶、RNA鸟苷酰基转移酶和RNA N7-鸟嘌呤甲基转移酶(Pena,Yanez等人1993,Jais 2011,Dixon,Chapman等人2013,Jais,Decroly等人2018)。
2.方法
a.质粒
表达质粒(pK1ERNAP或pT3RNAP)以及拴系形式的萤火虫荧光素酶报告质粒(pK1Ep-荧光素酶和pT3p-荧光素酶)或非拴系形式的萤火虫荧光素酶报告质粒(pK1Ep-荧光素酶-4xλBoxBr和pT3p-荧光素酶-4xλBoxBr)与如上所述相同。
如上所述，将由非洲猪瘟病毒NP868R加帽酶(NCBI ASFV基因组序列毒株BA71VNC_001659；UniProtKB/Swiss-Prot登录号P32094；SEQ ID N°8和SEQ ID N°9分别对应于非洲猪瘟病毒NP868R加帽酶的核苷酸序列和氨基酸序列)的编码序列组成的、带有Nλ拴系结构域(pNλ-NP868R)或不带有Nλ拴系结构域(pNP868R)的测试质粒亚克隆到pCMVScript质粒骨架中。
b.细胞培养和转染
与实施例1中所述相同。
c.萤火虫荧光素酶发光和SEAP比色测定
与实施例1中所述相同。
d.统计分析
与实施例1中所述相同。
3.结果
测定的设计与实施例1非常相似，不同之处在于其使用单亚基加帽酶NP868R代替异二聚体的D1R/D12L加帽酶。简而言之，通过噬菌体K1E或T3RNA聚合酶在细胞中合成了未加帽但聚腺苷酸化的萤火虫荧光素酶mRNA，并测定了其在NP868R存在下的表达(图7)。
下表显示了使用K1E驱动的系统进行的第一组实验的结果：
在这第一组实验中，用pK1ERNAP/pK1Ep-荧光素酶或pK1ERNAP/pK1Ep-荧光素酶-4xλBoxBr质粒共转染细胞，显示低水平的萤火虫荧光素酶报告基因表达(第1行和第2行)。与单独的pK1ERNAP/pK1Ep-荧光素酶-4xλBoxBr相比，pK1ERNAP/pK1Ep-荧光素酶-4xλBoxBr与非拴系的pNP868R质粒(pNP868R)共转染使表达增加了约29倍(第3行分别相对于第1行或第2行，p<0.05，双向学生t检验)，因此证实了mRNA翻译需要mRNA加帽。最后，与非拴系的NP868R情况相比，pK1ERNAP/pK1Ep-荧光素酶-4xλBoxBr与拴系的NP868R(pNλ-NP868R)的共转染甚至使萤火虫荧光素酶的表达增加了2.1倍，证明了通过拴系结构域将酶引导至目标mRNA对于有效的mRNA加帽的重要性(第4行相对于第3行，p<0.05，双向学生t检验)。
下表显示了用T3驱动的系统进行的第二组实验的结果：
在该第二组实验中，由噬菌体T3 RNA聚合酶产生萤火虫荧光素酶mRNA，并使其与非洲猪瘟病毒加帽酶拴系在一起。第二组实验给出了非常相似的结果，共转染结果顺序如下：不带有4xλBoxBr和加帽酶的RNA(第1行，pT3RNAP/pT3p-荧光素酶)≈4xλBoxBr-RNA，不带有加帽酶(第2行，pT3RNAP/pT3p-荧光素酶-4xλBoxBr)<4xλBoxBr-RNA，带有非拴系的NP868R加帽酶(第3行，pT3RNAP/pNP868R/pT3p-荧光素酶-4xλBoxBr)<<4xλBoxBr-RNA，带有拴系的NP868R加帽酶(第4行，pT3RNAP/pNλ-NP868R/pT3p-荧光素酶-4xλBoxBr)。最后一种情况的表达水平在统计学上高于所有其他条件，特别是比非拴系的NP868R加帽酶高2.9倍，因此证明了通过拴系结构域将NP868R拴系至靶mRNA对于有效的mRNA加帽的重要性(第4行相对于第3行，p<0.05，双向学生t检验)。
4.结论
这些实验表明，当通过Nλ-BoxBr拴系系统与未加帽且聚腺苷酸化的报告基因mRNA适当地拴系在一起时，不包含特定RNA序列的已知或预测的结合结构域的另一种加帽酶——非洲猪瘟病毒的NP868R——会增加萤火虫荧光素酶报告基因的表达。
实施例3：与非洲猪瘟病毒加帽酶NP868R偶联的各种蛋白质：RNA拴系系统可以增加宿主细胞的细胞质中通过K1E噬菌体RNA聚合酶产生的荧光素酶报告基因mRNA的表达
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1.目标
本实验的目的是研究除Nλ-4xBoxBr系统外的其他蛋白质：RNA拴系系统是否可以指导非洲猪瘟病毒加帽酶NP868R，以增加通过噬菌体K1E RNA聚合酶产生的适当拴系的荧光素酶报告基因mRNA的表达。
目前测试了以下拴系系统：i)MS2病毒的MS2蛋白和RNA茎环拴系序列(Valegard,Murray等人1994,Valegard,Murray等人1997)，ii)λ病毒的Nλ肽及其BoxB1 RNA拴系序列(Das 1993,Greenblatt,Nodwell等人1993,Friedman和Court 1995)，iii)P22λ病毒的Nλ肽及其BoxBr RNA拴系序列(Das 1993,Greenblatt,Nodwell等人1993,Friedman和Court1995)，iv)Φ21λ病毒的Nλ肽及其BoxBr RNA拴系序列(Das 1993,Greenblatt,Nodwell等人1993,Friedman和Court 1995)，v)人免疫缺陷病毒1(HIV-1)的TAT结合结构域(其包含经过生物学验证的核定位信号(Duconge和Toulme 1999))以及TAR RNA拴系序列(Dingwall,Ernberg等人1990,Weeks,Ampe等人1990,Karn,Dingwall等人1991,Puglisi,Tan等人1992,Frankel和Young 1998)，和vi)人小核核糖核蛋白U1亚基70(SNRNP70)蛋白质拴系序列(Romac,Graff等人1994)(其包含经过生物学验证的核定位信号(Keene,Query等人1999))及其U1snRNA-茎环拴系序列。
2.方法
a.质粒
pK1ERNAP表达质粒在实施例1中描述。
测试质粒由在其氨基末端融合有以下蛋白质或肽的非洲猪瘟病毒NP868R加帽酶的编码序列组成：i)噬菌体N-抗终止蛋白整个MS2蛋白的N-末端(pMS2-NP868R，NCBI登录号NC_001417.2，UniProtKB/Swiss-Prot P03612；SEQ ID N°36和SEQ ID N°37分别对应于核苷酸和氨基酸序列)，ii)前述λ噬菌体的N-末端肽，iii)P22噬菌体N-抗终止蛋白的N-末端肽(pP22N-NP868R，UniProtKB/Swiss-Prot P04891)，iv)Φ21噬菌体N-抗终止蛋白的N-末端肽(pNΦ21-NP868R，UniProtKB/Swiss-Prot P07243)，v)HIV-1分离株HXB2的TAT蛋白结合结构域(pTAT-NP868R，NCBI参考序列：AAB50256.1)，和vi)人小核核糖核蛋白U1亚基70(SNRNP70)RNA结合蛋白序列(pSNRNP70-NP868R，氨基酸92-202，NCBI登录号NM_003089.5)。
萤火虫荧光素酶报告质粒的RNA拴系结构域被以下的四个串联重复序列取代：i)MS2 RNA茎环(pK1Ep-荧光素酶-4xMS2sl质粒；肠噬菌体MS2分离株DL52的核苷酸1748-766，NCBI登录号JQ966307.1；SEQ ID N°38)，ii)λ病毒的BoxB1 RNA序列(pK1Ep-荧光素酶-4xλBoxB1；NCBI登录号J02459.1核苷酸35518-35534)，iii)P22λ病毒的λBoxBr RNA序列(pK1Ep-荧光素酶-4xP22BoxBr；NCBI登录号NC_002371.2，核苷酸31,953-31,971)，iv)Φ21λ病毒的λBoxBr RNA序列(pK1Ep-荧光素酶-4xΦ21BoxBr；NCBI登录号AH007390.1，核苷酸866-883)，v)人类免疫缺陷病毒1型分离株HXB2的TAR RNA序列(pK1Ep-荧光素酶-4xTAR；NCBI登录号K03455.1，核苷酸471-497)和vi)U1snRNA RNA茎环(pK1Ep-荧光素酶-4xU1snRNA；NCBI登录号M28013.1，核苷酸123-155)。
b.细胞培养和转染
与实施例1中所述相同。
c.萤火虫荧光素酶发光和SEAP比色测定
与实施例1中所述相同。
d.统计分析
与实施例1中所述相同。
3.结果
该测定的设计与实施例2非常相似，不同之处在于测试了各种蛋白质：RNA拴系系统以替代Nλ：BoxBr系统。简而言之，通过噬菌体K1E RNA聚合酶在细胞中合成了具有40个腺苷残基的短聚腺苷酸化尾的未加帽的萤火虫荧光素酶mRNA，并测定了其在被各种系统拴系的NP868R存在下的表达(图8)。
这些实验的结果如下表所示：
当与非拴系系统相比，pK1ERNAP与仅具有拴系系统的一个组分(即与测试质粒的NP868R加帽酶融合的蛋白结构域)的质粒或萤火虫荧光素酶报告质粒(在其3’UTR中引入了四个串联的RNA拴系重复序列)的共转染，对带有任何系统的萤火虫荧光素酶报告基因mRNA的表达没有明显影响(第2或3行相对于第1行；对于所有比较，p＝NS，双向学生t检验)。与非拴系加帽酶和/或非拴系的萤火虫荧光素酶质粒的条件相比，将pK1ERNAP与编码MS2-4xMS2sl的组分的质粒(即pMS2-NP868R/pK1Ep-荧光素酶-4xMS2sl)、编码Nλ-NP868R-4xλBoxBl的组分的质粒(即pNλ-NP868R/pK1Ep-荧光素酶-4xλBoxBl)、编码NP22-NP868R-4xP22BoxBr的组分的质粒(即pNP22-NP868R/pK1Ep-荧光素酶-4xP22BoxBr)、编码NΦ21-4xΦ21BoxBr的组分的质粒(即pNΦ21-NP868R/pK1Ep-荧光素酶-4xΦ21BoxBr)共转染，拴系系统使萤火虫荧光素酶报告基因的表达水平显著增加了3.8至4.6倍(第4行相对于第1-3行；对于所有比较，p<0.05，双向学生t检验)。相比之下，与具有非拴系加帽酶和/或非拴系的萤火虫荧光素酶质粒的条件相比，pK1ERNAP与TAT/4xTAR拴系系统(即pTAT-NP868R/pK1Ep-荧光素酶-4xTAR)或SNRNP70/4xU1snRNA拴系系统(即pSNRNP70-NP868R/pK1Ep-荧光素酶-4xU1snRNA)的共转染显示出非常低的萤火虫荧光素酶变化(第4行相对于第1-3行；对于所有比较，p＝NS，双向学生t检验)。
总之，用Nλ-4xλBoxB1拴系表达系统(即pNλ-NP868R/pK1Ep-荧光素酶-4xλBoxB1)获得了最佳性能，其他拴系系统的性能按以下顺序排列(即条件4与1的比例)：Nλ-4xλBoxBl>MS2-4xMS2sl>NΦ21-4xΦ21BoxBr>NP22-4xP22BoxBr>>TAT-4xTAR>SNRNP70-4xU1snRNA。
4.结论
本实验表明，当通过噬菌体蛋白质-RNA拴系系统与mRNA适当地拴系在一起时，非洲猪瘟病毒加帽酶NP868R可以提高萤火虫荧光素酶mRNA的表达。
实施例4：与痘苗病毒加帽的D12L亚基偶联的噬菌体蛋白：RNA拴系系统可以增加宿主细胞的细胞质中通过K1E噬菌体RNA聚合酶产生的荧光素酶报告基因mRNA的表达
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1.目标
本实验的目的是研究除Nλ-4xBoxBr系统以外的其他蛋白质：RNA拴系系统是否可以用于将异二聚体的加帽酶从痘苗病毒引导至目标mRNA，从而增加其表达。
下文中测试的蛋白质：RNA拴系系统与先前实施例中描述的相同。
2.方法
a.质粒
pK1ERNAP表达质粒在实施例1中描述。
测试质粒由在其氨基末端与上述拴系蛋白序列融合的痘苗病毒加帽酶的D12L亚基的编码序列组成。D1R质粒与先前描述的相同。
含有各种拴系的RNA序列的萤火虫荧光素酶报告质粒与先前实施例中所述相同。
b.细胞培养和转染
与实施例1中所述相同。
c.萤火虫荧光素酶发光和SEAP比色测定
与实施例1中所述相同。
d.统计分析
与实施例1中所述相同。
3.结果
测定的设计与实施例1非常相似，不同之处在于，测试了各种蛋白质：RNA拴系系统以替代Nλ：BoxBr系统(图9)。
这些实验的结果如下表所示：
当与非拴系系统相比，pK1ERNAP与仅具有拴系系统的两个组分中的一个(即与测试质粒的痘苗病毒加帽的D12L亚基融合的蛋白结构域)的质粒或萤火虫荧光素酶报告质粒(在其3’UTR中引入了四个串联的RNA拴系的重复序列)的共转染，对带有任何系统的萤火虫荧光素酶报告基因mRNA的表达没有明显影响(第2或3行相对于第1行；对于所有比较，p＝NS，双向学生t检验)。与以前的发现相似，与具有非拴系加帽酶和/或非拴系的萤火虫荧光素酶质粒的条件相比，将pK1ERNAP与具有MS2-4xMS2sl的所有组分和痘苗病毒加帽酶的D1R亚基的质粒、具有Nλ-D12L-4xλBoxBl的所有组分和痘苗病毒加帽酶的D1R亚基的质粒(即pNλ-D12L/pD1R/pK1Ep-荧光素酶-4xλBoxBl)、具有NP22-D12L-4xP22BoxBr的所有组分和痘苗病毒加帽酶的D1R亚基的质粒(即pNP22-D12L/pD1R/pK1Ep-荧光素酶-4xP22BoxBr)、具有NΦ21-4xΦ21BoxBr的所有组分和痘苗病毒加帽酶的D1R亚基的质粒(即pNΦ21-D12L/pD1R/pK1Ep-荧光素酶-4xΦ21BoxBr)共转染，拴系系统使萤火虫荧光素酶报告基因的表达水平显著增加了2至2.5倍(第4行相对于第1-3行；对于所有比较，p<0.05，双向学生t检验)。相比之下，与具有非拴系加帽酶和/或非拴系的萤火虫荧光素酶质粒的条件相比，pK1ERNAP与TAT/4xTAR拴系系统(即pTAT-D12L/pD1R/pK1Ep-荧光素酶-4xTAR)或SNRNP70/4xU1snRNA拴系系统(即pSNRNP70-D12L/pD1R/pK1Ep-荧光素酶-4xU1snRNA)的共转染显示出非常低的萤火虫荧光素酶变化(对于所有比较，p＝NS，双向学生t检验)。
最后，拴系系统的性能按与上一个实施例不同的顺序排列(即条件4与1的比例)：NΦ21-4xΦ21BoxBr>NP22-4xP22BoxBr>Nλ-4xλBoxBl>MS2-4xMS2sl>>TAT-4xTAR>SNRNP70-4xU1snRNA。
4.结论
本实验表明，当通过噬菌体蛋白质-RNA拴系系统的噬菌体RNA结合结构域与mRNA适当地拴系在一起时，来自痘苗病毒的异二聚体的D1R/D12L加帽酶也可以提高通过K1E噬菌体RNA聚合酶产生的萤火虫荧光素酶mRNA的表达。
实施例5：当与目标转录物适当地拴系在一起时，poly(A)聚合酶和非洲猪瘟病毒NP868R加帽酶之间的融合可增加通过K1E噬菌体RNA聚合酶产生的荧光素酶报告基因mRNA的表达
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1.目标
本实验旨在确定，当通过Nλ-BoxBl拴系系统与目标转录物适当地拴系在一起时，与NP868R非洲猪瘟病毒加帽酶融合的poly(A)聚合酶是否可以增加通过K1E噬菌体RNA聚合酶产生的萤火虫荧光素酶报告基因mRNA的表达。
2.方法
a.质粒
合成了以下poly(A)聚合酶的ORF：i)酿酒酵母的PAP1 poly(A)聚合酶，通过缺失含有核定位信号的42个羧基末端氨基酸而被分类到细胞质中(Zhelkovsky,Helmling等人1998)(NCBI登录号：P29468)；ii)痘苗病毒VP55 poly(A)聚合酶(UniProtKB/Swiss-Prot登录号P23371分别对应于核苷酸和氨基酸序列)，iii)棘阿米巴多噬模拟病毒的病毒性R341poly(A)聚合酶(UniProtKB/Swiss-Prot登录号：E3VZZ8)，iv)Megavirus chilensis的病毒MG561 poly(A)聚合酶(NCBI登录号：YP_004894612)，v)非洲猪瘟病毒的病毒性C475L poly(A)聚合酶(UniProtKB/Swiss-Prot登录号：A0A0A1E081)，vi)突变的PAPOLA(人PAPOLA的K656R-K657R突变，UniProtKB/Swiss-Prot登录号P51003)，在其核定位信号处发生了突变(Raabe,Murthy等人1994,Vethantham,Rao等人2008)，vii)野生型典型小家鼠睾丸特异性PAPOLB(UniProtKB/Swiss-Prot Q9WVP6)。
通过在以下位置框内亚克隆poly(A)聚合酶ORF生成了四种类型的测试质粒：i)仅在pCMV-Script骨架中(例如pPAP1)，ii)在Nλ拴系结构域的下游(例如pNλ-PAP1)，iii)通过G4柔性接头在Nλ-NP868R蛋白的下游(例如pNλ-NP868R-G4-PAP1)，导致单体蛋白的表达，或iv)通过G4柔性接头在λ噬菌体的N蛋白拴系结构域和NP868R之间(例如pNλ-PAP1-G4-NP868R)，也导致单体蛋白的表达。后两种结构的设计如图10所示。
萤火虫荧光素酶报告质粒的非拴系形式(pK1Ep-荧光素酶)或拴系形式(pK1Ep-荧光素酶-4xλBoxB1)与上述相同。
b.细胞培养和转染
与实施例1中所述相同。
c.萤火虫荧光素酶发光和SEAP比色测定
与实施例1中所述相同。
d.统计分析
与实施例1中所述相同。
3.结果
测定的设计与实施例2非常相似，不同之处在于，poly(A)聚合酶与非洲猪瘟病毒NP868R加帽酶融合(图11)。
这些实验的结果如下表所示：
在没有由pNλ-NP868R提供的mRNA加帽的情况下，当用非拴系的poly(A)聚合酶质粒(第2行相对于第1行)或拴系的poly(A)聚合酶质粒(第3行相对于第1行)转染时，观察到萤火虫荧光素酶mRNA的表达非统计学显著增加，分别约1.5倍和2.5倍(p＝NS，双向学生t检验)。当萤火虫荧光素酶mRNA被pNλ-NP868R的共转染加帽时，当用非拴系的或拴系的poly(A)聚合酶质粒转染时，观察到表达统计学显著增加，分别约1.5倍(第5行相对于第4行)和约2倍(第6行相对于第4行)(对于所有非拴系的poly(A)聚合酶，与无poly(A)聚合酶相比，p<0.05，双向学生t检验)。
将Poly(A)聚合酶与NP868R非洲猪瘟病毒加帽酶以及Nλ蛋白拴系结构域融合在一起。测试了两种融合类型，通过G4柔性接头将poly(A)聚合酶亚克隆到Nλ-NP868R下游或通过G4柔性接头将poly(A)聚合酶亚克隆到λ噬菌体的N蛋白拴系结构域和NP868R之间。与非连接的酶相比，两种类型的所有拴系的融合基因均增加了萤火虫荧光素酶mRNA的表达(第7和8行相对于第6行；对于所有比较，p<0.05，双向学生t检验)。融合蛋白的活性排列如下：Nλ-NP868R-G4-C475L<Nλ-NP868R-G4-R341<Nλ-MG561-G4-NP868R<Nλ-VP55-G4-NP868R<Nλ-C475L-G4-NP868R<Nλ-R341-G4-NP868R<Nλ-NP868R-G4-MG561<Nλ-PAPOLB-G4-NP868R<Nλ-NP868R-G4-PAPOLB<Nλ-NP868R-G4-VP55<Nλ-PAP1-G4-NP868R<Nλ-NP868R-G4-PAPOLA<Nλ-PAPOLA-G4-NP868R<Nλ-NP868R-G4-PAP1
4.结论
本实验表明，当适当地拴系至Nλ-4xBoxBl系统时，与非洲猪瘟病毒NP868R加帽酶融合的各种poly(A)聚合酶(包括哺乳动物、酵母、病毒和细菌的酶)均可以提高通过噬菌体RNA聚合酶产生的转录物的表达。令人惊讶的是，各种poly(A)聚合酶和NP868R加帽酶(它们在自然情况下没有物理连接并且不包含RNA结合结构域)之间的融合可以协同起作用，并且当这些融合蛋白被适当地拴系时，这种作用甚至更大(上表中的第7和8行)。这些结果确实令人惊讶，并且由于成分相同，本领域技术人员可能预期获得相同的表达率。
实施例6：当与目标转录物适当地拴系在一起时，poly(A)聚合酶和异二聚体的痘苗病毒加帽酶的D12亚基的融合可以增加通过K1E噬菌体RNA聚合酶产生的荧光素酶报告基因mRNA的表达。
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1.目标
本实验旨在确定，当与Nλ-4xBoxB1系统适当地拴系时，poly(A)聚合酶与异二聚体的痘苗病毒加帽酶的D12亚基的融合是否可以增加通过噬菌体RNA聚合酶产生的转录物的表达。
2.方法
a.质粒
pK1ERNAP表达质粒在实施例1中描述。
测试的poly(A)聚合酶在先前的实施例中进行了描述，并框内亚克隆至如下位置(图12)：i)通过G4柔性接头在Nλ-D12L蛋白的下游(例如pNλ-D12L-G4-PAP1)，或ii)通过G4柔性接头在λ噬菌体的N蛋白拴系结构域和D12L之间(例如pNλ-PAP1-G4-D12L)。
萤火虫荧光素酶报告质粒的非拴系形式(pK1Ep-荧光素酶)或拴系形式(pK1Ep-荧光素酶-4xλBoxB1)与上述相同。
b.细胞培养和转染
与实施例1中所述相同。
c.萤火虫荧光素酶发光和SEAP比色测定
与实施例1中所述相同。
d.统计分析
与实施例1中所述相同。
3.结果
实验的设计与先前的实施例相似，不同之处在于，加帽酶由痘苗病毒异二聚体D1R/D12组成(图13)。
这些实验的结果如下表所示：
在没有由pNλ-D12L/D1R提供的mRNA加帽的情况下，当用非拴系的poly(A)聚合酶质粒(第2行相对于第1行)或拴系的poly(A)聚合酶质粒(第3行相对于第1行)转染时，观察到萤火虫荧光素酶mRNA的表达无统计学显著变化，分别约1.5倍和2倍(p＝NS，双向学生t检验)。与以前的发现类似，当萤火虫荧光素酶mRNA被pNλ-D12L/D1R的共转染加帽时，当用非拴系的或拴系的poly(A)聚合酶质粒转染时，观察到表达统计学显著增加，分别约1.5倍(第5行相对于第4行)和约2倍(第6行相对于第4行)(对于所有非拴系的poly(A)聚合酶，与无poly(A)聚合酶相比，p<0.05，双向学生t检验)。
如上所述，将poly(A)聚合酶与异二聚体的痘苗病毒加帽酶的D12亚基以及Nλ-蛋白结构域融合在一起。与非连接的酶相比，两种类型的所有拴系的融合基因均增加了萤火虫荧光素酶mRNA的表达(第7和8行相对于第6行；对于所有比较，p<0.05，双向学生t检验)。融合复合物的活性排列如下：Nλ-D12L/D1R-G4-C475L<Nλ-C475L-G4-D12L/D1R<Nλ-D12L/D1R-G4-MG561<Nλ-PAPOLB-G4-D12L/D1R<Nλ-VP55-G4-D12L/D1R<Nλ-D12L/D1R-G4-R341<Nλ-D12L/D1R-G4-VP55<Nλ-MG561-G4-D12L/D1R<Nλ-D12L/D1R-G4-PAPOLB<Nλ-PAPOLA-G4-D12L/D1R<Nλ-D12L/D1R-G4-PAP1<Nλ-R341-G4-D12L/D1R<Nλ-D12L/D1R-G4-PAPOLA<Nλ-PAP1-G4-D12L/D1R。
4.结论
本实验显示，与异二聚体的痘苗病毒加帽酶的D12亚基以及D1R亚基融合的各种poly(A)聚合酶(它们在自然情况下没有物理连接并且不包含RNA结合结构域)可以协同起作用，并且当这些融合蛋白被适当地拴系时，这种作用甚至更大。
实施例7：棘阿米巴多噬模拟病毒R341 poly(A)聚合酶和非洲猪瘟病毒NP868R加帽酶之间的非共价拴系偶联，可以增加通过噬菌体RNA聚合酶产生的荧光素酶报告基因mRNA的表达
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1.目标
这组实验的目的是为了确定，当被适当地拴系时，R341 poly(A)聚合酶和NP868R加帽酶之间的非共价偶联是否能够形成一个能够增强未加帽的萤火虫荧光素酶报告基因mRNA的表达的活性表达系统。在本实验中，使用形成异二聚体的互补亮氨酸拉链生成非共价偶联。
2.方法
a.质粒
pK1ERNAP表达质粒和非拴系形式(pK1Ep-荧光素酶)或拴系形式(pK1Ep-荧光素酶-4xλBoxBr)的萤火虫荧光素酶报告质粒与上述相同。
由EE1234L和RR1234L互补亮氨酸拉链(SEQ ID N°28和SEQ ID N°29分别对应于G4-EE1234L亮氨酸拉链的核苷酸和氨基酸序列；SEQ ID N°30和SEQ ID N°31分别对应于RR1234L-G4亮氨酸拉链的核苷酸和氨基酸序列)诱导棘阿米巴多噬模拟病毒R341 poly(A)聚合酶与非洲猪瘟病毒NP868R加帽酶之间的非共价偶联。这些两亲性α-螺旋形成反向平行异二聚体，解离亲和力约10-15M(Moll,Ruvinov等人2001)。在NP868R加帽酶和R341 RNA聚合酶之间生成了两个非共价异二聚体复合物(图14)。首先，将EE1234L亮氨酸拉链与Nλ-R41的羧基末端框内融合，将RR1234L与NP868R的氨基末端框内融合。因此，这两种质粒(分别命名为pNλ-R341-EE1234L和RR1234L-NP868R)的共表达会产生具有由R341携带的Nλ拴系结构域的异二聚体。其次，将RR1234L亮氨酸拉链与Nλ-NP868R的羧基末端框内融合，将EE1234L与R341的氨基末端框内融合。因此，这两种质粒(分别命名为pNλ-NP868R-RR1234L和pEE1234L-R341)的共表达会产生具有由NP868R携带的Nλ拴系结构域的异二聚体。通过在内切核酸酶限制性酶位点将相应ORF亚克隆到pCMVScript质粒骨架中来合成这些质粒。
b.细胞培养和转染
与实施例1中所述相同。
c.萤火虫荧光素酶发光和SEAP比色测定
与实施例1中所述相同。
d.统计分析
与实施例1中所述相同。
3.结果
如前所述，使用pK1Ep-荧光素酶和pK1Ep-荧光素酶-4xλBoxBr质粒，通过K1E噬菌体RNA聚合酶生成了未加帽的萤火虫荧光素酶报告基因mRNA(图15)。
这些实验的结果如下表所示：
与非拴系的棘阿米巴吞噬拟象病毒R341 poly(A)聚合酶相比，带有或不带有亮氨酸拉链的棘阿米巴多吞噬菌拟象病毒R341 poly(A)聚合酶的Nλ-拴系适当地增加了通过噬菌体K1E RNA聚合酶产生的非加帽4xλBoxBr-萤火虫荧光素酶mRNA的表达(第1或3行相对于第2行)。与非拴系的NP868R加帽酶或无加帽酶相比，带有或不带有亮氨酸拉链的非洲猪瘟病毒NP868R加帽酶的Nλ拴系明显增加了通过噬菌体K1E RNA聚合酶产生的非加帽萤火虫荧光素酶4xλBoxBr-mRNA的表达(第4或6行相对于第5行)。
使用EE1234L和RR1234L互补性高亲和力亮氨酸拉链，在棘阿米巴多噬模拟病毒R341poly(A)聚合酶和非洲猪瘟病毒NP868R加帽酶之间产生了非共价偶联。与单独使用具有亮氨酸拉链的非拴系R341或NP868R的条件相比，这种没有Nλ-拴系结构域的异二聚体复合物产生了活性复合物，该活性复合物显著增加了通过噬菌体K1E RNA聚合酶产生的非加帽萤火虫荧光素酶mRNA的表达(第7行相对于第2或5行)。值得注意的是，与非拴系复合物相比，在此异二聚体复合物的棘阿米巴吞噬拟象病毒R341 poly(A)聚合酶或非洲猪瘟病毒NP868R加帽酶的氨基末端添加Nλ-拴系，使非加帽萤火虫荧光素酶mRNA的表达增加了2.80倍和2.24倍(第8或9行相对于第7行；对于两个比较，p<0.05，双向学生t检验)。
4.结论
本实验表明，R341 poly(A)聚合酶与非洲猪瘟病毒NP868R加帽酶之间的人工偶联(即通过亮氨酸拉链非共价偶联)也可产生协同活性的异二聚体，当这些融合蛋白被适当地拴系时，这种作用甚至更大。
实施例8：棘阿米巴多噬模拟病毒R341 poly(A)聚合酶、非洲猪瘟病毒NP868R加帽酶和噬菌体K1E RNA聚合酶之间的组装产生了活性表达复合物
-----------------------------------------------
1.目标
以下实验的目的是为了确定，当被适当地Nλ-拴系时，通过组装棘阿米巴多噬模拟病毒poly(A)聚合酶R341、非洲猪瘟病毒NP868R加帽酶和噬菌体K1E RNA聚合酶是否可以产生活性复合物。
以下测试的组装体是根据常见的Nλ-R341-[X1]-NP868R-[X2]-K1ERNAP蛋白支架设计的，其中[X1]和[X2]是可变的。
生成了以下开放阅读框架以测试此假设(图16)：
[X1]＝G4,[X2]＝G4；即Nλ-R341-G4-NP868R-G4-K1ERNAP构建体(NCBI登录号J02459-SEQ ID N°40和SEQ ID N°41分别对应于Nλ-R341-G4-NP868R-G4-K1ERNAP的核苷酸和氨基酸序列)，其特征在于，这三个酶亚基通过柔性接头与Nλ-拴系肽在其N-末端融合，
[X1]＝G4,[X2]＝F2A；即Nλ-R341-G4-NP868R-F2A-K1ERNAP构建体(SEQ ID N°42和SEQ ID N°43分别对应于Nλ-R341-G4-NP868R-F2A-K1ERNAP的核苷酸和氨基酸序列)，其特征在于，将非洲猪瘟病毒NP868R加帽酶和噬菌体K1E RNA聚合酶之间的G4柔性接头替换为来自小核糖核酸病毒口疮病毒的F2A核糖体跳跃序列(O型口蹄疫口疮病毒多蛋白，UniProtKB/Swiss-Prot登录号AAT01756，残基934-955)。核糖体跳跃导致蛋白质的明显共翻译切割(Donnelly,Luke等人2001)，
[X1]＝F2A,[X2]＝G4；即Nλ-R341-F2A-NP868R-G4-K1ERNAP构建体(SEQ ID N°44和SEQ ID N°45分别对应于Nλ-R341-F2A-NP868R-G4-K1ERNAP的核苷酸和氨基酸序列)，其特征在于，将R341 poly(A)聚合酶和非洲猪瘟病毒NP868R加帽酶之间的G4柔性接头替换为F2A核糖体跳跃序列。
[X1]＝T2A,[X2]＝G4；即Nλ-R341-T2A-NP868R-G4-K1ERNAP构建体(SEQ ID N°46和SEQ ID N°47分别对应于Nλ-R341-T2A-NP868R-G4-K1ERNAP的核苷酸和氨基酸序列)，其特征在于，将R341 poly(A)聚合酶和非洲猪瘟病毒NP868R加帽酶之间的G4柔性接头替换为来自1型猪捷申病毒的T2A核糖体跳跃序列(UniProtKB/Swiss-Prot登录号Q9WJ28，残基979-997)。
2.方法
a.质粒
萤火虫荧光素酶报告质粒的非拴系形式(pK1Ep-荧光素酶)或拴系形式(pK1Ep-荧光素酶-4xλBoxB1)与上述相同。
将先前描述的ORF亚克隆到pCMVScript骨架中，从而得到了pNλ-R341-G4-NP868R-G4-K1ERNAP、pNλ-R341-G4-NP868R-F2A-K1ERNAP、pNλ-R341-F2A-NP868R-G4-K1ERNAP和pNλ-R341-T2A-NP868R-G4-K1ERNAP质粒。
b.细胞培养和转染
与实施例1中所述相同。
c.萤火虫荧光素酶发光和SEAP比色测定
与实施例1中所述相同。
d.统计分析
与实施例1中所述相同。
3.结果
图17显示了该测定的图示。
这些实验的结果如下表所示：
将K1ERNAP编码序列与Nλ-R341-F2A-NP868R的各种融合与未连接的Nλ-R341-F2A-NP868R和K1ERNAP进行了比较。与未连接的Nλ-R341-F2A-NP868R和K1ERNAP相比，所有融合构建体的表达水平均明显更高(第2至5行相对于第1行；对于所有比较，p<0.05，双向学生t检验)。Nλ-R341-T2A-NP868R-G4-K1ERNAP和Nλ-R341-F2A-NP868R-G4-K1ERNAP融合获得了最佳结果，其他条件的排列顺序如下：Nλ-R341-T2A-NP868R-G4-K1ERNAP～Nλ-R341-F2A-NP868R-G4-K1ERNAP>>Nλ-R341-G4-NP868R-F2A-K1ERNAP>Nλ-R341-G4-NP868R-G4-K1ERNAP>pNλ-R341-G4-NP868R,pK1ERNAP。
4.结论
本实验表明，通过将poly(A)聚合酶R341、非洲猪瘟病毒NP868R加帽酶和噬菌体K1E RNA聚合酶组装在Nλ-R341-[X1]-NP868R-[X2]-K1ERNAP(优选地，其中[X1]＝T2A或F2A，并且[X2]＝G4)支架之下，可以产生活性拴系表达系统。出乎意料的是，与构建体Nλ-R341和NP868R-G4-K1ERNAP的联合相比，构建体Nλ-R341-F2A-NP868R-G4-K1ERNAP和Nλ-R341-T2A-NP868R-G4-K1ERNAP能够具有更高的表达率(第4和5行相对于第1行)。这些结果确实令人惊讶，并且由于成分相同，并且自然情况下没有物理连接，也不包含任何RNA结合结构域，本领域技术人员可能预期获得相同的表达率。
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序列表
<110> 优卡瑞斯
<120> 新型嵌合酶及其应用
<130> B00801-PCT
<150> EP 17306006.2
<151> 2017-07-27
<160> 64
<170> PatentIn version 3.5
<210> 1
<211> 66
<212> DNA
<213> λ噬菌体
<400> 1
atggatgctc agaccagacg cagagaacgg cgggcagaga agcaggcaca gtggaaggcc 60
gctaat 66
<210> 2
<211> 22
<212> PRT
<213> λ噬菌体
<400> 2
Met Asp Ala Gln Thr Arg Arg Arg Glu Arg Arg Ala Glu Lys Gln Ala
1 5 10 15
Gln Trp Lys Ala Ala Asn
20
<210> 3
<211> 870
<212> DNA
<213> 痘苗病毒
<400> 3
atggaattcg acgagatcgt gaagaacatc cgcgagggca cccacgtgct gctgcccttc 60
tacgagacac tgcccgagct gaacctgtcc ctgggcaagt cccccctgcc ctccctggaa 120
tacggcgcca actacttcct gcagatctcc agagtgaacg acctgaaccg gatgcccacc 180
gacatgctga agctgttcac ccacgacatc atgctgcccg agtccgacct ggacaaggtg 240
tacgagattc tgaagatcaa ctccgtgaag tactacggcc ggtccacaaa ggctgacgcc 300
gtggtggccg acctgtctgc ccggaacaag ctgttcaaga gagagcggga cgccatcaag 360
tccaacaacc acctgaccga gaacaacctg tacatctccg actacaagat gctgaccttc 420
gacgtgttcc ggcccctgtt cgacttcgtg aacgagaagt actgcatcat caagctgccc 480
accctgttcg gcagaggcgt gatcgacacc atgcggatct actgctccct gttcaagaac 540
gtgcggctgc tgaagtgcgt gtccgactcc tggctgaagg actccgccat catggtggcc 600
tccgacgtgt gcaagaagaa cctggacctg ttcatgtccc acgtgaagtc cgtgaccaag 660
tcatccagct ggaaggacgt gaactccgtg cagttctcca tcctgaacaa ccccgtggac 720
accgagttca tcaacaagtt tctggaattt tccaaccggg tgtacgaggc cctgtactac 780
gtgcactccc tgctgtactc ctccatgacc tccgactcca agtccatcga gaacaaacat 840
cagcgacggc tggtcaagct gctgctgtaa 870
<210> 4
<211> 287
<212> PRT
<213> 痘苗病毒
<400> 4
Met Asp Glu Ile Val Lys Asn Ile Arg Glu Gly Thr His Val Leu Leu
1 5 10 15
Pro Phe Tyr Glu Thr Leu Pro Glu Leu Asn Leu Ser Leu Gly Lys Ser
20 25 30
Pro Leu Pro Ser Leu Glu Tyr Gly Ala Asn Tyr Phe Leu Gln Ile Ser
35 40 45
Arg Val Asn Asp Leu Asn Arg Met Pro Thr Asp Met Leu Lys Leu Phe
50 55 60
Thr His Asp Ile Met Leu Pro Glu Ser Asp Leu Asp Lys Val Tyr Glu
65 70 75 80
Ile Leu Lys Ile Asn Ser Val Lys Tyr Tyr Gly Arg Ser Thr Lys Ala
85 90 95
Asp Ala Val Val Ala Asp Leu Ser Ala Arg Asn Lys Leu Phe Lys Arg
100 105 110
Glu Arg Asp Ala Ile Lys Ser Asn Asn His Leu Thr Glu Asn Asn Leu
115 120 125
Tyr Ile Ser Asp Tyr Lys Met Leu Thr Phe Asp Val Phe Arg Pro Leu
130 135 140
Phe Asp Phe Val Asn Glu Lys Tyr Cys Ile Ile Lys Leu Pro Thr Leu
145 150 155 160
Phe Gly Arg Gly Val Ile Asp Thr Met Arg Ile Tyr Cys Ser Leu Phe
165 170 175
Lys Asn Val Arg Leu Leu Lys Cys Val Ser Asp Ser Trp Leu Lys Asp
180 185 190
Ser Ala Ile Met Val Ala Ser Asp Val Cys Lys Lys Asn Leu Asp Leu
195 200 205
Phe Met Ser His Val Lys Ser Val Thr Lys Ser Ser Ser Trp Lys Asp
210 215 220
Val Asn Ser Val Gln Phe Ser Ile Leu Asn Asn Pro Val Asp Thr Glu
225 230 235 240
Phe Ile Asn Lys Phe Leu Glu Phe Ser Asn Arg Val Tyr Glu Ala Leu
245 250 255
Tyr Tyr Val His Ser Leu Leu Tyr Ser Ser Met Thr Ser Asp Ser Lys
260 265 270
Ser Ile Glu Asn Lys His Gln Arg Arg Leu Val Lys Leu Leu Leu
275 280 285
<210> 5
<211> 2541
<212> DNA
<213> 痘苗病毒
<400> 5
atggaattcg acgccaacgt ggtgtcctcc tccacaatcg ccacctacat cgacgccctg 60
gccaagaacg cctccgagct ggaacagcgg tccaccgcct acgagatcaa caatgagctg 120
gaactggtgt tcatcaagcc ccccctgatc accctgacca acgtggtcaa catcagcacc 180
atccaggaat ccttcatccg gttcaccgtg accaacaaag aaggcgtgaa gatccggacc 240
aagatccccc tgtccaaggt gcacggcctg gacgtgaaga acgtgcagct ggtggacgcc 300
atcgacaaca tcgtgtggga gaagaagtcc ctggtcaccg agaaccggct gcacaaagag 360
tgcctgctgc ggctgtccac cgaggaacgg cacatctttc tggactacaa gaagtacggc 420
tcctccatca gactggaact ggtcaacctg atccaggcca agaccaagaa cttcaccatc 480
gacttcaagc tgaagtactt cctgggctct ggcgcccagt ccaagtcctc tctgctgcac 540
gccatcaacc accccaagtc ccggcccaac acctccctgg aaatcgagtt cacccctcgg 600
gacaacgaga cagtgcccta cgacgagctg atcaaagagc tgaccaccct gtccagacac 660
atcttcatgg cctcccccga gaacgtgatc ctgtcccccc ccatcaacgc ccccatcaag 720
accttcatgc tgcccaagca ggacatcgtg ggcctggacc tggaaaacct gtacgccgtg 780
accaagaccg acggcatccc catcaccatc agagtgacct ccaacggcct gtactgctac 840
ttcacccacc tgggctacat catcagatac cccgtgaagc ggatcatcga ctccgaggtg 900
gtggtgttcg gcgaggccgt gaaggacaag aactggaccg tgtacctgat caagctgatc 960
gagcccgtga acgccatcaa tgaccggctg gaagagtcca aatacgtgga atccaagctg 1020
gtggatatct gcgaccggat cgtgttcaag tctaagaagt acgagggacc cttcactaca 1080
acttcagagg tggtcgacat gctgtccacc tacctgccta agcagcccga gggcgtcatc 1140
ctgttctact ccaagggacc caagtccaac atcgatttca agatcaagaa agagaacacc 1200
atcgaccaga ccgccaatgt ggtgttccgg tacatgtcct ccgagcccat catcttcggc 1260
gagtcctcca tcttcgtcga gtacaagaag ttctccaacg acaagggctt ccccaaagag 1320
tacggcagcg gcaagatcgt gctgtacaac ggcgtgaact acctgaacaa catctactgc 1380
ctggagtaca tcaacaccca caacgaagtg ggcatcaagt ccgtggtggt gcccatcaag 1440
tttatcgccg agttcctggt caacggcgag atcctgaagc cccggatcga caagaccatg 1500
aagtacatca attccgagga ctactacggc aaccagcaca acatcatcgt ggaacacctg 1560
agggaccagt ccatcaagat cggcgacatc ttcaacgagg acaagctgtc cgacgtgggc 1620
caccagtacg ccaacaacga caagttccgg ctgaaccccg aggtgtccta cttcaccaac 1680
aagagaaccc gaggcccact gggcatcctg tccaactacg tgaaaaccct gctgatctcc 1740
atgtactgct ccaagacctt cctggacgac tccaacaagc ggaaggtgct ggccatcgat 1800
ttcggcaacg gcgccgatct ggaaaagtac ttctatggcg agatcgccct gctggtggct 1860
accgaccctg acgccgacgc tatcgccaga ggcaacgagc ggtacaacaa gctgaactcc 1920
ggcatcaaga ccaagtacta caagttcgac tacatccagg aaaccatccg ctccgacacc 1980
ttcgtgtcct ccgtgcgcga ggtgttctat ttcggcaagt tcaatatcat cgactggcag 2040
ttcgccatcc actacagctt ccacccccgg cactacgcca ccgtgatgaa caacctgtcc 2100
gagctgaccg cctccggcgg caaggtgctg atcaccacca tggacggcga caagctgagc 2160
aagctgaccg acaagaaaac cttcatcatc cacaagaacc tgccctccag cgagaactac 2220
atgtccgtgg aaaagatcgc cgacgacaga atcgtggtgt acaatccctc caccatgtcc 2280
acccccatga ccgagtacat catcaagaag aacgacatcg tccgggtgtt caacgagtac 2340
ggcttcgtgc tggtggacaa cgtggacttc gccaccatca tcgagcggtc caaaaagttc 2400
atcaatggag ccagcaccat ggaagatcgg ccctctaccc ggaacttctt cgagctgaac 2460
agaggcgcca tcaagtgcga gggcctggat gtggaagatc tgctgagcta ctacgtggtg 2520
tacgtgttct ccaagagata a 2541
<210> 6
<211> 844
<212> PRT
<213> 痘苗病毒
<400> 6
Met Asp Ala Asn Val Val Ser Ser Ser Thr Ile Ala Thr Tyr Ile Asp
1 5 10 15
Ala Leu Ala Lys Asn Ala Ser Glu Leu Glu Gln Arg Ser Thr Ala Tyr
20 25 30
Glu Ile Asn Asn Glu Leu Glu Leu Val Phe Ile Lys Pro Pro Leu Ile
35 40 45
Thr Leu Thr Asn Val Val Asn Ile Ser Thr Ile Gln Glu Ser Phe Ile
50 55 60
Arg Phe Thr Val Thr Asn Lys Glu Gly Val Lys Ile Arg Thr Lys Ile
65 70 75 80
Pro Leu Ser Lys Val His Gly Leu Asp Val Lys Asn Val Gln Leu Val
85 90 95
Asp Ala Ile Asp Asn Ile Val Trp Glu Lys Lys Ser Leu Val Thr Glu
100 105 110
Asn Arg Leu His Lys Glu Cys Leu Leu Arg Leu Ser Thr Glu Glu Arg
115 120 125
His Ile Phe Leu Asp Tyr Lys Lys Tyr Gly Ser Ser Ile Arg Leu Glu
130 135 140
Leu Val Asn Leu Ile Gln Ala Lys Thr Lys Asn Phe Thr Ile Asp Phe
145 150 155 160
Lys Leu Lys Tyr Phe Leu Gly Ser Gly Ala Gln Ser Lys Ser Ser Leu
165 170 175
Leu His Ala Ile Asn His Pro Lys Ser Arg Pro Asn Thr Ser Leu Glu
180 185 190
Ile Glu Phe Thr Pro Arg Asp Asn Glu Thr Val Pro Tyr Asp Glu Leu
195 200 205
Ile Lys Glu Leu Thr Thr Leu Ser Arg His Ile Phe Met Ala Ser Pro
210 215 220
Glu Asn Val Ile Leu Ser Pro Pro Ile Asn Ala Pro Ile Lys Thr Phe
225 230 235 240
Met Leu Pro Lys Gln Asp Ile Val Gly Leu Asp Leu Glu Asn Leu Tyr
245 250 255
Ala Val Thr Lys Thr Asp Gly Ile Pro Ile Thr Ile Arg Val Thr Ser
260 265 270
Asn Gly Leu Tyr Cys Tyr Phe Thr His Leu Gly Tyr Ile Ile Arg Tyr
275 280 285
Pro Val Lys Arg Ile Ile Asp Ser Glu Val Val Val Phe Gly Glu Ala
290 295 300
Val Lys Asp Lys Asn Trp Thr Val Tyr Leu Ile Lys Leu Ile Glu Pro
305 310 315 320
Val Asn Ala Ile Asn Asp Arg Leu Glu Glu Ser Lys Tyr Val Glu Ser
325 330 335
Lys Leu Val Asp Ile Cys Asp Arg Ile Val Phe Lys Ser Lys Lys Tyr
340 345 350
Glu Gly Pro Phe Thr Thr Thr Ser Glu Val Val Asp Met Leu Ser Thr
355 360 365
Tyr Leu Pro Lys Gln Pro Glu Gly Val Ile Leu Phe Tyr Ser Lys Gly
370 375 380
Pro Lys Ser Asn Ile Asp Phe Lys Ile Lys Lys Glu Asn Thr Ile Asp
385 390 395 400
Gln Thr Ala Asn Val Val Phe Arg Tyr Met Ser Ser Glu Pro Ile Ile
405 410 415
Phe Gly Glu Ser Ser Ile Phe Val Glu Tyr Lys Lys Phe Ser Asn Asp
420 425 430
Lys Gly Phe Pro Lys Glu Tyr Gly Ser Gly Lys Ile Val Leu Tyr Asn
435 440 445
Gly Val Asn Tyr Leu Asn Asn Ile Tyr Cys Leu Glu Tyr Ile Asn Thr
450 455 460
His Asn Glu Val Gly Ile Lys Ser Val Val Val Pro Ile Lys Phe Ile
465 470 475 480
Ala Glu Phe Leu Val Asn Gly Glu Ile Leu Lys Pro Arg Ile Asp Lys
485 490 495
Thr Met Lys Tyr Ile Asn Ser Glu Asp Tyr Tyr Gly Asn Gln His Asn
500 505 510
Ile Ile Val Glu His Leu Arg Asp Gln Ser Ile Lys Ile Gly Asp Ile
515 520 525
Phe Asn Glu Asp Lys Leu Ser Asp Val Gly His Gln Tyr Ala Asn Asn
530 535 540
Asp Lys Phe Arg Leu Asn Pro Glu Val Ser Tyr Phe Thr Asn Lys Arg
545 550 555 560
Thr Arg Gly Pro Leu Gly Ile Leu Ser Asn Tyr Val Lys Thr Leu Leu
565 570 575
Ile Ser Met Tyr Cys Ser Lys Thr Phe Leu Asp Asp Ser Asn Lys Arg
580 585 590
Lys Val Leu Ala Ile Asp Phe Gly Asn Gly Ala Asp Leu Glu Lys Tyr
595 600 605
Phe Tyr Gly Glu Ile Ala Leu Leu Val Ala Thr Asp Pro Asp Ala Asp
610 615 620
Ala Ile Ala Arg Gly Asn Glu Arg Tyr Asn Lys Leu Asn Ser Gly Ile
625 630 635 640
Lys Thr Lys Tyr Tyr Lys Phe Asp Tyr Ile Gln Glu Thr Ile Arg Ser
645 650 655
Asp Thr Phe Val Ser Ser Val Arg Glu Val Phe Tyr Phe Gly Lys Phe
660 665 670
Asn Ile Ile Asp Trp Gln Phe Ala Ile His Tyr Ser Phe His Pro Arg
675 680 685
His Tyr Ala Thr Val Met Asn Asn Leu Ser Glu Leu Thr Ala Ser Gly
690 695 700
Gly Lys Val Leu Ile Thr Thr Met Asp Gly Asp Lys Leu Ser Lys Leu
705 710 715 720
Thr Asp Lys Lys Thr Phe Ile Ile His Lys Asn Leu Pro Ser Ser Glu
725 730 735
Asn Tyr Met Ser Val Glu Lys Ile Ala Asp Asp Arg Ile Val Val Tyr
740 745 750
Asn Pro Ser Thr Met Ser Thr Pro Met Thr Glu Tyr Ile Ile Lys Lys
755 760 765
Asn Asp Ile Val Arg Val Phe Asn Glu Tyr Gly Phe Val Leu Val Asp
770 775 780
Asn Val Asp Phe Ala Thr Ile Ile Glu Arg Ser Lys Lys Phe Ile Asn
785 790 795 800
Gly Ala Ser Thr Met Glu Asp Arg Pro Ser Thr Arg Asn Phe Phe Glu
805 810 815
Leu Asn Arg Gly Ala Ile Lys Cys Glu Gly Leu Asp Val Glu Asp Leu
820 825 830
Leu Ser Tyr Tyr Val Val Tyr Val Phe Ser Lys Arg
835 840
<210> 7
<211> 15
<212> DNA
<213> λ噬菌体
<400> 7
gccctgaaaa agggc 15
<210> 8
<211> 2613
<212> DNA
<213> 非洲猪瘟病毒
<400> 8
atggaattcg ccagcctgga caacctggtg gccagatacc agcggtgctt caacgaccag 60
agcctgaaga acagcaccat cgagctggaa atccggttcc agcagatcaa cttcctgctg 120
ttcaagaccg tgtacgaggc cctggtcgcc caggaaatcc ccagcaccat cagccacagc 180
atccggtgca tcaagaaggt gcaccacgag aaccactgcc gggagaagat cctgcccagc 240
gagaacctgt acttcaagaa acagcccctg atgttcttca agttcagcga gcccgccagc 300
ctgggctgta aagtgtccct ggccatcgag cagcccatcc ggaagttcat cctggacagc 360
agcgtgctgg tccggctgaa gaaccggacc accttccggg tgtccgagct gtggaagatc 420
gagctgacca tcgtgaagca gctgatgggc agcgaggtgt cagccaagct ggccgccttc 480
aagaccctgc tgttcgacac ccccgagcag cagaccacca agaacatgat gaccctgatc 540
aaccccgacg acgagtacct gtacgagatc gagatcgagt acaccggcaa gcctgagagc 600
ctgacagccg ccgacgtgat caagatcaag aacaccgtgc tgacactgat cagccccaac 660
cacctgatgc tgaccgccta ccaccaggcc atcgagttta tcgccagcca catcctgagc 720
agcgagatcc tgctggcccg gatcaagagc ggcaagtggg gcctgaagag actgctgccc 780
caggtcaagt ccatgaccaa ggccgactac atgaagttct acccccccgt gggctactac 840
gtgaccgaca aggccgacgg catccggggc attgccgtga tccaggacac ccagatctac 900
gtggtggccg accagctgta cagcctgggc accaccggca tcgagcccct gaagcccacc 960
atcctggacg gcgagttcat gcccgagaag aaagagttct acggctttga cgtgatcatg 1020
tacgagggca acctgctgac ccagcagggc ttcgagacac ggatcgagag cctgagcaag 1080
ggcatcaagg tgctgcaggc cttcaacatc aaggccgaga tgaagccctt catcagcctg 1140
acctccgccg accccaacgt gctgctgaag aatttcgaga gcatcttcaa gaagaaaacc 1200
cggccctaca gcatcgacgg catcatcctg gtggagcccg gcaacagcta cctgaacacc 1260
aacaccttca agtggaagcc cacctgggac aacaccctgg actttctggt ccggaagtgc 1320
cccgagtccc tgaacgtgcc cgagtacgcc cccaagaagg gcttcagcct gcatctgctg 1380
ttcgtgggca tcagcggcga gctgtttaag aagctggccc tgaactggtg ccccggctac 1440
accaagctgt tccccgtgac ccagcggaac cagaactact tccccgtgca gttccagccc 1500
agcgacttcc ccctggcctt cctgtactac caccccgaca ccagcagctt cagcaacatc 1560
gatggcaagg tgctggaaat gcggtgcctg aagcgggaga tcaactacgt gcgctgggag 1620
atcgtgaaga tccgggagga ccggcagcag gatctgaaaa ccggcggcta cttcggcaac 1680
gacttcaaga ccgccgagct gacctggctg aactacatgg accccttcag cttcgaggaa 1740
ctggccaagg gacccagcgg catgtacttc gctggcgcca agaccggcat ctacagagcc 1800
cagaccgccc tgatcagctt catcaagcag gaaatcatcc agaagatcag ccaccagagc 1860
tgggtgatcg acctgggcat cggcaagggc caggacctgg gcagatacct ggacgccggc 1920
gtgagacacc tggtcggcat cgataaggac cagacagccc tggccgagct ggtgtaccgg 1980
aagttctccc acgccaccac cagacagcac aagcacgcca ccaacatcta cgtgctgcac 2040
caggatctgg ccgagcctgc caaagaaatc agcgagaaag tgcaccagat ctatggcttc 2100
cccaaagagg gcgccagcag catcgtgtcc aacctgttca tccactacct gatgaagaac 2160
acccagcagg tcgagaacct ggctgtgctg tgccacaagc tgctgcagcc tggcggcatg 2220
gtctggttca ccaccatgct gggcgaacag gtgctggaac tgctgcacga gaaccggatc 2280
gaactgaacg aagtgtggga ggcccgggag aacgaggtgg tcaagttcgc catcaagcgg 2340
ctgttcaaag aggacatcct gcaggaaacc ggccaggaaa tcggcgtcct gctgcccttc 2400
agcaacggcg acttctacaa tgagtacctg gtcaacaccg cctttctgat caagattttc 2460
aagcaccatg gctttagcct cgtgcagaag cagagcttca aggactggat ccccgagttc 2520
cagaacttca gcaagagcct gtacaagatc ctgaccgagg ccgacaagac ctggaccagc 2580
ctgttcggct tcatctgcct gcggaagaac taa 2613
<210> 9
<211> 868
<212> PRT
<213> 非洲猪瘟病毒
<400> 9
Met Ala Ser Leu Asp Asn Leu Val Ala Arg Tyr Gln Arg Cys Phe Asn
1 5 10 15
Asp Gln Ser Leu Lys Asn Ser Thr Ile Glu Leu Glu Ile Arg Phe Gln
20 25 30
Gln Ile Asn Phe Leu Leu Phe Lys Thr Val Tyr Glu Ala Leu Val Ala
35 40 45
Gln Glu Ile Pro Ser Thr Ile Ser His Ser Ile Arg Cys Ile Lys Lys
50 55 60
Val His His Glu Asn His Cys Arg Glu Lys Ile Leu Pro Ser Glu Asn
65 70 75 80
Leu Tyr Phe Lys Lys Gln Pro Leu Met Phe Phe Lys Phe Ser Glu Pro
85 90 95
Ala Ser Leu Gly Cys Lys Val Ser Leu Ala Ile Glu Gln Pro Ile Arg
100 105 110
Lys Phe Ile Leu Asp Ser Ser Val Leu Val Arg Leu Lys Asn Arg Thr
115 120 125
Thr Phe Arg Val Ser Glu Leu Trp Lys Ile Glu Leu Thr Ile Val Lys
130 135 140
Gln Leu Met Gly Ser Glu Val Ser Ala Lys Leu Ala Ala Phe Lys Thr
145 150 155 160
Leu Leu Phe Asp Thr Pro Glu Gln Gln Thr Thr Lys Asn Met Met Thr
165 170 175
Leu Ile Asn Pro Asp Asp Glu Tyr Leu Tyr Glu Ile Glu Ile Glu Tyr
180 185 190
Thr Gly Lys Pro Glu Ser Leu Thr Ala Ala Asp Val Ile Lys Ile Lys
195 200 205
Asn Thr Val Leu Thr Leu Ile Ser Pro Asn His Leu Met Leu Thr Ala
210 215 220
Tyr His Gln Ala Ile Glu Phe Ile Ala Ser His Ile Leu Ser Ser Glu
225 230 235 240
Ile Leu Leu Ala Arg Ile Lys Ser Gly Lys Trp Gly Leu Lys Arg Leu
245 250 255
Leu Pro Gln Val Lys Ser Met Thr Lys Ala Asp Tyr Met Lys Phe Tyr
260 265 270
Pro Pro Val Gly Tyr Tyr Val Thr Asp Lys Ala Asp Gly Ile Arg Gly
275 280 285
Ile Ala Val Ile Gln Asp Thr Gln Ile Tyr Val Val Ala Asp Gln Leu
290 295 300
Tyr Ser Leu Gly Thr Thr Gly Ile Glu Pro Leu Lys Pro Thr Ile Leu
305 310 315 320
Asp Gly Glu Phe Met Pro Glu Lys Lys Glu Phe Tyr Gly Phe Asp Val
325 330 335
Ile Met Tyr Glu Gly Asn Leu Leu Thr Gln Gln Gly Phe Glu Thr Arg
340 345 350
Ile Glu Ser Leu Ser Lys Gly Ile Lys Val Leu Gln Ala Phe Asn Ile
355 360 365
Lys Ala Glu Met Lys Pro Phe Ile Ser Leu Thr Ser Ala Asp Pro Asn
370 375 380
Val Leu Leu Lys Asn Phe Glu Ser Ile Phe Lys Lys Lys Thr Arg Pro
385 390 395 400
Tyr Ser Ile Asp Gly Ile Ile Leu Val Glu Pro Gly Asn Ser Tyr Leu
405 410 415
Asn Thr Asn Thr Phe Lys Trp Lys Pro Thr Trp Asp Asn Thr Leu Asp
420 425 430
Phe Leu Val Arg Lys Cys Pro Glu Ser Leu Asn Val Pro Glu Tyr Ala
435 440 445
Pro Lys Lys Gly Phe Ser Leu His Leu Leu Phe Val Gly Ile Ser Gly
450 455 460
Glu Leu Phe Lys Lys Leu Ala Leu Asn Trp Cys Pro Gly Tyr Thr Lys
465 470 475 480
Leu Phe Pro Val Thr Gln Arg Asn Gln Asn Tyr Phe Pro Val Gln Phe
485 490 495
Gln Pro Ser Asp Phe Pro Leu Ala Phe Leu Tyr Tyr His Pro Asp Thr
500 505 510
Ser Ser Phe Ser Asn Ile Asp Gly Lys Val Leu Glu Met Arg Cys Leu
515 520 525
Lys Arg Glu Ile Asn Tyr Val Arg Trp Glu Ile Val Lys Ile Arg Glu
530 535 540
Asp Arg Gln Gln Asp Leu Lys Thr Gly Gly Tyr Phe Gly Asn Asp Phe
545 550 555 560
Lys Thr Ala Glu Leu Thr Trp Leu Asn Tyr Met Asp Pro Phe Ser Phe
565 570 575
Glu Glu Leu Ala Lys Gly Pro Ser Gly Met Tyr Phe Ala Gly Ala Lys
580 585 590
Thr Gly Ile Tyr Arg Ala Gln Thr Ala Leu Ile Ser Phe Ile Lys Gln
595 600 605
Glu Ile Ile Gln Lys Ile Ser His Gln Ser Trp Val Ile Asp Leu Gly
610 615 620
Ile Gly Lys Gly Gln Asp Leu Gly Arg Tyr Leu Asp Ala Gly Val Arg
625 630 635 640
His Leu Val Gly Ile Asp Lys Asp Gln Thr Ala Leu Ala Glu Leu Val
645 650 655
Tyr Arg Lys Phe Ser His Ala Thr Thr Arg Gln His Lys His Ala Thr
660 665 670
Asn Ile Tyr Val Leu His Gln Asp Leu Ala Glu Pro Ala Lys Glu Ile
675 680 685
Ser Glu Lys Val His Gln Ile Tyr Gly Phe Pro Lys Glu Gly Ala Ser
690 695 700
Ser Ile Val Ser Asn Leu Phe Ile His Tyr Leu Met Lys Asn Thr Gln
705 710 715 720
Gln Val Glu Asn Leu Ala Val Leu Cys His Lys Leu Leu Gln Pro Gly
725 730 735
Gly Met Val Trp Phe Thr Thr Met Leu Gly Glu Gln Val Leu Glu Leu
740 745 750
Leu His Glu Asn Arg Ile Glu Leu Asn Glu Val Trp Glu Ala Arg Glu
755 760 765
Asn Glu Val Val Lys Phe Ala Ile Lys Arg Leu Phe Lys Glu Asp Ile
770 775 780
Leu Gln Glu Thr Gly Gln Glu Ile Gly Val Leu Leu Pro Phe Ser Asn
785 790 795 800
Gly Asp Phe Tyr Asn Glu Tyr Leu Val Asn Thr Ala Phe Leu Ile Lys
805 810 815
Ile Phe Lys His His Gly Phe Ser Leu Val Gln Lys Gln Ser Phe Lys
820 825 830
Asp Trp Ile Pro Glu Phe Gln Asn Phe Ser Lys Ser Leu Tyr Lys Ile
835 840 845
Leu Thr Glu Ala Asp Lys Thr Trp Thr Ser Leu Phe Gly Phe Ile Cys
850 855 860
Leu Arg Lys Asn
865
<210> 10
<211> 5277
<212> DNA
<213> 人工序列
<220>
<223> C3P3-K1E嵌合酶的核苷酸序列，由非洲猪瘟病毒NP868R
加帽酶和突变型噬菌体K1E-RNA聚合酶融合构成
<400> 10
atggaattcg ccagcctgga caacctggtg gccagatacc agcggtgctt caacgaccag 60
agcctgaaga acagcaccat cgagctggaa atccggttcc agcagatcaa cttcctgctg 120
ttcaagaccg tgtacgaggc cctggtcgcc caggaaatcc ccagcaccat cagccacagc 180
atccggtgca tcaagaaggt gcaccacgag aaccactgcc gggagaagat cctgcccagc 240
gagaacctgt acttcaagaa acagcccctg atgttcttca agttcagcga gcccgccagc 300
ctgggctgta aagtgtccct ggccatcgag cagcccatcc ggaagttcat cctggacagc 360
agcgtgctgg tccggctgaa gaaccggacc accttccggg tgtccgagct gtggaagatc 420
gagctgacca tcgtgaagca gctgatgggc agcgaggtgt cagccaagct ggccgccttc 480
aagaccctgc tgttcgacac ccccgagcag cagaccacca agaacatgat gaccctgatc 540
aaccccgacg acgagtacct gtacgagatc gagatcgagt acaccggcaa gcctgagagc 600
ctgacagccg ccgacgtgat caagatcaag aacaccgtgc tgacactgat cagccccaac 660
cacctgatgc tgaccgccta ccaccaggcc atcgagttta tcgccagcca catcctgagc 720
agcgagatcc tgctggcccg gatcaagagc ggcaagtggg gcctgaagag actgctgccc 780
caggtcaagt ccatgaccaa ggccgactac atgaagttct acccccccgt gggctactac 840
gtgaccgaca aggccgacgg catccggggc attgccgtga tccaggacac ccagatctac 900
gtggtggccg accagctgta cagcctgggc accaccggca tcgagcccct gaagcccacc 960
atcctggacg gcgagttcat gcccgagaag aaagagttct acggctttga cgtgatcatg 1020
tacgagggca acctgctgac ccagcagggc ttcgagacac ggatcgagag cctgagcaag 1080
ggcatcaagg tgctgcaggc cttcaacatc aaggccgaga tgaagccctt catcagcctg 1140
acctccgccg accccaacgt gctgctgaag aatttcgaga gcatcttcaa gaagaaaacc 1200
cggccctaca gcatcgacgg catcatcctg gtggagcccg gcaacagcta cctgaacacc 1260
aacaccttca agtggaagcc cacctgggac aacaccctgg actttctggt ccggaagtgc 1320
cccgagtccc tgaacgtgcc cgagtacgcc cccaagaagg gcttcagcct gcatctgctg 1380
ttcgtgggca tcagcggcga gctgtttaag aagctggccc tgaactggtg ccccggctac 1440
accaagctgt tccccgtgac ccagcggaac cagaactact tccccgtgca gttccagccc 1500
agcgacttcc ccctggcctt cctgtactac caccccgaca ccagcagctt cagcaacatc 1560
gatggcaagg tgctggaaat gcggtgcctg aagcgggaga tcaactacgt gcgctgggag 1620
atcgtgaaga tccgggagga ccggcagcag gatctgaaaa ccggcggcta cttcggcaac 1680
gacttcaaga ccgccgagct gacctggctg aactacatgg accccttcag cttcgaggaa 1740
ctggccaagg gacccagcgg catgtacttc gctggcgcca agaccggcat ctacagagcc 1800
cagaccgccc tgatcagctt catcaagcag gaaatcatcc agaagatcag ccaccagagc 1860
tgggtgatcg acctgggcat cggcaagggc caggacctgg gcagatacct ggacgccggc 1920
gtgagacacc tggtcggcat cgataaggac cagacagccc tggccgagct ggtgtaccgg 1980
aagttctccc acgccaccac cagacagcac aagcacgcca ccaacatcta cgtgctgcac 2040
caggatctgg ccgagcctgc caaagaaatc agcgagaaag tgcaccagat ctatggcttc 2100
cccaaagagg gcgccagcag catcgtgtcc aacctgttca tccactacct gatgaagaac 2160
acccagcagg tcgagaacct ggctgtgctg tgccacaagc tgctgcagcc tggcggcatg 2220
gtctggttca ccaccatgct gggcgaacag gtgctggaac tgctgcacga gaaccggatc 2280
gaactgaacg aagtgtggga ggcccgggag aacgaggtgg tcaagttcgc catcaagcgg 2340
ctgttcaaag aggacatcct gcaggaaacc ggccaggaaa tcggcgtcct gctgcccttc 2400
agcaacggcg acttctacaa tgagtacctg gtcaacaccg cctttctgat caagattttc 2460
aagcaccatg gctttagcct cgtgcagaag cagagcttca aggactggat ccccgagttc 2520
cagaacttca gcaagagcct gtacaagatc ctgaccgagg ccgacaagac ctggaccagc 2580
ctgttcggct tcatctgcct gcggaagaac ctcgagggag gaggaggatc aggcggaggc 2640
ggaagtgtcg agcaggacct gcacgccatc cagctgcagc tcgaagagga aatgttcaac 2700
ggcggcatca gaagattcga ggccgaccag cagagacaga tcgcctctgg caacgagagc 2760
gacaccgcct ggaatagaag gctgctgtct gagctgatcg cccctatggc cgaaggcatc 2820
caggcctaca aagaggaata cgagggcaag agaggcagag cccctagagc cctggccttc 2880
atcaactgtg tgggcaatga ggtggccgcc tacatcacca tgaagatcgt gatggacatg 2940
ctgaacaccg acgtgaccct gcaggccatt gccatgaacg tggccgacag aatcgaggac 3000
caggtccgat tcagcaagct ggaaggacac gccgccaagt acttcgagaa agtgaagaag 3060
tccctgaagg ccagcaagac caagagctac agacacgccc acaacgtggc cgtggtggcc 3120
gaaaaatctg tggccgatag ggacgccgac ttctctagat gggaggcctg gcctaaggac 3180
accctgctgc agatcggcat gaccctgctg gaaatcctgg aaaacagcgt gttcttcaac 3240
ggccagcccg tgttcctgag aaccctgagg acaaatggcg gcaagcacgg cgtgtactac 3300
ctgcagacat ctgagcacgt gggcgagtgg atcaccgcct tcaaagaaca tgtggcccag 3360
ctgagccctg cctatgcccc ttgtgtgatc cctcctagac cctgggtgtc ccctttcaat 3420
ggcggctttc acaccgagaa ggtggccagc agaatcagac tggtcaaggg caaccgggaa 3480
cacgtgcgga agctgaccaa gaaacagatg cccgccgtgt acaaggccgt gaatgctctg 3540
caggccacca agtggcaggt caacaaagag gtgctgcagg tcgtcgagga cgtgatcaga 3600
ctggatctgg gctacggcgt gccaagcttt aagcccctga tcgacagaga gaacaagccc 3660
gccaaccctg tgcccctgga atttcagcac ctgagaggcc gcgagctgaa agagatgctg 3720
acacctgaac agtggcaggc ctttatcaat tggaagggcg agtgcaccaa gctgtacacc 3780
gccgagacaa agaggggctc taagtctgcc gccacagtgc gaatggtcgg acaggccaga 3840
aagtacagcc agttcgacgc catctacttc gtgtacgccc tggacagccg gtctagagtg 3900
tatgcccaga gcagcacact gagcccccag tctaacgatc tgggaaaggc cctgctgaga 3960
ttcaccgagg gccagagact ggattctgcc gaagccctga agtggttcct ggtcaacggc 4020
gccaacaact ggggctggga caagaaaacc ttcgatgtgc ggaccgccaa cgtgctggat 4080
agcgagttcc aggacatgtg cagagatatc gccgccgacc ctctgacctt tacccagtgg 4140
gtcaacgccg atagccccta tggattcctg gcctggtgct tcgagtacgc cagatacctg 4200
gacgccctgg atgagggaac ccaggatcag ttcatgaccc atctgcccgt gcaccaggat 4260
ggctcttgtt ctggcatcca gcactacagc gccatgctga gcgatgccgt gggagccaaa 4320
gccgtgaacc tgaagcctag cgacagcccc caggatatct atggcgctgt ggcccaggtg 4380
gtcatccaga aaaactacgc ctacatgaac gccgaggacg ccgagacatt cacaagcgga 4440
agcgtgacac tgacaggcgc cgagctgaga tctatggcct ctgcctggga catgatcggc 4500
atcacacggg gcctgaccaa aaagcctgtg atgacactgc cctacggcag caccagactg 4560
acctgtagag aaagcgtgat cgactacatc gtggacctgg aagagaaaga ggcccagaga 4620
gccattgccg agggcagaac agccaatcct gtgcacccct tcgacaacga ccggaaggat 4680
agcctgacac ctagcgccgc ctacaactac atgaccgccc tgatctggcc cagcatctct 4740
gaagtggtca aggcccctat cgtggccatg aagatgatca gacagctggc cagattcgcc 4800
gccaagagaa atgagggcct ggaataccct ctgcccaccg gctttatcct gcagcagaaa 4860
atcatggcca ccgacatgct gcgggtgtcc acatgtctga tgggcgagat caagatgagc 4920
ctgcagatcg agacagacgt ggtggacgag acagccatga tgggagccgc cgctcctaat 4980
tttgtgcacg gacacgatgc cagccacctg atcctgaccg tgtgcgatct ggtggacaag 5040
ggcatcacta gcgtggccgt gatccacgat agctttggaa cacacgccgg cagaaccgcc 5100
gacctgagag attctctgcg ggaagagatg gtcaagatgt accagaacca caacgccctg 5160
cagaacctgc tggacgtgca cgaagaaaga tggctggtgg acaccggcat ccaggtgcca 5220
gaacagggag agttcgacct gaacgagatc ctggtgtccg actactgctt cgcctaa 5277
<210> 11
<211> 1758
<212> PRT
<213> 人工序列
<220>
<223> C3P3-K1E嵌合酶的氨基酸序列，由非洲猪瘟病毒NP868R
加帽酶和突变型噬菌体K1E-RNA聚合酶融合构成
<400> 11
Met Glu Phe Ala Ser Leu Asp Asn Leu Val Ala Arg Tyr Gln Arg Cys
1 5 10 15
Phe Asn Asp Gln Ser Leu Lys Asn Ser Thr Ile Glu Leu Glu Ile Arg
20 25 30
Phe Gln Gln Ile Asn Phe Leu Leu Phe Lys Thr Val Tyr Glu Ala Leu
35 40 45
Val Ala Gln Glu Ile Pro Ser Thr Ile Ser His Ser Ile Arg Cys Ile
50 55 60
Lys Lys Val His His Glu Asn His Cys Arg Glu Lys Ile Leu Pro Ser
65 70 75 80
Glu Asn Leu Tyr Phe Lys Lys Gln Pro Leu Met Phe Phe Lys Phe Ser
85 90 95
Glu Pro Ala Ser Leu Gly Cys Lys Val Ser Leu Ala Ile Glu Gln Pro
100 105 110
Ile Arg Lys Phe Ile Leu Asp Ser Ser Val Leu Val Arg Leu Lys Asn
115 120 125
Arg Thr Thr Phe Arg Val Ser Glu Leu Trp Lys Ile Glu Leu Thr Ile
130 135 140
Val Lys Gln Leu Met Gly Ser Glu Val Ser Ala Lys Leu Ala Ala Phe
145 150 155 160
Lys Thr Leu Leu Phe Asp Thr Pro Glu Gln Gln Thr Thr Lys Asn Met
165 170 175
Met Thr Leu Ile Asn Pro Asp Asp Glu Tyr Leu Tyr Glu Ile Glu Ile
180 185 190
Glu Tyr Thr Gly Lys Pro Glu Ser Leu Thr Ala Ala Asp Val Ile Lys
195 200 205
Ile Lys Asn Thr Val Leu Thr Leu Ile Ser Pro Asn His Leu Met Leu
210 215 220
Thr Ala Tyr His Gln Ala Ile Glu Phe Ile Ala Ser His Ile Leu Ser
225 230 235 240
Ser Glu Ile Leu Leu Ala Arg Ile Lys Ser Gly Lys Trp Gly Leu Lys
245 250 255
Arg Leu Leu Pro Gln Val Lys Ser Met Thr Lys Ala Asp Tyr Met Lys
260 265 270
Phe Tyr Pro Pro Val Gly Tyr Tyr Val Thr Asp Lys Ala Asp Gly Ile
275 280 285
Arg Gly Ile Ala Val Ile Gln Asp Thr Gln Ile Tyr Val Val Ala Asp
290 295 300
Gln Leu Tyr Ser Leu Gly Thr Thr Gly Ile Glu Pro Leu Lys Pro Thr
305 310 315 320
Ile Leu Asp Gly Glu Phe Met Pro Glu Lys Lys Glu Phe Tyr Gly Phe
325 330 335
Asp Val Ile Met Tyr Glu Gly Asn Leu Leu Thr Gln Gln Gly Phe Glu
340 345 350
Thr Arg Ile Glu Ser Leu Ser Lys Gly Ile Lys Val Leu Gln Ala Phe
355 360 365
Asn Ile Lys Ala Glu Met Lys Pro Phe Ile Ser Leu Thr Ser Ala Asp
370 375 380
Pro Asn Val Leu Leu Lys Asn Phe Glu Ser Ile Phe Lys Lys Lys Thr
385 390 395 400
Arg Pro Tyr Ser Ile Asp Gly Ile Ile Leu Val Glu Pro Gly Asn Ser
405 410 415
Tyr Leu Asn Thr Asn Thr Phe Lys Trp Lys Pro Thr Trp Asp Asn Thr
420 425 430
Leu Asp Phe Leu Val Arg Lys Cys Pro Glu Ser Leu Asn Val Pro Glu
435 440 445
Tyr Ala Pro Lys Lys Gly Phe Ser Leu His Leu Leu Phe Val Gly Ile
450 455 460
Ser Gly Glu Leu Phe Lys Lys Leu Ala Leu Asn Trp Cys Pro Gly Tyr
465 470 475 480
Thr Lys Leu Phe Pro Val Thr Gln Arg Asn Gln Asn Tyr Phe Pro Val
485 490 495
Gln Phe Gln Pro Ser Asp Phe Pro Leu Ala Phe Leu Tyr Tyr His Pro
500 505 510
Asp Thr Ser Ser Phe Ser Asn Ile Asp Gly Lys Val Leu Glu Met Arg
515 520 525
Cys Leu Lys Arg Glu Ile Asn Tyr Val Arg Trp Glu Ile Val Lys Ile
530 535 540
Arg Glu Asp Arg Gln Gln Asp Leu Lys Thr Gly Gly Tyr Phe Gly Asn
545 550 555 560
Asp Phe Lys Thr Ala Glu Leu Thr Trp Leu Asn Tyr Met Asp Pro Phe
565 570 575
Ser Phe Glu Glu Leu Ala Lys Gly Pro Ser Gly Met Tyr Phe Ala Gly
580 585 590
Ala Lys Thr Gly Ile Tyr Arg Ala Gln Thr Ala Leu Ile Ser Phe Ile
595 600 605
Lys Gln Glu Ile Ile Gln Lys Ile Ser His Gln Ser Trp Val Ile Asp
610 615 620
Leu Gly Ile Gly Lys Gly Gln Asp Leu Gly Arg Tyr Leu Asp Ala Gly
625 630 635 640
Val Arg His Leu Val Gly Ile Asp Lys Asp Gln Thr Ala Leu Ala Glu
645 650 655
Leu Val Tyr Arg Lys Phe Ser His Ala Thr Thr Arg Gln His Lys His
660 665 670
Ala Thr Asn Ile Tyr Val Leu His Gln Asp Leu Ala Glu Pro Ala Lys
675 680 685
Glu Ile Ser Glu Lys Val His Gln Ile Tyr Gly Phe Pro Lys Glu Gly
690 695 700
Ala Ser Ser Ile Val Ser Asn Leu Phe Ile His Tyr Leu Met Lys Asn
705 710 715 720
Thr Gln Gln Val Glu Asn Leu Ala Val Leu Cys His Lys Leu Leu Gln
725 730 735
Pro Gly Gly Met Val Trp Phe Thr Thr Met Leu Gly Glu Gln Val Leu
740 745 750
Glu Leu Leu His Glu Asn Arg Ile Glu Leu Asn Glu Val Trp Glu Ala
755 760 765
Arg Glu Asn Glu Val Val Lys Phe Ala Ile Lys Arg Leu Phe Lys Glu
770 775 780
Asp Ile Leu Gln Glu Thr Gly Gln Glu Ile Gly Val Leu Leu Pro Phe
785 790 795 800
Ser Asn Gly Asp Phe Tyr Asn Glu Tyr Leu Val Asn Thr Ala Phe Leu
805 810 815
Ile Lys Ile Phe Lys His His Gly Phe Ser Leu Val Gln Lys Gln Ser
820 825 830
Phe Lys Asp Trp Ile Pro Glu Phe Gln Asn Phe Ser Lys Ser Leu Tyr
835 840 845
Lys Ile Leu Thr Glu Ala Asp Lys Thr Trp Thr Ser Leu Phe Gly Phe
850 855 860
Ile Cys Leu Arg Lys Asn Leu Glu Gly Gly Gly Gly Ser Gly Gly Gly
865 870 875 880
Gly Ser Val Glu Gln Asp Leu His Ala Ile Gln Leu Gln Leu Glu Glu
885 890 895
Glu Met Phe Asn Gly Gly Ile Arg Arg Phe Glu Ala Asp Gln Gln Arg
900 905 910
Gln Ile Ala Ser Gly Asn Glu Ser Asp Thr Ala Trp Asn Arg Arg Leu
915 920 925
Leu Ser Glu Leu Ile Ala Pro Met Ala Glu Gly Ile Gln Ala Tyr Lys
930 935 940
Glu Glu Tyr Glu Gly Lys Arg Gly Arg Ala Pro Arg Ala Leu Ala Phe
945 950 955 960
Ile Asn Cys Val Gly Asn Glu Val Ala Ala Tyr Ile Thr Met Lys Ile
965 970 975
Val Met Asp Met Leu Asn Thr Asp Val Thr Leu Gln Ala Ile Ala Met
980 985 990
Asn Val Ala Asp Arg Ile Glu Asp Gln Val Arg Phe Ser Lys Leu Glu
995 1000 1005
Gly His Ala Ala Lys Tyr Phe Glu Lys Val Lys Lys Ser Leu Lys
1010 1015 1020
Ala Ser Lys Thr Lys Ser Tyr Arg His Ala His Asn Val Ala Val
1025 1030 1035
Val Ala Glu Lys Ser Val Ala Asp Arg Asp Ala Asp Phe Ser Arg
1040 1045 1050
Trp Glu Ala Trp Pro Lys Asp Thr Leu Leu Gln Ile Gly Met Thr
1055 1060 1065
Leu Leu Glu Ile Leu Glu Asn Ser Val Phe Phe Asn Gly Gln Pro
1070 1075 1080
Val Phe Leu Arg Thr Leu Arg Thr Asn Gly Gly Lys His Gly Val
1085 1090 1095
Tyr Tyr Leu Gln Thr Ser Glu His Val Gly Glu Trp Ile Thr Ala
1100 1105 1110
Phe Lys Glu His Val Ala Gln Leu Ser Pro Ala Tyr Ala Pro Cys
1115 1120 1125
Val Ile Pro Pro Arg Pro Trp Val Ser Pro Phe Asn Gly Gly Phe
1130 1135 1140
His Thr Glu Lys Val Ala Ser Arg Ile Arg Leu Val Lys Gly Asn
1145 1150 1155
Arg Glu His Val Arg Lys Leu Thr Lys Lys Gln Met Pro Ala Val
1160 1165 1170
Tyr Lys Ala Val Asn Ala Leu Gln Ala Thr Lys Trp Gln Val Asn
1175 1180 1185
Lys Glu Val Leu Gln Val Val Glu Asp Val Ile Arg Leu Asp Leu
1190 1195 1200
Gly Tyr Gly Val Pro Ser Phe Lys Pro Leu Ile Asp Arg Glu Asn
1205 1210 1215
Lys Pro Ala Asn Pro Val Pro Leu Glu Phe Gln His Leu Arg Gly
1220 1225 1230
Arg Glu Leu Lys Glu Met Leu Thr Pro Glu Gln Trp Gln Ala Phe
1235 1240 1245
Ile Asn Trp Lys Gly Glu Cys Thr Lys Leu Tyr Thr Ala Glu Thr
1250 1255 1260
Lys Arg Gly Ser Lys Ser Ala Ala Thr Val Arg Met Val Gly Gln
1265 1270 1275
Ala Arg Lys Tyr Ser Gln Phe Asp Ala Ile Tyr Phe Val Tyr Ala
1280 1285 1290
Leu Asp Ser Arg Ser Arg Val Tyr Ala Gln Ser Ser Thr Leu Ser
1295 1300 1305
Pro Gln Ser Asn Asp Leu Gly Lys Ala Leu Leu Arg Phe Thr Glu
1310 1315 1320
Gly Gln Arg Leu Asp Ser Ala Glu Ala Leu Lys Trp Phe Leu Val
1325 1330 1335
Asn Gly Ala Asn Asn Trp Gly Trp Asp Lys Lys Thr Phe Asp Val
1340 1345 1350
Arg Thr Ala Asn Val Leu Asp Ser Glu Phe Gln Asp Met Cys Arg
1355 1360 1365
Asp Ile Ala Ala Asp Pro Leu Thr Phe Thr Gln Trp Val Asn Ala
1370 1375 1380
Asp Ser Pro Tyr Gly Phe Leu Ala Trp Cys Phe Glu Tyr Ala Arg
1385 1390 1395
Tyr Leu Asp Ala Leu Asp Glu Gly Thr Gln Asp Gln Phe Met Thr
1400 1405 1410
His Leu Pro Val His Gln Asp Gly Ser Cys Ser Gly Ile Gln His
1415 1420 1425
Tyr Ser Ala Met Leu Ser Asp Ala Val Gly Ala Lys Ala Val Asn
1430 1435 1440
Leu Lys Pro Ser Asp Ser Pro Gln Asp Ile Tyr Gly Ala Val Ala
1445 1450 1455
Gln Val Val Ile Gln Lys Asn Tyr Ala Tyr Met Asn Ala Glu Asp
1460 1465 1470
Ala Glu Thr Phe Thr Ser Gly Ser Val Thr Leu Thr Gly Ala Glu
1475 1480 1485
Leu Arg Ser Met Ala Ser Ala Trp Asp Met Ile Gly Ile Thr Arg
1490 1495 1500
Gly Leu Thr Lys Lys Pro Val Met Thr Leu Pro Tyr Gly Ser Thr
1505 1510 1515
Arg Leu Thr Cys Arg Glu Ser Val Ile Asp Tyr Ile Val Asp Leu
1520 1525 1530
Glu Glu Lys Glu Ala Gln Arg Ala Ile Ala Glu Gly Arg Thr Ala
1535 1540 1545
Asn Pro Val His Pro Phe Asp Asn Asp Arg Lys Asp Ser Leu Thr
1550 1555 1560
Pro Ser Ala Ala Tyr Asn Tyr Met Thr Ala Leu Ile Trp Pro Ser
1565 1570 1575
Ile Ser Glu Val Val Lys Ala Pro Ile Val Ala Met Lys Met Ile
1580 1585 1590
Arg Gln Leu Ala Arg Phe Ala Ala Lys Arg Asn Glu Gly Leu Glu
1595 1600 1605
Tyr Pro Leu Pro Thr Gly Phe Ile Leu Gln Gln Lys Ile Met Ala
1610 1615 1620
Thr Asp Met Leu Arg Val Ser Thr Cys Leu Met Gly Glu Ile Lys
1625 1630 1635
Met Ser Leu Gln Ile Glu Thr Asp Val Val Asp Glu Thr Ala Met
1640 1645 1650
Met Gly Ala Ala Ala Pro Asn Phe Val His Gly His Asp Ala Ser
1655 1660 1665
His Leu Ile Leu Thr Val Cys Asp Leu Val Asp Lys Gly Ile Thr
1670 1675 1680
Ser Val Ala Val Ile His Asp Ser Phe Gly Thr His Ala Gly Arg
1685 1690 1695
Thr Ala Asp Leu Arg Asp Ser Leu Arg Glu Glu Met Val Lys Met
1700 1705 1710
Tyr Gln Asn His Asn Ala Leu Gln Asn Leu Leu Asp Val His Glu
1715 1720 1725
Glu Arg Trp Leu Val Asp Thr Gly Ile Gln Val Pro Glu Gln Gly
1730 1735 1740
Glu Phe Asp Leu Asn Glu Ile Leu Val Ser Asp Tyr Cys Phe Ala
1745 1750 1755
<210> 12
<211> 7929
<212> DNA
<213> 人工序列
<220>
<223> C3P3-T3嵌合酶的核苷酸序列，由非洲猪瘟病毒NP868R
加帽酶和突变型噬菌体T3-RNA聚合酶融合构成
<400> 12
atggaattcg ccagcctgga caacctggtg gccagatacc agcggtgctt caacgaccag 60
agcctgaaga acagcaccat cgagctggaa atccggttcc agcagatcaa cttcctgctg 120
ttcaagaccg tgtacgaggc cctggtcgcc caggaaatcc ccagcaccat cagccacagc 180
atccggtgca tcaagaaggt gcaccacgag aaccactgcc gggagaagat cctgcccagc 240
gagaacctgt acttcaagaa acagcccctg atgttcttca agttcagcga gcccgccagc 300
ctgggctgta aagtgtccct ggccatcgag cagcccatcc ggaagttcat cctggacagc 360
agcgtgctgg tccggctgaa gaaccggacc accttccggg tgtccgagct gtggaagatc 420
gagctgacca tcgtgaagca gctgatgggc agcgaggtgt cagccaagct ggccgccttc 480
aagaccctgc tgttcgacac ccccgagcag cagaccacca agaacatgat gaccctgatc 540
aaccccgacg acgagtacct gtacgagatc gagatcgagt acaccggcaa gcctgagagc 600
ctgacagccg ccgacgtgat caagatcaag aacaccgtgc tgacactgat cagccccaac 660
cacctgatgc tgaccgccta ccaccaggcc atcgagttta tcgccagcca catcctgagc 720
agcgagatcc tgctggcccg gatcaagagc ggcaagtggg gcctgaagag actgctgccc 780
caggtcaagt ccatgaccaa ggccgactac atgaagttct acccccccgt gggctactac 840
gtgaccgaca aggccgacgg catccggggc attgccgtga tccaggacac ccagatctac 900
gtggtggccg accagctgta cagcctgggc accaccggca tcgagcccct gaagcccacc 960
atcctggacg gcgagttcat gcccgagaag aaagagttct acggctttga cgtgatcatg 1020
tacgagggca acctgctgac ccagcagggc ttcgagacac ggatcgagag cctgagcaag 1080
ggcatcaagg tgctgcaggc cttcaacatc aaggccgaga tgaagccctt catcagcctg 1140
acctccgccg accccaacgt gctgctgaag aatttcgaga gcatcttcaa gaagaaaacc 1200
cggccctaca gcatcgacgg catcatcctg gtggagcccg gcaacagcta cctgaacacc 1260
aacaccttca agtggaagcc cacctgggac aacaccctgg actttctggt ccggaagtgc 1320
cccgagtccc tgaacgtgcc cgagtacgcc cccaagaagg gcttcagcct gcatctgctg 1380
ttcgtgggca tcagcggcga gctgtttaag aagctggccc tgaactggtg ccccggctac 1440
accaagctgt tccccgtgac ccagcggaac cagaactact tccccgtgca gttccagccc 1500
agcgacttcc ccctggcctt cctgtactac caccccgaca ccagcagctt cagcaacatc 1560
gatggcaagg tgctggaaat gcggtgcctg aagcgggaga tcaactacgt gcgctgggag 1620
atcgtgaaga tccgggagga ccggcagcag gatctgaaaa ccggcggcta cttcggcaac 1680
gacttcaaga ccgccgagct gacctggctg aactacatgg accccttcag cttcgaggaa 1740
ctggccaagg gacccagcgg catgtacttc gctggcgcca agaccggcat ctacagagcc 1800
cagaccgccc tgatcagctt catcaagcag gaaatcatcc agaagatcag ccaccagagc 1860
tgggtgatcg acctgggcat cggcaagggc caggacctgg gcagatacct ggacgccggc 1920
gtgagacacc tggtcggcat cgataaggac cagacagccc tggccgagct ggtgtaccgg 1980
aagttctccc acgccaccac cagacagcac aagcacgcca ccaacatcta cgtgctgcac 2040
caggatctgg ccgagcctgc caaagaaatc agcgagaaag tgcaccagat ctatggcttc 2100
cccaaagagg gcgccagcag catcgtgtcc aacctgttca tccactacct gatgaagaac 2160
acccagcagg tcgagaacct ggctgtgctg tgccacaagc tgctgcagcc tggcggcatg 2220
gtctggttca ccaccatgct gggcgaacag gtgctggaac tgctgcacga gaaccggatc 2280
gaactgaacg aagtgtggga ggcccgggag aacgaggtgg tcaagttcgc catcaagcgg 2340
ctgttcaaag aggacatcct gcaggaaacc ggccaggaaa tcggcgtcct gctgcccttc 2400
agcaacggcg acttctacaa tgagtacctg gtcaacaccg cctttctgat caagattttc 2460
aagcaccatg gctttagcct cgtgcagaag cagagcttca aggactggat ccccgagttc 2520
cagaacttca gcaagagcct gtacaagatc ctgaccgagg ccgacaagac ctggaccagc 2580
ctgttcggct tcatctgcct gcggaagaac ctcgagggag gaggaggatc aggcggaggc 2640
ggaagtgtcg agatggaatt cgccagcctg gacaacctgg tggccagata ccagcggtgc 2700
ttcaacgacc agagcctgaa gaacagcacc atcgagctgg aaatccggtt ccagcagatc 2760
aacttcctgc tgttcaagac cgtgtacgag gccctggtcg cccaggaaat ccccagcacc 2820
atcagccaca gcatccggtg catcaagaag gtgcaccacg agaaccactg ccgggagaag 2880
atcctgccca gcgagaacct gtacttcaag aaacagcccc tgatgttctt caagttcagc 2940
gagcccgcca gcctgggctg taaagtgtcc ctggccatcg agcagcccat ccggaagttc 3000
atcctggaca gcagcgtgct ggtccggctg aagaaccgga ccaccttccg ggtgtccgag 3060
ctgtggaaga tcgagctgac catcgtgaag cagctgatgg gcagcgaggt gtcagccaag 3120
ctggccgcct tcaagaccct gctgttcgac acccccgagc agcagaccac caagaacatg 3180
atgaccctga tcaaccccga cgacgagtac ctgtacgaga tcgagatcga gtacaccggc 3240
aagcctgaga gcctgacagc cgccgacgtg atcaagatca agaacaccgt gctgacactg 3300
atcagcccca accacctgat gctgaccgcc taccaccagg ccatcgagtt tatcgccagc 3360
cacatcctga gcagcgagat cctgctggcc cggatcaaga gcggcaagtg gggcctgaag 3420
agactgctgc cccaggtcaa gtccatgacc aaggccgact acatgaagtt ctaccccccc 3480
gtgggctact acgtgaccga caaggccgac ggcatccggg gcattgccgt gatccaggac 3540
acccagatct acgtggtggc cgaccagctg tacagcctgg gcaccaccgg catcgagccc 3600
ctgaagccca ccatcctgga cggcgagttc atgcccgaga agaaagagtt ctacggcttt 3660
gacgtgatca tgtacgaggg caacctgctg acccagcagg gcttcgagac acggatcgag 3720
agcctgagca agggcatcaa ggtgctgcag gccttcaaca tcaaggccga gatgaagccc 3780
ttcatcagcc tgacctccgc cgaccccaac gtgctgctga agaatttcga gagcatcttc 3840
aagaagaaaa cccggcccta cagcatcgac ggcatcatcc tggtggagcc cggcaacagc 3900
tacctgaaca ccaacacctt caagtggaag cccacctggg acaacaccct ggactttctg 3960
gtccggaagt gccccgagtc cctgaacgtg cccgagtacg cccccaagaa gggcttcagc 4020
ctgcatctgc tgttcgtggg catcagcggc gagctgttta agaagctggc cctgaactgg 4080
tgccccggct acaccaagct gttccccgtg acccagcgga accagaacta cttccccgtg 4140
cagttccagc ccagcgactt ccccctggcc ttcctgtact accaccccga caccagcagc 4200
ttcagcaaca tcgatggcaa ggtgctggaa atgcggtgcc tgaagcggga gatcaactac 4260
gtgcgctggg agatcgtgaa gatccgggag gaccggcagc aggatctgaa aaccggcggc 4320
tacttcggca acgacttcaa gaccgccgag ctgacctggc tgaactacat ggaccccttc 4380
agcttcgagg aactggccaa gggacccagc ggcatgtact tcgctggcgc caagaccggc 4440
atctacagag cccagaccgc cctgatcagc ttcatcaagc aggaaatcat ccagaagatc 4500
agccaccaga gctgggtgat cgacctgggc atcggcaagg gccaggacct gggcagatac 4560
ctggacgccg gcgtgagaca cctggtcggc atcgataagg accagacagc cctggccgag 4620
ctggtgtacc ggaagttctc ccacgccacc accagacagc acaagcacgc caccaacatc 4680
tacgtgctgc accaggatct ggccgagcct gccaaagaaa tcagcgagaa agtgcaccag 4740
atctatggct tccccaaaga gggcgccagc agcatcgtgt ccaacctgtt catccactac 4800
ctgatgaaga acacccagca ggtcgagaac ctggctgtgc tgtgccacaa gctgctgcag 4860
cctggcggca tggtctggtt caccaccatg ctgggcgaac aggtgctgga actgctgcac 4920
gagaaccgga tcgaactgaa cgaagtgtgg gaggcccggg agaacgaggt ggtcaagttc 4980
gccatcaagc ggctgttcaa agaggacatc ctgcaggaaa ccggccagga aatcggcgtc 5040
ctgctgccct tcagcaacgg cgacttctac aatgagtacc tggtcaacac cgcctttctg 5100
atcaagattt tcaagcacca tggctttagc ctcgtgcaga agcagagctt caaggactgg 5160
atccccgagt tccagaactt cagcaagagc ctgtacaaga tcctgaccga ggccgacaag 5220
acctggacca gcctgttcgg cttcatctgc ctgcggaaga acctcgaggg aggaggagga 5280
tcaggcggag gcggaagtgt cgagcaggac ctgcacgcca tccagctgca gctcgaagag 5340
gaaatgttca acggcggcat cagaagattc gaggccgacc agcagagaca gatcgcctct 5400
ggcaacgaga gcgacaccgc ctggaataga aggctgctgt ctgagctgat cgcccctatg 5460
gccgaaggca tccaggccta caaagaggaa tacgagggca agagaggcag agcccctaga 5520
gccctggcct tcatcaactg tgtgggcaat gaggtggccg cctacatcac catgaagatc 5580
gtgatggaca tgctgaacac cgacgtgacc ctgcaggcca ttgccatgaa cgtggccgac 5640
agaatcgagg accaggtccg attcagcaag ctggaaggac acgccgccaa gtacttcgag 5700
aaagtgaaga agtccctgaa ggccagcaag accaagagct acagacacgc ccacaacgtg 5760
gccgtggtgg ccgaaaaatc tgtggccgat agggacgccg acttctctag atgggaggcc 5820
tggcctaagg acaccctgct gcagatcggc atgaccctgc tggaaatcct ggaaaacagc 5880
gtgttcttca acggccagcc cgtgttcctg agaaccctga ggacaaatgg cggcaagcac 5940
ggcgtgtact acctgcagac atctgagcac gtgggcgagt ggatcaccgc cttcaaagaa 6000
catgtggccc agctgagccc tgcctatgcc ccttgtgtga tccctcctag accctgggtg 6060
tcccctttca atggcggctt tcacaccgag aaggtggcca gcagaatcag actggtcaag 6120
ggcaaccggg aacacgtgcg gaagctgacc aagaaacaga tgcccgccgt gtacaaggcc 6180
gtgaatgctc tgcaggccac caagtggcag gtcaacaaag aggtgctgca ggtcgtcgag 6240
gacgtgatca gactggatct gggctacggc gtgccaagct ttaagcccct gatcgacaga 6300
gagaacaagc ccgccaaccc tgtgcccctg gaatttcagc acctgagagg ccgcgagctg 6360
aaagagatgc tgacacctga acagtggcag gcctttatca attggaaggg cgagtgcacc 6420
aagctgtaca ccgccgagac aaagaggggc tctaagtctg ccgccacagt gcgaatggtc 6480
ggacaggcca gaaagtacag ccagttcgac gccatctact tcgtgtacgc cctggacagc 6540
cggtctagag tgtatgccca gagcagcaca ctgagccccc agtctaacga tctgggaaag 6600
gccctgctga gattcaccga gggccagaga ctggattctg ccgaagccct gaagtggttc 6660
ctggtcaacg gcgccaacaa ctggggctgg gacaagaaaa ccttcgatgt gcggaccgcc 6720
aacgtgctgg atagcgagtt ccaggacatg tgcagagata tcgccgccga ccctctgacc 6780
tttacccagt gggtcaacgc cgatagcccc tatggattcc tggcctggtg cttcgagtac 6840
gccagatacc tggacgccct ggatgaggga acccaggatc agttcatgac ccatctgccc 6900
gtgcaccagg atggctcttg ttctggcatc cagcactaca gcgccatgct gagcgatgcc 6960
gtgggagcca aagccgtgaa cctgaagcct agcgacagcc cccaggatat ctatggcgct 7020
gtggcccagg tggtcatcca gaaaaactac gcctacatga acgccgagga cgccgagaca 7080
ttcacaagcg gaagcgtgac actgacaggc gccgagctga gatctatggc ctctgcctgg 7140
gacatgatcg gcatcacacg gggcctgacc aaaaagcctg tgatgacact gccctacggc 7200
agcaccagac tgacctgtag agaaagcgtg atcgactaca tcgtggacct ggaagagaaa 7260
gaggcccaga gagccattgc cgagggcaga acagccaatc ctgtgcaccc cttcgacaac 7320
gaccggaagg atagcctgac acctagcgcc gcctacaact acatgaccgc cctgatctgg 7380
cccagcatct ctgaagtggt caaggcccct atcgtggcca tgaagatgat cagacagctg 7440
gccagattcg ccgccaagag aaatgagggc ctggaatacc ctctgcccac cggctttatc 7500
ctgcagcaga aaatcatggc caccgacatg ctgcgggtgt ccacatgtct gatgggcgag 7560
atcaagatga gcctgcagat cgagacagac gtggtggacg agacagccat gatgggagcc 7620
gccgctccta attttgtgca cggacacgat gccagccacc tgatcctgac cgtgtgcgat 7680
ctggtggaca agggcatcac tagcgtggcc gtgatccacg atagctttgg aacacacgcc 7740
ggcagaaccg ccgacctgag agattctctg cgggaagaga tggtcaagat gtaccagaac 7800
cacaacgccc tgcagaacct gctggacgtg cacgaagaaa gatggctggt ggacaccggc 7860
atccaggtgc cagaacaggg agagttcgac ctgaacgaga tcctggtgtc cgactactgc 7920
ttcgcctaa 7929
<210> 13
<211> 2642
<212> PRT
<213> 人工序列
<220>
<223> C3P3-T3嵌合酶的氨基酸序列，由非洲猪瘟病毒NP868R
加帽酶和突变型噬菌体T3-RNA聚合酶融合构成
<400> 13
Met Glu Phe Ala Ser Leu Asp Asn Leu Val Ala Arg Tyr Gln Arg Cys
1 5 10 15
Phe Asn Asp Gln Ser Leu Lys Asn Ser Thr Ile Glu Leu Glu Ile Arg
20 25 30
Phe Gln Gln Ile Asn Phe Leu Leu Phe Lys Thr Val Tyr Glu Ala Leu
35 40 45
Val Ala Gln Glu Ile Pro Ser Thr Ile Ser His Ser Ile Arg Cys Ile
50 55 60
Lys Lys Val His His Glu Asn His Cys Arg Glu Lys Ile Leu Pro Ser
65 70 75 80
Glu Asn Leu Tyr Phe Lys Lys Gln Pro Leu Met Phe Phe Lys Phe Ser
85 90 95
Glu Pro Ala Ser Leu Gly Cys Lys Val Ser Leu Ala Ile Glu Gln Pro
100 105 110
Ile Arg Lys Phe Ile Leu Asp Ser Ser Val Leu Val Arg Leu Lys Asn
115 120 125
Arg Thr Thr Phe Arg Val Ser Glu Leu Trp Lys Ile Glu Leu Thr Ile
130 135 140
Val Lys Gln Leu Met Gly Ser Glu Val Ser Ala Lys Leu Ala Ala Phe
145 150 155 160
Lys Thr Leu Leu Phe Asp Thr Pro Glu Gln Gln Thr Thr Lys Asn Met
165 170 175
Met Thr Leu Ile Asn Pro Asp Asp Glu Tyr Leu Tyr Glu Ile Glu Ile
180 185 190
Glu Tyr Thr Gly Lys Pro Glu Ser Leu Thr Ala Ala Asp Val Ile Lys
195 200 205
Ile Lys Asn Thr Val Leu Thr Leu Ile Ser Pro Asn His Leu Met Leu
210 215 220
Thr Ala Tyr His Gln Ala Ile Glu Phe Ile Ala Ser His Ile Leu Ser
225 230 235 240
Ser Glu Ile Leu Leu Ala Arg Ile Lys Ser Gly Lys Trp Gly Leu Lys
245 250 255
Arg Leu Leu Pro Gln Val Lys Ser Met Thr Lys Ala Asp Tyr Met Lys
260 265 270
Phe Tyr Pro Pro Val Gly Tyr Tyr Val Thr Asp Lys Ala Asp Gly Ile
275 280 285
Arg Gly Ile Ala Val Ile Gln Asp Thr Gln Ile Tyr Val Val Ala Asp
290 295 300
Gln Leu Tyr Ser Leu Gly Thr Thr Gly Ile Glu Pro Leu Lys Pro Thr
305 310 315 320
Ile Leu Asp Gly Glu Phe Met Pro Glu Lys Lys Glu Phe Tyr Gly Phe
325 330 335
Asp Val Ile Met Tyr Glu Gly Asn Leu Leu Thr Gln Gln Gly Phe Glu
340 345 350
Thr Arg Ile Glu Ser Leu Ser Lys Gly Ile Lys Val Leu Gln Ala Phe
355 360 365
Asn Ile Lys Ala Glu Met Lys Pro Phe Ile Ser Leu Thr Ser Ala Asp
370 375 380
Pro Asn Val Leu Leu Lys Asn Phe Glu Ser Ile Phe Lys Lys Lys Thr
385 390 395 400
Arg Pro Tyr Ser Ile Asp Gly Ile Ile Leu Val Glu Pro Gly Asn Ser
405 410 415
Tyr Leu Asn Thr Asn Thr Phe Lys Trp Lys Pro Thr Trp Asp Asn Thr
420 425 430
Leu Asp Phe Leu Val Arg Lys Cys Pro Glu Ser Leu Asn Val Pro Glu
435 440 445
Tyr Ala Pro Lys Lys Gly Phe Ser Leu His Leu Leu Phe Val Gly Ile
450 455 460
Ser Gly Glu Leu Phe Lys Lys Leu Ala Leu Asn Trp Cys Pro Gly Tyr
465 470 475 480
Thr Lys Leu Phe Pro Val Thr Gln Arg Asn Gln Asn Tyr Phe Pro Val
485 490 495
Gln Phe Gln Pro Ser Asp Phe Pro Leu Ala Phe Leu Tyr Tyr His Pro
500 505 510
Asp Thr Ser Ser Phe Ser Asn Ile Asp Gly Lys Val Leu Glu Met Arg
515 520 525
Cys Leu Lys Arg Glu Ile Asn Tyr Val Arg Trp Glu Ile Val Lys Ile
530 535 540
Arg Glu Asp Arg Gln Gln Asp Leu Lys Thr Gly Gly Tyr Phe Gly Asn
545 550 555 560
Asp Phe Lys Thr Ala Glu Leu Thr Trp Leu Asn Tyr Met Asp Pro Phe
565 570 575
Ser Phe Glu Glu Leu Ala Lys Gly Pro Ser Gly Met Tyr Phe Ala Gly
580 585 590
Ala Lys Thr Gly Ile Tyr Arg Ala Gln Thr Ala Leu Ile Ser Phe Ile
595 600 605
Lys Gln Glu Ile Ile Gln Lys Ile Ser His Gln Ser Trp Val Ile Asp
610 615 620
Leu Gly Ile Gly Lys Gly Gln Asp Leu Gly Arg Tyr Leu Asp Ala Gly
625 630 635 640
Val Arg His Leu Val Gly Ile Asp Lys Asp Gln Thr Ala Leu Ala Glu
645 650 655
Leu Val Tyr Arg Lys Phe Ser His Ala Thr Thr Arg Gln His Lys His
660 665 670
Ala Thr Asn Ile Tyr Val Leu His Gln Asp Leu Ala Glu Pro Ala Lys
675 680 685
Glu Ile Ser Glu Lys Val His Gln Ile Tyr Gly Phe Pro Lys Glu Gly
690 695 700
Ala Ser Ser Ile Val Ser Asn Leu Phe Ile His Tyr Leu Met Lys Asn
705 710 715 720
Thr Gln Gln Val Glu Asn Leu Ala Val Leu Cys His Lys Leu Leu Gln
725 730 735
Pro Gly Gly Met Val Trp Phe Thr Thr Met Leu Gly Glu Gln Val Leu
740 745 750
Glu Leu Leu His Glu Asn Arg Ile Glu Leu Asn Glu Val Trp Glu Ala
755 760 765
Arg Glu Asn Glu Val Val Lys Phe Ala Ile Lys Arg Leu Phe Lys Glu
770 775 780
Asp Ile Leu Gln Glu Thr Gly Gln Glu Ile Gly Val Leu Leu Pro Phe
785 790 795 800
Ser Asn Gly Asp Phe Tyr Asn Glu Tyr Leu Val Asn Thr Ala Phe Leu
805 810 815
Ile Lys Ile Phe Lys His His Gly Phe Ser Leu Val Gln Lys Gln Ser
820 825 830
Phe Lys Asp Trp Ile Pro Glu Phe Gln Asn Phe Ser Lys Ser Leu Tyr
835 840 845
Lys Ile Leu Thr Glu Ala Asp Lys Thr Trp Thr Ser Leu Phe Gly Phe
850 855 860
Ile Cys Leu Arg Lys Asn Leu Glu Gly Gly Gly Gly Ser Gly Gly Gly
865 870 875 880
Gly Ser Val Glu Met Glu Phe Ala Ser Leu Asp Asn Leu Val Ala Arg
885 890 895
Tyr Gln Arg Cys Phe Asn Asp Gln Ser Leu Lys Asn Ser Thr Ile Glu
900 905 910
Leu Glu Ile Arg Phe Gln Gln Ile Asn Phe Leu Leu Phe Lys Thr Val
915 920 925
Tyr Glu Ala Leu Val Ala Gln Glu Ile Pro Ser Thr Ile Ser His Ser
930 935 940
Ile Arg Cys Ile Lys Lys Val His His Glu Asn His Cys Arg Glu Lys
945 950 955 960
Ile Leu Pro Ser Glu Asn Leu Tyr Phe Lys Lys Gln Pro Leu Met Phe
965 970 975
Phe Lys Phe Ser Glu Pro Ala Ser Leu Gly Cys Lys Val Ser Leu Ala
980 985 990
Ile Glu Gln Pro Ile Arg Lys Phe Ile Leu Asp Ser Ser Val Leu Val
995 1000 1005
Arg Leu Lys Asn Arg Thr Thr Phe Arg Val Ser Glu Leu Trp Lys
1010 1015 1020
Ile Glu Leu Thr Ile Val Lys Gln Leu Met Gly Ser Glu Val Ser
1025 1030 1035
Ala Lys Leu Ala Ala Phe Lys Thr Leu Leu Phe Asp Thr Pro Glu
1040 1045 1050
Gln Gln Thr Thr Lys Asn Met Met Thr Leu Ile Asn Pro Asp Asp
1055 1060 1065
Glu Tyr Leu Tyr Glu Ile Glu Ile Glu Tyr Thr Gly Lys Pro Glu
1070 1075 1080
Ser Leu Thr Ala Ala Asp Val Ile Lys Ile Lys Asn Thr Val Leu
1085 1090 1095
Thr Leu Ile Ser Pro Asn His Leu Met Leu Thr Ala Tyr His Gln
1100 1105 1110
Ala Ile Glu Phe Ile Ala Ser His Ile Leu Ser Ser Glu Ile Leu
1115 1120 1125
Leu Ala Arg Ile Lys Ser Gly Lys Trp Gly Leu Lys Arg Leu Leu
1130 1135 1140
Pro Gln Val Lys Ser Met Thr Lys Ala Asp Tyr Met Lys Phe Tyr
1145 1150 1155
Pro Pro Val Gly Tyr Tyr Val Thr Asp Lys Ala Asp Gly Ile Arg
1160 1165 1170
Gly Ile Ala Val Ile Gln Asp Thr Gln Ile Tyr Val Val Ala Asp
1175 1180 1185
Gln Leu Tyr Ser Leu Gly Thr Thr Gly Ile Glu Pro Leu Lys Pro
1190 1195 1200
Thr Ile Leu Asp Gly Glu Phe Met Pro Glu Lys Lys Glu Phe Tyr
1205 1210 1215
Gly Phe Asp Val Ile Met Tyr Glu Gly Asn Leu Leu Thr Gln Gln
1220 1225 1230
Gly Phe Glu Thr Arg Ile Glu Ser Leu Ser Lys Gly Ile Lys Val
1235 1240 1245
Leu Gln Ala Phe Asn Ile Lys Ala Glu Met Lys Pro Phe Ile Ser
1250 1255 1260
Leu Thr Ser Ala Asp Pro Asn Val Leu Leu Lys Asn Phe Glu Ser
1265 1270 1275
Ile Phe Lys Lys Lys Thr Arg Pro Tyr Ser Ile Asp Gly Ile Ile
1280 1285 1290
Leu Val Glu Pro Gly Asn Ser Tyr Leu Asn Thr Asn Thr Phe Lys
1295 1300 1305
Trp Lys Pro Thr Trp Asp Asn Thr Leu Asp Phe Leu Val Arg Lys
1310 1315 1320
Cys Pro Glu Ser Leu Asn Val Pro Glu Tyr Ala Pro Lys Lys Gly
1325 1330 1335
Phe Ser Leu His Leu Leu Phe Val Gly Ile Ser Gly Glu Leu Phe
1340 1345 1350
Lys Lys Leu Ala Leu Asn Trp Cys Pro Gly Tyr Thr Lys Leu Phe
1355 1360 1365
Pro Val Thr Gln Arg Asn Gln Asn Tyr Phe Pro Val Gln Phe Gln
1370 1375 1380
Pro Ser Asp Phe Pro Leu Ala Phe Leu Tyr Tyr His Pro Asp Thr
1385 1390 1395
Ser Ser Phe Ser Asn Ile Asp Gly Lys Val Leu Glu Met Arg Cys
1400 1405 1410
Leu Lys Arg Glu Ile Asn Tyr Val Arg Trp Glu Ile Val Lys Ile
1415 1420 1425
Arg Glu Asp Arg Gln Gln Asp Leu Lys Thr Gly Gly Tyr Phe Gly
1430 1435 1440
Asn Asp Phe Lys Thr Ala Glu Leu Thr Trp Leu Asn Tyr Met Asp
1445 1450 1455
Pro Phe Ser Phe Glu Glu Leu Ala Lys Gly Pro Ser Gly Met Tyr
1460 1465 1470
Phe Ala Gly Ala Lys Thr Gly Ile Tyr Arg Ala Gln Thr Ala Leu
1475 1480 1485
Ile Ser Phe Ile Lys Gln Glu Ile Ile Gln Lys Ile Ser His Gln
1490 1495 1500
Ser Trp Val Ile Asp Leu Gly Ile Gly Lys Gly Gln Asp Leu Gly
1505 1510 1515
Arg Tyr Leu Asp Ala Gly Val Arg His Leu Val Gly Ile Asp Lys
1520 1525 1530
Asp Gln Thr Ala Leu Ala Glu Leu Val Tyr Arg Lys Phe Ser His
1535 1540 1545
Ala Thr Thr Arg Gln His Lys His Ala Thr Asn Ile Tyr Val Leu
1550 1555 1560
His Gln Asp Leu Ala Glu Pro Ala Lys Glu Ile Ser Glu Lys Val
1565 1570 1575
His Gln Ile Tyr Gly Phe Pro Lys Glu Gly Ala Ser Ser Ile Val
1580 1585 1590
Ser Asn Leu Phe Ile His Tyr Leu Met Lys Asn Thr Gln Gln Val
1595 1600 1605
Glu Asn Leu Ala Val Leu Cys His Lys Leu Leu Gln Pro Gly Gly
1610 1615 1620
Met Val Trp Phe Thr Thr Met Leu Gly Glu Gln Val Leu Glu Leu
1625 1630 1635
Leu His Glu Asn Arg Ile Glu Leu Asn Glu Val Trp Glu Ala Arg
1640 1645 1650
Glu Asn Glu Val Val Lys Phe Ala Ile Lys Arg Leu Phe Lys Glu
1655 1660 1665
Asp Ile Leu Gln Glu Thr Gly Gln Glu Ile Gly Val Leu Leu Pro
1670 1675 1680
Phe Ser Asn Gly Asp Phe Tyr Asn Glu Tyr Leu Val Asn Thr Ala
1685 1690 1695
Phe Leu Ile Lys Ile Phe Lys His His Gly Phe Ser Leu Val Gln
1700 1705 1710
Lys Gln Ser Phe Lys Asp Trp Ile Pro Glu Phe Gln Asn Phe Ser
1715 1720 1725
Lys Ser Leu Tyr Lys Ile Leu Thr Glu Ala Asp Lys Thr Trp Thr
1730 1735 1740
Ser Leu Phe Gly Phe Ile Cys Leu Arg Lys Asn Leu Glu Gly Gly
1745 1750 1755
Gly Gly Ser Gly Gly Gly Gly Ser Val Glu Gln Asp Leu His Ala
1760 1765 1770
Ile Gln Leu Gln Leu Glu Glu Glu Met Phe Asn Gly Gly Ile Arg
1775 1780 1785
Arg Phe Glu Ala Asp Gln Gln Arg Gln Ile Ala Ser Gly Asn Glu
1790 1795 1800
Ser Asp Thr Ala Trp Asn Arg Arg Leu Leu Ser Glu Leu Ile Ala
1805 1810 1815
Pro Met Ala Glu Gly Ile Gln Ala Tyr Lys Glu Glu Tyr Glu Gly
1820 1825 1830
Lys Arg Gly Arg Ala Pro Arg Ala Leu Ala Phe Ile Asn Cys Val
1835 1840 1845
Gly Asn Glu Val Ala Ala Tyr Ile Thr Met Lys Ile Val Met Asp
1850 1855 1860
Met Leu Asn Thr Asp Val Thr Leu Gln Ala Ile Ala Met Asn Val
1865 1870 1875
Ala Asp Arg Ile Glu Asp Gln Val Arg Phe Ser Lys Leu Glu Gly
1880 1885 1890
His Ala Ala Lys Tyr Phe Glu Lys Val Lys Lys Ser Leu Lys Ala
1895 1900 1905
Ser Lys Thr Lys Ser Tyr Arg His Ala His Asn Val Ala Val Val
1910 1915 1920
Ala Glu Lys Ser Val Ala Asp Arg Asp Ala Asp Phe Ser Arg Trp
1925 1930 1935
Glu Ala Trp Pro Lys Asp Thr Leu Leu Gln Ile Gly Met Thr Leu
1940 1945 1950
Leu Glu Ile Leu Glu Asn Ser Val Phe Phe Asn Gly Gln Pro Val
1955 1960 1965
Phe Leu Arg Thr Leu Arg Thr Asn Gly Gly Lys His Gly Val Tyr
1970 1975 1980
Tyr Leu Gln Thr Ser Glu His Val Gly Glu Trp Ile Thr Ala Phe
1985 1990 1995
Lys Glu His Val Ala Gln Leu Ser Pro Ala Tyr Ala Pro Cys Val
2000 2005 2010
Ile Pro Pro Arg Pro Trp Val Ser Pro Phe Asn Gly Gly Phe His
2015 2020 2025
Thr Glu Lys Val Ala Ser Arg Ile Arg Leu Val Lys Gly Asn Arg
2030 2035 2040
Glu His Val Arg Lys Leu Thr Lys Lys Gln Met Pro Ala Val Tyr
2045 2050 2055
Lys Ala Val Asn Ala Leu Gln Ala Thr Lys Trp Gln Val Asn Lys
2060 2065 2070
Glu Val Leu Gln Val Val Glu Asp Val Ile Arg Leu Asp Leu Gly
2075 2080 2085
Tyr Gly Val Pro Ser Phe Lys Pro Leu Ile Asp Arg Glu Asn Lys
2090 2095 2100
Pro Ala Asn Pro Val Pro Leu Glu Phe Gln His Leu Arg Gly Arg
2105 2110 2115
Glu Leu Lys Glu Met Leu Thr Pro Glu Gln Trp Gln Ala Phe Ile
2120 2125 2130
Asn Trp Lys Gly Glu Cys Thr Lys Leu Tyr Thr Ala Glu Thr Lys
2135 2140 2145
Arg Gly Ser Lys Ser Ala Ala Thr Val Arg Met Val Gly Gln Ala
2150 2155 2160
Arg Lys Tyr Ser Gln Phe Asp Ala Ile Tyr Phe Val Tyr Ala Leu
2165 2170 2175
Asp Ser Arg Ser Arg Val Tyr Ala Gln Ser Ser Thr Leu Ser Pro
2180 2185 2190
Gln Ser Asn Asp Leu Gly Lys Ala Leu Leu Arg Phe Thr Glu Gly
2195 2200 2205
Gln Arg Leu Asp Ser Ala Glu Ala Leu Lys Trp Phe Leu Val Asn
2210 2215 2220
Gly Ala Asn Asn Trp Gly Trp Asp Lys Lys Thr Phe Asp Val Arg
2225 2230 2235
Thr Ala Asn Val Leu Asp Ser Glu Phe Gln Asp Met Cys Arg Asp
2240 2245 2250
Ile Ala Ala Asp Pro Leu Thr Phe Thr Gln Trp Val Asn Ala Asp
2255 2260 2265
Ser Pro Tyr Gly Phe Leu Ala Trp Cys Phe Glu Tyr Ala Arg Tyr
2270 2275 2280
Leu Asp Ala Leu Asp Glu Gly Thr Gln Asp Gln Phe Met Thr His
2285 2290 2295
Leu Pro Val His Gln Asp Gly Ser Cys Ser Gly Ile Gln His Tyr
2300 2305 2310
Ser Ala Met Leu Ser Asp Ala Val Gly Ala Lys Ala Val Asn Leu
2315 2320 2325
Lys Pro Ser Asp Ser Pro Gln Asp Ile Tyr Gly Ala Val Ala Gln
2330 2335 2340
Val Val Ile Gln Lys Asn Tyr Ala Tyr Met Asn Ala Glu Asp Ala
2345 2350 2355
Glu Thr Phe Thr Ser Gly Ser Val Thr Leu Thr Gly Ala Glu Leu
2360 2365 2370
Arg Ser Met Ala Ser Ala Trp Asp Met Ile Gly Ile Thr Arg Gly
2375 2380 2385
Leu Thr Lys Lys Pro Val Met Thr Leu Pro Tyr Gly Ser Thr Arg
2390 2395 2400
Leu Thr Cys Arg Glu Ser Val Ile Asp Tyr Ile Val Asp Leu Glu
2405 2410 2415
Glu Lys Glu Ala Gln Arg Ala Ile Ala Glu Gly Arg Thr Ala Asn
2420 2425 2430
Pro Val His Pro Phe Asp Asn Asp Arg Lys Asp Ser Leu Thr Pro
2435 2440 2445
Ser Ala Ala Tyr Asn Tyr Met Thr Ala Leu Ile Trp Pro Ser Ile
2450 2455 2460
Ser Glu Val Val Lys Ala Pro Ile Val Ala Met Lys Met Ile Arg
2465 2470 2475
Gln Leu Ala Arg Phe Ala Ala Lys Arg Asn Glu Gly Leu Glu Tyr
2480 2485 2490
Pro Leu Pro Thr Gly Phe Ile Leu Gln Gln Lys Ile Met Ala Thr
2495 2500 2505
Asp Met Leu Arg Val Ser Thr Cys Leu Met Gly Glu Ile Lys Met
2510 2515 2520
Ser Leu Gln Ile Glu Thr Asp Val Val Asp Glu Thr Ala Met Met
2525 2530 2535
Gly Ala Ala Ala Pro Asn Phe Val His Gly His Asp Ala Ser His
2540 2545 2550
Leu Ile Leu Thr Val Cys Asp Leu Val Asp Lys Gly Ile Thr Ser
2555 2560 2565
Val Ala Val Ile His Asp Ser Phe Gly Thr His Ala Gly Arg Thr
2570 2575 2580
Ala Asp Leu Arg Asp Ser Leu Arg Glu Glu Met Val Lys Met Tyr
2585 2590 2595
Gln Asn His Asn Ala Leu Gln Asn Leu Leu Asp Val His Glu Glu
2600 2605 2610
Arg Trp Leu Val Asp Thr Gly Ile Gln Val Pro Glu Gln Gly Glu
2615 2620 2625
Phe Asp Leu Asn Glu Ile Leu Val Ser Asp Tyr Cys Phe Ala
2630 2635 2640
<210> 14
<211> 1578
<212> DNA
<213> Saccharomyces cerevisiae
<400> 14
atgagtagtc agaaggtgtt tggcatcact ggaccagtaa gtacggtcgg tgctactgcc 60
gctgaaaaca agctgaatga ttctttaatc caggagctga aaaaggaagg ctcatttgaa 120
actgaacagg agaccgccaa tagggtccaa gtgctgaaaa ttctgcaaga actggctcag 180
cgcttcgttt acgaggttag taagaaaaag aatatgagcg acggcatggc ccgcgacgca 240
ggaggcaaaa ttttcaccta tggtagctat agactggggg tccatggtcc cggaagcgac 300
atcgatacct tagtcgtggt tcctaagcac gttacccgcg aggacttctt cacagtgttc 360
gactcactac tgagggaaag aaaagaatta gatgaaatcg cgcccgtgcc tgatgcgttt 420
gtgcccatta tcaagattaa gttctcaggc attagtattg atctgatatg cgcacgtctc 480
gatcagccac aggttccact gagcctgacc ttgagcgata aaaacctgtt aaggaaccta 540
gacgaaaagg accttagggc actgaatggt actagagtga ctgatgagat actggagcta 600
gtccccaagc ccaatgtttt caggatagcc ctccgcgcta tcaagctctg ggcacagaga 660
cgggcggtat acgctaatat atttggcttc cccggaggcg tcgcctgggc aatgctagtc 720
gctcgaattt gtcagctcta ccccaatgcc tgctcggcgg tcattctgaa tcgtttcttt 780
ataatcctgt cagaatggaa ttggccccaa cctgtgattc tcaagcctat tgaggatggc 840
ccactgcaag tgagagtctg gaaccctaaa atatacgccc aggacagatc ccatcggatg 900
cccgttatca cccctgctta cccttccatg tgcgctacac acaatattac ggaatccact 960
aaaaaagtga tattgcagga atttgtgaga ggtgtgcaaa tcaccaacga catcttttcc 1020
aacaaaaaat cttgggcaaa cctttttgag aagaatgact ttttctttcg ctacaagttt 1080
tatctggaaa tcaccgctta tacacgcggg tccgacgagc agcatctcaa atggtctggc 1140
ctggtagagt ctaaggtccg cttgttagtg atgaaactcg aggtgctagc tggcattaag 1200
atcgcccatc cctttaccaa gccctttgag tcttcctact gctgcccgac tgaggacgat 1260
tatgagatga tccaagacaa gtacggttcc cataaaactg aaacagctct gaacgccctg 1320
aaactcgtta ccgatgaaaa caaagaggaa gaatcaatta aggatgctcc caaggcgtac 1380
ctcagcacta tgtacatcgg gctcgacttc aatatcgaga ataagaagga gaaggtcgac 1440
atccatatcc cctgtaccga gttcgtgaac ctctgtcgga gctttaatga ggactacggg 1500
gatcacaagg tgttcaatct ggccctccgg ttcgtcaaag gatacgatct gcccgacgag 1560
gtattcgacg aaaatgag 1578
<210> 15
<211> 526
<212> PRT
<213> Saccharomyces cerevisiae
<400> 15
Met Ser Ser Gln Lys Val Phe Gly Ile Thr Gly Pro Val Ser Thr Val
1 5 10 15
Gly Ala Thr Ala Ala Glu Asn Lys Leu Asn Asp Ser Leu Ile Gln Glu
20 25 30
Leu Lys Lys Glu Gly Ser Phe Glu Thr Glu Gln Glu Thr Ala Asn Arg
35 40 45
Val Gln Val Leu Lys Ile Leu Gln Glu Leu Ala Gln Arg Phe Val Tyr
50 55 60
Glu Val Ser Lys Lys Lys Asn Met Ser Asp Gly Met Ala Arg Asp Ala
65 70 75 80
Gly Gly Lys Ile Phe Thr Tyr Gly Ser Tyr Arg Leu Gly Val His Gly
85 90 95
Pro Gly Ser Asp Ile Asp Thr Leu Val Val Val Pro Lys His Val Thr
100 105 110
Arg Glu Asp Phe Phe Thr Val Phe Asp Ser Leu Leu Arg Glu Arg Lys
115 120 125
Glu Leu Asp Glu Ile Ala Pro Val Pro Asp Ala Phe Val Pro Ile Ile
130 135 140
Lys Ile Lys Phe Ser Gly Ile Ser Ile Asp Leu Ile Cys Ala Arg Leu
145 150 155 160
Asp Gln Pro Gln Val Pro Leu Ser Leu Thr Leu Ser Asp Lys Asn Leu
165 170 175
Leu Arg Asn Leu Asp Glu Lys Asp Leu Arg Ala Leu Asn Gly Thr Arg
180 185 190
Val Thr Asp Glu Ile Leu Glu Leu Val Pro Lys Pro Asn Val Phe Arg
195 200 205
Ile Ala Leu Arg Ala Ile Lys Leu Trp Ala Gln Arg Arg Ala Val Tyr
210 215 220
Ala Asn Ile Phe Gly Phe Pro Gly Gly Val Ala Trp Ala Met Leu Val
225 230 235 240
Ala Arg Ile Cys Gln Leu Tyr Pro Asn Ala Cys Ser Ala Val Ile Leu
245 250 255
Asn Arg Phe Phe Ile Ile Leu Ser Glu Trp Asn Trp Pro Gln Pro Val
260 265 270
Ile Leu Lys Pro Ile Glu Asp Gly Pro Leu Gln Val Arg Val Trp Asn
275 280 285
Pro Lys Ile Tyr Ala Gln Asp Arg Ser His Arg Met Pro Val Ile Thr
290 295 300
Pro Ala Tyr Pro Ser Met Cys Ala Thr His Asn Ile Thr Glu Ser Thr
305 310 315 320
Lys Lys Val Ile Leu Gln Glu Phe Val Arg Gly Val Gln Ile Thr Asn
325 330 335
Asp Ile Phe Ser Asn Lys Lys Ser Trp Ala Asn Leu Phe Glu Lys Asn
340 345 350
Asp Phe Phe Phe Arg Tyr Lys Phe Tyr Leu Glu Ile Thr Ala Tyr Thr
355 360 365
Arg Gly Ser Asp Glu Gln His Leu Lys Trp Ser Gly Leu Val Glu Ser
370 375 380
Lys Val Arg Leu Leu Val Met Lys Leu Glu Val Leu Ala Gly Ile Lys
385 390 395 400
Ile Ala His Pro Phe Thr Lys Pro Phe Glu Ser Ser Tyr Cys Cys Pro
405 410 415
Thr Glu Asp Asp Tyr Glu Met Ile Gln Asp Lys Tyr Gly Ser His Lys
420 425 430
Thr Glu Thr Ala Leu Asn Ala Leu Lys Leu Val Thr Asp Glu Asn Lys
435 440 445
Glu Glu Glu Ser Ile Lys Asp Ala Pro Lys Ala Tyr Leu Ser Thr Met
450 455 460
Tyr Ile Gly Leu Asp Phe Asn Ile Glu Asn Lys Lys Glu Lys Val Asp
465 470 475 480
Ile His Ile Pro Cys Thr Glu Phe Val Asn Leu Cys Arg Ser Phe Asn
485 490 495
Glu Asp Tyr Gly Asp His Lys Val Phe Asn Leu Ala Leu Arg Phe Val
500 505 510
Lys Gly Tyr Asp Leu Pro Asp Glu Val Phe Asp Glu Asn Glu
515 520 525
<210> 16
<211> 1932
<212> DNA
<213> 小家鼠（Mus musculus）
<400> 16
atgcctttcg cagtcacaac ccagggagcc cagcagccag ccccagcacc taaacaattc 60
ggaatctcca gcccgatttc tttagctgcc cccaaagaca ccgatagaga attaacacag 120
aagctgattg aaactttgca gccctttggc gtcttcgagg aagaggagga gcttcaacgc 180
cgaatcttaa ttctgcagaa gttgaataac ctggtcaagg agtggatccg ggaaatatcc 240
gagtctagaa atctacctca ggcggtgatc gaaaacgtgg gaggcaaaat attcactttc 300
ggctcttacc gcctgggggt gcacacaaaa ggggcagaca ttgacgccct ttgcgtggct 360
cctaggcatg tcgatcggaa cgacttcttt actagtttct atgataaact taaacttcaa 420
gaggaggtga aagatctgcg ggcggtggaa gaggcgttcg tacctgttat caaactctgt 480
tttgacggga tcgagattga catccttttc gcccgactcg cattgcagac aatacctgag 540
gatcttgatc tgcgtgatga ttcactgtta aaaaatctgg atattaggtg cattcgcagc 600
ctgaatggat gtcgggttac agacgagatt ttgcacctgg tccctaacat cgattctttt 660
cgactgaccc ttagggctat caaactatgg gctaaatgtc ataatatcta tagcaatata 720
ctgggctttc ttggaggcgt cagctgggcc atgctggtcg ctagaacttg tcagctctat 780
cccaacgcca tcgcatctac tcttgtccgt aaattctttc tggtgttttc cgaatgggag 840
tggcctaatc cggtgctcct caaagagcca gaagagcgta accttaacct gccggtctgg 900
gacccccgtg taaatccttc cgataggtat cacctgatgc cgataattac ccctgcgtac 960
ccccagcaga acagtactta caacgtcagt gtgtctacaa ggatggtgat gatcgaggaa 1020
tttaagcaag gcctggctat cacgcacgag attctactta ataaggccga gtggtccaag 1080
ctgttcgagg cccctagttt ctttcagaag tataagcact acatcgtcct tttggcatct 1140
gcacccacag aaaagcaaca cctggaatgg gttgggctgg ttgagagcaa gattcgcatt 1200
ctggttggga gcttggagaa aaatgagttc attaccctgg ctcacgttaa cccccagtcc 1260
tttcccgccc caaaggagac cgccgacaag gaggagtttc gtacgatgtg ggtgataggt 1320
ctcgtgctga aaaaacccga gaacagcgag attctttcca tagacctgac ctatgacatc 1380
caaagtttca ccgacaccgt gtacagacag gcaataaact cgaaaatgtt tgagatggat 1440
atgaagattg ccgccatgca ccttagaaga aaggagctgc atcagctgct gcctaaccat 1500
gtactgcaga agaaagaaac ccacctgacc gagtccgtgc gactgaccgc cgtcaccgat 1560
agttccctgc ttctgtccat cgacagcgag aactcgatga ccgctccgtc accaacagga 1620
accatgaaaa ccgggcctct gacaggcaat ccacagggac gtaacagtcc cgctttggcc 1680
gttatggccg cgtcggtgac taacattcag ttccctgatg tgtctctgca acatgtaaat 1740
ccaatcgaaa gctctggcat cgccctgtct gagagcatac ctcaaatacc tagtcagcct 1800
acaatatccc ccccacccaa gcctactatg accagagtgg tgagttctac tcacttagtg 1860
aatcacccat cgagaccctc cggcaacacc gctacaaaca tccccaaccc cattctgggc 1920
gtcgggccct aa 1932
<210> 17
<211> 641
<212> PRT
<213> 小家鼠
<400> 17
Met Pro Phe Ala Val Thr Thr Gln Gly Ala Gln Gln Pro Ala Pro Ala
1 5 10 15
Pro Lys Gln Phe Gly Ile Ser Ser Pro Ile Ser Leu Ala Ala Pro Lys
20 25 30
Asp Thr Asp Arg Glu Leu Thr Gln Lys Leu Ile Glu Thr Leu Gln Pro
35 40 45
Phe Gly Val Phe Glu Glu Glu Glu Glu Leu Gln Arg Arg Ile Leu Ile
50 55 60
Leu Gln Lys Leu Asn Asn Leu Val Lys Glu Trp Ile Arg Glu Ile Ser
65 70 75 80
Glu Ser Arg Asn Leu Pro Gln Ala Val Ile Glu Asn Val Gly Gly Lys
85 90 95
Ile Phe Thr Phe Gly Ser Tyr Arg Leu Gly Val His Thr Lys Gly Ala
100 105 110
Asp Ile Asp Ala Leu Cys Val Ala Pro Arg His Val Asp Arg Asn Asp
115 120 125
Phe Phe Thr Ser Phe Tyr Asp Lys Leu Lys Leu Gln Glu Glu Val Lys
130 135 140
Asp Leu Arg Ala Val Glu Glu Ala Phe Val Pro Val Ile Lys Leu Cys
145 150 155 160
Phe Asp Gly Ile Glu Ile Asp Ile Leu Phe Ala Arg Leu Ala Leu Gln
165 170 175
Thr Ile Pro Glu Asp Leu Asp Leu Arg Asp Asp Ser Leu Leu Lys Asn
180 185 190
Leu Asp Ile Arg Cys Ile Arg Ser Leu Asn Gly Cys Arg Val Thr Asp
195 200 205
Glu Ile Leu His Leu Val Pro Asn Ile Asp Ser Phe Arg Leu Thr Leu
210 215 220
Arg Ala Ile Lys Leu Trp Ala Lys Cys His Asn Ile Tyr Ser Asn Ile
225 230 235 240
Leu Gly Phe Leu Gly Gly Val Ser Trp Ala Met Leu Val Ala Arg Thr
245 250 255
Cys Gln Leu Tyr Pro Asn Ala Ile Ala Ser Thr Leu Val Arg Lys Phe
260 265 270
Phe Leu Val Phe Ser Glu Trp Glu Trp Pro Asn Pro Val Leu Leu Lys
275 280 285
Glu Pro Glu Glu Arg Asn Leu Asn Leu Pro Val Trp Asp Pro Arg Val
290 295 300
Asn Pro Ser Asp Arg Tyr His Leu Met Pro Ile Ile Thr Pro Ala Tyr
305 310 315 320
Pro Gln Gln Asn Ser Thr Tyr Asn Val Ser Val Ser Thr Arg Met Val
325 330 335
Met Ile Glu Glu Phe Lys Gln Gly Leu Ala Ile Thr His Glu Ile Leu
340 345 350
Leu Asn Lys Ala Glu Trp Ser Lys Leu Phe Glu Ala Pro Ser Phe Phe
355 360 365
Gln Lys Tyr Lys His Tyr Ile Val Leu Leu Ala Ser Ala Pro Thr Glu
370 375 380
Lys Gln His Leu Glu Trp Val Gly Leu Val Glu Ser Lys Ile Arg Ile
385 390 395 400
Leu Val Gly Ser Leu Glu Lys Asn Glu Phe Ile Thr Leu Ala His Val
405 410 415
Asn Pro Gln Ser Phe Pro Ala Pro Lys Glu Thr Ala Asp Lys Glu Glu
420 425 430
Phe Arg Thr Met Trp Val Ile Gly Leu Val Leu Lys Lys Pro Glu Asn
435 440 445
Ser Glu Ile Leu Ser Ile Asp Leu Thr Tyr Asp Ile Gln Ser Phe Thr
450 455 460
Asp Thr Val Tyr Arg Gln Ala Ile Asn Ser Lys Met Phe Glu Met Asp
465 470 475 480
Met Lys Ile Ala Ala Met His Leu Arg Arg Lys Glu Leu His Gln Leu
485 490 495
Leu Pro Asn His Val Leu Gln Lys Lys Glu Thr His Leu Thr Glu Ser
500 505 510
Val Arg Leu Thr Ala Val Thr Asp Ser Ser Leu Leu Leu Ser Ile Asp
515 520 525
Ser Glu Asn Ser Met Thr Ala Pro Ser Pro Thr Gly Thr Met Lys Thr
530 535 540
Gly Pro Leu Thr Gly Asn Pro Gln Gly Arg Asn Ser Pro Ala Leu Ala
545 550 555 560
Val Met Ala Ala Ser Val Thr Asn Ile Gln Phe Pro Asp Val Ser Leu
565 570 575
Gln His Val Asn Pro Ile Glu Ser Ser Gly Ile Ala Leu Ser Glu Ser
580 585 590
Ile Pro Gln Ile Pro Ser Gln Pro Thr Ile Ser Pro Pro Pro Lys Pro
595 600 605
Thr Met Thr Arg Val Val Ser Ser Thr His Leu Val Asn His Pro Ser
610 615 620
Arg Pro Ser Gly Asn Thr Ala Thr Asn Ile Pro Asn Pro Ile Leu Gly
625 630 635 640
Val
<210> 18
<211> 1365
<212> DNA
<213> Escherichia coli
<400> 18
aaggttttgt cgagagaaga gtccgaggct gagcaggcgg tcgcccgacc tcaagtgaca 60
gtgattccga gggagcaaca tgcgatcagc cgcaaagata tctccgaaaa cgctctgaaa 120
gtgatgtata ggctcaataa agccggatac gaagcctggc ttgttggcgg cggtgtgcgc 180
gatctcctcc tgggaaagaa acccaaagat tttgacgtaa ccacaaacgc taccccagag 240
caggtgagaa aactgttccg aaattgccgt cttgtgggac gcagatttag actcgctcac 300
gtcatgttcg gcccagagat cattgaggtg gctactttta ggggtcatca tgagggcaat 360
gtttcggacc gaacgacttc acaacgaggg cagaacggta tgctcctccg ggataatatt 420
ttcggttcca tcgaggagga cgcccaacgg cgcgacttta ccattaacag cctgtactac 480
agcgtcgccg actttactgt cagggactat gtcgggggca tgaaagacct gaaggacggc 540
gtgatccgcc taattggtaa tcctgagact cggtatagag aggacccagt gcgtatgttg 600
agggctgtga ggttcgcagc aaagcttggt atgagaattt ccccagagac agctgagccc 660
attcctcgac tagcgacact gctgaacgac ataccacctg cccgattgtt tgaagaatcg 720
ttgaagctgc tacaggccgg gtatggctat gagacataca agctgctctg cgaataccac 780
ctgttccaac cgttgttccc aacaatcact agatacttta ccgagaatgg ggactctcca 840
atggagcgga ttatcgagca ggtgctgaag aacactgaca cacgtattca caatgacatg 900
cgcgtgaatc cagccttcct attcgccgcc atgttttggt atcctttact agaaacagcc 960
cagaagatcg cccaggagag cggtctgacc taccatgatg catttgccct cgcaatgaac 1020
gacgtgcttg acgaggcatg cagatccctg gccatcccta aacgtttgac tacactgaca 1080
agagacatct ggcaattgca attacggatg agtcgaagac aaggcaaacg cgcttggaag 1140
cttctggaac atcctaaatt tcgggccgcc tacgacctgt tagccttaag ggcagaagtc 1200
gaacgaaacg ctgagttaca gaggcttgtc aaatggtggg gtgaatttca ggtgtcagcc 1260
ccccctgatc agaaggggat gctaaacgag ctagatgaag aaccatcacc gcggcggaga 1320
acgcgtcgcc ctagaaaacg ggcgccaagg agagagggta cagcc 1365
<210> 19
<211> 455
<212> PRT
<213> Escherichia coli
<400> 19
Lys Val Leu Ser Arg Glu Glu Ser Glu Ala Glu Gln Ala Val Ala Arg
1 5 10 15
Pro Gln Val Thr Val Ile Pro Arg Glu Gln His Ala Ile Ser Arg Lys
20 25 30
Asp Ile Ser Glu Asn Ala Leu Lys Val Met Tyr Arg Leu Asn Lys Ala
35 40 45
Gly Tyr Glu Ala Trp Leu Val Gly Gly Gly Val Arg Asp Leu Leu Leu
50 55 60
Gly Lys Lys Pro Lys Asp Phe Asp Val Thr Thr Asn Ala Thr Pro Glu
65 70 75 80
Gln Val Arg Lys Leu Phe Arg Asn Cys Arg Leu Val Gly Arg Arg Phe
85 90 95
Arg Leu Ala His Val Met Phe Gly Pro Glu Ile Ile Glu Val Ala Thr
100 105 110
Phe Arg Gly His His Glu Gly Asn Val Ser Asp Arg Thr Thr Ser Gln
115 120 125
Arg Gly Gln Asn Gly Met Leu Leu Arg Asp Asn Ile Phe Gly Ser Ile
130 135 140
Glu Glu Asp Ala Gln Arg Arg Asp Phe Thr Ile Asn Ser Leu Tyr Tyr
145 150 155 160
Ser Val Ala Asp Phe Thr Val Arg Asp Tyr Val Gly Gly Met Lys Asp
165 170 175
Leu Lys Asp Gly Val Ile Arg Leu Ile Gly Asn Pro Glu Thr Arg Tyr
180 185 190
Arg Glu Asp Pro Val Arg Met Leu Arg Ala Val Arg Phe Ala Ala Lys
195 200 205
Leu Gly Met Arg Ile Ser Pro Glu Thr Ala Glu Pro Ile Pro Arg Leu
210 215 220
Ala Thr Leu Leu Asn Asp Ile Pro Pro Ala Arg Leu Phe Glu Glu Ser
225 230 235 240
Leu Lys Leu Leu Gln Ala Gly Tyr Gly Tyr Glu Thr Tyr Lys Leu Leu
245 250 255
Cys Glu Tyr His Leu Phe Gln Pro Leu Phe Pro Thr Ile Thr Arg Tyr
260 265 270
Phe Thr Glu Asn Gly Asp Ser Pro Met Glu Arg Ile Ile Glu Gln Val
275 280 285
Leu Lys Asn Thr Asp Thr Arg Ile His Asn Asp Met Arg Val Asn Pro
290 295 300
Ala Phe Leu Phe Ala Ala Met Phe Trp Tyr Pro Leu Leu Glu Thr Ala
305 310 315 320
Gln Lys Ile Ala Gln Glu Ser Gly Leu Thr Tyr His Asp Ala Phe Ala
325 330 335
Leu Ala Met Asn Asp Val Leu Asp Glu Ala Cys Arg Ser Leu Ala Ile
340 345 350
Pro Lys Arg Leu Thr Thr Leu Thr Arg Asp Ile Trp Gln Leu Gln Leu
355 360 365
Arg Met Ser Arg Arg Gln Gly Lys Arg Ala Trp Lys Leu Leu Glu His
370 375 380
Pro Lys Phe Arg Ala Ala Tyr Asp Leu Leu Ala Leu Arg Ala Glu Val
385 390 395 400
Glu Arg Asn Ala Glu Leu Gln Arg Leu Val Lys Trp Trp Gly Glu Phe
405 410 415
Gln Val Ser Ala Pro Pro Asp Gln Lys Gly Met Leu Asn Glu Leu Asp
420 425 430
Glu Glu Pro Ser Pro Arg Arg Arg Thr Arg Arg Pro Arg Lys Arg Ala
435 440 445
Pro Arg Arg Glu Gly Thr Ala
450 455
<210> 20
<211> 1440
<212> DNA
<213> 痘苗病毒
<400> 20
atgaatcgga atcccgacca aaatacattt cctaacatca cactgaaaat tatcgaaaca 60
tacctgggcc gcgtcccctc tgtgaatgaa taccacatgc tcaagctcca ggccagaaat 120
attcagaaga tcaccgtgtt caataaagac attttcgtct cactggtcaa aaaaaacaag 180
aagagatttt tttctgacgt tgacacctcc gccagtgaaa ttaaagacag gatactgagc 240
tatttctcta agcaaactca gacatacaac atcgggaagc tgtttacgat catagagctg 300
caaagcgtcc tagtgacaac atacaccgat atactcggtg tgctcacgat taaagccccc 360
aacgttatct cctcgaagat cagctacaat gttacaagca tggaggagct cgcaagagat 420
atgctcaata gcatgaatgt ggccgtgatc gataaggcga aagtcatggg gaggcacaat 480
gtcagctccc tcgtgaaaaa cgtcaacaag ctgatggagg agtacttgcg gcggcacaat 540
aagtcctgca tttgctatgg ctcctactcc ttgtacctca tcaatcccaa cattctttac 600
ggggacattg acatcctgca gaccaattct cgcacatttt tgattgatct cgcgttcctg 660
atcaagttca taaccggcaa caacattatc ctctccaaga tcccatatct gcgaaattat 720
atggtcatta aggacgagaa cgataaccac attatagact cgttcaacat ccgccaggac 780
acgatgaatg tggtgccgaa aattttcata gacaatatct atatcgtaga ccccactttt 840
cagctcctga acatgattaa aatgttttct cagattgaca ggctagaaga tctgtccaaa 900
gatccagaaa agtttaacgc tcgtatggct accatgctgg agtacgtccg gtacactcat 960
ggtatagttt ttgacggcaa gcggaataat atgcccatga aatgtatcat cgatgaaaat 1020
aaccgcatcg taacagtaac gacaaaagat tactttagtt tcaagaaatg cctggtatac 1080
cttgacgaga atgttctgtc atccgacatt ttagacctta acgcggacac atcatgcgat 1140
tttgagagtg tgactaactc cgtctatctt atacacgata atattatgta tacctacttt 1200
tccaacacga tcctgctttc cgacaagggt aaagtccatg aaatcagcgc caggggactc 1260
tgcgcccata tattgcttta tcagatgttg acatctggag aatacaaaca gtgtctgtcc 1320
gaccttctca attccatgat gaaccgagac aagatcccta tctactccca cacagagagg 1380
gataaaaagc ccggacggca tggattcatc aatatcgaga aggatatcat cgtgttttaa 1440
<210> 21
<211> 479
<212> PRT
<213> 痘苗病毒
<400> 21
Met Asn Arg Asn Pro Asp Gln Asn Thr Leu Pro Asn Ile Thr Leu Lys
1 5 10 15
Ile Ile Glu Thr Tyr Leu Gly Arg Val Pro Ser Val Asn Glu Tyr His
20 25 30
Met Leu Lys Leu Gln Ala Arg Asn Ile Gln Lys Ile Thr Val Phe Asn
35 40 45
Lys Asp Ile Phe Val Ser Leu Val Lys Lys Asn Lys Lys Arg Phe Phe
50 55 60
Ser Asp Val Asn Thr Ser Ala Ser Glu Ile Lys Asp Arg Ile Leu Ser
65 70 75 80
Tyr Phe Ser Lys Gln Thr Gln Thr Tyr Asn Ile Gly Lys Leu Phe Thr
85 90 95
Ile Ile Glu Leu Gln Ser Val Leu Val Thr Thr Tyr Thr Asp Ile Leu
100 105 110
Gly Val Leu Thr Ile Lys Ala Pro Asn Val Ile Ser Ser Lys Ile Ser
115 120 125
Tyr Asn Val Thr Ser Met Glu Glu Leu Ala Arg Asp Met Leu Asn Ser
130 135 140
Met Asn Val Ala Val Ile Asp Lys Ala Lys Val Met Gly Arg His Asn
145 150 155 160
Val Ser Ser Leu Val Lys Asn Val Asn Lys Leu Met Glu Glu Tyr Leu
165 170 175
Arg Arg His Asn Lys Ser Cys Ile Cys Tyr Gly Ser Tyr Ser Leu Tyr
180 185 190
Leu Ile Asn Pro Asn Ile Arg Tyr Gly Asp Ile Asp Ile Leu Gln Thr
195 200 205
Asn Ser Arg Thr Phe Leu Ile Asp Leu Ala Phe Leu Ile Lys Phe Ile
210 215 220
Thr Gly Asn Asn Ile Ile Leu Ser Lys Ile Pro Tyr Leu Arg Asn Tyr
225 230 235 240
Met Val Ile Lys Asp Glu Asn Asp Asn His Ile Ile Asp Ser Phe Asn
245 250 255
Ile Arg Gln Asp Thr Met Asn Val Val Pro Lys Ile Phe Ile Asp Asn
260 265 270
Ile Tyr Ile Val Asp Pro Thr Phe Gln Leu Leu Asn Met Ile Lys Met
275 280 285
Phe Ser Gln Ile Asp Arg Leu Glu Asp Leu Ser Lys Asp Pro Glu Lys
290 295 300
Phe Asn Ala Arg Met Ala Thr Met Leu Glu Tyr Val Arg Tyr Thr His
305 310 315 320
Gly Ile Val Phe Asp Gly Lys Arg Asn Asn Met Pro Met Lys Cys Ile
325 330 335
Ile Asp Glu Asn Asn Arg Ile Val Thr Val Thr Thr Lys Asp Tyr Phe
340 345 350
Ser Phe Lys Lys Cys Leu Val Tyr Leu Asp Glu Asn Val Leu Ser Ser
355 360 365
Asp Ile Leu Asp Leu Asn Ala Asp Thr Ser Cys Asp Phe Glu Ser Val
370 375 380
Thr Asn Ser Val Tyr Leu Ile His Asp Asn Ile Met Tyr Thr Tyr Phe
385 390 395 400
Ser Asn Thr Ile Leu Leu Ser Asp Lys Gly Lys Val His Glu Ile Ser
405 410 415
Ala Arg Gly Leu Cys Ala His Ile Leu Leu Tyr Gln Met Leu Thr Ser
420 425 430
Gly Glu Tyr Lys Gln Cys Leu Ser Asp Leu Leu Asn Ser Met Met Asn
435 440 445
Arg Asp Lys Ile Pro Ile Tyr Ser His Thr Glu Arg Asp Lys Lys Pro
450 455 460
Gly Arg His Gly Phe Ile Asn Ile Glu Lys Asp Ile Ile Val Phe
465 470 475
<210> 22
<211> 1761
<212> DNA
<213> Acanthamoeba polyphaga
<400> 22
atgctcaaaa acaagaccag ggctgagaag tatcagacct actacaccac caatgaatat 60
cagatcgtta aagaaaaact accagacatt ataagagacg cggaaattaa ggcgtctgaa 120
gtgctcgagc caaccatcta cgagaagcgc gcaatcatgg aagtcattaa ggatttcatt 180
cgggatcatc aaaggaaagt gtatggcgga acagccctga atgaggcatt gaaacaggtg 240
aatcccaagg atgccatcta tgataactat tccttcagcg acatcgagtt ttattcccct 300
acccccgtgc aggatctcgt ggatctctgc aacatcctgt atagaaaagg gtataagttc 360
gtccagggga aggacgctca gcatgaggaa acctattcta tctttgtaaa tttccagctc 420
tactgtgaca ttacctattc gccaacccgg gtcttttatg gtattaaaac gatagaaatt 480
gacggcatta actataccga tcctcatttc atgctcatag attacctccg aatggtgaac 540
cagcccttga ctgccgccgg ccagcgctgg gagaaagcgt tcgaacggat gtacaggctg 600
ctcaaagact atcccattga ggattttgac aagaggctgg atattcctga gccacccgaa 660
gaaatccaga gttatatttc tcggattaag accgagtttc tgagcgataa caagctgaat 720
gaaagcttcc tcatctccgg catcgaggct tacaacttct acattcgcca tgctgcctct 780
agcaaagatg aagaacagat ggcccggaca aaccgcaatg tggtcaatct taataacttt 840
attgcaaatg tcccctttag cgagctgatc tccgtgaact atcgcgaaga tgtcaagaat 900
acctataact tcctgcggat gatcgtcgag gataaagaga aaatcagtgt tgacgaatat 960
ttccctctct ttcaattcac tggctattcc actgtcatca aatacgatga tcaccccata 1020
attaggatct acgagggcga cggttattgt attcctaacg tcaagaccgt taaaacggtg 1080
gagaatgaca acggaacgaa gacaaagtac gagtacaagt acgtatcctt ccagtacgtc 1140
ctcatgattc tatatatcaa caaatttcgt gcgcacttgg acaagaataa gcctatgtat 1200
tttaactacg gtattgccat atccaatctg gtcaaagctc gcaatatata cctggaccag 1260
accgggaaaa gcgtccttga caacactgtg tttaaggagt tccgcactaa ctgtaccgga 1320
aatacgatct ctttcacacg gatgaacaga ctgagattac tcgagaaaag aaagcagggc 1380
aagcagactt cgttcgttta cacccctgaa gacttcttta agaaggatct ggaaacccaa 1440
gccaagcttg acccgtcgaa agcgagattc aaaaatacca gtggtaacaa gattatggtg 1500
ccaaagtacc tgctgttcaa aatagataac aacggaaata ttgaagataa catacatagc 1560
gaagaggcag aaatctcaga gaaagaagaa acttccggtg gctcttctat atccactgat 1620
aaatcattcg aagaatcacc taattcctcc cctaacagct ctcctaacaa ctcgttgaat 1680
aattctattg atatcagtac aaataattac gacgaccgct cggaaaacag cctggactca 1740
ctcacgtctg atgggcccta a 1761
<210> 23
<211> 584
<212> PRT
<213> Acanthamoeba polyphaga
<400> 23
Met Leu Lys Asn Lys Thr Arg Ala Glu Lys Tyr Gln Thr Tyr Tyr Thr
1 5 10 15
Thr Asn Glu Tyr Gln Ile Val Lys Glu Lys Leu Pro Asp Ile Ile Arg
20 25 30
Asp Ala Glu Ile Lys Ala Ser Glu Val Leu Glu Pro Thr Ile Tyr Glu
35 40 45
Lys Arg Ala Ile Met Glu Val Ile Lys Asp Phe Ile Arg Asp His Gln
50 55 60
Arg Lys Val Tyr Gly Gly Thr Ala Leu Asn Glu Ala Leu Lys Gln Val
65 70 75 80
Asn Pro Lys Asp Ala Ile Tyr Asp Asn Tyr Ser Phe Ser Asp Ile Glu
85 90 95
Phe Tyr Ser Pro Thr Pro Val Gln Asp Leu Val Asp Leu Cys Asn Ile
100 105 110
Leu Tyr Arg Lys Gly Tyr Lys Phe Val Gln Gly Lys Asp Ala Gln His
115 120 125
Glu Glu Thr Tyr Ser Ile Phe Val Asn Phe Gln Leu Tyr Cys Asp Ile
130 135 140
Thr Tyr Ser Pro Thr Arg Val Phe Tyr Gly Ile Lys Thr Ile Glu Ile
145 150 155 160
Asp Gly Ile Asn Tyr Thr Asp Pro His Phe Met Leu Ile Asp Tyr Leu
165 170 175
Arg Met Val Asn Gln Pro Leu Thr Ala Ala Gly Gln Arg Trp Glu Lys
180 185 190
Ala Phe Glu Arg Met Tyr Arg Leu Leu Lys Asp Tyr Pro Ile Glu Asp
195 200 205
Phe Asp Lys Arg Leu Asp Ile Pro Glu Pro Pro Glu Glu Ile Gln Ser
210 215 220
Tyr Ile Ser Arg Ile Lys Thr Glu Phe Leu Ser Asp Asn Lys Leu Asn
225 230 235 240
Glu Ser Phe Leu Ile Ser Gly Ile Glu Ala Tyr Asn Phe Tyr Ile Arg
245 250 255
His Ala Ala Ser Ser Lys Asp Glu Glu Gln Met Ala Arg Thr Asn Arg
260 265 270
Asn Val Val Asn Leu Asn Asn Phe Ile Ala Asn Val Pro Phe Ser Glu
275 280 285
Leu Ile Ser Val Asn Tyr Arg Glu Asp Val Lys Asn Thr Tyr Asn Phe
290 295 300
Leu Arg Met Ile Val Glu Asp Lys Glu Lys Ile Ser Val Asp Glu Tyr
305 310 315 320
Phe Pro Leu Phe Gln Phe Thr Gly Tyr Ser Thr Val Ile Lys Tyr Asp
325 330 335
Asp His Pro Ile Ile Arg Ile Tyr Glu Gly Asp Gly Tyr Cys Ile Pro
340 345 350
Asn Val Lys Thr Val Lys Thr Val Glu Asn Asp Asn Gly Thr Lys Thr
355 360 365
Lys Tyr Glu Tyr Lys Tyr Val Ser Phe Gln Tyr Val Leu Met Ile Leu
370 375 380
Tyr Ile Asn Lys Phe Arg Ala His Leu Asp Lys Asn Lys Pro Met Tyr
385 390 395 400
Phe Asn Tyr Gly Ile Ala Ile Ser Asn Leu Val Lys Ala Arg Asn Ile
405 410 415
Tyr Leu Asp Gln Thr Gly Lys Ser Val Leu Asp Asn Thr Val Phe Lys
420 425 430
Glu Phe Arg Thr Asn Cys Thr Gly Asn Thr Ile Ser Phe Thr Arg Met
435 440 445
Asn Arg Leu Arg Leu Leu Glu Lys Arg Lys Gln Gly Lys Gln Thr Ser
450 455 460
Phe Val Tyr Thr Pro Glu Asp Phe Phe Lys Lys Asp Leu Glu Thr Gln
465 470 475 480
Ala Lys Leu Asp Pro Ser Lys Ala Arg Phe Lys Asn Thr Ser Gly Asn
485 490 495
Lys Ile Met Val Pro Lys Tyr Leu Leu Phe Lys Ile Asp Asn Asn Gly
500 505 510
Asn Ile Glu Asp Asn Ile His Ser Glu Glu Ala Glu Ile Ser Glu Lys
515 520 525
Glu Glu Thr Ser Gly Gly Ser Ser Ile Ser Thr Asp Lys Ser Phe Glu
530 535 540
Glu Ser Pro Asn Ser Ser Pro Asn Ser Ser Pro Asn Asn Ser Leu Asn
545 550 555 560
Asn Ser Ile Asp Ile Ser Thr Asn Asn Tyr Asp Asp Arg Ser Glu Asn
565 570 575
Ser Leu Asp Ser Leu Thr Ser Asp
580
<210> 24
<211> 1593
<212> DNA
<213> Megavirus chilensis
<400> 24
atgtctgaaa ataggaatag gaagctctca taccaagaat attatgtaga cggggattac 60
gaggaggtcc gtaaaaagct gcccgagata attaaacagg ctagaatcaa agcctcccaa 120
gtgatggagc ccacaatcta tgagaagcgg gtggtgatgg aaataatcaa ggattttatt 180
cgggacaagg ggagaaaggt ctatgggggg accgcactca acgagaccat taagaagaag 240
aatcccgaag acgcgatcta cgacagctac ctgttttcag atatcgagtt ttactcacca 300
acacctgtgc ctgatctaaa ggagctgtgt gacattttat accacaaagg ctatgacccg 360
gtacagggaa aggaggcgca gcacgaggaa acttattcta tcttcgtcaa tctgcagttg 420
tattgtgaca tcacatacgt ccctactaaa gtatatcatg ggattaagac cattgaaatc 480
gatgggatca actacacaca cccgcatttc atgctgatcg attacctgcg gatgatcaac 540
cagccattaa cagcagcgga acagaggtgg gaaaaggcct tcgacaggat gtacgtgctg 600
ctgaagaatt atccgatgga gaaatatgat aattcaatgc gaatcaccag tccgcgggac 660
gatatccaaa tgtacatcgg aaaagtcaag tccgagttca tgaagatacc ggaaattcag 720
gaaagttgct taatctcagg ctttgacgca tacaactttt ttatcagaca cgcaatgggg 780
gacaggaagg tcgaacagat ggcccgcatc aaaaatgact acaacagcct gaagaacttc 840
attactgtgc tgccttttat ggagttaatc agtgtaaagt acaaagacac tgtggaaaag 900
ctctataatt ttctccggga gaaagtcgtt aacccagact tgatcactat cgatgagtac 960
ttcccgttgt tccagttcac tgggtattct gtctctatca actatgacgg cattccaatc 1020
gtgaaggttt acgaggcgga tggctattgc gtcccggata ttaagacgac ctcaggatac 1080
cgatacgtct cctatcagta tatcctgatg attatgtata tctccaaatt caaagctcac 1140
ctggataaga acaaggaaat gtattttaat tatggcatcg ccatttccaa tctggtacaa 1200
gctcgcaact cctatctgaa ccagaaaaat atcggtgtta ttaacgacac cgtttttagc 1260
gagttcagga taggctgtat aggcacaact gtctcctata caagaatgtc aagacttaga 1320
atgttggaga aaaagaaaca aggaaaggtt atccaatttg tctatacccc taaacagtac 1380
tttagtcaga cccccgagca gcagaacaat ttcgacgaga gcatgaagaa gtaccgcttt 1440
aaaaacactt ccggcaataa gattactatt cctaagaacc tgctgtttaa gatagacgag 1500
agaggaaata tttctgagga gatctcaacc gaggaagcct atataacaga agatactacc 1560
agcatcaata caaccacgga tatcaatact aat 1593
<210> 25
<211> 531
<212> PRT
<213> Megavirus chilensis
<400> 25
Met Ser Glu Asn Arg Asn Arg Lys Leu Ser Tyr Gln Glu Tyr Tyr Val
1 5 10 15
Asp Gly Asp Tyr Glu Glu Val Arg Lys Lys Leu Pro Glu Ile Ile Lys
20 25 30
Gln Ala Arg Ile Lys Ala Ser Gln Val Met Glu Pro Thr Ile Tyr Glu
35 40 45
Lys Arg Val Val Met Glu Ile Ile Lys Asp Phe Ile Arg Asp Lys Gly
50 55 60
Arg Lys Val Tyr Gly Gly Thr Ala Leu Asn Glu Thr Ile Lys Lys Lys
65 70 75 80
Asn Pro Glu Asp Ala Ile Tyr Asp Ser Tyr Leu Phe Ser Asp Ile Glu
85 90 95
Phe Tyr Ser Pro Thr Pro Val Pro Asp Leu Lys Glu Leu Cys Asp Ile
100 105 110
Leu Tyr His Lys Gly Tyr Asp Pro Val Gln Gly Lys Glu Ala Gln His
115 120 125
Glu Glu Thr Tyr Ser Ile Phe Val Asn Leu Gln Leu Tyr Cys Asp Ile
130 135 140
Thr Tyr Val Pro Thr Lys Val Tyr His Gly Ile Lys Thr Ile Glu Ile
145 150 155 160
Asp Gly Ile Asn Tyr Thr His Pro His Phe Met Leu Ile Asp Tyr Leu
165 170 175
Arg Met Ile Asn Gln Pro Leu Thr Ala Ala Glu Gln Arg Trp Glu Lys
180 185 190
Ala Phe Asp Arg Met Tyr Val Leu Leu Lys Asn Tyr Pro Met Glu Lys
195 200 205
Tyr Asp Asn Ser Met Arg Ile Thr Ser Pro Arg Asp Asp Ile Gln Met
210 215 220
Tyr Ile Gly Lys Val Lys Ser Glu Phe Met Lys Ile Pro Glu Ile Gln
225 230 235 240
Glu Ser Cys Leu Ile Ser Gly Phe Asp Ala Tyr Asn Phe Phe Ile Arg
245 250 255
His Ala Met Gly Asp Arg Lys Val Glu Gln Met Ala Arg Ile Lys Asn
260 265 270
Asp Tyr Asn Ser Leu Lys Asn Phe Ile Thr Val Leu Pro Phe Met Glu
275 280 285
Leu Ile Ser Val Lys Tyr Lys Asp Thr Val Glu Lys Leu Tyr Asn Phe
290 295 300
Leu Arg Glu Lys Val Val Asn Pro Asp Leu Ile Thr Ile Asp Glu Tyr
305 310 315 320
Phe Pro Leu Phe Gln Phe Thr Gly Tyr Ser Val Ser Ile Asn Tyr Asp
325 330 335
Gly Ile Pro Ile Val Lys Val Tyr Glu Ala Asp Gly Tyr Cys Val Pro
340 345 350
Asp Ile Lys Thr Thr Ser Gly Tyr Arg Tyr Val Ser Tyr Gln Tyr Ile
355 360 365
Leu Met Ile Met Tyr Ile Ser Lys Phe Lys Ala His Leu Asp Lys Asn
370 375 380
Lys Glu Met Tyr Phe Asn Tyr Gly Ile Ala Ile Ser Asn Leu Val Gln
385 390 395 400
Ala Arg Asn Ser Tyr Leu Asn Gln Lys Asn Ile Gly Val Ile Asn Asp
405 410 415
Thr Val Phe Ser Glu Phe Arg Ile Gly Cys Ile Gly Thr Thr Val Ser
420 425 430
Tyr Thr Arg Met Ser Arg Leu Arg Met Leu Glu Lys Lys Lys Gln Gly
435 440 445
Lys Val Ile Gln Phe Val Tyr Thr Pro Lys Gln Tyr Phe Ser Gln Thr
450 455 460
Pro Glu Gln Gln Asn Asn Phe Asp Glu Ser Met Lys Lys Tyr Arg Phe
465 470 475 480
Lys Asn Thr Ser Gly Asn Lys Ile Thr Ile Pro Lys Asn Leu Leu Phe
485 490 495
Lys Ile Asp Glu Arg Gly Asn Ile Ser Glu Glu Ile Ser Thr Glu Glu
500 505 510
Ala Tyr Ile Thr Glu Asp Thr Thr Ser Ile Asn Thr Thr Thr Asp Ile
515 520 525
Asn Thr Asn
530
<210> 26
<211> 1434
<212> DNA
<213> 非洲猪瘟病毒
<400> 26
atgagtagtc tcctcaagac tgatttcaac gtgtctaagt accgcctgat cgcgcagaag 60
cgggaggcaa atgccgttga aatcgaggct gcactcgaag tagtgaggga atttataatt 120
aagaagaaac tgatcctgta cggtgggatc gccatcgatt acgccctcca ccttaaagga 180
tcttccattt acccagaagg tgaaaggccc gacttcgaca tgttcagccc aaaccacgtc 240
gaggacgctt acgagctggc agacatactg tacgagaagg ggtttaaaca ggtgggtacg 300
gtgcgtgcaa tacacgttca gacgatgcgg gtgcgtactg acttcgtgtg ggtcgcagac 360
ttatcctaca tgccaccaaa tatctttaat acaatcccga cactgaccta caaaaacttg 420
aagattattc accccgacta tcagagggca gggctacatc ttgctttctg ctttccgttt 480
gataaccctc ctagggagga cgtttttagt cgatttaaga aagacttgca acgctataac 540
ctcattgaaa agtactaccc tatccccgtc gtgcctgtaa agtcgatcta cgaatcaaag 600
acattttcta tccccttcaa acaggttgcc atacacggtt ttgctgcata cgcgctcctg 660
tatcagactc tgaacgagtt acgtatcacc tgtaaagtac ctgagtggaa gaccgagttt 720
ccccagccta gctactccta tcacaagaac gacaaaaaca tcacacttac agtggacatg 780
cctaaagcct accctgcgct ggtgctggca acctacaatc ccgaagaggt gatcaaggag 840
atgggactgc atctgactga gatatgcgaa ccatacatgg actactcacc gcctattttc 900
aagacaaacg acattcactt tttttcgact atgtttaaag agctggccat cagcattatc 960
caagataacc tgattgtggt gagccctcaa tatttactgt tgtatttcct gtatggcgca 1020
ttcgccactc cagcggataa aagtctgttt ttattttatt ataacgccac actgtggatc 1080
ctggagaaag ccgactccct gttgaacatc attcagaagc agaccagccc cgaggagttc 1140
accagatttg ccaatacctc accattcgtc ctgaccacga gagtgctatc atgctcgcaa 1200
gagcgctgca cattctctcc agcatacaga atctctctgg ccaatgacgt acagcagtcc 1260
cagctcccat tgccgaagac acacttcctg agtaacagtt tgccggatgt gtcaactttg 1320
ccatataatt attaccctgg aaaagggaag gaccgaccca caaatttctc gtacgaaaag 1380
aatctgttat ttaatattgg gggaaagtgc actccctccg ctatggggcc ctaa 1434
<210> 27
<211> 475
<212> PRT
<213> 非洲猪瘟病毒
<400> 27
Met Ser Ser Leu Leu Lys Thr Asp Phe Asn Val Ser Lys Tyr Arg Leu
1 5 10 15
Ile Ala Gln Lys Arg Glu Ala Asn Ala Val Glu Ile Glu Ala Ala Leu
20 25 30
Glu Val Val Arg Glu Phe Ile Ile Lys Lys Lys Leu Ile Leu Tyr Gly
35 40 45
Gly Ile Ala Ile Asp Tyr Ala Leu His Leu Lys Gly Ser Ser Ile Tyr
50 55 60
Pro Glu Gly Glu Arg Pro Asp Phe Asp Met Phe Ser Pro Asn His Val
65 70 75 80
Glu Asp Ala Tyr Glu Leu Ala Asp Ile Leu Tyr Glu Lys Gly Phe Lys
85 90 95
Gln Val Gly Thr Val Arg Ala Ile His Val Gln Thr Met Arg Val Arg
100 105 110
Thr Asp Phe Val Trp Val Ala Asp Leu Ser Tyr Met Pro Pro Asn Ile
115 120 125
Phe Asn Thr Ile Pro Thr Leu Thr Tyr Lys Asn Leu Lys Ile Ile His
130 135 140
Pro Asp Tyr Gln Arg Ala Gly Leu His Leu Ala Phe Cys Phe Pro Phe
145 150 155 160
Asp Asn Pro Pro Arg Glu Asp Val Phe Ser Arg Phe Lys Lys Asp Leu
165 170 175
Gln Arg Tyr Asn Leu Ile Glu Lys Tyr Tyr Pro Ile Pro Val Val Pro
180 185 190
Val Lys Ser Ile Tyr Glu Ser Lys Thr Phe Ser Ile Pro Phe Lys Gln
195 200 205
Val Ala Ile His Gly Phe Ala Ala Tyr Ala Leu Leu Tyr Gln Thr Leu
210 215 220
Asn Glu Leu Arg Ile Thr Cys Lys Val Pro Glu Trp Lys Thr Glu Phe
225 230 235 240
Pro Gln Pro Ser Tyr Ser Tyr His Lys Asn Asp Lys Asn Ile Thr Leu
245 250 255
Thr Val Asp Met Pro Lys Ala Tyr Pro Ala Leu Val Leu Ala Thr Tyr
260 265 270
Asn Pro Glu Glu Val Ile Lys Glu Met Gly Leu His Leu Thr Glu Ile
275 280 285
Cys Glu Pro Tyr Met Asp Tyr Ser Pro Pro Ile Phe Lys Thr Asn Asp
290 295 300
Ile His Phe Phe Ser Thr Met Phe Lys Glu Leu Ala Ile Ser Ile Ile
305 310 315 320
Gln Asp Asn Leu Ile Val Val Ser Pro Gln Tyr Leu Leu Leu Tyr Phe
325 330 335
Leu Tyr Gly Ala Phe Ala Thr Pro Ala Asp Lys Ser Leu Phe Leu Phe
340 345 350
Tyr Tyr Asn Ala Thr Leu Trp Ile Leu Glu Lys Ala Asp Ser Leu Leu
355 360 365
Asn Ile Ile Gln Lys Gln Thr Ser Pro Glu Glu Phe Thr Arg Phe Ala
370 375 380
Asn Thr Ser Pro Phe Val Leu Thr Thr Arg Val Leu Ser Cys Ser Gln
385 390 395 400
Glu Arg Cys Thr Phe Ser Pro Ala Tyr Arg Ile Ser Leu Ala Asn Asp
405 410 415
Val Gln Gln Ser Gln Leu Pro Leu Pro Lys Thr His Phe Leu Ser Asn
420 425 430
Ser Leu Pro Asp Val Ser Thr Leu Pro Tyr Asn Tyr Tyr Pro Gly Lys
435 440 445
Gly Lys Asp Arg Pro Thr Asn Phe Ser Tyr Glu Lys Asn Leu Leu Phe
450 455 460
Asn Ile Gly Gly Lys Cys Thr Pro Ser Ala Met
465 470 475
<210> 28
<211> 141
<212> DNA
<213> 人工序列
<220>
<223> G4-EE1234L亮氨酸拉链的核苷酸序列
<400> 28
ggcggcggag gcctggaaat cgaggccgcc ttcctggaac aggaaaacac cgccctggaa 60
accgaggtgg ccgagctgga acaggaagtg cagcggctgg aaaacatcgt gtcccagtac 120
gagacaagat acggccccct g 141
<210> 29
<211> 47
<212> PRT
<213> 人工序列
<220>
<223> G4-EE1234L亮氨酸拉链的氨基酸序列
<400> 29
Gly Gly Gly Gly Leu Glu Ile Glu Ala Ala Phe Leu Glu Gln Glu Asn
1 5 10 15
Thr Ala Leu Glu Thr Glu Val Ala Glu Leu Glu Gln Glu Val Gln Arg
20 25 30
Leu Glu Asn Ile Val Ser Gln Tyr Glu Thr Arg Tyr Gly Pro Leu
35 40 45
<210> 30
<211> 141
<212> DNA
<213> 人工序列
<220>
<223> RR1234L-G4亮氨酸拉链的核苷酸序列
<400> 30
ctggaaatca gggccgcctt cctgcggaga agaaacaccg ccctgcggac cagagtggcc 60
gagctgagac agcgggtgca gcggctgcgg aacatcgtgt cccagtacga gacaagatac 120
ggccccctgg gcggaggcgg c 141
<210> 31
<211> 47
<212> PRT
<213> 人工序列
<220>
<223> RR1234L-G4亮氨酸拉链的氨基酸序列
<400> 31
Leu Glu Ile Arg Ala Ala Phe Leu Arg Arg Arg Asn Thr Ala Leu Arg
1 5 10 15
Thr Arg Val Ala Glu Leu Arg Gln Arg Val Gln Arg Leu Arg Asn Ile
20 25 30
Val Ser Gln Tyr Glu Thr Arg Tyr Gly Pro Leu Gly Gly Gly Gly
35 40 45
<210> 32
<211> 4467
<212> DNA
<213> 人工序列
<220>
<223> λ N-R341-G4-NP868R的核苷酸序列
<400> 32
atggatgctc agaccagacg cagagaacgg cgggcagaga agcaggcaca gtggaaggcc 60
gctaatggcg gtggcggcac cggtctcaaa aacaagacca gggctgagaa gtatcagacc 120
tactacacca ccaatgaata tcagatcgtt aaagaaaaac taccagacat tataagagac 180
gcggaaatta aggcgtctga agtgctcgag ccaaccatct acgagaagcg cgcaatcatg 240
gaagtcatta aggatttcat tcgggatcat caaaggaaag tgtatggcgg aacagccctg 300
aatgaggcat tgaaacaggt gaatcccaag gatgccatct atgataacta ttccttcagc 360
gacatcgagt tttattcccc tacccccgtg caggatctcg tggatctctg caacatcctg 420
tatagaaaag ggtataagtt cgtccagggg aaggacgctc agcatgagga aacctattct 480
atctttgtaa atttccagct ctactgtgac attacctatt cgccaacccg ggtcttttat 540
ggtattaaaa cgatagaaat tgacggcatt aactataccg atcctcattt catgctcata 600
gattacctcc gaatggtgaa ccagcccttg actgccgccg gccagcgctg ggagaaagcg 660
ttcgaacgga tgtacaggct gctcaaagac tatcccattg aggattttga caagaggctg 720
gatattcctg agccacccga agaaatccag agttatattt ctcggattaa gaccgagttt 780
ctgagcgata acaagctgaa tgaaagcttc ctcatctccg gcatcgaggc ttacaacttc 840
tacattcgcc atgctgcctc tagcaaagat gaagaacaga tggcccggac aaaccgcaat 900
gtggtcaatc ttaataactt tattgcaaat gtccccttta gcgagctgat ctccgtgaac 960
tatcgcgaag atgtcaagaa tacctataac ttcctgcgga tgatcgtcga ggataaagag 1020
aaaatcagtg ttgacgaata tttccctctc tttcaattca ctggctattc cactgtcatc 1080
aaatacgatg atcaccccat aattaggatc tacgagggcg acggttattg tattcctaac 1140
gtcaagaccg ttaaaacggt ggagaatgac aacggaacga agacaaagta cgagtacaag 1200
tacgtatcct tccagtacgt cctcatgatt ctatatatca acaaatttcg tgcgcacttg 1260
gacaagaata agcctatgta ttttaactac ggtattgcca tatccaatct ggtcaaagct 1320
cgcaatatat acctggacca gaccgggaaa agcgtccttg acaacactgt gtttaaggag 1380
ttccgcacta actgtaccgg aaatacgatc tctttcacac ggatgaacag actgagatta 1440
ctcgagaaaa gaaagcaggg caagcagact tcgttcgttt acacccctga agacttcttt 1500
aagaaggatc tggaaaccca agccaagctt gacccgtcga aagcgagatt caaaaatacc 1560
agtggtaaca agattatggt gccaaagtac ctgctgttca aaatagataa caacggaaat 1620
attgaagata acatacatag cgaagaggca gaaatctcag agaaagaaga aacttccggt 1680
ggctcttcta tatccactga taaatcattc gaagaatcac ctaattcctc ccctaacagc 1740
tctcctaaca actcgttgaa taattctatt gatatcagta caaataatta cgacgaccgc 1800
tcggaaaaca gcctggactc actcacgtct gatgggcccg gcggtggcgg cgaattcgcc 1860
agcctggaca acctggtggc cagataccag cggtgcttca acgaccagag cctgaagaac 1920
agcaccatcg agctggaaat ccggttccag cagatcaact tcctgctgtt caagaccgtg 1980
tacgaggccc tggtcgccca ggaaatcccc agcaccatca gccacagcat ccggtgcatc 2040
aagaaggtgc accacgagaa ccactgccgg gagaagatcc tgcccagcga gaacctgtac 2100
ttcaagaaac agcccctgat gttcttcaag ttcagcgagc ccgccagcct gggctgtaaa 2160
gtgtccctgg ccatcgagca gcccatccgg aagttcatcc tggacagcag cgtgctggtc 2220
cggctgaaga accggaccac cttccgggtg tccgagctgt ggaagatcga gctgaccatc 2280
gtgaagcagc tgatgggcag cgaggtgtca gccaagctgg ccgccttcaa gaccctgctg 2340
ttcgacaccc ccgagcagca gaccaccaag aacatgatga ccctgatcaa ccccgacgac 2400
gagtacctgt acgagatcga gatcgagtac accggcaagc ctgagagcct gacagccgcc 2460
gacgtgatca agatcaagaa caccgtgctg acactgatca gccccaacca cctgatgctg 2520
accgcctacc accaggccat cgagtttatc gccagccaca tcctgagcag cgagatcctg 2580
ctggcccgga tcaagagcgg caagtggggc ctgaagagac tgctgcccca ggtcaagtcc 2640
atgaccaagg ccgactacat gaagttctac ccccccgtgg gctactacgt gaccgacaag 2700
gccgacggca tccggggcat tgccgtgatc caggacaccc agatctacgt ggtggccgac 2760
cagctgtaca gcctgggcac caccggcatc gagcccctga agcccaccat cctggacggc 2820
gagttcatgc ccgagaagaa agagttctac ggctttgacg tgatcatgta cgagggcaac 2880
ctgctgaccc agcagggctt cgagacacgg atcgagagcc tgagcaaggg catcaaggtg 2940
ctgcaggcct tcaacatcaa ggccgagatg aagcccttca tcagcctgac ctccgccgac 3000
cccaacgtgc tgctgaagaa tttcgagagc atcttcaaga agaaaacccg gccctacagc 3060
atcgacggca tcatcctggt ggagcccggc aacagctacc tgaacaccaa caccttcaag 3120
tggaagccca cctgggacaa caccctggac tttctggtcc ggaagtgccc cgagtccctg 3180
aacgtgcccg agtacgcccc caagaagggc ttcagcctgc atctgctgtt cgtgggcatc 3240
agcggcgagc tgtttaagaa gctggccctg aactggtgcc ccggctacac caagctgttc 3300
cccgtgaccc agcggaacca gaactacttc cccgtgcagt tccagcccag cgacttcccc 3360
ctggccttcc tgtactacca ccccgacacc agcagcttca gcaacatcga tggcaaggtg 3420
ctggaaatgc ggtgcctgaa gcgggagatc aactacgtgc gctgggagat cgtgaagatc 3480
cgggaggacc ggcagcagga tctgaaaacc ggcggctact tcggcaacga cttcaagacc 3540
gccgagctga cctggctgaa ctacatggac cccttcagct tcgaggaact ggccaaggga 3600
cccagcggca tgtacttcgc tggcgccaag accggcatct acagagccca gaccgccctg 3660
atcagcttca tcaagcagga aatcatccag aagatcagcc accagagctg ggtgatcgac 3720
ctgggcatcg gcaagggcca ggacctgggc agatacctgg acgccggcgt gagacacctg 3780
gtcggcatcg ataaggacca gacagccctg gccgagctgg tgtaccggaa gttctcccac 3840
gccaccacca gacagcacaa gcacgccacc aacatctacg tgctgcacca ggatctggcc 3900
gagcctgcca aagaaatcag cgagaaagtg caccagatct atggcttccc caaagagggc 3960
gccagcagca tcgtgtccaa cctgttcatc cactacctga tgaagaacac ccagcaggtc 4020
gagaacctgg ctgtgctgtg ccacaagctg ctgcagcctg gcggcatggt ctggttcacc 4080
accatgctgg gcgaacaggt gctggaactg ctgcacgaga accggatcga actgaacgaa 4140
gtgtgggagg cccgggagaa cgaggtggtc aagttcgcca tcaagcggct gttcaaagag 4200
gacatcctgc aggaaaccgg ccaggaaatc ggcgtcctgc tgcccttcag caacggcgac 4260
ttctacaatg agtacctggt caacaccgcc tttctgatca agattttcaa gcaccatggc 4320
tttagcctcg tgcagaagca gagcttcaag gactggatcc ccgagttcca gaacttcagc 4380
aagagcctgt acaagatcct gaccgaggcc gacaagacct ggaccagcct gttcggcttc 4440
atctgcctgc ggaagaacgg gccctga 4467
<210> 33
<211> 1488
<212> PRT
<213> 人工序列
<220>
<223> λ N-R341-G4-NP868R的氨基酸序列
<400> 33
Met Asp Ala Gln Thr Arg Arg Arg Glu Arg Arg Ala Glu Lys Gln Ala
1 5 10 15
Gln Trp Lys Ala Ala Asn Gly Gly Gly Gly Thr Gly Leu Lys Asn Lys
20 25 30
Thr Arg Ala Glu Lys Tyr Gln Thr Tyr Tyr Thr Thr Asn Glu Tyr Gln
35 40 45
Ile Val Lys Glu Lys Leu Pro Asp Ile Ile Arg Asp Ala Glu Ile Lys
50 55 60
Ala Ser Glu Val Leu Glu Pro Thr Ile Tyr Glu Lys Arg Ala Ile Met
65 70 75 80
Glu Val Ile Lys Asp Phe Ile Arg Asp His Gln Arg Lys Val Tyr Gly
85 90 95
Gly Thr Ala Leu Asn Glu Ala Leu Lys Gln Val Asn Pro Lys Asp Ala
100 105 110
Ile Tyr Asp Asn Tyr Ser Phe Ser Asp Ile Glu Phe Tyr Ser Pro Thr
115 120 125
Pro Val Gln Asp Leu Val Asp Leu Cys Asn Ile Leu Tyr Arg Lys Gly
130 135 140
Tyr Lys Phe Val Gln Gly Lys Asp Ala Gln His Glu Glu Thr Tyr Ser
145 150 155 160
Ile Phe Val Asn Phe Gln Leu Tyr Cys Asp Ile Thr Tyr Ser Pro Thr
165 170 175
Arg Val Phe Tyr Gly Ile Lys Thr Ile Glu Ile Asp Gly Ile Asn Tyr
180 185 190
Thr Asp Pro His Phe Met Leu Ile Asp Tyr Leu Arg Met Val Asn Gln
195 200 205
Pro Leu Thr Ala Ala Gly Gln Arg Trp Glu Lys Ala Phe Glu Arg Met
210 215 220
Tyr Arg Leu Leu Lys Asp Tyr Pro Ile Glu Asp Phe Asp Lys Arg Leu
225 230 235 240
Asp Ile Pro Glu Pro Pro Glu Glu Ile Gln Ser Tyr Ile Ser Arg Ile
245 250 255
Lys Thr Glu Phe Leu Ser Asp Asn Lys Leu Asn Glu Ser Phe Leu Ile
260 265 270
Ser Gly Ile Glu Ala Tyr Asn Phe Tyr Ile Arg His Ala Ala Ser Ser
275 280 285
Lys Asp Glu Glu Gln Met Ala Arg Thr Asn Arg Asn Val Val Asn Leu
290 295 300
Asn Asn Phe Ile Ala Asn Val Pro Phe Ser Glu Leu Ile Ser Val Asn
305 310 315 320
Tyr Arg Glu Asp Val Lys Asn Thr Tyr Asn Phe Leu Arg Met Ile Val
325 330 335
Glu Asp Lys Glu Lys Ile Ser Val Asp Glu Tyr Phe Pro Leu Phe Gln
340 345 350
Phe Thr Gly Tyr Ser Thr Val Ile Lys Tyr Asp Asp His Pro Ile Ile
355 360 365
Arg Ile Tyr Glu Gly Asp Gly Tyr Cys Ile Pro Asn Val Lys Thr Val
370 375 380
Lys Thr Val Glu Asn Asp Asn Gly Thr Lys Thr Lys Tyr Glu Tyr Lys
385 390 395 400
Tyr Val Ser Phe Gln Tyr Val Leu Met Ile Leu Tyr Ile Asn Lys Phe
405 410 415
Arg Ala His Leu Asp Lys Asn Lys Pro Met Tyr Phe Asn Tyr Gly Ile
420 425 430
Ala Ile Ser Asn Leu Val Lys Ala Arg Asn Ile Tyr Leu Asp Gln Thr
435 440 445
Gly Lys Ser Val Leu Asp Asn Thr Val Phe Lys Glu Phe Arg Thr Asn
450 455 460
Cys Thr Gly Asn Thr Ile Ser Phe Thr Arg Met Asn Arg Leu Arg Leu
465 470 475 480
Leu Glu Lys Arg Lys Gln Gly Lys Gln Thr Ser Phe Val Tyr Thr Pro
485 490 495
Glu Asp Phe Phe Lys Lys Asp Leu Glu Thr Gln Ala Lys Leu Asp Pro
500 505 510
Ser Lys Ala Arg Phe Lys Asn Thr Ser Gly Asn Lys Ile Met Val Pro
515 520 525
Lys Tyr Leu Leu Phe Lys Ile Asp Asn Asn Gly Asn Ile Glu Asp Asn
530 535 540
Ile His Ser Glu Glu Ala Glu Ile Ser Glu Lys Glu Glu Thr Ser Gly
545 550 555 560
Gly Ser Ser Ile Ser Thr Asp Lys Ser Phe Glu Glu Ser Pro Asn Ser
565 570 575
Ser Pro Asn Ser Ser Pro Asn Asn Ser Leu Asn Asn Ser Ile Asp Ile
580 585 590
Ser Thr Asn Asn Tyr Asp Asp Arg Ser Glu Asn Ser Leu Asp Ser Leu
595 600 605
Thr Ser Asp Gly Pro Gly Gly Gly Gly Glu Phe Ala Ser Leu Asp Asn
610 615 620
Leu Val Ala Arg Tyr Gln Arg Cys Phe Asn Asp Gln Ser Leu Lys Asn
625 630 635 640
Ser Thr Ile Glu Leu Glu Ile Arg Phe Gln Gln Ile Asn Phe Leu Leu
645 650 655
Phe Lys Thr Val Tyr Glu Ala Leu Val Ala Gln Glu Ile Pro Ser Thr
660 665 670
Ile Ser His Ser Ile Arg Cys Ile Lys Lys Val His His Glu Asn His
675 680 685
Cys Arg Glu Lys Ile Leu Pro Ser Glu Asn Leu Tyr Phe Lys Lys Gln
690 695 700
Pro Leu Met Phe Phe Lys Phe Ser Glu Pro Ala Ser Leu Gly Cys Lys
705 710 715 720
Val Ser Leu Ala Ile Glu Gln Pro Ile Arg Lys Phe Ile Leu Asp Ser
725 730 735
Ser Val Leu Val Arg Leu Lys Asn Arg Thr Thr Phe Arg Val Ser Glu
740 745 750
Leu Trp Lys Ile Glu Leu Thr Ile Val Lys Gln Leu Met Gly Ser Glu
755 760 765
Val Ser Ala Lys Leu Ala Ala Phe Lys Thr Leu Leu Phe Asp Thr Pro
770 775 780
Glu Gln Gln Thr Thr Lys Asn Met Met Thr Leu Ile Asn Pro Asp Asp
785 790 795 800
Glu Tyr Leu Tyr Glu Ile Glu Ile Glu Tyr Thr Gly Lys Pro Glu Ser
805 810 815
Leu Thr Ala Ala Asp Val Ile Lys Ile Lys Asn Thr Val Leu Thr Leu
820 825 830
Ile Ser Pro Asn His Leu Met Leu Thr Ala Tyr His Gln Ala Ile Glu
835 840 845
Phe Ile Ala Ser His Ile Leu Ser Ser Glu Ile Leu Leu Ala Arg Ile
850 855 860
Lys Ser Gly Lys Trp Gly Leu Lys Arg Leu Leu Pro Gln Val Lys Ser
865 870 875 880
Met Thr Lys Ala Asp Tyr Met Lys Phe Tyr Pro Pro Val Gly Tyr Tyr
885 890 895
Val Thr Asp Lys Ala Asp Gly Ile Arg Gly Ile Ala Val Ile Gln Asp
900 905 910
Thr Gln Ile Tyr Val Val Ala Asp Gln Leu Tyr Ser Leu Gly Thr Thr
915 920 925
Gly Ile Glu Pro Leu Lys Pro Thr Ile Leu Asp Gly Glu Phe Met Pro
930 935 940
Glu Lys Lys Glu Phe Tyr Gly Phe Asp Val Ile Met Tyr Glu Gly Asn
945 950 955 960
Leu Leu Thr Gln Gln Gly Phe Glu Thr Arg Ile Glu Ser Leu Ser Lys
965 970 975
Gly Ile Lys Val Leu Gln Ala Phe Asn Ile Lys Ala Glu Met Lys Pro
980 985 990
Phe Ile Ser Leu Thr Ser Ala Asp Pro Asn Val Leu Leu Lys Asn Phe
995 1000 1005
Glu Ser Ile Phe Lys Lys Lys Thr Arg Pro Tyr Ser Ile Asp Gly
1010 1015 1020
Ile Ile Leu Val Glu Pro Gly Asn Ser Tyr Leu Asn Thr Asn Thr
1025 1030 1035
Phe Lys Trp Lys Pro Thr Trp Asp Asn Thr Leu Asp Phe Leu Val
1040 1045 1050
Arg Lys Cys Pro Glu Ser Leu Asn Val Pro Glu Tyr Ala Pro Lys
1055 1060 1065
Lys Gly Phe Ser Leu His Leu Leu Phe Val Gly Ile Ser Gly Glu
1070 1075 1080
Leu Phe Lys Lys Leu Ala Leu Asn Trp Cys Pro Gly Tyr Thr Lys
1085 1090 1095
Leu Phe Pro Val Thr Gln Arg Asn Gln Asn Tyr Phe Pro Val Gln
1100 1105 1110
Phe Gln Pro Ser Asp Phe Pro Leu Ala Phe Leu Tyr Tyr His Pro
1115 1120 1125
Asp Thr Ser Ser Phe Ser Asn Ile Asp Gly Lys Val Leu Glu Met
1130 1135 1140
Arg Cys Leu Lys Arg Glu Ile Asn Tyr Val Arg Trp Glu Ile Val
1145 1150 1155
Lys Ile Arg Glu Asp Arg Gln Gln Asp Leu Lys Thr Gly Gly Tyr
1160 1165 1170
Phe Gly Asn Asp Phe Lys Thr Ala Glu Leu Thr Trp Leu Asn Tyr
1175 1180 1185
Met Asp Pro Phe Ser Phe Glu Glu Leu Ala Lys Gly Pro Ser Gly
1190 1195 1200
Met Tyr Phe Ala Gly Ala Lys Thr Gly Ile Tyr Arg Ala Gln Thr
1205 1210 1215
Ala Leu Ile Ser Phe Ile Lys Gln Glu Ile Ile Gln Lys Ile Ser
1220 1225 1230
His Gln Ser Trp Val Ile Asp Leu Gly Ile Gly Lys Gly Gln Asp
1235 1240 1245
Leu Gly Arg Tyr Leu Asp Ala Gly Val Arg His Leu Val Gly Ile
1250 1255 1260
Asp Lys Asp Gln Thr Ala Leu Ala Glu Leu Val Tyr Arg Lys Phe
1265 1270 1275
Ser His Ala Thr Thr Arg Gln His Lys His Ala Thr Asn Ile Tyr
1280 1285 1290
Val Leu His Gln Asp Leu Ala Glu Pro Ala Lys Glu Ile Ser Glu
1295 1300 1305
Lys Val His Gln Ile Tyr Gly Phe Pro Lys Glu Gly Ala Ser Ser
1310 1315 1320
Ile Val Ser Asn Leu Phe Ile His Tyr Leu Met Lys Asn Thr Gln
1325 1330 1335
Gln Val Glu Asn Leu Ala Val Leu Cys His Lys Leu Leu Gln Pro
1340 1345 1350
Gly Gly Met Val Trp Phe Thr Thr Met Leu Gly Glu Gln Val Leu
1355 1360 1365
Glu Leu Leu His Glu Asn Arg Ile Glu Leu Asn Glu Val Trp Glu
1370 1375 1380
Ala Arg Glu Asn Glu Val Val Lys Phe Ala Ile Lys Arg Leu Phe
1385 1390 1395
Lys Glu Asp Ile Leu Gln Glu Thr Gly Gln Glu Ile Gly Val Leu
1400 1405 1410
Leu Pro Phe Ser Asn Gly Asp Phe Tyr Asn Glu Tyr Leu Val Asn
1415 1420 1425
Thr Ala Phe Leu Ile Lys Ile Phe Lys His His Gly Phe Ser Leu
1430 1435 1440
Val Gln Lys Gln Ser Phe Lys Asp Trp Ile Pro Glu Phe Gln Asn
1445 1450 1455
Phe Ser Lys Ser Leu Tyr Lys Ile Leu Thr Glu Ala Asp Lys Thr
1460 1465 1470
Trp Thr Ser Leu Phe Gly Phe Ile Cys Leu Arg Lys Asn Gly Pro
1475 1480 1485
<210> 34
<211> 4467
<212> DNA
<213> 人工序列
<220>
<223> λ N-NP868R-G4-R341的核苷酸序列
<400> 34
atggatgctc agaccagacg cagagaacgg cgggcagaga agcaggcaca gtggaaggcc 60
gctaatggcg gtggcggcac cggtgccagc ctggacaacc tggtggccag ataccagcgg 120
tgcttcaacg accagagcct gaagaacagc accatcgagc tggaaatccg gttccagcag 180
atcaacttcc tgctgttcaa gaccgtgtac gaggccctgg tcgcccagga aatccccagc 240
accatcagcc acagcatccg gtgcatcaag aaggtgcacc acgagaacca ctgccgggag 300
aagatcctgc ccagcgagaa cctgtacttc aagaaacagc ccctgatgtt cttcaagttc 360
agcgagcccg ccagcctggg ctgtaaagtg tccctggcca tcgagcagcc catccggaag 420
ttcatcctgg acagcagcgt gctggtccgg ctgaagaacc ggaccacctt ccgggtgtcc 480
gagctgtgga agatcgagct gaccatcgtg aagcagctga tgggcagcga ggtgtcagcc 540
aagctggccg ccttcaagac cctgctgttc gacacccccg agcagcagac caccaagaac 600
atgatgaccc tgatcaaccc cgacgacgag tacctgtacg agatcgagat cgagtacacc 660
ggcaagcctg agagcctgac agccgccgac gtgatcaaga tcaagaacac cgtgctgaca 720
ctgatcagcc ccaaccacct gatgctgacc gcctaccacc aggccatcga gtttatcgcc 780
agccacatcc tgagcagcga gatcctgctg gcccggatca agagcggcaa gtggggcctg 840
aagagactgc tgccccaggt caagtccatg accaaggccg actacatgaa gttctacccc 900
cccgtgggct actacgtgac cgacaaggcc gacggcatcc ggggcattgc cgtgatccag 960
gacacccaga tctacgtggt ggccgaccag ctgtacagcc tgggcaccac cggcatcgag 1020
cccctgaagc ccaccatcct ggacggcgag ttcatgcccg agaagaaaga gttctacggc 1080
tttgacgtga tcatgtacga gggcaacctg ctgacccagc agggcttcga gacacggatc 1140
gagagcctga gcaagggcat caaggtgctg caggccttca acatcaaggc cgagatgaag 1200
cccttcatca gcctgacctc cgccgacccc aacgtgctgc tgaagaattt cgagagcatc 1260
ttcaagaaga aaacccggcc ctacagcatc gacggcatca tcctggtgga gcccggcaac 1320
agctacctga acaccaacac cttcaagtgg aagcccacct gggacaacac cctggacttt 1380
ctggtccgga agtgccccga gtccctgaac gtgcccgagt acgcccccaa gaagggcttc 1440
agcctgcatc tgctgttcgt gggcatcagc ggcgagctgt ttaagaagct ggccctgaac 1500
tggtgccccg gctacaccaa gctgttcccc gtgacccagc ggaaccagaa ctacttcccc 1560
gtgcagttcc agcccagcga cttccccctg gccttcctgt actaccaccc cgacaccagc 1620
agcttcagca acatcgatgg caaggtgctg gaaatgcggt gcctgaagcg ggagatcaac 1680
tacgtgcgct gggagatcgt gaagatccgg gaggaccggc agcaggatct gaaaaccggc 1740
ggctacttcg gcaacgactt caagaccgcc gagctgacct ggctgaacta catggacccc 1800
ttcagcttcg aggaactggc caagggaccc agcggcatgt acttcgctgg cgccaagacc 1860
ggcatctaca gagcccagac cgccctgatc agcttcatca agcaggaaat catccagaag 1920
atcagccacc agagctgggt gatcgacctg ggcatcggca agggccagga cctgggcaga 1980
tacctggacg ccggcgtgag acacctggtc ggcatcgata aggaccagac agccctggcc 2040
gagctggtgt accggaagtt ctcccacgcc accaccagac agcacaagca cgccaccaac 2100
atctacgtgc tgcaccagga tctggccgag cctgccaaag aaatcagcga gaaagtgcac 2160
cagatctatg gcttccccaa agagggcgcc agcagcatcg tgtccaacct gttcatccac 2220
tacctgatga agaacaccca gcaggtcgag aacctggctg tgctgtgcca caagctgctg 2280
cagcctggcg gcatggtctg gttcaccacc atgctgggcg aacaggtgct ggaactgctg 2340
cacgagaacc ggatcgaact gaacgaagtg tgggaggccc gggagaacga ggtggtcaag 2400
ttcgccatca agcggctgtt caaagaggac atcctgcagg aaaccggcca ggaaatcggc 2460
gtcctgctgc ccttcagcaa cggcgacttc tacaatgagt acctggtcaa caccgccttt 2520
ctgatcaaga ttttcaagca ccatggcttt agcctcgtgc agaagcagag cttcaaggac 2580
tggatccccg agttccagaa cttcagcaag agcctgtaca agatcctgac cgaggccgac 2640
aagacctgga ccagcctgtt cggcttcatc tgcctgcgga agaacgggcc cggcggtggc 2700
ggcgaattcc tcaaaaacaa gaccagggct gagaagtatc agacctacta caccaccaat 2760
gaatatcaga tcgttaaaga aaaactacca gacattataa gagacgcgga aattaaggcg 2820
tctgaagtgc tcgagccaac catctacgag aagcgcgcaa tcatggaagt cattaaggat 2880
ttcattcggg atcatcaaag gaaagtgtat ggcggaacag ccctgaatga ggcattgaaa 2940
caggtgaatc ccaaggatgc catctatgat aactattcct tcagcgacat cgagttttat 3000
tcccctaccc ccgtgcagga tctcgtggat ctctgcaaca tcctgtatag aaaagggtat 3060
aagttcgtcc aggggaagga cgctcagcat gaggaaacct attctatctt tgtaaatttc 3120
cagctctact gtgacattac ctattcgcca acccgggtct tttatggtat taaaacgata 3180
gaaattgacg gcattaacta taccgatcct catttcatgc tcatagatta cctccgaatg 3240
gtgaaccagc ccttgactgc cgccggccag cgctgggaga aagcgttcga acggatgtac 3300
aggctgctca aagactatcc cattgaggat tttgacaaga ggctggatat tcctgagcca 3360
cccgaagaaa tccagagtta tatttctcgg attaagaccg agtttctgag cgataacaag 3420
ctgaatgaaa gcttcctcat ctccggcatc gaggcttaca acttctacat tcgccatgct 3480
gcctctagca aagatgaaga acagatggcc cggacaaacc gcaatgtggt caatcttaat 3540
aactttattg caaatgtccc ctttagcgag ctgatctccg tgaactatcg cgaagatgtc 3600
aagaatacct ataacttcct gcggatgatc gtcgaggata aagagaaaat cagtgttgac 3660
gaatatttcc ctctctttca attcactggc tattccactg tcatcaaata cgatgatcac 3720
cccataatta ggatctacga gggcgacggt tattgtattc ctaacgtcaa gaccgttaaa 3780
acggtggaga atgacaacgg aacgaagaca aagtacgagt acaagtacgt atccttccag 3840
tacgtcctca tgattctata tatcaacaaa tttcgtgcgc acttggacaa gaataagcct 3900
atgtatttta actacggtat tgccatatcc aatctggtca aagctcgcaa tatatacctg 3960
gaccagaccg ggaaaagcgt ccttgacaac actgtgttta aggagttccg cactaactgt 4020
accggaaata cgatctcttt cacacggatg aacagactga gattactcga gaaaagaaag 4080
cagggcaagc agacttcgtt cgtttacacc cctgaagact tctttaagaa ggatctggaa 4140
acccaagcca agcttgaccc gtcgaaagcg agattcaaaa ataccagtgg taacaagatt 4200
atggtgccaa agtacctgct gttcaaaata gataacaacg gaaatattga agataacata 4260
catagcgaag aggcagaaat ctcagagaaa gaagaaactt ccggtggctc ttctatatcc 4320
actgataaat cattcgaaga atcacctaat tcctccccta acagctctcc taacaactcg 4380
ttgaataatt ctattgatat cagtacaaat aattacgacg accgctcgga aaacagcctg 4440
gactcactca cgtctgatgg gccctga 4467
<210> 35
<211> 1488
<212> PRT
<213> 人工序列
<220>
<223> λ N-NP868R-G4-R341的氨基酸序列
<400> 35
Met Asp Ala Gln Thr Arg Arg Arg Glu Arg Arg Ala Glu Lys Gln Ala
1 5 10 15
Gln Trp Lys Ala Ala Asn Gly Gly Gly Gly Thr Gly Ala Ser Leu Asp
20 25 30
Asn Leu Val Ala Arg Tyr Gln Arg Cys Phe Asn Asp Gln Ser Leu Lys
35 40 45
Asn Ser Thr Ile Glu Leu Glu Ile Arg Phe Gln Gln Ile Asn Phe Leu
50 55 60
Leu Phe Lys Thr Val Tyr Glu Ala Leu Val Ala Gln Glu Ile Pro Ser
65 70 75 80
Thr Ile Ser His Ser Ile Arg Cys Ile Lys Lys Val His His Glu Asn
85 90 95
His Cys Arg Glu Lys Ile Leu Pro Ser Glu Asn Leu Tyr Phe Lys Lys
100 105 110
Gln Pro Leu Met Phe Phe Lys Phe Ser Glu Pro Ala Ser Leu Gly Cys
115 120 125
Lys Val Ser Leu Ala Ile Glu Gln Pro Ile Arg Lys Phe Ile Leu Asp
130 135 140
Ser Ser Val Leu Val Arg Leu Lys Asn Arg Thr Thr Phe Arg Val Ser
145 150 155 160
Glu Leu Trp Lys Ile Glu Leu Thr Ile Val Lys Gln Leu Met Gly Ser
165 170 175
Glu Val Ser Ala Lys Leu Ala Ala Phe Lys Thr Leu Leu Phe Asp Thr
180 185 190
Pro Glu Gln Gln Thr Thr Lys Asn Met Met Thr Leu Ile Asn Pro Asp
195 200 205
Asp Glu Tyr Leu Tyr Glu Ile Glu Ile Glu Tyr Thr Gly Lys Pro Glu
210 215 220
Ser Leu Thr Ala Ala Asp Val Ile Lys Ile Lys Asn Thr Val Leu Thr
225 230 235 240
Leu Ile Ser Pro Asn His Leu Met Leu Thr Ala Tyr His Gln Ala Ile
245 250 255
Glu Phe Ile Ala Ser His Ile Leu Ser Ser Glu Ile Leu Leu Ala Arg
260 265 270
Ile Lys Ser Gly Lys Trp Gly Leu Lys Arg Leu Leu Pro Gln Val Lys
275 280 285
Ser Met Thr Lys Ala Asp Tyr Met Lys Phe Tyr Pro Pro Val Gly Tyr
290 295 300
Tyr Val Thr Asp Lys Ala Asp Gly Ile Arg Gly Ile Ala Val Ile Gln
305 310 315 320
Asp Thr Gln Ile Tyr Val Val Ala Asp Gln Leu Tyr Ser Leu Gly Thr
325 330 335
Thr Gly Ile Glu Pro Leu Lys Pro Thr Ile Leu Asp Gly Glu Phe Met
340 345 350
Pro Glu Lys Lys Glu Phe Tyr Gly Phe Asp Val Ile Met Tyr Glu Gly
355 360 365
Asn Leu Leu Thr Gln Gln Gly Phe Glu Thr Arg Ile Glu Ser Leu Ser
370 375 380
Lys Gly Ile Lys Val Leu Gln Ala Phe Asn Ile Lys Ala Glu Met Lys
385 390 395 400
Pro Phe Ile Ser Leu Thr Ser Ala Asp Pro Asn Val Leu Leu Lys Asn
405 410 415
Phe Glu Ser Ile Phe Lys Lys Lys Thr Arg Pro Tyr Ser Ile Asp Gly
420 425 430
Ile Ile Leu Val Glu Pro Gly Asn Ser Tyr Leu Asn Thr Asn Thr Phe
435 440 445
Lys Trp Lys Pro Thr Trp Asp Asn Thr Leu Asp Phe Leu Val Arg Lys
450 455 460
Cys Pro Glu Ser Leu Asn Val Pro Glu Tyr Ala Pro Lys Lys Gly Phe
465 470 475 480
Ser Leu His Leu Leu Phe Val Gly Ile Ser Gly Glu Leu Phe Lys Lys
485 490 495
Leu Ala Leu Asn Trp Cys Pro Gly Tyr Thr Lys Leu Phe Pro Val Thr
500 505 510
Gln Arg Asn Gln Asn Tyr Phe Pro Val Gln Phe Gln Pro Ser Asp Phe
515 520 525
Pro Leu Ala Phe Leu Tyr Tyr His Pro Asp Thr Ser Ser Phe Ser Asn
530 535 540
Ile Asp Gly Lys Val Leu Glu Met Arg Cys Leu Lys Arg Glu Ile Asn
545 550 555 560
Tyr Val Arg Trp Glu Ile Val Lys Ile Arg Glu Asp Arg Gln Gln Asp
565 570 575
Leu Lys Thr Gly Gly Tyr Phe Gly Asn Asp Phe Lys Thr Ala Glu Leu
580 585 590
Thr Trp Leu Asn Tyr Met Asp Pro Phe Ser Phe Glu Glu Leu Ala Lys
595 600 605
Gly Pro Ser Gly Met Tyr Phe Ala Gly Ala Lys Thr Gly Ile Tyr Arg
610 615 620
Ala Gln Thr Ala Leu Ile Ser Phe Ile Lys Gln Glu Ile Ile Gln Lys
625 630 635 640
Ile Ser His Gln Ser Trp Val Ile Asp Leu Gly Ile Gly Lys Gly Gln
645 650 655
Asp Leu Gly Arg Tyr Leu Asp Ala Gly Val Arg His Leu Val Gly Ile
660 665 670
Asp Lys Asp Gln Thr Ala Leu Ala Glu Leu Val Tyr Arg Lys Phe Ser
675 680 685
His Ala Thr Thr Arg Gln His Lys His Ala Thr Asn Ile Tyr Val Leu
690 695 700
His Gln Asp Leu Ala Glu Pro Ala Lys Glu Ile Ser Glu Lys Val His
705 710 715 720
Gln Ile Tyr Gly Phe Pro Lys Glu Gly Ala Ser Ser Ile Val Ser Asn
725 730 735
Leu Phe Ile His Tyr Leu Met Lys Asn Thr Gln Gln Val Glu Asn Leu
740 745 750
Ala Val Leu Cys His Lys Leu Leu Gln Pro Gly Gly Met Val Trp Phe
755 760 765
Thr Thr Met Leu Gly Glu Gln Val Leu Glu Leu Leu His Glu Asn Arg
770 775 780
Ile Glu Leu Asn Glu Val Trp Glu Ala Arg Glu Asn Glu Val Val Lys
785 790 795 800
Phe Ala Ile Lys Arg Leu Phe Lys Glu Asp Ile Leu Gln Glu Thr Gly
805 810 815
Gln Glu Ile Gly Val Leu Leu Pro Phe Ser Asn Gly Asp Phe Tyr Asn
820 825 830
Glu Tyr Leu Val Asn Thr Ala Phe Leu Ile Lys Ile Phe Lys His His
835 840 845
Gly Phe Ser Leu Val Gln Lys Gln Ser Phe Lys Asp Trp Ile Pro Glu
850 855 860
Phe Gln Asn Phe Ser Lys Ser Leu Tyr Lys Ile Leu Thr Glu Ala Asp
865 870 875 880
Lys Thr Trp Thr Ser Leu Phe Gly Phe Ile Cys Leu Arg Lys Asn Gly
885 890 895
Pro Gly Gly Gly Gly Glu Phe Leu Lys Asn Lys Thr Arg Ala Glu Lys
900 905 910
Tyr Gln Thr Tyr Tyr Thr Thr Asn Glu Tyr Gln Ile Val Lys Glu Lys
915 920 925
Leu Pro Asp Ile Ile Arg Asp Ala Glu Ile Lys Ala Ser Glu Val Leu
930 935 940
Glu Pro Thr Ile Tyr Glu Lys Arg Ala Ile Met Glu Val Ile Lys Asp
945 950 955 960
Phe Ile Arg Asp His Gln Arg Lys Val Tyr Gly Gly Thr Ala Leu Asn
965 970 975
Glu Ala Leu Lys Gln Val Asn Pro Lys Asp Ala Ile Tyr Asp Asn Tyr
980 985 990
Ser Phe Ser Asp Ile Glu Phe Tyr Ser Pro Thr Pro Val Gln Asp Leu
995 1000 1005
Val Asp Leu Cys Asn Ile Leu Tyr Arg Lys Gly Tyr Lys Phe Val
1010 1015 1020
Gln Gly Lys Asp Ala Gln His Glu Glu Thr Tyr Ser Ile Phe Val
1025 1030 1035
Asn Phe Gln Leu Tyr Cys Asp Ile Thr Tyr Ser Pro Thr Arg Val
1040 1045 1050
Phe Tyr Gly Ile Lys Thr Ile Glu Ile Asp Gly Ile Asn Tyr Thr
1055 1060 1065
Asp Pro His Phe Met Leu Ile Asp Tyr Leu Arg Met Val Asn Gln
1070 1075 1080
Pro Leu Thr Ala Ala Gly Gln Arg Trp Glu Lys Ala Phe Glu Arg
1085 1090 1095
Met Tyr Arg Leu Leu Lys Asp Tyr Pro Ile Glu Asp Phe Asp Lys
1100 1105 1110
Arg Leu Asp Ile Pro Glu Pro Pro Glu Glu Ile Gln Ser Tyr Ile
1115 1120 1125
Ser Arg Ile Lys Thr Glu Phe Leu Ser Asp Asn Lys Leu Asn Glu
1130 1135 1140
Ser Phe Leu Ile Ser Gly Ile Glu Ala Tyr Asn Phe Tyr Ile Arg
1145 1150 1155
His Ala Ala Ser Ser Lys Asp Glu Glu Gln Met Ala Arg Thr Asn
1160 1165 1170
Arg Asn Val Val Asn Leu Asn Asn Phe Ile Ala Asn Val Pro Phe
1175 1180 1185
Ser Glu Leu Ile Ser Val Asn Tyr Arg Glu Asp Val Lys Asn Thr
1190 1195 1200
Tyr Asn Phe Leu Arg Met Ile Val Glu Asp Lys Glu Lys Ile Ser
1205 1210 1215
Val Asp Glu Tyr Phe Pro Leu Phe Gln Phe Thr Gly Tyr Ser Thr
1220 1225 1230
Val Ile Lys Tyr Asp Asp His Pro Ile Ile Arg Ile Tyr Glu Gly
1235 1240 1245
Asp Gly Tyr Cys Ile Pro Asn Val Lys Thr Val Lys Thr Val Glu
1250 1255 1260
Asn Asp Asn Gly Thr Lys Thr Lys Tyr Glu Tyr Lys Tyr Val Ser
1265 1270 1275
Phe Gln Tyr Val Leu Met Ile Leu Tyr Ile Asn Lys Phe Arg Ala
1280 1285 1290
His Leu Asp Lys Asn Lys Pro Met Tyr Phe Asn Tyr Gly Ile Ala
1295 1300 1305
Ile Ser Asn Leu Val Lys Ala Arg Asn Ile Tyr Leu Asp Gln Thr
1310 1315 1320
Gly Lys Ser Val Leu Asp Asn Thr Val Phe Lys Glu Phe Arg Thr
1325 1330 1335
Asn Cys Thr Gly Asn Thr Ile Ser Phe Thr Arg Met Asn Arg Leu
1340 1345 1350
Arg Leu Leu Glu Lys Arg Lys Gln Gly Lys Gln Thr Ser Phe Val
1355 1360 1365
Tyr Thr Pro Glu Asp Phe Phe Lys Lys Asp Leu Glu Thr Gln Ala
1370 1375 1380
Lys Leu Asp Pro Ser Lys Ala Arg Phe Lys Asn Thr Ser Gly Asn
1385 1390 1395
Lys Ile Met Val Pro Lys Tyr Leu Leu Phe Lys Ile Asp Asn Asn
1400 1405 1410
Gly Asn Ile Glu Asp Asn Ile His Ser Glu Glu Ala Glu Ile Ser
1415 1420 1425
Glu Lys Glu Glu Thr Ser Gly Gly Ser Ser Ile Ser Thr Asp Lys
1430 1435 1440
Ser Phe Glu Glu Ser Pro Asn Ser Ser Pro Asn Ser Ser Pro Asn
1445 1450 1455
Asn Ser Leu Asn Asn Ser Ile Asp Ile Ser Thr Asn Asn Tyr Asp
1460 1465 1470
Asp Arg Ser Glu Asn Ser Leu Asp Ser Leu Thr Ser Asp Gly Pro
1475 1480 1485
<210> 36
<211> 390
<212> DNA
<213> Bacteriophage MS2
<400> 36
atggcttcaa acttcacgca gtttgttctg gtcgacaacg gcgggaccgg agacgtgaca 60
gtcgcccctt ctaactttgc taatggagtc gctgaatgga tctccagtaa ttcccggtca 120
caagcctata aggttacatg ctccgtccgc caatccagcg cgcagaatcg taagtacact 180
ataaaggtag aggtaccgaa ggtagcgact cagaccgttg gaggggaaga actgcccgtc 240
gcggggtggc gctcctacct gaacatggag ttgaccatcc cgattttcgc gacaaactct 300
gactgtgaac tgattgtgaa ggccatgcag ggactgctga aggatgggaa ccctattccc 360
agcgccattg cggcaaattc agggatatat 390
<210> 37
<211> 130
<212> PRT
<213> Bacteriophage MS2
<400> 37
Met Ala Ser Asn Phe Thr Gln Phe Val Leu Val Asp Asn Gly Gly Thr
1 5 10 15
Gly Asp Val Thr Val Ala Pro Ser Asn Phe Ala Asn Gly Val Ala Glu
20 25 30
Trp Ile Ser Ser Asn Ser Arg Ser Gln Ala Tyr Lys Val Thr Cys Ser
35 40 45
Val Arg Gln Ser Ser Ala Gln Asn Arg Lys Tyr Thr Ile Lys Val Glu
50 55 60
Val Pro Lys Val Ala Thr Gln Thr Val Gly Gly Val Glu Leu Pro Val
65 70 75 80
Ala Ala Trp Arg Ser Tyr Leu Asn Met Glu Leu Thr Ile Pro Ile Phe
85 90 95
Ala Thr Asn Ser Asp Cys Glu Leu Ile Val Lys Ala Met Gln Gly Leu
100 105 110
Leu Lys Asp Gly Asn Pro Ile Pro Ser Ala Ile Ala Ala Asn Ser Gly
115 120 125
Ile Tyr
130
<210> 38
<211> 15
<212> DNA
<213> λ噬菌体
<400> 38
gccctgaaga agggc 15
<210> 39
<211> 19
<212> DNA
<213> Bacteriophage MS2
<400> 39
acatgaggat cacccatgt 19
<210> 40
<211> 7125
<212> DNA
<213> 人工序列
<220>
<223> λ N-R341-G4-NP868R-(G4S)2-K1ERNAP的核苷酸序列
<400> 40
atggatgctc agaccagacg cagagaacgg cgggcagaga agcaggcaca gtggaaggcc 60
gctaatggcg gtggcggcac cggtctcaaa aacaagacca gggctgagaa gtatcagacc 120
tactacacca ccaatgaata tcagatcgtt aaagaaaaac taccagacat tataagagac 180
gcggaaatta aggcgtctga agtgctcgag ccaaccatct acgagaagcg cgcaatcatg 240
gaagtcatta aggatttcat tcgggatcat caaaggaaag tgtatggcgg aacagccctg 300
aatgaggcat tgaaacaggt gaatcccaag gatgccatct atgataacta ttccttcagc 360
gacatcgagt tttattcccc tacccccgtg caggatctcg tggatctctg caacatcctg 420
tatagaaaag ggtataagtt cgtccagggg aaggacgctc agcatgagga aacctattct 480
atctttgtaa atttccagct ctactgtgac attacctatt cgccaacccg ggtcttttat 540
ggtattaaaa cgatagaaat tgacggcatt aactataccg atcctcattt catgctcata 600
gattacctcc gaatggtgaa ccagcccttg actgccgccg gccagcgctg ggagaaagcg 660
ttcgaacgga tgtacaggct gctcaaagac tatcccattg aggattttga caagaggctg 720
gatattcctg agccacccga agaaatccag agttatattt ctcggattaa gaccgagttt 780
ctgagcgata acaagctgaa tgaaagcttc ctcatctccg gcatcgaggc ttacaacttc 840
tacattcgcc atgctgcctc tagcaaagat gaagaacaga tggcccggac aaaccgcaat 900
gtggtcaatc ttaataactt tattgcaaat gtccccttta gcgagctgat ctccgtgaac 960
tatcgcgaag atgtcaagaa tacctataac ttcctgcgga tgatcgtcga ggataaagag 1020
aaaatcagtg ttgacgaata tttccctctc tttcaattca ctggctattc cactgtcatc 1080
aaatacgatg atcaccccat aattaggatc tacgagggcg acggttattg tattcctaac 1140
gtcaagaccg ttaaaacggt ggagaatgac aacggaacga agacaaagta cgagtacaag 1200
tacgtatcct tccagtacgt cctcatgatt ctatatatca acaaatttcg tgcgcacttg 1260
gacaagaata agcctatgta ttttaactac ggtattgcca tatccaatct ggtcaaagct 1320
cgcaatatat acctggacca gaccgggaaa agcgtccttg acaacactgt gtttaaggag 1380
ttccgcacta actgtaccgg aaatacgatc tctttcacac ggatgaacag actgagatta 1440
ctcgagaaaa gaaagcaggg caagcagact tcgttcgttt acacccctga agacttcttt 1500
aagaaggatc tggaaaccca agccaagctt gacccgtcga aagcgagatt caaaaatacc 1560
agtggtaaca agattatggt gccaaagtac ctgctgttca aaatagataa caacggaaat 1620
attgaagata acatacatag cgaagaggca gaaatctcag agaaagaaga aacttccggt 1680
ggctcttcta tatccactga taaatcattc gaagaatcac ctaattcctc ccctaacagc 1740
tctcctaaca actcgttgaa taattctatt gatatcagta caaataatta cgacgaccgc 1800
tcggaaaaca gcctggactc actcacgtct gatgggcccg gcggtggcgg cgaattcgcc 1860
agcctggaca acctggtggc cagataccag cggtgcttca acgaccagag cctgaagaac 1920
agcaccatcg agctggaaat ccggttccag cagatcaact tcctgctgtt caagaccgtg 1980
tacgaggccc tggtcgccca ggaaatcccc agcaccatca gccacagcat ccggtgcatc 2040
aagaaggtgc accacgagaa ccactgccgg gagaagatcc tgcccagcga gaacctgtac 2100
ttcaagaaac agcccctgat gttcttcaag ttcagcgagc ccgccagcct gggctgtaaa 2160
gtgtccctgg ccatcgagca gcccatccgg aagttcatcc tggacagcag cgtgctggtc 2220
cggctgaaga accggaccac cttccgggtg tccgagctgt ggaagatcga gctgaccatc 2280
gtgaagcagc tgatgggcag cgaggtgtca gccaagctgg ccgccttcaa gaccctgctg 2340
ttcgacaccc ccgagcagca gaccaccaag aacatgatga ccctgatcaa ccccgacgac 2400
gagtacctgt acgagatcga gatcgagtac accggcaagc ctgagagcct gacagccgcc 2460
gacgtgatca agatcaagaa caccgtgctg acactgatca gccccaacca cctgatgctg 2520
accgcctacc accaggccat cgagtttatc gccagccaca tcctgagcag cgagatcctg 2580
ctggcccgga tcaagagcgg caagtggggc ctgaagagac tgctgcccca ggtcaagtcc 2640
atgaccaagg ccgactacat gaagttctac ccccccgtgg gctactacgt gaccgacaag 2700
gccgacggca tccggggcat tgccgtgatc caggacaccc agatctacgt ggtggccgac 2760
cagctgtaca gcctgggcac caccggcatc gagcccctga agcccaccat cctggacggc 2820
gagttcatgc ccgagaagaa agagttctac ggctttgacg tgatcatgta cgagggcaac 2880
ctgctgaccc agcagggctt cgagacacgg atcgagagcc tgagcaaggg catcaaggtg 2940
ctgcaggcct tcaacatcaa ggccgagatg aagcccttca tcagcctgac ctccgccgac 3000
cccaacgtgc tgctgaagaa tttcgagagc atcttcaaga agaaaacccg gccctacagc 3060
atcgacggca tcatcctggt ggagcccggc aacagctacc tgaacaccaa caccttcaag 3120
tggaagccca cctgggacaa caccctggac tttctggtcc ggaagtgccc cgagtccctg 3180
aacgtgcccg agtacgcccc caagaagggc ttcagcctgc atctgctgtt cgtgggcatc 3240
agcggcgagc tgtttaagaa gctggccctg aactggtgcc ccggctacac caagctgttc 3300
cccgtgaccc agcggaacca gaactacttc cccgtgcagt tccagcccag cgacttcccc 3360
ctggccttcc tgtactacca ccccgacacc agcagcttca gcaacatcga tggcaaggtg 3420
ctggaaatgc ggtgcctgaa gcgggagatc aactacgtgc gctgggagat cgtgaagatc 3480
cgggaggacc ggcagcagga tctgaaaacc ggcggctact tcggcaacga cttcaagacc 3540
gccgagctga cctggctgaa ctacatggac cccttcagct tcgaggaact ggccaaggga 3600
cccagcggca tgtacttcgc tggcgccaag accggcatct acagagccca gaccgccctg 3660
atcagcttca tcaagcagga aatcatccag aagatcagcc accagagctg ggtgatcgac 3720
ctgggcatcg gcaagggcca ggacctgggc agatacctgg acgccggcgt gagacacctg 3780
gtcggcatcg ataaggacca gacagccctg gccgagctgg tgtaccggaa gttctcccac 3840
gccaccacca gacagcacaa gcacgccacc aacatctacg tgctgcacca ggatctggcc 3900
gagcctgcca aagaaatcag cgagaaagtg caccagatct atggcttccc caaagagggc 3960
gccagcagca tcgtgtccaa cctgttcatc cactacctga tgaagaacac ccagcaggtc 4020
gagaacctgg ctgtgctgtg ccacaagctg ctgcagcctg gcggcatggt ctggttcacc 4080
accatgctgg gcgaacaggt gctggaactg ctgcacgaga accggatcga actgaacgaa 4140
gtgtgggagg cccgggagaa cgaggtggtc aagttcgcca tcaagcggct gttcaaagag 4200
gacatcctgc aggaaaccgg ccaggaaatc ggcgtcctgc tgcccttcag caacggcgac 4260
ttctacaatg agtacctggt caacaccgcc tttctgatca agattttcaa gcaccatggc 4320
tttagcctcg tgcagaagca gagcttcaag gactggatcc ccgagttcca gaacttcagc 4380
aagagcctgt acaagatcct gaccgaggcc gacaagacct ggaccagcct gttcggcttc 4440
atctgcctgc ggaagaacct cgagggagga ggaggatcag gcggaggcgg aagtgtcgag 4500
caggacctgc acgccatcca gctgcagctc gaagaggaaa tgttcaacgg cggcatcaga 4560
agattcgagg ccgaccagca gagacagatc gcctctggca acgagagcga caccgcctgg 4620
aatagaaggc tgctgtctga gctgatcgcc cctatggccg aaggcatcca ggcctacaaa 4680
gaggaatacg agggcaagag aggcagagcc cctagagccc tggccttcat caactgtgtg 4740
ggcaatgagg tggccgccta catcaccatg aagatcgtga tggacatgct gaacaccgac 4800
gtgaccctgc aggccattgc catgaacgtg gccgacagaa tcgaggacca ggtccgattc 4860
agcaagctgg aaggacacgc cgccaagtac ttcgagaaag tgaagaagtc cctgaaggcc 4920
agcaagacca agagctacag acacgcccac aacgtggccg tggtggccga aaaatctgtg 4980
gccgataggg acgccgactt ctctagatgg gaggcctggc ctaaggacac cctgctgcag 5040
atcggcatga ccctgctgga aatcctggaa aacagcgtgt tcttcaacgg ccagcccgtg 5100
ttcctgagaa ccctgaggac aaatggcggc aagcacggcg tgtactacct gcagacatct 5160
gagcacgtgg gcgagtggat caccgccttc aaagaacatg tggcccagct gagccctgcc 5220
tatgcccctt gtgtgatccc tcctagaccc tgggtgtccc ctttcaatgg cggctttcac 5280
accgagaagg tggccagcag aatcagactg gtcaagggca accgggaaca cgtgcggaag 5340
ctgaccaaga aacagatgcc cgccgtgtac aaggccgtga atgctctgca ggccaccaag 5400
tggcaggtca acaaagaggt gctgcaggtc gtcgaggacg tgatcagact ggatctgggc 5460
tacggcgtgc caagctttaa gcccctgatc gacagagaga acaagcccgc caaccctgtg 5520
cccctggaat ttcagcacct gagaggccgc gagctgaaag agatgctgac acctgaacag 5580
tggcaggcct ttatcaattg gaagggcgag tgcaccaagc tgtacaccgc cgagacaaag 5640
aggggctcta agtctgccgc cacagtgcga atggtcggac aggccagaaa gtacagccag 5700
ttcgacgcca tctacttcgt gtacgccctg gacagccggt ctagagtgta tgcccagagc 5760
agcacactga gcccccagtc taacgatctg ggaaaggccc tgctgagatt caccgagggc 5820
cagagactgg attctgccga agccctgaag tggttcctgg tcaacggcgc caacaactgg 5880
ggctgggaca agaaaacctt cgatgtgcgg accgccaacg tgctggatag cgagttccag 5940
gacatgtgca gagatatcgc cgccgaccct ctgaccttta cccagtgggt caacgccgat 6000
agcccctatg gattcctggc ctggtgcttc gagtacgcca gatacctgga cgccctggat 6060
gagggaaccc aggatcagtt catgacccat ctgcccgtgc accaggatgg ctcttgttct 6120
ggcatccagc actacagcgc catgctgagc gatgccgtgg gagccaaagc cgtgaacctg 6180
aagcctagcg acagccccca ggatatctat ggcgctgtgg cccaggtggt catccagaaa 6240
aactacgcct acatgaacgc cgaggacgcc gagacattca caagcggaag cgtgacactg 6300
acaggcgccg agctgagatc tatggcctct gcctgggaca tgatcggcat cacacggggc 6360
ctgaccaaaa agcctgtgat gacactgccc tacggcagca ccagactgac ctgtagagaa 6420
agcgtgatcg actacatcgt ggacctggaa gagaaagagg cccagagagc cattgccgag 6480
ggcagaacag ccaatcctgt gcaccccttc gacaacgacc ggaaggatag cctgacacct 6540
agcgccgcct acaactacat gaccgccctg atctggccca gcatctctga agtggtcaag 6600
gcccctatcg tggccatgaa gatgatcaga cagctggcca gattcgccgc caagagaaat 6660
gagggcctgg aataccctct gcccaccggc tttatcctgc agcagaaaat catggccacc 6720
gacatgctgc gggtgtccac atgtctgatg ggcgagatca agatgagcct gcagatcgag 6780
acagacgtgg tggacgagac agccatgatg ggagccgccg ctcctaattt tgtgcacgga 6840
cacgatgcca gccacctgat cctgaccgtg tgcgatctgg tggacaaggg catcactagc 6900
gtggccgtga tccacgatag ctttggaaca cacgccggca gaaccgccga cctgagagat 6960
tctctgcggg aagagatggt caagatgtac cagaaccaca acgccctgca gaacctgctg 7020
gacgtgcacg aagaaagatg gctggtggac accggcatcc aggtgccaga acagggagag 7080
ttcgacctga acgagatcct ggtgtccgac tactgcttcg cctga 7125
<210> 41
<211> 2374
<212> PRT
<213> 人工序列
<220>
<223> λ N-R341-G4-NP868R-(G4S)2-K1ERNAP的氨基酸序列
<400> 41
Met Asp Ala Gln Thr Arg Arg Arg Glu Arg Arg Ala Glu Lys Gln Ala
1 5 10 15
Gln Trp Lys Ala Ala Asn Gly Gly Gly Gly Thr Gly Leu Lys Asn Lys
20 25 30
Thr Arg Ala Glu Lys Tyr Gln Thr Tyr Tyr Thr Thr Asn Glu Tyr Gln
35 40 45
Ile Val Lys Glu Lys Leu Pro Asp Ile Ile Arg Asp Ala Glu Ile Lys
50 55 60
Ala Ser Glu Val Leu Glu Pro Thr Ile Tyr Glu Lys Arg Ala Ile Met
65 70 75 80
Glu Val Ile Lys Asp Phe Ile Arg Asp His Gln Arg Lys Val Tyr Gly
85 90 95
Gly Thr Ala Leu Asn Glu Ala Leu Lys Gln Val Asn Pro Lys Asp Ala
100 105 110
Ile Tyr Asp Asn Tyr Ser Phe Ser Asp Ile Glu Phe Tyr Ser Pro Thr
115 120 125
Pro Val Gln Asp Leu Val Asp Leu Cys Asn Ile Leu Tyr Arg Lys Gly
130 135 140
Tyr Lys Phe Val Gln Gly Lys Asp Ala Gln His Glu Glu Thr Tyr Ser
145 150 155 160
Ile Phe Val Asn Phe Gln Leu Tyr Cys Asp Ile Thr Tyr Ser Pro Thr
165 170 175
Arg Val Phe Tyr Gly Ile Lys Thr Ile Glu Ile Asp Gly Ile Asn Tyr
180 185 190
Thr Asp Pro His Phe Met Leu Ile Asp Tyr Leu Arg Met Val Asn Gln
195 200 205
Pro Leu Thr Ala Ala Gly Gln Arg Trp Glu Lys Ala Phe Glu Arg Met
210 215 220
Tyr Arg Leu Leu Lys Asp Tyr Pro Ile Glu Asp Phe Asp Lys Arg Leu
225 230 235 240
Asp Ile Pro Glu Pro Pro Glu Glu Ile Gln Ser Tyr Ile Ser Arg Ile
245 250 255
Lys Thr Glu Phe Leu Ser Asp Asn Lys Leu Asn Glu Ser Phe Leu Ile
260 265 270
Ser Gly Ile Glu Ala Tyr Asn Phe Tyr Ile Arg His Ala Ala Ser Ser
275 280 285
Lys Asp Glu Glu Gln Met Ala Arg Thr Asn Arg Asn Val Val Asn Leu
290 295 300
Asn Asn Phe Ile Ala Asn Val Pro Phe Ser Glu Leu Ile Ser Val Asn
305 310 315 320
Tyr Arg Glu Asp Val Lys Asn Thr Tyr Asn Phe Leu Arg Met Ile Val
325 330 335
Glu Asp Lys Glu Lys Ile Ser Val Asp Glu Tyr Phe Pro Leu Phe Gln
340 345 350
Phe Thr Gly Tyr Ser Thr Val Ile Lys Tyr Asp Asp His Pro Ile Ile
355 360 365
Arg Ile Tyr Glu Gly Asp Gly Tyr Cys Ile Pro Asn Val Lys Thr Val
370 375 380
Lys Thr Val Glu Asn Asp Asn Gly Thr Lys Thr Lys Tyr Glu Tyr Lys
385 390 395 400
Tyr Val Ser Phe Gln Tyr Val Leu Met Ile Leu Tyr Ile Asn Lys Phe
405 410 415
Arg Ala His Leu Asp Lys Asn Lys Pro Met Tyr Phe Asn Tyr Gly Ile
420 425 430
Ala Ile Ser Asn Leu Val Lys Ala Arg Asn Ile Tyr Leu Asp Gln Thr
435 440 445
Gly Lys Ser Val Leu Asp Asn Thr Val Phe Lys Glu Phe Arg Thr Asn
450 455 460
Cys Thr Gly Asn Thr Ile Ser Phe Thr Arg Met Asn Arg Leu Arg Leu
465 470 475 480
Leu Glu Lys Arg Lys Gln Gly Lys Gln Thr Ser Phe Val Tyr Thr Pro
485 490 495
Glu Asp Phe Phe Lys Lys Asp Leu Glu Thr Gln Ala Lys Leu Asp Pro
500 505 510
Ser Lys Ala Arg Phe Lys Asn Thr Ser Gly Asn Lys Ile Met Val Pro
515 520 525
Lys Tyr Leu Leu Phe Lys Ile Asp Asn Asn Gly Asn Ile Glu Asp Asn
530 535 540
Ile His Ser Glu Glu Ala Glu Ile Ser Glu Lys Glu Glu Thr Ser Gly
545 550 555 560
Gly Ser Ser Ile Ser Thr Asp Lys Ser Phe Glu Glu Ser Pro Asn Ser
565 570 575
Ser Pro Asn Ser Ser Pro Asn Asn Ser Leu Asn Asn Ser Ile Asp Ile
580 585 590
Ser Thr Asn Asn Tyr Asp Asp Arg Ser Glu Asn Ser Leu Asp Ser Leu
595 600 605
Thr Ser Asp Gly Pro Gly Gly Gly Gly Glu Phe Ala Ser Leu Asp Asn
610 615 620
Leu Val Ala Arg Tyr Gln Arg Cys Phe Asn Asp Gln Ser Leu Lys Asn
625 630 635 640
Ser Thr Ile Glu Leu Glu Ile Arg Phe Gln Gln Ile Asn Phe Leu Leu
645 650 655
Phe Lys Thr Val Tyr Glu Ala Leu Val Ala Gln Glu Ile Pro Ser Thr
660 665 670
Ile Ser His Ser Ile Arg Cys Ile Lys Lys Val His His Glu Asn His
675 680 685
Cys Arg Glu Lys Ile Leu Pro Ser Glu Asn Leu Tyr Phe Lys Lys Gln
690 695 700
Pro Leu Met Phe Phe Lys Phe Ser Glu Pro Ala Ser Leu Gly Cys Lys
705 710 715 720
Val Ser Leu Ala Ile Glu Gln Pro Ile Arg Lys Phe Ile Leu Asp Ser
725 730 735
Ser Val Leu Val Arg Leu Lys Asn Arg Thr Thr Phe Arg Val Ser Glu
740 745 750
Leu Trp Lys Ile Glu Leu Thr Ile Val Lys Gln Leu Met Gly Ser Glu
755 760 765
Val Ser Ala Lys Leu Ala Ala Phe Lys Thr Leu Leu Phe Asp Thr Pro
770 775 780
Glu Gln Gln Thr Thr Lys Asn Met Met Thr Leu Ile Asn Pro Asp Asp
785 790 795 800
Glu Tyr Leu Tyr Glu Ile Glu Ile Glu Tyr Thr Gly Lys Pro Glu Ser
805 810 815
Leu Thr Ala Ala Asp Val Ile Lys Ile Lys Asn Thr Val Leu Thr Leu
820 825 830
Ile Ser Pro Asn His Leu Met Leu Thr Ala Tyr His Gln Ala Ile Glu
835 840 845
Phe Ile Ala Ser His Ile Leu Ser Ser Glu Ile Leu Leu Ala Arg Ile
850 855 860
Lys Ser Gly Lys Trp Gly Leu Lys Arg Leu Leu Pro Gln Val Lys Ser
865 870 875 880
Met Thr Lys Ala Asp Tyr Met Lys Phe Tyr Pro Pro Val Gly Tyr Tyr
885 890 895
Val Thr Asp Lys Ala Asp Gly Ile Arg Gly Ile Ala Val Ile Gln Asp
900 905 910
Thr Gln Ile Tyr Val Val Ala Asp Gln Leu Tyr Ser Leu Gly Thr Thr
915 920 925
Gly Ile Glu Pro Leu Lys Pro Thr Ile Leu Asp Gly Glu Phe Met Pro
930 935 940
Glu Lys Lys Glu Phe Tyr Gly Phe Asp Val Ile Met Tyr Glu Gly Asn
945 950 955 960
Leu Leu Thr Gln Gln Gly Phe Glu Thr Arg Ile Glu Ser Leu Ser Lys
965 970 975
Gly Ile Lys Val Leu Gln Ala Phe Asn Ile Lys Ala Glu Met Lys Pro
980 985 990
Phe Ile Ser Leu Thr Ser Ala Asp Pro Asn Val Leu Leu Lys Asn Phe
995 1000 1005
Glu Ser Ile Phe Lys Lys Lys Thr Arg Pro Tyr Ser Ile Asp Gly
1010 1015 1020
Ile Ile Leu Val Glu Pro Gly Asn Ser Tyr Leu Asn Thr Asn Thr
1025 1030 1035
Phe Lys Trp Lys Pro Thr Trp Asp Asn Thr Leu Asp Phe Leu Val
1040 1045 1050
Arg Lys Cys Pro Glu Ser Leu Asn Val Pro Glu Tyr Ala Pro Lys
1055 1060 1065
Lys Gly Phe Ser Leu His Leu Leu Phe Val Gly Ile Ser Gly Glu
1070 1075 1080
Leu Phe Lys Lys Leu Ala Leu Asn Trp Cys Pro Gly Tyr Thr Lys
1085 1090 1095
Leu Phe Pro Val Thr Gln Arg Asn Gln Asn Tyr Phe Pro Val Gln
1100 1105 1110
Phe Gln Pro Ser Asp Phe Pro Leu Ala Phe Leu Tyr Tyr His Pro
1115 1120 1125
Asp Thr Ser Ser Phe Ser Asn Ile Asp Gly Lys Val Leu Glu Met
1130 1135 1140
Arg Cys Leu Lys Arg Glu Ile Asn Tyr Val Arg Trp Glu Ile Val
1145 1150 1155
Lys Ile Arg Glu Asp Arg Gln Gln Asp Leu Lys Thr Gly Gly Tyr
1160 1165 1170
Phe Gly Asn Asp Phe Lys Thr Ala Glu Leu Thr Trp Leu Asn Tyr
1175 1180 1185
Met Asp Pro Phe Ser Phe Glu Glu Leu Ala Lys Gly Pro Ser Gly
1190 1195 1200
Met Tyr Phe Ala Gly Ala Lys Thr Gly Ile Tyr Arg Ala Gln Thr
1205 1210 1215
Ala Leu Ile Ser Phe Ile Lys Gln Glu Ile Ile Gln Lys Ile Ser
1220 1225 1230
His Gln Ser Trp Val Ile Asp Leu Gly Ile Gly Lys Gly Gln Asp
1235 1240 1245
Leu Gly Arg Tyr Leu Asp Ala Gly Val Arg His Leu Val Gly Ile
1250 1255 1260
Asp Lys Asp Gln Thr Ala Leu Ala Glu Leu Val Tyr Arg Lys Phe
1265 1270 1275
Ser His Ala Thr Thr Arg Gln His Lys His Ala Thr Asn Ile Tyr
1280 1285 1290
Val Leu His Gln Asp Leu Ala Glu Pro Ala Lys Glu Ile Ser Glu
1295 1300 1305
Lys Val His Gln Ile Tyr Gly Phe Pro Lys Glu Gly Ala Ser Ser
1310 1315 1320
Ile Val Ser Asn Leu Phe Ile His Tyr Leu Met Lys Asn Thr Gln
1325 1330 1335
Gln Val Glu Asn Leu Ala Val Leu Cys His Lys Leu Leu Gln Pro
1340 1345 1350
Gly Gly Met Val Trp Phe Thr Thr Met Leu Gly Glu Gln Val Leu
1355 1360 1365
Glu Leu Leu His Glu Asn Arg Ile Glu Leu Asn Glu Val Trp Glu
1370 1375 1380
Ala Arg Glu Asn Glu Val Val Lys Phe Ala Ile Lys Arg Leu Phe
1385 1390 1395
Lys Glu Asp Ile Leu Gln Glu Thr Gly Gln Glu Ile Gly Val Leu
1400 1405 1410
Leu Pro Phe Ser Asn Gly Asp Phe Tyr Asn Glu Tyr Leu Val Asn
1415 1420 1425
Thr Ala Phe Leu Ile Lys Ile Phe Lys His His Gly Phe Ser Leu
1430 1435 1440
Val Gln Lys Gln Ser Phe Lys Asp Trp Ile Pro Glu Phe Gln Asn
1445 1450 1455
Phe Ser Lys Ser Leu Tyr Lys Ile Leu Thr Glu Ala Asp Lys Thr
1460 1465 1470
Trp Thr Ser Leu Phe Gly Phe Ile Cys Leu Arg Lys Asn Leu Glu
1475 1480 1485
Gly Gly Gly Gly Ser Gly Gly Gly Gly Ser Val Glu Gln Asp Leu
1490 1495 1500
His Ala Ile Gln Leu Gln Leu Glu Glu Glu Met Phe Asn Gly Gly
1505 1510 1515
Ile Arg Arg Phe Glu Ala Asp Gln Gln Arg Gln Ile Ala Ser Gly
1520 1525 1530
Asn Glu Ser Asp Thr Ala Trp Asn Arg Arg Leu Leu Ser Glu Leu
1535 1540 1545
Ile Ala Pro Met Ala Glu Gly Ile Gln Ala Tyr Lys Glu Glu Tyr
1550 1555 1560
Glu Gly Lys Arg Gly Arg Ala Pro Arg Ala Leu Ala Phe Ile Asn
1565 1570 1575
Cys Val Gly Asn Glu Val Ala Ala Tyr Ile Thr Met Lys Ile Val
1580 1585 1590
Met Asp Met Leu Asn Thr Asp Val Thr Leu Gln Ala Ile Ala Met
1595 1600 1605
Asn Val Ala Asp Arg Ile Glu Asp Gln Val Arg Phe Ser Lys Leu
1610 1615 1620
Glu Gly His Ala Ala Lys Tyr Phe Glu Lys Val Lys Lys Ser Leu
1625 1630 1635
Lys Ala Ser Lys Thr Lys Ser Tyr Arg His Ala His Asn Val Ala
1640 1645 1650
Val Val Ala Glu Lys Ser Val Ala Asp Arg Asp Ala Asp Phe Ser
1655 1660 1665
Arg Trp Glu Ala Trp Pro Lys Asp Thr Leu Leu Gln Ile Gly Met
1670 1675 1680
Thr Leu Leu Glu Ile Leu Glu Asn Ser Val Phe Phe Asn Gly Gln
1685 1690 1695
Pro Val Phe Leu Arg Thr Leu Arg Thr Asn Gly Gly Lys His Gly
1700 1705 1710
Val Tyr Tyr Leu Gln Thr Ser Glu His Val Gly Glu Trp Ile Thr
1715 1720 1725
Ala Phe Lys Glu His Val Ala Gln Leu Ser Pro Ala Tyr Ala Pro
1730 1735 1740
Cys Val Ile Pro Pro Arg Pro Trp Val Ser Pro Phe Asn Gly Gly
1745 1750 1755
Phe His Thr Glu Lys Val Ala Ser Arg Ile Arg Leu Val Lys Gly
1760 1765 1770
Asn Arg Glu His Val Arg Lys Leu Thr Lys Lys Gln Met Pro Ala
1775 1780 1785
Val Tyr Lys Ala Val Asn Ala Leu Gln Ala Thr Lys Trp Gln Val
1790 1795 1800
Asn Lys Glu Val Leu Gln Val Val Glu Asp Val Ile Arg Leu Asp
1805 1810 1815
Leu Gly Tyr Gly Val Pro Ser Phe Lys Pro Leu Ile Asp Arg Glu
1820 1825 1830
Asn Lys Pro Ala Asn Pro Val Pro Leu Glu Phe Gln His Leu Arg
1835 1840 1845
Gly Arg Glu Leu Lys Glu Met Leu Thr Pro Glu Gln Trp Gln Ala
1850 1855 1860
Phe Ile Asn Trp Lys Gly Glu Cys Thr Lys Leu Tyr Thr Ala Glu
1865 1870 1875
Thr Lys Arg Gly Ser Lys Ser Ala Ala Thr Val Arg Met Val Gly
1880 1885 1890
Gln Ala Arg Lys Tyr Ser Gln Phe Asp Ala Ile Tyr Phe Val Tyr
1895 1900 1905
Ala Leu Asp Ser Arg Ser Arg Val Tyr Ala Gln Ser Ser Thr Leu
1910 1915 1920
Ser Pro Gln Ser Asn Asp Leu Gly Lys Ala Leu Leu Arg Phe Thr
1925 1930 1935
Glu Gly Gln Arg Leu Asp Ser Ala Glu Ala Leu Lys Trp Phe Leu
1940 1945 1950
Val Asn Gly Ala Asn Asn Trp Gly Trp Asp Lys Lys Thr Phe Asp
1955 1960 1965
Val Arg Thr Ala Asn Val Leu Asp Ser Glu Phe Gln Asp Met Cys
1970 1975 1980
Arg Asp Ile Ala Ala Asp Pro Leu Thr Phe Thr Gln Trp Val Asn
1985 1990 1995
Ala Asp Ser Pro Tyr Gly Phe Leu Ala Trp Cys Phe Glu Tyr Ala
2000 2005 2010
Arg Tyr Leu Asp Ala Leu Asp Glu Gly Thr Gln Asp Gln Phe Met
2015 2020 2025
Thr His Leu Pro Val His Gln Asp Gly Ser Cys Ser Gly Ile Gln
2030 2035 2040
His Tyr Ser Ala Met Leu Ser Asp Ala Val Gly Ala Lys Ala Val
2045 2050 2055
Asn Leu Lys Pro Ser Asp Ser Pro Gln Asp Ile Tyr Gly Ala Val
2060 2065 2070
Ala Gln Val Val Ile Gln Lys Asn Tyr Ala Tyr Met Asn Ala Glu
2075 2080 2085
Asp Ala Glu Thr Phe Thr Ser Gly Ser Val Thr Leu Thr Gly Ala
2090 2095 2100
Glu Leu Arg Ser Met Ala Ser Ala Trp Asp Met Ile Gly Ile Thr
2105 2110 2115
Arg Gly Leu Thr Lys Lys Pro Val Met Thr Leu Pro Tyr Gly Ser
2120 2125 2130
Thr Arg Leu Thr Cys Arg Glu Ser Val Ile Asp Tyr Ile Val Asp
2135 2140 2145
Leu Glu Glu Lys Glu Ala Gln Arg Ala Ile Ala Glu Gly Arg Thr
2150 2155 2160
Ala Asn Pro Val His Pro Phe Asp Asn Asp Arg Lys Asp Ser Leu
2165 2170 2175
Thr Pro Ser Ala Ala Tyr Asn Tyr Met Thr Ala Leu Ile Trp Pro
2180 2185 2190
Ser Ile Ser Glu Val Val Lys Ala Pro Ile Val Ala Met Lys Met
2195 2200 2205
Ile Arg Gln Leu Ala Arg Phe Ala Ala Lys Arg Asn Glu Gly Leu
2210 2215 2220
Glu Tyr Pro Leu Pro Thr Gly Phe Ile Leu Gln Gln Lys Ile Met
2225 2230 2235
Ala Thr Asp Met Leu Arg Val Ser Thr Cys Leu Met Gly Glu Ile
2240 2245 2250
Lys Met Ser Leu Gln Ile Glu Thr Asp Val Val Asp Glu Thr Ala
2255 2260 2265
Met Met Gly Ala Ala Ala Pro Asn Phe Val His Gly His Asp Ala
2270 2275 2280
Ser His Leu Ile Leu Thr Val Cys Asp Leu Val Asp Lys Gly Ile
2285 2290 2295
Thr Ser Val Ala Val Ile His Asp Ser Phe Gly Thr His Ala Gly
2300 2305 2310
Arg Thr Ala Asp Leu Arg Asp Ser Leu Arg Glu Glu Met Val Lys
2315 2320 2325
Met Tyr Gln Asn His Asn Ala Leu Gln Asn Leu Leu Asp Val His
2330 2335 2340
Glu Glu Arg Trp Leu Val Asp Thr Gly Ile Gln Val Pro Glu Gln
2345 2350 2355
Gly Glu Phe Asp Leu Asn Glu Ile Leu Val Ser Asp Tyr Cys Phe
2360 2365 2370
Ala
<210> 42
<211> 7161
<212> DNA
<213> 人工序列
<220>
<223> λ N-R341-G4-NP868R-F2A-K1ERNAP的核苷酸序列
<400> 42
atggatgctc agaccagacg cagagaacgg cgggcagaga agcaggcaca gtggaaggcc 60
gctaatggcg gtggcggcac cggtctcaaa aacaagacca gggctgagaa gtatcagacc 120
tactacacca ccaatgaata tcagatcgtt aaagaaaaac taccagacat tataagagac 180
gcggaaatta aggcgtctga agtgctcgag ccaaccatct acgagaagcg cgcaatcatg 240
gaagtcatta aggatttcat tcgggatcat caaaggaaag tgtatggcgg aacagccctg 300
aatgaggcat tgaaacaggt gaatcccaag gatgccatct atgataacta ttccttcagc 360
gacatcgagt tttattcccc tacccccgtg caggatctcg tggatctctg caacatcctg 420
tatagaaaag ggtataagtt cgtccagggg aaggacgctc agcatgagga aacctattct 480
atctttgtaa atttccagct ctactgtgac attacctatt cgccaacccg ggtcttttat 540
ggtattaaaa cgatagaaat tgacggcatt aactataccg atcctcattt catgctcata 600
gattacctcc gaatggtgaa ccagcccttg actgccgccg gccagcgctg ggagaaagcg 660
ttcgaacgga tgtacaggct gctcaaagac tatcccattg aggattttga caagaggctg 720
gatattcctg agccacccga agaaatccag agttatattt ctcggattaa gaccgagttt 780
ctgagcgata acaagctgaa tgaaagcttc ctcatctccg gcatcgaggc ttacaacttc 840
tacattcgcc atgctgcctc tagcaaagat gaagaacaga tggcccggac aaaccgcaat 900
gtggtcaatc ttaataactt tattgcaaat gtccccttta gcgagctgat ctccgtgaac 960
tatcgcgaag atgtcaagaa tacctataac ttcctgcgga tgatcgtcga ggataaagag 1020
aaaatcagtg ttgacgaata tttccctctc tttcaattca ctggctattc cactgtcatc 1080
aaatacgatg atcaccccat aattaggatc tacgagggcg acggttattg tattcctaac 1140
gtcaagaccg ttaaaacggt ggagaatgac aacggaacga agacaaagta cgagtacaag 1200
tacgtatcct tccagtacgt cctcatgatt ctatatatca acaaatttcg tgcgcacttg 1260
gacaagaata agcctatgta ttttaactac ggtattgcca tatccaatct ggtcaaagct 1320
cgcaatatat acctggacca gaccgggaaa agcgtccttg acaacactgt gtttaaggag 1380
ttccgcacta actgtaccgg aaatacgatc tctttcacac ggatgaacag actgagatta 1440
ctcgagaaaa gaaagcaggg caagcagact tcgttcgttt acacccctga agacttcttt 1500
aagaaggatc tggaaaccca agccaagctt gacccgtcga aagcgagatt caaaaatacc 1560
agtggtaaca agattatggt gccaaagtac ctgctgttca aaatagataa caacggaaat 1620
attgaagata acatacatag cgaagaggca gaaatctcag agaaagaaga aacttccggt 1680
ggctcttcta tatccactga taaatcattc gaagaatcac ctaattcctc ccctaacagc 1740
tctcctaaca actcgttgaa taattctatt gatatcagta caaataatta cgacgaccgc 1800
tcggaaaaca gcctggactc actcacgtct gatgggcccg gcggtggcgg cgaattcgcc 1860
agcctggaca acctggtggc cagataccag cggtgcttca acgaccagag cctgaagaac 1920
agcaccatcg agctggaaat ccggttccag cagatcaact tcctgctgtt caagaccgtg 1980
tacgaggccc tggtcgccca ggaaatcccc agcaccatca gccacagcat ccggtgcatc 2040
aagaaggtgc accacgagaa ccactgccgg gagaagatcc tgcccagcga gaacctgtac 2100
ttcaagaaac agcccctgat gttcttcaag ttcagcgagc ccgccagcct gggctgtaaa 2160
gtgtccctgg ccatcgagca gcccatccgg aagttcatcc tggacagcag cgtgctggtc 2220
cggctgaaga accggaccac cttccgggtg tccgagctgt ggaagatcga gctgaccatc 2280
gtgaagcagc tgatgggcag cgaggtgtca gccaagctgg ccgccttcaa gaccctgctg 2340
ttcgacaccc ccgagcagca gaccaccaag aacatgatga ccctgatcaa ccccgacgac 2400
gagtacctgt acgagatcga gatcgagtac accggcaagc ctgagagcct gacagccgcc 2460
gacgtgatca agatcaagaa caccgtgctg acactgatca gccccaacca cctgatgctg 2520
accgcctacc accaggccat cgagtttatc gccagccaca tcctgagcag cgagatcctg 2580
ctggcccgga tcaagagcgg caagtggggc ctgaagagac tgctgcccca ggtcaagtcc 2640
atgaccaagg ccgactacat gaagttctac ccccccgtgg gctactacgt gaccgacaag 2700
gccgacggca tccggggcat tgccgtgatc caggacaccc agatctacgt ggtggccgac 2760
cagctgtaca gcctgggcac caccggcatc gagcccctga agcccaccat cctggacggc 2820
gagttcatgc ccgagaagaa agagttctac ggctttgacg tgatcatgta cgagggcaac 2880
ctgctgaccc agcagggctt cgagacacgg atcgagagcc tgagcaaggg catcaaggtg 2940
ctgcaggcct tcaacatcaa ggccgagatg aagcccttca tcagcctgac ctccgccgac 3000
cccaacgtgc tgctgaagaa tttcgagagc atcttcaaga agaaaacccg gccctacagc 3060
atcgacggca tcatcctggt ggagcccggc aacagctacc tgaacaccaa caccttcaag 3120
tggaagccca cctgggacaa caccctggac tttctggtcc ggaagtgccc cgagtccctg 3180
aacgtgcccg agtacgcccc caagaagggc ttcagcctgc atctgctgtt cgtgggcatc 3240
agcggcgagc tgtttaagaa gctggccctg aactggtgcc ccggctacac caagctgttc 3300
cccgtgaccc agcggaacca gaactacttc cccgtgcagt tccagcccag cgacttcccc 3360
ctggccttcc tgtactacca ccccgacacc agcagcttca gcaacatcga tggcaaggtg 3420
ctggaaatgc ggtgcctgaa gcgggagatc aactacgtgc gctgggagat cgtgaagatc 3480
cgggaggacc ggcagcagga tctgaaaacc ggcggctact tcggcaacga cttcaagacc 3540
gccgagctga cctggctgaa ctacatggac cccttcagct tcgaggaact ggccaaggga 3600
cccagcggca tgtacttcgc tggcgccaag accggcatct acagagccca gaccgccctg 3660
atcagcttca tcaagcagga aatcatccag aagatcagcc accagagctg ggtgatcgac 3720
ctgggcatcg gcaagggcca ggacctgggc agatacctgg acgccggcgt gagacacctg 3780
gtcggcatcg ataaggacca gacagccctg gccgagctgg tgtaccggaa gttctcccac 3840
gccaccacca gacagcacaa gcacgccacc aacatctacg tgctgcacca ggatctggcc 3900
gagcctgcca aagaaatcag cgagaaagtg caccagatct atggcttccc caaagagggc 3960
gccagcagca tcgtgtccaa cctgttcatc cactacctga tgaagaacac ccagcaggtc 4020
gagaacctgg ctgtgctgtg ccacaagctg ctgcagcctg gcggcatggt ctggttcacc 4080
accatgctgg gcgaacaggt gctggaactg ctgcacgaga accggatcga actgaacgaa 4140
gtgtgggagg cccgggagaa cgaggtggtc aagttcgcca tcaagcggct gttcaaagag 4200
gacatcctgc aggaaaccgg ccaggaaatc ggcgtcctgc tgcccttcag caacggcgac 4260
ttctacaatg agtacctggt caacaccgcc tttctgatca agattttcaa gcaccatggc 4320
tttagcctcg tgcagaagca gagcttcaag gactggatcc ccgagttcca gaacttcagc 4380
aagagcctgt acaagatcct gaccgaggcc gacaagacct ggaccagcct gttcggcttc 4440
atctgcctgc ggaagaaccc gcgggtgaag cagactctga actttgactt gttgaaactt 4500
gcgggtgacg tggaaagcaa cccaggcccc actagtcagg acctgcacgc catccagctg 4560
cagctcgaag aggaaatgtt caacggcggc atcagaagat tcgaggccga ccagcagaga 4620
cagatcgcct ctggcaacga gagcgacacc gcctggaata gaaggctgct gtctgagctg 4680
atcgccccta tggccgaagg catccaggcc tacaaagagg aatacgaggg caagagaggc 4740
agagccccta gagccctggc cttcatcaac tgtgtgggca atgaggtggc cgcctacatc 4800
accatgaaga tcgtgatgga catgctgaac accgacgtga ccctgcaggc cattgccatg 4860
aacgtggccg acagaatcga ggaccaggtc cgattcagca agctggaagg acacgccgcc 4920
aagtacttcg agaaagtgaa gaagtccctg aaggccagca agaccaagag ctacagacac 4980
gcccacaacg tggccgtggt ggccgaaaaa tctgtggccg atagggacgc cgacttctct 5040
agatgggagg cctggcctaa ggacaccctg ctgcagatcg gcatgaccct gctggaaatc 5100
ctggaaaaca gcgtgttctt caacggccag cccgtgttcc tgagaaccct gaggacaaat 5160
ggcggcaagc acggcgtgta ctacctgcag acatctgagc acgtgggcga gtggatcacc 5220
gccttcaaag aacatgtggc ccagctgagc cctgcctatg ccccttgtgt gatccctcct 5280
agaccctggg tgtccccttt caatggcggc tttcacaccg agaaggtggc cagcagaatc 5340
agactggtca agggcaaccg ggaacacgtg cggaagctga ccaagaaaca gatgcccgcc 5400
gtgtacaagg ccgtgaatgc tctgcaggcc accaagtggc aggtcaacaa agaggtgctg 5460
caggtcgtcg aggacgtgat cagactggat ctgggctacg gcgtgccaag ctttaagccc 5520
ctgatcgaca gagagaacaa gcccgccaac cctgtgcccc tggaatttca gcacctgaga 5580
ggccgcgagc tgaaagagat gctgacacct gaacagtggc aggcctttat caattggaag 5640
ggcgagtgca ccaagctgta caccgccgag acaaagaggg gctctaagtc tgccgccaca 5700
gtgcgaatgg tcggacaggc cagaaagtac agccagttcg acgccatcta cttcgtgtac 5760
gccctggaca gccggtctag agtgtatgcc cagagcagca cactgagccc ccagtctaac 5820
gatctgggaa aggccctgct gagattcacc gagggccaga gactggattc tgccgaagcc 5880
ctgaagtggt tcctggtcaa cggcgccaac aactggggct gggacaagaa aaccttcgat 5940
gtgcggaccg ccaacgtgct ggatagcgag ttccaggaca tgtgcagaga tatcgccgcc 6000
gaccctctga cctttaccca gtgggtcaac gccgatagcc cctatggatt cctggcctgg 6060
tgcttcgagt acgccagata cctggacgcc ctggatgagg gaacccagga tcagttcatg 6120
acccatctgc ccgtgcacca ggatggctct tgttctggca tccagcacta cagcgccatg 6180
ctgagcgatg ccgtgggagc caaagccgtg aacctgaagc ctagcgacag cccccaggat 6240
atctatggcg ctgtggccca ggtggtcatc cagaaaaact acgcctacat gaacgccgag 6300
gacgccgaga cattcacaag cggaagcgtg acactgacag gcgccgagct gagatctatg 6360
gcctctgcct gggacatgat cggcatcaca cggggcctga ccaaaaagcc tgtgatgaca 6420
ctgccctacg gcagcaccag actgacctgt agagaaagcg tgatcgacta catcgtggac 6480
ctggaagaga aagaggccca gagagccatt gccgagggca gaacagccaa tcctgtgcac 6540
cccttcgaca acgaccggaa ggatagcctg acacctagcg ccgcctacaa ctacatgacc 6600
gccctgatct ggcccagcat ctctgaagtg gtcaaggccc ctatcgtggc catgaagatg 6660
atcagacagc tggccagatt cgccgccaag agaaatgagg gcctggaata ccctctgccc 6720
accggcttta tcctgcagca gaaaatcatg gccaccgaca tgctgcgggt gtccacatgt 6780
ctgatgggcg agatcaagat gagcctgcag atcgagacag acgtggtgga cgagacagcc 6840
atgatgggag ccgccgctcc taattttgtg cacggacacg atgccagcca cctgatcctg 6900
accgtgtgcg atctggtgga caagggcatc actagcgtgg ccgtgatcca cgatagcttt 6960
ggaacacacg ccggcagaac cgccgacctg agagattctc tgcgggaaga gatggtcaag 7020
atgtaccaga accacaacgc cctgcagaac ctgctggacg tgcacgaaga aagatggctg 7080
gtggacaccg gcatccaggt gccagaacag ggagagttcg acctgaacga gatcctggtg 7140
tccgactact gcttcgcctg a 7161
<210> 43
<211> 2386
<212> PRT
<213> 人工序列
<220>
<223> λ N-R341-G4-NP868R-F2A-K1ERNAP的氨基酸序列
<400> 43
Met Asp Ala Gln Thr Arg Arg Arg Glu Arg Arg Ala Glu Lys Gln Ala
1 5 10 15
Gln Trp Lys Ala Ala Asn Gly Gly Gly Gly Thr Gly Leu Lys Asn Lys
20 25 30
Thr Arg Ala Glu Lys Tyr Gln Thr Tyr Tyr Thr Thr Asn Glu Tyr Gln
35 40 45
Ile Val Lys Glu Lys Leu Pro Asp Ile Ile Arg Asp Ala Glu Ile Lys
50 55 60
Ala Ser Glu Val Leu Glu Pro Thr Ile Tyr Glu Lys Arg Ala Ile Met
65 70 75 80
Glu Val Ile Lys Asp Phe Ile Arg Asp His Gln Arg Lys Val Tyr Gly
85 90 95
Gly Thr Ala Leu Asn Glu Ala Leu Lys Gln Val Asn Pro Lys Asp Ala
100 105 110
Ile Tyr Asp Asn Tyr Ser Phe Ser Asp Ile Glu Phe Tyr Ser Pro Thr
115 120 125
Pro Val Gln Asp Leu Val Asp Leu Cys Asn Ile Leu Tyr Arg Lys Gly
130 135 140
Tyr Lys Phe Val Gln Gly Lys Asp Ala Gln His Glu Glu Thr Tyr Ser
145 150 155 160
Ile Phe Val Asn Phe Gln Leu Tyr Cys Asp Ile Thr Tyr Ser Pro Thr
165 170 175
Arg Val Phe Tyr Gly Ile Lys Thr Ile Glu Ile Asp Gly Ile Asn Tyr
180 185 190
Thr Asp Pro His Phe Met Leu Ile Asp Tyr Leu Arg Met Val Asn Gln
195 200 205
Pro Leu Thr Ala Ala Gly Gln Arg Trp Glu Lys Ala Phe Glu Arg Met
210 215 220
Tyr Arg Leu Leu Lys Asp Tyr Pro Ile Glu Asp Phe Asp Lys Arg Leu
225 230 235 240
Asp Ile Pro Glu Pro Pro Glu Glu Ile Gln Ser Tyr Ile Ser Arg Ile
245 250 255
Lys Thr Glu Phe Leu Ser Asp Asn Lys Leu Asn Glu Ser Phe Leu Ile
260 265 270
Ser Gly Ile Glu Ala Tyr Asn Phe Tyr Ile Arg His Ala Ala Ser Ser
275 280 285
Lys Asp Glu Glu Gln Met Ala Arg Thr Asn Arg Asn Val Val Asn Leu
290 295 300
Asn Asn Phe Ile Ala Asn Val Pro Phe Ser Glu Leu Ile Ser Val Asn
305 310 315 320
Tyr Arg Glu Asp Val Lys Asn Thr Tyr Asn Phe Leu Arg Met Ile Val
325 330 335
Glu Asp Lys Glu Lys Ile Ser Val Asp Glu Tyr Phe Pro Leu Phe Gln
340 345 350
Phe Thr Gly Tyr Ser Thr Val Ile Lys Tyr Asp Asp His Pro Ile Ile
355 360 365
Arg Ile Tyr Glu Gly Asp Gly Tyr Cys Ile Pro Asn Val Lys Thr Val
370 375 380
Lys Thr Val Glu Asn Asp Asn Gly Thr Lys Thr Lys Tyr Glu Tyr Lys
385 390 395 400
Tyr Val Ser Phe Gln Tyr Val Leu Met Ile Leu Tyr Ile Asn Lys Phe
405 410 415
Arg Ala His Leu Asp Lys Asn Lys Pro Met Tyr Phe Asn Tyr Gly Ile
420 425 430
Ala Ile Ser Asn Leu Val Lys Ala Arg Asn Ile Tyr Leu Asp Gln Thr
435 440 445
Gly Lys Ser Val Leu Asp Asn Thr Val Phe Lys Glu Phe Arg Thr Asn
450 455 460
Cys Thr Gly Asn Thr Ile Ser Phe Thr Arg Met Asn Arg Leu Arg Leu
465 470 475 480
Leu Glu Lys Arg Lys Gln Gly Lys Gln Thr Ser Phe Val Tyr Thr Pro
485 490 495
Glu Asp Phe Phe Lys Lys Asp Leu Glu Thr Gln Ala Lys Leu Asp Pro
500 505 510
Ser Lys Ala Arg Phe Lys Asn Thr Ser Gly Asn Lys Ile Met Val Pro
515 520 525
Lys Tyr Leu Leu Phe Lys Ile Asp Asn Asn Gly Asn Ile Glu Asp Asn
530 535 540
Ile His Ser Glu Glu Ala Glu Ile Ser Glu Lys Glu Glu Thr Ser Gly
545 550 555 560
Gly Ser Ser Ile Ser Thr Asp Lys Ser Phe Glu Glu Ser Pro Asn Ser
565 570 575
Ser Pro Asn Ser Ser Pro Asn Asn Ser Leu Asn Asn Ser Ile Asp Ile
580 585 590
Ser Thr Asn Asn Tyr Asp Asp Arg Ser Glu Asn Ser Leu Asp Ser Leu
595 600 605
Thr Ser Asp Gly Pro Gly Gly Gly Gly Glu Phe Ala Ser Leu Asp Asn
610 615 620
Leu Val Ala Arg Tyr Gln Arg Cys Phe Asn Asp Gln Ser Leu Lys Asn
625 630 635 640
Ser Thr Ile Glu Leu Glu Ile Arg Phe Gln Gln Ile Asn Phe Leu Leu
645 650 655
Phe Lys Thr Val Tyr Glu Ala Leu Val Ala Gln Glu Ile Pro Ser Thr
660 665 670
Ile Ser His Ser Ile Arg Cys Ile Lys Lys Val His His Glu Asn His
675 680 685
Cys Arg Glu Lys Ile Leu Pro Ser Glu Asn Leu Tyr Phe Lys Lys Gln
690 695 700
Pro Leu Met Phe Phe Lys Phe Ser Glu Pro Ala Ser Leu Gly Cys Lys
705 710 715 720
Val Ser Leu Ala Ile Glu Gln Pro Ile Arg Lys Phe Ile Leu Asp Ser
725 730 735
Ser Val Leu Val Arg Leu Lys Asn Arg Thr Thr Phe Arg Val Ser Glu
740 745 750
Leu Trp Lys Ile Glu Leu Thr Ile Val Lys Gln Leu Met Gly Ser Glu
755 760 765
Val Ser Ala Lys Leu Ala Ala Phe Lys Thr Leu Leu Phe Asp Thr Pro
770 775 780
Glu Gln Gln Thr Thr Lys Asn Met Met Thr Leu Ile Asn Pro Asp Asp
785 790 795 800
Glu Tyr Leu Tyr Glu Ile Glu Ile Glu Tyr Thr Gly Lys Pro Glu Ser
805 810 815
Leu Thr Ala Ala Asp Val Ile Lys Ile Lys Asn Thr Val Leu Thr Leu
820 825 830
Ile Ser Pro Asn His Leu Met Leu Thr Ala Tyr His Gln Ala Ile Glu
835 840 845
Phe Ile Ala Ser His Ile Leu Ser Ser Glu Ile Leu Leu Ala Arg Ile
850 855 860
Lys Ser Gly Lys Trp Gly Leu Lys Arg Leu Leu Pro Gln Val Lys Ser
865 870 875 880
Met Thr Lys Ala Asp Tyr Met Lys Phe Tyr Pro Pro Val Gly Tyr Tyr
885 890 895
Val Thr Asp Lys Ala Asp Gly Ile Arg Gly Ile Ala Val Ile Gln Asp
900 905 910
Thr Gln Ile Tyr Val Val Ala Asp Gln Leu Tyr Ser Leu Gly Thr Thr
915 920 925
Gly Ile Glu Pro Leu Lys Pro Thr Ile Leu Asp Gly Glu Phe Met Pro
930 935 940
Glu Lys Lys Glu Phe Tyr Gly Phe Asp Val Ile Met Tyr Glu Gly Asn
945 950 955 960
Leu Leu Thr Gln Gln Gly Phe Glu Thr Arg Ile Glu Ser Leu Ser Lys
965 970 975
Gly Ile Lys Val Leu Gln Ala Phe Asn Ile Lys Ala Glu Met Lys Pro
980 985 990
Phe Ile Ser Leu Thr Ser Ala Asp Pro Asn Val Leu Leu Lys Asn Phe
995 1000 1005
Glu Ser Ile Phe Lys Lys Lys Thr Arg Pro Tyr Ser Ile Asp Gly
1010 1015 1020
Ile Ile Leu Val Glu Pro Gly Asn Ser Tyr Leu Asn Thr Asn Thr
1025 1030 1035
Phe Lys Trp Lys Pro Thr Trp Asp Asn Thr Leu Asp Phe Leu Val
1040 1045 1050
Arg Lys Cys Pro Glu Ser Leu Asn Val Pro Glu Tyr Ala Pro Lys
1055 1060 1065
Lys Gly Phe Ser Leu His Leu Leu Phe Val Gly Ile Ser Gly Glu
1070 1075 1080
Leu Phe Lys Lys Leu Ala Leu Asn Trp Cys Pro Gly Tyr Thr Lys
1085 1090 1095
Leu Phe Pro Val Thr Gln Arg Asn Gln Asn Tyr Phe Pro Val Gln
1100 1105 1110
Phe Gln Pro Ser Asp Phe Pro Leu Ala Phe Leu Tyr Tyr His Pro
1115 1120 1125
Asp Thr Ser Ser Phe Ser Asn Ile Asp Gly Lys Val Leu Glu Met
1130 1135 1140
Arg Cys Leu Lys Arg Glu Ile Asn Tyr Val Arg Trp Glu Ile Val
1145 1150 1155
Lys Ile Arg Glu Asp Arg Gln Gln Asp Leu Lys Thr Gly Gly Tyr
1160 1165 1170
Phe Gly Asn Asp Phe Lys Thr Ala Glu Leu Thr Trp Leu Asn Tyr
1175 1180 1185
Met Asp Pro Phe Ser Phe Glu Glu Leu Ala Lys Gly Pro Ser Gly
1190 1195 1200
Met Tyr Phe Ala Gly Ala Lys Thr Gly Ile Tyr Arg Ala Gln Thr
1205 1210 1215
Ala Leu Ile Ser Phe Ile Lys Gln Glu Ile Ile Gln Lys Ile Ser
1220 1225 1230
His Gln Ser Trp Val Ile Asp Leu Gly Ile Gly Lys Gly Gln Asp
1235 1240 1245
Leu Gly Arg Tyr Leu Asp Ala Gly Val Arg His Leu Val Gly Ile
1250 1255 1260
Asp Lys Asp Gln Thr Ala Leu Ala Glu Leu Val Tyr Arg Lys Phe
1265 1270 1275
Ser His Ala Thr Thr Arg Gln His Lys His Ala Thr Asn Ile Tyr
1280 1285 1290
Val Leu His Gln Asp Leu Ala Glu Pro Ala Lys Glu Ile Ser Glu
1295 1300 1305
Lys Val His Gln Ile Tyr Gly Phe Pro Lys Glu Gly Ala Ser Ser
1310 1315 1320
Ile Val Ser Asn Leu Phe Ile His Tyr Leu Met Lys Asn Thr Gln
1325 1330 1335
Gln Val Glu Asn Leu Ala Val Leu Cys His Lys Leu Leu Gln Pro
1340 1345 1350
Gly Gly Met Val Trp Phe Thr Thr Met Leu Gly Glu Gln Val Leu
1355 1360 1365
Glu Leu Leu His Glu Asn Arg Ile Glu Leu Asn Glu Val Trp Glu
1370 1375 1380
Ala Arg Glu Asn Glu Val Val Lys Phe Ala Ile Lys Arg Leu Phe
1385 1390 1395
Lys Glu Asp Ile Leu Gln Glu Thr Gly Gln Glu Ile Gly Val Leu
1400 1405 1410
Leu Pro Phe Ser Asn Gly Asp Phe Tyr Asn Glu Tyr Leu Val Asn
1415 1420 1425
Thr Ala Phe Leu Ile Lys Ile Phe Lys His His Gly Phe Ser Leu
1430 1435 1440
Val Gln Lys Gln Ser Phe Lys Asp Trp Ile Pro Glu Phe Gln Asn
1445 1450 1455
Phe Ser Lys Ser Leu Tyr Lys Ile Leu Thr Glu Ala Asp Lys Thr
1460 1465 1470
Trp Thr Ser Leu Phe Gly Phe Ile Cys Leu Arg Lys Asn Pro Arg
1475 1480 1485
Val Lys Gln Thr Leu Asn Phe Asp Leu Leu Lys Leu Ala Gly Asp
1490 1495 1500
Val Glu Ser Asn Pro Gly Pro Thr Ser Gln Asp Leu His Ala Ile
1505 1510 1515
Gln Leu Gln Leu Glu Glu Glu Met Phe Asn Gly Gly Ile Arg Arg
1520 1525 1530
Phe Glu Ala Asp Gln Gln Arg Gln Ile Ala Ser Gly Asn Glu Ser
1535 1540 1545
Asp Thr Ala Trp Asn Arg Arg Leu Leu Ser Glu Leu Ile Ala Pro
1550 1555 1560
Met Ala Glu Gly Ile Gln Ala Tyr Lys Glu Glu Tyr Glu Gly Lys
1565 1570 1575
Arg Gly Arg Ala Pro Arg Ala Leu Ala Phe Ile Asn Cys Val Gly
1580 1585 1590
Asn Glu Val Ala Ala Tyr Ile Thr Met Lys Ile Val Met Asp Met
1595 1600 1605
Leu Asn Thr Asp Val Thr Leu Gln Ala Ile Ala Met Asn Val Ala
1610 1615 1620
Asp Arg Ile Glu Asp Gln Val Arg Phe Ser Lys Leu Glu Gly His
1625 1630 1635
Ala Ala Lys Tyr Phe Glu Lys Val Lys Lys Ser Leu Lys Ala Ser
1640 1645 1650
Lys Thr Lys Ser Tyr Arg His Ala His Asn Val Ala Val Val Ala
1655 1660 1665
Glu Lys Ser Val Ala Asp Arg Asp Ala Asp Phe Ser Arg Trp Glu
1670 1675 1680
Ala Trp Pro Lys Asp Thr Leu Leu Gln Ile Gly Met Thr Leu Leu
1685 1690 1695
Glu Ile Leu Glu Asn Ser Val Phe Phe Asn Gly Gln Pro Val Phe
1700 1705 1710
Leu Arg Thr Leu Arg Thr Asn Gly Gly Lys His Gly Val Tyr Tyr
1715 1720 1725
Leu Gln Thr Ser Glu His Val Gly Glu Trp Ile Thr Ala Phe Lys
1730 1735 1740
Glu His Val Ala Gln Leu Ser Pro Ala Tyr Ala Pro Cys Val Ile
1745 1750 1755
Pro Pro Arg Pro Trp Val Ser Pro Phe Asn Gly Gly Phe His Thr
1760 1765 1770
Glu Lys Val Ala Ser Arg Ile Arg Leu Val Lys Gly Asn Arg Glu
1775 1780 1785
His Val Arg Lys Leu Thr Lys Lys Gln Met Pro Ala Val Tyr Lys
1790 1795 1800
Ala Val Asn Ala Leu Gln Ala Thr Lys Trp Gln Val Asn Lys Glu
1805 1810 1815
Val Leu Gln Val Val Glu Asp Val Ile Arg Leu Asp Leu Gly Tyr
1820 1825 1830
Gly Val Pro Ser Phe Lys Pro Leu Ile Asp Arg Glu Asn Lys Pro
1835 1840 1845
Ala Asn Pro Val Pro Leu Glu Phe Gln His Leu Arg Gly Arg Glu
1850 1855 1860
Leu Lys Glu Met Leu Thr Pro Glu Gln Trp Gln Ala Phe Ile Asn
1865 1870 1875
Trp Lys Gly Glu Cys Thr Lys Leu Tyr Thr Ala Glu Thr Lys Arg
1880 1885 1890
Gly Ser Lys Ser Ala Ala Thr Val Arg Met Val Gly Gln Ala Arg
1895 1900 1905
Lys Tyr Ser Gln Phe Asp Ala Ile Tyr Phe Val Tyr Ala Leu Asp
1910 1915 1920
Ser Arg Ser Arg Val Tyr Ala Gln Ser Ser Thr Leu Ser Pro Gln
1925 1930 1935
Ser Asn Asp Leu Gly Lys Ala Leu Leu Arg Phe Thr Glu Gly Gln
1940 1945 1950
Arg Leu Asp Ser Ala Glu Ala Leu Lys Trp Phe Leu Val Asn Gly
1955 1960 1965
Ala Asn Asn Trp Gly Trp Asp Lys Lys Thr Phe Asp Val Arg Thr
1970 1975 1980
Ala Asn Val Leu Asp Ser Glu Phe Gln Asp Met Cys Arg Asp Ile
1985 1990 1995
Ala Ala Asp Pro Leu Thr Phe Thr Gln Trp Val Asn Ala Asp Ser
2000 2005 2010
Pro Tyr Gly Phe Leu Ala Trp Cys Phe Glu Tyr Ala Arg Tyr Leu
2015 2020 2025
Asp Ala Leu Asp Glu Gly Thr Gln Asp Gln Phe Met Thr His Leu
2030 2035 2040
Pro Val His Gln Asp Gly Ser Cys Ser Gly Ile Gln His Tyr Ser
2045 2050 2055
Ala Met Leu Ser Asp Ala Val Gly Ala Lys Ala Val Asn Leu Lys
2060 2065 2070
Pro Ser Asp Ser Pro Gln Asp Ile Tyr Gly Ala Val Ala Gln Val
2075 2080 2085
Val Ile Gln Lys Asn Tyr Ala Tyr Met Asn Ala Glu Asp Ala Glu
2090 2095 2100
Thr Phe Thr Ser Gly Ser Val Thr Leu Thr Gly Ala Glu Leu Arg
2105 2110 2115
Ser Met Ala Ser Ala Trp Asp Met Ile Gly Ile Thr Arg Gly Leu
2120 2125 2130
Thr Lys Lys Pro Val Met Thr Leu Pro Tyr Gly Ser Thr Arg Leu
2135 2140 2145
Thr Cys Arg Glu Ser Val Ile Asp Tyr Ile Val Asp Leu Glu Glu
2150 2155 2160
Lys Glu Ala Gln Arg Ala Ile Ala Glu Gly Arg Thr Ala Asn Pro
2165 2170 2175
Val His Pro Phe Asp Asn Asp Arg Lys Asp Ser Leu Thr Pro Ser
2180 2185 2190
Ala Ala Tyr Asn Tyr Met Thr Ala Leu Ile Trp Pro Ser Ile Ser
2195 2200 2205
Glu Val Val Lys Ala Pro Ile Val Ala Met Lys Met Ile Arg Gln
2210 2215 2220
Leu Ala Arg Phe Ala Ala Lys Arg Asn Glu Gly Leu Glu Tyr Pro
2225 2230 2235
Leu Pro Thr Gly Phe Ile Leu Gln Gln Lys Ile Met Ala Thr Asp
2240 2245 2250
Met Leu Arg Val Ser Thr Cys Leu Met Gly Glu Ile Lys Met Ser
2255 2260 2265
Leu Gln Ile Glu Thr Asp Val Val Asp Glu Thr Ala Met Met Gly
2270 2275 2280
Ala Ala Ala Pro Asn Phe Val His Gly His Asp Ala Ser His Leu
2285 2290 2295
Ile Leu Thr Val Cys Asp Leu Val Asp Lys Gly Ile Thr Ser Val
2300 2305 2310
Ala Val Ile His Asp Ser Phe Gly Thr His Ala Gly Arg Thr Ala
2315 2320 2325
Asp Leu Arg Asp Ser Leu Arg Glu Glu Met Val Lys Met Tyr Gln
2330 2335 2340
Asn His Asn Ala Leu Gln Asn Leu Leu Asp Val His Glu Glu Arg
2345 2350 2355
Trp Leu Val Asp Thr Gly Ile Gln Val Pro Glu Gln Gly Glu Phe
2360 2365 2370
Asp Leu Asn Glu Ile Leu Val Ser Asp Tyr Cys Phe Ala
2375 2380 2385
<210> 44
<211> 7179
<212> DNA
<213> 人工序列
<220>
<223> λ N-R341-F2A-NP868R-(G4S)2-K1ERNAP的核苷酸序列
<400> 44
atggatgctc agaccagacg cagagaacgg cgggcagaga agcaggcaca gtggaaggcc 60
gctaatggcg gtggcggcac cggtctcaaa aacaagacca gggctgagaa gtatcagacc 120
tactacacca ccaatgaata tcagatcgtt aaagaaaaac taccagacat tataagagac 180
gcggaaatta aggcgtctga agtgctcgag ccaaccatct acgagaagcg cgcaatcatg 240
gaagtcatta aggatttcat tcgggatcat caaaggaaag tgtatggcgg aacagccctg 300
aatgaggcat tgaaacaggt gaatcccaag gatgccatct atgataacta ttccttcagc 360
gacatcgagt tttattcccc tacccccgtg caggatctcg tggatctctg caacatcctg 420
tatagaaaag ggtataagtt cgtccagggg aaggacgctc agcatgagga aacctattct 480
atctttgtaa atttccagct ctactgtgac attacctatt cgccaacccg ggtcttttat 540
ggtattaaaa cgatagaaat tgacggcatt aactataccg atcctcattt catgctcata 600
gattacctcc gaatggtgaa ccagcccttg actgccgccg gccagcgctg ggagaaagcg 660
ttcgaacgga tgtacaggct gctcaaagac tatcccattg aggattttga caagaggctg 720
gatattcctg agccacccga agaaatccag agttatattt ctcggattaa gaccgagttt 780
ctgagcgata acaagctgaa tgaaagcttc ctcatctccg gcatcgaggc ttacaacttc 840
tacattcgcc atgctgcctc tagcaaagat gaagaacaga tggcccggac aaaccgcaat 900
gtggtcaatc ttaataactt tattgcaaat gtccccttta gcgagctgat ctccgtgaac 960
tatcgcgaag atgtcaagaa tacctataac ttcctgcgga tgatcgtcga ggataaagag 1020
aaaatcagtg ttgacgaata tttccctctc tttcaattca ctggctattc cactgtcatc 1080
aaatacgatg atcaccccat aattaggatc tacgagggcg acggttattg tattcctaac 1140
gtcaagaccg ttaaaacggt ggagaatgac aacggaacga agacaaagta cgagtacaag 1200
tacgtatcct tccagtacgt cctcatgatt ctatatatca acaaatttcg tgcgcacttg 1260
gacaagaata agcctatgta ttttaactac ggtattgcca tatccaatct ggtcaaagct 1320
cgcaatatat acctggacca gaccgggaaa agcgtccttg acaacactgt gtttaaggag 1380
ttccgcacta actgtaccgg aaatacgatc tctttcacac ggatgaacag actgagatta 1440
ctcgagaaaa gaaagcaggg caagcagact tcgttcgttt acacccctga agacttcttt 1500
aagaaggatc tggaaaccca agccaagctt gacccgtcga aagcgagatt caaaaatacc 1560
agtggtaaca agattatggt gccaaagtac ctgctgttca aaatagataa caacggaaat 1620
attgaagata acatacatag cgaagaggca gaaatctcag agaaagaaga aacttccggt 1680
ggctcttcta tatccactga taaatcattc gaagaatcac ctaattcctc ccctaacagc 1740
tctcctaaca actcgttgaa taattctatt gatatcagta caaataatta cgacgaccgc 1800
tcggaaaaca gcctggactc actcacgtct gatgggcccg tgaagcagac tctgaacttt 1860
gacttgttga aacttgcggg tgacgtggaa agcaacccag gccccgaatt cgccagcctg 1920
gacaacctgg tggccagata ccagcggtgc ttcaacgacc agagcctgaa gaacagcacc 1980
atcgagctgg aaatccggtt ccagcagatc aacttcctgc tgttcaagac cgtgtacgag 2040
gccctggtcg cccaggaaat ccccagcacc atcagccaca gcatccggtg catcaagaag 2100
gtgcaccacg agaaccactg ccgggagaag atcctgccca gcgagaacct gtacttcaag 2160
aaacagcccc tgatgttctt caagttcagc gagcccgcca gcctgggctg taaagtgtcc 2220
ctggccatcg agcagcccat ccggaagttc atcctggaca gcagcgtgct ggtccggctg 2280
aagaaccgga ccaccttccg ggtgtccgag ctgtggaaga tcgagctgac catcgtgaag 2340
cagctgatgg gcagcgaggt gtcagccaag ctggccgcct tcaagaccct gctgttcgac 2400
acccccgagc agcagaccac caagaacatg atgaccctga tcaaccccga cgacgagtac 2460
ctgtacgaga tcgagatcga gtacaccggc aagcctgaga gcctgacagc cgccgacgtg 2520
atcaagatca agaacaccgt gctgacactg atcagcccca accacctgat gctgaccgcc 2580
taccaccagg ccatcgagtt tatcgccagc cacatcctga gcagcgagat cctgctggcc 2640
cggatcaaga gcggcaagtg gggcctgaag agactgctgc cccaggtcaa gtccatgacc 2700
aaggccgact acatgaagtt ctaccccccc gtgggctact acgtgaccga caaggccgac 2760
ggcatccggg gcattgccgt gatccaggac acccagatct acgtggtggc cgaccagctg 2820
tacagcctgg gcaccaccgg catcgagccc ctgaagccca ccatcctgga cggcgagttc 2880
atgcccgaga agaaagagtt ctacggcttt gacgtgatca tgtacgaggg caacctgctg 2940
acccagcagg gcttcgagac acggatcgag agcctgagca agggcatcaa ggtgctgcag 3000
gccttcaaca tcaaggccga gatgaagccc ttcatcagcc tgacctccgc cgaccccaac 3060
gtgctgctga agaatttcga gagcatcttc aagaagaaaa cccggcccta cagcatcgac 3120
ggcatcatcc tggtggagcc cggcaacagc tacctgaaca ccaacacctt caagtggaag 3180
cccacctggg acaacaccct ggactttctg gtccggaagt gccccgagtc cctgaacgtg 3240
cccgagtacg cccccaagaa gggcttcagc ctgcatctgc tgttcgtggg catcagcggc 3300
gagctgttta agaagctggc cctgaactgg tgccccggct acaccaagct gttccccgtg 3360
acccagcgga accagaacta cttccccgtg cagttccagc ccagcgactt ccccctggcc 3420
ttcctgtact accaccccga caccagcagc ttcagcaaca tcgatggcaa ggtgctggaa 3480
atgcggtgcc tgaagcggga gatcaactac gtgcgctggg agatcgtgaa gatccgggag 3540
gaccggcagc aggatctgaa aaccggcggc tacttcggca acgacttcaa gaccgccgag 3600
ctgacctggc tgaactacat ggaccccttc agcttcgagg aactggccaa gggacccagc 3660
ggcatgtact tcgctggcgc caagaccggc atctacagag cccagaccgc cctgatcagc 3720
ttcatcaagc aggaaatcat ccagaagatc agccaccaga gctgggtgat cgacctgggc 3780
atcggcaagg gccaggacct gggcagatac ctggacgccg gcgtgagaca cctggtcggc 3840
atcgataagg accagacagc cctggccgag ctggtgtacc ggaagttctc ccacgccacc 3900
accagacagc acaagcacgc caccaacatc tacgtgctgc accaggatct ggccgagcct 3960
gccaaagaaa tcagcgagaa agtgcaccag atctatggct tccccaaaga gggcgccagc 4020
agcatcgtgt ccaacctgtt catccactac ctgatgaaga acacccagca ggtcgagaac 4080
ctggctgtgc tgtgccacaa gctgctgcag cctggcggca tggtctggtt caccaccatg 4140
ctgggcgaac aggtgctgga actgctgcac gagaaccgga tcgaactgaa cgaagtgtgg 4200
gaggcccggg agaacgaggt ggtcaagttc gccatcaagc ggctgttcaa agaggacatc 4260
ctgcaggaaa ccggccagga aatcggcgtc ctgctgccct tcagcaacgg cgacttctac 4320
aatgagtacc tggtcaacac cgcctttctg atcaagattt tcaagcacca tggctttagc 4380
ctcgtgcaga agcagagctt caaggactgg atccccgagt tccagaactt cagcaagagc 4440
ctgtacaaga tcctgaccga ggccgacaag acctggacca gcctgttcgg cttcatctgc 4500
ctgcggaaga acctcgaggg aggaggagga tcaggcggag gcggaagtgt cgagcaggac 4560
ctgcacgcca tccagctgca gctcgaagag gaaatgttca acggcggcat cagaagattc 4620
gaggccgacc agcagagaca gatcgcctct ggcaacgaga gcgacaccgc ctggaataga 4680
aggctgctgt ctgagctgat cgcccctatg gccgaaggca tccaggccta caaagaggaa 4740
tacgagggca agagaggcag agcccctaga gccctggcct tcatcaactg tgtgggcaat 4800
gaggtggccg cctacatcac catgaagatc gtgatggaca tgctgaacac cgacgtgacc 4860
ctgcaggcca ttgccatgaa cgtggccgac agaatcgagg accaggtccg attcagcaag 4920
ctggaaggac acgccgccaa gtacttcgag aaagtgaaga agtccctgaa ggccagcaag 4980
accaagagct acagacacgc ccacaacgtg gccgtggtgg ccgaaaaatc tgtggccgat 5040
agggacgccg acttctctag atgggaggcc tggcctaagg acaccctgct gcagatcggc 5100
atgaccctgc tggaaatcct ggaaaacagc gtgttcttca acggccagcc cgtgttcctg 5160
agaaccctga ggacaaatgg cggcaagcac ggcgtgtact acctgcagac atctgagcac 5220
gtgggcgagt ggatcaccgc cttcaaagaa catgtggccc agctgagccc tgcctatgcc 5280
ccttgtgtga tccctcctag accctgggtg tcccctttca atggcggctt tcacaccgag 5340
aaggtggcca gcagaatcag actggtcaag ggcaaccggg aacacgtgcg gaagctgacc 5400
aagaaacaga tgcccgccgt gtacaaggcc gtgaatgctc tgcaggccac caagtggcag 5460
gtcaacaaag aggtgctgca ggtcgtcgag gacgtgatca gactggatct gggctacggc 5520
gtgccaagct ttaagcccct gatcgacaga gagaacaagc ccgccaaccc tgtgcccctg 5580
gaatttcagc acctgagagg ccgcgagctg aaagagatgc tgacacctga acagtggcag 5640
gcctttatca attggaaggg cgagtgcacc aagctgtaca ccgccgagac aaagaggggc 5700
tctaagtctg ccgccacagt gcgaatggtc ggacaggcca gaaagtacag ccagttcgac 5760
gccatctact tcgtgtacgc cctggacagc cggtctagag tgtatgccca gagcagcaca 5820
ctgagccccc agtctaacga tctgggaaag gccctgctga gattcaccga gggccagaga 5880
ctggattctg ccgaagccct gaagtggttc ctggtcaacg gcgccaacaa ctggggctgg 5940
gacaagaaaa ccttcgatgt gcggaccgcc aacgtgctgg atagcgagtt ccaggacatg 6000
tgcagagata tcgccgccga ccctctgacc tttacccagt gggtcaacgc cgatagcccc 6060
tatggattcc tggcctggtg cttcgagtac gccagatacc tggacgccct ggatgaggga 6120
acccaggatc agttcatgac ccatctgccc gtgcaccagg atggctcttg ttctggcatc 6180
cagcactaca gcgccatgct gagcgatgcc gtgggagcca aagccgtgaa cctgaagcct 6240
agcgacagcc cccaggatat ctatggcgct gtggcccagg tggtcatcca gaaaaactac 6300
gcctacatga acgccgagga cgccgagaca ttcacaagcg gaagcgtgac actgacaggc 6360
gccgagctga gatctatggc ctctgcctgg gacatgatcg gcatcacacg gggcctgacc 6420
aaaaagcctg tgatgacact gccctacggc agcaccagac tgacctgtag agaaagcgtg 6480
atcgactaca tcgtggacct ggaagagaaa gaggcccaga gagccattgc cgagggcaga 6540
acagccaatc ctgtgcaccc cttcgacaac gaccggaagg atagcctgac acctagcgcc 6600
gcctacaact acatgaccgc cctgatctgg cccagcatct ctgaagtggt caaggcccct 6660
atcgtggcca tgaagatgat cagacagctg gccagattcg ccgccaagag aaatgagggc 6720
ctggaatacc ctctgcccac cggctttatc ctgcagcaga aaatcatggc caccgacatg 6780
ctgcgggtgt ccacatgtct gatgggcgag atcaagatga gcctgcagat cgagacagac 6840
gtggtggacg agacagccat gatgggagcc gccgctccta attttgtgca cggacacgat 6900
gccagccacc tgatcctgac cgtgtgcgat ctggtggaca agggcatcac tagcgtggcc 6960
gtgatccacg atagctttgg aacacacgcc ggcagaaccg ccgacctgag agattctctg 7020
cgggaagaga tggtcaagat gtaccagaac cacaacgccc tgcagaacct gctggacgtg 7080
cacgaagaaa gatggctggt ggacaccggc atccaggtgc cagaacaggg agagttcgac 7140
ctgaacgaga tcctggtgtc cgactactgc ttcgcctga 7179
<210> 45
<211> 2392
<212> PRT
<213> 人工序列
<220>
<223> λ N-R341-F2A-NP868R-(G4S)2-K1ERNAP的氨基酸序列
<400> 45
Met Asp Ala Gln Thr Arg Arg Arg Glu Arg Arg Ala Glu Lys Gln Ala
1 5 10 15
Gln Trp Lys Ala Ala Asn Gly Gly Gly Gly Thr Gly Leu Lys Asn Lys
20 25 30
Thr Arg Ala Glu Lys Tyr Gln Thr Tyr Tyr Thr Thr Asn Glu Tyr Gln
35 40 45
Ile Val Lys Glu Lys Leu Pro Asp Ile Ile Arg Asp Ala Glu Ile Lys
50 55 60
Ala Ser Glu Val Leu Glu Pro Thr Ile Tyr Glu Lys Arg Ala Ile Met
65 70 75 80
Glu Val Ile Lys Asp Phe Ile Arg Asp His Gln Arg Lys Val Tyr Gly
85 90 95
Gly Thr Ala Leu Asn Glu Ala Leu Lys Gln Val Asn Pro Lys Asp Ala
100 105 110
Ile Tyr Asp Asn Tyr Ser Phe Ser Asp Ile Glu Phe Tyr Ser Pro Thr
115 120 125
Pro Val Gln Asp Leu Val Asp Leu Cys Asn Ile Leu Tyr Arg Lys Gly
130 135 140
Tyr Lys Phe Val Gln Gly Lys Asp Ala Gln His Glu Glu Thr Tyr Ser
145 150 155 160
Ile Phe Val Asn Phe Gln Leu Tyr Cys Asp Ile Thr Tyr Ser Pro Thr
165 170 175
Arg Val Phe Tyr Gly Ile Lys Thr Ile Glu Ile Asp Gly Ile Asn Tyr
180 185 190
Thr Asp Pro His Phe Met Leu Ile Asp Tyr Leu Arg Met Val Asn Gln
195 200 205
Pro Leu Thr Ala Ala Gly Gln Arg Trp Glu Lys Ala Phe Glu Arg Met
210 215 220
Tyr Arg Leu Leu Lys Asp Tyr Pro Ile Glu Asp Phe Asp Lys Arg Leu
225 230 235 240
Asp Ile Pro Glu Pro Pro Glu Glu Ile Gln Ser Tyr Ile Ser Arg Ile
245 250 255
Lys Thr Glu Phe Leu Ser Asp Asn Lys Leu Asn Glu Ser Phe Leu Ile
260 265 270
Ser Gly Ile Glu Ala Tyr Asn Phe Tyr Ile Arg His Ala Ala Ser Ser
275 280 285
Lys Asp Glu Glu Gln Met Ala Arg Thr Asn Arg Asn Val Val Asn Leu
290 295 300
Asn Asn Phe Ile Ala Asn Val Pro Phe Ser Glu Leu Ile Ser Val Asn
305 310 315 320
Tyr Arg Glu Asp Val Lys Asn Thr Tyr Asn Phe Leu Arg Met Ile Val
325 330 335
Glu Asp Lys Glu Lys Ile Ser Val Asp Glu Tyr Phe Pro Leu Phe Gln
340 345 350
Phe Thr Gly Tyr Ser Thr Val Ile Lys Tyr Asp Asp His Pro Ile Ile
355 360 365
Arg Ile Tyr Glu Gly Asp Gly Tyr Cys Ile Pro Asn Val Lys Thr Val
370 375 380
Lys Thr Val Glu Asn Asp Asn Gly Thr Lys Thr Lys Tyr Glu Tyr Lys
385 390 395 400
Tyr Val Ser Phe Gln Tyr Val Leu Met Ile Leu Tyr Ile Asn Lys Phe
405 410 415
Arg Ala His Leu Asp Lys Asn Lys Pro Met Tyr Phe Asn Tyr Gly Ile
420 425 430
Ala Ile Ser Asn Leu Val Lys Ala Arg Asn Ile Tyr Leu Asp Gln Thr
435 440 445
Gly Lys Ser Val Leu Asp Asn Thr Val Phe Lys Glu Phe Arg Thr Asn
450 455 460
Cys Thr Gly Asn Thr Ile Ser Phe Thr Arg Met Asn Arg Leu Arg Leu
465 470 475 480
Leu Glu Lys Arg Lys Gln Gly Lys Gln Thr Ser Phe Val Tyr Thr Pro
485 490 495
Glu Asp Phe Phe Lys Lys Asp Leu Glu Thr Gln Ala Lys Leu Asp Pro
500 505 510
Ser Lys Ala Arg Phe Lys Asn Thr Ser Gly Asn Lys Ile Met Val Pro
515 520 525
Lys Tyr Leu Leu Phe Lys Ile Asp Asn Asn Gly Asn Ile Glu Asp Asn
530 535 540
Ile His Ser Glu Glu Ala Glu Ile Ser Glu Lys Glu Glu Thr Ser Gly
545 550 555 560
Gly Ser Ser Ile Ser Thr Asp Lys Ser Phe Glu Glu Ser Pro Asn Ser
565 570 575
Ser Pro Asn Ser Ser Pro Asn Asn Ser Leu Asn Asn Ser Ile Asp Ile
580 585 590
Ser Thr Asn Asn Tyr Asp Asp Arg Ser Glu Asn Ser Leu Asp Ser Leu
595 600 605
Thr Ser Asp Gly Pro Val Lys Gln Thr Leu Asn Phe Asp Leu Leu Lys
610 615 620
Leu Ala Gly Asp Val Glu Ser Asn Pro Gly Pro Glu Phe Ala Ser Leu
625 630 635 640
Asp Asn Leu Val Ala Arg Tyr Gln Arg Cys Phe Asn Asp Gln Ser Leu
645 650 655
Lys Asn Ser Thr Ile Glu Leu Glu Ile Arg Phe Gln Gln Ile Asn Phe
660 665 670
Leu Leu Phe Lys Thr Val Tyr Glu Ala Leu Val Ala Gln Glu Ile Pro
675 680 685
Ser Thr Ile Ser His Ser Ile Arg Cys Ile Lys Lys Val His His Glu
690 695 700
Asn His Cys Arg Glu Lys Ile Leu Pro Ser Glu Asn Leu Tyr Phe Lys
705 710 715 720
Lys Gln Pro Leu Met Phe Phe Lys Phe Ser Glu Pro Ala Ser Leu Gly
725 730 735
Cys Lys Val Ser Leu Ala Ile Glu Gln Pro Ile Arg Lys Phe Ile Leu
740 745 750
Asp Ser Ser Val Leu Val Arg Leu Lys Asn Arg Thr Thr Phe Arg Val
755 760 765
Ser Glu Leu Trp Lys Ile Glu Leu Thr Ile Val Lys Gln Leu Met Gly
770 775 780
Ser Glu Val Ser Ala Lys Leu Ala Ala Phe Lys Thr Leu Leu Phe Asp
785 790 795 800
Thr Pro Glu Gln Gln Thr Thr Lys Asn Met Met Thr Leu Ile Asn Pro
805 810 815
Asp Asp Glu Tyr Leu Tyr Glu Ile Glu Ile Glu Tyr Thr Gly Lys Pro
820 825 830
Glu Ser Leu Thr Ala Ala Asp Val Ile Lys Ile Lys Asn Thr Val Leu
835 840 845
Thr Leu Ile Ser Pro Asn His Leu Met Leu Thr Ala Tyr His Gln Ala
850 855 860
Ile Glu Phe Ile Ala Ser His Ile Leu Ser Ser Glu Ile Leu Leu Ala
865 870 875 880
Arg Ile Lys Ser Gly Lys Trp Gly Leu Lys Arg Leu Leu Pro Gln Val
885 890 895
Lys Ser Met Thr Lys Ala Asp Tyr Met Lys Phe Tyr Pro Pro Val Gly
900 905 910
Tyr Tyr Val Thr Asp Lys Ala Asp Gly Ile Arg Gly Ile Ala Val Ile
915 920 925
Gln Asp Thr Gln Ile Tyr Val Val Ala Asp Gln Leu Tyr Ser Leu Gly
930 935 940
Thr Thr Gly Ile Glu Pro Leu Lys Pro Thr Ile Leu Asp Gly Glu Phe
945 950 955 960
Met Pro Glu Lys Lys Glu Phe Tyr Gly Phe Asp Val Ile Met Tyr Glu
965 970 975
Gly Asn Leu Leu Thr Gln Gln Gly Phe Glu Thr Arg Ile Glu Ser Leu
980 985 990
Ser Lys Gly Ile Lys Val Leu Gln Ala Phe Asn Ile Lys Ala Glu Met
995 1000 1005
Lys Pro Phe Ile Ser Leu Thr Ser Ala Asp Pro Asn Val Leu Leu
1010 1015 1020
Lys Asn Phe Glu Ser Ile Phe Lys Lys Lys Thr Arg Pro Tyr Ser
1025 1030 1035
Ile Asp Gly Ile Ile Leu Val Glu Pro Gly Asn Ser Tyr Leu Asn
1040 1045 1050
Thr Asn Thr Phe Lys Trp Lys Pro Thr Trp Asp Asn Thr Leu Asp
1055 1060 1065
Phe Leu Val Arg Lys Cys Pro Glu Ser Leu Asn Val Pro Glu Tyr
1070 1075 1080
Ala Pro Lys Lys Gly Phe Ser Leu His Leu Leu Phe Val Gly Ile
1085 1090 1095
Ser Gly Glu Leu Phe Lys Lys Leu Ala Leu Asn Trp Cys Pro Gly
1100 1105 1110
Tyr Thr Lys Leu Phe Pro Val Thr Gln Arg Asn Gln Asn Tyr Phe
1115 1120 1125
Pro Val Gln Phe Gln Pro Ser Asp Phe Pro Leu Ala Phe Leu Tyr
1130 1135 1140
Tyr His Pro Asp Thr Ser Ser Phe Ser Asn Ile Asp Gly Lys Val
1145 1150 1155
Leu Glu Met Arg Cys Leu Lys Arg Glu Ile Asn Tyr Val Arg Trp
1160 1165 1170
Glu Ile Val Lys Ile Arg Glu Asp Arg Gln Gln Asp Leu Lys Thr
1175 1180 1185
Gly Gly Tyr Phe Gly Asn Asp Phe Lys Thr Ala Glu Leu Thr Trp
1190 1195 1200
Leu Asn Tyr Met Asp Pro Phe Ser Phe Glu Glu Leu Ala Lys Gly
1205 1210 1215
Pro Ser Gly Met Tyr Phe Ala Gly Ala Lys Thr Gly Ile Tyr Arg
1220 1225 1230
Ala Gln Thr Ala Leu Ile Ser Phe Ile Lys Gln Glu Ile Ile Gln
1235 1240 1245
Lys Ile Ser His Gln Ser Trp Val Ile Asp Leu Gly Ile Gly Lys
1250 1255 1260
Gly Gln Asp Leu Gly Arg Tyr Leu Asp Ala Gly Val Arg His Leu
1265 1270 1275
Val Gly Ile Asp Lys Asp Gln Thr Ala Leu Ala Glu Leu Val Tyr
1280 1285 1290
Arg Lys Phe Ser His Ala Thr Thr Arg Gln His Lys His Ala Thr
1295 1300 1305
Asn Ile Tyr Val Leu His Gln Asp Leu Ala Glu Pro Ala Lys Glu
1310 1315 1320
Ile Ser Glu Lys Val His Gln Ile Tyr Gly Phe Pro Lys Glu Gly
1325 1330 1335
Ala Ser Ser Ile Val Ser Asn Leu Phe Ile His Tyr Leu Met Lys
1340 1345 1350
Asn Thr Gln Gln Val Glu Asn Leu Ala Val Leu Cys His Lys Leu
1355 1360 1365
Leu Gln Pro Gly Gly Met Val Trp Phe Thr Thr Met Leu Gly Glu
1370 1375 1380
Gln Val Leu Glu Leu Leu His Glu Asn Arg Ile Glu Leu Asn Glu
1385 1390 1395
Val Trp Glu Ala Arg Glu Asn Glu Val Val Lys Phe Ala Ile Lys
1400 1405 1410
Arg Leu Phe Lys Glu Asp Ile Leu Gln Glu Thr Gly Gln Glu Ile
1415 1420 1425
Gly Val Leu Leu Pro Phe Ser Asn Gly Asp Phe Tyr Asn Glu Tyr
1430 1435 1440
Leu Val Asn Thr Ala Phe Leu Ile Lys Ile Phe Lys His His Gly
1445 1450 1455
Phe Ser Leu Val Gln Lys Gln Ser Phe Lys Asp Trp Ile Pro Glu
1460 1465 1470
Phe Gln Asn Phe Ser Lys Ser Leu Tyr Lys Ile Leu Thr Glu Ala
1475 1480 1485
Asp Lys Thr Trp Thr Ser Leu Phe Gly Phe Ile Cys Leu Arg Lys
1490 1495 1500
Asn Leu Glu Gly Gly Gly Gly Ser Gly Gly Gly Gly Ser Val Glu
1505 1510 1515
Gln Asp Leu His Ala Ile Gln Leu Gln Leu Glu Glu Glu Met Phe
1520 1525 1530
Asn Gly Gly Ile Arg Arg Phe Glu Ala Asp Gln Gln Arg Gln Ile
1535 1540 1545
Ala Ser Gly Asn Glu Ser Asp Thr Ala Trp Asn Arg Arg Leu Leu
1550 1555 1560
Ser Glu Leu Ile Ala Pro Met Ala Glu Gly Ile Gln Ala Tyr Lys
1565 1570 1575
Glu Glu Tyr Glu Gly Lys Arg Gly Arg Ala Pro Arg Ala Leu Ala
1580 1585 1590
Phe Ile Asn Cys Val Gly Asn Glu Val Ala Ala Tyr Ile Thr Met
1595 1600 1605
Lys Ile Val Met Asp Met Leu Asn Thr Asp Val Thr Leu Gln Ala
1610 1615 1620
Ile Ala Met Asn Val Ala Asp Arg Ile Glu Asp Gln Val Arg Phe
1625 1630 1635
Ser Lys Leu Glu Gly His Ala Ala Lys Tyr Phe Glu Lys Val Lys
1640 1645 1650
Lys Ser Leu Lys Ala Ser Lys Thr Lys Ser Tyr Arg His Ala His
1655 1660 1665
Asn Val Ala Val Val Ala Glu Lys Ser Val Ala Asp Arg Asp Ala
1670 1675 1680
Asp Phe Ser Arg Trp Glu Ala Trp Pro Lys Asp Thr Leu Leu Gln
1685 1690 1695
Ile Gly Met Thr Leu Leu Glu Ile Leu Glu Asn Ser Val Phe Phe
1700 1705 1710
Asn Gly Gln Pro Val Phe Leu Arg Thr Leu Arg Thr Asn Gly Gly
1715 1720 1725
Lys His Gly Val Tyr Tyr Leu Gln Thr Ser Glu His Val Gly Glu
1730 1735 1740
Trp Ile Thr Ala Phe Lys Glu His Val Ala Gln Leu Ser Pro Ala
1745 1750 1755
Tyr Ala Pro Cys Val Ile Pro Pro Arg Pro Trp Val Ser Pro Phe
1760 1765 1770
Asn Gly Gly Phe His Thr Glu Lys Val Ala Ser Arg Ile Arg Leu
1775 1780 1785
Val Lys Gly Asn Arg Glu His Val Arg Lys Leu Thr Lys Lys Gln
1790 1795 1800
Met Pro Ala Val Tyr Lys Ala Val Asn Ala Leu Gln Ala Thr Lys
1805 1810 1815
Trp Gln Val Asn Lys Glu Val Leu Gln Val Val Glu Asp Val Ile
1820 1825 1830
Arg Leu Asp Leu Gly Tyr Gly Val Pro Ser Phe Lys Pro Leu Ile
1835 1840 1845
Asp Arg Glu Asn Lys Pro Ala Asn Pro Val Pro Leu Glu Phe Gln
1850 1855 1860
His Leu Arg Gly Arg Glu Leu Lys Glu Met Leu Thr Pro Glu Gln
1865 1870 1875
Trp Gln Ala Phe Ile Asn Trp Lys Gly Glu Cys Thr Lys Leu Tyr
1880 1885 1890
Thr Ala Glu Thr Lys Arg Gly Ser Lys Ser Ala Ala Thr Val Arg
1895 1900 1905
Met Val Gly Gln Ala Arg Lys Tyr Ser Gln Phe Asp Ala Ile Tyr
1910 1915 1920
Phe Val Tyr Ala Leu Asp Ser Arg Ser Arg Val Tyr Ala Gln Ser
1925 1930 1935
Ser Thr Leu Ser Pro Gln Ser Asn Asp Leu Gly Lys Ala Leu Leu
1940 1945 1950
Arg Phe Thr Glu Gly Gln Arg Leu Asp Ser Ala Glu Ala Leu Lys
1955 1960 1965
Trp Phe Leu Val Asn Gly Ala Asn Asn Trp Gly Trp Asp Lys Lys
1970 1975 1980
Thr Phe Asp Val Arg Thr Ala Asn Val Leu Asp Ser Glu Phe Gln
1985 1990 1995
Asp Met Cys Arg Asp Ile Ala Ala Asp Pro Leu Thr Phe Thr Gln
2000 2005 2010
Trp Val Asn Ala Asp Ser Pro Tyr Gly Phe Leu Ala Trp Cys Phe
2015 2020 2025
Glu Tyr Ala Arg Tyr Leu Asp Ala Leu Asp Glu Gly Thr Gln Asp
2030 2035 2040
Gln Phe Met Thr His Leu Pro Val His Gln Asp Gly Ser Cys Ser
2045 2050 2055
Gly Ile Gln His Tyr Ser Ala Met Leu Ser Asp Ala Val Gly Ala
2060 2065 2070
Lys Ala Val Asn Leu Lys Pro Ser Asp Ser Pro Gln Asp Ile Tyr
2075 2080 2085
Gly Ala Val Ala Gln Val Val Ile Gln Lys Asn Tyr Ala Tyr Met
2090 2095 2100
Asn Ala Glu Asp Ala Glu Thr Phe Thr Ser Gly Ser Val Thr Leu
2105 2110 2115
Thr Gly Ala Glu Leu Arg Ser Met Ala Ser Ala Trp Asp Met Ile
2120 2125 2130
Gly Ile Thr Arg Gly Leu Thr Lys Lys Pro Val Met Thr Leu Pro
2135 2140 2145
Tyr Gly Ser Thr Arg Leu Thr Cys Arg Glu Ser Val Ile Asp Tyr
2150 2155 2160
Ile Val Asp Leu Glu Glu Lys Glu Ala Gln Arg Ala Ile Ala Glu
2165 2170 2175
Gly Arg Thr Ala Asn Pro Val His Pro Phe Asp Asn Asp Arg Lys
2180 2185 2190
Asp Ser Leu Thr Pro Ser Ala Ala Tyr Asn Tyr Met Thr Ala Leu
2195 2200 2205
Ile Trp Pro Ser Ile Ser Glu Val Val Lys Ala Pro Ile Val Ala
2210 2215 2220
Met Lys Met Ile Arg Gln Leu Ala Arg Phe Ala Ala Lys Arg Asn
2225 2230 2235
Glu Gly Leu Glu Tyr Pro Leu Pro Thr Gly Phe Ile Leu Gln Gln
2240 2245 2250
Lys Ile Met Ala Thr Asp Met Leu Arg Val Ser Thr Cys Leu Met
2255 2260 2265
Gly Glu Ile Lys Met Ser Leu Gln Ile Glu Thr Asp Val Val Asp
2270 2275 2280
Glu Thr Ala Met Met Gly Ala Ala Ala Pro Asn Phe Val His Gly
2285 2290 2295
His Asp Ala Ser His Leu Ile Leu Thr Val Cys Asp Leu Val Asp
2300 2305 2310
Lys Gly Ile Thr Ser Val Ala Val Ile His Asp Ser Phe Gly Thr
2315 2320 2325
His Ala Gly Arg Thr Ala Asp Leu Arg Asp Ser Leu Arg Glu Glu
2330 2335 2340
Met Val Lys Met Tyr Gln Asn His Asn Ala Leu Gln Asn Leu Leu
2345 2350 2355
Asp Val His Glu Glu Arg Trp Leu Val Asp Thr Gly Ile Gln Val
2360 2365 2370
Pro Glu Gln Gly Glu Phe Asp Leu Asn Glu Ile Leu Val Ser Asp
2375 2380 2385
Tyr Cys Phe Ala
2390
<210> 46
<211> 7170
<212> DNA
<213> 人工序列
<220>
<223> λ N-R341-T2A-NP868R-(G4S)2-K1ERNAP的核苷酸序列
<400> 46
atggatgctc agaccagacg cagagaacgg cgggcagaga agcaggcaca gtggaaggcc 60
gctaatggcg gtggcggcac cggtctcaaa aacaagacca gggctgagaa gtatcagacc 120
tactacacca ccaatgaata tcagatcgtt aaagaaaaac taccagacat tataagagac 180
gcggaaatta aggcgtctga agtgctcgag ccaaccatct acgagaagcg cgcaatcatg 240
gaagtcatta aggatttcat tcgggatcat caaaggaaag tgtatggcgg aacagccctg 300
aatgaggcat tgaaacaggt gaatcccaag gatgccatct atgataacta ttccttcagc 360
gacatcgagt tttattcccc tacccccgtg caggatctcg tggatctctg caacatcctg 420
tatagaaaag ggtataagtt cgtccagggg aaggacgctc agcatgagga aacctattct 480
atctttgtaa atttccagct ctactgtgac attacctatt cgccaacccg ggtcttttat 540
ggtattaaaa cgatagaaat tgacggcatt aactataccg atcctcattt catgctcata 600
gattacctcc gaatggtgaa ccagcccttg actgccgccg gccagcgctg ggagaaagcg 660
ttcgaacgga tgtacaggct gctcaaagac tatcccattg aggattttga caagaggctg 720
gatattcctg agccacccga agaaatccag agttatattt ctcggattaa gaccgagttt 780
ctgagcgata acaagctgaa tgaaagcttc ctcatctccg gcatcgaggc ttacaacttc 840
tacattcgcc atgctgcctc tagcaaagat gaagaacaga tggcccggac aaaccgcaat 900
gtggtcaatc ttaataactt tattgcaaat gtccccttta gcgagctgat ctccgtgaac 960
tatcgcgaag atgtcaagaa tacctataac ttcctgcgga tgatcgtcga ggataaagag 1020
aaaatcagtg ttgacgaata tttccctctc tttcaattca ctggctattc cactgtcatc 1080
aaatacgatg atcaccccat aattaggatc tacgagggcg acggttattg tattcctaac 1140
gtcaagaccg ttaaaacggt ggagaatgac aacggaacga agacaaagta cgagtacaag 1200
tacgtatcct tccagtacgt cctcatgatt ctatatatca acaaatttcg tgcgcacttg 1260
gacaagaata agcctatgta ttttaactac ggtattgcca tatccaatct ggtcaaagct 1320
cgcaatatat acctggacca gaccgggaaa agcgtccttg acaacactgt gtttaaggag 1380
ttccgcacta actgtaccgg aaatacgatc tctttcacac ggatgaacag actgagatta 1440
ctcgagaaaa gaaagcaggg caagcagact tcgttcgttt acacccctga agacttcttt 1500
aagaaggatc tggaaaccca agccaagctt gacccgtcga aagcgagatt caaaaatacc 1560
agtggtaaca agattatggt gccaaagtac ctgctgttca aaatagataa caacggaaat 1620
attgaagata acatacatag cgaagaggca gaaatctcag agaaagaaga aacttccggt 1680
ggctcttcta tatccactga taaatcattc gaagaatcac ctaattcctc ccctaacagc 1740
tctcctaaca actcgttgaa taattctatt gatatcagta caaataatta cgacgaccgc 1800
tcggaaaaca gcctggactc actcacgtct gatgggcccg caaccaactt ttctctgctg 1860
aagcaggcag gggacgttga ggagaaccct ggtcccgaat tcgccagcct ggacaacctg 1920
gtggccagat accagcggtg cttcaacgac cagagcctga agaacagcac catcgagctg 1980
gaaatccggt tccagcagat caacttcctg ctgttcaaga ccgtgtacga ggccctggtc 2040
gcccaggaaa tccccagcac catcagccac agcatccggt gcatcaagaa ggtgcaccac 2100
gagaaccact gccgggagaa gatcctgccc agcgagaacc tgtacttcaa gaaacagccc 2160
ctgatgttct tcaagttcag cgagcccgcc agcctgggct gtaaagtgtc cctggccatc 2220
gagcagccca tccggaagtt catcctggac agcagcgtgc tggtccggct gaagaaccgg 2280
accaccttcc gggtgtccga gctgtggaag atcgagctga ccatcgtgaa gcagctgatg 2340
ggcagcgagg tgtcagccaa gctggccgcc ttcaagaccc tgctgttcga cacccccgag 2400
cagcagacca ccaagaacat gatgaccctg atcaaccccg acgacgagta cctgtacgag 2460
atcgagatcg agtacaccgg caagcctgag agcctgacag ccgccgacgt gatcaagatc 2520
aagaacaccg tgctgacact gatcagcccc aaccacctga tgctgaccgc ctaccaccag 2580
gccatcgagt ttatcgccag ccacatcctg agcagcgaga tcctgctggc ccggatcaag 2640
agcggcaagt ggggcctgaa gagactgctg ccccaggtca agtccatgac caaggccgac 2700
tacatgaagt tctacccccc cgtgggctac tacgtgaccg acaaggccga cggcatccgg 2760
ggcattgccg tgatccagga cacccagatc tacgtggtgg ccgaccagct gtacagcctg 2820
ggcaccaccg gcatcgagcc cctgaagccc accatcctgg acggcgagtt catgcccgag 2880
aagaaagagt tctacggctt tgacgtgatc atgtacgagg gcaacctgct gacccagcag 2940
ggcttcgaga cacggatcga gagcctgagc aagggcatca aggtgctgca ggccttcaac 3000
atcaaggccg agatgaagcc cttcatcagc ctgacctccg ccgaccccaa cgtgctgctg 3060
aagaatttcg agagcatctt caagaagaaa acccggccct acagcatcga cggcatcatc 3120
ctggtggagc ccggcaacag ctacctgaac accaacacct tcaagtggaa gcccacctgg 3180
gacaacaccc tggactttct ggtccggaag tgccccgagt ccctgaacgt gcccgagtac 3240
gcccccaaga agggcttcag cctgcatctg ctgttcgtgg gcatcagcgg cgagctgttt 3300
aagaagctgg ccctgaactg gtgccccggc tacaccaagc tgttccccgt gacccagcgg 3360
aaccagaact acttccccgt gcagttccag cccagcgact tccccctggc cttcctgtac 3420
taccaccccg acaccagcag cttcagcaac atcgatggca aggtgctgga aatgcggtgc 3480
ctgaagcggg agatcaacta cgtgcgctgg gagatcgtga agatccggga ggaccggcag 3540
caggatctga aaaccggcgg ctacttcggc aacgacttca agaccgccga gctgacctgg 3600
ctgaactaca tggacccctt cagcttcgag gaactggcca agggacccag cggcatgtac 3660
ttcgctggcg ccaagaccgg catctacaga gcccagaccg ccctgatcag cttcatcaag 3720
caggaaatca tccagaagat cagccaccag agctgggtga tcgacctggg catcggcaag 3780
ggccaggacc tgggcagata cctggacgcc ggcgtgagac acctggtcgg catcgataag 3840
gaccagacag ccctggccga gctggtgtac cggaagttct cccacgccac caccagacag 3900
cacaagcacg ccaccaacat ctacgtgctg caccaggatc tggccgagcc tgccaaagaa 3960
atcagcgaga aagtgcacca gatctatggc ttccccaaag agggcgccag cagcatcgtg 4020
tccaacctgt tcatccacta cctgatgaag aacacccagc aggtcgagaa cctggctgtg 4080
ctgtgccaca agctgctgca gcctggcggc atggtctggt tcaccaccat gctgggcgaa 4140
caggtgctgg aactgctgca cgagaaccgg atcgaactga acgaagtgtg ggaggcccgg 4200
gagaacgagg tggtcaagtt cgccatcaag cggctgttca aagaggacat cctgcaggaa 4260
accggccagg aaatcggcgt cctgctgccc ttcagcaacg gcgacttcta caatgagtac 4320
ctggtcaaca ccgcctttct gatcaagatt ttcaagcacc atggctttag cctcgtgcag 4380
aagcagagct tcaaggactg gatccccgag ttccagaact tcagcaagag cctgtacaag 4440
atcctgaccg aggccgacaa gacctggacc agcctgttcg gcttcatctg cctgcggaag 4500
aacctcgagg gaggaggagg atcaggcgga ggcggaagtg tcgagcagga cctgcacgcc 4560
atccagctgc agctcgaaga ggaaatgttc aacggcggca tcagaagatt cgaggccgac 4620
cagcagagac agatcgcctc tggcaacgag agcgacaccg cctggaatag aaggctgctg 4680
tctgagctga tcgcccctat ggccgaaggc atccaggcct acaaagagga atacgagggc 4740
aagagaggca gagcccctag agccctggcc ttcatcaact gtgtgggcaa tgaggtggcc 4800
gcctacatca ccatgaagat cgtgatggac atgctgaaca ccgacgtgac cctgcaggcc 4860
attgccatga acgtggccga cagaatcgag gaccaggtcc gattcagcaa gctggaagga 4920
cacgccgcca agtacttcga gaaagtgaag aagtccctga aggccagcaa gaccaagagc 4980
tacagacacg cccacaacgt ggccgtggtg gccgaaaaat ctgtggccga tagggacgcc 5040
gacttctcta gatgggaggc ctggcctaag gacaccctgc tgcagatcgg catgaccctg 5100
ctggaaatcc tggaaaacag cgtgttcttc aacggccagc ccgtgttcct gagaaccctg 5160
aggacaaatg gcggcaagca cggcgtgtac tacctgcaga catctgagca cgtgggcgag 5220
tggatcaccg ccttcaaaga acatgtggcc cagctgagcc ctgcctatgc cccttgtgtg 5280
atccctccta gaccctgggt gtcccctttc aatggcggct ttcacaccga gaaggtggcc 5340
agcagaatca gactggtcaa gggcaaccgg gaacacgtgc ggaagctgac caagaaacag 5400
atgcccgccg tgtacaaggc cgtgaatgct ctgcaggcca ccaagtggca ggtcaacaaa 5460
gaggtgctgc aggtcgtcga ggacgtgatc agactggatc tgggctacgg cgtgccaagc 5520
tttaagcccc tgatcgacag agagaacaag cccgccaacc ctgtgcccct ggaatttcag 5580
cacctgagag gccgcgagct gaaagagatg ctgacacctg aacagtggca ggcctttatc 5640
aattggaagg gcgagtgcac caagctgtac accgccgaga caaagagggg ctctaagtct 5700
gccgccacag tgcgaatggt cggacaggcc agaaagtaca gccagttcga cgccatctac 5760
ttcgtgtacg ccctggacag ccggtctaga gtgtatgccc agagcagcac actgagcccc 5820
cagtctaacg atctgggaaa ggccctgctg agattcaccg agggccagag actggattct 5880
gccgaagccc tgaagtggtt cctggtcaac ggcgccaaca actggggctg ggacaagaaa 5940
accttcgatg tgcggaccgc caacgtgctg gatagcgagt tccaggacat gtgcagagat 6000
atcgccgccg accctctgac ctttacccag tgggtcaacg ccgatagccc ctatggattc 6060
ctggcctggt gcttcgagta cgccagatac ctggacgccc tggatgaggg aacccaggat 6120
cagttcatga cccatctgcc cgtgcaccag gatggctctt gttctggcat ccagcactac 6180
agcgccatgc tgagcgatgc cgtgggagcc aaagccgtga acctgaagcc tagcgacagc 6240
ccccaggata tctatggcgc tgtggcccag gtggtcatcc agaaaaacta cgcctacatg 6300
aacgccgagg acgccgagac attcacaagc ggaagcgtga cactgacagg cgccgagctg 6360
agatctatgg cctctgcctg ggacatgatc ggcatcacac ggggcctgac caaaaagcct 6420
gtgatgacac tgccctacgg cagcaccaga ctgacctgta gagaaagcgt gatcgactac 6480
atcgtggacc tggaagagaa agaggcccag agagccattg ccgagggcag aacagccaat 6540
cctgtgcacc ccttcgacaa cgaccggaag gatagcctga cacctagcgc cgcctacaac 6600
tacatgaccg ccctgatctg gcccagcatc tctgaagtgg tcaaggcccc tatcgtggcc 6660
atgaagatga tcagacagct ggccagattc gccgccaaga gaaatgaggg cctggaatac 6720
cctctgccca ccggctttat cctgcagcag aaaatcatgg ccaccgacat gctgcgggtg 6780
tccacatgtc tgatgggcga gatcaagatg agcctgcaga tcgagacaga cgtggtggac 6840
gagacagcca tgatgggagc cgccgctcct aattttgtgc acggacacga tgccagccac 6900
ctgatcctga ccgtgtgcga tctggtggac aagggcatca ctagcgtggc cgtgatccac 6960
gatagctttg gaacacacgc cggcagaacc gccgacctga gagattctct gcgggaagag 7020
atggtcaaga tgtaccagaa ccacaacgcc ctgcagaacc tgctggacgt gcacgaagaa 7080
agatggctgg tggacaccgg catccaggtg ccagaacagg gagagttcga cctgaacgag 7140
atcctggtgt ccgactactg cttcgcctga 7170
<210> 47
<211> 2389
<212> PRT
<213> 人工序列
<220>
<223> λ N-R341-T2A-NP868R-(G4S)2-K1ERNAP的氨基酸序列
<400> 47
Met Asp Ala Gln Thr Arg Arg Arg Glu Arg Arg Ala Glu Lys Gln Ala
1 5 10 15
Gln Trp Lys Ala Ala Asn Gly Gly Gly Gly Thr Gly Leu Lys Asn Lys
20 25 30
Thr Arg Ala Glu Lys Tyr Gln Thr Tyr Tyr Thr Thr Asn Glu Tyr Gln
35 40 45
Ile Val Lys Glu Lys Leu Pro Asp Ile Ile Arg Asp Ala Glu Ile Lys
50 55 60
Ala Ser Glu Val Leu Glu Pro Thr Ile Tyr Glu Lys Arg Ala Ile Met
65 70 75 80
Glu Val Ile Lys Asp Phe Ile Arg Asp His Gln Arg Lys Val Tyr Gly
85 90 95
Gly Thr Ala Leu Asn Glu Ala Leu Lys Gln Val Asn Pro Lys Asp Ala
100 105 110
Ile Tyr Asp Asn Tyr Ser Phe Ser Asp Ile Glu Phe Tyr Ser Pro Thr
115 120 125
Pro Val Gln Asp Leu Val Asp Leu Cys Asn Ile Leu Tyr Arg Lys Gly
130 135 140
Tyr Lys Phe Val Gln Gly Lys Asp Ala Gln His Glu Glu Thr Tyr Ser
145 150 155 160
Ile Phe Val Asn Phe Gln Leu Tyr Cys Asp Ile Thr Tyr Ser Pro Thr
165 170 175
Arg Val Phe Tyr Gly Ile Lys Thr Ile Glu Ile Asp Gly Ile Asn Tyr
180 185 190
Thr Asp Pro His Phe Met Leu Ile Asp Tyr Leu Arg Met Val Asn Gln
195 200 205
Pro Leu Thr Ala Ala Gly Gln Arg Trp Glu Lys Ala Phe Glu Arg Met
210 215 220
Tyr Arg Leu Leu Lys Asp Tyr Pro Ile Glu Asp Phe Asp Lys Arg Leu
225 230 235 240
Asp Ile Pro Glu Pro Pro Glu Glu Ile Gln Ser Tyr Ile Ser Arg Ile
245 250 255
Lys Thr Glu Phe Leu Ser Asp Asn Lys Leu Asn Glu Ser Phe Leu Ile
260 265 270
Ser Gly Ile Glu Ala Tyr Asn Phe Tyr Ile Arg His Ala Ala Ser Ser
275 280 285
Lys Asp Glu Glu Gln Met Ala Arg Thr Asn Arg Asn Val Val Asn Leu
290 295 300
Asn Asn Phe Ile Ala Asn Val Pro Phe Ser Glu Leu Ile Ser Val Asn
305 310 315 320
Tyr Arg Glu Asp Val Lys Asn Thr Tyr Asn Phe Leu Arg Met Ile Val
325 330 335
Glu Asp Lys Glu Lys Ile Ser Val Asp Glu Tyr Phe Pro Leu Phe Gln
340 345 350
Phe Thr Gly Tyr Ser Thr Val Ile Lys Tyr Asp Asp His Pro Ile Ile
355 360 365
Arg Ile Tyr Glu Gly Asp Gly Tyr Cys Ile Pro Asn Val Lys Thr Val
370 375 380
Lys Thr Val Glu Asn Asp Asn Gly Thr Lys Thr Lys Tyr Glu Tyr Lys
385 390 395 400
Tyr Val Ser Phe Gln Tyr Val Leu Met Ile Leu Tyr Ile Asn Lys Phe
405 410 415
Arg Ala His Leu Asp Lys Asn Lys Pro Met Tyr Phe Asn Tyr Gly Ile
420 425 430
Ala Ile Ser Asn Leu Val Lys Ala Arg Asn Ile Tyr Leu Asp Gln Thr
435 440 445
Gly Lys Ser Val Leu Asp Asn Thr Val Phe Lys Glu Phe Arg Thr Asn
450 455 460
Cys Thr Gly Asn Thr Ile Ser Phe Thr Arg Met Asn Arg Leu Arg Leu
465 470 475 480
Leu Glu Lys Arg Lys Gln Gly Lys Gln Thr Ser Phe Val Tyr Thr Pro
485 490 495
Glu Asp Phe Phe Lys Lys Asp Leu Glu Thr Gln Ala Lys Leu Asp Pro
500 505 510
Ser Lys Ala Arg Phe Lys Asn Thr Ser Gly Asn Lys Ile Met Val Pro
515 520 525
Lys Tyr Leu Leu Phe Lys Ile Asp Asn Asn Gly Asn Ile Glu Asp Asn
530 535 540
Ile His Ser Glu Glu Ala Glu Ile Ser Glu Lys Glu Glu Thr Ser Gly
545 550 555 560
Gly Ser Ser Ile Ser Thr Asp Lys Ser Phe Glu Glu Ser Pro Asn Ser
565 570 575
Ser Pro Asn Ser Ser Pro Asn Asn Ser Leu Asn Asn Ser Ile Asp Ile
580 585 590
Ser Thr Asn Asn Tyr Asp Asp Arg Ser Glu Asn Ser Leu Asp Ser Leu
595 600 605
Thr Ser Asp Gly Pro Ala Thr Asn Phe Ser Leu Leu Lys Gln Ala Gly
610 615 620
Asp Val Glu Glu Asn Pro Gly Pro Glu Phe Ala Ser Leu Asp Asn Leu
625 630 635 640
Val Ala Arg Tyr Gln Arg Cys Phe Asn Asp Gln Ser Leu Lys Asn Ser
645 650 655
Thr Ile Glu Leu Glu Ile Arg Phe Gln Gln Ile Asn Phe Leu Leu Phe
660 665 670
Lys Thr Val Tyr Glu Ala Leu Val Ala Gln Glu Ile Pro Ser Thr Ile
675 680 685
Ser His Ser Ile Arg Cys Ile Lys Lys Val His His Glu Asn His Cys
690 695 700
Arg Glu Lys Ile Leu Pro Ser Glu Asn Leu Tyr Phe Lys Lys Gln Pro
705 710 715 720
Leu Met Phe Phe Lys Phe Ser Glu Pro Ala Ser Leu Gly Cys Lys Val
725 730 735
Ser Leu Ala Ile Glu Gln Pro Ile Arg Lys Phe Ile Leu Asp Ser Ser
740 745 750
Val Leu Val Arg Leu Lys Asn Arg Thr Thr Phe Arg Val Ser Glu Leu
755 760 765
Trp Lys Ile Glu Leu Thr Ile Val Lys Gln Leu Met Gly Ser Glu Val
770 775 780
Ser Ala Lys Leu Ala Ala Phe Lys Thr Leu Leu Phe Asp Thr Pro Glu
785 790 795 800
Gln Gln Thr Thr Lys Asn Met Met Thr Leu Ile Asn Pro Asp Asp Glu
805 810 815
Tyr Leu Tyr Glu Ile Glu Ile Glu Tyr Thr Gly Lys Pro Glu Ser Leu
820 825 830
Thr Ala Ala Asp Val Ile Lys Ile Lys Asn Thr Val Leu Thr Leu Ile
835 840 845
Ser Pro Asn His Leu Met Leu Thr Ala Tyr His Gln Ala Ile Glu Phe
850 855 860
Ile Ala Ser His Ile Leu Ser Ser Glu Ile Leu Leu Ala Arg Ile Lys
865 870 875 880
Ser Gly Lys Trp Gly Leu Lys Arg Leu Leu Pro Gln Val Lys Ser Met
885 890 895
Thr Lys Ala Asp Tyr Met Lys Phe Tyr Pro Pro Val Gly Tyr Tyr Val
900 905 910
Thr Asp Lys Ala Asp Gly Ile Arg Gly Ile Ala Val Ile Gln Asp Thr
915 920 925
Gln Ile Tyr Val Val Ala Asp Gln Leu Tyr Ser Leu Gly Thr Thr Gly
930 935 940
Ile Glu Pro Leu Lys Pro Thr Ile Leu Asp Gly Glu Phe Met Pro Glu
945 950 955 960
Lys Lys Glu Phe Tyr Gly Phe Asp Val Ile Met Tyr Glu Gly Asn Leu
965 970 975
Leu Thr Gln Gln Gly Phe Glu Thr Arg Ile Glu Ser Leu Ser Lys Gly
980 985 990
Ile Lys Val Leu Gln Ala Phe Asn Ile Lys Ala Glu Met Lys Pro Phe
995 1000 1005
Ile Ser Leu Thr Ser Ala Asp Pro Asn Val Leu Leu Lys Asn Phe
1010 1015 1020
Glu Ser Ile Phe Lys Lys Lys Thr Arg Pro Tyr Ser Ile Asp Gly
1025 1030 1035
Ile Ile Leu Val Glu Pro Gly Asn Ser Tyr Leu Asn Thr Asn Thr
1040 1045 1050
Phe Lys Trp Lys Pro Thr Trp Asp Asn Thr Leu Asp Phe Leu Val
1055 1060 1065
Arg Lys Cys Pro Glu Ser Leu Asn Val Pro Glu Tyr Ala Pro Lys
1070 1075 1080
Lys Gly Phe Ser Leu His Leu Leu Phe Val Gly Ile Ser Gly Glu
1085 1090 1095
Leu Phe Lys Lys Leu Ala Leu Asn Trp Cys Pro Gly Tyr Thr Lys
1100 1105 1110
Leu Phe Pro Val Thr Gln Arg Asn Gln Asn Tyr Phe Pro Val Gln
1115 1120 1125
Phe Gln Pro Ser Asp Phe Pro Leu Ala Phe Leu Tyr Tyr His Pro
1130 1135 1140
Asp Thr Ser Ser Phe Ser Asn Ile Asp Gly Lys Val Leu Glu Met
1145 1150 1155
Arg Cys Leu Lys Arg Glu Ile Asn Tyr Val Arg Trp Glu Ile Val
1160 1165 1170
Lys Ile Arg Glu Asp Arg Gln Gln Asp Leu Lys Thr Gly Gly Tyr
1175 1180 1185
Phe Gly Asn Asp Phe Lys Thr Ala Glu Leu Thr Trp Leu Asn Tyr
1190 1195 1200
Met Asp Pro Phe Ser Phe Glu Glu Leu Ala Lys Gly Pro Ser Gly
1205 1210 1215
Met Tyr Phe Ala Gly Ala Lys Thr Gly Ile Tyr Arg Ala Gln Thr
1220 1225 1230
Ala Leu Ile Ser Phe Ile Lys Gln Glu Ile Ile Gln Lys Ile Ser
1235 1240 1245
His Gln Ser Trp Val Ile Asp Leu Gly Ile Gly Lys Gly Gln Asp
1250 1255 1260
Leu Gly Arg Tyr Leu Asp Ala Gly Val Arg His Leu Val Gly Ile
1265 1270 1275
Asp Lys Asp Gln Thr Ala Leu Ala Glu Leu Val Tyr Arg Lys Phe
1280 1285 1290
Ser His Ala Thr Thr Arg Gln His Lys His Ala Thr Asn Ile Tyr
1295 1300 1305
Val Leu His Gln Asp Leu Ala Glu Pro Ala Lys Glu Ile Ser Glu
1310 1315 1320
Lys Val His Gln Ile Tyr Gly Phe Pro Lys Glu Gly Ala Ser Ser
1325 1330 1335
Ile Val Ser Asn Leu Phe Ile His Tyr Leu Met Lys Asn Thr Gln
1340 1345 1350
Gln Val Glu Asn Leu Ala Val Leu Cys His Lys Leu Leu Gln Pro
1355 1360 1365
Gly Gly Met Val Trp Phe Thr Thr Met Leu Gly Glu Gln Val Leu
1370 1375 1380
Glu Leu Leu His Glu Asn Arg Ile Glu Leu Asn Glu Val Trp Glu
1385 1390 1395
Ala Arg Glu Asn Glu Val Val Lys Phe Ala Ile Lys Arg Leu Phe
1400 1405 1410
Lys Glu Asp Ile Leu Gln Glu Thr Gly Gln Glu Ile Gly Val Leu
1415 1420 1425
Leu Pro Phe Ser Asn Gly Asp Phe Tyr Asn Glu Tyr Leu Val Asn
1430 1435 1440
Thr Ala Phe Leu Ile Lys Ile Phe Lys His His Gly Phe Ser Leu
1445 1450 1455
Val Gln Lys Gln Ser Phe Lys Asp Trp Ile Pro Glu Phe Gln Asn
1460 1465 1470
Phe Ser Lys Ser Leu Tyr Lys Ile Leu Thr Glu Ala Asp Lys Thr
1475 1480 1485
Trp Thr Ser Leu Phe Gly Phe Ile Cys Leu Arg Lys Asn Leu Glu
1490 1495 1500
Gly Gly Gly Gly Ser Gly Gly Gly Gly Ser Val Glu Gln Asp Leu
1505 1510 1515
His Ala Ile Gln Leu Gln Leu Glu Glu Glu Met Phe Asn Gly Gly
1520 1525 1530
Ile Arg Arg Phe Glu Ala Asp Gln Gln Arg Gln Ile Ala Ser Gly
1535 1540 1545
Asn Glu Ser Asp Thr Ala Trp Asn Arg Arg Leu Leu Ser Glu Leu
1550 1555 1560
Ile Ala Pro Met Ala Glu Gly Ile Gln Ala Tyr Lys Glu Glu Tyr
1565 1570 1575
Glu Gly Lys Arg Gly Arg Ala Pro Arg Ala Leu Ala Phe Ile Asn
1580 1585 1590
Cys Val Gly Asn Glu Val Ala Ala Tyr Ile Thr Met Lys Ile Val
1595 1600 1605
Met Asp Met Leu Asn Thr Asp Val Thr Leu Gln Ala Ile Ala Met
1610 1615 1620
Asn Val Ala Asp Arg Ile Glu Asp Gln Val Arg Phe Ser Lys Leu
1625 1630 1635
Glu Gly His Ala Ala Lys Tyr Phe Glu Lys Val Lys Lys Ser Leu
1640 1645 1650
Lys Ala Ser Lys Thr Lys Ser Tyr Arg His Ala His Asn Val Ala
1655 1660 1665
Val Val Ala Glu Lys Ser Val Ala Asp Arg Asp Ala Asp Phe Ser
1670 1675 1680
Arg Trp Glu Ala Trp Pro Lys Asp Thr Leu Leu Gln Ile Gly Met
1685 1690 1695
Thr Leu Leu Glu Ile Leu Glu Asn Ser Val Phe Phe Asn Gly Gln
1700 1705 1710
Pro Val Phe Leu Arg Thr Leu Arg Thr Asn Gly Gly Lys His Gly
1715 1720 1725
Val Tyr Tyr Leu Gln Thr Ser Glu His Val Gly Glu Trp Ile Thr
1730 1735 1740
Ala Phe Lys Glu His Val Ala Gln Leu Ser Pro Ala Tyr Ala Pro
1745 1750 1755
Cys Val Ile Pro Pro Arg Pro Trp Val Ser Pro Phe Asn Gly Gly
1760 1765 1770
Phe His Thr Glu Lys Val Ala Ser Arg Ile Arg Leu Val Lys Gly
1775 1780 1785
Asn Arg Glu His Val Arg Lys Leu Thr Lys Lys Gln Met Pro Ala
1790 1795 1800
Val Tyr Lys Ala Val Asn Ala Leu Gln Ala Thr Lys Trp Gln Val
1805 1810 1815
Asn Lys Glu Val Leu Gln Val Val Glu Asp Val Ile Arg Leu Asp
1820 1825 1830
Leu Gly Tyr Gly Val Pro Ser Phe Lys Pro Leu Ile Asp Arg Glu
1835 1840 1845
Asn Lys Pro Ala Asn Pro Val Pro Leu Glu Phe Gln His Leu Arg
1850 1855 1860
Gly Arg Glu Leu Lys Glu Met Leu Thr Pro Glu Gln Trp Gln Ala
1865 1870 1875
Phe Ile Asn Trp Lys Gly Glu Cys Thr Lys Leu Tyr Thr Ala Glu
1880 1885 1890
Thr Lys Arg Gly Ser Lys Ser Ala Ala Thr Val Arg Met Val Gly
1895 1900 1905
Gln Ala Arg Lys Tyr Ser Gln Phe Asp Ala Ile Tyr Phe Val Tyr
1910 1915 1920
Ala Leu Asp Ser Arg Ser Arg Val Tyr Ala Gln Ser Ser Thr Leu
1925 1930 1935
Ser Pro Gln Ser Asn Asp Leu Gly Lys Ala Leu Leu Arg Phe Thr
1940 1945 1950
Glu Gly Gln Arg Leu Asp Ser Ala Glu Ala Leu Lys Trp Phe Leu
1955 1960 1965
Val Asn Gly Ala Asn Asn Trp Gly Trp Asp Lys Lys Thr Phe Asp
1970 1975 1980
Val Arg Thr Ala Asn Val Leu Asp Ser Glu Phe Gln Asp Met Cys
1985 1990 1995
Arg Asp Ile Ala Ala Asp Pro Leu Thr Phe Thr Gln Trp Val Asn
2000 2005 2010
Ala Asp Ser Pro Tyr Gly Phe Leu Ala Trp Cys Phe Glu Tyr Ala
2015 2020 2025
Arg Tyr Leu Asp Ala Leu Asp Glu Gly Thr Gln Asp Gln Phe Met
2030 2035 2040
Thr His Leu Pro Val His Gln Asp Gly Ser Cys Ser Gly Ile Gln
2045 2050 2055
His Tyr Ser Ala Met Leu Ser Asp Ala Val Gly Ala Lys Ala Val
2060 2065 2070
Asn Leu Lys Pro Ser Asp Ser Pro Gln Asp Ile Tyr Gly Ala Val
2075 2080 2085
Ala Gln Val Val Ile Gln Lys Asn Tyr Ala Tyr Met Asn Ala Glu
2090 2095 2100
Asp Ala Glu Thr Phe Thr Ser Gly Ser Val Thr Leu Thr Gly Ala
2105 2110 2115
Glu Leu Arg Ser Met Ala Ser Ala Trp Asp Met Ile Gly Ile Thr
2120 2125 2130
Arg Gly Leu Thr Lys Lys Pro Val Met Thr Leu Pro Tyr Gly Ser
2135 2140 2145
Thr Arg Leu Thr Cys Arg Glu Ser Val Ile Asp Tyr Ile Val Asp
2150 2155 2160
Leu Glu Glu Lys Glu Ala Gln Arg Ala Ile Ala Glu Gly Arg Thr
2165 2170 2175
Ala Asn Pro Val His Pro Phe Asp Asn Asp Arg Lys Asp Ser Leu
2180 2185 2190
Thr Pro Ser Ala Ala Tyr Asn Tyr Met Thr Ala Leu Ile Trp Pro
2195 2200 2205
Ser Ile Ser Glu Val Val Lys Ala Pro Ile Val Ala Met Lys Met
2210 2215 2220
Ile Arg Gln Leu Ala Arg Phe Ala Ala Lys Arg Asn Glu Gly Leu
2225 2230 2235
Glu Tyr Pro Leu Pro Thr Gly Phe Ile Leu Gln Gln Lys Ile Met
2240 2245 2250
Ala Thr Asp Met Leu Arg Val Ser Thr Cys Leu Met Gly Glu Ile
2255 2260 2265
Lys Met Ser Leu Gln Ile Glu Thr Asp Val Val Asp Glu Thr Ala
2270 2275 2280
Met Met Gly Ala Ala Ala Pro Asn Phe Val His Gly His Asp Ala
2285 2290 2295
Ser His Leu Ile Leu Thr Val Cys Asp Leu Val Asp Lys Gly Ile
2300 2305 2310
Thr Ser Val Ala Val Ile His Asp Ser Phe Gly Thr His Ala Gly
2315 2320 2325
Arg Thr Ala Asp Leu Arg Asp Ser Leu Arg Glu Glu Met Val Lys
2330 2335 2340
Met Tyr Gln Asn His Asn Ala Leu Gln Asn Leu Leu Asp Val His
2345 2350 2355
Glu Glu Arg Trp Leu Val Asp Thr Gly Ile Gln Val Pro Glu Gln
2360 2365 2370
Gly Glu Phe Asp Leu Asn Glu Ile Leu Val Ser Asp Tyr Cys Phe
2375 2380 2385
Ala
<210> 48
<211> 15
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 48
Gly Gly Gly Gly Ile Ala Pro Ser Met Val Gly Gly Gly Gly Ser
1 5 10 15
<210> 49
<211> 21
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 49
Ser Pro Asn Gly Ala Ser Asn Ser Gly Ser Ala Pro Asp Thr Ser Ser
1 5 10 15
Ala Pro Gly Ser Gln
20
<210> 50
<211> 15
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 50
Glu Gly Lys Ser Ser Gly Ser Gly Ser Glu Ser Lys Ser Thr Glu
1 5 10 15
<210> 51
<211> 14
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 51
Glu Gly Lys Ser Ser Gly Ser Gly Ser Glu Ser Lys Glu Phe
1 5 10
<210> 52
<211> 19
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 52
Gly Gly Gly Ser Gly Gly Gly Ser Gly Gly Gly Thr Gly Gly Gly Ser
1 5 10 15
Gly Gly Gly
<210> 53
<211> 14
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 53
Gly Ser Thr Ser Gly Ser Gly Lys Ser Ser Glu Gly Lys Gly
1 5 10
<210> 54
<211> 18
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 54
Tyr Pro Arg Ser Ile Tyr Ile Arg Arg Arg His Pro Ser Pro Ser Leu
1 5 10 15
Thr Thr
<210> 55
<211> 15
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 55
Gly Ser Thr Ser Gly Ser Gly Lys Pro Gly Ser Gly Glu Gly Ser
1 5 10 15
<210> 56
<211> 25
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 56
Ser Ser Ala Asp Asp Ala Lys Lys Asp Ala Ala Lys Lys Asp Asp Ala
1 5 10 15
Lys Lys Asp Asp Ala Lys Lys Asp Ala
20 25
<210> 57
<211> 26
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<220>
<221> misc_feature
<222> (7)..(8)
<223> Xaa可以是任何天然氨基酸
<220>
<221> misc_feature
<222> (12)..(12)
<223> Xaa可以是任何天然氨基酸
<400> 57
Gly Ser Ala Asp Asp Ala Xaa Xaa Asp Ala Ala Xaa Lys Asp Asp Ala
1 5 10 15
Lys Lys Asp Asp Ala Lys Lys Asp Gly Ser
20 25
<210> 58
<211> 25
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 58
Leu Ser Ala Asp Asp Ala Lys Lys Asp Ala Ala Lys Lys Asp Asp Ala
1 5 10 15
Lys Lys Asp Asp Ala Lys Lys Asp Leu
20 25
<210> 59
<211> 17
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 59
Ala Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys Glu Ala Ala Ala Lys
1 5 10 15
Ala
<210> 60
<211> 34
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 60
Gly Ser Thr Ser Gly Ser Gly Lys Pro Gly Ser Gly Glu Gly Ser Thr
1 5 10 15
Gly Ala Gly Gly Ala Gly Ser Thr Ser Gly Ser Gly Lys Pro Ser Gly
20 25 30
Glu Gly
<210> 61
<211> 16
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 61
Leu Ser Leu Glu Val Ala Glu Glu Ile Ala Arg Leu Glu Ala Glu Val
1 5 10 15
<210> 62
<211> 13
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 62
Gly Thr Pro Thr Pro Thr Pro Thr Pro Thr Gly Glu Phe
1 5 10
<210> 63
<211> 18
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 63
Gly Ser Thr Ser Gly Ser Gly Lys Pro Gly Ser Gly Glu Gly Ser Thr
1 5 10 15
Lys Gly
<210> 64
<211> 9
<212> PRT
<213> 人工序列
<220>
<223> 肽接头
<400> 64
Gly Ser His Ser Gly Ser Gly Lys Pro
1 5
Claims (22)
1.一种细胞质嵌合酶，其包含或由以下组成：
-至少一个加帽酶催化结构域，特别地，所述加帽酶选自由以下组成的组：cap-0典型加帽酶、cap-0非典型加帽酶、cap-1加帽酶和cap-2加帽酶；和
-至少一个蛋白质-RNA拴系系统RNA结合结构域；
其中所述RNA结合结构域与所述蛋白质-RNA拴系系统的RNA元件特异性结合，所述RNA元件由特定RNA序列和/或结构组成。
2.根据权利要求1所述的嵌合酶，其特征在于，其包含或由以下组成：
-至少一个RNA三磷酸酶催化结构域；
-至少一个鸟苷酰基转移酶催化结构域；
-至少一个N7-鸟嘌呤甲基转移酶催化结构域；和
-至少一个蛋白质-RNA拴系系统RNA结合结构域；
特别地，其中至少一个所述催化结构域是cap-0典型加帽酶的催化结构域，更特别地是病毒cap-0典型加帽酶的催化结构域。
3.根据权利要求1或2所述的嵌合酶，其中所述RNA结合结构域是噬菌体蛋白质-RNA拴系系统的噬菌体RNA结合结构域。
4.根据权利要求3所述的嵌合酶，其中所述RNA结合结构域是选自由以下组成的组的噬菌体蛋白的噬菌体RNA结合结构域：MS2外壳蛋白、R17外壳蛋白和λ形噬菌体N抗终止蛋白。
5.根据权利要求1至4中任一项所述的嵌合酶，其特征在于，其进一步包含至少一个poly(A)聚合酶催化结构域。
6.根据权利要求1至5中任一项所述的嵌合酶，其特征在于，其进一步包含至少一个DNA依赖性RNA聚合酶催化结构域，特别是噬菌体DNA依赖性RNA聚合酶催化结构域。
7.一种分离的核酸分子或一组分离的核酸分子，所述核酸分子编码根据权利要求1至6中任一项所述的嵌合酶，或者分离的核酸分子编码嵌合酶，其特征在于，其序列包含编码蛋白质-RNA拴系系统的RNA结合结构域的核酸序列，所述核酸序列与以下核酸序列框内融合，特别是按顺序框内融合：
-编码至少一个poly(A)聚合酶催化结构域的核酸序列；和
-编码至少一个加帽酶催化结构域的核酸序列；
其中所述RNA结合结构域与所述蛋白质-RNA拴系系统的RNA元件特异性结合，所述RNA元件由特定RNA序列和/或结构组成。
8.根据权利要求7所述的分离的核酸分子，其特征在于，其序列进一步包含编码至少一个DNA依赖性RNA聚合酶催化结构域，特别是噬菌体DNA依赖性RNA聚合酶催化结构域的核酸序列。
9.根据权利要求7或8所述的分离的核酸分子，其特征在于，其序列进一步包含编码核糖体跳跃基序的核酸序列，所述编码核糖体跳跃基序的核酸序列位于所述编码poly(A)聚合酶催化结构域的核酸序列与所述编码至少一个加帽酶催化结构域的核酸序列之间。
10.根据权利要求7至9中任一项所述的分离的核酸分子，其与至少一个选自由以下组成的组的启动子可操作地连接：
-RNA聚合酶II的启动子；
-噬菌体DNA依赖性RNA聚合酶的启动子；和
-权利要求6或8中所定义的所述DNA依赖性RNA聚合酶的启动子。
11.一种载体，其包含根据权利要求7至10中任一项所述的核酸分子。
12.一种宿主细胞，其包含根据权利要求7至10中任一项所述的核酸分子或根据权利要求7至10中任一项所述的一组分离的核酸分子或根据权利要求11所述的载体。
13.一种基因工程改造的非人类真核生物，其表达由根据权利要求7至10中任一项所述的核酸分子或根据权利要求7至10中任一项所述的一组分离的核酸分子编码的嵌合酶。
14.根据权利要求1至6中任一项所述的嵌合酶或根据权利要求7至10中任一项所述的分离的核酸分子或根据权利要求7至10中任一项所述的一组分离的核酸分子用于生产RNA分子的体外或离体用途，所述RNA分子具有5'-末端帽并且任选地包含至少一种化学修饰。
15.根据权利要求14所述的用途，其中所述RNA分子通过噬菌体DNA依赖性RNA聚合酶合成。
16.一种在宿主细胞中生产由DNA序列编码的具有5'-末端帽的RNA分子的体外或离体方法，所述方法包括以下步骤：在宿主细胞中表达根据权利要求7至10中任一项所述的核酸分子或根据权利要求7至10中任一项所述的一组分离的核酸分子，其中所述DNA序列与至少一个编码所述蛋白质-RNA拴系系统的RNA元件的序列共价连接，所述RNA元件与所述RNA结合结构域特异性结合。
17.根据权利要求16所述的方法，其中所述DNA序列与噬菌体DNA依赖性RNA聚合酶的启动子或与权利要求6或8中所定义的所述DNA依赖性RNA聚合酶的启动子可操作地连接。
18.一种用于生产具有5'-末端帽的RNA分子的试剂盒，其包含至少一种根据权利要求1至6中任一项所述的嵌合酶，和/或根据权利要求7至10中任一项所述的分离的核酸分子和/或根据权利要求7至10中任一项所述的一组分离的核酸分子，和/或根据权利要求11所述的载体，以及任选的，编码所述RNA分子的DNA序列，所述DNA序列与至少一个编码所述蛋白质-RNA拴系系统的RNA元件的序列共价连接，所述RNA元件与所述RNA结合结构域特异性结合。
19.根据权利要求18所述的试剂盒，其包含编码所述RNA分子的DNA序列，所述DNA序列与噬菌体DNA依赖性RNA聚合酶的启动子或与权利要求6或8中所定义的所述DNA依赖性RNA聚合酶的启动子可操作地连接。
20.根据权利要求18或19所述的试剂盒，其进一步包含分离的核酸分子，所述分离的核酸分子编码至少一个DNA依赖性RNA聚合酶催化结构域，和/或至少一个DNA依赖性RNA聚合酶催化结构域，特别是噬菌体DNA依赖性RNA聚合酶催化结构域。
21.一种药物组合物，其包含根据权利要求1至6中任一项所述的嵌合酶，和/或根据权利要求7至10中任一项所述的分离的核酸分子和/或根据权利要求7至10中任一项所述的一组分离的核酸分子，和/或根据权利要求11所述的载体。
22.根据权利要求1至6中任一项所述的嵌合酶，或根据权利要求7至10中任一项所述的分离的核酸分子和/或根据权利要求7至10中任一项所述的一组分离的核酸分子，或根据权利要求11所述的载体，其用作药物。
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| CN113862235A (zh) * | 2021-09-24 | 2021-12-31 | 苏州近岸蛋白质科技股份有限公司 | 一种嵌合酶及其在体外一步反应合成Cap0 mRNA的用途和方法 |
| CN114316053A (zh) * | 2021-12-20 | 2022-04-12 | 南京诺唯赞生物科技股份有限公司 | 一种vce的单克隆抗体及其制备方法 |
| CN114574466A (zh) * | 2022-05-09 | 2022-06-03 | 翌圣生物科技(上海)股份有限公司 | 一种嵌合加帽酶及其制备方法与应用 |
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| CN114574466A (zh) * | 2022-05-09 | 2022-06-03 | 翌圣生物科技(上海)股份有限公司 | 一种嵌合加帽酶及其制备方法与应用 |
| CN115927246A (zh) * | 2022-08-01 | 2023-04-07 | 恺佧生物科技(上海)有限公司 | 一种具有高加帽效率的牛痘加帽酶突变体 |
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| CN116083392A (zh) * | 2023-02-13 | 2023-05-09 | 深圳蓝晶生物科技有限公司 | 哺乳动物定量表达系统 |
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