CN111954683A - 感兴趣的蛋白质的展示系统 - Google Patents
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Abstract
本文描述一种蛋白质展示选择方法,其将感兴趣的蛋白质(POI)文库与展示选择系统解偶联。POI的展示可以通过表达后POI和锚蛋白之间形成共价键来实现,所述共价键可以通过酶促蛋白连接(例如SpyLigase、SnoopLigase、分选酶、蝶豆粘酶、肽基连接酶等)或自发的共价键形成(例如SpyTag/SpyCatcher,SnoopTag/SnoopCatcher等)来实现。对于最常见的噬菌体展示形式,POI文库在POI的C端融合至系连序列(例如SpyTag),然后与锚蛋白(例如,SpyCatcher融合的锚蛋白,例如SpyCatcher‑基因III蛋白(SpyCatcher‑pIII)融合体)上的捕获序列形成共价键。本文还提供了核酸构建体、宿主细胞系统和产生蛋白质展示系统的方法。
Description
此申请的序列表被记为“Seq-List.txt”,其于2019年4月1日创建,大小为70KB。序列表的全部内容通过引用整体并入本文。
背景技术
噬菌体展示
从大型文库中选择具有特定特性的蛋白质是一个熟知的过程。噬菌体展示是最古老、最常用的选择方法。它基于感兴趣的蛋白质(POI)在包含POI(例如抗体)相应DNA的噬菌体表面上的呈递。对于噬菌体展示,POI与噬菌体包衣蛋白共价连接。最常见的是,使用与丝状噬菌体M13的基因III蛋白(pIII)的遗传融合体,但文献中也有与其他包衣蛋白的融合体。通常将POI-pIII融合体克隆到噬菌粒载体中,该载体包含抗生素抗性标志物、噬菌体和细菌DNA聚合酶的复制起点以及用于将噬菌粒包装入噬菌体的噬菌体形态发生信号(Qi等,2012)。
噬菌体生产的缺失基因是通过感染辅助噬菌体(一种带有突变包装信号的全长噬菌体)来提供的。由于这种突变,被包装到噬菌体包衣中的主要是噬菌粒而不是辅助噬菌体DNA。这些噬菌体通常在表面上仅包含一个POI-pIII融合体,因为来自辅助噬菌体的野生型pIII的掺入比来自噬菌粒的融合蛋白的掺入更快。所得的单价蛋白质展示允许选择蛋白质结合强度,因为它不受任何亲合力影响。多价展示可通过使用超噬菌体实现,超噬菌体是一种修饰的辅助噬菌体,不含野生型基因III(gIII),因此,pIII的所有5个拷贝都必须来自噬菌粒(Rondot等,2001)。
除了POI和pIII之间的遗传融合,还可以通过二硫键将POI连接到噬菌体。在称为的系统中,噬菌粒上的pIII在N末端带有一个半胱氨酸,而POI在重链的C末端带有另一个半胱氨酸(Rothe等,2008)。一旦输出到周质,它们形成二硫键并结合到噬菌体包衣中。的优点是通过用还原剂(如DTT)裂解二硫键来简单、定量地洗脱特定的噬菌体POI,该方法与POI亲和力无关。这避免了偏向低亲和力POI的倾向,而低亲和力POI更容易用常规的pH转移洗脱(shift elution)来洗脱。通过包含蛋白酶可切割的接头的遗传融合体可以实现类似的优点(Ward等,1996)。
在pIII蛋白上展示POI的非共价方式是使用两个二聚化域(Wang等,2010)或蛋白AZZ域(Mazor等,2010)。
在噬菌体展示中,选择通常进行2至4轮,以富集含有所需特性(如结合靶标或酶活性)的克隆。然后对选择输出进行筛选以鉴定所需的克隆。在此步骤中,最常见的是将POI以不含pIII的方式进行表达,方法是将其亚克隆到表达载体中、通过限制性内切酶消化和再连接从载体上去除基因III、或在POI和gIII之间使用琥珀终止密码子(与缺乏抑制性tRNA的大肠杆菌菌株结合使用)。然而,为了大量表达,POI通常被亚克隆到表达载体中,该表达载体的产量高于琥珀终止密码子系统。该亚克隆步骤是费力且费时的步骤,尤其是对于并行针对多个目标的高通量选择而言。
蛋白质连接
近年来已经开发了几种使多肽在特定的预定位点共价偶联的技术。一个例子是分选酶系统(Schmohl等,2014),其中短肽(分选基序)遗传融合到一个多肽的C末端,两个甘氨酸残基遗传融合到第二个肽的N末端(或反之亦然)。在分选酶的存在下,两个修饰的多肽融合在一起。该系统的缺点是酶促反应缓慢,并且目前可获得的终产物的产率相对较低。其他酶促蛋白连接酶系统是蝶豆粘酶(Nguyen等,2014)或肽基连接酶(Toplak等,2016)。
另一个例子是编码一个或多个半胱氨酸的核苷酸向两个多肽的C-或N末端的框内添加。当这种游离的含半胱氨酸的多肽在氧化条件下混合时,它们将形成二硫键。但是,这样的系统会出现许多副产物。
第三个例子是所谓的SpyTag/SpyCatcher(Reddington等,2015)系统。在此,自然存在的蛋白质中自发的异肽形成的概念被用于将一个多肽共价连接到另一个多肽上。包含这样的异肽键的酿脓链球菌(Streptococcus pyogenes)蛋白FbaB的一个结构域被分为两部分。一个部分是SpyTag,其是一种13个氨基酸的肽,其中包含自催化中心的一部分。另一部分是SpyCatcher,其是一个116个氨基酸的蛋白质结构域,包含中心的其他部分。结果表明,将这两种多肽混合会恢复自催化中心并导致异肽键的形成,从而将SpyTag与SpyCatcher共价结合(Zakeri等,2012)。进一步的工程改造获得SpyCatcher的缩短版本,其只有84个氨基酸,以及优化的版本SpyTag002和SpyCatcher002,具有加快的反应(Keeble等,2017;在此通过引用将其全文并入本文)。该系统的另一种改进是SpyLigase的发明(Fierer等,2014),其是通过将FbaB结构域分为SpyTag、K-tag和SpyLigase三个部分实现的。SpyTag和K-tag都是通过添加SpyLigase而共价融合的短肽。
附图简要说明
图1A-1D提供了噬菌体和细胞展示(图1A-1B)和所公开的发明(图1C-1D)的图示。
图2提供了示例性的SpyCatcher-pIII序列(SEQ ID NO:29和30)。
图3A-3B示出了与VCSM13噬菌体相比较的SpyCatcher噬菌体的Western印迹分析。图3A显示了用抗SpyCatcher抗体的检测,该抗体只能检测修饰的SpyCatcher噬菌体,而不能检测VCSM13噬菌体。图3b显示了具有抗pIII检测的Western印迹,可同时检测两个噬菌体的信号,但显示SpyCatcher噬菌体的较大产物(SpyCatcher-pIII融合体)。
图4是Fab噬菌体的Western印迹。用抗pIII抗体通过Western印迹分析表达Fab的噬菌体。可见两个主要条带,Fab-SpyTag-SpyCatcher-pIII和SpyCatcher-pIII融合蛋白(图3A-3B)。这两个条带的强度之比表明平均展示率约为每个噬菌体2Fab(每个噬菌体在包衣末端均带有约5个pIII蛋白)。
图5证明了Fab在噬菌体表面上的适当折叠。通过ELISA分析展示Fab的噬菌体的抗原结合能力(TNFα用于阿达木单抗(Adalimumab),ErbB2用于曲妥单抗(Trastuzumab))。两种Fab噬菌体均显示出超过背景值的强信号,这证实了Fab在噬菌体表面上的功能。
发明内容
本文描述的发明是一种蛋白质展示选择方法,其将POI文库与展示选择系统解偶联。这是通过从文库载体中去除融合伴侣(例如,在噬菌体展示的情况下为gIIII蛋白)来实现的。POI的展示可以通过表达后POI和锚蛋白之间的共价键来实现,所述共价键可以通过酶促蛋白连接(例如SpyLigase、SnoopLigase、分选酶(sortase)、蝶豆粘酶(butelase)、肽基连接酶(peptiligase)等)或自发的共价键形成(例如SpyTag/SpyCatcher,SnoopTag/SnoopCatcher等)来实现。对于最常见的噬菌体展示形式,POI文库在POI的C端融合至系连序列(例如SpyTag),然后与锚蛋白(例如,SpyCatcher融合的锚蛋白,例如SpyCatcher-geneIII蛋白(SpyCatcher-pIII)融合体)上的捕获序列形成共价键。或者,将POI融合到捕获蛋白上,将锚蛋白融合到系连序列上。图1提供了所公开的发明的示意图。
为了使文库与锚蛋白分开,捕获-锚蛋白基因不是在噬菌粒文库上编码的,而是由修饰的辅助噬菌体、修饰的大肠杆菌基因组或其他质粒编码(用于噬菌体展示或细菌展示),或者由修饰的表达宿主或用于其他展示技术的其他质粒编码。
本发明的展示系统表达用于选择步骤的POI,而无需修饰,即不与锚蛋白融合。因此,在选择过程中表达的POI与用于以下筛选和生产步骤的蛋白质相同。规避常规展示选择系统中所需的人工融合体(例如POI-pIII)可以避免POI锚融合蛋白的不同特性(表达、溶解度、活性、稳定性等)在选择中产生的偏差。此外,由于POI未作为锚定融合蛋白表达,因此在筛选和表达之前不需要亚克隆步骤。这样可以节省时间和精力,尤其是在高通量选择设置中。
通过使文库载体与选择系统保持无关,该文库可用于多种展示选择系统,即用于噬菌体展示的不同包衣蛋白(pIII以外的包衣蛋白)以及其他展示系统(例如,酵母展示、哺乳动物或细菌展示),只要载体元件与另一宿主相容。用于从原核表达系统切换到真核表达系统的双重宿主载体的开发已被描述(例如Tesar等,2013;Batonick等,2016)。展示系统还可以在不同的选择轮次之间的选择中或在选择之后进行切换以筛选在例如其他宿主上展示的POI。
对于噬菌体展示,在第一轮选择中,在噬菌体上高POI展示率通常是有利的。在标准噬菌体展示中,这可以通过使用超噬菌体来实现,而无需使用gIII,该gIII不将POI-pIII作为pIII的唯一来源,因而产生在噬菌体表面带有多个POI的噬菌体。在这里描述的发明中,通过用携带SpyCatcher-gIII而不是gIII的修饰的辅助噬菌体来感染包含POI文库的大肠杆菌,可以实现高展示率。因此,噬菌体包衣中所有pIII的拷贝都带有SpyCatcher,它可以实现每个噬菌体的多个POI蛋白质连接。为了在接下来的轮次中实现单价展示,针对亲和力而不是亲合力进行选择,第一轮选择的噬菌体用于感染修饰的大肠杆菌(E.coli)菌株,该菌株在其基因组中携带SpyCatcher-gIII融合体或携带另外的质粒用于SpyCatcher-gIII的共表达。SpyCatcher可以与完整的gIII(N1-N2-Ct)融合或与截短或修饰的gIII(N2-Ct或Ct)融合。感染了辅助噬菌体后,细菌会产生新的噬菌体,这些噬菌体主要携带来自辅助噬菌体的野生型pIII,它被掺入噬菌体包衣的速度比来自大肠杆菌基因组/第二质粒的修饰pIII更快。或者,可以共表达另外的SpyTag或SpyTag融合蛋白,其与POI-SpyTag融合体竞争与由修饰的SpyCatcher-pIII辅助噬菌体引入的SpyCatcher的连接。可以使用其他噬菌体包衣蛋白来代替gIII。
在最后一轮选择后,不携带SpyCatcher-融合基因的未修饰大肠杆菌被噬菌体输出所感染,可用于筛选和表达。POI的C末端的SpyTag可用于检测(带有针对标签的合适抗体)以及与生物素化的、荧光或酶标记的SpyCatcher偶联。
因此,本发明的一个实施方式由POI-SpyTag融合蛋白和SpyCatcher-噬菌体包衣蛋白融合体组成,其由修饰的大肠杆菌菌株、修饰的辅助噬菌体产生,或由第二质粒共表达以在噬菌体表面展示POI。
在一个实施方式中,将POI系连序列融合蛋白与捕获序列-锚定融合蛋白连接,以在生物实体的表面上展示POI。捕获序列-锚蛋白融合体是生物实体外表面的一部分或与其相关联,因此通过在系连序列和捕获序列之间形成共价键将POI连接到生物实体表面。此外,该生物实体包含编码POI的DNA,因此提供了蛋白质选择系统所需的POI及其DNA的物理连接。在一个非限制性实施方式中,生物实体可以是具有与噬菌体包衣蛋白(例如pIII,pVI,pVII,pVIII或pIX蛋白)或这些包衣蛋白的优化、修饰或截短形式融合的捕获序列和噬菌体包衣内部的编码POI的噬菌粒的噬菌体。在其他实施方式中,生物实体可以是酵母,例如酿酒酵母(Saccharomyces cerevisiae)或毕赤酵母(Pichia pastoris)(其捕获序列与例如α-凝集素酵母粘附受体融合),细菌或哺乳动物细胞。POI可以是蛋白质或蛋白质文库,例如抗体、抗体片段、单链抗体(例如scFv、scFab)、单结构域抗体、蛋白质支架(例如基于纤连蛋白III、半胱氨酸蛋白酶抑制剂、脂蛋白、锚蛋白重复结构域、蛋白A的Z结构域等)等。该系统可用于选择具有某些特性(例如亲和力、活性或某些生物学或物理特性)的蛋白质。取决于生物实体,选择系统将被称为噬菌体展示、酵母展示、细菌展示或哺乳动物展示。
在其他实施方式中,将POI文库融合到捕获蛋白上,将锚蛋白融合到系连序列上。
在其他实施方式中,第一实施方式中所描述的系统使用SpyTag/SpyCatcher系统自发形成共价键并将POI连接至锚蛋白。SpyTag或其修饰或优化版本(例如SpyTag002)是系连序列,而SpyCatcher或其修饰或优化版本(例如SpyCatcher002)是捕获序列。POI与SpyTag融合、锚蛋白与SpyCatcher融合,或者POI与SpyCatcher融合、SpyTag与锚蛋白融合。
在其他实施方式中,第一实施方式中描述的系统使用与SpyTag/SpyCatcher不同的蛋白质连接系统来将POI连接至锚蛋白,例如SnoopTag/SnoopCatcher(Veggiani等,2016)、SpyTag/KTag/SpyLigase(Fierer等,2014)、分选酶系统(Schmohl等,2014)、分裂内含蛋白(Thiel等,2014)、蝶豆粘酶1(Nguyen等,2014)、肽基连接酶(Toplak等,2016)或其他类似方法。
在又一个实施方式中,表达了POI,例如在重链或轻链的C末端具有系连序列、以及一个或多个可用于抗体纯化或检测的其他标签的抗体片段或单链抗体文库。丝状噬菌体,例如M13,是生物实体。通过用修饰的辅助噬菌体对携带带有POI-系连序列融合基因的噬菌粒的细菌的感染来实现在噬菌体表面上展示有POI的噬菌体的生产。修饰的辅助噬菌体产生捕获序列-pIII融合体,而不是野生型pIII。修饰的pIII可能在捕获序列和pIII之间包含蛋白酶切割位点,例如TEV、鼻病毒3C蛋白酶或胰蛋白酶,用于通过蛋白酶切割洗脱选择的噬菌体。此外,携带有编码捕获序列-pIII融合蛋白的质粒的细菌或在其基因组中携带捕获序列-pIII基因的经修饰的细菌可以与辅助噬菌体感染结合使用,以产生新的POI展示噬菌体。可以使用优化的、修饰的或截短的pIII融合体来代替完整的pIII融合体。
在又一个实施方式中,表达了POI,例如在重链或轻链的C末端具有捕获序列、以及一个或多个可用于抗体纯化或检测的其他标签的抗体片段或单链抗体文库。丝状噬菌体,例如M13,是生物实体。通过用修饰的辅助噬菌体对携带带有POI-捕获序列融合基因的噬菌粒的细菌的感染来实现在噬菌体表面上展示有POI的噬菌体的生产。修饰的辅助噬菌体产生系连序列-pIII融合体,而不是野生型pIII。修饰的pIII可能在系连序列和pIII之间包含蛋白酶切割位点,例如TEV、鼻病毒3C蛋白酶或胰蛋白酶,用于通过蛋白酶切割洗脱选择的噬菌体。此外,携带有编码系连序列-pIII融合蛋白的质粒的细菌或在其基因组中携带系连序列-pIII基因的经修饰的细菌可以与辅助噬菌体感染结合使用,以产生新的POI展示噬菌体。可以使用优化的、修饰的或截短的pIII融合体来代替完整的pIII融合体。
在本说明书和权利要求中,所用的单数形式“一个”、“一种”和“该”包括多个指示物,除非上下文中有明显的表示。
如本文所用,术语“遗传构建体”是指编码感兴趣的蛋白质(POI)的多核苷酸序列,该感兴趣的蛋白质在框内与“系连序列”(TS)或捕获序列(CS)或蛋白质(也称为“锚蛋白”)融合,其在表面上表达或与原核细胞、真核细胞、原核病毒或噬菌体的表面连接,或真核病毒,其在框内与捕获序列或系连序列融合。为了便于参考,GC1是指包含融合至系连序列或捕获序列的POI的遗传构建体,GC2是指包含融合至锚蛋白的捕获序列或系连序列的遗传构建体。
术语“系连序列”或“系链序列”是指连接至POI或锚蛋白并促进与在生物实体表面上表达或与POI融合的捕获部分的共价键形成的序列。系连序列的非限制性实例包括SpyTag序列,包括SpyTag002、SnoopTag序列、分选酶基序、C肽、蝶豆粘酶底物和肽基连接酶底物。系连序列可以在这样的蛋白质或多肽的N-或C-末端或内部环中与POI或锚蛋白融合。特别地,间隔序列(例如,富含甘氨酸/丝氨酸的间隔子)可以侧接系连序列,以增强反应的可及性。间隔子可以进一步包括特异性蛋白水解的位点(例如,通过因子X、凝血酶、肠激酶、烟草蚀刻病毒(TEV)NIa蛋白酶、鼻病毒3C蛋白酶或胰蛋白酶),从而允许从系连序列中特异性释放。
术语“捕获序列”是指与锚蛋白或POI融合的多肽,通过其与系连序列形成共价肽键,例如用于SpyTag系连序列的SpyCatcher。捕获序列可以在这样的蛋白质或多肽的N-或C-末端或内部环中与锚蛋白或POI融合。特别地,间隔序列(例如,富含甘氨酸/丝氨酸的间隔子)可以侧接捕获序列,以增强反应的可及性。间隔子可以进一步包括特异性蛋白水解的位点(例如,通过因子X、凝血酶、肠激酶、烟草蚀刻病毒NIa蛋白酶、鼻病毒3C蛋白酶或胰蛋白酶),从而允许从捕获序列中特异性释放。
术语“原核系统”是指原核细胞,例如细菌细胞或原核病毒、原核噬菌体或细菌孢子。术语“真核系统”是指真核细胞,包括动物、植物、真菌和原生生物的细胞,以及真核病毒,例如逆转录病毒、腺病毒、杆状病毒。原核和真核系统可以统称为“表达系统”。
术语“表达盒”在本文中是指出于表达重组蛋白/肽的目的而构建在载体中的功能单元。表达盒包括一个或多个启动子、转录终止子序列、核糖体结合位点或核糖体结合位点,以及编码系连序列或捕获序列的cDNA。取决于表达系统(例如,用于真核表达系统的增强子和聚腺苷酸化信号),可以将其他遗传组件添加至表达盒。
如本文所用,术语“载体”是指核酸分子,优选自我复制的核酸分子,其将插入的核酸分子转移至宿主细胞内和/或之间。通常,载体是包含复制起点、选择标志物和/或病毒包装信号以及其他调控元件的环状DNA。在本发明的描述中,载体、载体DNA、质粒DNA、噬菌粒DNA是可互换的术语。该术语包括主要起到将DNA或RNA插入细胞的功能的载体,主要起到DNA或RNA的复制的功能的复制载体,以及起到DNA或RNA的转录和/或翻译的功能的表达载体。还包括提供超过一种上述功能的载体。
如本文所用,术语“表达载体”是多核苷酸,当引入合适的宿主细胞中时,其可以被转录并翻译成多肽。术语“表达载体”是指指导在框内与系连序列融合的感兴趣的蛋白质的可溶性表达的载体,所述系连序列的特征在于能够与本文所述的辅助载体产生的捕获序列形成与捕获序列的肽键。POI还可以与捕获序列框内融合,该捕获序列的特征在于能够与本文所述的辅助载体产生的系连序列形成与系连序列的肽键。
术语“辅助载体”是指遗传系统或宿主细胞特异性载体,其设计用于产生融合蛋白,所述融合蛋白包含在框内与捕获序列融合的锚蛋白。锚蛋白也可以在框内与系连序列融合。辅助载体可以与表达载体结合、通过瞬时共转化、永久整合入宿主基因组,或通过宿主细胞的病毒或噬菌体感染而被引入表达系统。
如本文所用,术语“展示载体组”是指表达载体(其提供GC1构建体)和辅助载体(其提供GC2构建体)的特定组合。GC1和GC2的共表达导致POI在表达系统的表面上展示。
如本文所用,术语“锚蛋白”是指其一部分位于表达系统的外表面或细胞膜之外的多肽或蛋白质。捕获序列或系连序列与在细胞外表面上发现的锚蛋白的一部分(通常是锚蛋白的N-末端部分)融合(参见例如图1)。锚蛋白的非限制性实例在表1中提供或已经在其他出版物中公开,例如Little等,其通过引用整体并入本文。
如果将噬菌体展示系统(例如噬菌体M13病毒颗粒)用作表达系统,则捕获序列或系连序列可以在框内融合至包衣蛋白,例如在基因III、VI、VII、VIII和IX中,修饰或截短的包衣蛋白例如为长、短或超短版本的pIII。或者,捕获序列或系连序列可以在细菌细胞如大肠杆菌的表面上表达。例如,捕获序列可以融合至锚蛋白,例如外膜蛋白(Chang和Lo,2000;Lee等,2004),菌毛和鞭毛(Westerlund-Wikstrom等,1997),修饰的脂蛋白(Georgiou等,1996),冰核蛋白(Jung等,1998)或自转运蛋白(Veiga等,2003)。
捕获序列也可以展示在真核宿主细胞如酿酒酵母的表面上。例如,酿酒酵母(S.cerevisiae)中的细胞壁蛋白α-凝集素1和α-凝集素2、Cwp1、Cwp2、Gas1p、Yap3p、Flo1p、Crh2p、Pir1、Pir2、Pir4和Icwp;多形汉逊酵母(Hansenula polymorpha)中的HpSEDI、HpGASI、HpTIPI、HPWPI和白色念珠菌(Candida albicans)中的HwpIp、Als3p、Rbt5p可用于在酵母细胞表面展示捕获序列。
同样,可以操纵哺乳动物细胞以在细胞表面表达捕获序列。例如,捕获序列可以在膜锚蛋白上表达,例如细胞表面受体(Chesnut等,1996,J Immunological Methods;Ho等,2006,PNAS,103:9637-9642),GPI锚序列(美国专利第6,838,446号),不可切割的11型信号锚序列(美国专利第7,125,973号)。
如本文所用,术语“多核苷酸”、“核酸”、“核苷酸”和“寡核苷酸”可互换使用。它们指任何长度的核苷酸聚合形式,不论是脱氧核糖核苷酸或核糖核苷酸或它们的类似物。以下是多核苷酸的非限制性例子:基因或基因片段的编码或非编码区域、由连锁分析定义的基因座、外显子、内含子、信使RNA(mRNA)、转移RNA、核糖体RNA、核酶、cDNA、重组多核苷酸、支链多核苷酸、质粒、载体、任意序列的分离DNA、任意序列的分离RNA、核酸探针和引物。多核苷酸可包括修饰的核苷酸,如甲基化的核苷酸和核苷酸类似物。如果存在,对核苷酸结构的修饰可在核苷酸聚合物的组装之前或之后赋予。
本文所用术语“氨基酸”指天然和/或非天然或合成的氨基酸,包括甘氨酸,以及D或L光学异构体,以及氨基酸类似物和肽模拟物。
如本文所用,术语“多肽”、“肽”和“蛋白质”在本文中可互换使用,指任何长度的氨基酸的聚合物。因此,POI可以是多肽、肽或蛋白质。
“感兴趣的蛋白质”(POI)是任何所需的多肽、肽或蛋白质。POI的非限制性实例包括抗体,例如全长抗体、抗体片段、单链抗体(例如,scFv,scFab)、或单结构域抗体,蛋白支架(例如基于纤连蛋白III、半胱氨酸蛋白酶抑制剂、脂蛋白、锚蛋白重复结构域、蛋白A的Z结构域等),激素,白介素,用于开发疫苗的抗原,酶等。其他实例包括但不限于:人生长激素(hGH),N-甲硫基人生长激素,牛生长激素,甲状旁腺激素,甲状腺素,胰岛素A链,胰岛素B链,胰岛素原,松弛素A链,松弛素B链,前松弛素,糖蛋白激素例如卵泡刺激素(FSH)、甲状腺刺激素(TSH)、黄体化激素(LH),糖蛋白激素受体,降钙素,胰高血糖素,VIII因子,抗体,肺表面活性剂,尿激酶,链激酶,人类组织型纤溶酶原激活物(t-PA),蛙皮素,IX因子,凝血酶,造血生长因子,肿瘤坏死因子-α和-β,脑啡肽酶,人血清白蛋白,穆勒氏抑制物,小鼠促性腺激素相关肽,微生物蛋白,例如β-内酰胺酶,组织因子蛋白,抑制素,激活素,血管内皮生长因子,激素或生长因子的受体;整联蛋白,血小板生成素,蛋白A或D,类风湿因子,神经生长因子例如NGF-β,血小板生长因子,转化生长因子(TGF),例如TGF-α和TGF-β,胰岛素样生长因子I和-II,胰岛素样生长因子结合蛋白,CD-4,DNA酶,潜伏期相关肽,促红细胞生成素,骨诱导因子,干扰素例如干扰素-α、-β和-γ,集落刺激因子(CSF)例如M-CSF、GF-CSF和G-CSF,白介素(IL)例如IL-1、IL-2、IL-3、IL-4,超氧化物歧化酶;衰变促进因子,病毒抗原,HIV包膜蛋白例如GP120、GP140,心房利钠肽A、B或C,免疫球蛋白以及上述任何蛋白的片段。
如本文所用,术语“宿主细胞”包括可以是或已经是所公开载体的受体的单个细胞或细胞培养物。宿主细胞包括单个宿主细胞的后代。由于自然、偶然或故意的突变,后代不一定与原始亲代细胞完全相同。
如本文所用,如果多肽或多核苷酸序列与比较序列均显示出实质性的氨基酸或核苷酸序列同源性,则这两个序列“基本上相同”或“基本上相似”。通常,在与同源区域比对后,基本相同或大致相同的序列彼此至少约60%相同。优选地,序列至少约70%相同,更优选地,至少约80%相同,更优选地,至少约90%相同,更优选地,序列至少约95%相同。
系连和捕获序列
系连和捕获序列是蛋白质或肽序列,其可以在添加或涉及或不添加或不涉及其他因素(蛋白质、酶、催化剂、盐等)的情况下自发或酶促地形成共价键。下面列出了一些非限制性实例。
SpyTag/SpyCatcher
美国专利第9,547,003号(其公开内容通过引用整体并入本文)公开了SpyTag/SpyCatcher系统的组件(用作系连)。如本文中所讨论的,在这方面,系连序列的长度(例如,长度约为10、20、30、40或50个氨基酸)为约5-50个氨基酸,并且衍生自SEQ ID NO:1、3、5或6。捕获序列也衍生自SEQ ID NO:1、3、5或6,并且可以具有任何长度。
系连序列和捕获序列可以在这样的蛋白质或多肽的N-或C-末端或内部环中与POI融合。特别地,间隔序列(例如,富含甘氨酸/丝氨酸的间隔子)可以侧接系连序列或捕获序列,以增强反应的可及性。间隔子可以进一步包括特异性蛋白水解的位点(例如,通过因子X、凝血酶、肠激酶或烟草蚀刻病毒NIa蛋白酶),从而允许从系连或捕获序列中特异性释放。因此,系连序列包含SEQ ID NO:1所示序列的残基302-308、SEQ ID NO:25(mgsshhhhhhssglvprgsv ptivmvdayk rykgsgesgk)、SEQ ID NO:27(VPTIVMVDAYKRYKS),或与SEQ IDNO:1、25或27具有至少50%同一性的序列,其中所述肽标签的长度小于50个氨基酸。在某些实施方式中,系连序列与SEQ ID NO:1具有至少约55%,约60%,约65%,约70%,约75%,约80%,约85%,约90%,或约95%的同一性,并且长度小于50个氨基酸。更特别地,系连序列可包含SEQ ID NO:1的残基301-308、300-308、299-308、298-308、297-308、296-308、295-308、294-308、293-308、292-308、291-308或290-308或与SEQ ID NO:1或25的残基302-308具有至少约50%至95%同一性的序列。优选地,系连序列包含SEQ ID NO:1中303位的反应性天冬酰胺,即,该残基优选是不变的。此外,系连序列可以是SEQ ID NO:1或25的片段,并且在一个优选的实施方式中,使用少于50个氨基酸的系连序列,其包含SEQ ID NO:1所示序列的残基293-308,或包含与其具有至少50%同一性的序列。肽标签受长度限制并且包含少于50个氨基酸残基。因此,肽标签不包含SEQ ID NO:1的序列而仅包含其特定片段,或包含与此类特定片段具有至少50%同一性(例如75、80、85、90或95%同一性)的序列。其他实施方式利用SEQ ID NO:25或与其具有至少50%序列同一性的序列作为系连序列。捕获序列包含以下或由以下组成:SEQ ID NO:1所示的序列的残基31-291,SEQ ID NO:26(msyyhhhhhhdydipttenl yfqgamvttl sglsgeqgps gdmtteedsa thikfskrde dgrelagatm elrdssgktistwisdghvk dfylypgkyt fvetaapdgy evataitftv neqgqvtvng eatkgdaht),SEQ ID NO:28(yfqgamvttl sglsgeqgps gdmtteedsa thikfskrde dgrelagatm elrdssgktistwisdghvk dfylypgkyt fvetaapdgy evataitftv neqgqvtvng eatkgdaht),或与SEQ IDNO:1的残基32-291、SEQ ID NO:26或SEQ ID NO:28(yfqgamvttl sglsgeqgps gdmtteedsathikfskrde dgrelagatm elrdssgkti stwisdghvk dfylypgkyt fvetaapdgy evataitftvneqgqvtvng eatkgdaht)具有至少50%同一性(例如75、80、85、90、95、96、97、98或99%同一性)的序列。明确排除的是SEQ ID NO:1中列出的完整序列,但是,捕获序列应包含与SEQ IDNO:1的179位相对应的反应性赖氨酸。特别地,结合伴侣包含SEQ ID NO:1所示序列的残基31-292、31-293、31-294、31-295、31-296、31-297、31-298、31-299、31-300、31-301或31-302或与其具有至少70%同一性的序列,但SEQ ID NO:1的序列除外。
另外,可以使用N末端的替代异肽键,从主要的菌毛蛋白Spy0128设计系连序列。因此,系连序列可以被设计或可以从异肽蛋白质的N-末端片段获得,并且剩余的、截短的或重叠的蛋白质片段可以构成捕获序列。在SEQ ID NO:1的36位发现了与结合于N-末端的异肽键有关的反应性赖氨酸,并且在SEQ ID NO:1的168位发现了与该异肽键有关的反应性天冬酰胺。因此,在一个优选的实施方式中,在这种情况下,系连序列包含反应性赖氨酸残基,而捕获序列包含反应性天冬酰胺。特别地,系连序列包含SEQ ID NO:1所示序列的残基31-40或与其具有至少70%同一性并且长度小于50个氨基酸的序列。上述系连序列的相应捕获序列包含SEQ ID NO:1所示序列的残基37-304或具有与其至少70%同一性的序列,但SEQ IDNO:1的序列除外。优选地,肽标签和结合伴侣中的反应性残基不被突变。
另一个系连序列包含SEQ ID NO:3所示序列的残基179-184(例如173-185)或与其具有至少50%同一性并且长度小于50个氨基酸的序列。捕获序列包含SEQ ID NO:3的残基191-317(例如186-318)或与其具有至少50%同一性的序列,但SEQ ID NO:3除外。SEQ IDNO:3的全长序列被特别排除作为系连序列或捕获序列。
另一系连序列包含在266位包括天冬酰胺的SEQ ID NO:5的片段(或与其具有至少50%同一性的序列)以及捕获序列,该捕获序列包含SEQ ID NO:5的片段,与它们具有至少50%的序列同一性,并且在149位包含赖氨酸残基,但在266位不包含天冬酰胺。系连序列或捕获序列均不包含SEQ ID NO:5。
在另一个实施方式中,系连序列包含在101位包含天冬氨酸残基的SEQ ID NO:6的片段(或至少70%与之相同的序列),并且捕获序列包含包含15位的反应性赖氨酸的SEQ IDNO:6的片段(或至少50%与之相同的序列)。系连序列或捕获序列均不包含SEQ ID NO:6。
另一个实施方式提供了包含SEQ ID NO:25(mgsshhhhhh ssglvprgsv ptivmvdaykrykgsgesgk)、SEQ ID NO:27(VPTIVMVDAYKRYKS),或与SEQ ID NO:25或27具有至少50%同一性的序列的系连序列,其中所述肽标签的长度为15至40或50个氨基酸。在某些实施方式中,系连序列与SEQ ID NO:25或27具有至少约55%,约60%,约65%,约70%,约75%,约80%,约85%,约90%,或约95%的同一性,并且长度小于50个氨基酸。优选地,系连序列包含SEQ ID NO:27中8位的反应性天冬氨酸,即,该残基优选是不变的。
为SEQ ID NO:25、27或与SEQ ID NO:25或27具有至少50%同一性的序列的捕获序列长度为15至40或50个氨基酸,并包含SEQ ID NO:27中8位的天冬氨酸,包含以下或由以下组成:SEQ ID NO:26(msyyhhhhhh dydipttenl yfqgamvttl sglsgeqgps gdmtteedsathikfskrde dgrelagatm elrdssgkti stwisdghvk dfylypgkyt fvetaapdgy evataitftvneqgqvtvng eatkgdaht),或yfqgamvttl sglsgeqgps gdmtteedsa thikfskrdedgrelagatm elrdssgkti stwisdghvk dfylypgkyt fvetaapdgy evataitftv neqgqvtvngeatkgdaht,或与其具有至少50%同一性,例如与SEQ ID NO:26或SEQ ID NO:28(yfqgamvttl sglsgeqgps gdmtteedsa thikfskrde dgrelagatm elrdssgkti stwisdghvkdfylypgkyt fvetaapdgy evataitftv neqgqvtvng eatkgdaht)具有75、75、80、85、90、95、96、97、98或99%同一性的序列。变体具有至少50%的序列同一性并保留SEQ ID NO:26的57位的赖氨酸。
SpyLigase/SnoopLigase
或者,可以使用WO2016/193746(其通过引用整体并入本文)中描述的系统将POI附着到暴露的表面蛋白上。在这样的实施方式中,系连序列任选地通过接头序列(例如富含甘氨酸/丝氨酸的间隔子)附接到POI和锚蛋白两者上。然后将这些系连序列由宿主细胞同样编码的连接酶连接。在一些实施方式中,系连序列的长度可以为6-50个氨基酸,例如7-45、8-40、9-35、10-30、11-25个氨基酸,例如其可以包含6、7、8、9、10、11、12、13、14、15、16、17、18、19或20个氨基酸或由6、7、8、9、10、11、12、13、14、15、16、17、18、19或20个氨基酸组成。其他实施方式提供了长度为约20-300个氨基酸的系连序列(例如,约10、20、30、40、50、60、70等氨基酸长度)。在一些实施方式中,肽连接酶的长度可以在50-300个氨基酸之间(例如长度为60-250、70-225、80-200个氨基酸),例如,它可以包含60、65、70、75、80、85、90、95、100、110、120、130、140、150、160、170、180、190或200个氨基酸或由60、,65、70、75、80、85、90、95、100、110、120、130、140、150、160、170、180、190或200个氨基酸组成,条件是其满足以下针对连接酶提出的定义。附着于锚蛋白的系连序列对应于本发明该方面的捕获序列。
一对系连序列可以衍生自任何合适的异肽蛋白。例如,系连序列可衍生自主要菌毛蛋白Spy0128,其具有如SEQ ID NO:7所示的氨基酸序列并且由如SEQ ID NO:8所示的核苷酸序列编码。在蛋白质中形成两个异肽键。在SEQ ID NO:7的179位的赖氨酸和SEQ IDNO:7的303位的天冬酰胺(反应残基)之间形成有一个异肽键。在SEQ ID NO:7的258位发现了诱导自发异肽键的谷氨酸残基。因此,由SEQ ID NO:7所示的异肽蛋白质形成的一对系连序列将优选地包含含有在303位包含反应性天冬酰胺的蛋白质的片段的系连序列、以及含有在179位包含反应性赖氨酸的蛋白质的片段的系连序列。可以分别提供在258位上包含谷氨酸残基的蛋白质的片段,即,作为形成异肽键的肽连接酶。
主要菌毛蛋白Spy0128中的另一个异肽键出现在SEQ ID NO:7的36位赖氨酸残基和SEQ ID NO:7的168位天冬酰胺残基之间。在SEQ ID NO:7的117位发现了诱导异肽形成的谷氨酸残基。因此,由SEQ ID NO:7所示的异肽蛋白质形成的一对系连序列将优选地包含含有在36位包含反应性赖氨酸残基的蛋白质的片段的系连序列、以及含有在168位包含反应性天冬酰胺的蛋白质的片段的系连序列。在117位包含谷氨酸残基的蛋白质的片段可以作为肽连接酶单独提供。
在SEQ ID NO:9的181位的赖氨酸残基(ACE19,粪肠球菌(E.faecalis)的粘附素蛋白的结构域)和SEQ ID NO:9的294位的天冬酰胺残基之间存在异肽键。该键由SEQ ID NO:9中213位的天冬氨酸残基诱导。因此,由SEQ ID NO:9所示的异肽蛋白质形成的一对系连序列将优选地包含含有在294位包含反应性天冬酰胺的蛋白质的片段的系连序列、以及含有在181位包含反应性赖氨酸残基的蛋白质的片段的系连序列。在213位包含天冬氨酸残基的蛋白质的片段可以作为肽连接酶单独提供。
也可以使用具有SEQ ID NO:10所示氨基酸序列的金黄色葡萄球菌(S.aureus)的胶原结合域。异肽键出现在SEQ ID NO:10的176位的赖氨酸和SEQ ID NO:10的308位的天冬酰胺之间。诱导异肽键的天冬氨酸残基在SEQ ID NO:10的209位。因此,由SEQ ID NO:10所示的异肽蛋白质形成的一对系连序列将优选地包含含有在176位包含反应性赖氨酸的蛋白质的片段的系连序列、以及含有在308位包含反应性天冬酰胺的蛋白质的片段的系连序列。在209位包含天冬氨酸残基的蛋白质的片段可以作为肽连接酶单独提供。
化脓性链球菌(Streptococcus pyogenes)的FbaB也可用于提供系连序列,并包含具有由SEQ ID NO:11所示的氨基酸序列,由SEQ ID NO:12所示的核苷酸序列编码的结构域CnaB2。CnaB2域中的异肽键在SEQ ID NO:11的15位赖氨酸和SEQ ID NO:11的101位天冬氨酸残基之间形成。诱导异肽键的谷氨酸残基在SEQ ID NO:11的61位。因此,由SEQ ID NO:11所示的异肽蛋白质形成的一对系连序列将优选地包含含有在15位包含反应性赖氨酸的蛋白质的片段的系连序列、以及含有在101位包含反应性天冬氨酸的蛋白质的片段的系连序列。在61位包含谷氨酸残基的蛋白质的片段可以作为肽连接酶单独提供。
RrgA蛋白是肺炎链球菌(Streptococcus pneumoniae)的粘附蛋白,其具有如SEQID NO:13所示的氨基酸序列并且由如SEQ ID NO:14所示的核苷酸序列编码。在SEQ ID NO:13的742位的赖氨酸和SEQ ID NO:13的854位的天冬酰胺之间形成有一个异肽键。该键由SEQ ID NO:13中803位的谷氨酸残基诱导。因此,由SEQ ID NO:13所示的异肽蛋白质形成的一对系连序列将优选地包含含有在854位包含反应性天冬酰胺的蛋白质的片段的系连序列、以及含有在742位包含反应性赖氨酸的蛋白质的片段的系连序列。如上所述,在803位包含谷氨酸残基的蛋白质的片段可以作为肽连接酶单独提供。
PsCs蛋白是中间链球菌(Streptococcus intermedius)的por分泌系统C末端分选域蛋白的片段,其具有如SEQ ID NO:15所示的氨基酸序列并且由如SEQ ID NO:16所示的核苷酸序列编码。在SEQ ID NO:15的405位的赖氨酸和SEQ ID NO:15的496位的天冬氨酸之间形成有一个异肽键。因此,由SEQ ID NO:15所示的异肽蛋白质形成的一对系连序列将优选地包含含有在496位包含反应性天冬氨酸的蛋白质的片段的系连序列、以及含有在405位包含反应性赖氨酸的蛋白质的片段的系连序列。
在各种实施方式中,系连序列可以衍生自包含SEQ ID NO:21、23、25或27中任一项所示的氨基酸序列的异肽蛋白,或与SEQ ID NO:21、23、25或27中任一项所示的氨基酸序列具有至少70%序列同一性的蛋白质。在一些实施方案中,上述所述异肽蛋白质序列与其比较的序列(SEQ ID NO:21、23、25或27)至少75、80、85、90、95、96、97、98、99或100%相同。
分选酶
将POI系连到锚蛋白的另一种方法包括使用分选酶和分选酶识别和桥接结构域。Schmohl等(2014年)讨论了分选酶介导的针对蛋白质的位点特异性修饰的连接,在此全文引入作为参考。在本发明的这个方面,分选酶识别和桥接结构域被认为是系连序列,并且与锚蛋白融合的系连序列对应于本发明的这个方面的捕获序列。分选酶是由革兰氏阳性细菌产生的转肽酶,以将细胞表面蛋白共价锚定在细胞壁上。金黄色葡萄球菌(Staphylococcusaureus)分选酶A(SrtA)切割短的C末端识别基序(LPXTG(SEQ ID NO:17))(在本文中称为分选酶识别结构域)。分选酶识别结构域是分选酶A识别结构域或分选酶B识别结构域。在特定的实施方式中,分选酶识别结构域包含以下氨基酸序列或由其组成:LPTGAA(SEQ ID NO:18),LPTGGG(SEQ ID NO:19),LPKTGG(SEQ ID NO:20),LPETG(SEQ ID NO:21),LPXTG(SEQID NO:22)或LPXTG(X)n(SEQ ID NO:23),其中X是任何氨基酸,n为0、1、2、3、4、5、7、8、9、10,在0-5或0-10的范围内,或不超过100的任何整数。可以任选地通过富含甘氨酸/丝氨酸的间隔子在框内将分选酶识别结构域与POI或锚蛋白融合。如果它与POI相连,则被视为系连序列;当它与锚蛋白相连时,则被视为捕获序列。
分选酶A桥接结构域在其末端之一包含一个或多个甘氨酸残基。在某些实施方式中,一个或多个甘氨酸残基可以任选为:Gly,(Gly)2,(Gly)3,(Gly)4,或(Gly)x,其中x为1-20的整数。可以任选地通过富含甘氨酸/丝氨酸的间隔子连接分选酶桥接结构域与POI或锚蛋白。如果它与POI相连,则被视为系连序列;当它与锚蛋白相连时,则被视为捕获序列。
分选酶B识别域包含氨基酸序列NPX1TX2,其中X1是谷氨酰胺或赖氨酸;X2是天冬酰胺或甘氨酸;N是天冬酰胺;P是脯氨酸,T是苏氨酸。分选酶B。可以任选地通过富含甘氨酸/丝氨酸的间隔子在框内将分选酶识别结构域与POI或锚蛋白融合。如果它与POI相连,则被视为系连序列;当它与锚蛋白相连时,则被视为捕获序列。
分选酶B桥接结构域在其末端之一包含一个或多个甘氨酸残基。在某些实施方式中,一个或多个甘氨酸残基可以任选为:Gly,(Gly)2,(Gly)3,(Gly)4,或(Gly)x,其中x为1-20的整数。可以任选地通过富含甘氨酸/丝氨酸的间隔子连接分选酶桥接结构域与POI或锚蛋白。如果它与POI相连,则被视为系连序列;当它与锚蛋白相连时,则被视为捕获序列。
蝶豆粘酶1
将POI系连到锚蛋白的另一种方法包括使用蝶豆粘酶1在蝶豆粘酶识别基序(其中Asx是Asn或Asp)和另一多肽的氨基末端之间形成肽键。在该情况下,可以任选地通过富含甘氨酸/丝氨酸的间隔子在框内将Asx-His-Val基序与POI或锚蛋白融合。然后,可使用蝶豆粘酶在Asx-His-Val基序和锚蛋白的N端氨基酸之间形成肽键。WO 2017/058114公开了用于蝶豆粘酶介导的肽连接的方法和材料,其通过引用整体并入本文。
分裂内含蛋白
将POI系连到锚蛋白的另一种方法包括使用分裂内含蛋白。内含蛋白可以由两个分别转录和翻译的基因编码的两个片段存在。这些所谓的分裂内含蛋白以反式自缔合并催化蛋白质剪接活性。已在多种蓝细菌和古细菌中鉴定了分裂内含蛋白(Caspi等,2003;ChoiJ.等,2006;Dassa B.等,2007;Liu X.和Yang J.,2003;Wu H.等,1998;和Zettler J.等,2009,其公开内容通过引用整体并入本文)。Thiel等(2014)和WO 2013/045632也公开了可用于融合异源蛋白质的分裂内含蛋白的用途,其均以整体通过引用并入本文。
载体
本发明的载体通常包含表达POI和锚蛋白所需的转录或翻译控制序列。合适的转录或翻译控制序列包括但不限于复制起点、启动子、增强子、阻遏物结合区、转录起始位点、核糖体结合位点、翻译起始位点和针对转录和翻译的终止位点。
复制起点(通常称为ori序列)允许载体在合适的宿主细胞中复制。ori的选择将取决于所用宿主细胞和/或基因包装的类型。在宿主细胞是原核生物的情况下,表达载体通常包含两个ori序列,一个指导载体在原核细胞内的自主复制,另一个ori支持噬菌体颗粒的包装。优选的原核ori能够指导细菌细胞中的载体复制。此类ori的非限制性实例包括pMB1、pUC以及其他大肠杆菌来源。支持噬菌体颗粒包装的优选的ori包括但不限于f1 ori、Pf3噬菌体复制ori。
在真核生物系统中,高级真核生物包含DNA复制的多个起点,但是ori序列并未清楚定义。哺乳动物载体的合适复制起点通常来自真核病毒。优选的真核ori包括但不限于SV40 ori、EBV ori或HSV ori。
如本文所用,“启动子”是在某些条件下能够结合RNA聚合酶并启动位于启动子下游(在3′方向)的编码区的转录的DNA区域。它可以是组成型的或诱导型的。通常,启动子序列在其3′末端与转录起始位点结合,并向上游(5′方向)延伸,以包括在高于背景的可检测水平启动转录所必需的最小数目的碱基或元件。启动子序列内是转录起始位点以及负责RNA聚合酶结合的蛋白质结合结构域。真核启动子通常但并不总是包含“TATA”盒和“CAT”盒。
启动子的选择将在很大程度上取决于引入载体的宿主细胞。对于原核细胞,本领域已知多种稳健的启动子。优选的启动子是lac启动子、Trc启动子、T7启动子和pBAD启动子。通常,为了获得外源序列在多种物种中的表达,可以将原核启动子直接置于真核启动子之后,或置于真核启动子下游的内含子序列内部。
真核细胞的合适启动子序列包括3-磷酸甘油酸激酶或其他糖酵解酶,如烯醇酶,3-磷酸甘油醛脱氢酶,己糖激酶,丙酮酸脱羧酶,磷酸果糖激酶,葡萄糖-6磷酸异构酶,3-磷酸甘油酸突变酶,丙酮酸激酶,三糖磷酸异构酶,磷酸葡萄糖异构酶和葡萄糖激酶的启动子。具有由生长条件控制的转录的额外优势的其他启动子是用于醇脱氢酶2、异细胞色素C、酸性磷酸酶,与氮代谢相关的降解酶和上述的3-磷酸甘油醛脱氢酶,以及负责麦芽糖和半乳糖利用的酶的启动子区域。哺乳动物细胞的优选启动子是SV40启动子、CMV启动子、β-肌动蛋白启动子及其杂合体。酵母细胞的优选启动子包括但不限于酿酒酵母(S.cerevisiae)中的GAL 10、GAL I、TEFI,以及巴斯德毕赤酵母(P.pastoris)中的GAP、AOX1。
在构建本发明载体中,与外源序列相关的终止序列也被插入需要转录的序列的3′端,以提供mRNA和/或转录终止信号的多腺苷酸化。终止子序列优选包含一个或多个转录终止序列(如聚腺苷酸化序列),并且还可以通过包含额外的DNA序列而被延长,从而进一步破坏转录通读。本发明优选的终止子序列(或终止位点)具有一个基因,其后是转录终止序列,为其自身的终止序列或异源终止序列。这样的终止序列的实例包括与各种酵母转录终止序列、或本领域已知的并且广泛可得的哺乳动物聚腺苷酸化序列偶联的终止密码子。在终止子包含基因的情况下,使用编码可检测或选择性标志物的基因可能是有利的;从而提供了一种手段,通过该手段可以检测和/或选择终止子序列的存在和/或不存在(以及由此转录单位的相应的失活和/或激活)。
除了上述元件之外,载体可以包含选择性标志物(例如,编码用载体转化的宿主细胞存活或生长所必需的蛋白质的基因),尽管这种标志物基因可以携带在共同引入宿主细胞中的另一个多核苷酸序列上。只有那些已导入选择性基因的宿主细胞才能在选择条件下存活和/或生长。典型的选择基因编码蛋白质,其将(a)对抗生素或其他毒素例如氨苄青霉素、卡那霉素、新霉素、博来霉素、G418,甲氨蝶呤等赋予抗性;(b)弥补营养缺陷型缺陷;或(c)提供无法从复杂培养基中获得的关键营养素。合适的标志物基因的选择将取决于宿主细胞,并且不同宿主的合适基因是本领域已知的。
在一个实施方案中,表达载体是穿梭载体,其能够在至少两个不相关的宿主系统中复制。为了促进这种复制,载体通常包含至少两个复制起点,一个在每个宿主系统中有效。通常,穿梭载体能够在真核宿主系统和原核宿主系统中复制。这使得能够检测真核宿主(表达细胞类型)中的蛋白质表达和原核宿主(扩增细胞类型)中的载体扩增。尽管可以使用本领域已知的任何合适的起点,只要其指导载体的复制即可,优选地,一个复制起点源自SV40或2u,一个复制起点源自pUC。当载体是穿梭载体时,该载体优选包含至少两种选择标志物,一种用于表达细胞类型,一种用于扩增细胞类型。可以使用本领域已知的任何选择标志物或本文所述的那些,只要其在所利用的表达系统中起作用即可。
可以使用重组克隆方法和/或通过化学合成获得本发明涵盖的载体。大量的重组克隆技术如PCR、限制性核酸内切酶消化和连接在本领域中是众所周知的,在此无需详细描述。本领域技术人员还可以使用本文提供的序列数据或公共数据库或专有数据库中的序列数据,以通过本领域可用的任何合成手段获得所需载体。另外,可以使用众所周知的限制和连接技术从各种DNA来源中切出合适的序列,并与根据本发明待表达的外源序列可操作地整合在一起。
本文所引用或引用的所有专利、专利申请、临时申请和出版物(包括所有附图和表格)均通过引用全文并入本文,只要它们与本说明书的明确教导不矛盾即可。
以下是说明实施本发明的方法的实施例。这些示例不应被解释为限制性的。
实施例1-共价捕获展示
该实施例描述了人Fab片段、阿达木单抗(adalimumab)和曲妥珠单抗(trastuzumab)的共价展示,其在M13噬菌体表面重链的C末端带有SpyTag,所述M13噬菌体具有与pIII的N末端融合的SpyCatcher。SpyTag和SpyCatcher在大肠杆菌内形成共价键,导致Fab与pIII噬菌体包衣蛋白共价连接。
修饰辅助噬菌体以产生SpyCatcher-pIII
修饰了VCSM13辅助噬菌体,用SpyCatcher-gIII基因融合体替代gIII。通过限制性消化的PCR产物的连接进行克隆。噬菌体基因组在gIII的开始处(带有XbaI位点的正向引物)和gIII的起始密码子之前(带有EcoRI位点的反向引物)用引物进行PCR扩增并插入限制性酶切位点,从而省略gIII信号肽。具有匹配的限制性位点的插入序列由DsbA信号肽、SpyCatcher、TEV蛋白酶切割位点和短接头(GGGGSGGGGS,图2)组成。转化入大肠杆菌XL1Blue F′感受态细胞后,通过集落PCR鉴定出具有正确插入片段的克隆,以将较短的wt gIII与SpyCatcher-gIII区分开来。通过对PCR扩增的噬菌体DNA的相关区域进行测序来确认正确插入。
SpyCatcher-pIII辅助噬菌体的生产
携带经修饰的辅助噬菌体基因组的XL1F’在含卡那霉素的50ml2xYT中于37℃过夜生长。第二天,接种来自过夜培养物的六个300ml培养物至OD600为0.1,并在37℃下在含卡那霉素的2xYT中生长至OD600为约0.6,然后转移至18℃过夜进行辅助噬菌体生产。如下所述旋转细菌,通过PEG沉淀从上清液中纯化并浓缩噬菌体。将噬菌体沉淀溶于1000μ1PBS中,加入20%甘油等分试样后,于-80℃保存。
将XL1 Blue F′、MC1061 F′或TG1F′细胞在2xYT中生长至OD600为0.6-0.8,然后用修饰的辅助噬菌体感染45分钟(不振荡)和45分钟(振荡)。将培养物在振荡器上于37℃孵育3-4小时,然后转移至含有50μg/ml卡那霉素的较大体积的2xYT(例如50ml培养物和350ml2xYT)中,以在18℃过夜生产噬菌体。第二天,如下所述沉淀噬菌体,将其溶解于400μ lPBS中,并在添加20%甘油后储存在-80℃。
噬菌体PEG沉淀
通过加入1/4体积+10%的冰冷的PEG/NaCl溶液(20%PEG 6000,2.5M NaCl),从过夜培养上清液中沉淀出噬菌体。将上清液在振荡器上冰上孵育30分钟,并将噬菌体在2-8℃下以13,000x g离心60分钟。弃去上清液,将沉淀重悬于10ml PBS中。通过用0.22μm的过滤器进行无菌过滤,从噬菌体溶液中除去残留的细菌,并如上所述重复进行PEG/NaCl沉淀。将噬菌体沉淀溶解在PBS中。
SpyCatcher-pIII辅助噬菌体分析
1.斑点滴定
通过点滴定和ELISA测定噬菌体滴度和感染性。为了进行斑点滴定,用M9最小琼脂平板上的TG1F′集落接种培养物,并在37℃下生长到OD600为0.6-0.8。制备了2xYT中1∶10辅助噬菌体稀释系列,并将10μl稀释液与90μlTG1F′培养物混合,在37℃下孵育30分钟,然后将5μl铺板在含有1%葡萄糖和50μg/ml卡那霉素的LB琼脂平板上,并在37℃下孵育过夜。未修饰的VCSM13辅助噬菌体用作参照。对少于30个集落的斑点中的集落进行计数,并根据稀释度和使用量计算滴度。滴度为约1011cfu/ml。
2.ELISA
通过ELISA确定噬菌体滴度和在噬菌体表面上展示SpyCatcher的证据。将ELISA微量滴定板在4℃包被一抗M13抗体(抗pVIII,在PBS中为1μl/ml)和抗SpyCatcher Fab(在PBS中为5μg/ml)过夜。在室温下在TBST中用化学阻滞剂封闭1-2小时后,将具有已知滴度的1∶2稀释系列的修饰辅助噬菌体和wt VCSM13辅助噬菌体添加到ELISA板中,并在室温下孵育1小时。用抗M13 HRP检测抗体和QuantaBlu过氧化物酶荧光底物进行检测。通过该方法估计的滴度为约5×1013噬菌体/ml。用抗SpyCatcher捕获抗体的ELISA给出了一个强信号,因此证实了噬菌体的存在,其表面上展示了一个或多个SpyCatcher。
3.Western印迹
通过分别用抗-SpyCatcher抗体和抗-pIII抗体的变性噬菌体的Western印迹也证实了SpyCatcher在噬菌体表面上的存在。将Bio-Rad Mini-PROTEANTM垂直电泳池与Mini-Protean TGX 4-20%凝胶和Bio-Rad Precision Plus蛋白质标准分子量标志物一起使用。通过使用Bio-Rad Trans-Blot Turbo转移系统将蛋白质印迹到PVDF膜上。将膜用牛奶在TBST中在振荡器上摇动1小时封闭,然后与两种一抗之一孵育1小时。抗SpyCatcher一抗的二抗是HRP偶联的抗hFab抗体,对于抗pIII则使用HRP偶联的抗小鼠IgG抗体。使用ClarityWestern ECL底物进行检测。
具有抗SpyCatcher检测的Western印迹仅显示SpyCatcher噬菌体的条带,而不显示VCSM13噬菌体的条带(阴性对照;图3A)。使用抗pIII检测的印迹显示出SpyCatcher噬菌体中的pIII带比VCSM13噬菌体高。该位移等于融合到pIII的SpyCatcher的分子量增加(图3B)。
Fab-SpyTag构建体
将在重链C末端带有SpyTag(Spy)和六组氨酸标签(H)的人Fab抗体片段(曲妥珠单抗和阿达木单抗)克隆到双顺反子表达载体pBBx1中,该双顺反子表达载体由lac启动子、Shine-Dalgarno(SD)序列、起始密码子、pelB前导序列、Fab轻链终止密码子、SD序列、起始密码子、ompA前导序列Fab重链、SpyTag、His标签和终止密码子组成。该载体还带有ColE1和f1复制起点,氯霉素抗性和lac阻遏物(LacI)。
共价捕获展示
将上述质粒转化到TG1F′细胞中。将新鲜接种的培养物在含氯霉素和1%葡萄糖的5ml 2xYT中于振荡器上于37℃过夜生长。第二天,用该预培养物接种100ml培养物至OD 600为0.05,并在37℃下生长至OD 600为0.5。在37℃下将50ml的这种培养物用SpyCatcher-pIII辅助噬菌体感染45分钟(不振荡)和45分钟(振荡)。离心分离细菌,弃去上清液。将沉淀重悬于含氯霉素、卡那霉素和0.25mM IPTG的300ml 2xYT中,并在30℃下培养过夜以产生噬菌体。如前所述,离心分离细菌并从上清液中PEG沉淀出噬菌体。将噬菌体沉淀溶解于500-1000μlPBS中,并在添加20%甘油后储存在-80℃。
共价捕获展示Fab噬菌体的分析
1.斑点滴定
如对SpyCatcher-pIII噬菌体所述进行斑点滴定。滴度为1-3×1013 cfu。
2.ELISA
如对SpyCatcher-pIII噬菌体所述进行噬菌体ELISA。具有已知滴度的Fab展示噬菌体用作确定噬菌体滴度的标准。用这种方法测定的滴度在1×1014噬菌体/ml的范围内。
使用抗Fd捕获抗体来代替抗SpyCatcher抗体用于第二ELISA。该抗体可捕获在噬菌体表面带有Fab重链(Fd)的噬菌体,因此该测定证实了成功的Fab共价捕获展示。该ELISA给出强信号,因此表明每个噬菌体最可能有几个Fab的良好展示率。
3.Western印迹
用抗pIII抗体通过Western印迹分析Fab噬菌体。如SpyCatcher-pIII噬菌体所述进行Western印迹。可见两个主要条带,Fab-SpyTag-SpyCatcher-pIII和SpyCatcher-pIII融合蛋白(图4)。这两个条带的强度之比表明平均展示率约为每个噬菌体2个Fab(每个噬菌体在包衣末端均带有约5个pIII蛋白)。
4.功能-抗原结合
为了确认Fab在噬菌体表面上的正确折叠,还通过ELISA分析了Fab展示噬菌体的抗原结合能力。Fab抗原(阿达木单抗为TNF-α和曲妥珠单抗为ErbB2)在ELISA板上以5μg/ml的浓度在PBS中于4℃包被过夜。用TBST中的化学阻滞剂封闭1小时后,将板与TBST中的1∶500Fab噬菌体稀释液一起孵育1小时。用偶联HRP的抗M13抗体和QuantaBlu荧光底物检测噬菌体。对于两种Fab噬菌体,超过背景值的强信号证实了Fab在噬菌体表面上的功能(图5)。
5.噬菌体感染后的噬菌体DNA回收
为了证实Fab展示噬菌体携带噬菌粒TG1F’被Fab噬菌体感染,并通过PCR分析单个集落的DNA以检测噬菌粒。
用10μl Fab噬菌体感染OD600为0.4-0.6的2xYT中的1ml TG1F′培养物,并在37℃下孵育45分钟(不振荡),然后45分钟(振荡)。将不同量的该培养物(10μl,1μl和0.1μl)铺板在含氯霉素和葡萄糖的LB琼脂平板上。挑出单个集落用于集落PCR,该集落PCR用在pBBx1噬菌粒中退火的引物进行。通过琼脂糖凝胶电泳确认所有集落均具有正确大小的PCR产物。
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序列表
<110> 生物辐射ABD瑟罗泰克有限公司(Bio-Rad ABD Serotec GmbH)
<120> 感兴趣的蛋白质的展示系统
<130> BRL.119XC1PCT
<150> US 62/652,938
<151> 2018-04-05
<160> 30
<170> PatentIn version 3.5
<210> 1
<211> 340
<212> PRT
<213> 化脓性链球菌(Streptococcus pyogenes)
<400> 1
Met Lys Leu Arg His Leu Leu Leu Thr Gly Ala Ala Leu Thr Ser Phe
1 5 10 15
Ala Ala Thr Thr Val His Gly Glu Thr Val Val Asn Gly Ala Lys Leu
20 25 30
Thr Val Thr Lys Asn Leu Asp Leu Val Asn Ser Asn Ala Leu Ile Pro
35 40 45
Asn Thr Asp Phe Thr Phe Lys Ile Glu Pro Asp Thr Thr Val Asn Glu
50 55 60
Asp Gly Asn Lys Phe Lys Gly Val Ala Leu Asn Thr Pro Met Thr Lys
65 70 75 80
Val Thr Tyr Thr Asn Ser Asp Lys Gly Gly Ser Asn Thr Lys Thr Ala
85 90 95
Glu Phe Asp Phe Ser Glu Val Thr Phe Glu Lys Pro Gly Val Tyr Tyr
100 105 110
Tyr Lys Val Thr Glu Glu Lys Ile Asp Lys Val Pro Gly Val Ser Tyr
115 120 125
Asp Thr Thr Ser Tyr Thr Val Gln Val His Val Leu Trp Asn Glu Glu
130 135 140
Gln Gln Lys Pro Val Ala Thr Tyr Ile Val Gly Tyr Lys Glu Gly Ser
145 150 155 160
Lys Val Pro Ile Gln Phe Lys Asn Ser Leu Asp Ser Thr Thr Leu Thr
165 170 175
Val Lys Lys Lys Val Ser Gly Thr Gly Gly Asp Arg Ser Lys Asp Phe
180 185 190
Asn Phe Gly Leu Thr Leu Lys Ala Asn Gln Tyr Tyr Lys Ala Ser Glu
195 200 205
Lys Val Met Ile Glu Lys Thr Thr Lys Gly Gly Gln Ala Pro Val Gln
210 215 220
Thr Glu Ala Ser Ile Asp Gln Leu Tyr His Phe Thr Leu Lys Asp Gly
225 230 235 240
Glu Ser Ile Lys Val Thr Asn Leu Pro Val Gly Val Asp Tyr Val Val
245 250 255
Thr Glu Asp Asp Tyr Lys Ser Glu Lys Tyr Thr Thr Asn Val Glu Val
260 265 270
Ser Pro Gln Asp Gly Ala Val Lys Asn Ile Ala Gly Asn Ser Thr Glu
275 280 285
Gln Glu Thr Ser Thr Asp Lys Asp Met Thr Ile Thr Phe Thr Asn Lys
290 295 300
Lys Asp Phe Glu Val Pro Thr Gly Val Ala Met Thr Val Ala Pro Tyr
305 310 315 320
Ile Ala Leu Gly Ile Val Ala Val Gly Gly Ala Leu Tyr Phe Val Lys
325 330 335
Lys Lys Asn Ala
340
<210> 2
<211> 1023
<212> DNA
<213> 化脓性链球菌(Streptococcus pyogenes)
<400> 2
atgaaattac gtcacttact attaacggga gcagccctaa ctagttttgc tgctacaaca 60
gttcacgggg agactgttgt aaacggagcc aaactaacag ttacaaaaaa ccttgattta 120
gttaatagca atgcattaat tccaaataca gattttacat ttaaaatcga acctgatact 180
actgtcaacg aagacggaaa taagtttaaa ggtgtagctt tgaacacacc gatgactaaa 240
gtcacttaca ccaattcaga taaaggtgga tcaaatacga aaactgcaga atttgatttt 300
tcagaagtta cttttgaaaa accaggtgtt tattattaca aagtaactga ggagaagata 360
gataaagttc ctggtgtttc ttatgataca acatcttaca ctgttcaagt tcatgtcttg 420
tggaatgaag agcaacaaaa accagtagct acttatattg ttggttataa agaaggtagt 480
aaggtgccaa ttcagttcaa aaatagctta gattctacta cattaacggt gaagaaaaaa 540
gtttcaggta ccggtggaga tcgctctaaa gattttaatt ttggtctgac tttaaaagca 600
aatcagtatt ataaggcgtc agaaaaagtc atgattgaga agacaactaa aggtggtcaa 660
gctcctgttc aaacagaggc tagtatagat caactctatc attttacctt gaaagatggt 720
gaatcaatca aagtcacaaa tcttccagta ggtgtggatt atgttgtcac tgaagacgat 780
tacaaatcag aaaaatatac aaccaacgtg gaagttagtc ctcaagatgg agctgtaaaa 840
aatatcgcag gtaattcaac tgaacaagag acatctactg ataaagatat gaccattact 900
tttacaaata aaaaagactt tgaagtgcca acaggagtag caatgactgt ggcaccatat 960
attgctttag gaattgtagc agttggtgga gctctttact ttgttaaaaa gaaaaatgct 1020
taa 1023
<210> 3
<211> 674
<212> PRT
<213> 粪肠球菌(Enterococcus faecalis)
<400> 3
Met Thr Lys Ser Val Lys Phe Leu Val Leu Leu Leu Val Met Ile Leu
1 5 10 15
Pro Ile Ala Gly Ala Leu Leu Ile Gly Pro Ile Ser Phe Gly Ala Glu
20 25 30
Leu Ser Lys Ser Ser Ile Val Asp Lys Val Glu Leu Asp His Thr Thr
35 40 45
Leu Tyr Gln Gly Glu Met Thr Ser Ile Lys Val Ser Phe Ser Asp Lys
50 55 60
Glu Asn Gln Lys Ile Lys Pro Gly Asp Thr Ile Thr Leu Thr Leu Pro
65 70 75 80
Asp Ala Leu Val Gly Met Thr Glu Asn Asp Ser Ser Pro Arg Lys Ile
85 90 95
Asn Leu Asn Gly Leu Gly Glu Val Phe Ile Tyr Lys Asp His Val Val
100 105 110
Ala Thr Phe Asn Glu Lys Val Glu Ser Leu His Asn Val Asn Gly His
115 120 125
Phe Ser Phe Gly Ile Lys Thr Leu Ile Thr Asn Ser Ser Gln Pro Asn
130 135 140
Val Ile Glu Thr Asp Phe Gly Thr Ala Thr Ala Thr Gln Arg Leu Thr
145 150 155 160
Ile Glu Gly Val Thr Asn Thr Glu Thr Gly Gln Ile Glu Arg Asp Tyr
165 170 175
Pro Phe Phe Tyr Lys Val Gly Asp Leu Ala Gly Glu Ser Asn Gln Val
180 185 190
Arg Trp Phe Leu Asn Val Asn Leu Asn Lys Ser Asp Val Thr Glu Asp
195 200 205
Ile Ser Ile Ala Asp Arg Gln Gly Ser Gly Gln Gln Leu Asn Lys Glu
210 215 220
Ser Phe Thr Phe Asp Ile Val Asn Asp Lys Glu Thr Lys Tyr Ile Ser
225 230 235 240
Leu Ala Glu Phe Glu Gln Gln Gly Tyr Gly Lys Ile Asp Phe Val Thr
245 250 255
Asp Asn Asp Phe Asn Leu Arg Phe Tyr Arg Asp Lys Ala Arg Phe Thr
260 265 270
Ser Phe Ile Val Arg Tyr Thr Ser Thr Ile Thr Glu Ala Gly Gln His
275 280 285
Gln Ala Thr Phe Glu Asn Ser Tyr Asp Ile Asn Tyr Gln Leu Asn Asn
290 295 300
Gln Asp Ala Thr Asn Glu Lys Asn Thr Ser Gln Val Lys Asn Val Phe
305 310 315 320
Val Glu Gly Glu Ala Ser Gly Asn Gln Asn Val Glu Met Pro Thr Glu
325 330 335
Glu Ser Leu Asp Ile Pro Leu Glu Thr Ile Asp Glu Trp Glu Pro Lys
340 345 350
Thr Pro Thr Ser Glu Gln Ala Thr Glu Thr Ser Glu Lys Thr Asp Thr
355 360 365
Thr Glu Thr Ala Glu Ser Ser Gln Pro Glu Val His Val Ser Pro Thr
370 375 380
Glu Glu Glu Asn Pro Asp Glu Gly Glu Thr Leu Gly Thr Ile Glu Pro
385 390 395 400
Ile Ile Pro Glu Lys Pro Ser Val Thr Thr Glu Glu Asn Gly Thr Thr
405 410 415
Glu Thr Ala Glu Ser Ser Gln Pro Glu Val His Val Ser Pro Thr Glu
420 425 430
Glu Glu Asn Pro Asp Glu Ser Glu Thr Leu Gly Thr Ile Glu Pro Ile
435 440 445
Ile Pro Glu Lys Pro Ser Val Thr Thr Glu Glu Asn Gly Thr Thr Glu
450 455 460
Thr Ala Glu Ser Ser Gln Pro Glu Val His Val Ser Pro Ala Glu Glu
465 470 475 480
Glu Asn Pro Asp Glu Ser Glu Thr Leu Gly Thr Ile Leu Pro Ile Leu
485 490 495
Pro Glu Lys Pro Ser Val Thr Thr Glu Glu Asn Gly Thr Thr Glu Thr
500 505 510
Ala Glu Ser Ser Gln Pro Glu Val His Val Ser Pro Thr Glu Glu Glu
515 520 525
Asn Pro Asp Glu Ser Glu Thr Leu Gly Thr Ile Ala Pro Ile Ile Pro
530 535 540
Glu Lys Pro Ser Val Thr Thr Glu Glu Asn Gly Ile Thr Glu Thr Ala
545 550 555 560
Glu Ser Ser Gln Pro Glu Val His Val Ser Pro Thr Lys Glu Ile Thr
565 570 575
Thr Thr Glu Lys Lys Gln Pro Ser Thr Glu Thr Thr Val Glu Lys Asn
580 585 590
Lys Asn Val Thr Ser Lys Asn Gln Pro Gln Ile Leu Asn Ala Pro Leu
595 600 605
Asn Thr Leu Lys Asn Glu Gly Ser Pro Gln Leu Ala Pro Gln Leu Leu
610 615 620
Ser Glu Pro Ile Gln Lys Leu Asn Glu Ala Asn Gly Gln Arg Glu Leu
625 630 635 640
Pro Lys Thr Gly Thr Thr Lys Thr Pro Phe Met Leu Ile Ala Gly Ile
645 650 655
Leu Ala Ser Thr Phe Ala Val Leu Gly Val Ser Tyr Leu Gln Ile Arg
660 665 670
Lys Asn
<210> 4
<211> 2025
<212> DNA
<213> 粪肠球菌(Enterococcus faecalis)
<400> 4
atgacaaaaa gtgtaaaatt tttagtgtta ctgttggtaa tgattctacc aattgcgggg 60
gcgttattga ttggtccaat ttcgtttggc gccgaattga gcaaaagttc aatcgttgac 120
aaagtagaat tagatcacac tactttatat caaggagaga tgacctcaat taaagtatct 180
tttagtgaca aagaaaatca gaaaataaaa cctggagata ctattacttt aactttacca 240
gacgcactag ttggaatgac cgagaacgat agttcaccac gaaaaatcaa tttaaatggt 300
ttaggggaag tttttatcta taaagatcat gttgtagcaa catttaacga aaaagttgaa 360
tctttacata atgtgaatgg gcatttttct ttcgggatta aaacgcttat caccaatagt 420
tctcaaccga atgtgataga aacggatttc ggaacagcaa cggcgactca acgtttgacg 480
attgaaggag tgactaacac agagactggc caaattgagc gagactatcc gtttttttat 540
aaagtaggcg atttggctgg agagtcaaat caagtacgtt ggtttttaaa tgtgaacctc 600
aataaatccg atgtcacaga agatatttca attgcggatc gacaaggaag tggtcaacaa 660
ttaaataaag agagttttac atttgatatt gtgaatgaca aagaaactaa atatatttca 720
cttgccgagt ttgagcaaca aggttatggc aaaattgact tcgtaacaga taatgacttt 780
aacttacgtt tttatcggga taaagcacgc tttacttcct ttatcgtccg ttacacttcg 840
acaatcacag aagcaggcca acatcaagca acatttgaaa atagttatga catcaattat 900
caactaaaca atcaagacgc aacgaatgaa aaaaatacat cacaggttaa aaatgttttt 960
gtagaaggcg aggcaagcgg caatcaaaat gtggaaatgc caacagaaga aagtctagac 1020
attcctttag agacaataga tgaatgggaa ccaaagacac ctacttcgga acaggcaaca 1080
gaaacaagtg aaaagacaga cacaacagaa accgcagaaa gcagccaacc agaagttcat 1140
gtctcaccaa cagaagaaga aaatccagat gaaggtgaaa cactaggcac gattgagcca 1200
atcatacctg aaaaaccaag tgtgacaact gaagagaatg gcacgacaga aactgcagaa 1260
agcagccaac cagaagttca tgtctcacca acagaagaag aaaatccaga tgaaagtgaa 1320
acactaggca cgattgagcc aatcatacct gaaaaaccaa gtgtgacaac tgaagagaac 1380
ggcacaacag aaaccgcaga aagcagccaa ccagaagttc atgtctcacc agcggaagaa 1440
gaaaatccag atgaaagtga aacgttaggt acaattttac caatcctacc tgaaaaacca 1500
agtgtgacaa ctgaagagaa tggcacaacg gaaactgcag aaagcagtca accagaagtc 1560
catgtgtcgc caacggaaga agaaaatcca gatgaaagtg aaacactagg cacgattgca 1620
ccaatcatac ctgaaaaacc aagcgtaaca actgaagaga atggtataac ggaaacggca 1680
gaaagcagcc agccagaagt tcatgtctca ccaacaaaag aaattactac aactgagaaa 1740
aaacagccat ccacagaaac aactgtggag aaaaataaaa atgttacatc aaaaaatcaa 1800
ccacaaatac taaacgctcc attaaataca ttgaaaaatg aaggaagccc acagttggct 1860
ccccaactgc ttagtgaacc aattcaaaaa ttaaatgaag caaacgggca acgagaactt 1920
cccaaaacag gcacaacaaa aacaccgttt atgctaatag caggaatact ggcaagtaca 1980
tttgccgttt taggtgtaag ttatctacaa atcagaaaga attaa 2025
<210> 5
<211> 303
<212> PRT
<213> 金黄色葡萄球菌(Staphylococcus aureus)
<400> 5
Gly Ser Ala Arg Asp Ile Ser Ser Thr Asn Val Thr Asp Leu Thr Val
1 5 10 15
Ser Pro Ser Lys Ile Glu Asp Gly Gly Lys Thr Thr Val Lys Met Thr
20 25 30
Phe Asp Asp Lys Asn Gly Lys Ile Gln Asn Gly Asp Met Ile Lys Val
35 40 45
Ala Trp Pro Thr Ser Gly Thr Val Lys Ile Glu Gly Tyr Ser Lys Thr
50 55 60
Val Pro Leu Thr Val Lys Gly Glu Gln Val Gly Gln Ala Val Ile Thr
65 70 75 80
Pro Asp Gly Ala Thr Ile Thr Phe Asn Asp Lys Val Glu Lys Leu Ser
85 90 95
Asp Val Ser Gly Phe Ala Glu Phe Glu Val Gln Gly Arg Asn Leu Thr
100 105 110
Gln Thr Asn Thr Ser Asp Asp Lys Val Ala Thr Ile Thr Ser Gly Asn
115 120 125
Lys Ser Thr Asn Val Thr Val His Lys Ser Glu Ala Gly Thr Ser Ser
130 135 140
Val Phe Tyr Tyr Lys Thr Gly Asp Met Leu Pro Glu Asp Thr Thr His
145 150 155 160
Val Arg Trp Phe Leu Asn Ile Asn Asn Glu Lys Ser Tyr Val Ser Lys
165 170 175
Asp Ile Thr Ile Lys Asp Gln Ile Gln Gly Gly Gln Gln Leu Asp Leu
180 185 190
Ser Thr Leu Asn Ile Asn Val Thr Gly Thr His Ser Asn Tyr Tyr Ser
195 200 205
Gly Gln Ser Ala Ile Thr Asp Phe Glu Lys Ala Phe Pro Gly Ser Lys
210 215 220
Ile Thr Val Asp Asn Thr Lys Asn Thr Ile Asp Val Thr Ile Pro Gln
225 230 235 240
Gly Tyr Gly Ser Tyr Asn Ser Phe Ser Ile Asn Tyr Lys Thr Lys Ile
245 250 255
Thr Asn Glu Gln Gln Lys Glu Phe Val Asn Asn Ser Gln Ala Trp Tyr
260 265 270
Gln Glu His Gly Lys Glu Glu Val Asn Gly Lys Ser Phe Asn His Thr
275 280 285
Val His Asn Ile Asn Ala Asn Ala Gly Ile Glu Gly Thr Val Lys
290 295 300
<210> 6
<211> 102
<212> PRT
<213> 化脓性链球菌(Streptococcus pyogenes)
<400> 6
Met Thr Ile Glu Glu Asp Ser Ala Thr His Ile Lys Phe Ser Lys Arg
1 5 10 15
Asp Ile Asp Gly Lys Glu Leu Ala Gly Ala Thr Met Glu Leu Arg Asp
20 25 30
Ser Ser Gly Lys Thr Ile Ser Thr Trp Ile Ser Asp Gly Gln Val Lys
35 40 45
Asp Phe Tyr Leu Met Pro Gly Lys Tyr Thr Phe Val Glu Thr Ala Ala
50 55 60
Pro Asp Gly Tyr Glu Val Ala Thr Ala Ile Thr Phe Thr Val Asn Glu
65 70 75 80
Gln Gly Gln Val Thr Val Asn Gly Lys Ala Thr Lys Gly Asp Ala His
85 90 95
Ile Val Met Val Asp Ala
100
<210> 7
<211> 340
<212> PRT
<213> 化脓性链球菌(Streptococcus pyogenes)
<400> 7
Met Lys Leu Arg His Leu Leu Leu Thr Gly Ala Ala Leu Thr Ser Phe
1 5 10 15
Ala Ala Thr Thr Val His Gly Glu Thr Val Val Asn Gly Ala Lys Leu
20 25 30
Thr Val Thr Lys Asn Leu Asp Leu Val Asn Ser Asn Ala Leu Ile Pro
35 40 45
Asn Thr Asp Phe Thr Phe Lys Ile Glu Pro Asp Thr Thr Val Asn Glu
50 55 60
Asp Gly Asn Lys Phe Lys Gly Val Ala Leu Asn Thr Pro Met Thr Lys
65 70 75 80
Val Thr Tyr Thr Asn Ser Asp Lys Gly Gly Ser Asn Thr Lys Thr Ala
85 90 95
Glu Phe Asp Phe Ser Glu Val Thr Phe Glu Lys Pro Gly Val Tyr Tyr
100 105 110
Tyr Lys Val Thr Glu Glu Lys Ile Asp Lys Val Pro Gly Val Ser Tyr
115 120 125
Asp Thr Thr Ser Tyr Thr Val Gln Val His Val Leu Trp Asn Glu Glu
130 135 140
Gln Gln Lys Pro Val Ala Thr Tyr Ile Val Gly Tyr Lys Glu Gly Ser
145 150 155 160
Lys Val Pro Ile Gln Phe Lys Asn Ser Leu Asp Ser Thr Thr Leu Thr
165 170 175
Val Lys Lys Lys Val Ser Gly Thr Gly Gly Asp Arg Ser Lys Asp Phe
180 185 190
Asn Phe Gly Leu Thr Leu Lys Ala Asn Gln Tyr Tyr Lys Ala Ser Glu
195 200 205
Lys Val Met Ile Glu Lys Thr Thr Lys Gly Gly Gln Ala Pro Val Gln
210 215 220
Thr Glu Ala Ser Ile Asp Gln Leu Tyr His Phe Thr Leu Lys Asp Gly
225 230 235 240
Glu Ser Ile Lys Val Thr Asn Leu Pro Val Gly Val Asp Tyr Val Val
245 250 255
Thr Glu Asp Asp Tyr Lys Ser Glu Lys Tyr Thr Thr Asn Val Glu Val
260 265 270
Ser Pro Gln Asp Gly Ala Val Lys Asn Ile Ala Gly Asn Ser Thr Glu
275 280 285
Gln Glu Thr Ser Thr Asp Lys Asp Met Thr Ile Thr Phe Thr Asn Lys
290 295 300
Lys Asp Phe Glu Val Pro Thr Gly Val Ala Met Thr Val Ala Pro Tyr
305 310 315 320
Ile Ala Leu Gly Ile Val Ala Val Gly Gly Ala Leu Tyr Phe Val Lys
325 330 335
Lys Lys Asn Ala
340
<210> 8
<211> 1023
<212> DNA
<213> 化脓性链球菌(Streptococcus pyogenes)
<400> 8
atgaaattac gtcacttact attaacggga gcagccctaa ctagttttgc tgctacaaca 60
gttcacgggg agactgttgt aaacggagcc aaactaacag ttacaaaaaa ccttgattta 120
gttaatagca atgcattaat tccaaataca gattttacat ttaaaatcga acctgatact 180
actgtcaacg aagacggaaa taagtttaaa ggtgtagctt tgaacacacc gatgactaaa 240
gtcacttaca ccaattcaga taaaggtgga tcaaatacga aaactgcaga atttgatttt 300
tcagaagtta cttttgaaaa accaggtgtt tattattaca aagtaactga ggagaagata 360
gataaagttc ctggtgtttc ttatgataca acatcttaca ctgttcaagt tcatgtcttg 420
tggaatgaag agcaacaaaa accagtagct acttatattg ttggttataa agaaggtagt 480
aaggtgccaa ttcagttcaa aaatagctta gattctacta cattaacggt gaagaaaaaa 540
gtttcaggta ccggtggaga tcgctctaaa gattttaatt ttggtctgac tttaaaagca 600
aatcagtatt ataaggcgtc agaaaaagtc atgattgaga agacaactaa aggtggtcaa 660
gctcctgttc aaacagaggc tagtatagat caactctatc attttacctt gaaagatggt 720
gaatcaatca aagtcacaaa tcttccagta ggtgtggatt atgttgtcac tgaagacgat 780
tacaaatcag aaaaatatac aaccaacgtg gaagttagtc ctcaagatgg agctgtaaaa 840
aatatcgcag gtaattcaac tgaacaagag acatctactg ataaagatat gaccattact 900
tttacaaata aaaaagactt tgaagtgcca acaggagtag caatgactgt ggcaccatat 960
attgctttag gaattgtagc agttggtgga gctctttact ttgttaaaaa gaaaaatgct 1020
taa 1023
<210> 9
<211> 674
<212> PRT
<213> 粪肠球菌(Enterococcus faecalis)
<400> 9
Met Thr Lys Ser Val Lys Phe Leu Val Leu Leu Leu Val Met Ile Leu
1 5 10 15
Pro Ile Ala Gly Ala Leu Leu Ile Gly Pro Ile Ser Phe Gly Ala Glu
20 25 30
Leu Ser Lys Ser Ser Ile Val Asp Lys Val Glu Leu Asp His Thr Thr
35 40 45
Leu Tyr Gln Gly Glu Met Thr Ser Ile Lys Val Ser Phe Ser Asp Lys
50 55 60
Glu Asn Gln Lys Ile Lys Pro Gly Asp Thr Ile Thr Leu Thr Leu Pro
65 70 75 80
Asp Ala Leu Val Gly Met Thr Glu Asn Asp Ser Ser Pro Arg Lys Ile
85 90 95
Asn Leu Asn Gly Leu Gly Glu Val Phe Ile Tyr Lys Asp His Val Val
100 105 110
Ala Thr Phe Asn Glu Lys Val Glu Ser Leu His Asn Val Asn Gly His
115 120 125
Phe Ser Phe Gly Ile Lys Thr Leu Ile Thr Asn Ser Ser Gln Pro Asn
130 135 140
Val Ile Glu Thr Asp Phe Gly Thr Ala Thr Ala Thr Gln Arg Leu Thr
145 150 155 160
Ile Glu Gly Val Thr Asn Thr Glu Thr Gly Gln Ile Glu Arg Asp Tyr
165 170 175
Pro Phe Phe Tyr Lys Val Gly Asp Leu Ala Gly Glu Ser Asn Gln Val
180 185 190
Arg Trp Phe Leu Asn Val Asn Leu Asn Lys Ser Asp Val Thr Glu Asp
195 200 205
Ile Ser Ile Ala Asp Arg Gln Gly Ser Gly Gln Gln Leu Asn Lys Glu
210 215 220
Ser Phe Thr Phe Asp Ile Val Asn Asp Lys Glu Thr Lys Tyr Ile Ser
225 230 235 240
Leu Ala Glu Phe Glu Gln Gln Gly Tyr Gly Lys Ile Asp Phe Val Thr
245 250 255
Asp Asn Asp Phe Asn Leu Arg Phe Tyr Arg Asp Lys Ala Arg Phe Thr
260 265 270
Ser Phe Ile Val Arg Tyr Thr Ser Thr Ile Thr Glu Ala Gly Gln His
275 280 285
Gln Ala Thr Phe Glu Asn Ser Tyr Asp Ile Asn Tyr Gln Leu Asn Asn
290 295 300
Gln Asp Ala Thr Asn Glu Lys Asn Thr Ser Gln Val Lys Asn Val Phe
305 310 315 320
Val Glu Gly Glu Ala Ser Gly Asn Gln Asn Val Glu Met Pro Thr Glu
325 330 335
Glu Ser Leu Asp Ile Pro Leu Glu Thr Ile Asp Glu Trp Glu Pro Lys
340 345 350
Thr Pro Thr Ser Glu Gln Ala Thr Glu Thr Ser Glu Lys Thr Asp Thr
355 360 365
Thr Glu Thr Ala Glu Ser Ser Gln Pro Glu Val His Val Ser Pro Thr
370 375 380
Glu Glu Glu Asn Pro Asp Glu Gly Glu Thr Leu Gly Thr Ile Glu Pro
385 390 395 400
Ile Ile Pro Glu Lys Pro Ser Val Thr Thr Glu Glu Asn Gly Thr Thr
405 410 415
Glu Thr Ala Glu Ser Ser Gln Pro Glu Val His Val Ser Pro Thr Glu
420 425 430
Glu Glu Asn Pro Asp Glu Ser Glu Thr Leu Gly Thr Ile Glu Pro Ile
435 440 445
Ile Pro Glu Lys Pro Ser Val Thr Thr Glu Glu Asn Gly Thr Thr Glu
450 455 460
Thr Ala Glu Ser Ser Gln Pro Glu Val His Val Ser Pro Ala Glu Glu
465 470 475 480
Glu Asn Pro Asp Glu Ser Glu Thr Leu Gly Thr Ile Leu Pro Ile Leu
485 490 495
Pro Glu Lys Pro Ser Val Thr Thr Glu Glu Asn Gly Thr Thr Glu Thr
500 505 510
Ala Glu Ser Ser Gln Pro Glu Val His Val Ser Pro Thr Glu Glu Glu
515 520 525
Asn Pro Asp Glu Ser Glu Thr Leu Gly Thr Ile Ala Pro Ile Ile Pro
530 535 540
Glu Lys Pro Ser Val Thr Thr Glu Glu Asn Gly Ile Thr Glu Thr Ala
545 550 555 560
Glu Ser Ser Gln Pro Glu Val His Val Ser Pro Thr Lys Glu Ile Thr
565 570 575
Thr Thr Glu Lys Lys Gln Pro Ser Thr Glu Thr Thr Val Glu Lys Asn
580 585 590
Lys Asn Val Thr Ser Lys Asn Gln Pro Gln Ile Leu Asn Ala Pro Leu
595 600 605
Asn Thr Leu Lys Asn Glu Gly Ser Pro Gln Leu Ala Pro Gln Leu Leu
610 615 620
Ser Glu Pro Ile Gln Lys Leu Asn Glu Ala Asn Gly Gln Arg Glu Leu
625 630 635 640
Pro Lys Thr Gly Thr Thr Lys Thr Pro Phe Met Leu Ile Ala Gly Ile
645 650 655
Leu Ala Ser Thr Phe Ala Val Leu Gly Val Ser Tyr Leu Gln Ile Arg
660 665 670
Lys Asn
<210> 10
<211> 331
<212> PRT
<213> 金黄色葡萄球菌(Staphylococcus aureus)
<400> 10
Met Asn Lys Asn Val Leu Lys Phe Met Val Phe Ile Met Leu Leu Asn
1 5 10 15
Ile Ile Thr Pro Leu Phe Asn Lys Asn Glu Ala Phe Ala Ala Arg Asp
20 25 30
Ile Ser Ser Thr Asn Val Thr Asp Leu Thr Val Ser Pro Ser Lys Ile
35 40 45
Glu Asp Gly Gly Lys Thr Thr Val Lys Met Thr Phe Asp Asp Lys Asn
50 55 60
Gly Lys Ile Gln Asn Gly Asp Met Ile Lys Val Ala Trp Pro Thr Ser
65 70 75 80
Gly Thr Val Lys Ile Glu Gly Tyr Ser Lys Thr Val Pro Leu Thr Val
85 90 95
Lys Gly Glu Gln Val Gly Gln Ala Val Ile Thr Pro Asp Gly Ala Thr
100 105 110
Ile Thr Phe Asn Asp Lys Val Glu Lys Leu Ser Asp Val Ser Gly Phe
115 120 125
Ala Glu Phe Glu Val Gln Gly Arg Asn Leu Thr Gln Thr Asn Thr Ser
130 135 140
Asp Asp Lys Val Ala Thr Ile Thr Ser Gly Asn Lys Ser Thr Asn Val
145 150 155 160
Thr Val His Lys Ser Glu Ala Gly Thr Ser Ser Val Phe Tyr Tyr Lys
165 170 175
Thr Gly Asp Met Leu Pro Glu Asp Thr Thr His Val Arg Trp Phe Leu
180 185 190
Asn Ile Asn Asn Glu Lys Ser Tyr Val Ser Lys Asp Ile Thr Ile Lys
195 200 205
Asp Gln Ile Gln Gly Gly Gln Gln Leu Asp Leu Ser Thr Leu Asn Ile
210 215 220
Asn Val Thr Gly Thr His Ser Asn Tyr Tyr Ser Gly Gln Ser Ala Ile
225 230 235 240
Thr Asp Phe Glu Lys Ala Phe Pro Gly Ser Lys Ile Thr Val Asp Asn
245 250 255
Thr Lys Asn Thr Ile Asp Val Thr Ile Pro Gln Gly Tyr Gly Ser Tyr
260 265 270
Asn Ser Phe Ser Ile Asn Tyr Lys Thr Lys Ile Thr Asn Glu Gln Gln
275 280 285
Lys Glu Phe Val Asn Asn Ser Gln Ala Trp Tyr Gln Glu His Gly Lys
290 295 300
Glu Glu Val Asn Gly Lys Ser Phe Asn His Thr Val His Asn Ile Asn
305 310 315 320
Ala Asn Ala Gly Ile Glu Gly Thr Val Lys Gly
325 330
<210> 11
<211> 102
<212> PRT
<213> 化脓性链球菌(Streptococcus pyogenes)
<400> 11
Met Thr Ile Glu Glu Asp Ser Ala Thr His Ile Lys Phe Ser Lys Arg
1 5 10 15
Asp Ile Asp Gly Lys Glu Leu Ala Gly Ala Thr Met Glu Leu Arg Asp
20 25 30
Ser Ser Gly Lys Thr Ile Ser Thr Trp Ile Ser Asp Gly Gln Val Lys
35 40 45
Asp Phe Tyr Leu Met Pro Gly Lys Tyr Thr Phe Val Glu Thr Ala Ala
50 55 60
Pro Asp Gly Tyr Glu Val Ala Thr Ala Ile Thr Phe Thr Val Asn Glu
65 70 75 80
Gln Gly Gln Val Thr Val Asn Gly Lys Ala Thr Lys Gly Asp Ala His
85 90 95
Ile Val Met Val Asp Ala
100
<210> 12
<211> 309
<212> DNA
<213> 化脓性链球菌(Streptococcus pyogenes)
<400> 12
atgacaattg aagaagatag tgctacccat attaaattct caaaacgtga tattgacggc 60
aaagagttag ctggtgcaac tatggagttg cgtgattcat ctggtaaaac tattagtaca 120
tggatttcag atggacaagt gaaagatttc tacctgatgc caggaaaata tacatttgtc 180
gaaaccgcag caccagacgg ttatgaggta gcaactgcta ttacctttac agttaatgag 240
caaggtcagg ttactgtaaa tggcaaagca actaaaggtg acgctcatat tgtcatggtt 300
gatgcttga 309
<210> 13
<211> 893
<212> PRT
<213> 肺炎链球菌(Streptococcus pneumoniae)
<400> 13
Met Leu Asn Arg Glu Thr His Met Lys Lys Val Arg Lys Ile Phe Gln
1 5 10 15
Lys Ala Val Ala Gly Leu Cys Cys Ile Ser Gln Leu Thr Ala Phe Ser
20 25 30
Ser Ile Val Ala Leu Ala Glu Thr Pro Glu Thr Ser Pro Ala Ile Gly
35 40 45
Lys Val Val Ile Lys Glu Thr Gly Glu Gly Gly Ala Leu Leu Gly Asp
50 55 60
Ala Val Phe Glu Leu Lys Asn Asn Thr Asp Gly Thr Thr Val Ser Gln
65 70 75 80
Arg Thr Glu Ala Gln Thr Gly Glu Ala Ile Phe Ser Asn Ile Lys Pro
85 90 95
Gly Thr Tyr Thr Leu Thr Glu Ala Gln Pro Pro Val Gly Tyr Lys Pro
100 105 110
Ser Thr Lys Gln Trp Thr Val Glu Val Glu Lys Asn Gly Arg Thr Thr
115 120 125
Val Gln Gly Glu Gln Val Glu Asn Arg Glu Glu Ala Leu Ser Asp Gln
130 135 140
Tyr Pro Gln Thr Gly Thr Tyr Pro Asp Val Gln Thr Pro Tyr Gln Ile
145 150 155 160
Ile Lys Val Asp Gly Ser Glu Lys Asn Gly Gln His Lys Ala Leu Asn
165 170 175
Pro Asn Pro Tyr Glu Arg Val Ile Pro Glu Gly Thr Leu Ser Lys Arg
180 185 190
Ile Tyr Gln Val Asn Asn Leu Asp Asp Asn Gln Tyr Gly Ile Glu Leu
195 200 205
Thr Val Ser Gly Lys Thr Val Tyr Glu Gln Lys Asp Lys Ser Val Pro
210 215 220
Leu Asp Val Val Ile Leu Leu Asp Asn Ser Asn Ser Met Ser Asn Ile
225 230 235 240
Arg Asn Lys Asn Ala Arg Arg Ala Glu Arg Ala Gly Glu Ala Thr Arg
245 250 255
Ser Leu Ile Asp Lys Ile Thr Ser Asp Ser Glu Asn Arg Val Ala Leu
260 265 270
Val Thr Tyr Ala Ser Thr Ile Phe Asp Gly Thr Glu Phe Thr Val Glu
275 280 285
Lys Gly Val Ala Asp Lys Asn Gly Lys Arg Leu Asn Asp Ser Leu Phe
290 295 300
Trp Asn Tyr Asp Gln Thr Ser Phe Thr Thr Asn Thr Lys Asp Tyr Ser
305 310 315 320
Tyr Leu Lys Leu Thr Asn Asp Lys Asn Asp Ile Val Glu Leu Lys Asn
325 330 335
Lys Val Pro Thr Glu Ala Glu Asp His Asp Gly Asn Arg Leu Met Tyr
340 345 350
Gln Phe Gly Ala Thr Phe Thr Gln Lys Ala Leu Met Lys Ala Asp Glu
355 360 365
Ile Leu Thr Gln Gln Ala Arg Gln Asn Ser Gln Lys Val Ile Phe His
370 375 380
Ile Thr Asp Gly Val Pro Thr Met Ser Tyr Pro Ile Asn Phe Asn His
385 390 395 400
Ala Thr Phe Ala Pro Ser Tyr Gln Asn Gln Leu Asn Ala Phe Phe Ser
405 410 415
Lys Ser Pro Asn Lys Asp Gly Ile Leu Leu Ser Asp Phe Ile Thr Gln
420 425 430
Ala Thr Ser Gly Glu His Thr Ile Val Arg Gly Asp Gly Gln Ser Tyr
435 440 445
Gln Met Phe Thr Asp Lys Thr Val Tyr Glu Lys Gly Ala Pro Ala Ala
450 455 460
Phe Pro Val Lys Pro Glu Lys Tyr Ser Glu Met Lys Ala Ala Gly Tyr
465 470 475 480
Ala Val Ile Gly Asp Pro Ile Asn Gly Gly Tyr Ile Trp Leu Asn Trp
485 490 495
Arg Glu Ser Ile Leu Ala Tyr Pro Phe Asn Ser Asn Thr Ala Lys Ile
500 505 510
Thr Asn His Gly Asp Pro Thr Arg Trp Tyr Tyr Asn Gly Asn Ile Ala
515 520 525
Pro Asp Gly Tyr Asp Val Phe Thr Val Gly Ile Gly Ile Asn Gly Asp
530 535 540
Pro Gly Thr Asp Glu Ala Thr Ala Thr Ser Phe Met Gln Ser Ile Ser
545 550 555 560
Ser Lys Pro Glu Asn Tyr Thr Asn Val Thr Asp Thr Thr Lys Ile Leu
565 570 575
Glu Gln Leu Asn Arg Tyr Phe His Thr Ile Val Thr Glu Lys Lys Ser
580 585 590
Ile Glu Asn Gly Thr Ile Thr Asp Pro Met Gly Glu Leu Ile Asp Leu
595 600 605
Gln Leu Gly Thr Asp Gly Arg Phe Asp Pro Ala Asp Tyr Thr Leu Thr
610 615 620
Ala Asn Asp Gly Ser Arg Leu Glu Asn Gly Gln Ala Val Gly Gly Pro
625 630 635 640
Gln Asn Asp Gly Gly Leu Leu Lys Asn Ala Lys Val Leu Tyr Asp Thr
645 650 655
Thr Glu Lys Arg Ile Arg Val Thr Gly Leu Tyr Leu Gly Thr Asp Glu
660 665 670
Lys Val Thr Leu Thr Tyr Asn Val Arg Leu Asn Asp Glu Phe Val Ser
675 680 685
Asn Lys Phe Tyr Asp Thr Asn Gly Arg Thr Thr Leu His Pro Lys Glu
690 695 700
Val Glu Gln Asn Thr Val Arg Asp Phe Pro Ile Pro Lys Ile Arg Asp
705 710 715 720
Val Arg Lys Tyr Pro Glu Ile Thr Ile Ser Lys Glu Lys Lys Leu Gly
725 730 735
Asp Ile Glu Phe Ile Lys Val Asn Lys Asn Asp Lys Lys Pro Leu Arg
740 745 750
Gly Ala Val Phe Ser Leu Gln Lys Gln His Pro Asp Tyr Pro Asp Ile
755 760 765
Tyr Gly Ala Ile Asp Gln Asn Gly Thr Tyr Gln Asn Val Arg Thr Gly
770 775 780
Glu Asp Gly Lys Leu Thr Phe Lys Asn Leu Ser Asp Gly Lys Tyr Arg
785 790 795 800
Leu Phe Glu Asn Ser Glu Pro Ala Gly Tyr Lys Pro Val Gln Asn Lys
805 810 815
Pro Ile Val Ala Phe Gln Ile Val Asn Gly Glu Val Arg Asp Val Thr
820 825 830
Ser Ile Val Pro Gln Asp Ile Pro Ala Gly Tyr Glu Phe Thr Asn Asp
835 840 845
Lys His Tyr Ile Thr Asn Glu Pro Ile Pro Pro Lys Arg Glu Tyr Pro
850 855 860
Arg Thr Gly Gly Ile Gly Met Leu Pro Phe Tyr Leu Ile Gly Cys Met
865 870 875 880
Met Met Gly Gly Val Leu Leu Tyr Thr Arg Lys His Pro
885 890
<210> 14
<211> 2682
<212> DNA
<213> 肺炎链球菌(Streptococcus pneumoniae)
<400> 14
atgctgaacc gcgaaaccca tatgaaaaaa gtaagaaaga tatttcagaa ggcagttgca 60
ggactgtgct gtatatctca gttgacagct ttttcttcga tagttgcttt agcagaaacg 120
cctgaaacca gtccagcgat aggaaaagta gtgattaagg agacaggcga aggaggagcg 180
cttctaggag atgccgtctt tgagttgaaa aacaatacgg atggcacaac tgtttcgcaa 240
aggacagagg cgcaaacagg agaagcgata ttttcaaaca taaaacctgg gacatacacc 300
ttgacagaag cccaacctcc agttggttat aaaccctcta ctaaacaatg gactgttgaa 360
gttgagaaga atggtcggac gactgtccaa ggtgaacagg tagaaaatcg agaagaggct 420
ctatctgacc agtatccaca aacagggact tatccagatg ttcaaacacc ttatcagatt 480
attaaggtag atggttcgga aaaaaacgga cagcacaagg cgttgaatcc gaatccatat 540
gaacgtgtga ttccagaagg tacactttca aagagaattt atcaagtgaa taatttggat 600
gataaccaat atggaatcga attgacggtt agtgggaaaa cagtgtatga acaaaaagat 660
aagtctgtgc cgctggatgt cgttatcttg ctcgataact caaatagtat gagtaacatt 720
cgaaacaaga atgctcgacg tgcggaaaga gctggtgagg cgacacgttc tcttattgat 780
aaaattacat ctgattcaga aaatagggta gcgcttgtga cttatgcttc cactatcttt 840
gatgggaccg agtttacagt agaaaaaggg gtagcagata aaaacggaaa gcgattgaat 900
gattctcttt tttggaatta tgatcagacg agttttacaa ccaataccaa agattatagt 960
tatttaaagc tgactaatga taagaatgac attgtagaat taaaaaataa ggtacctacc 1020
gaggcagaag accatgatgg aaatagattg atgtaccaat tcggtgccac ttttactcag 1080
aaagctttga tgaaggcaga tgagattttg acacaacaag cgagacaaaa tagtcaaaaa 1140
gtcattttcc atattacgga tggtgtccca actatgtcgt atccgattaa ttttaatcat 1200
gctacgtttg ctccatcata tcaaaatcaa ctaaatgcat tttttagtaa atctcctaat 1260
aaagatggaa tactattaag tgattttatt acgcaagcaa ctagtggaga acatacaatt 1320
gtacgcggag atgggcaaag ttaccagatg tttacagata agacagttta tgaaaaaggt 1380
gctcctgcag ctttcccagt taaacctgaa aaatattctg aaatgaaggc ggctggttat 1440
gcagttatag gcgatccaat taatggtgga tatatttggc ttaattggag agagagtatt 1500
ctggcttatc cgtttaattc taatactgct aaaattacca atcatggtga ccctacaaga 1560
tggtactata acgggaatat tgctcctgat gggtatgatg tctttacggt aggtattggt 1620
attaacggag atcctggtac ggatgaagca acggctacta gttttatgca aagtatttct 1680
agtaaacctg aaaactatac caatgttact gacacgacaa aaatattgga acagttgaat 1740
cgttatttcc acaccatcgt aactgaaaag aaatcaattg agaatggtac gattacagat 1800
ccgatgggtg agttaattga tttgcaattg ggcacagatg gaagatttga tccagcagat 1860
tacactttaa ctgcaaacga tggtagtcgc ttggagaatg gacaagctgt aggtggtcca 1920
caaaatgatg gtggtttgtt aaaaaatgca aaagtgctct atgatacgac tgagaaaagg 1980
attcgtgtaa caggtctgta ccttggaacg gatgaaaaag ttacgttgac ctacaatgtt 2040
cgtttgaatg atgagtttgt aagcaataaa ttttatgata ccaatggtcg aacaacctta 2100
catcctaagg aagtagaaca gaacacagtg cgcgacttcc cgattcctaa gattcgtgat 2160
gtgcggaagt atccagaaat cacaatttca aaagagaaaa aacttggtga cattgagttt 2220
attaaggtca ataaaaatga taaaaaacca ctgagaggtg cggtctttag tcttcaaaaa 2280
caacatccgg attatccaga tatttatgga gctattgatc aaaatggcac ttatcaaaat 2340
gtgagaacag gtgaagatgg taagttgacc tttaaaaatc tgtcagatgg gaaatatcga 2400
ttatttgaaa attctgaacc agctggttat aaacccgttc aaaataagcc tatcgttgcc 2460
ttccaaatag taaatggaga agtcagagat gtgacttcaa tcgttccaca agatatacca 2520
gcgggttacg agtttacgaa tgataagcac tatattacca atgaacctat tcctccaaag 2580
agagaatatc ctcgaactgg tggtatcgga atgttgccat tctatctgat aggttgcatg 2640
atgatgggag gagttctatt atacacacgg aaacatccgt aa 2682
<210> 15
<211> 738
<212> PRT
<213> 中间链球菌(Streptococcus intermedius)
<400> 15
Met Lys Lys Arg Arg Gly Gln Phe Phe Lys Ser Ala Ile Ser Phe Leu
1 5 10 15
Val Val Phe Leu Met Val Met Val Ser Ile Ile Tyr Pro Ser Ser Lys
20 25 30
Ile Lys Ala Asp Gly Phe Pro Asn Asp Ala Thr Gly Val Ser Pro Asn
35 40 45
Gly Lys Tyr Tyr Ser Ala Gly Arg Glu Asn Arg Leu Gly Met Val Thr
50 55 60
Ser Asp Glu Leu His Thr Ala Thr Glu Leu Phe Gly Phe Cys Met Ala
65 70 75 80
Asn Ser Lys Lys Tyr Pro Gly Tyr Asp Ser Lys Lys Asp Glu Tyr Phe
85 90 95
Gly Val Tyr Glu Gln Ile Leu Asn Leu Asn Lys Glu Ser Phe Asn Lys
100 105 110
Leu Val Arg Asp Asn His Thr Tyr Gly Asn Ile Pro Thr Ser Pro Glu
115 120 125
Glu Leu Trp Asp Lys Val Ser Lys Leu Ile Tyr Ile Tyr Leu Lys Asp
130 135 140
Pro Thr Asn Val Ile Gly Gln Ala Gly Trp Thr Asn Pro Gln Asp Ala
145 150 155 160
Met Tyr Glu Phe Tyr Thr Val Val Gln Gln Glu Ile Trp Arg Tyr Thr
165 170 175
Asp Gly Gln Lys Val Asp Lys Asp Thr Asn Ser Tyr Leu Tyr Tyr Lys
180 185 190
Tyr Ser Lys Gln Gly Gln Lys Ala Val Tyr Leu Leu Arg Asp Ala Val
195 200 205
Asn Ser Ile Ser Ile Pro Ser Asn Phe Glu Leu Arg Gly Tyr Lys Pro
210 215 220
Glu Trp Val Gln Gly Gln Lys Gly Tyr Gln Ala Ile Val Thr Gly Arg
225 230 235 240
Leu Lys Val Asp Gln Pro Val Gly Glu Ile Lys Thr Thr Val Thr Ala
245 250 255
Gly Gly Lys Thr Ser Ser Glu Asn Asp Ile Ala Thr Leu Lys Ala Gln
260 265 270
Asp Val Ile Gly Gly Val Glu Val Ser Asp Lys Ile Thr Tyr Ser Gly
275 280 285
Leu Tyr Pro Asn Thr Glu Tyr Asp Val Ile Gly Glu Ile Tyr Glu Val
290 295 300
Lys Asp Gly Glu Leu Val Asn Pro Gly Arg Pro Val Ser Val Val Asn
305 310 315 320
Ser Gly Asp Asp Leu Lys Thr Asp Ala Thr Gly Lys Gly Lys Trp Thr
325 330 335
Leu Asn Phe Gly Lys Leu Asp Leu Glu Ala Gly Lys Ser Tyr Val Val
340 345 350
Phe Glu Lys Val Val Ser Leu Lys Asn Val Ile Asp Thr Asp Gly Asp
355 360 365
Gly Lys Pro Asp Lys Lys Gln Glu Leu Ser His Asn Asp Pro Lys Asp
370 375 380
Lys Ser Gln Thr Phe Thr Ile Leu Pro Lys Glu Ile Val Glu Gln Asp
385 390 395 400
Val Val Phe Ser Lys Val Asn Val Ala Gly Glu Glu Ile Ala Gly Ala
405 410 415
Lys Ile Gln Leu Lys Asp Ala Gln Gly Gln Val Val His Ser Trp Thr
420 425 430
Ser Lys Ala Gly Gln Ser Glu Thr Val Lys Leu Lys Ala Gly Thr Tyr
435 440 445
Thr Phe His Glu Ala Ser Ala Pro Thr Gly Tyr Leu Ala Val Thr Asp
450 455 460
Ile Thr Phe Glu Val Asp Val Gln Gly Lys Val Thr Val Lys Asp Ala
465 470 475 480
Asn Gly Asn Gly Val Lys Ala Asp Gly Asn Lys Leu Thr Val Thr Asp
485 490 495
Gln Ala Ala Pro Ser Val Pro Asn Glu Gln Asp Val Val Phe Ser Lys
500 505 510
Val Asn Val Ala Gly Glu Glu Ile Ala Gly Ala Lys Ile Gln Leu Lys
515 520 525
Asp Ala Gln Gly Gln Val Val His Ser Trp Thr Ser Lys Ala Gly Gln
530 535 540
Ser Glu Thr Val Lys Leu Lys Ala Gly Thr Tyr Thr Phe His Glu Ala
545 550 555 560
Ser Ala Pro Thr Gly Tyr Leu Ala Val Thr Asp Ile Thr Phe Glu Val
565 570 575
Asp Val Gln Gly Lys Val Thr Val Lys Asp Ala Asn Gly Asn Gly Val
580 585 590
Lys Ala Asp Gly Asn Lys Leu Thr Val Thr Asp Gln Ala Ala Pro Ser
595 600 605
Val Pro Asn Glu Gln Asp Val Val Phe Ser Lys Val Asn Val Ala Gly
610 615 620
Glu Glu Ile Ala Gly Ala Lys Ile Gln Leu Lys Asp Ala Gln Gly Gln
625 630 635 640
Val Val His Ser Trp Thr Ser Lys Ala Gly Gln Ser Glu Thr Val Lys
645 650 655
Leu Lys Ala Gly Thr Tyr Thr Phe His Glu Ala Ser Ala Pro Thr Gly
660 665 670
Tyr Leu Ala Val Thr Asp Ile Thr Phe Glu Val Asp Val Gln Gly Lys
675 680 685
Val Thr Val Lys Asp Ala Asn Gly Asn Gly Val Lys Ala Asp Gly Asn
690 695 700
Lys Leu Thr Val Thr Asp Gln Ala Ala Pro Ser Val Pro Asn Glu Gln
705 710 715 720
Asp Val Val Phe Ser Lys Val Asn Val Ala Gly Glu Glu Ile Ala Gly
725 730 735
Ala Lys
<210> 16
<211> 2215
<212> DNA
<213> 中间链球菌(Streptococcus intermedius)
<400> 16
atgaaaaaga gaagaggaca atttttcaaa agtgcaattt cgtttttggt tgtatttttg 60
atggtaatgg taagtatcat ttacccatct tcaaaaatta aagcagatgg atttcctaat 120
gatgctacgg gagtatcgcc aaatggtaaa tattactcgg cagggagaga aaaccgttta 180
ggaatggtta catcagatga attgcataca gctacagaat tattcggttt ttgtatggca 240
aatagcaaga aatatccagg atatgattca aaaaaggatg agtattttgg ggtgtatgaa 300
caaatcttaa accttaataa agaaagcttt aataagcttg ttagagataa tcatacgtat 360
ggtaacattc ctacaagtcc agaggaactt tgggataaag tatctaaact gatttatatt 420
tatttgaaag accctacaaa tgttattgga caagctgggt ggacgaatcc acaggatgca 480
atgtatgaat tttatactgt tgtacaacag gaaatatggc gttatacaga tggacaaaag 540
gtggataaag acaccaattc atatttgtat tataaatatt caaaacaagg tcaaaaagca 600
gtgtacttac tgcgtgacgc tgtgaatagc atcagtatac ctagtaattt tgaacttcgt 660
ggctataaac ctgaatgggt tcaaggtcaa aaaggatacc aagctattgt aactggtaga 720
ttgaaagtag atcaacctgt cggggaaata aagactacag taacagcagg tggaaaaacc 780
tcaagtgaaa acgacattgc tacattgaag gcgcaagacg ttataggtgg ggttgaagtc 840
tctgataaga taacatatag tggtctttat ccaaatacag aatatgatgt tataggtgaa 900
atttacgaag taaaagatgg agaacttgtt aatccaggac gaccggtttc tgtagtcaat 960
agtggtgacg atttaaaaac agatgcaaca ggaaaaggga aatggacatt aaactttgga 1020
aagcttgatt tagaagcagg aaaatcctat gtggtctttg aaaaagttgt ttcattaaaa 1080
aacgtgatag atacagatgg agatggaaaa ccggataaaa aacaagaact atcgcataat 1140
gatccaaaag ataaatcgca aacatttaca attttaccta aggaaatagt tgaacaagac 1200
gttgtcttca gtaaggtgaa tgtggctggt gaagaaatcg ctggtgcgaa gatccaactg 1260
aaggatgcgc aaggtcaagt tgttcattcc tggacttcta aagcgggtca aagtgaaacg 1320
gtcaaattga aagctggcac ctatactttc catgaagcat ccgctccgac tggttacttg 1380
gccgtaacgg atatcacatt cgaagtagat gttcaaggaa aagtgacggt taaggatgcc 1440
aacggcaatg gtgttaaggc ggatggtaat aagttaacgg tgaccgatca agctgctcct 1500
agcgtaccga atgaacaaga cgttgtcttc agtaaggtga atgtggctgg tgaagaaatc 1560
gctggtgcga agatccaact gaaggatgcg caaggtcaag ttgttcattc ctggacttct 1620
aaagcgggtc aaagtgaaac ggtcaaattg aaagctggca cctatacttt ccatgaagca 1680
tccgctccga ctggttactt ggccgtaacg gatatcacat tcgaagtaga tgttcaagga 1740
aaagtgacgg ttaaggatgc caacggcaat ggtgttaagg cggatggtaa taagttaacg 1800
gtgaccgatc aagctgctcc tagcgtaccg aatgaacaag acgttgtctt cagtaaggtg 1860
aatgtggctg gtgaagaaat cgctggtgcg aagatccaac tgaaggatgc gcaaggtcaa 1920
gttgttcatt cctggacttc taaagcgggt caaagtgaaa cggtcaaatt gaaagctggc 1980
acctatactt tccatgaagc atccgctccg actggttact tggccgtaac ggatatcaca 2040
ttcgaagtag atgttcaagg aaaagtgacg gttaaggatg ccaacggcaa tggtgttaag 2100
gcggatggta ataagttaac ggtgaccgat caagctgctc ctagcgtacc gaatgaacaa 2160
gacgttgtct tcagtaaggt gaatgtggct ggtgaagaaa tcgctggtgc gaaga 2215
<210> 17
<211> 5
<212> PRT
<213> 人工序列
<220>
<223> 合成的
<220>
<221> MISC_FEATURE
<222> (3)..(3)
<223> X是任意氨基酸
<400> 17
Leu Pro Xaa Thr Gly
1 5
<210> 18
<211> 6
<212> PRT
<213> 人工序列
<220>
<223> 合成的
<400> 18
Leu Pro Thr Gly Ala Ala
1 5
<210> 19
<211> 6
<212> PRT
<213> 人工序列
<220>
<223> 合成的
<400> 19
Leu Pro Thr Gly Gly Gly
1 5
<210> 20
<211> 6
<212> PRT
<213> 人工序列
<220>
<223> 合成的
<400> 20
Leu Pro Lys Thr Gly Gly
1 5
<210> 21
<211> 5
<212> PRT
<213> 人工序列
<220>
<223> 合成的
<400> 21
Leu Pro Glu Thr Gly
1 5
<210> 22
<211> 5
<212> PRT
<213> 人工序列
<220>
<223> 合成的
<220>
<221> MISC_FEATURE
<222> (3)..(3)
<223> X是任意氨基酸
<400> 22
Leu Pro Xaa Thr Gly
1 5
<210> 23
<211> 6
<212> PRT
<213> 人工序列
<220>
<223> 合成的
<220>
<221> MISC_FEATURE
<222> (3)..(3)
<223> X是任意氨基酸
<220>
<221> MISC_FEATURE
<222> (6)..(6)
<223> X是任意氨基酸
<400> 23
Leu Pro Xaa Thr Gly Xaa
1 5
<210> 24
<211> 5
<212> PRT
<213> 人工序列
<220>
<223> 合成的
<220>
<221> MISC_FEATURE
<222> (3)..(3)
<223> X是谷氨酰胺或赖氨酸
<220>
<221> MISC_FEATURE
<222> (5)..(5)
<223> X是天冬酰胺或甘氨酸
<400> 24
Asn Pro Xaa Thr Xaa
1 5
<210> 25
<211> 40
<212> PRT
<213> 人工序列
<220>
<223> 合成的
<400> 25
Met Gly Ser Ser His His His His His His Ser Ser Gly Leu Val Pro
1 5 10 15
Arg Gly Ser Val Pro Thr Ile Val Met Val Asp Ala Tyr Lys Arg Tyr
20 25 30
Lys Gly Ser Gly Glu Ser Gly Lys
35 40
<210> 26
<211> 139
<212> PRT
<213> 人工序列
<220>
<223> 合成的
<400> 26
Met Ser Tyr Tyr His His His His His His Asp Tyr Asp Ile Pro Thr
1 5 10 15
Thr Glu Asn Leu Tyr Phe Gln Gly Ala Met Val Thr Thr Leu Ser Gly
20 25 30
Leu Ser Gly Glu Gln Gly Pro Ser Gly Asp Met Thr Thr Glu Glu Asp
35 40 45
Ser Ala Thr His Ile Lys Phe Ser Lys Arg Asp Glu Asp Gly Arg Glu
50 55 60
Leu Ala Gly Ala Thr Met Glu Leu Arg Asp Ser Ser Gly Lys Thr Ile
65 70 75 80
Ser Thr Trp Ile Ser Asp Gly His Val Lys Asp Phe Tyr Leu Tyr Pro
85 90 95
Gly Lys Tyr Thr Phe Val Glu Thr Ala Ala Pro Asp Gly Tyr Glu Val
100 105 110
Ala Thr Ala Ile Thr Phe Thr Val Asn Glu Gln Gly Gln Val Thr Val
115 120 125
Asn Gly Glu Ala Thr Lys Gly Asp Ala His Thr
130 135
<210> 27
<211> 15
<212> PRT
<213> 人工序列
<220>
<223> 合成的
<400> 27
Val Pro Thr Ile Val Met Val Asp Ala Tyr Lys Arg Tyr Lys Ser
1 5 10 15
<210> 28
<211> 119
<212> PRT
<213> 人工序列
<220>
<223> 合成的
<400> 28
Tyr Phe Gln Gly Ala Met Val Thr Thr Leu Ser Gly Leu Ser Gly Glu
1 5 10 15
Gln Gly Pro Ser Gly Asp Met Thr Thr Glu Glu Asp Ser Ala Thr His
20 25 30
Ile Lys Phe Ser Lys Arg Asp Glu Asp Gly Arg Glu Leu Ala Gly Ala
35 40 45
Thr Met Glu Leu Arg Asp Ser Ser Gly Lys Thr Ile Ser Thr Trp Ile
50 55 60
Ser Asp Gly His Val Lys Asp Phe Tyr Leu Tyr Pro Gly Lys Tyr Thr
65 70 75 80
Phe Val Glu Thr Ala Ala Pro Asp Gly Tyr Glu Val Ala Thr Ala Ile
85 90 95
Thr Phe Thr Val Asn Glu Gln Gly Gln Val Thr Val Asn Gly Glu Ala
100 105 110
Thr Lys Gly Asp Ala His Thr
115
<210> 29
<211> 1689
<212> DNA
<213> 人工序列
<220>
<223> 示例性SpyCatcher-pIII序列
<220>
<221> CDS
<222> (1)..(1689)
<220>
<221> misc_feature
<222> (1)..(6)
<223> EcoRI (2)
<220>
<221> misc_feature
<222> (7)..(63)
<223> DsbA信号肽
<220>
<221> misc_feature
<222> (64)..(411)
<223> SpyCatcher
<220>
<221> misc_feature
<222> (412)..(432)
<223> TEV切割位点
<220>
<221> misc_feature
<222> (433)..(468)
<223> 接头
<220>
<221> misc_feature
<222> (463)..(468)
<223> Xba1
<220>
<221> misc_feature
<222> (469)..(1686)
<223> gIII
<400> 29
gaa ttc atg aag aag atc tgg tta gcg tta gct ggt ctt gtg ttg gcg 48
Glu Phe Met Lys Lys Ile Trp Leu Ala Leu Ala Gly Leu Val Leu Ala
1 5 10 15
ttt agt gct agt gcc ggg gca atg gtg gac acg ttg tca ggt ctg agc 96
Phe Ser Ala Ser Ala Gly Ala Met Val Asp Thr Leu Ser Gly Leu Ser
20 25 30
tca gaa cag ggt cag tcg ggt gat atg acc att gag gaa gat agc gca 144
Ser Glu Gln Gly Gln Ser Gly Asp Met Thr Ile Glu Glu Asp Ser Ala
35 40 45
acc cac atc aaa ttt agc aag cgc gat gaa gat ggc aaa gag ctt gcc 192
Thr His Ile Lys Phe Ser Lys Arg Asp Glu Asp Gly Lys Glu Leu Ala
50 55 60
ggc gcg aca atg gag tta cgg gat tct tcc ggc aag acc att agc acc 240
Gly Ala Thr Met Glu Leu Arg Asp Ser Ser Gly Lys Thr Ile Ser Thr
65 70 75 80
tgg att tcc gat ggc cag gta aaa gac ttc tat ctg tat ccg ggg aaa 288
Trp Ile Ser Asp Gly Gln Val Lys Asp Phe Tyr Leu Tyr Pro Gly Lys
85 90 95
tat acc ttt gtc gaa acg gct gca cct gac ggc tat gag gtt gca acc 336
Tyr Thr Phe Val Glu Thr Ala Ala Pro Asp Gly Tyr Glu Val Ala Thr
100 105 110
gcc atc aca ttc acg gtt aac gaa cag ggt cag gta acc gtc aat ggc 384
Ala Ile Thr Phe Thr Val Asn Glu Gln Gly Gln Val Thr Val Asn Gly
115 120 125
aaa gcg act aaa ggc gat gcg cat att gag aat ctg tac ttt caa ggc 432
Lys Ala Thr Lys Gly Asp Ala His Ile Glu Asn Leu Tyr Phe Gln Gly
130 135 140
ggc gga ggc ggg tct ggc ggt ggt ggt tcg tct aga gct gaa act gtt 480
Gly Gly Gly Gly Ser Gly Gly Gly Gly Ser Ser Arg Ala Glu Thr Val
145 150 155 160
gaa agt tgt tta gca aaa ccc cat aca gaa aat tca ttt act aac gtc 528
Glu Ser Cys Leu Ala Lys Pro His Thr Glu Asn Ser Phe Thr Asn Val
165 170 175
tgg aaa gac gac aaa act tta gat cgt tac gct aac tat gag ggc tgt 576
Trp Lys Asp Asp Lys Thr Leu Asp Arg Tyr Ala Asn Tyr Glu Gly Cys
180 185 190
ctg tgg aat gct aca ggc gtt gta gtt tgt act ggt gac gaa act cag 624
Leu Trp Asn Ala Thr Gly Val Val Val Cys Thr Gly Asp Glu Thr Gln
195 200 205
tgt tac ggt aca tgg gtt cct att ggg ctt gct atc cct gaa aat gag 672
Cys Tyr Gly Thr Trp Val Pro Ile Gly Leu Ala Ile Pro Glu Asn Glu
210 215 220
ggt ggt ggc tct gag ggt ggc ggt tct gag ggt ggc ggt tct gag ggt 720
Gly Gly Gly Ser Glu Gly Gly Gly Ser Glu Gly Gly Gly Ser Glu Gly
225 230 235 240
ggc ggt act aaa cct cct gag tac ggt gat aca cct att ccg ggc tat 768
Gly Gly Thr Lys Pro Pro Glu Tyr Gly Asp Thr Pro Ile Pro Gly Tyr
245 250 255
act tat atc aac cct ctc gac ggc act tat ccg cct ggt act gag caa 816
Thr Tyr Ile Asn Pro Leu Asp Gly Thr Tyr Pro Pro Gly Thr Glu Gln
260 265 270
aac ccc gct aat cct aat cct tct ctt gag gag tct cag cct ctt aat 864
Asn Pro Ala Asn Pro Asn Pro Ser Leu Glu Glu Ser Gln Pro Leu Asn
275 280 285
act ttc atg ttt cag aat aat agg ttc cga aat agg cag ggg gca tta 912
Thr Phe Met Phe Gln Asn Asn Arg Phe Arg Asn Arg Gln Gly Ala Leu
290 295 300
act gtt tat acg ggc act gtt act caa ggc act gac ccc gtt aaa act 960
Thr Val Tyr Thr Gly Thr Val Thr Gln Gly Thr Asp Pro Val Lys Thr
305 310 315 320
tat tac cag tac act cct gta tca tca aaa gcc atg tat gac gct tac 1008
Tyr Tyr Gln Tyr Thr Pro Val Ser Ser Lys Ala Met Tyr Asp Ala Tyr
325 330 335
tgg aac ggt aaa ttc aga gac tgc gct ttc cat tct ggc ttt aat gag 1056
Trp Asn Gly Lys Phe Arg Asp Cys Ala Phe His Ser Gly Phe Asn Glu
340 345 350
gat cca ttc gtt tgt gaa tat caa ggc caa tcg tct gac ctg cct caa 1104
Asp Pro Phe Val Cys Glu Tyr Gln Gly Gln Ser Ser Asp Leu Pro Gln
355 360 365
cct cct gtc aat gct ggc ggc ggc tct ggt ggt ggt tct ggt ggc ggc 1152
Pro Pro Val Asn Ala Gly Gly Gly Ser Gly Gly Gly Ser Gly Gly Gly
370 375 380
tct gag ggt ggt ggc tct gag ggt ggc ggt tct gag ggt ggc ggc tct 1200
Ser Glu Gly Gly Gly Ser Glu Gly Gly Gly Ser Glu Gly Gly Gly Ser
385 390 395 400
gag gga ggc ggt tcc ggt ggt ggc tct ggt tcc ggt gat ttt gat tat 1248
Glu Gly Gly Gly Ser Gly Gly Gly Ser Gly Ser Gly Asp Phe Asp Tyr
405 410 415
gaa aag atg gca aac gct aat aag ggg gct atg acc gaa aat gcc gat 1296
Glu Lys Met Ala Asn Ala Asn Lys Gly Ala Met Thr Glu Asn Ala Asp
420 425 430
gaa aac gcg cta cag tct gac gct aaa ggc aaa ctt gat tct gtc gct 1344
Glu Asn Ala Leu Gln Ser Asp Ala Lys Gly Lys Leu Asp Ser Val Ala
435 440 445
act gat tac ggt gct gct atc gat ggt ttc att ggt gac gtt tcc ggc 1392
Thr Asp Tyr Gly Ala Ala Ile Asp Gly Phe Ile Gly Asp Val Ser Gly
450 455 460
ctt gct aat ggt aat ggt gct act ggt gat ttt gct ggc tct aat tcc 1440
Leu Ala Asn Gly Asn Gly Ala Thr Gly Asp Phe Ala Gly Ser Asn Ser
465 470 475 480
caa atg gct caa gtc ggt gac ggt gat aat tca cct tta atg aat aat 1488
Gln Met Ala Gln Val Gly Asp Gly Asp Asn Ser Pro Leu Met Asn Asn
485 490 495
ttc cgt caa tat tta cct tcc ctc cct caa tcg gtt gaa tgt cgc cct 1536
Phe Arg Gln Tyr Leu Pro Ser Leu Pro Gln Ser Val Glu Cys Arg Pro
500 505 510
ttt gtc ttt ggc gct ggt aaa cca tat gaa ttt tct att gat tgt gac 1584
Phe Val Phe Gly Ala Gly Lys Pro Tyr Glu Phe Ser Ile Asp Cys Asp
515 520 525
aaa ata aac tta ttc cgt ggt gtc ttt gcg ttt ctt tta tat gtt gcc 1632
Lys Ile Asn Leu Phe Arg Gly Val Phe Ala Phe Leu Leu Tyr Val Ala
530 535 540
acc ttt atg tat gta ttt tct acg ttt gct aac ata ctg cgt aat aag 1680
Thr Phe Met Tyr Val Phe Ser Thr Phe Ala Asn Ile Leu Arg Asn Lys
545 550 555 560
gag tct taa 1689
Glu Ser
<210> 30
<211> 562
<212> PRT
<213> 人工序列
<220>
<223> 合成的构建体
<400> 30
Glu Phe Met Lys Lys Ile Trp Leu Ala Leu Ala Gly Leu Val Leu Ala
1 5 10 15
Phe Ser Ala Ser Ala Gly Ala Met Val Asp Thr Leu Ser Gly Leu Ser
20 25 30
Ser Glu Gln Gly Gln Ser Gly Asp Met Thr Ile Glu Glu Asp Ser Ala
35 40 45
Thr His Ile Lys Phe Ser Lys Arg Asp Glu Asp Gly Lys Glu Leu Ala
50 55 60
Gly Ala Thr Met Glu Leu Arg Asp Ser Ser Gly Lys Thr Ile Ser Thr
65 70 75 80
Trp Ile Ser Asp Gly Gln Val Lys Asp Phe Tyr Leu Tyr Pro Gly Lys
85 90 95
Tyr Thr Phe Val Glu Thr Ala Ala Pro Asp Gly Tyr Glu Val Ala Thr
100 105 110
Ala Ile Thr Phe Thr Val Asn Glu Gln Gly Gln Val Thr Val Asn Gly
115 120 125
Lys Ala Thr Lys Gly Asp Ala His Ile Glu Asn Leu Tyr Phe Gln Gly
130 135 140
Gly Gly Gly Gly Ser Gly Gly Gly Gly Ser Ser Arg Ala Glu Thr Val
145 150 155 160
Glu Ser Cys Leu Ala Lys Pro His Thr Glu Asn Ser Phe Thr Asn Val
165 170 175
Trp Lys Asp Asp Lys Thr Leu Asp Arg Tyr Ala Asn Tyr Glu Gly Cys
180 185 190
Leu Trp Asn Ala Thr Gly Val Val Val Cys Thr Gly Asp Glu Thr Gln
195 200 205
Cys Tyr Gly Thr Trp Val Pro Ile Gly Leu Ala Ile Pro Glu Asn Glu
210 215 220
Gly Gly Gly Ser Glu Gly Gly Gly Ser Glu Gly Gly Gly Ser Glu Gly
225 230 235 240
Gly Gly Thr Lys Pro Pro Glu Tyr Gly Asp Thr Pro Ile Pro Gly Tyr
245 250 255
Thr Tyr Ile Asn Pro Leu Asp Gly Thr Tyr Pro Pro Gly Thr Glu Gln
260 265 270
Asn Pro Ala Asn Pro Asn Pro Ser Leu Glu Glu Ser Gln Pro Leu Asn
275 280 285
Thr Phe Met Phe Gln Asn Asn Arg Phe Arg Asn Arg Gln Gly Ala Leu
290 295 300
Thr Val Tyr Thr Gly Thr Val Thr Gln Gly Thr Asp Pro Val Lys Thr
305 310 315 320
Tyr Tyr Gln Tyr Thr Pro Val Ser Ser Lys Ala Met Tyr Asp Ala Tyr
325 330 335
Trp Asn Gly Lys Phe Arg Asp Cys Ala Phe His Ser Gly Phe Asn Glu
340 345 350
Asp Pro Phe Val Cys Glu Tyr Gln Gly Gln Ser Ser Asp Leu Pro Gln
355 360 365
Pro Pro Val Asn Ala Gly Gly Gly Ser Gly Gly Gly Ser Gly Gly Gly
370 375 380
Ser Glu Gly Gly Gly Ser Glu Gly Gly Gly Ser Glu Gly Gly Gly Ser
385 390 395 400
Glu Gly Gly Gly Ser Gly Gly Gly Ser Gly Ser Gly Asp Phe Asp Tyr
405 410 415
Glu Lys Met Ala Asn Ala Asn Lys Gly Ala Met Thr Glu Asn Ala Asp
420 425 430
Glu Asn Ala Leu Gln Ser Asp Ala Lys Gly Lys Leu Asp Ser Val Ala
435 440 445
Thr Asp Tyr Gly Ala Ala Ile Asp Gly Phe Ile Gly Asp Val Ser Gly
450 455 460
Leu Ala Asn Gly Asn Gly Ala Thr Gly Asp Phe Ala Gly Ser Asn Ser
465 470 475 480
Gln Met Ala Gln Val Gly Asp Gly Asp Asn Ser Pro Leu Met Asn Asn
485 490 495
Phe Arg Gln Tyr Leu Pro Ser Leu Pro Gln Ser Val Glu Cys Arg Pro
500 505 510
Phe Val Phe Gly Ala Gly Lys Pro Tyr Glu Phe Ser Ile Asp Cys Asp
515 520 525
Lys Ile Asn Leu Phe Arg Gly Val Phe Ala Phe Leu Leu Tyr Val Ala
530 535 540
Thr Phe Met Tyr Val Phe Ser Thr Phe Ala Asn Ile Leu Arg Asn Lys
545 550 555 560
Glu Ser
Claims (28)
1.一种载体的组合,包括:
a)第一载体,其包含包括编码在框内与系连序列融合的感兴趣的蛋白质(POI)的多核苷酸的遗传构建体(GC1);和
b)第二载体,其包含包括编码在框内与捕获序列融合的锚蛋白的多核苷酸的遗传构建体(GC2),
系连序列和捕获序列当自发地或借助于酶相互接触时形成共价键,或一种载体的组合,包括:
a)第一载体,其包含包括编码在框内与捕获序列融合的感兴趣的蛋白质(POI)的多核苷酸的遗传构建体(GC1);和
b)第二载体,其包含包括编码在框内与系连序列融合的锚蛋白的多核苷酸的遗传构建体(GC2),
系连序列和捕获序列当自发地或借助于酶相互接触时形成共价键。
2.如权利要求1所述的载体的组合,其中,每个载体包含用于表达POI和锚蛋白的转录或翻译控制序列。
3.如权利要求1所述的载体的组合,其中,所述转录或翻译控制序列选自复制起点、启动子、增强子、阻遏物结合区、转录起始位点、核糖体结合位点、翻译起始位点和针对转录和翻译的终止位点。
4.如权利要求1-3中任一项所述的载体的组合,其中,所述POI选自抗体、抗体片段、单链抗体、scFv、scFab、单结构域抗体、蛋白质支架(例如基于纤连蛋白III、半胱氨酸蛋白酶抑制剂、脂蛋白、锚蛋白重复结构域、蛋白A的Z结构域等)、酶、激素、白介素、用于开发疫苗的抗原、人生长激素(hGH)、N-甲硫基人生长激素、牛生长激素、甲状旁腺激素、甲状腺素、胰岛素A链、胰岛素B链、胰岛素原、松弛素A链、松弛素B链、前松弛素、卵泡刺激素(FSH)、甲状腺刺激素(TSH)、黄体化激素(LH)、糖蛋白激素受体、降钙素、胰高血糖素、VIII因子、肺表面活性剂、尿激酶、链激酶、人类组织型纤溶酶原激活物(t-PA)、蛙皮素、IX因子、凝血酶、造血生长因子、肿瘤坏死因子-α和-β、脑啡肽酶、人血清白蛋白、穆勒氏抑制物、小鼠促性腺激素相关肽、微生物蛋白、β-内酰胺酶、组织因子蛋白、抑制素、激活素、血管内皮生长因子、激素或生长因子的受体;整联蛋白、血小板生成素、蛋白A或D、类风湿因子、神经生长因子、NGF-β、血小板生长因子、转化生长因子(TGF)、TGF-α、TGF-β、胰岛素样生长因子-I或-II、胰岛素样生长因子结合蛋白、DNA酶、潜伏期相关肽、促红细胞生成素、骨诱导因子、干扰素、干扰素-α、干扰素-β、干扰素-γ、集落刺激因子(CSF)、M-CSF、GF-CSF和G-CSF、白介素(IL)、IL-1、IL-2、IL-3、IL-4、超氧化物歧化酶;衰变促进因子、病毒抗原、HIV包膜蛋白、GP120、GP140、心房利钠肽A、B或C。
5.如权利要求1-4中任一项所述的载体的组合,其中,所述系连序列选自:选自SEQ IDNO:1、3、5、6、25或27中的任一项的SpyTag序列,与SEQ ID NO:1、3、5、6、25或27中任一项所述的氨基酸序列具有至少50%序列同一性的蛋白质,选自SEQ ID NO:13、7、11、9、10或15中的任一项的SnoopTag序列,或与SEQ ID NO:13、7、11、9、10或15中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,分选酶识别结构域或分选酶桥接结构域,蝶豆粘酶识别基序或分裂内含蛋白。
6.如权利要求1-5中任一项所述的载体的组合,其中,所述捕获序列是选自SEQ ID NO:1、3、5、6、26或yfqgamvttlsglsgeqgpsgdmtteedsathikfskrdedgrelagatmelrdssgktistwisdghvkdfylypgkytfvetaapdgyevataitftvneqgqvtvngeatkgdaht(SEQ ID NO:28)中任一项的SpyCatcher序列,与SEQ ID NO:1、3、5、6、26或yfqgamvttlsglsgeqgpsgdmtteedsathikfskrdedgrelagatmelrdssgktistwisdghvkdfylypgkytfvetaapdgyevataitftvneqgqvtvngeatkgdaht(SEQ ID NO:28)中任一项所述的氨基酸序列具有至少50%序列同一性的蛋白质,选自SEQ ID NO:13、7、11、9、10或15中的任一项的SnoopCatcher序列,或与SEQ ID NO:13、7、11、9、10或15中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,分选酶识别结构域或分选酶桥接结构域,蝶豆粘酶识别基序或分裂内含蛋白。
7.如权利要求1-6中任一项所述的载体的组合,其中,所述锚蛋白选自pIII、pVI、pVII、pVIII、pIX、细菌细胞外膜蛋白、菌毛、鞭毛、修饰的细菌脂蛋白、冰核蛋白、自转运蛋白、LamB、OmpA、Lpp-OmpA、INP、FliC、FimA、蛋白A、LysM、FnBPB、M6、SbpA、RsaA、细胞壁蛋白α-凝集素1、α-凝集素2、Cwp1、Cwp2、Gas1p、Yap3p、Flo1p、Crh2p、Pir1、Pir2、Pir4、Icwp、HpSEDI、HpGASI、HpTIPI、HPWPI、HwpIp、Als3p、Rbt5p、表面受体或GPI锚序列。
8.一种原核或真核宿主细胞,其包含如权利要求1-7中任一项所述的载体的组合。
9.如权利要求8所述的原核或真核宿主细胞,所述宿主细胞包含:包含与编码衍生自SEQ ID NO:13、7、11、9、10或15的连接酶或与衍生自SEQ ID NO:13、7、11、9、10或15的连接酶具有至少70%序列同一性的连接酶的多核苷酸可操作地连接的启动子的载体;包含与编码分选酶的多核苷酸可操作地连接的启动子的载体;或包含与编码蝶豆粘酶的多核苷酸可操纵地连接的启动子的载体;或者所述宿主细胞包含掺入其基因组中的启动子,所述启动子可操作地连接至:编码衍生自SEQ ID NO:13、7、11、9、10或15的连接酶或与衍生自SEQ IDNO:13、7、11、9、10或15的连接酶具有至少70%序列同一性的连接酶的多核苷酸;编码分选酶的多核苷酸;或编码蝶豆粘酶的多核苷酸。
10.一种在噬菌体颗粒表面展示(多)肽/蛋白质的方法,包括:引起或允许将感兴趣的蛋白质(POI)附着到所述噬菌体颗粒的蛋白质包衣中的锚蛋白上,其中所述附着是由于与所述POI融合的系连序列和与所述锚蛋白融合的捕获序列之间形成肽键所致。
11.如权利要求10所述的方法,其中,所述系连序列选自:选自SEQ ID NO:1、3、5、6、25或27中的任一项的SpyTag序列,或与SEQ ID NO:1、3、5、6、25或27中任一项所述的氨基酸序列具有至少50%序列同一性的蛋白质,选自SEQ ID NO:13、7、11、9、10或15中的任一项的SnoopTag序列,或与SEQ ID NO:13、7、11、9、10或15中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,分选酶识别结构域或分选酶桥接结构域,蝶豆粘酶识别基序或分裂内含蛋白。
12.如权利要求10所述的方法,其中,所述捕获序列是选自SEQ ID NO:1、3、5、6、26或28中任一项的SpyCatcher序列,与SEQ ID NO:1、3、5、6、26或28中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,选自SEQ ID NO:13、7、11、9、10或15中的任一项的SnoopCatcher序列,或与SEQ ID NO:13、7、11、9、10或15中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,分选酶识别结构域或分选酶桥接结构域,蝶豆粘酶识别基序或分裂内含蛋白。
13.如权利要求10-12中任一项所述的方法,其中,所述锚蛋白选自pIII、pVI、pVII、pVIII、或pIX。
14.如权利要求10-13中任一项所述的方法,其中,所述POI选自抗体、抗体片段、单链抗体、scFv、scFab、单结构域抗体、蛋白质支架(例如基于纤连蛋白III、半胱氨酸蛋白酶抑制剂、脂蛋白、锚蛋白重复结构域、蛋白A的Z结构域等)、酶、激素、白介素、用于开发疫苗的抗原、人生长激素(hGH)、N-甲硫基人生长激素、牛生长激素、甲状旁腺激素、甲状腺素、胰岛素A链、胰岛素B链、胰岛素原、松弛素A链、松弛素B链、前松弛素、卵泡刺激素(FSH)、甲状腺刺激素(TSH)、黄体化激素(LH)、糖蛋白激素受体、降钙素、胰高血糖素、VIII因子、肺表面活性剂、尿激酶、链激酶、人类组织型纤溶酶原激活物(t-PA)、蛙皮素、IX因子、凝血酶、造血生长因子、肿瘤坏死因子-d和-β、脑啡肽酶、人血清白蛋白、穆勒氏抑制物、小鼠促性腺激素相关肽、微生物蛋白、β-内酰胺酶、组织因子蛋白、抑制素、激活素、血管内皮生长因子、激素或生长因子的受体;整联蛋白、血小板生成素、蛋白A或D、类风湿因子、神经生长因子、NGF-β、血小板生长因子、转化生长因子(TGF)、TGF-α、TGF-β、胰岛素样生长因子-I或-II、胰岛素样生长因子结合蛋白、DNA酶、潜伏期相关肽、促红细胞生成素、骨诱导因子、干扰素、干扰素-α、干扰素-β、干扰素-γ、集落刺激因子(CSF)、M-CSF、GF-CSF和G-CSF、白介素(IL)、IL-1、IL-2、IL-3、IL-4、超氧化物歧化酶;衰变促进因子、病毒抗原、HIV包膜蛋白、GP120、GP140、和心房利钠肽A、B或C。
15.如权利要求14所述的方法,其中,所述POI是抗体、抗体片段、单链抗体、scFv、scFab或单结构域抗体。
16.一种在原核或真核细胞表面展示(多)肽/蛋白质的方法,包括:引起或允许将感兴趣的蛋白质(POI)附着到表达于所述原核或真核细胞的外表面的锚蛋白上,其中所述附着是由于与所述POI融合的系连序列和与所述锚蛋白融合的捕获序列之间形成肽键所致。
17.如权利要求16所述的方法,其中,所述系连序列选自:选自SEQ ID NO:1、3、5、6、25或27中的任一项的SpyTag序列,与SEQ ID NO:1、3、5、6、25或27中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,选自SEQ ID NO:13、7、11、9、10或15中的任一项的SnoopTag序列,或与SEQ ID NO:13、7、11、9、10或15中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,分选酶识别结构域或分选酶桥接结构域,蝶豆粘酶识别基序或分裂内含蛋白。
18.如权利要求16所述的方法,其中,所述捕获序列是选自SEQ ID NO:1、3、5、6、26或28中任一项的SpyCatcher序列,与SEQ ID NO:1、3、5、6、26或28中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,选自SEQ ID NO:13、7、11、9、10或15中的任一项的SnoopCatcher序列,或与SEQ ID NO:13、7、11、9、10或15中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,分选酶识别结构域或分选酶桥接结构域,蝶豆粘酶识别基序或分裂内含蛋白。
19.如权利要求16-18中任一项所述的方法,其中,所述方法包括在原核细胞表面上表达所述POI,并且所述锚蛋白选自外膜蛋白、菌毛、鞭毛、修饰的脂蛋白、冰核蛋白、自转运蛋白、LamB、OmpA、Lpp-OmpA、INP、FliC、FimA、蛋白A、LysM、FnBPB、M6、SbpA、或RsaA。
20.如权利要求16-18中任一项所述的方法,其中,所述方法包括在真核细胞表面上表达所述POI,并且所述锚蛋白选自细胞壁蛋白α-凝集素1、α-凝集素2、Cwp1、Cwp2、Gas1p、Yap3p、Flo1p、Crh2p、Pir1、Pir2、Pir4、Icwp、HpSEDI、HpGASI、HpTIPI、HPWPI、HwpIp、Als3p、Rbt5p、表面受体或GPI锚序列。
21.如权利要求16-19中任一项所述的方法,其中,所述POI选自抗体、抗体片段、单链抗体、scFv、scFab、单结构域抗体、蛋白质支架(例如基于纤连蛋白III、半胱氨酸蛋白酶抑制剂、脂蛋白、锚蛋白重复结构域、蛋白A的Z结构域等)、酶、激素、白介素、用于开发疫苗的抗原、人生长激素(hGH)、N-甲硫基人生长激素、牛生长激素、甲状旁腺激素、甲状腺素、胰岛素A链、胰岛素B链、胰岛素原、松弛素A链、松弛素B链、前松弛素、卵泡刺激素(FSH)、甲状腺刺激素(TSH)、黄体化激素(LH)、糖蛋白激素受体、降钙素、胰高血糖素、VIII因子、肺表面活性剂、尿激酶、链激酶、人类组织型纤溶酶原激活物(t-PA)、蛙皮素、IX因子、凝血酶、造血生长因子、肿瘤坏死因子-α和-β、脑啡肽酶、人血清白蛋白、穆勒氏抑制物、小鼠促性腺激素相关肽、微生物蛋白、β-内酰胺酶、组织因子蛋白、抑制素、激活素、血管内皮生长因子、激素或生长因子的受体;整联蛋白、血小板生成素、蛋白A或D、类风湿因子、神经生长因子、NGF-β、血小板生长因子、转化生长因子(TGF)、TGF-α、TGF-β、胰岛素样生长因子-I或-II、胰岛素样生长因子结合蛋白、DNA酶、潜伏期相关肽、促红细胞生成素、骨诱导因子、干扰素、干扰素-α、干扰素-β、干扰素-γ、集落刺激因子(CSF)、M-CSF、GF-CSF和G-CSF、白介素(IL)、IL-1、IL-2、IL-3、IL-4、超氧化物歧化酶;衰变促进因子、病毒抗原、HIV包膜蛋白、GP120、GP140、和心房利钠肽A、B或C。
22.如权利要求21所述的方法,其中,所述POI是抗体、抗体片段、单链抗体、scFv、scFab或单结构域抗体。
23.一种辅助噬菌体如VCSM13或M13KO7,其具有噬菌体DNA修饰以表达捕获序列-锚序列融合蛋白或系连序列-锚序列融合蛋白,其可用于感染原核细胞以产生噬菌体,该噬菌体在其表面带有捕获序列或系连序列。
24.如权利要求23所述的噬菌体,其中,所述系连序列选自:选自SEQ ID NO:1、3、5、6、25或27中的任一项的SpyTag序列,与SEQ ID NO:1、3、5、6、25或27中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,选自SEQ ID NO:13、7、11、9、10或15中的任一项的SnoopTag序列,或与SEQ ID NO:13、7、11、9、10或15中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,分选酶识别结构域或分选酶桥接结构域,蝶豆粘酶识别基序或分裂内含蛋白。
25.如权利要求23所述的噬菌体,其中,所述捕获序列是选自SEQ ID NO:1、3、5、6、26或28中任一项的SpyCatcher序列,与SEQ ID NO:1、3、5、6、26或28中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,选自SEQ ID NO:13、7、11、9、10或15中的任一项的SnoopCatcher序列,或与SEQ ID NO:13、7、11、9、10或15中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,分选酶识别结构域或分选酶桥接结构域,蝶豆粘酶识别基序或分裂内含蛋白。
26.一种原核或真核宿主细胞,其具有:经修饰的基因组,以表达捕获序列-锚序列融合蛋白或系连序列-锚序列融合蛋白,其可用于在其表面上或在宿主细胞产生的病毒或噬菌体表面上展示捕获或系连序列;和/或经修饰的基因组,以表达衍生自SEQ ID NO:13、7、11、9、10或15的连接酶或与衍生自SEQ ID NO:13、7、11、9、10或15的连接酶具有至少70%序列同一性的连接酶;包含与编码分选酶的多核苷酸可操作连接的启动子的载体;或包含与编码蝶豆粘酶的多核苷酸可操作连接的启动子的载体。
27.如权利要求26所述的原核或真核宿主细胞,其中,所述系连序列选自:选自SEQ IDNO:1、3、5、6、25或27中的任一项的SpyTag序列,与SEQ ID NO:1、3、5、6、25或27中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,选自SEQ ID NO:13、7、11、9、10或15中的任一项的SnoopTag序列,或与SEQ ID NO:13、7、11、9、10或15中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,分选酶识别结构域或分选酶桥接结构域,蝶豆粘酶识别基序或分裂内含蛋白。
28.如权利要求26所述的原核或真核宿主细胞,其中,所述捕获序列是选自SEQ ID NO:1、3、5、6、26或28中任一项的SpyCatcher序列,与SEQ ID NO:1、3、5、6、26或28中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,选自SEQ ID NO:13、7、11、9、10或15中的任一项的SnoopCatcher序列,或与SEQ ID NO:13、7、11、9、10或15中任一项所述的氨基酸序列具有至少70%序列同一性的蛋白质,分选酶识别结构域或分选酶桥接结构域,蝶豆粘酶识别基序或分裂内含蛋白。
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