CN113880953A - T细胞抗原受体、其多聚体复合物及其制备方法和应用 - Google Patents
T细胞抗原受体、其多聚体复合物及其制备方法和应用 Download PDFInfo
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Abstract
本发明提供了一种T细胞抗原受体、表达T细胞抗原受体(TCR)的免疫细胞,及其制备方法和应用。本发明所述的TCR能够特异性地被病毒抗原肽提呈细胞激活,提高胞外细胞因子IFNγ、IL2的释放水平及乳酸脱氢酶释放量,且显著地杀伤靶细胞。
Description
技术领域
本发明涉及能够识别CMV pp65的T细胞抗原受体技术领域,具体涉及多种TCR以及在治疗和/或预防CMV相关疾病中的用途。
背景技术
T细胞抗原受体(T cell receptor,TCR)对免疫系统的细胞免疫功能是至关重要的, TCR是呈递在主要组织相容性复合体(MHC)上的特异性抗原肽的唯一受体,通过抗原特异性TCR与pMHC复合物结合引发T细胞与抗原呈递细胞直接的物理接触,相互作用, 导致一系列后续细胞信号传递和其他生理反应,从而使得不同抗原特异性的T细胞对其 靶细胞发挥免疫效应。
CN107827959A公开了一种分离抗原特异性T细胞以及抗原特异性T细胞中克隆编码抗原特异性TCR基因的方法。该TCR能够与HBVS183-91抗原短肽复合物 FLLTRILTI-HLA*A0201结合,同时转导了本发明TCR的细胞能够被特异性激活并且对 靶细胞具有很强的杀伤作用。CN106279404A公开了一种可溶且稳定的异质二聚TCR, 其在α链可变区与β链恒定区之间含有人工链间二硫键的异质二聚TCR,该TCR可溶且 稳定,能够被很好地复性、重折叠、纯化同时能够与原配体特异性结合。但上述专利均未 公开针对巨细胞病毒的特异性TCR。
巨细胞病毒(Cytomegalocirus,CMV)是一种疱疹病毒组DNA病毒,也称细胞包涵体病毒。巨细胞病毒分布广泛,在人群中感染非常普遍,中国成人感染率达95%以上, 通常呈阴性感染,多数感染者无临床症状,但在一定条件下侵袭多个器官和系统可产生炎 症疾病。CN102656188A公开了能够识别来自巨细胞病毒的抗原的T细胞受体,其与通过 主要组织相容性复合物(MHC)呈递时的来自巨细胞病毒(CMV)磷蛋白pp65的肽特 异性结合,所述肽具有氨基酸序列NLVPMVATV。但该申请中T细胞受体对靶细胞的杀 伤作用不明显。
因此,本发明提供了一种T细胞抗原受体,能够特异性地被病毒抗原肽提呈细胞激活,提高胞外细胞因子IFNγ、IL2的释放水平及乳酸脱氢酶释放量,且显著地杀伤靶细 胞。
发明内容
本发明提供了多种T细胞抗原受体(TCR),其特异性结合MHC分子提呈的巨细胞 病毒(CMV)磷蛋白pp65抗原肽,所述的巨细胞病毒(CMV)磷蛋白pp65包含SEQ ID NO:34所示的氨基酸序列(其核苷酸序列如SEQ ID NO:35所示)。本发明所述的TCR 能够特异性地识别对应的CMV pp65抗原肽-MHC分子复合物,激活TCR-T细胞,进而 产生高水平的细胞因子IFNγ、IL2、TNFα,体内体外试验均显著杀伤肿瘤细胞。具体如 下:
本发明的第一方面,提供了一种T细胞抗原受体,所述的T细胞抗原受体特异 性结合CMV pp65。
优选的,结合表位包含SEQ ID NO:16。
优选的,所述的T细胞抗原受体通过主要组织相容性复合物分子(MHC)的呈 递,与来自巨细胞病毒(CMV)磷蛋白pp65的抗原肽特异性结合。
优选的,所述的T细胞抗原受体至少包含一个α链可变区和/或β链可变区。
优选的,所述的T细胞抗原受体是αβ异二聚体。
优选的,所述的T细胞抗原受体包含α链的CDR1α-CDR3α和β链的 CDR1β-CDR3β。
进一步优选的,CDR1α的氨基酸序列包含SEQ ID NO:1-2中的任一种或包含与 SEQID NO:1-2中任一种所示氨基酸序列具有至少80%同源性,CDR2α的氨基酸序 列包含SEQID NO:3-4中的任一种或包含与SEQ ID NO:3-4中任一种所示氨基酸 序列具有至少80%同源性,CDR3α的氨基酸序列包含SEQ ID NO:5-6中的任一种或 包含与SEQ ID NO:5-6中任一种所示氨基酸序列具有至少80%同源性,CDR1β的氨 基酸序列包含SEQ ID NO:7-8中的任一种或包含与SEQ ID NO:7-8中任一种所示 氨基酸序列具有至少80%同源性,CDR2β的氨基酸序列包含SEQ ID NO:9-10中的 任一种或包含与SEQ ID NO:9-10中任一种所示氨基酸序列具有至少80%同源性,CDR3β的氨基酸序列包含SEQ ID NO:11-12中的任一种或包含与SEQ ID NO:11-12 中任一种所示氨基酸序列具有至少80%同源性。
更进一步优选的,所述的CDR1α-CDR3α及CDR1β-CDR3β包含下列任一组:
优选的,其β链的氨基酸序列包含SEQ ID NO:43-44中的任一种或包含与SEQ IDNO:43-44任一种所示氨基酸序列具有至少80%同源性,其α链的氨基酸序列包 含SEQ IDNO:46-47中的任一种或包含与SEQ ID NO:46-47任一种所示氨基酸序 列具有至少80%同源性。
优选的,其α链与β链的氨基酸直接连接或间接连接,优选为间接连接,更优选 通过fp2A(furin-SGSG-p2A)连接。
进一步优选的,所述fp2A的氨基酸序列包含SEQ ID NO:54。
在本发明的一个具体实施方式中,所述的α链与β链的连接顺序,可以为α链、 fp2A与β链,或者,β链、fp2A与α链。
在本发明的一个具体实施方式中,其氨基酸序列包含SEQ ID NO:36或38或包 含与SEQ ID NO:36或38所示氨基酸序列具有至少80%同源性。
本发明的第二方面,提供了一种抗体或其抗原结合片段,所述的抗体或其抗原结合片段特异性结合CMV pp65。优选的,结合表位包含SEQ ID NO:16。
优选的,所述的抗体或其抗原结合片段包含α链的CDR1α-CDR3α和β链的 CDR1β-CDR3β。
在本发明的一个具体实施方式中,所述的抗体或其抗原结合片段的氨基酸序列包含SEQ ID NO:36或38或包含与SEQ ID NO:36或38所示氨基酸序列具有至少80% 同源性。
优选的,所述的抗体或其抗原结合片段还可以包含如Fab、Fab’、Fab’-SH、Fv、scFv、(Fab’)2、单结构域抗体、双抗体(dAb)或线性抗体的片段。
本发明的第三方面,提供了一种编码本发明所述的T细胞抗原受体的β链的核酸。
优选的,编码T细胞抗原受体的β链的核酸序列包含SEQ ID NO:49-50中任一 种或包含与SEQ ID NO:49-50中任一种所示核酸序列具有至少80%同源性。
本发明的第四方面,提供了一种编码本发明所述的T细胞抗原受体的α链的核酸。
优选的,编码T细胞抗原受体的α链的核酸序列包含SEQ ID NO:52-53中任一 种或包含与SEQ ID NO:52-53中任一种所示核酸序列具有至少80%同源性。
本发明的第五方面,提供了一种编码本发明所述的T细胞抗原受体或抗体或其抗原结合片段的核酸。
优选的,所述的编码T细胞抗原受体的α链与β链之间采用编码fp2A的核酸序 列连接。进一步优选的,所述的编码fp2A的核酸序列包含SEQ ID NO:55。
优选的,编码T细胞抗原受体的核酸序列包含SEQ ID NO:37或39或包含与 SEQ IDNO:37或39所示核酸序列具有至少80%同源性。
优选的,编码抗体或其抗原结合片段的核酸序列包含SEQ ID NO:37或39或包 含与SEQ ID NO:37或39所示核酸序列具有至少80%同源性。
本发明所述的核酸序列可以是单链或双链,可以是DNA也可以是RNA。
优选的,所述的核酸序列可以是经过密码子优化的。进一步优选的,所述的密码子优化包括将病毒等使用的大量稀有密码子变为对应的哺乳动物密码子和/或移除 mRNA不稳定基序和/或隐藏的剪接位点。
本发明的第六方面,提供了一种表达载体,所述的表达载体包含本发明任一所述的核酸。
优选的,所述的表达载体能够在体内或体外或离体条件下表达。进一步优选的,所述的表达载体在体内细胞中持续高水平表达。
优选的,所述的表达载体可以是原核表达载体或逆转录病毒载体。
进一步优选的,所述的原核表达载体为大肠杆菌系列。在本发明的一个具体实施方式中,所述的表达载体为pET-26b或pET28a+。
进一步优选可以是劳氏肉瘤病毒(RSV)、慢病毒、人免疫缺陷病毒(HIV)、 鼠科白血病病毒(MLV)、马传染性贫血病毒(EIAV)、小鼠乳腺癌病毒(MMTV)、 Fujinami肉瘤病毒(FuSV)、FBR鼠骨肉瘤病毒(FBR MSV)、莫洛尼氏鼠白血病 病毒(Mo-MLV)、莫洛尼氏鼠肉瘤病毒(Mo-MSV)、Abelson鼠白血病病毒(A-MLV)、 禽髓细胞增生病毒29(MC29)或禽骨髓成红细胞增多症病毒(AEV)等等。更进一 步优选的,所述的表达载体是慢病毒表达载体。
在本发明的一个具体实施方式中,所述的表达载体为pHAGE-IRES-RFP。
进一步优选的,所述的表达载体中β链、α链与骨架载体的连接顺序为启动子、 β链、fp2A、α链、IRES和RFP序列。
本发明的第七方面,提供了一种宿主细胞,所述的宿主细胞包含本发明任一所述的核酸或表达载体。
优选的,所述的宿主细胞可以是真核的或原核的。更优选的,所述的宿主细胞为酵母细胞、293细胞、CHO细胞、大肠杆菌等。
在本发明的一个具体实施方式中,所述的宿主细胞为Stbl3、BL21或transetta。
本发明的第八方面,提供了一种免疫细胞,所述的免疫细胞表达本发明所述的T细胞抗原受体或者抗体或其抗原结合片段。
优选的,所述的免疫细胞包含一个或多个本发明任一所述的异源的核酸序列。
优选的,所述的免疫细胞包括但不限于干细胞、淋巴细胞(包括T细胞、B细胞)。 进一步,所述免疫细胞是B细胞,所述B细胞表达上述的抗体或其抗原结合片段。 所述免疫细胞是T细胞,所述T细胞的T细胞抗原受体结构如上所限定。
优选的,所述的T细胞可以是CD4+T、CD8+T等。
进一步优选的,所述的免疫细胞分离自受试者的T细胞衍生。
优选的,所述的免疫细胞是来自CMV血清阴性供体的T细胞。
本发明的第九方面,提供了一种免疫细胞的制备方法,包括将编码上述抗体或其抗原结合片段,或者所述的T细胞抗原受体的核酸序列转染至免疫细胞中表达获得。
优选的,所述的免疫细胞包括但不限于干细胞、淋巴细胞(包括T细胞、B细胞)。 进一步,所述免疫细胞是B细胞,所述B细胞表达上述的抗体或其抗原结合片段。 所述免疫细胞是T细胞,所述T细胞的T细胞抗原受体结构如上所限定。
优选的,还包括敲除细胞内源性TCR的步骤。具体的,可以为将靶向内源TCR 的guide构建至慢病毒载体,与包装质粒、转染试剂共转至T细胞。
本发明的第十方面,提供了一种重组T细胞的制备方法,包括如下步骤:
1)从阳性T细胞克隆得到本发明任一所述的核酸;
2)分离、培养原代T细胞;
3)将步骤1)得到的核酸递送至步骤2)所述的原代T细胞中,获得表达本发明 任一所述T细胞抗原受体的重组T细胞。
优选的,所述的T细胞选自造血干细胞或外周血淋巴细胞(PBL)源T细胞。
优选的,还包括敲除细胞内源性TCR的步骤。具体的,可以为将靶向内源TCR 的guide构建至慢病毒载体,与包装质粒、转染试剂共转至T细胞。
本发明的第十一方面,提供了一种抗体或其抗原结合片段或T细胞抗原受体的制备方法,包括如下步骤:
(1)从阳性T细胞克隆得到本发明任一所述的核酸;
(2)将步骤(1)得到的核酸连接至载体骨架,获得表达载体;
(3)将步骤(2)获得的表达载体转化至宿主细胞,然后诱导其表达;
(4)获得抗体或其抗原结合片段或者T细胞抗原受体。
优选的,所述的阳性T细胞为与MHC呈递的巨细胞病毒(CMV)磷蛋白pp65 抗原肽特异性结合。进一步优选的,所述的MHC呈递的巨细胞病毒(CMV)磷蛋白 pp65抗原肽复合物为单体或多聚体复合物。
本发明的第十二方面,提供了一种多聚体复合物,所述多聚体复合物包含本发明任一所述的T细胞抗原受体。
优选的,所述多聚体复合物还包括单体,生物素分子,以及链霉亲和素分子或亲和素分子,其中,所述单体包括MHC分子α链胞外区和β2m链及抗原肽,所述单体 与所述生物素分子偶联,所述生物素分子与所述链霉亲和素或亲和素分子结合。
优选的,所述的MHC分子α链胞外区的C端连接avi-tag序列。
优选的,所述的MHC分子α链胞外区不含信号肽序列。并在成熟肽序列前面添 加M氨基酸。
优选的,所述的β2m链不含有信号肽序列。且在成熟肽序列前面添加M和A两 个氨基酸。
在本发明的一个具体实施方式中,所述的β2m链为不含信号肽且在成熟肽序列 前加了两个氨基酸,优选为M、A。
优选的,所述的抗原肽为SEQ ID NO:16。
优选的,所述的MHC分子为HLA-A*0201。
在本发明的一个具体实施方式中,所述的多聚体复合物包含:
(1)T细胞抗原受体;优选包含SEQ ID NO:36或38或包含与SEQ ID NO:36或38所示氨基酸序列具有至少80%同源性。
(3)单体,所述的单体包括抗原肽、C端连接avi-tag序列的MHC分子α链胞 外区和不含信号肽的β2m链;所述的抗原肽为SEQ ID NO:16;
(4)生物素分子;以及
(5)链霉亲和素分子或亲和素分子;其中,所述单体与所述生物素分子偶联, 所述生物素分子与所述链霉亲和素或亲和素结合。
优选的,所述的MHC分子为MHC I类分子或MHC II类分子。更优选的,所述 的MHC分子为MHC I类分子。
优选的,所述的MHC分子为HLA-A*0201。
在本发明的一个具体实施方式中,所述的MHC分子α链的氨基酸序列如SEQ ID NO:30所示或SEQ ID NO:30所示氨基酸序列具有至少80%同源性。
在本发明的一个具体实施方式中,所述的MHC分子α链的核苷酸序列如SEQ ID NO:31所示或SEQ ID NO:31所示核苷酸序列具有至少80%同源性。
在本发明的一个具体实施方式中,所述的MHC分子的β2m链的氨基酸序列如 SEQID NO:32所示或SEQ ID NO:32所示氨基酸序列具有至少80%同源性。
在本发明的一个具体实施方式中,所述的MHC分子的β2m链的核苷酸序列如 SEQID NO:33所示或SEQ ID NO:33所示氨基酸序列具有至少80%同源性。
为提高抗原肽-MHC四聚体与T细胞抗原受体结合的特异性,所述的单体还包括 至少一个氨基酸的化学修饰、突变、插入和/或缺失。
优选的,所述的MHC分子α链胞外区和β2m链为非共价结合。
优选的,所述的多聚体复合物至少包含一个单体。
优选的,每个单体至少偶联一个生物素分子。
本发明的第十三方面,提供了一种本发明任一所述的多聚体复合物的制备方法,包括如下步骤:
I)表达和纯化在C端连接avi-tag序列的MHC分子α链胞外区和β2m链;
II)将抗原肽、步骤I)获得的β2m链和在C端连接avi-tag序列的MHC分子α 链胞外区复性折叠,制备单体;
III)将步骤II)制备的单体生物素化,获得生物素化的单体;
IV)将步骤III)获得的生物素化的单体与带荧光标记的链霉亲和素或亲和素反应,制备抗原肽-MHC分子四聚体;
V)将步骤IV)得到的抗原肽-MHC分子四聚体与T细胞共孵育,形成T细胞抗 原受体与抗原肽-MHC分子四聚体复合物。
优选的,所述步骤I)中包括分别克隆在编码C端连接avi-tag序列的MHC分子 α链胞外区的核苷酸序列和MHC分子β2m链的核苷酸序列,连接至载体后,转化到 表达菌中培养,加入诱导剂,提取包涵体。
进一步优选的,所述表达菌培养至OD600值在0.2-0.4之间。
进一步优选的,所述诱导剂加入后的最终摩尔浓度为0.5-1mM。
优选的,诱导表达4-6h。
优选的,所述步骤II)包括β2m链的复性折叠,即将抗原肽、MHC分子β2m链、 C端连接avi-tag序列的MHC分子α链胞外区依次加入稀释缓冲液中避光水浴,其中, 所述的β2m链的复性折叠包括包涵体变性、加入蛋白酶抑制剂,而后透析。
优选的,所述的抗原肽、不含信号肽的β2m链与C端连接avi-tag序列的α链的 摩尔比为(30-50):(2-2.5):1。进一步优选为40:2:1。
优选的,所述步骤II)中还包含纯化单体的步骤。
优选的,所述步骤III)中的生物素化是在BirA酶的催化下,将单体与BiomixA、BiomixB结合。
优选的,所述步骤III)中还包括纯化生物素化单体的步骤。
优选的,所述步骤IV)中的单体与链霉亲和素反应的摩尔比为(4-7):1。
本发明的第十四方面,提供了一种上述多聚体复合物在制备、筛选或检测本发明所述抗体或其抗原结合片段或T细胞抗原受体中的应用。
本发明的第十五方面,提供了一种T细胞抗原受体的制备方法,包括如下步骤:
I)表达和纯化在C端连接avi-tag序列的MHC分子α链胞外区和β2m链;
II)将抗原肽、步骤I)获得的在C端连接avi-tag序列的MHC分子α链胞外区 和β2m链复性折叠,制备单体;
III)将步骤II)制备的单体生物素化,获得生物素化的单体;
IV)将步骤III)获得的生物素化的单体与带荧光标记的链霉亲和素或亲和素反应,制备抗原肽-MHC分子四聚体;
V)将步骤IV)得到的抗原肽-MHC分子四聚体与T细胞共孵育,形成T细胞抗 原受体与抗原肽-MHC分子四聚体复合物,分离获得T细胞抗原受体。
本发明的第十六方面,提供了本发明任一所述的T细胞抗原受体、本发明任一所述的抗体或其抗原结合片段、本发明任一所述的核酸、本发明所述的表达载体、本发 明所述的宿主细胞、本发明所述的免疫细胞、本发明任一所述的多聚体复合物在制备 诊断或治疗肿瘤或与CMV相关疾病的产品中的应用。
优选的,所述的CMV相关疾病选自新生儿CMV包涵体病、急性获得性CMV 感染或免疫妥协人员CMV感染导致的疾病。
在本发明的一个具体实施方式中,所述的CMV相关疾病选自肝、脾或中枢神经 系统疾病,残疾,传染性单核细胞增多症,骨骼肌疼痛,CMV视网膜炎,胃肠CMV 或脑炎等等。
本发明的第十七方面,提供了本发明任一所述的T细胞抗原受体、本发明任一所述的抗体或其抗原结合片段、本发明任一所述的核酸、本发明所述的表达载体、本发 明所述的宿主细胞、本发明所述的免疫细胞、本发明任一所述的多聚体复合物在T 细胞标记、检测、细胞分选或活化中的应用。
本发明的第十八方面,提供了一种药物组合物,所述的药物组合物包含下列任一组:
i)本发明所述的T细胞抗原受体;
ii)本发明所述的核酸;
iii)本发明所述的表达载体;
iv)本发明所述的宿主细胞;
v)本发明所述的免疫细胞;
vi)本发明所述的多聚体复合物;或
vii)本发明所述的抗体或其抗原结合片段。
优选的,所述的药物组合物还可以包含药学上可接受的辅料。
优选的,所述的药物组合物还可以与其他治疗剂共同使用。进一步优选的,所述的治疗剂可以为免疫调节剂。
本发明的第十九方面,提供了一种试剂盒,所述的试剂盒包含下列任一组:
i)本发明所述的T细胞抗原受体;
ii)本发明所述的核酸;
iii)本发明所述的表达载体;
iv)本发明所述的宿主细胞;
v)本发明所述的免疫细胞;
vi)本发明所述的多聚体复合物;或
vii)本发明所述的抗体或其抗原结合片段。
本发明的第二十方面,提供了一种检测巨细胞病毒(CMV)磷蛋白pp65的方法,所述的方法包括将待检测样品与本发明所述的抗体或其抗原结合片段或者T细胞抗原受体接触,然后检测巨细胞病毒(CMV)磷蛋白pp65与抗体或其抗原结合片段或者T细胞抗 原受体形成的复合物。
优选的,所述检测巨细胞病毒(CMV)磷蛋白pp65为检测巨细胞病毒(CMV)磷 蛋白pp65的存在或含量。其中,所述的存在表示有无,所述的含量可以为表达量或蛋白 浓度等。
优选的,所述的抗体或其抗原结合片段或者T细胞抗原受体包括可检测的标记物。
在本发明的一个具体实施方式中,所述的标记物可以是His和/或HA。
本发明所述的检测巨细胞病毒(CMV)磷蛋白pp65的方法,不是疾病的诊断方法。首先,待检测样品并非生物体或其离体组织或细胞,其次,即便生物体中存在巨细胞病毒(CMV)磷蛋白pp65或者包含一定浓度或表达水平的巨细胞病毒(CMV)磷蛋白pp65 也并非确定是疾病,只是一种可能性。
本发明的第二十一方面,提供了一种治疗和/或预防与CMV相关疾病的方法,所 述的方法包括给予个体有效量的本发明所述抗体或其抗原结合片段、所述的T细胞抗原 受体、所述的核酸、所述的表达载体、所述的宿主细胞、所述的免疫细胞或所述的药物组 合物。
优选的,所述的方法包括将表达本发明所述T细胞抗原受体的T细胞过继性转 移至受试者的步骤。
优选的,所述的方法包括将本发明所述的T细胞抗原受体定位在与CMV相关疾病(优选为肿瘤或转移性肿瘤)的附近,以提高毒素或免疫刺激剂的效力。
优选的,用于治疗和/或预防同种异体造血干细胞移植后的CMV再活化。
优选的,用于治疗和/或预防器官移植(例如肾、肝、胰、肠、角膜)、组织移植、 细胞移植(胰岛细胞、角膜缘干细胞)或干细胞治疗后的CMV再活化。
优选的,所述的表达本发明所述T细胞抗原受体的T细胞衍生自受试者。
优选的,所述的表达本发明所述T细胞抗原受体的T细胞与所述造血干细胞、 器官、组织、细胞或干细胞衍生自相同的供体。
本发明的第二十二方面,提供了一种诊断与CMV相关疾病的方法,所述的方法包括取样,将样品与本发明所述的抗体或其抗原结合片段或T细胞抗原受体接触,然后检测 CMVpp65与抗体或其抗原结合片段或T细胞抗原受体形成的复合物。
优选的,所述的抗体或其抗原结合片段或T细胞抗原受体包括可检测的标记物。
本发明所述的TCR能够特异性地识别对应的CMV pp65抗原肽-MHC分子复合物, 激活TCR-T细胞,进而产生高水平的细胞因子IFNγ、IL2、TNFα,体内体外试验均显著 杀伤肿瘤细胞。
本发明所述的“T细胞抗原受体”是能够识别当由MHC分子或其四聚体递呈时 的肽的分子。
本发明所述的“肿瘤”包括但不限于胰腺癌、肝癌、结肠癌、直肠癌、胃癌、淋巴瘤、基底细胞癌、非小细胞肺癌、白血病、卵巢癌、鼻咽癌、乳腺癌、子宫内膜癌、、膀胱癌、 肺癌、支气管癌、骨癌、前列腺癌、、胆管癌、食管癌、肾癌、甲状腺癌、头颈部癌、睾 丸癌、胶质母细胞瘤、星形细胞瘤、黑色素瘤、骨髓增生异常综合征、以及肉瘤。其中, 所述的白血病选自急性淋巴细胞性(成淋巴细胞性)白血病、急性骨髓性白血病、髓性白 血病、慢性淋巴细胞性白血病、多发性骨髓瘤、浆细胞白血病、以及慢性骨髓性白血病; 所述淋巴瘤选自霍奇金淋巴瘤和非霍奇金淋巴瘤,包括B细胞淋巴瘤、弥漫性大B细胞 淋巴瘤、滤泡性淋巴瘤、套细胞淋巴瘤、边缘区B细胞淋巴瘤、T细胞淋巴瘤、和瓦尔 登斯特伦巨球蛋白血症;所述肉瘤选自骨肉瘤、尤文肉瘤、平滑肌肉瘤、滑膜肉瘤、软组 织肉瘤、血管肉瘤、脂肪肉瘤、纤维肉瘤、横纹肌肉瘤、以及软骨肉瘤。优选的,所述的 肿瘤选自胰腺癌、肝癌、口腔鳞状上皮癌、结肠癌、卵巢癌、胃癌。在本发明的一个具体 实施方式中,所述的肿瘤为淋巴瘤。
本发明所述的“与CMV相关疾病”包含CMV pp65感染所导致的疾病,包括新 生儿CMV包涵体病,严重可影响肝、脾和中枢神经系统以及残疾。还包括急性获得 性CMV感染,类似于传染性单核细胞增多症,包括发烧、骨骼肌疼痛等等。还包括 免疫妥协人员,例如移植过器官的人,或患有HIV的人,感染可导致CMV视网膜炎、 胃肠CMV和脑炎等等。
本发明所述的“包含”在本申请中用于描述蛋白质或核酸的序列时,所述蛋白质或核 酸可以是由所述序列组成,或者在所述蛋白质或核酸的一端或两端可以具有额外的氨基酸 或核苷酸,但仍然具有本发明所述的活性。
本发明所述的“预防”是指通过施用本发明所述的产品来抑制症状或者延缓特定症状 紧张的所有行为。
本发明所述的“诊断”是指以查明患者过去、诊断时或将来是否患有疾病或病症,或者是查明疾病的进展或将来可能的进展,或者是评估患者对治疗的反应。
本发明所述的“治疗”表示减缓、中断、阻止、控制、停止、减轻、或逆转一种体征、症状、失调、病症、或疾病的进展或严重性,但不一定涉及所有疾病相关体征、症状、病 症、或失调的完全消除,且是指在疾病已开始发展后改善疾病或病理状态的体征、症状等 等的治疗干预。
本发明所述的“有效量”是指在以单个或多个剂量给予至患者或器官之后提供所希望 的治疗或预防的本发明所述的产品的量或剂量。
本发明所述的“产品”包括但不限于本发明所述的抗体或其抗原结合片段、所述的T 细胞抗原受体、所述的核酸、所述的表达载体、所述的宿主细胞、所述的免疫细胞或者所述的多聚体复合物,以及其他辅助或与上述上述产品协同的试剂。
本发明所述的“产品”可以为试剂盒、芯片、抗体偶联物或多功能抗体等药物组合物。
本发明所述的“受试者”包括但不限于人或非人哺乳动物。优选的,所述的非人哺乳动物包括但不限于小鼠、大鼠、猴子、猪或兔子等。
本发明所述的“同源性”是指在使用氨基酸序列或核苷酸序列的方面,本领域技术人员可以在不改变原序列主要结构或功能的前提下,根据实际工作需要对序列进行 调整,使使用序列与本发明所述的具体序列相比,具有(包括但不限于)1%,2%, 3%,4%,5%,6%,7%,8%,9%,10%,11%,12%,13%,14%,15%,16%,17%, 18%,19%,20%,21%,22%,23%,24%,25%,26%,27%,28%,29%,30%,31%,32%,33%,34%,35%,36%,37%,38%,39%,40%,41%,42%,43%, 44%,45%,46%,47%,48%,49%,50%,51%,52%,53%,54%,55%,56%, 57%,58%,59%,60%,70%,80%,81%,82%,83%,84%,85%,86%,87%, 88%,89%,90%,91%,92%,93%,94%,95%,96%,97%,98%,99%或100% 的同源性。例如,本发明所述的“与SEQ ID NO:1-2中任一种所示氨基酸序列具有至 少80%同源性”即为在保留与巨细胞病毒(CMV)磷蛋白pp65肽表位:MHC复合物 结合功能的前提下,可以根据实际工作需要对SEQ ID NO:1或2进行调整,包含一 个或多个改变例如取代、缺失和/或插入一个或多个氨基酸等,截短,或者一端或两 端的加长,只要其保持80%以上同源性并且保留与pp65表位:MHC复合物或pp65 表位:MHC分子四聚体结合的能力。所述的改变可以是取代、添加或缺失。可以是 相同极性的氨基酸之间的取代、相同电荷的氨基酸之间的取代、不带电荷氨基酸之间 的取代、脂肪族氨基酸之间的取代、芳香族氨基酸之间的取代、无极性氨基酸之间的 取代或者具有相关性质的侧链部分的氨基酸之间的取代。其中,碱性侧链包括但不限 于赖氨酸、精氨酸或组氨酸。酸性侧链包括但不限于天冬氨酸或谷氨酸。不带电荷的 氨基酸包括但不限于天冬氨酰、谷氨酰胺、丝氨酸、苏氨酸或酪氨酸。非极性侧链包 括但不限于甘氨酸、丙氨酸、缬氨酸、亮氨酸、异亮氨酸、脯氨酸、苯丙氨酸、甲硫 氨酸、色氨酸或半胱氨酸。其中,所述的至少80%包括但不限于80%,81%,82%, 83%,84%,85%,86%,87%,88%,89%,90%,91%,92%,93%,94%,95%, 96%,97%,98%,99%或100%。
附图说明
以下,结合附图来详细说明本发明的实施例,其中:
图1:抗原肽-MHC单体的SDS PAGE检测结果,其中,M为蛋白Marker,条带1、 2为单体;
图2:四聚体钓取特异性T细胞检测结果,具体为自主研发A0201-NLV四聚体与商业化四聚体钓取特异性T细胞的对比结果;
图3:TCRβ链和α链在pHAGE载体中连接示意图,其中连接顺序依次为启动子、 β链、furin-p2A、α链、IRES和RFP序列;
图4:流式细胞检测TCR的膜表面表达情况;具体为HLA-A*A0201 NLVPMVATV 特异性TCR(C2,C22,C29)的膜表面表达情况,其中,BV421是系列荧光素中的一种, BV的全称是Brilliant Violet;
图5:流式细胞检测TCR与CMV pp65四聚体探针结合的亲和力。具体为 HLA-A*A0201 NLVPMVATV特异性TCR(C2,C22,C29)与CMV pp65四聚体探针结合 的亲和力检测结果;
图6:C2-TCR-T、C22-TCR-T、C29-TCR-T细胞因子IL2、TNFα及IFNγ的释放水 平及靶细胞荧光素酶水平的检测结果图,其中,C2、C22、C29为实施例2制备的TCR, NE代表空白对照组,NT代表只有T细胞组,Raji为未转pp65的Raji细胞,1G4代表针 对NY-ESO-1特异的TCR;
图7:淋巴瘤动物模型中评价C22-TCR在小鼠体内抑制肿瘤生长情况;
图8:淋巴瘤动物模型,无T细胞注射组(PBS)、对照T细胞注射组(NC)、及CMV TCR-T注射组(C22-TCR)的小鼠存活率情况统计结果;
图9:淋巴瘤动物模型,无T细胞注射组(PBS)、对照T细胞注射组(NC)、及CMV TCR-T注射组(C22-TCR)的小鼠体重统计结果;
图10:淋巴瘤动物模型,无T细胞注射组(PBS)、对照T细胞注射组(NC)、及 CMVTCR-T注射组(C22-TCR)的小鼠体内TCR-T细胞特异增殖统计结果,结果显示, PBS组和NC组结果均为0,图10中6条折线为C22-TCR组6个不同小鼠体内TCR-T 细胞特异增殖统计结果。
具体实施方式
下面将结合本发明实施例中的附图,对本发明实施例中的技术方案进行清楚、完整地 描述,显然,所描述的实施例仅是本发明的部分实施例,而不是全部。基于本发明中的实 施例,本领域普通技术人员在没有做出创造性劳动前提下所获得的所有其他实施例,都属 于本发明保护的范围。
实施例1:CMV抗原表位四聚体的构建和效果检测
一、CMV抗原表位四聚体的构建
1)表达序列优化的HLA-A*0201(其氨基酸序列如SEQ ID NO:30所示,核苷酸序 列如SEQ ID NO:31所示)的α链和β2m链(其氨基酸序列如SEQ ID NO:32所示, 核苷酸序列如SEQ ID NO:33所示)。其中,α链的结构为相应HLA型α链的胞外区序 列链接了Avi-tag序列,以BamHI酶切位点间隔,以提供生物素化的位点。β2m链去掉了 信号肽序列,在成熟肽序列前面加了两个氨基酸(M和A)。表达载体为PET28a+,表达 菌为transetta或BL21。IPTG浓度为0.5mM,诱导表达4h。提取α链和β2m链蛋白包涵 体。
2)CMV抗原表位(抗原肽)的选择:HLA-A*0201型对应抗原表位NLVPMVATV (SEQ IDNO:16)。
3)pMHC I单体折叠及纯化:将步骤2)所述抗原肽、相应的步骤1)所述β2m链复 性蛋白和α链蛋白以摩尔比40:2:1的比例按顺序加入还原体系中,折叠反应72h。将得到 的产物过Superdex75 10/300GL柱子纯化。收集纯化产物用avidity试剂盒生物素化,再次 纯化得到生物素化的单体,凝胶电泳检测单体纯度。
4)将步骤3)所述生物素化的单体与APC标记的链霉亲和素结合反应,得到对应的四聚体,命名A0201–NLVPMVATV-四聚体(简称A0201–NLV四聚体)。
二、CMV抗原表位四聚体效果检测
1.分离人外周血单核细胞(PBMC)或准备TCR-T细胞,制备细胞悬液,细胞密度 为1×106细胞/mL。
2.将细胞离心,3000rpm,5min。去除上清,用50μL含有1%血清的PBS重悬。
3.加入2μL四聚体室温孵育30min。
4.加入2μL CD8抗体,冰上孵育20min。
5.加入1mL PBS,3000rpm,5min。
6.去除上清,加入1mL PBS,3000rpm,5min。
7.去除上清,用500μL的4%多聚甲醛重悬细胞,用滤膜过滤细胞悬液。
8.用流式细胞仪检测阳性细胞。
三、实验结果
单体的SDS PAGE检测结果见图1。如图1所示,经复性折叠和HPLC纯化,得到的 单体清晰地显示重链(C端连接Avi-tag序列的α链胞外区)和轻链(去除信号肽区的β2m 链)对应大小的蛋白,且纯度较高。
将构建好的四聚体分别与对应HLA型PBMC细胞共孵育钓取特异性T细胞。如图2 所示,自主研发四聚体较商业化四聚体(MBL公司)在钓取特异性T细胞方面具有明显 优势。具体的,自主研发A0201-NLV四聚体分选特异性T细胞是商业化四聚体(MBL, TS-0010-2C)的两倍。且不同批次分选效率相似,显示了极高的分选效率和稳定性。用此 A0201-NLV四聚体钓取到了C22-TCR和C29-TCR。
实施例2:pHAGE-TCR-RFP载体的构建
一、获取CMV pp65表位特异的TCR的β和α基因片段
1)用从MBL公司购买的或本公司自制的HLA-A*0201-NLVPMVATV-四聚体,根据 产品说明书与外周血染色,将四聚体染色阳性的T细胞进行流式单细胞分选,反转录获 得cDNA(IV Reverse Transcriptase,Invitrogen)。根据多重PCR(MultiplexPCR) 原理,通过两轮PCR(KOD-Plus-Neo,TOYOBO)扩增出TCRβ基因的可变区片段,。
反转录引物为:TRBC1-TCAGGCAGTATCTGGAGTCATTG(SEQ ID NO:136)
PCR扩增引物为:
上游引物1:T细胞受体β可变区--TRBV_F1(SEQ ID NO:56至95,其中SEQ ID NO:92、66、80也可以单独分别作为制备C2、C22、C29的β基因的可变区片段的TRBV_F1)
上游引物2:T细胞受体β可变区--TRBV_F2(SEQ ID NO:96至135,其中SEQ ID NO:133、106、120也可以单独分别作为制备C2、C22、C29的β基因的可变区片段的TRBV_F2)
下游引物1:T细胞受体β恒定区--TRBC2-GCACCTCCTTCCCATTCACC(SEQ ID NO:137)
下游引物2:T细胞受体β恒定区-TRBC3-GCTTCTGATGGCTCAAACACAG(SEQ ID NO:138)
具体的,根据PCR聚合酶KOD-Plus-Neo产品说明书,一轮PCR体系为20μL,退火 温度为60℃,反应30个循环。取第一轮PCR反应的产物1μL,作为二轮PCR的模板, 二轮PCR体系为30μL,退火温度为60℃,反应30个循环。然后将第二轮PCR产物跑琼 脂糖胶,将对应大小的条带进行切胶回收(天根胶回收试剂盒),送测序,测序引物为下 游引物2。得到TCRβ基因序列。C2、C22、C29的β链核苷酸序列如SEQ ID NO:48-50 所示。
2)如上,对四聚体染色阳性T细胞的进行反转录获得cDNA(IVReverse Transcriptase,Invitrogen)。根据产品说明书通过两轮PCR(KOD-Plus-Neo,TOYOBO)扩 增出TCRα基因片段。
反转录引物为:T细胞受体α恒定区-TRAC1-CGACCAGCTTGACATCACAG(SEQ ID NO:139)
PCR扩增引物为:
上游引物3:T细胞受体α可变区-TRAV_F1(SEQ ID NO:142至186,其中SEQ ID NO:172、172、159也可以单独分别作为制备C2、C22、C29的α基因的可变区片段的 TRAV_F1)
上游引物4:T细胞受体α可变区-TRAV_F2(SEQ ID NO:29、187至227,其中SEQ IDNO:215、215、202也可以单独分别作为制备C2、C22、C29的α基因的可变区片段 的TRAV_F2)
下游引物3:T细胞受体α恒定区-TRAC2-GTTGCTCTTGAAGTCCATAGACCTC(SEQ ID NO:140)
下游引物4:T细胞受体α恒定区-TRAC3-CAGGGTCAGGGTTCTGGATA(SEQ ID NO:141)
具体的,根据PCR聚合酶KOD-Plus-Neo产品说明书,一轮PCR体系为20μL,退火 温度为60℃,反应30个循环。取第一轮PCR反应的产物1μL,作为二轮PCR的模板, 二轮PCR体系为30μL,退火温度为60℃,反应30个循环。然后将第二轮PCR产物跑琼 脂糖胶,将对应大小的条带进行切胶回收(天根胶回收试剂盒),送测序,测序引物为下 游引物4。得到TCRα基因序列。具体的,C2、C22、C29的α链核苷酸序列如SEQ ID NO: 51-53所示。
二、pHAGE-TCR载体的构建
将TCRβ,fp2A,TCRα经长引物(包含fp2A序列)overlap-PCR扩增(KOD-Plus-Neo,TOYOBO)得到TCRβ-fp2A-TCRα片段,分别命名为C2、C22、C29。
扩增的引物包含上游引物5(C2、C22、C29分别对应SEQ ID NO:17-19)、下游引 物5(SEQ ID NO:20),上游引物6(C2、C22、C29分别对应SEQ ID NO:21-23)、下 游引物6(SEQ IDNO:24)。
具体的,先用引物5和引物6分别将TCRβ和TCRα扩增出来,PCR体系为50μL, 退火温度为60℃,反应30个循环。将PCR产物跑胶回收(天根胶回收试剂盒),分别取 回收产物1μL作为模板,用上游引物5和下游引物6进行overlap PCR,PCR体系为50μL, 退火温度为60℃,反应30个循环。跑琼脂糖胶得到约1800bp的条带,进行切胶回收。
将慢病毒载体pHAGE-IRES-RFP用NotI和NheI进行双酶切。酶切体系为40μL,其 中含NotI和NheI分别1.5μL,质粒2-3μg,37℃酶切6h,然后在体系里加入1μL碱性磷 酸酶(NEB),处理1h以减少质粒自连,将酶切后的质粒跑胶回收,用nanodrop测浓度, 作为骨架用于构建质粒。
根据Clone Express II One Step Cloning kit产品说明书,将TCR通过overlap与酶切后 线性化的pHAGE-IRES-RFP载体进行连接(见图3),转化至Stbl3菌株,在含氨苄的LB 平板上培养12-16h,挑取单克隆菌进行测序,测序引物选用pHAGE载体上的引物seq-pHAGE-F和seq-pHAGE-R,以及下游引物4。获得相应TCR,简称分别为C22(其 核苷酸序列如SEQ ID NO:37所示,其氨基酸序列如SEQ ID NO:36所示)、C29(其核 苷酸序列如SEQ IDNO:39所示,其氨基酸序列如SEQ ID NO:38所示)、C2(其核苷 酸序列如SEQ ID NO:41所示,其氨基酸序列如SEQ ID NO:40所示,其β链的氨基酸 序列如SEQ ID NO:42所示,其α链的氨基酸序列如SEQ ID NO:45所示)。
其中,制备的C2作为对照,其与C22的CDR区仅几个氨基酸不同,具体见表1。
表1
实施例3:pMHC四聚体染色法检测TCR的膜表达和亲和力
1、构建内源性TCR敲除的JurkatT细胞系
基于Jurkat细胞TCR的序列特征,在α链和β链的恒定区设计guide序列 (TRA_oligo1-CACCGTCTCTCAGCTGGTACACGGC(SEQ ID NO:25), TRA_oligo2-AAACGCCGTGTACCAGCTGAGAGAC(SEQ ID NO:26), TRB_oligo1-CACCGGGCTCAAACACAGCGACCTC(SEQ ID NO:27), TRB_oligo2-AAACGAGGTCGCTGTGTTTGAGCCC(SEQ ID NO:28))。
将合成的α链和β链的guide序列分别构建到sgRNA-LentiCRISPR-puro和 sgRNA-LentiCRISPR-BSD慢病毒载体中,与包装质粒psPAX2,pMD2.G和PEI转染试剂 按照一定的比例共转293T细胞,收取48h和72h的细胞培养上清,浓缩后的两个病毒同 时感染人类JurkatT细胞系。感染48h后用合适浓度的嘌呤霉素和杀稻瘟菌素进行药杀, 直至两个药物各自的对照组细胞全部死亡。存活的细胞用流式分选单细胞到96孔板中进 行培养。对于获得的单克隆细胞系,用TCRα链和β链的抗体分别对其表达进行鉴定,两 条链都缺陷的细胞株即为获得的内源性TCR敲除的Jurkat T细胞,命名为JC5。
2、构建稳定整合CMV TCR的JC5细胞系
将实施例2构建的上述C22、C29的pHAGE-TCR质粒分别与包装质粒psPAX2, pMD2.G以及转染试剂PEI按照一定的比例混合,转染293T细胞。收取48h和72h的细 胞培养上清,浓缩后感染对数生长期的JC5细胞(MOI=0.3)。感染3天后,用抗人CD3 和抗人TCRαβ流式抗体对细胞进行染色,将相同TCR表达水平的细胞分选后进行培养, 即为JC5-TCR细胞系。
3、TCR上膜情况及亲和力检测
取1×106JC5-TCR细胞,用Brilliant Violet 421TM anti-human TCRαβ(Biolegend)以 及相应的CMV pp65pMHC四聚体-APC(Tetramer-APC)染色,之后进行流式分析。
由图4、5可知,所制备的针对CMV pp65 HLA-A*A0201 NLVPMVATV的特异 C22-TCR、C29-TCR,均可以正确表达并且展示在细胞膜外,且与对应四聚体探针具有一 定的亲和力。其中,C2-TCR与C22-TCR相比,CDR区仅几个氨基酸不同,上膜情况(图 4)及与对应四聚体探针亲和力(图5)的效果差距却相当明显。
实施例4:人原代TCR-T细胞的构建及体外功能检测
1、人类原代T细胞的分离、培养和慢病毒感染
为了进一步验证所筛选到的TCR对CMV pp65抗原的识别和杀伤功能,采用淋巴细胞分离液Ficoll从志愿者的外周血中分离得到单个核细胞(PBMC),然后根据EasySepHuman T cell isolation kit(stem cell technologies)产品说明书,从PBMC中通过阴选的方法 获得T细胞,用含有100U/mL IL2的1640完全培养基将细胞重悬至1×106细胞/mL,培 养在抗CD3/CD28抗体包被的培养皿中进行激活。激活48h后,用慢病毒体系将装载有TCR的病毒颗粒(由实施例2制备)感染T细胞,感染方法为32℃下1500rpm离心2h, 取出后放在37℃细胞培养箱培养10h,然后通过添换培养基终止感染,并继续放在37℃ 细胞培养箱进行培养。感染三天后可用流式细胞仪将TCR阳性的细胞分选出来,获得 TCR-T细胞(包括上述C2、C22、C29)。
2.靶细胞的构建
用慢病毒体系将分别装载有pp65-PURO、HLA-A*0201-BSD以及荧光素酶-GFP的病毒颗粒感染对数生长期的Raji细胞。通过药物筛选及流式分选,得到同时稳定表达pp65、HLA-A*0201分子以及荧光素酶-GFP的Raji细胞,命名为Raji-A0201-pp65-荧光素酶。
3.TCR在人原代T细胞中的体外功能验证
将Raji/HLA-A0201/pp65以及未转pp65的Raji细胞(命名为Raji)与1G4-TCR-T、TCR-C2-T、TCR-C22-T,TCR-C29-T细胞按照1:1的数量比例进行共培养后,共培养 24小时后,分别收集细胞和上清,初步检测了TCR-C22-T和TCR-C29-T细胞的激活和 靶细胞的死亡情况。从胞外细胞因子TNFα、IL2和IFNγ(见图6)的释放水平来看, TCR-C22-T及TCR-C29-T同靶细胞共孵育后,与对照组1G4-TCR-T、TCR-C2-T相比, 可以显著地引起T细胞的激活。从靶细胞裂解后释放的荧光素酶量可反映出靶细胞死亡 情况(见图6),与CDR结构类似的TCR-C2-T相比,本申请实施例制备的TCR-C22-T 和TCR-C29-T具有显著的杀伤靶细胞的效果。实验结果证明,本发明实施例构建的 TCR-C22-T及TCR-C29-T细胞可以特异性地被CMVpp65抗原肽提呈细胞激活,且能够 显著地杀伤靶细胞。
同时,TCRα链和β链的CDR3高变区对TCR的功能影响最大。已有实验证实,CDR3 高变区即便高度相似,只相差一个氨基酸,其功能也会相差甚远。例如,将TCR E141的 α链和β链的CDR3区首位后的氨基酸依次突变成丙氨酸,构建在慢病毒载体上,制备相 应的TCR-T细胞。将其与靶细胞1:1共培养,检测荧光素酶量以及细胞因子TNFα、IL2 和IFNγ的分泌量。结果显示,未突变的E141遇到HLA matched-Raji后能有效的清除靶 细胞,并且特异的产生大量的细胞因子IL2。并且,在a3、a5、a6以及b6、b7、b8这些 位点,单个氨基酸的突变足以让TCR-T细胞彻底失活(具体参见专利CN202010373100.4)。 相应的,突变后的TCR,虽然仅一个氨基酸不同,但足以让TCR-T细胞彻底失活。由此 可见,虽然靶点相同,但仅有一个氨基酸不同就会使两个近乎相同的TCR产生完全不同 的技术效果,更不用说多个氨基酸不同。
实施例5:动物模型构建及CMV TCR-T体内功能检测
将3×105Raji-HLA-A*0201-pp65-荧光素酶肿瘤细胞通过尾静脉接种给5-6周的NOD/Scid IL-2Rγnull(NCG)雌鼠,构建淋巴瘤模型(见图7),记为第1天,第6天时, 将小鼠分为3个小组,分别为A:PBS注射组(注射等体积PBS);B:对照NC细胞注 射组(NC T细胞);C:CMV TCR-T注射组(C22-TCR-T细胞),B组小鼠尾静脉注射1×107 NC T细胞,C组小鼠尾静脉注射1×107TCR-T细胞,A组注射等体积200μL PBS。在接 下来的几周监测小鼠体内肿瘤细胞生长情况,T细胞体内增殖情况以及小鼠存活情况,如 图7-10所示,与对照组比较,本发明实施例构建的CMV特异性C22-TCR-T细胞能够显 著地杀伤小鼠体内肿瘤细胞(图7),提高小鼠的存活率(图8,图9),并且回输的TCR-T 细胞在体内进行了特异的增殖(图10)。
以上详细描述了本发明的优选实施方式,但是,本发明并不限于上述实施方式中的具 体细节,在本发明的技术构思范围内,可以对本发明的技术方案进行多种简单变型,这些 简单变型均属于本发明的保护范围。
另外需要说明的是,在上述具体实施方式中所描述的各个具体技术特征,在不矛盾的情况下,可以通过任何合适的方式进行组合,为了避免不必要的重复,本发明对 各种可能的组合方式不再另行说明。
序列表
<110> 华夏英泰(北京)生物技术有限公司、清华大学
<120> T细胞抗原受体、其多聚体复合物及其制备方法和应用
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<213> 人工序列(Artificial Sequence)
<400> 10
Phe Asn Asn Asn Val Pro
1 5
<210> 11
<211> 14
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 11
Ala Ser Ser Phe Ala Thr Gly Val Gly Ser Tyr Gly Tyr Thr
1 5 10
<210> 12
<211> 11
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 12
Ala Ser Ser Phe Ser Leu Pro Tyr Glu Gln Tyr
1 5 10
<210> 13
<211> 9
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 13
Ala Ser Pro Tyr Phe Asn Lys Phe Tyr
1 5
<210> 14
<211> 6
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 14
Ser Met Asn Val Glu Val
1 5
<210> 15
<211> 14
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 15
Ala Ser Ser Pro Thr Leu Gly Thr Gly Ala Glu Thr Gln Tyr
1 5 10
<210> 16
<211> 9
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 16
Asn Leu Val Pro Met Val Ala Thr Val
1 5
<210> 17
<211> 48
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 17
atttcaggtg tcgtgaagcg gccgcgccac catgggcccc cagctcct 48
<210> 18
<211> 48
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 18
atttcaggtg tcgtgaagcg gccgcgccac catgagcatc ggcctcct 48
<210> 19
<211> 48
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 19
atttcaggtg tcgtgaagcg gccgcgccac catggactcc tggacctt 48
<210> 20
<211> 81
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 20
tctccagcct gcttcagcag gctgaagtta gtagctccgc ttccgctccg tttccgccgg 60
aaatcctttc tcttgaccat g 81
<210> 21
<211> 71
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 21
tcagcctgct gaagcaggct ggagacgtgg aggagaaccc tggacctatg atggagaaga 60
atcctttggc a 71
<210> 22
<211> 71
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 22
tcagcctgct gaagcaggct ggagacgtgg aggagaaccc tggacctatg gagaagaatc 60
ctttggcagc c 71
<210> 23
<211> 71
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 23
tcagcctgct gaagcaggct ggagacgtgg aggagaaccc tggacctatg atgaaatcct 60
tgagagtttt a 71
<210> 24
<211> 45
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 24
agggatcctc tagactcgag ctagctcagc tggaccacag ccgca 45
<210> 25
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 25
caccgtctct cagctggtac acggc 25
<210> 26
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 26
aaacgccgtg taccagctga gagac 25
<210> 27
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 27
caccgggctc aaacacagcg acctc 25
<210> 28
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 28
aaacgaggtc gctgtgtttg agccc 25
<210> 29
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 29
gccacaataa acatacagga aaagc 25
<210> 30
<211> 294
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 30
Met Gly Ser His Ser Met Arg Tyr Phe Phe Thr Ser Val Ser Arg Pro
1 5 10 15
Gly Arg Gly Glu Pro Arg Phe Ile Ala Val Gly Tyr Val Asp Asp Thr
20 25 30
Gln Phe Val Arg Phe Asp Ser Asp Ala Ala Ser Gln Arg Met Glu Pro
35 40 45
Arg Ala Pro Trp Ile Glu Gln Glu Gly Pro Glu Tyr Trp Asp Gly Glu
50 55 60
Thr Arg Lys Val Lys Ala His Ser Gln Thr His Arg Val Asp Leu Gly
65 70 75 80
Thr Leu Arg Gly Tyr Tyr Asn Gln Ser Glu Ala Gly Ser His Thr Val
85 90 95
Gln Arg Met Tyr Gly Cys Asp Val Gly Ser Asp Trp Arg Phe Leu Arg
100 105 110
Gly Tyr His Gln Tyr Ala Tyr Asp Gly Lys Asp Tyr Ile Ala Leu Lys
115 120 125
Glu Asp Leu Arg Ser Trp Thr Ala Ala Asp Met Ala Ala Gln Thr Thr
130 135 140
Lys His Lys Trp Glu Ala Ala His Val Ala Glu Gln Leu Arg Ala Tyr
145 150 155 160
Leu Glu Gly Thr Cys Val Glu Trp Leu Arg Arg Tyr Leu Glu Asn Gly
165 170 175
Lys Glu Thr Leu Gln Arg Thr Asp Ala Pro Lys Thr His Met Thr His
180 185 190
His Ala Val Ser Asp His Glu Ala Thr Leu Arg Cys Trp Ala Leu Ser
195 200 205
Phe Tyr Pro Ala Glu Ile Thr Leu Thr Trp Gln Arg Asp Gly Glu Asp
210 215 220
Gln Thr Gln Asp Thr Glu Leu Val Glu Thr Arg Pro Ala Gly Asp Gly
225 230 235 240
Thr Phe Gln Lys Trp Val Ala Val Val Val Pro Ser Gly Gln Glu Gln
245 250 255
Arg Tyr Thr Cys His Val Gln His Glu Gly Leu Pro Lys Pro Leu Thr
260 265 270
Leu Arg Trp Glu Pro Gly Ser Gly Leu Asn Asp Ile Phe Glu Ala Gln
275 280 285
Lys Ile Glu Trp His Glu
290
<210> 31
<211> 885
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 31
atgggcagcc atagcatgcg ttacttcttc accagcgtta gtcgccccgg tcgcggtgaa 60
ccgcgcttta tcgcagtggg ttacgttgac gacacccagt ttgtgcgttt tgacagcgat 120
gccgcaagcc agcgtatgga accgcgtgcc ccgtggattg aacaagaagg cccggaatac 180
tgggatggcg aaacccgcaa agttaaagcc cacagccaga cccatcgcgt ggatctgggt 240
actttacgtg gctactataa ccagagcgaa gccggtagcc ataccgtgca gcgcatgtat 300
ggctgtgatg tgggcagtga ttggcgcttt ctgcgcggct atcatcagta cgcctatgac 360
ggcaaggact acatcgcttt aaaagaagat ttacgtagtt ggaccgccgc cgatatggca 420
gcccagacca ccaaacataa atgggaagcc gcccatgtgg cagagcagct gcgcgcatat 480
ctggaaggca catgcgtgga atggctgcgt cgctatctgg agaacggcaa agaaacttta 540
cagcgcaccg acgcaccgaa aacccatatg acccaccatg ccgtgagtga ccatgaagcc 600
actttacgct gttgggcttt aagcttctat ccggccgaaa ttactttaac ttggcagcgc 660
gatggtgaag accagaccca agataccgag ctggtggaaa cccgtccggc tggtgatggc 720
acctttcaga aatgggtggc cgttgtggtg ccgagcggtc aagaacagcg ctatacttgt 780
catgtgcagc atgagggctt accgaaaccg ctgactttac gctgggaacc gggatccggt 840
ttgaacgaca tcttcgaagc tcagaaaatc gaatggcacg aataa 885
<210> 32
<211> 101
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 32
Met Ala Ile Gln Arg Thr Pro Lys Ile Gln Val Tyr Ser Arg His Pro
1 5 10 15
Ala Glu Asn Gly Lys Ser Asn Phe Leu Asn Cys Tyr Val Ser Gly Phe
20 25 30
His Pro Ser Asp Ile Glu Val Asp Leu Leu Lys Asn Gly Glu Arg Ile
35 40 45
Glu Lys Val Glu His Ser Asp Leu Ser Phe Ser Lys Asp Trp Ser Phe
50 55 60
Tyr Leu Leu Tyr Tyr Thr Glu Phe Thr Pro Thr Glu Lys Asp Glu Tyr
65 70 75 80
Ala Cys Arg Val Asn His Val Thr Leu Ser Gln Pro Lys Ile Val Lys
85 90 95
Trp Asp Arg Asp Met
100
<210> 33
<211> 306
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 33
atggcaatcc agcgtactcc aaagattcag gtttactcac gtcatccagc agagaatgga 60
aagtcaaatt tcctgaattg ctatgtgtct gggtttcatc catccgacat tgaagttgac 120
ttactgaaga atggagagag aattgaaaaa gtggagcatt cagacttgtc tttcagcaag 180
gactggtctt tctatctctt gtactacact gaattcaccc ccactgaaaa agatgagtat 240
gcctgccgtg tgaaccatgt gactttgtca cagcccaaga tagttaagtg ggatcgagac 300
atgtaa 306
<210> 34
<211> 561
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 34
Met Glu Ser Arg Gly Arg Arg Cys Pro Glu Met Ile Ser Val Leu Gly
1 5 10 15
Pro Ile Ser Gly His Val Leu Lys Ala Val Phe Ser Arg Gly Asp Thr
20 25 30
Pro Val Leu Pro His Glu Thr Arg Leu Leu Gln Thr Gly Ile His Val
35 40 45
Arg Val Ser Gln Pro Ser Leu Ile Leu Val Ser Gln Tyr Thr Pro Asp
50 55 60
Ser Thr Pro Cys His Arg Gly Asp Asn Gln Leu Gln Val Gln His Thr
65 70 75 80
Tyr Phe Thr Gly Ser Glu Val Glu Asn Val Ser Val Asn Val His Asn
85 90 95
Pro Thr Gly Arg Ser Ile Cys Pro Ser Gln Glu Pro Met Ser Ile Tyr
100 105 110
Val Tyr Ala Leu Pro Leu Lys Met Leu Asn Ile Pro Ser Ile Asn Val
115 120 125
His His Tyr Pro Ser Ala Ala Glu Arg Lys His Arg His Leu Pro Val
130 135 140
Ala Asp Ala Val Ile His Ala Ser Gly Lys Gln Met Trp Gln Ala Arg
145 150 155 160
Leu Thr Val Ser Gly Leu Ala Trp Thr Arg Gln Gln Asn Gln Trp Lys
165 170 175
Glu Pro Asp Val Tyr Tyr Thr Ser Ala Phe Val Phe Pro Thr Lys Asp
180 185 190
Val Ala Leu Arg His Val Val Cys Ala His Glu Leu Val Cys Ser Met
195 200 205
Glu Asn Thr Arg Ala Thr Lys Met Gln Val Ile Gly Asp Gln Tyr Val
210 215 220
Lys Val Tyr Leu Glu Ser Phe Cys Glu Asp Val Pro Ser Gly Lys Leu
225 230 235 240
Phe Met His Val Thr Leu Gly Ser Asp Val Glu Glu Asp Leu Thr Met
245 250 255
Thr Arg Asn Pro Gln Pro Phe Met Arg Pro His Glu Arg Asn Gly Phe
260 265 270
Thr Val Leu Cys Pro Lys Asn Met Ile Ile Lys Pro Gly Lys Ile Ser
275 280 285
His Ile Met Leu Asp Val Ala Phe Thr Ser His Glu His Phe Gly Leu
290 295 300
Leu Cys Pro Lys Ser Ile Pro Gly Leu Ser Ile Ser Gly Asn Leu Leu
305 310 315 320
Met Asn Gly Gln Gln Ile Phe Leu Glu Val Gln Ala Ile Arg Glu Thr
325 330 335
Val Glu Leu Arg Gln Tyr Asp Pro Val Ala Ala Leu Phe Phe Phe Asp
340 345 350
Ile Asp Leu Leu Leu Gln Arg Gly Pro Gln Tyr Ser Glu His Pro Thr
355 360 365
Phe Thr Ser Gln Tyr Arg Ile Gln Gly Lys Leu Glu Tyr Arg His Thr
370 375 380
Trp Asp Arg His Asp Glu Gly Ala Ala Gln Gly Asp Asp Asp Val Trp
385 390 395 400
Thr Ser Gly Ser Asp Ser Asp Glu Glu Leu Val Thr Thr Glu Arg Lys
405 410 415
Thr Pro Arg Val Thr Gly Gly Gly Ala Met Ala Ser Ala Ser Thr Ser
420 425 430
Ala Gly Arg Lys Arg Lys Ser Ala Ser Ser Ala Thr Ala Cys Thr Ala
435 440 445
Gly Val Met Thr Arg Gly Arg Leu Lys Ala Glu Ser Thr Val Ala Pro
450 455 460
Glu Glu Asp Thr Asp Glu Asp Ser Asp Asn Glu Ile His Asn Pro Ala
465 470 475 480
Val Phe Thr Trp Pro Pro Trp Gln Ala Gly Ile Leu Ala Arg Asn Leu
485 490 495
Val Pro Met Val Ala Thr Val Gln Gly Gln Asn Leu Lys Tyr Gln Glu
500 505 510
Phe Phe Trp Asp Ala Asn Asp Ile Tyr Arg Ile Phe Ala Glu Leu Glu
515 520 525
Gly Val Trp Gln Pro Ala Ala Gln Pro Lys Arg Arg Arg His Arg Gln
530 535 540
Asp Ala Leu Pro Gly Pro Cys Ile Ala Ser Thr Pro Lys Lys His Arg
545 550 555 560
Gly
<210> 35
<211> 1686
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 35
atggagtcgc gcggtcgccg ttgtcccgaa atgatatccg tactgggtcc catttcgggg 60
cacgtgctga aagccgtgtt tagtcgcggc gatacgccgg tgctgccgca cgagacgcga 120
ctcctgcaga cgggtatcca cgtacgcgtg agccagccct cgctgatcct ggtgtcgcag 180
tacacgcccg actcgacgcc atgccaccgc ggcgacaatc agctgcaggt gcagcacacg 240
tactttacgg gcagcgaggt ggagaacgtg tcggtcaacg tgcacaaccc cacgggccga 300
agcatctgcc ccagccaaga gcccatgtcg atctatgtgt acgcgctgcc gctcaagatg 360
ctgaacatcc ccagcatcaa cgtgcaccac tacccgtcgg cggccgagcg caaacaccga 420
cacctgcccg tagccgacgc tgttattcac gcgtcgggca agcagatgtg gcaggcgcgt 480
ctcacggtct cgggactggc ctggacgcgt cagcagaacc agtggaaaga gcccgacgtc 540
tactacacgt cagcgttcgt gtttcccacc aaggacgtgg cactgcggca cgtggtgtgc 600
gcgcacgagc tggtttgctc catggagaac acgcgcgcaa ccaagatgca ggtgataggt 660
gaccagtacg tcaaggtgta cctggagtcc ttctgcgagg acgtgccctc cggcaagctc 720
tttatgcacg tcacgctggg ctctgacgtg gaagaggacc taacgatgac ccgcaacccg 780
caacccttca tgcgccccca cgagcgcaac ggctttacgg tgttgtgtcc caaaaatatg 840
ataatcaaac cgggcaagat ctcgcacatc atgctggatg tggcttttac ctcacacgag 900
cattttgggc tgctgtgtcc caagagcatc ccgggcctga gcatctcagg taacctgttg 960
atgaacgggc agcaaatctt cctggaggta caagcgatac gcgagaccgt ggaactgcgt 1020
cagtacgatc ccgtggctgc gctcttcttt ttcgatatcg acttgttgct gcagcgcggg 1080
cctcagtaca gcgagcaccc caccttcacc agccagtatc gcatccaggg caagcttgag 1140
taccgacaca cctgggaccg gcacgacgag ggtgccgccc agggcgacga cgacgtctgg 1200
accagcggat cggactccga cgaagaactc gtaaccaccg agcgtaagac gccccgcgtc 1260
accggcggcg gcgccatggc gagcgcctcc acttccgcgg gccgcaaacg caaatcagca 1320
tcctcggcga cggcgtgcac ggcgggcgtt atgacacgcg gccgccttaa ggccgagtcc 1380
accgtcgcgc ccgaagagga caccgacgag gattccgaca acgaaatcca caatccggcc 1440
gtgttcacct ggccgccctg gcaggccggc atcctggccc gcaacctggt gcccatggtg 1500
gctacggttc agggtcagaa tctgaagtac caggagttct tctgggacgc caacgacatc 1560
taccgcatct tcgccgaatt ggaaggcgta tggcagcccg ctgcgcaacc caaacgtcgc 1620
cgccaccggc aagacgcctt gcccgggcca tgcatcgcct cgacgcccaa aaagcaccga 1680
ggttga 1686
<210> 36
<211> 611
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 36
Met Ser Ile Gly Leu Leu Cys Cys Ala Ala Leu Ser Leu Leu Trp Ala
1 5 10 15
Gly Pro Val Asn Ala Gly Val Thr Gln Thr Pro Lys Phe Gln Val Leu
20 25 30
Lys Thr Gly Gln Ser Met Thr Leu Gln Cys Ala Gln Asp Met Asn His
35 40 45
Glu Tyr Met Ser Trp Tyr Arg Gln Asp Pro Gly Met Gly Leu Arg Leu
50 55 60
Ile His Tyr Ser Val Gly Ala Gly Ile Thr Asp Gln Gly Glu Val Pro
65 70 75 80
Asn Gly Tyr Asn Val Ser Arg Ser Thr Thr Glu Asp Phe Pro Leu Arg
85 90 95
Leu Leu Ser Ala Ala Pro Ser Gln Thr Ser Val Tyr Phe Cys Ala Ser
100 105 110
Ser Phe Ala Thr Gly Val Gly Ser Tyr Gly Tyr Thr Phe Gly Ser Gly
115 120 125
Thr Arg Leu Thr Val Val Glu Asp Leu Lys Asn Val Phe Pro Pro Glu
130 135 140
Val Ala Val Phe Glu Pro Ser Glu Ala Glu Ile Ser His Thr Gln Lys
145 150 155 160
Ala Thr Leu Val Cys Leu Ala Thr Gly Phe Phe Pro Asp His Val Glu
165 170 175
Leu Ser Trp Trp Val Asn Gly Lys Glu Val His Ser Gly Val Ser Thr
180 185 190
Asp Pro Gln Pro Leu Lys Glu Gln Pro Ala Leu Asn Asp Ser Arg Tyr
195 200 205
Cys Leu Ser Ser Arg Leu Arg Val Ser Ala Thr Phe Trp Gln Asn Pro
210 215 220
Arg Asn His Phe Arg Cys Gln Val Gln Phe Tyr Gly Leu Ser Glu Asn
225 230 235 240
Asp Glu Trp Thr Gln Asp Arg Ala Lys Pro Val Thr Gln Ile Val Ser
245 250 255
Ala Glu Ala Trp Gly Arg Ala Asp Cys Gly Phe Thr Ser Val Ser Tyr
260 265 270
Gln Gln Gly Val Leu Ser Ala Thr Ile Leu Tyr Glu Ile Leu Leu Gly
275 280 285
Lys Ala Thr Leu Tyr Ala Val Leu Val Ser Ala Leu Val Leu Met Ala
290 295 300
Met Val Lys Arg Lys Asp Phe Arg Arg Lys Arg Ser Gly Ser Gly Ala
305 310 315 320
Thr Asn Phe Ser Leu Leu Lys Gln Ala Gly Asp Val Glu Glu Asn Pro
325 330 335
Gly Pro Met Glu Lys Asn Pro Leu Ala Ala Pro Leu Leu Ile Leu Trp
340 345 350
Phe His Leu Asp Cys Val Ser Ser Ile Leu Asn Val Glu Gln Ser Pro
355 360 365
Gln Ser Leu His Val Gln Glu Gly Asp Ser Thr Asn Phe Thr Cys Ser
370 375 380
Phe Pro Ser Ser Asn Phe Tyr Ala Leu His Trp Tyr Arg Trp Glu Thr
385 390 395 400
Ala Lys Ser Pro Glu Ala Leu Phe Val Met Thr Leu Asn Gly Asp Glu
405 410 415
Lys Lys Lys Gly Arg Ile Ser Ala Thr Leu Asn Thr Lys Glu Gly Tyr
420 425 430
Ser Tyr Leu Tyr Ile Lys Gly Ser Gln Pro Glu Asp Ser Ala Thr Tyr
435 440 445
Leu Cys Ala Arg Asn Thr Gly Asn Gln Phe Tyr Phe Gly Thr Gly Thr
450 455 460
Ser Leu Thr Val Ile Pro Asn Ile Gln Asn Pro Asp Pro Ala Val Tyr
465 470 475 480
Gln Leu Arg Asp Ser Lys Ser Ser Asp Lys Ser Val Cys Leu Phe Thr
485 490 495
Asp Phe Asp Ser Gln Thr Asn Val Ser Gln Ser Lys Asp Ser Asp Val
500 505 510
Tyr Ile Thr Asp Lys Thr Val Leu Asp Met Arg Ser Met Asp Phe Lys
515 520 525
Ser Asn Ser Ala Val Ala Trp Ser Asn Lys Ser Asp Phe Ala Cys Ala
530 535 540
Asn Ala Phe Asn Asn Ser Ile Ile Pro Glu Asp Thr Phe Phe Pro Ser
545 550 555 560
Pro Glu Ser Ser Cys Asp Val Lys Leu Val Glu Lys Ser Phe Glu Thr
565 570 575
Asp Thr Asn Leu Asn Phe Gln Asn Leu Ser Val Ile Gly Phe Arg Ile
580 585 590
Leu Leu Leu Lys Val Ala Gly Phe Asn Leu Leu Met Thr Leu Arg Leu
595 600 605
Trp Ser Ser
610
<210> 37
<211> 1836
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 37
atgagcatcg gcctcctgtg ctgtgcagcc ttgtctctcc tgtgggcagg tccagtgaat 60
gctggtgtca ctcagacccc aaaattccag gtcctgaaga caggacagag catgacactg 120
cagtgtgccc aggatatgaa ccatgaatac atgtcctggt atcgacaaga cccaggcatg 180
gggctgaggc tgattcatta ctcagttggt gctggtatca ctgaccaagg agaagtcccc 240
aatggctaca atgtctccag atcaaccaca gaggatttcc cgctcaggct gctgtcggct 300
gctccctccc agacatctgt gtacttctgt gccagcagtt tcgcgacagg ggtggggagc 360
tatggctaca ccttcggttc ggggaccagg ttaaccgttg tagaggacct gaaaaacgtg 420
ttcccacccg aggtcgctgt gtttgagcca tcagaagcag agatctccca cacccaaaag 480
gccacactgg tgtgcctggc cacaggcttc ttccccgacc acgtggagct gagctggtgg 540
gtgaatggga aggaggtgca cagtggggtc agcacagacc cgcagcccct caaggagcag 600
cccgccctca atgactccag atactgcctg agcagccgcc tgagggtctc ggccaccttc 660
tggcagaacc cccgcaacca cttccgctgt caagtccagt tctacgggct ctcggagaat 720
gacgagtgga cccaggatag ggccaaaccc gtcacccaga tcgtcagcgc cgaggcctgg 780
ggtagagcag actgtggctt tacctcggtg tcctaccagc aaggggtcct gtctgccacc 840
atcctctatg agatcctgct agggaaggcc accctgtatg ctgtgctggt cagcgccctt 900
gtgttgatgg ccatggtcaa gagaaaggat ttccggcgga aacggagcgg aagcggagct 960
actaacttca gcctgctgaa gcaggctgga gacgtggagg agaaccctgg acctatggag 1020
aagaatcctt tggcagcccc attactaatc ctctggtttc atcttgactg cgtgagcagc 1080
atactgaacg tggaacaaag tcctcagtca ctgcatgttc aggagggaga cagcaccaat 1140
ttcacctgca gcttcccttc cagcaatttt tatgccttac actggtacag atgggaaact 1200
gcaaaaagcc ccgaggcctt gtttgtaatg actttaaatg gggatgaaaa gaagaaagga 1260
cgaataagtg ccactcttaa taccaaggag ggttacagct atttgtacat caaaggatcc 1320
cagcctgaag actcagccac atacctctgt gcccggaaca ccggtaacca gttctatttt 1380
gggacaggga caagtttgac ggtcattcca aatatccaga accctgaccc tgccgtgtac 1440
cagctgagag actctaaatc cagtgacaag tctgtctgcc tattcaccga ttttgattct 1500
caaacaaatg tgtcacaaag taaggattct gatgtgtata tcacagacaa aactgtgcta 1560
gacatgaggt ctatggactt caagagcaac agtgctgtgg cctggagcaa caaatctgac 1620
tttgcatgtg caaacgcctt caacaacagc attattccag aagacacctt cttccccagc 1680
ccagaaagtt cctgtgatgt caagctggtc gagaaaagct ttgaaacaga tacgaaccta 1740
aactttcaaa acctgtcagt gattgggttc cgaatcctcc tcctgaaagt ggccgggttt 1800
aatctgctca tgacgctgcg gctgtggtcc agctga 1836
<210> 38
<211> 609
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 38
Met Asp Ser Trp Thr Phe Cys Cys Val Ser Leu Cys Ile Leu Val Ala
1 5 10 15
Lys His Thr Asp Ala Gly Val Ile Gln Ser Pro Arg His Glu Val Thr
20 25 30
Glu Met Gly Gln Glu Val Thr Leu Arg Cys Lys Pro Ile Ser Gly His
35 40 45
Asp Tyr Leu Phe Trp Tyr Arg Gln Thr Met Met Arg Gly Leu Glu Leu
50 55 60
Leu Ile Tyr Phe Asn Asn Asn Val Pro Ile Asp Asp Ser Gly Met Pro
65 70 75 80
Glu Asp Arg Phe Ser Ala Lys Met Pro Asn Ala Ser Phe Ser Thr Leu
85 90 95
Lys Ile Gln Pro Ser Glu Pro Arg Asp Ser Ala Val Tyr Phe Cys Ala
100 105 110
Ser Ser Phe Ser Leu Pro Tyr Glu Gln Tyr Phe Gly Pro Gly Thr Arg
115 120 125
Leu Thr Val Thr Glu Asp Leu Lys Asn Val Phe Pro Pro Glu Val Ala
130 135 140
Val Phe Glu Pro Ser Glu Ala Glu Ile Ser His Thr Gln Lys Ala Thr
145 150 155 160
Leu Val Cys Leu Ala Thr Gly Phe Phe Pro Asp His Val Glu Leu Ser
165 170 175
Trp Trp Val Asn Gly Lys Glu Val His Ser Gly Val Ser Thr Asp Pro
180 185 190
Gln Pro Leu Lys Glu Gln Pro Ala Leu Asn Asp Ser Arg Tyr Cys Leu
195 200 205
Ser Ser Arg Leu Arg Val Ser Ala Thr Phe Trp Gln Asn Pro Arg Asn
210 215 220
His Phe Arg Cys Gln Val Gln Phe Tyr Gly Leu Ser Glu Asn Asp Glu
225 230 235 240
Trp Thr Gln Asp Arg Ala Lys Pro Val Thr Gln Ile Val Ser Ala Glu
245 250 255
Ala Trp Gly Arg Ala Asp Cys Gly Phe Thr Ser Val Ser Tyr Gln Gln
260 265 270
Gly Val Leu Ser Ala Thr Ile Leu Tyr Glu Ile Leu Leu Gly Lys Ala
275 280 285
Thr Leu Tyr Ala Val Leu Val Ser Ala Leu Val Leu Met Ala Met Val
290 295 300
Lys Arg Lys Asp Phe Arg Arg Lys Arg Ser Gly Ser Gly Ala Thr Asn
305 310 315 320
Phe Ser Leu Leu Lys Gln Ala Gly Asp Val Glu Glu Asn Pro Gly Pro
325 330 335
Met Met Lys Ser Leu Arg Val Leu Leu Val Ile Leu Trp Leu Gln Leu
340 345 350
Ser Trp Val Trp Ser Gln Gln Lys Glu Val Glu Gln Asp Pro Gly Pro
355 360 365
Leu Ser Val Pro Glu Gly Ala Ile Val Ser Leu Asn Cys Thr Tyr Ser
370 375 380
Asn Ser Ala Phe Gln Tyr Phe Met Trp Tyr Arg Gln Tyr Ser Arg Lys
385 390 395 400
Gly Pro Glu Leu Leu Met Tyr Thr Tyr Ser Ser Gly Asn Lys Glu Asp
405 410 415
Gly Arg Phe Thr Ala Gln Val Asp Lys Ser Ser Lys Tyr Ile Ser Leu
420 425 430
Phe Ile Arg Asp Ser Gln Pro Ser Asp Ser Ala Thr Tyr Leu Cys Ala
435 440 445
Thr Phe Leu Thr Gly Asn Gln Phe Tyr Phe Gly Thr Gly Thr Ser Leu
450 455 460
Thr Val Ile Pro Asn Ile Gln Asn Pro Asp Pro Ala Val Tyr Gln Leu
465 470 475 480
Arg Asp Ser Lys Ser Ser Asp Lys Ser Val Cys Leu Phe Thr Asp Phe
485 490 495
Asp Ser Gln Thr Asn Val Ser Gln Ser Lys Asp Ser Asp Val Tyr Ile
500 505 510
Thr Asp Lys Thr Val Leu Asp Met Arg Ser Met Asp Phe Lys Ser Asn
515 520 525
Ser Ala Val Ala Trp Ser Asn Lys Ser Asp Phe Ala Cys Ala Asn Ala
530 535 540
Phe Asn Asn Ser Ile Ile Pro Glu Asp Thr Phe Phe Pro Ser Pro Glu
545 550 555 560
Ser Ser Cys Asp Val Lys Leu Val Glu Lys Ser Phe Glu Thr Asp Thr
565 570 575
Asn Leu Asn Phe Gln Asn Leu Ser Val Ile Gly Phe Arg Ile Leu Leu
580 585 590
Leu Lys Val Ala Gly Phe Asn Leu Leu Met Thr Leu Arg Leu Trp Ser
595 600 605
Ser
<210> 39
<211> 1830
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 39
atggactcct ggaccttctg ctgtgtgtcc ctttgcatcc tggtagcaaa gcacacagat 60
gctggagtta tccagtcacc ccggcacgag gtgacagaga tgggacaaga agtgactctg 120
agatgtaaac caatttcagg acatgactac cttttctggt acagacagac catgatgcgg 180
ggactggagt tgctcattta ctttaacaac aacgttccga tagatgattc agggatgccc 240
gaggatcgat tctcagctaa gatgcctaat gcatcattct ccactctgaa gatccagccc 300
tcagaaccca gggactcagc tgtgtacttc tgtgccagca gttttagcct tccctacgag 360
cagtacttcg ggccgggcac caggctcacg gtcacagagg acctgaaaaa cgtgttccca 420
cccgaggtcg ctgtgtttga gccatcagaa gcagagatct cccacaccca aaaggccaca 480
ctggtgtgcc tggccacagg cttcttcccc gaccacgtgg agctgagctg gtgggtgaat 540
gggaaggagg tgcacagtgg ggtcagcaca gacccgcagc ccctcaagga gcagcccgcc 600
ctcaatgact ccagatactg cctgagcagc cgcctgaggg tctcggccac cttctggcag 660
aacccccgca accacttccg ctgtcaagtc cagttctacg ggctctcgga gaatgacgag 720
tggacccagg atagggccaa acccgtcacc cagatcgtca gcgccgaggc ctggggtaga 780
gcagactgtg gctttacctc ggtgtcctac cagcaagggg tcctgtctgc caccatcctc 840
tatgagatcc tgctagggaa ggccaccctg tatgctgtgc tggtcagcgc ccttgtgttg 900
atggccatgg tcaagagaaa ggatttccgg cggaaacgga gcggaagcgg agctactaac 960
ttcagcctgc tgaagcaggc tggagacgtg gaggagaacc ctggacctat gatgaaatcc 1020
ttgagagttt tactggtgat cctgtggctt cagttaagct gggtttggag ccaacagaag 1080
gaggtggagc aggatcctgg accactcagt gttccagagg gagccattgt ttctctcaac 1140
tgcacttaca gcaacagtgc ttttcaatac ttcatgtggt acagacagta ttccagaaaa 1200
ggccctgagt tgctgatgta cacatactcc agtggtaaca aagaagatgg aaggtttaca 1260
gcacaggtcg ataaatccag caagtatatc tccttgttca tcagagactc acagcccagt 1320
gattcagcca cctacctctg tgcaactttt ttaaccggta accagttcta ttttgggaca 1380
gggacaagtt tgacggtcat tccaaatatc cagaaccctg accctgccgt gtaccagctg 1440
agagactcta aatccagtga caagtctgtc tgcctattca ccgattttga ttctcaaaca 1500
aatgtgtcac aaagtaagga ttctgatgtg tatatcacag acaaaactgt gctagacatg 1560
aggtctatgg acttcaagag caacagtgct gtggcctgga gcaacaaatc tgactttgca 1620
tgtgcaaacg ccttcaacaa cagcattatt ccagaagaca ccttcttccc cagcccagaa 1680
agttcctgtg atgtcaagct ggtcgagaaa agctttgaaa cagatacgaa cctaaacttt 1740
caaaacctgt cagtgattgg gttccgaatc ctcctcctga aagtggccgg gtttaatctg 1800
ctcatgacgc tgcggctgtg gtccagctga 1830
<210> 40
<211> 612
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 40
Met Gly Pro Gln Leu Leu Gly Tyr Val Val Leu Cys Leu Leu Gly Ala
1 5 10 15
Gly Pro Leu Glu Ala Gln Val Thr Gln Asn Pro Arg Tyr Leu Ile Thr
20 25 30
Val Thr Gly Lys Lys Leu Thr Val Thr Cys Ser Gln Asn Met Asn His
35 40 45
Glu Tyr Met Ser Trp Tyr Arg Gln Asp Pro Gly Leu Gly Leu Arg Gln
50 55 60
Ile Tyr Tyr Ser Met Asn Val Glu Val Thr Asp Lys Gly Asp Val Pro
65 70 75 80
Glu Gly Tyr Lys Val Ser Arg Lys Glu Lys Arg Asn Phe Pro Leu Ile
85 90 95
Leu Glu Ser Pro Ser Pro Asn Gln Thr Ser Leu Tyr Phe Cys Ala Ser
100 105 110
Ser Pro Thr Leu Gly Thr Gly Ala Glu Thr Gln Tyr Phe Gly Pro Gly
115 120 125
Thr Arg Leu Leu Val Leu Glu Asp Leu Lys Asn Val Phe Pro Pro Glu
130 135 140
Val Ala Val Phe Glu Pro Ser Glu Ala Glu Ile Ser His Thr Gln Lys
145 150 155 160
Ala Thr Leu Val Cys Leu Ala Thr Gly Phe Phe Pro Asp His Val Glu
165 170 175
Leu Ser Trp Trp Val Asn Gly Lys Glu Val His Ser Gly Val Ser Thr
180 185 190
Asp Pro Gln Pro Leu Lys Glu Gln Pro Ala Leu Asn Asp Ser Arg Tyr
195 200 205
Cys Leu Ser Ser Arg Leu Arg Val Ser Ala Thr Phe Trp Gln Asn Pro
210 215 220
Arg Asn His Phe Arg Cys Gln Val Gln Phe Tyr Gly Leu Ser Glu Asn
225 230 235 240
Asp Glu Trp Thr Gln Asp Arg Ala Lys Pro Val Thr Gln Ile Val Ser
245 250 255
Ala Glu Ala Trp Gly Arg Ala Asp Cys Gly Phe Thr Ser Val Ser Tyr
260 265 270
Gln Gln Gly Val Leu Ser Ala Thr Ile Leu Tyr Glu Ile Leu Leu Gly
275 280 285
Lys Ala Thr Leu Tyr Ala Val Leu Val Ser Ala Leu Val Leu Met Ala
290 295 300
Met Val Lys Arg Lys Asp Phe Arg Arg Lys Arg Ser Gly Ser Gly Ala
305 310 315 320
Thr Asn Phe Ser Leu Leu Lys Gln Ala Gly Asp Val Glu Glu Asn Pro
325 330 335
Gly Pro Met Met Glu Lys Asn Pro Leu Ala Ala Pro Leu Leu Ile Leu
340 345 350
Trp Phe His Leu Asp Cys Val Ser Ser Ile Leu Asn Val Glu Gln Ser
355 360 365
Pro Gln Ser Leu His Val Gln Glu Gly Asp Ser Thr Asn Phe Thr Cys
370 375 380
Ser Phe Pro Ser Ser Asn Phe Tyr Ala Leu His Trp Tyr Arg Trp Glu
385 390 395 400
Thr Ala Lys Ser Pro Glu Ala Leu Phe Val Met Thr Leu Asn Gly Asp
405 410 415
Glu Lys Lys Lys Gly Arg Ile Ser Ala Thr Leu Asn Thr Lys Glu Gly
420 425 430
Tyr Ser Tyr Leu Tyr Ile Lys Gly Ser Gln Pro Glu Asp Ser Ala Thr
435 440 445
Tyr Leu Cys Ala Ser Pro Tyr Phe Asn Lys Phe Tyr Phe Gly Ser Gly
450 455 460
Thr Lys Leu Asn Val Lys Pro Asn Ile Gln Asn Pro Asp Pro Ala Val
465 470 475 480
Tyr Gln Leu Arg Asp Ser Lys Ser Ser Asp Lys Ser Val Cys Leu Phe
485 490 495
Thr Asp Phe Asp Ser Gln Thr Asn Val Ser Gln Ser Lys Asp Ser Asp
500 505 510
Val Tyr Ile Thr Asp Lys Thr Val Leu Asp Met Arg Ser Met Asp Phe
515 520 525
Lys Ser Asn Ser Ala Val Ala Trp Ser Asn Lys Ser Asp Phe Ala Cys
530 535 540
Ala Asn Ala Phe Asn Asn Ser Ile Ile Pro Glu Asp Thr Phe Phe Pro
545 550 555 560
Ser Pro Glu Ser Ser Cys Asp Val Lys Leu Val Glu Lys Ser Phe Glu
565 570 575
Thr Asp Thr Asn Leu Asn Phe Gln Asn Leu Ser Val Ile Gly Phe Arg
580 585 590
Ile Leu Leu Leu Lys Val Ala Gly Phe Asn Leu Leu Met Thr Leu Arg
595 600 605
Leu Trp Ser Ser
610
<210> 41
<211> 1839
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 41
atgggccccc agctccttgg ctatgtggtc ctttgccttc taggagcagg ccccctggaa 60
gcccaagtga cccagaaccc aagatacctc atcacagtga ctggaaagaa gttaacagtg 120
acttgttctc agaatatgaa ccatgagtat atgtcctggt atcgacaaga cccagggctg 180
ggcttaaggc agatctacta ttcaatgaat gttgaggtga ctgataaggg agatgttcct 240
gaagggtaca aagtctctcg aaaagagaag aggaatttcc ccctgatcct ggagtcgccc 300
agccccaacc agacctctct gtacttctgt gccagcagcc ctaccctagg gacaggggcc 360
gagacccagt acttcgggcc aggcacgcgg ctcctggtgc tcgaggacct gaaaaacgtg 420
ttcccacccg aggtcgctgt gtttgagcca tcagaagcag agatctccca cacccaaaag 480
gccacactgg tgtgcctggc cacaggcttc ttccccgacc acgtggagct gagctggtgg 540
gtgaatggga aggaggtgca cagtggggtc agcacagacc cgcagcccct caaggagcag 600
cccgccctca atgactccag atactgcctg agcagccgcc tgagggtctc ggccaccttc 660
tggcagaacc cccgcaacca cttccgctgt caagtccagt tctacgggct ctcggagaat 720
gacgagtgga cccaggatag ggccaaaccc gtcacccaga tcgtcagcgc cgaggcctgg 780
ggtagagcag actgtggctt tacctcggtg tcctaccagc aaggggtcct gtctgccacc 840
atcctctatg agatcctgct agggaaggcc accctgtatg ctgtgctggt cagcgccctt 900
gtgttgatgg ccatggtcaa gagaaaggat ttccggcgga aacggagcgg aagcggagct 960
actaacttca gcctgctgaa gcaggctgga gacgtggagg agaaccctgg acctatgatg 1020
gagaagaatc ctttggcagc cccattacta atcctctggt ttcatcttga ctgcgtgagc 1080
agcatactga acgtggaaca aagtcctcag tcactgcatg ttcaggaggg agacagcacc 1140
aatttcacct gcagcttccc ttccagcaat ttttatgcct tacactggta cagatgggaa 1200
actgcaaaaa gccccgaggc cttgtttgta atgactttaa atggggatga aaagaagaaa 1260
ggacgaataa gtgccactct taataccaag gagggttaca gctatttgta catcaaagga 1320
tcccagcctg aagactcagc cacatacctc tgtgcctcac cgtacttcaa caaattttac 1380
tttggatctg ggaccaaact caatgtaaaa ccaaatatcc agaaccctga ccctgccgtg 1440
taccagctga gagactctaa atccagtgac aagtctgtct gcctattcac cgattttgat 1500
tctcaaacaa atgtgtcaca aagtaaggat tctgatgtgt atatcacaga caaaactgtg 1560
ctagacatga ggtctatgga cttcaagagc aacagtgctg tggcctggag caacaaatct 1620
gactttgcat gtgcaaacgc cttcaacaac agcattattc cagaagacac cttcttcccc 1680
agcccagaaa gttcctgtga tgtcaagctg gtcgagaaaa gctttgaaac agatacgaac 1740
ctaaactttc aaaacctgtc agtgattggg ttccgaatcc tcctcctgaa agtggccggg 1800
tttaatctgc tcatgacgct gcggctgtgg tccagctga 1839
<210> 42
<211> 311
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 42
Met Gly Pro Gln Leu Leu Gly Tyr Val Val Leu Cys Leu Leu Gly Ala
1 5 10 15
Gly Pro Leu Glu Ala Gln Val Thr Gln Asn Pro Arg Tyr Leu Ile Thr
20 25 30
Val Thr Gly Lys Lys Leu Thr Val Thr Cys Ser Gln Asn Met Asn His
35 40 45
Glu Tyr Met Ser Trp Tyr Arg Gln Asp Pro Gly Leu Gly Leu Arg Gln
50 55 60
Ile Tyr Tyr Ser Met Asn Val Glu Val Thr Asp Lys Gly Asp Val Pro
65 70 75 80
Glu Gly Tyr Lys Val Ser Arg Lys Glu Lys Arg Asn Phe Pro Leu Ile
85 90 95
Leu Glu Ser Pro Ser Pro Asn Gln Thr Ser Leu Tyr Phe Cys Ala Ser
100 105 110
Ser Pro Thr Leu Gly Thr Gly Ala Glu Thr Gln Tyr Phe Gly Pro Gly
115 120 125
Thr Arg Leu Leu Val Leu Glu Asp Leu Lys Asn Val Phe Pro Pro Glu
130 135 140
Val Ala Val Phe Glu Pro Ser Glu Ala Glu Ile Ser His Thr Gln Lys
145 150 155 160
Ala Thr Leu Val Cys Leu Ala Thr Gly Phe Phe Pro Asp His Val Glu
165 170 175
Leu Ser Trp Trp Val Asn Gly Lys Glu Val His Ser Gly Val Ser Thr
180 185 190
Asp Pro Gln Pro Leu Lys Glu Gln Pro Ala Leu Asn Asp Ser Arg Tyr
195 200 205
Cys Leu Ser Ser Arg Leu Arg Val Ser Ala Thr Phe Trp Gln Asn Pro
210 215 220
Arg Asn His Phe Arg Cys Gln Val Gln Phe Tyr Gly Leu Ser Glu Asn
225 230 235 240
Asp Glu Trp Thr Gln Asp Arg Ala Lys Pro Val Thr Gln Ile Val Ser
245 250 255
Ala Glu Ala Trp Gly Arg Ala Asp Cys Gly Phe Thr Ser Val Ser Tyr
260 265 270
Gln Gln Gly Val Leu Ser Ala Thr Ile Leu Tyr Glu Ile Leu Leu Gly
275 280 285
Lys Ala Thr Leu Tyr Ala Val Leu Val Ser Ala Leu Val Leu Met Ala
290 295 300
Met Val Lys Arg Lys Asp Phe
305 310
<210> 43
<211> 311
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 43
Met Ser Ile Gly Leu Leu Cys Cys Ala Ala Leu Ser Leu Leu Trp Ala
1 5 10 15
Gly Pro Val Asn Ala Gly Val Thr Gln Thr Pro Lys Phe Gln Val Leu
20 25 30
Lys Thr Gly Gln Ser Met Thr Leu Gln Cys Ala Gln Asp Met Asn His
35 40 45
Glu Tyr Met Ser Trp Tyr Arg Gln Asp Pro Gly Met Gly Leu Arg Leu
50 55 60
Ile His Tyr Ser Val Gly Ala Gly Ile Thr Asp Gln Gly Glu Val Pro
65 70 75 80
Asn Gly Tyr Asn Val Ser Arg Ser Thr Thr Glu Asp Phe Pro Leu Arg
85 90 95
Leu Leu Ser Ala Ala Pro Ser Gln Thr Ser Val Tyr Phe Cys Ala Ser
100 105 110
Ser Phe Ala Thr Gly Val Gly Ser Tyr Gly Tyr Thr Phe Gly Ser Gly
115 120 125
Thr Arg Leu Thr Val Val Glu Asp Leu Lys Asn Val Phe Pro Pro Glu
130 135 140
Val Ala Val Phe Glu Pro Ser Glu Ala Glu Ile Ser His Thr Gln Lys
145 150 155 160
Ala Thr Leu Val Cys Leu Ala Thr Gly Phe Phe Pro Asp His Val Glu
165 170 175
Leu Ser Trp Trp Val Asn Gly Lys Glu Val His Ser Gly Val Ser Thr
180 185 190
Asp Pro Gln Pro Leu Lys Glu Gln Pro Ala Leu Asn Asp Ser Arg Tyr
195 200 205
Cys Leu Ser Ser Arg Leu Arg Val Ser Ala Thr Phe Trp Gln Asn Pro
210 215 220
Arg Asn His Phe Arg Cys Gln Val Gln Phe Tyr Gly Leu Ser Glu Asn
225 230 235 240
Asp Glu Trp Thr Gln Asp Arg Ala Lys Pro Val Thr Gln Ile Val Ser
245 250 255
Ala Glu Ala Trp Gly Arg Ala Asp Cys Gly Phe Thr Ser Val Ser Tyr
260 265 270
Gln Gln Gly Val Leu Ser Ala Thr Ile Leu Tyr Glu Ile Leu Leu Gly
275 280 285
Lys Ala Thr Leu Tyr Ala Val Leu Val Ser Ala Leu Val Leu Met Ala
290 295 300
Met Val Lys Arg Lys Asp Phe
305 310
<210> 44
<211> 309
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 44
Met Asp Ser Trp Thr Phe Cys Cys Val Ser Leu Cys Ile Leu Val Ala
1 5 10 15
Lys His Thr Asp Ala Gly Val Ile Gln Ser Pro Arg His Glu Val Thr
20 25 30
Glu Met Gly Gln Glu Val Thr Leu Arg Cys Lys Pro Ile Ser Gly His
35 40 45
Asp Tyr Leu Phe Trp Tyr Arg Gln Thr Met Met Arg Gly Leu Glu Leu
50 55 60
Leu Ile Tyr Phe Asn Asn Asn Val Pro Ile Asp Asp Ser Gly Met Pro
65 70 75 80
Glu Asp Arg Phe Ser Ala Lys Met Pro Asn Ala Ser Phe Ser Thr Leu
85 90 95
Lys Ile Gln Pro Ser Glu Pro Arg Asp Ser Ala Val Tyr Phe Cys Ala
100 105 110
Ser Ser Phe Ser Leu Pro Tyr Glu Gln Tyr Phe Gly Pro Gly Thr Arg
115 120 125
Leu Thr Val Thr Glu Asp Leu Lys Asn Val Phe Pro Pro Glu Val Ala
130 135 140
Val Phe Glu Pro Ser Glu Ala Glu Ile Ser His Thr Gln Lys Ala Thr
145 150 155 160
Leu Val Cys Leu Ala Thr Gly Phe Phe Pro Asp His Val Glu Leu Ser
165 170 175
Trp Trp Val Asn Gly Lys Glu Val His Ser Gly Val Ser Thr Asp Pro
180 185 190
Gln Pro Leu Lys Glu Gln Pro Ala Leu Asn Asp Ser Arg Tyr Cys Leu
195 200 205
Ser Ser Arg Leu Arg Val Ser Ala Thr Phe Trp Gln Asn Pro Arg Asn
210 215 220
His Phe Arg Cys Gln Val Gln Phe Tyr Gly Leu Ser Glu Asn Asp Glu
225 230 235 240
Trp Thr Gln Asp Arg Ala Lys Pro Val Thr Gln Ile Val Ser Ala Glu
245 250 255
Ala Trp Gly Arg Ala Asp Cys Gly Phe Thr Ser Val Ser Tyr Gln Gln
260 265 270
Gly Val Leu Ser Ala Thr Ile Leu Tyr Glu Ile Leu Leu Gly Lys Ala
275 280 285
Thr Leu Tyr Ala Val Leu Val Ser Ala Leu Val Leu Met Ala Met Val
290 295 300
Lys Arg Lys Asp Phe
305
<210> 45
<211> 274
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 45
Met Met Glu Lys Asn Pro Leu Ala Ala Pro Leu Leu Ile Leu Trp Phe
1 5 10 15
His Leu Asp Cys Val Ser Ser Ile Leu Asn Val Glu Gln Ser Pro Gln
20 25 30
Ser Leu His Val Gln Glu Gly Asp Ser Thr Asn Phe Thr Cys Ser Phe
35 40 45
Pro Ser Ser Asn Phe Tyr Ala Leu His Trp Tyr Arg Trp Glu Thr Ala
50 55 60
Lys Ser Pro Glu Ala Leu Phe Val Met Thr Leu Asn Gly Asp Glu Lys
65 70 75 80
Lys Lys Gly Arg Ile Ser Ala Thr Leu Asn Thr Lys Glu Gly Tyr Ser
85 90 95
Tyr Leu Tyr Ile Lys Gly Ser Gln Pro Glu Asp Ser Ala Thr Tyr Leu
100 105 110
Cys Ala Ser Pro Tyr Phe Asn Lys Phe Tyr Phe Gly Ser Gly Thr Lys
115 120 125
Leu Asn Val Lys Pro Asn Ile Gln Asn Pro Asp Pro Ala Val Tyr Gln
130 135 140
Leu Arg Asp Ser Lys Ser Ser Asp Lys Ser Val Cys Leu Phe Thr Asp
145 150 155 160
Phe Asp Ser Gln Thr Asn Val Ser Gln Ser Lys Asp Ser Asp Val Tyr
165 170 175
Ile Thr Asp Lys Thr Val Leu Asp Met Arg Ser Met Asp Phe Lys Ser
180 185 190
Asn Ser Ala Val Ala Trp Ser Asn Lys Ser Asp Phe Ala Cys Ala Asn
195 200 205
Ala Phe Asn Asn Ser Ile Ile Pro Glu Asp Thr Phe Phe Pro Ser Pro
210 215 220
Glu Ser Ser Cys Asp Val Lys Leu Val Glu Lys Ser Phe Glu Thr Asp
225 230 235 240
Thr Asn Leu Asn Phe Gln Asn Leu Ser Val Ile Gly Phe Arg Ile Leu
245 250 255
Leu Leu Lys Val Ala Gly Phe Asn Leu Leu Met Thr Leu Arg Leu Trp
260 265 270
Ser Ser
<210> 46
<211> 273
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 46
Met Glu Lys Asn Pro Leu Ala Ala Pro Leu Leu Ile Leu Trp Phe His
1 5 10 15
Leu Asp Cys Val Ser Ser Ile Leu Asn Val Glu Gln Ser Pro Gln Ser
20 25 30
Leu His Val Gln Glu Gly Asp Ser Thr Asn Phe Thr Cys Ser Phe Pro
35 40 45
Ser Ser Asn Phe Tyr Ala Leu His Trp Tyr Arg Trp Glu Thr Ala Lys
50 55 60
Ser Pro Glu Ala Leu Phe Val Met Thr Leu Asn Gly Asp Glu Lys Lys
65 70 75 80
Lys Gly Arg Ile Ser Ala Thr Leu Asn Thr Lys Glu Gly Tyr Ser Tyr
85 90 95
Leu Tyr Ile Lys Gly Ser Gln Pro Glu Asp Ser Ala Thr Tyr Leu Cys
100 105 110
Ala Arg Asn Thr Gly Asn Gln Phe Tyr Phe Gly Thr Gly Thr Ser Leu
115 120 125
Thr Val Ile Pro Asn Ile Gln Asn Pro Asp Pro Ala Val Tyr Gln Leu
130 135 140
Arg Asp Ser Lys Ser Ser Asp Lys Ser Val Cys Leu Phe Thr Asp Phe
145 150 155 160
Asp Ser Gln Thr Asn Val Ser Gln Ser Lys Asp Ser Asp Val Tyr Ile
165 170 175
Thr Asp Lys Thr Val Leu Asp Met Arg Ser Met Asp Phe Lys Ser Asn
180 185 190
Ser Ala Val Ala Trp Ser Asn Lys Ser Asp Phe Ala Cys Ala Asn Ala
195 200 205
Phe Asn Asn Ser Ile Ile Pro Glu Asp Thr Phe Phe Pro Ser Pro Glu
210 215 220
Ser Ser Cys Asp Val Lys Leu Val Glu Lys Ser Phe Glu Thr Asp Thr
225 230 235 240
Asn Leu Asn Phe Gln Asn Leu Ser Val Ile Gly Phe Arg Ile Leu Leu
245 250 255
Leu Lys Val Ala Gly Phe Asn Leu Leu Met Thr Leu Arg Leu Trp Ser
260 265 270
Ser
<210> 47
<211> 273
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 47
Met Met Lys Ser Leu Arg Val Leu Leu Val Ile Leu Trp Leu Gln Leu
1 5 10 15
Ser Trp Val Trp Ser Gln Gln Lys Glu Val Glu Gln Asp Pro Gly Pro
20 25 30
Leu Ser Val Pro Glu Gly Ala Ile Val Ser Leu Asn Cys Thr Tyr Ser
35 40 45
Asn Ser Ala Phe Gln Tyr Phe Met Trp Tyr Arg Gln Tyr Ser Arg Lys
50 55 60
Gly Pro Glu Leu Leu Met Tyr Thr Tyr Ser Ser Gly Asn Lys Glu Asp
65 70 75 80
Gly Arg Phe Thr Ala Gln Val Asp Lys Ser Ser Lys Tyr Ile Ser Leu
85 90 95
Phe Ile Arg Asp Ser Gln Pro Ser Asp Ser Ala Thr Tyr Leu Cys Ala
100 105 110
Thr Phe Leu Thr Gly Asn Gln Phe Tyr Phe Gly Thr Gly Thr Ser Leu
115 120 125
Thr Val Ile Pro Asn Ile Gln Asn Pro Asp Pro Ala Val Tyr Gln Leu
130 135 140
Arg Asp Ser Lys Ser Ser Asp Lys Ser Val Cys Leu Phe Thr Asp Phe
145 150 155 160
Asp Ser Gln Thr Asn Val Ser Gln Ser Lys Asp Ser Asp Val Tyr Ile
165 170 175
Thr Asp Lys Thr Val Leu Asp Met Arg Ser Met Asp Phe Lys Ser Asn
180 185 190
Ser Ala Val Ala Trp Ser Asn Lys Ser Asp Phe Ala Cys Ala Asn Ala
195 200 205
Phe Asn Asn Ser Ile Ile Pro Glu Asp Thr Phe Phe Pro Ser Pro Glu
210 215 220
Ser Ser Cys Asp Val Lys Leu Val Glu Lys Ser Phe Glu Thr Asp Thr
225 230 235 240
Asn Leu Asn Phe Gln Asn Leu Ser Val Ile Gly Phe Arg Ile Leu Leu
245 250 255
Leu Lys Val Ala Gly Phe Asn Leu Leu Met Thr Leu Arg Leu Trp Ser
260 265 270
Ser
<210> 48
<211> 933
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 48
atgggccccc agctccttgg ctatgtggtc ctttgccttc taggagcagg ccccctggaa 60
gcccaagtga cccagaaccc aagatacctc atcacagtga ctggaaagaa gttaacagtg 120
acttgttctc agaatatgaa ccatgagtat atgtcctggt atcgacaaga cccagggctg 180
ggcttaaggc agatctacta ttcaatgaat gttgaggtga ctgataaggg agatgttcct 240
gaagggtaca aagtctctcg aaaagagaag aggaatttcc ccctgatcct ggagtcgccc 300
agccccaacc agacctctct gtacttctgt gccagcagcc ctaccctagg gacaggggcc 360
gagacccagt acttcgggcc aggcacgcgg ctcctggtgc tcgaggacct gaaaaacgtg 420
ttcccacccg aggtcgctgt gtttgagcca tcagaagcag agatctccca cacccaaaag 480
gccacactgg tgtgcctggc cacaggcttc ttccccgacc acgtggagct gagctggtgg 540
gtgaatggga aggaggtgca cagtggggtc agcacagacc cgcagcccct caaggagcag 600
cccgccctca atgactccag atactgcctg agcagccgcc tgagggtctc ggccaccttc 660
tggcagaacc cccgcaacca cttccgctgt caagtccagt tctacgggct ctcggagaat 720
gacgagtgga cccaggatag ggccaaaccc gtcacccaga tcgtcagcgc cgaggcctgg 780
ggtagagcag actgtggctt tacctcggtg tcctaccagc aaggggtcct gtctgccacc 840
atcctctatg agatcctgct agggaaggcc accctgtatg ctgtgctggt cagcgccctt 900
gtgttgatgg ccatggtcaa gagaaaggat ttc 933
<210> 49
<211> 933
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 49
atgagcatcg gcctcctgtg ctgtgcagcc ttgtctctcc tgtgggcagg tccagtgaat 60
gctggtgtca ctcagacccc aaaattccag gtcctgaaga caggacagag catgacactg 120
cagtgtgccc aggatatgaa ccatgaatac atgtcctggt atcgacaaga cccaggcatg 180
gggctgaggc tgattcatta ctcagttggt gctggtatca ctgaccaagg agaagtcccc 240
aatggctaca atgtctccag atcaaccaca gaggatttcc cgctcaggct gctgtcggct 300
gctccctccc agacatctgt gtacttctgt gccagcagtt tcgcgacagg ggtggggagc 360
tatggctaca ccttcggttc ggggaccagg ttaaccgttg tagaggacct gaaaaacgtg 420
ttcccacccg aggtcgctgt gtttgagcca tcagaagcag agatctccca cacccaaaag 480
gccacactgg tgtgcctggc cacaggcttc ttccccgacc acgtggagct gagctggtgg 540
gtgaatggga aggaggtgca cagtggggtc agcacagacc cgcagcccct caaggagcag 600
cccgccctca atgactccag atactgcctg agcagccgcc tgagggtctc ggccaccttc 660
tggcagaacc cccgcaacca cttccgctgt caagtccagt tctacgggct ctcggagaat 720
gacgagtgga cccaggatag ggccaaaccc gtcacccaga tcgtcagcgc cgaggcctgg 780
ggtagagcag actgtggctt tacctcggtg tcctaccagc aaggggtcct gtctgccacc 840
atcctctatg agatcctgct agggaaggcc accctgtatg ctgtgctggt cagcgccctt 900
gtgttgatgg ccatggtcaa gagaaaggat ttc 933
<210> 50
<211> 927
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 50
atggactcct ggaccttctg ctgtgtgtcc ctttgcatcc tggtagcaaa gcacacagat 60
gctggagtta tccagtcacc ccggcacgag gtgacagaga tgggacaaga agtgactctg 120
agatgtaaac caatttcagg acatgactac cttttctggt acagacagac catgatgcgg 180
ggactggagt tgctcattta ctttaacaac aacgttccga tagatgattc agggatgccc 240
gaggatcgat tctcagctaa gatgcctaat gcatcattct ccactctgaa gatccagccc 300
tcagaaccca gggactcagc tgtgtacttc tgtgccagca gttttagcct tccctacgag 360
cagtacttcg ggccgggcac caggctcacg gtcacagagg acctgaaaaa cgtgttccca 420
cccgaggtcg ctgtgtttga gccatcagaa gcagagatct cccacaccca aaaggccaca 480
ctggtgtgcc tggccacagg cttcttcccc gaccacgtgg agctgagctg gtgggtgaat 540
gggaaggagg tgcacagtgg ggtcagcaca gacccgcagc ccctcaagga gcagcccgcc 600
ctcaatgact ccagatactg cctgagcagc cgcctgaggg tctcggccac cttctggcag 660
aacccccgca accacttccg ctgtcaagtc cagttctacg ggctctcgga gaatgacgag 720
tggacccagg atagggccaa acccgtcacc cagatcgtca gcgccgaggc ctggggtaga 780
gcagactgtg gctttacctc ggtgtcctac cagcaagggg tcctgtctgc caccatcctc 840
tatgagatcc tgctagggaa ggccaccctg tatgctgtgc tggtcagcgc ccttgtgttg 900
atggccatgg tcaagagaaa ggatttc 927
<210> 51
<211> 825
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 51
atgatggaga agaatccttt ggcagcccca ttactaatcc tctggtttca tcttgactgc 60
gtgagcagca tactgaacgt ggaacaaagt cctcagtcac tgcatgttca ggagggagac 120
agcaccaatt tcacctgcag cttcccttcc agcaattttt atgccttaca ctggtacaga 180
tgggaaactg caaaaagccc cgaggccttg tttgtaatga ctttaaatgg ggatgaaaag 240
aagaaaggac gaataagtgc cactcttaat accaaggagg gttacagcta tttgtacatc 300
aaaggatccc agcctgaaga ctcagccaca tacctctgtg cctcaccgta cttcaacaaa 360
ttttactttg gatctgggac caaactcaat gtaaaaccaa atatccagaa ccctgaccct 420
gccgtgtacc agctgagaga ctctaaatcc agtgacaagt ctgtctgcct attcaccgat 480
tttgattctc aaacaaatgt gtcacaaagt aaggattctg atgtgtatat cacagacaaa 540
actgtgctag acatgaggtc tatggacttc aagagcaaca gtgctgtggc ctggagcaac 600
aaatctgact ttgcatgtgc aaacgccttc aacaacagca ttattccaga agacaccttc 660
ttccccagcc cagaaagttc ctgtgatgtc aagctggtcg agaaaagctt tgaaacagat 720
acgaacctaa actttcaaaa cctgtcagtg attgggttcc gaatcctcct cctgaaagtg 780
gccgggttta atctgctcat gacgctgcgg ctgtggtcca gctga 825
<210> 52
<211> 822
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 52
atggagaaga atcctttggc agccccatta ctaatcctct ggtttcatct tgactgcgtg 60
agcagcatac tgaacgtgga acaaagtcct cagtcactgc atgttcagga gggagacagc 120
accaatttca cctgcagctt cccttccagc aatttttatg ccttacactg gtacagatgg 180
gaaactgcaa aaagccccga ggccttgttt gtaatgactt taaatgggga tgaaaagaag 240
aaaggacgaa taagtgccac tcttaatacc aaggagggtt acagctattt gtacatcaaa 300
ggatcccagc ctgaagactc agccacatac ctctgtgccc ggaacaccgg taaccagttc 360
tattttggga cagggacaag tttgacggtc attccaaata tccagaaccc tgaccctgcc 420
gtgtaccagc tgagagactc taaatccagt gacaagtctg tctgcctatt caccgatttt 480
gattctcaaa caaatgtgtc acaaagtaag gattctgatg tgtatatcac agacaaaact 540
gtgctagaca tgaggtctat ggacttcaag agcaacagtg ctgtggcctg gagcaacaaa 600
tctgactttg catgtgcaaa cgccttcaac aacagcatta ttccagaaga caccttcttc 660
cccagcccag aaagttcctg tgatgtcaag ctggtcgaga aaagctttga aacagatacg 720
aacctaaact ttcaaaacct gtcagtgatt gggttccgaa tcctcctcct gaaagtggcc 780
gggtttaatc tgctcatgac gctgcggctg tggtccagct ga 822
<210> 53
<211> 822
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 53
atgatgaaat ccttgagagt tttactggtg atcctgtggc ttcagttaag ctgggtttgg 60
agccaacaga aggaggtgga gcaggatcct ggaccactca gtgttccaga gggagccatt 120
gtttctctca actgcactta cagcaacagt gcttttcaat acttcatgtg gtacagacag 180
tattccagaa aaggccctga gttgctgatg tacacatact ccagtggtaa caaagaagat 240
ggaaggttta cagcacaggt cgataaatcc agcaagtata tctccttgtt catcagagac 300
tcacagccca gtgattcagc cacctacctc tgtgcaactt ttttaaccgg taaccagttc 360
tattttggga cagggacaag tttgacggtc attccaaata tccagaaccc tgaccctgcc 420
gtgtaccagc tgagagactc taaatccagt gacaagtctg tctgcctatt caccgatttt 480
gattctcaaa caaatgtgtc acaaagtaag gattctgatg tgtatatcac agacaaaact 540
gtgctagaca tgaggtctat ggacttcaag agcaacagtg ctgtggcctg gagcaacaaa 600
tctgactttg catgtgcaaa cgccttcaac aacagcatta ttccagaaga caccttcttc 660
cccagcccag aaagttcctg tgatgtcaag ctggtcgaga aaagctttga aacagatacg 720
aacctaaact ttcaaaacct gtcagtgatt gggttccgaa tcctcctcct gaaagtggcc 780
gggtttaatc tgctcatgac gctgcggctg tggtccagct ga 822
<210> 54
<211> 27
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 54
Arg Arg Lys Arg Ser Gly Ser Gly Ala Thr Asn Phe Ser Leu Leu Lys
1 5 10 15
Gln Ala Gly Asp Val Glu Glu Asn Pro Gly Pro
20 25
<210> 55
<211> 81
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 55
cggcggaaac ggagcggaag cggagctact aacttcagcc tgctgaagca ggctggagac 60
gtggaggaga accctggacc t 81
<210> 56
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 56
atggatacct ggctcgtatg c 21
<210> 57
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 57
atgggctgca ggctcctc 18
<210> 58
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 58
atgggctgca ggctgctc 18
<210> 59
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 59
atgggctcca ggctgctct 19
<210> 60
<211> 14
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 60
atgggccccg ggct 14
<210> 61
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 61
atgggccctg ggctcct 17
<210> 62
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 62
atgggaccca ggctcctct 19
<210> 63
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 63
atgagcatcg ggctcctgtg 20
<210> 64
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 64
atgagcctcg ggctcctg 18
<210> 65
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 65
atgagaatca ggctcctgtg c 21
<210> 66
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 66
atgagcatcg gcctcctgtg 20
<210> 67
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 67
atgagcatca gcctcctgtg 20
<210> 68
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 68
atgggcacaa ggctcctctg 20
<210> 69
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 69
atgggcacca ggctcctc 18
<210> 70
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 70
atgggcacca gtctcctatg c 21
<210> 71
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 71
atgggtacca gtctcctatg ctg 23
<210> 72
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 72
atgggcacca gcctcctc 18
<210> 73
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 73
atgggcttca ggctcctctg 20
<210> 74
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 74
atgggcacga ggctcttctt c 21
<210> 75
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 75
atgggcacca ggctcttctt c 21
<210> 76
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 76
atgggcacaa ggttgttctt c 21
<210> 77
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 77
atgagcacca ggcttctctg c 21
<210> 78
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 78
atgggtacca ggctcctctg 20
<210> 79
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 79
atggactcct ggaccttctg ct 22
<210> 80
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 80
atggactcct ggaccctctg 20
<210> 81
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 81
atggccacca ggctcctc 18
<210> 82
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 82
atgcttagtc ctgacctgcc tg 22
<210> 83
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 83
atggtttcca ggcttctcag tt 22
<210> 84
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 84
atgggtcctg ggcttctcc 19
<210> 85
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 85
atgagcccaa tattcacctg ca 22
<210> 86
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 86
atggatatct ggctcctctg ct 22
<210> 87
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 87
atggacacca gagtactctg ctg 23
<210> 88
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 88
atgagcaacc aggtgctctg 20
<210> 89
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 89
atgctgctgc ttctgctgc 19
<210> 90
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 90
atggcctccc tgctcttctt c 21
<210> 91
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 91
atgactatca ggctcctctg ctac 24
<210> 92
<211> 16
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 92
atgggccccc agctcc 16
<210> 93
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 93
atgggaatca ggctcctctg tc 22
<210> 94
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 94
atgctgagtc ttctgctcct tctc 24
<210> 95
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 95
atgctctgct ctctccttgc 20
<210> 96
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 96
tcactctgaa gatccggtcc ac 22
<210> 97
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 97
cttcacatca attccctgga gc 22
<210> 98
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 98
cacctgaatg ccccaacag 19
<210> 99
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 99
agtcgcttct cacctgaatg c 21
<210> 100
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 100
gagatgaatg tgagcacctt ggag 24
<210> 101
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 101
gagatgaatg tgagtgcctt ggag 24
<210> 102
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 102
gctctgagct gaatgtgaac gc 22
<210> 103
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 103
gaagtcccca atggctacaa tg 22
<210> 104
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 104
aggtccctga tggctacaat g 21
<210> 105
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 105
tggcgtctgc tgtaccctct 20
<210> 106
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 106
acagaggatt tcccgctcag 20
<210> 107
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 107
cccctcaagc tggagtcagc 20
<210> 108
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 108
acacagagga tttcccactc agg 23
<210> 109
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 109
ttctctgcac agaggtctga gg 22
<210> 110
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 110
gctgcccagt gatcgcttct 20
<210> 111
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 111
actctgaaga tccagcgcac ag 22
<210> 112
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 112
cgtctccact ctgaagatcc agc 23
<210> 113
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 113
ggcctgaggg atccatctcc 20
<210> 114
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 114
ggctgctcag tgatcggttc 20
<210> 115
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 115
gaacgattct ccgcacaaca gt 22
<210> 116
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 116
acaaaggaga agtctcagat ggct 24
<210> 117
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 117
ccctcactct ggagtcagct ac 22
<210> 118
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 118
cctcactctg gagtccgcta c 21
<210> 119
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 119
gcagagaggc tcaaaggagt agac 24
<210> 120
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 120
atccagccct cagaacccag 20
<210> 121
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 121
atcattctga actgaacatg agctc 25
<210> 122
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 122
tggagggacg tattctactc tgaag 25
<210> 123
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 123
gacacccctg ataacttcca atcc 24
<210> 124
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 124
tgcctcccaa attcaccctg 20
<210> 125
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 125
catcccgcag agccgag 17
<210> 126
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 126
gcaggtagtg cgaggagatt c 21
<210> 127
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 127
ctgtgacatc ggcccaaaag 20
<210> 128
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 128
acagtgacca gtgcccatcc 20
<210> 129
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 129
caatgcccca agaacgcac 19
<210> 130
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 130
tagagtctgc catccccaac c 21
<210> 131
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 131
gacggagcat tttcccctga c 21
<210> 132
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 132
gccagcacca accagacatc 20
<210> 133
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 133
ctgatcctgg agtcgcccag 20
<210> 134
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 134
atcagccgcc caaacctaac 20
<210> 135
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 135
cggcagttca tcctgagttc taag 24
<210> 136
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 136
tcaggcagta tctggagtca ttg 23
<210> 137
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 137
gcacctcctt cccattcacc 20
<210> 138
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 138
gcttctgatg gctcaaacac ag 22
<210> 139
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 139
cgaccagctt gacatcacag 20
<210> 140
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 140
gttgctcttg aagtccatag acctc 25
<210> 141
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 141
cagggtcagg gttctggata 20
<210> 142
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 142
atgtggggag ctttccttct ct 22
<210> 143
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 143
atgtggggag ttttccttct ttatg 25
<210> 144
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 144
atggctttgc agagcactct g 21
<210> 145
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 145
atggcctctg cacccatctc 20
<210> 146
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 146
atgaggcaag tggcgagagt 20
<210> 147
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 147
atgaagacat ttgctggatt ttcg 24
<210> 148
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 148
atggagtcat tcctgggagg tg 22
<210> 149
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 149
atggagaaga tgcggagacc tg 22
<210> 150
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 150
atgctcctgt tgctcatacc ag 22
<210> 151
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 151
atgctcctgc tgctcgtcc 19
<210> 152
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 152
atgctcctgg agcttatccc a 21
<210> 153
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 153
atgctcttag tggtcattct gctg 24
<210> 154
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 154
atgaattctt ctccaggacc agc 23
<210> 155
<211> 28
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 155
atgaactatt ctccaggctt agtatctc 28
<210> 156
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 156
atgaaaaagc atctgacgac cttc 24
<210> 157
<211> 27
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 157
atgatatcct tgagagtttt actggtg 27
<210> 158
<211> 30
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 158
atgaaatcct tgagagtttt actagtgatc 30
<210> 159
<211> 28
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 159
atgatgaaat ccttgagagt tttactgg 28
<210> 160
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 160
atgacatcca ttcgagctgt atttat 26
<210> 161
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 161
atggcaggca ttcgagctt 19
<210> 162
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 162
atgtcacttt ctagcctgct gaag 24
<210> 163
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 163
atgaagccca ccctcatctc ag 22
<210> 164
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 164
atggaaactc tcctgggagt gtc 23
<210> 165
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 165
atgctgtctg cttcctgctc ag 22
<210> 166
<211> 19
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 166
atgctgactg ccagcctgt 19
<210> 167
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 167
atggagaaaa tgttggagtg tgc 23
<210> 168
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 168
atggagaccc tcttgggcct 20
<210> 169
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 169
atgaagagga tattgggagc tctg 24
<210> 170
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 170
atggacaaga tcttaggagc atcat 25
<210> 171
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 171
atgctgttct ccagcctgct 20
<210> 172
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 172
atggagaaga atcctttggc ag 22
<210> 173
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 173
atgctactca tcacatcaat gttgg 25
<210> 174
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 174
atgaggctgg tggcaagagt aac 23
<210> 175
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 175
atgaagttgg tgacaagcat tactg 25
<210> 176
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 176
atggtcctga aattctccgt gtc 23
<210> 177
<211> 17
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 177
atggccatgc tcctggg 17
<210> 178
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 178
atggagactc tcctgaaagt gct 23
<210> 179
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 179
atggagactg ttctgcaagt actcc 25
<210> 180
<211> 28
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 180
atgctccttg aacatttatt aataatct 28
<210> 181
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 181
atgatgaagt gtccacaggc tttac 25
<210> 182
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 182
atgacacgag ttagcttgct gtg 23
<210> 183
<211> 18
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 183
atggcatgcc ctggcttc 18
<210> 184
<211> 27
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 184
atgaagaagc tactagcaat gattctg 27
<210> 185
<211> 28
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 185
atgaactcct ctctggactt tctaattc 28
<210> 186
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 186
atggtgaaga tccggcaatt 20
<210> 187
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 187
tccttagtcg ctctgatagt tatggt 26
<210> 188
<211> 27
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 188
tttcttcatt ccttagtcgg tctaaag 27
<210> 189
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 189
ctcttcatcg ctgctcatcc tc 22
<210> 190
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 190
gccaaacctc cttccacctg 20
<210> 191
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 191
cctgccgaca gaaagtccag 20
<210> 192
<211> 20
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 192
cgcattgcag acacccagac 20
<210> 193
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 193
ttgataccac ccttaaacag agtttg 26
<210> 194
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 194
attactgaag aatggaagca gcttg 25
<210> 195
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 195
tcctttaatc tgaggaaacc ctctg 25
<210> 196
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 196
gtgaaacctc cttccacctg ac 22
<210> 197
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 197
aggctttgag gctgaattta agag 24
<210> 198
<211> 27
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 198
ggctgaattt aacaagagtc aaacttc 27
<210> 199
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 199
gagcgaaacc tccttctacc tg 22
<210> 200
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 200
aaaccacttc tttccacttg gag 23
<210> 201
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 201
caaagcaaag ctctctgcac atc 23
<210> 202
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 202
cagtgattca gccacctacc tc 22
<210> 203
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 203
cgaattgctg ttacattgaa caagac 26
<210> 204
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 204
ttgcagctac tcaacctgga gac 23
<210> 205
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 205
atccgccaac cttgtcatct c 21
<210> 206
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 206
aaggcgagac atctttccac c 21
<210> 207
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 207
agattaagag tcacgcttga cacttc 26
<210> 208
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 208
gccagtccta tcaagagtga cagttc 26
<210> 209
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 209
caacttcacc atcacagcct cac 23
<210> 210
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 210
cacatcacag cccctaaacc tg 22
<210> 211
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 211
tggaagactt aatgcctcgc tg 22
<210> 212
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 212
tgtcgctacg gaacgctaca g 21
<210> 213
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 213
cttcaataaa agtgccaagc agttc 25
<210> 214
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 214
aaaaagtggt cgctattctg tcaac 25
<210> 215
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 215
gaagaaagga cgaataagtg ccac 24
<210> 216
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 216
acatcacagc cacccagact ac 22
<210> 217
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 217
cacagaagac agaaagtcca gcac 24
<210> 218
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 218
gcaatcgctg aagacagaaa gtc 23
<210> 219
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 219
gaaaggacag ttctctccac atcac 25
<210> 220
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 220
aaaagtgcca agcacctctc tc 22
<210> 221
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 221
gcaaagctcc ctgtacctta cg 22
<210> 222
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 222
aaagataact gccaagttgg atgag 25
<210> 223
<211> 26
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 223
cttcctgaat atctcagcat ccatac 26
<210> 224
<211> 22
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 224
gcatcctgaa catcacagcc ac 22
<210> 225
<211> 25
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 225
gaatcgtttc tctgtgaact tccag 25
<210> 226
<211> 21
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 226
tctcagcacc ctccacatca c 21
<210> 227
<211> 24
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 227
caggtatcag actcagccgt gtac 24
Claims (12)
1.一种T细胞抗原受体,其特征在于,所述的T细胞抗原受体特异性结合CMV pp65,所述的T细胞抗原受体包含α链的CDR1α-CDR3α和β链的CDR1β-CDR3β,其中,CDR1α的氨基酸序列包含SEQ ID NO:1-2中的任一种或包含与SEQ ID NO:1-2中任一种所示氨基酸序列具有至少80%同源性,CDR2α的氨基酸序列包含SEQ ID NO:3-4中的任一种或包含与SEQ ID NO:3-4中任一种所示氨基酸序列具有至少80%同源性,CDR3α的氨基酸序列包含SEQ ID NO:5-6中的任一种或包含与SEQ ID NO:5-6中任一种所示氨基酸序列具有至少80%同源性,CDR1β的氨基酸序列包含SEQ ID NO:7-8中的任一种或包含与SEQ ID NO:7-8中任一种所示氨基酸序列具有至少80%同源性,CDR2β的氨基酸序列包含SEQ ID NO:9-10中的任一种或包含与SEQ ID NO:9-10中任一种所示氨基酸序列具有至少80%同源性,CDR3β的氨基酸序列包含SEQ ID NO:11-12中的任一种或包含与SEQ ID NO:11-12中任一种所示氨基酸序列具有至少80%同源性;
优选的,所述的CDR1α-CDR3α及CDR1β-CDR3β包含下列任一组:
2.根据权利要求1所述的T细胞抗原受体,其特征在于,其β链的氨基酸序列包含SEQ IDNO:43-44中的任一种或包含与SEQ ID NO:43-44任一种所示氨基酸序列具有至少80%同源性,其α链的氨基酸序列包含SEQ ID NO:46-47中的任一种或包含与SEQ ID NO:46-47任一种所示氨基酸序列具有至少80%同源性,
优选的,α链与β链的氨基酸直接连接或间接连接,优选为间接连接,更优选通过fp2A连接,
进一步优选的,所述的T细胞抗原受体的氨基酸序列包含SEQ ID NO:36或38或包含与SEQ ID NO:36或38所示氨基酸序列具有至少80%同源性。
3.一种编码权利要求1-2任一所述的T细胞抗原受体的核酸。
4.一种表达载体,其特征在于,所述的表达载体包含权利要求3所述的核酸。
5.一种宿主细胞或免疫细胞,其特征在于,所述的宿主细胞包含权利要求3所述的核酸或权利要求4所述的表达载体;所述的免疫细胞表达权利要求1-2任一所述的T细胞抗原受体,优选的,所述的免疫细胞选自T细胞或干细胞,进一步优选的,所述的免疫细胞分离自受试者的T细胞衍生。
6.一种重组T细胞的制备方法,其特征在于,包括如下步骤:
1)从阳性T细胞克隆得到权利要求3所述的核酸;
2)分离、培养原代T细胞;
3)将步骤1)得到的核酸递送至步骤2)所述的原代T细胞中,获得表达权利要求1-2任一所述T细胞抗原受体的重组T细胞;
优选的,所述的制备方法还包括敲除细胞内源性TCR的步骤。
7.一种T细胞抗原受体的制备方法,其特征在于,包括如下步骤:
(1)从阳性T细胞克隆得到权利要求3所述的核酸;
(2)将步骤(1)得到的核酸连接至载体骨架,获得表达载体;
(3)将步骤(2)获得的表达载体转化至宿主细胞,然后诱导其表达;
(4)获得T细胞抗原受体。
8.一种多聚体复合物,其特征在于,所述多聚体复合物包含权利要求1-2任一所述的T细胞抗原受体。
9.根据权利要求8所述的多聚体复合物,其特征在于,所述多聚体复合物还包括单体,生物素分子,以及链霉亲和素分子或亲和素分子,其中,所述单体包括MHC分子α链胞外区和β2m链及抗原肽,所述单体与所述生物素分子偶联,所述生物素分子与所述链霉亲和素或亲和素分子结合;
优选的,所述的抗原肽包含SEQ ID NO:16;
优选的,所述的MHC分子为HLA-A*0201。
10.一种权利要求8-9任一所述的多聚体复合物的制备方法,其特征在于,包括如下步骤:
I)表达和纯化在C端连接avi-tag序列的MHC分子α链胞外区和β2m链;
II)将抗原肽、步骤I)获得的β2m链和在C端连接avi-tag序列的MHC分子α链胞外区复性折叠,制备单体;
III)将步骤II)制备的单体生物素化,获得生物素化的单体;
IV)将步骤III)获得的生物素化的单体与带荧光标记的链霉亲和素或亲和素反应,制备抗原肽-MHC分子四聚体;
V)将步骤IV)得到的抗原肽-MHC分子四聚体与T细胞共孵育,形成T细胞抗原受体与抗原肽-MHC分子四聚体复合物。
11.权利要求1-2任一所述的T细胞抗原受体、权利要求3所述的核酸、权利要求4所述的表达载体、权利要求5所述的宿主细胞、权利要求5所述的免疫细胞、权利要求8-9任一所述的多聚体复合物在制备诊断或治疗肿瘤或与CMV相关疾病的产品中的应用,或者在T细胞标记、检测、细胞分选或活化中的应用。
12.一种药物组合物或试剂盒,所述的药物组合物或试剂盒包含下列任一组:
i)权利要求1-2任一所述的T细胞抗原受体;
ii)权利要求3所述的核酸;
iii)权利要求4所述的表达载体;
iv)权利要求5所述的宿主细胞;
v)权利要求5所述的免疫细胞;或
vi)权利要求8-9任一所述的多聚体复合物。
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CN202110133762.9A Pending CN113881680A (zh) | 2020-07-01 | 2021-02-01 | T细胞抗原受体、其多聚体复合物及其制备方法和应用 |
CN202110240703.1A Pending CN113880937A (zh) | 2020-07-01 | 2021-03-04 | T细胞抗原受体、其多聚体复合物及其制备方法和应用 |
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CN202110240703.1A Pending CN113880937A (zh) | 2020-07-01 | 2021-03-04 | T细胞抗原受体、其多聚体复合物及其制备方法和应用 |
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EP (1) | EP4163297A4 (zh) |
JP (1) | JP2023532108A (zh) |
KR (1) | KR20230061336A (zh) |
CN (3) | CN113880953A (zh) |
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WO2023238125A1 (en) * | 2022-06-08 | 2023-12-14 | Sheba Impact Ltd. | Virus-specific recombinant t cell receptors and t cells comprising them |
CN117736300A (zh) * | 2022-12-26 | 2024-03-22 | 上海市第一人民医院 | 靶向巨细胞病毒pp65的T细胞受体和表达其的T细胞及应用 |
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GB0917094D0 (en) * | 2009-09-29 | 2009-11-11 | Ucl Biomedica Plc | T-cell receptor |
US11390921B2 (en) * | 2014-04-01 | 2022-07-19 | Adaptive Biotechnologies Corporation | Determining WT-1 specific T cells and WT-1 specific T cell receptors (TCRs) |
CN106279404A (zh) | 2015-05-20 | 2017-01-04 | 广州市香雪制药股份有限公司 | 一种可溶且稳定的异质二聚tcr |
CN106632660B (zh) * | 2015-11-04 | 2021-01-29 | 广东香雪精准医疗技术有限公司 | 识别ny-eso-1抗原短肽的tcr |
CN107827959B (zh) | 2017-11-09 | 2018-10-30 | 杭州续缓生物科技有限公司 | 识别乙肝病毒(hbv)表面抗原s183-91表位的tcr及其用途 |
EP3731861A1 (en) * | 2017-12-29 | 2020-11-04 | Max-Delbrück-Centrum für Molekulare Medizin in der Helmholtz-Gemeinschaft | T cell receptors for tumor specific proteasome splice variants and uses thereof |
BR112020016330A8 (pt) * | 2018-02-12 | 2023-04-18 | Hutchinson Fred Cancer Res | Receptores de células t específicas a ciclina a1 e usos dos mesmos |
US20210393684A1 (en) * | 2018-05-18 | 2021-12-23 | The Council Of The Queensland Institute Of Medical Research (Qimr) | Adoptive T-Cell Therapy for CMV Infection and CMV-Associated Diseases |
KR20210095157A (ko) * | 2018-11-08 | 2021-07-30 | 넥스이뮨, 인크. | 개선된 표현형 속성을 갖는 t 세포 조성물 |
US20220144916A1 (en) * | 2019-02-12 | 2022-05-12 | Board Of Regents, The University Of Texas System | High affinity engineered t-cell receptors targeting cmv infected cells |
CN110357953B (zh) * | 2019-07-17 | 2022-05-03 | 深圳市因诺转化医学研究院 | 识别人巨细胞病毒pp65抗原的TCR |
CN113121676B (zh) * | 2020-01-15 | 2022-05-31 | 深圳华大生命科学研究院 | 一种靶向巨细胞病毒抗原的特异性t细胞受体及其应用 |
EP4121079A1 (en) * | 2020-03-16 | 2023-01-25 | University Of Southern California | Novel antigen binding domains and synthetic antigen receptors incorporating the same |
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2021
- 2021-01-20 CN CN202110072284.5A patent/CN113880953A/zh active Pending
- 2021-02-01 CN CN202110133762.9A patent/CN113881680A/zh active Pending
- 2021-03-04 CN CN202110240703.1A patent/CN113880937A/zh active Pending
- 2021-06-29 JP JP2022581319A patent/JP2023532108A/ja active Pending
- 2021-06-29 CA CA3184695A patent/CA3184695A1/en active Pending
- 2021-06-29 WO PCT/CN2021/102954 patent/WO2022002014A1/zh unknown
- 2021-06-29 EP EP21831677.6A patent/EP4163297A4/en active Pending
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KR20230061336A (ko) | 2023-05-08 |
JP2023532108A (ja) | 2023-07-26 |
US20230257447A1 (en) | 2023-08-17 |
EP4163297A4 (en) | 2024-05-22 |
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CA3184695A1 (en) | 2022-01-06 |
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