CN114045277A - 碱基编辑器及其构建方法与应用 - Google Patents
碱基编辑器及其构建方法与应用 Download PDFInfo
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Abstract
本发明涉及基因编辑技术领域，尤其是涉及一种碱基编辑器及其构建方法与应用，用于DNA水平特定位点特定碱基的突变。本发明构建得到编辑特异性更高且RNA/DNA脱靶风险低的ABE，编辑精度高且RNA/DNA脱靶风险低的CBE以及体积更小的ACBE工具。本发明中开发的ABE工具在介导A·T‑‑G·C突变时有更高的特异性，其介导C·G‑‑T·A突变的能力被明显抑制，甚至清除，且在RNA和DNA水平的脱靶效应有明显降低，甚至消除；本发明开发的CBE工具在介导C·G‑‑T·A突变时有更高的精确度，且RNA和DNA水平的脱靶效应有明显降低，甚至消除；本发明中开发的ACBE工具，只需要单一的脱氨酶，其体积更小，且在RNA和DNA水平的脱靶效应有明显降低，甚至消除，具有更高的应用价值。
Description
技术领域
本发明涉及基因编辑技术领域，尤其是涉及碱基编辑器及其构建方法。
背景技术
CRISPR/Cas9为代表的新型基因编辑技术给生物医学研究带来了变革，这一技术让高效精准的基因编辑成为可能。2016年以来，研究者开发出多种DNA碱基编辑工具可以在DNA水平高效诱导精准的点突变，且此过程中不会产生DNA双链断裂。目前已报道的碱基编辑器有很多种，其中最主要的是两种：胞嘧啶碱基编辑器(CBE，可介导C·G--T·A突变)和腺嘌呤碱基编辑器(ABE，可介导A·T--G·C突变)。其中，CBE是将特定的胞嘧啶核苷脱氨酶(cytidine deaminase)与突变型核酸酶(如携带有D10A或突变的spCas9，称为nspCas9)以及尿嘧啶糖基化酶抑制因子(UGI)融合，得到的融合蛋白可在sgRNA的引导下，介导C·G--T·A突变。ABE则是将定向演化后的大肠杆菌来源的腺苷脱氨酶(ecTadA*)二聚体或单体与突变型核酸酶(如携带有D10A或突变的spCas9，称为nCas9)融合，得到的融合蛋白可在sgRNA的引导下，介导A·T--G·C突变。在ABE和CBE的基础上，研究者还将特定的胞嘧啶核苷脱氨酶(cytidine deaminase)、定向演化后的大肠杆菌来源的腺苷脱氨酶(ecTadA*)、突变型核酸酶(如携带有D10A或突变的spCas9，称为nCas9)以及尿嘧啶糖基化酶抑制因子(UGI)融合构建出ACBE，得到的融合蛋白可在sgRNA的引导下，可介导特定位点中C·G--T·A突变和A·T--G·C突变的同时发生(Nat Biotechnol,2020.38(7):861-864；BMC Biol,2020.18(1):131；Nat Biotechnol,2020.38(7):856-860；Nat Biotechnol,2020.38(7):865-869)。现有的ABE、CBE和ACBE工具依然存在不足，限制了其应用前景。
现有ABE工具除能介导A·T--G·C突变外，在一定程度上还能介导C·G--T·A突变，这降低了ABE工具编辑的特异性和精确度，不利于ABE的应用(Nat Biotechnol,2021(https://doi.org/10.1038/s41587-021-00943-2)；Nat Biotechnol,2019.37(10):1145-1148；Nat Commun,2020.11(1):5827)。虽然有研究者根据ABE工具能介导C·G--T·A突变的特性，通过点突变构建出能特异性介导TC-TT突变的CBE工具，但其活性非常低，应用场景有限(Nat Biotechnol,2021，https://doi.org/10.1038/s41587-021-00943-2)。此外，目前适用于ABE构建的脱氨基酶均来自定向演化后的大肠杆菌来源的腺苷脱氨酶(ecTadA*)，脱氨基酶种类非常局限，这也限制了ABE碱基编辑特性的多样性，不利于工具的应用推广。此外，目前也缺少RNA/DNA脱靶效应以及编辑精度等多个方面进行协同优化的ABE工具。
现有CBE工具中的脱氨酶均属于胞嘧啶核苷脱氨酶(cytidine deaminase)家族，这类工具目前在碱基编辑精度(活性窗口特异性)、RNA脱靶效应以及DNA脱靶效应中均存在一系列问题，目前依然缺少在RNA/DNA脱靶效应以及编辑精度等多个方面进行协同优化的CBE工具(Nat Biotechnol,2020.38(5):620-628；Nat Commun,2020.11(1):2052；NatMethods,2020.17(6):600-604)。
现有ACBE工具需要在nCas9上同时融合两种脱氨酶(胞嘧啶核苷脱氨酶(cytidinedeaminase)、定向演化后的大肠杆菌来源的腺苷脱氨酶(ecTadA*))，这导致ACBE工具的体积过大，不利于应用(Nat Biotechnol,2020.38(7):861-864；BMC Biol,2020.18(1):131；Nat Biotechnol,2020.38(7):856-860；Nat Biotechnol,2020.38(7):865-869)。
发明内容
为了对碱基编辑器ABE、CBE和ACBE的不足进行优化，以开发出编辑特异性更高且RNA/DNA脱靶风险低的ABE，编辑精度高且RNA/DNA脱靶风险低的CBE以及体积更小的ACBE工具，本发明的目的是提供新型碱基编辑器及其构建方法与应用，用于DNA水平特定位点特定碱基的突变。
可实现DNA水平特定位点C·G碱基对向T·A碱基对的转换(C·G--T·A突变，CBE)，实现A·T碱基对向G·C碱基对的突变(A·T--G·C突变，ABE)，或同时实现特定位点C·G碱基对向T·A碱基对和A·T碱基对向G·C碱基对的转换(C·G--T·A和A·T--G·C同时突变，ACBE)。
本发明提供一种新的腺嘌呤碱基编辑器(ABE)、胞嘧啶碱基编辑器(CBE)和ACBE的构建方法，以开发出编辑特异性更高且RNA/DNA脱靶风险低的ABE，编辑精度高且RNA/DNA脱靶风险低的CBE以及体积更小的ACBE工具。
本发明的基本原理是，针对定向演化后的大肠杆菌来源的腺苷脱氨酶(TadA-8e)(Nat Biotechnol,2020.38(7):883-891)，对其关键氨基酸位点进行饱和突变筛选。最终通过特定点突变或是点突变组合，实现ABE、CBE和ACBE的构建和各种碱基编辑器之间的转换；或针对自然界的多种腺苷脱氨酶(TadA)进行突变和系统筛选，寻找新的功能性TadA，构建多样化的ABE、CBE和ACBE。
大肠杆菌来源的腺苷脱氨酶(ecTadA)经蛋白定向演化后(演化后称为ecTadA*)，已经具备对单链DNA中腺嘌呤(A)进行脱氨基化，实现A·T--G·C突变的能力。在此基础上构建的碱基编辑器ABE可以实现基因组目标位点中的A·T--G·C突变。研究者还发现，一定条件下，ABE能对单链DNA中胞嘧啶(C)进行脱氨基化，一定程度上还能介导C·G--T·A突变，具有一定的CBE功能。这种介导C·G--T·A突变的能力增加了ABE工具的脱靶风险(NatBiotechnol,2019.37(10):1145-1148；Nat Commun,2020.11(1):5827)，因此，对腺苷脱氨酶(TadA)的特定位点进行饱和突变筛选，或是对自然界存在的腺苷脱氨酶(TadA)进行点突变和系统筛选，或能构建出新的ABE、CBE和ACBE。特定的点突变或点突变组合可以实现以下目的：
(1)降低或消除ABE工具介导C·G--T·A突变的能力，并改善RNA脱靶效应以及DNA脱靶效应等不足，增强其安全性；
(2)将ABE转变为CBE，获得新的CBE工具，新工具在碱基编辑精度(活性窗口特异性)、RNA脱靶效应以及DNA脱靶效应等方面具有不同于已有工具的新特性；
(3)将ABE转变为ACBE，利用单一的脱氨酶构建出新型的ACBE，减小ACBE的体积，改善其碱基编辑精度(活性窗口特异性)、RNA脱靶效应以及DNA脱靶效应。
本发明人经过系统性的研究和饱和式筛选，首次发现，脱氨酶TadA-8e中氨基酸位点为第28、30、46、48、49、82、84、108、110或111位的氨基酸中一个或几个突变后对TadA-8e的脱氨基活性及碱基偏好性有显著影响。不同类型的点突变，或是多种点突变的组合，可以显著改变TadA-8e为基础的ABE工具的特性。通过特定氨基酸位点的不同点突变，或是多个氨基酸位点点突变的组合，最终获得编辑特异性更高的ABE，编辑精度高且RNA/DNA脱靶风险低的CBE以及体积更小的ACBE工具，最终完成本发明。
本发明的目的可以通过以下技术方案来实现：
本发明的第一个目的是提供一种碱基编辑器，为突变型TadA-8e置于Cas核酸酶的N-末端、C-末端或内部的融合位点融合形成的融合蛋白；
突变型TadA-8e是指TadA-8e中距离腺嘌呤(A)类似物8-azanebularine(8Az)小于5埃的氨基酸中一个或几个突变后的脱氨酶，尤其是指TadA-8e中氨基酸位点为第28、30、46、48、49、82、84、108、110或111位的氨基酸中一个或几个突变后的脱氨酶；
TadA-8e的氨基酸序列如SEQ ID NO.1所示。
在本发明的一个实施方式中，所述Cas核酸酶选自Cas9、Cas12、Cas13蛋白家族；优选地，所述Cas核酸酶选自nSpCas9及其突变体、SaCas9及其突变体、Cas12a及其突变体或Cas12f及其突变体，优选为nSpCas9。
在本发明的一个实施方式中，突变型TadA-8e是指TadA-8e中氨基酸位点为第28、30、46、48、49、82、84、108、110或111位的氨基酸中一个突变为不同于所在位点自身原来氨基酸的另外19种氨基酸中的一种以后所得的脱氨酶；
优选地，突变型TadA-8e是指TadA-8e中氨基酸进行以下单点突变后所得的脱氨酶：V28W、V28G、N46L、N46P、N46C、A48C、A48G、A48N、A48S、I49D、I49E、I49H、I49N、I49P、I49Q、I49V、I49W、I49Y、I49R、I49A、I49T、I49G、I49L、I49S、I49M、I49C、I49F、I49K、V82G、V82A、V82T、V82C、V82S、V82D、F84M、N108H、N108M、N108C、N108D、N108F、N108Q、N108S、N108Y、V28A、V28D、V28E、V28K、V28M、V28R、V28L、V28Q、V30A、V30D、V30G、V30W、V30E、V30N、V30Q、V30Y、V30T、V30S、N46H、N46K、N46Y、N46F、N46E、N46Q、N46D、N46G、N46M、N46V、N46S、N46T、A48D、A48E、A48Y、A48W、A48F、A48Q、V82F、F84D、F84R、F84A、F84E、F84P、F84K、F84C、F84H、F84I、F84S、F84T、F84V、F84Q、F84N、F84G、F84Y、N108V、N108I、N108L、N108A、N108T、N108R、K110D、K110W、K110E、R111P、R111E、R111T、R111A、R111V、R111G、R111I、R111W、R111L、R111D。
上述类似“V28W”的表述中，28表示TadA-8e中第28位氨基酸位点，V表示TadA-8e中第28位氨基酸位点突变前的氨基酸，W表示TadA-8e中第28位氨基酸位点突变后的氨基酸，V、W均为氨基酸的缩写表达方式，即V28W表示第28位氨基酸位点由缬氨酸突变为色氨酸；上文或下文中其他类似“V28W”的表达方式均做与此相似的解释，即N46L表示第46位氨基酸位点由天冬酰胺突变为亮氨酸。
通过上述单点突变得到的碱基编辑器介导C·G--T·A突变。
在本发明的一个实施方式中，突变型TadA-8e是指TadA-8e中氨基酸进行以下双点突变后所得的脱氨酶：
(1)氨基酸位点第28位的氨基酸突变的同时，第46、48、49、82、84、108、110或111位的氨基酸中一个突变后的脱氨酶，或，
(2)氨基酸位点第46位的氨基酸突变的同时，第28、82、84、108位的氨基酸中一个突变后的脱氨酶，或，
(3)氨基酸位点第48、108位的氨基酸同时突变后的脱氨酶，或，
(4)氨基酸位点第49位的氨基酸突变的同时，第82、84、108位的氨基酸中一个突变后的脱氨酶，或，
(5)氨基酸位点第108位的氨基酸突变的同时，第82或84位的氨基酸中一个突变后的脱氨酶。
优选地，突变型TadA-8e是指TadA-8e中氨基酸进行以下双点突变后所得的脱氨酶：
V28G-N108Y，V28G-N108C，V28G-N108H，V28G-N46L，V28G-N46C，V28G-N46P，V28W-N46L，V28W-N46C，V28W-N46P，N46P-N108Y，N46P-N108S，N46P-N108Q，N46P-N108D，N46P-N108C，N46P-N108H，N46P-F84M，N46P-I49A，N46P-I49G，N46P-A48G，N46C-N108Y，N46C-N108S，N46C-N108Q，N46C-N108M，N46C-N108C，N46C-N108H，N46L-N108Y，N46L-N108Q，N46L-N108D，N46L-N108M，N46L-N108H，N46L-F84M，N46L-I49A，N46L-I49G，N46L-V82G，I49A-N108Y、V28G-A48G、V28G-I49A、V28G-I49G、V28G-V82T、V28G-F84Y、V28W-N108H、N46C-V82G、N46C-V82T、N46C-F84M、N46L-V82T、N46P-F84M、N46P-N108M、A48G-N108H、I49A-N108H、I49G-N108H、I49G-N108Y、I49G-V82G、I49G-V82T、I49G-F84Y、V82G-N108Y、V82T-N108H、V82T-N108C、F84M-N108H、F84M-N108M、F84M-N108Q。
所述V28G-N46C表示第28位氨基酸位点由缬氨酸突变为甘氨酸以及第46位氨基酸位点由天冬酰胺变为半胱氨酸，上文或下文中其他类似的表达方式均做与此相似的解释。
通过上述双点突变得到的碱基编辑器介导C·G--T·A突变。
在本发明的一个实施方式中，所述突变型TadA-8e是指TadA-8e中氨基酸进行以下三点突变后所得的脱氨酶：
(1)氨基酸位点第28、48、49位的氨基酸同时突变后的脱氨酶，或，
(2)氨基酸位点第28、48、82位的氨基酸同时突变后的脱氨酶，或，
(3)氨基酸位点第28、48、108位的氨基酸同时突变后的脱氨酶，或，
(4)氨基酸位点第28、49、82位的氨基酸同时突变后的脱氨酶，或，
(5)氨基酸位点第28、49、108位的氨基酸同时突变后的脱氨酶，或，
(6)氨基酸位点第49、82、108位的氨基酸同时突变后的脱氨酶；
优选地，所述突变型TadA-8e是指TadA-8e中氨基酸进行以下三点突变后所得的脱氨酶：
I49A-V82T-N108Y、I49A-V82G1-N08Y、I49G-V82T-N108Y、I49G-V82G-N108Y、A48G-V82G-N108Y、A48G-I49G-N108Y、V28G-V82G-N108Y、V28G-V82G-N108H、V28G-I49A-N108Y、V28G-I49A-N108H、V28G-I49G-N108H、V28G-A48G-N108Y、V28G-A48G-N108H、V28G-A48G-I49G、V28G-A48G-I49A、V28G-A48G-V82G、V28G-A48G-V82T、V28G-I49G-V82G、V28G-I49G-V82T、V28G-I49G-N108Y、V28G-I49A-V82G、V28G-I49A-V82T、I49G-V82G-N108H、I49G-V82T-N108H。
所述V28G-A48G-I49G表示第28位氨基酸位点由缬氨酸突变为甘氨酸、第48位氨基酸位点由丙氨酸变为甘氨酸以及第49位由异亮氨酸突变为甘氨酸，上文或下文中其他类似的表达方式均做与此相似的解释。
通过上述三点突变得到的碱基编辑器介导C·G--T·A突变。
在本发明的一个实施方式中，所述突变型TadA-8e是指TadA-8e中氨基酸位点第108位的氨基酸突变的同时，TadA-8e中氨基酸位点第28、48、49、82中还有三个位点突变后的脱氨酶；
优选地，所述突变型TadA-8e是指TadA-8e中氨基酸位点第108位进行突变N108Y或N108H的同时，TadA-8e中氨基酸位点第28、48、49、82中还有三个位点进行如下突变后的脱氨酶；
TadA-8e中氨基酸位点第28位进行突变V28G，
TadA-8e中氨基酸位点第48位进行突变A48G，
TadA-8e中氨基酸位点第49位进行突变I49A或I49G，
TadA-8e中氨基酸位点第82位进行突变V82T或V82G。
所述V28G-I49G-V82T-N108Y表示第28位氨基酸由缬氨酸突变为甘氨酸、第49位氨基酸由异亮氨酸突变为甘氨酸、第82位氨基酸由缬氨酸突变为苏氨酸以及第108位氨基酸由天冬酰胺突变为酪氨酸，上文或下文中其他类似的表达方式均做与此相似的解释。
通过上述四点突变得到的碱基编辑器介导C·G--T·A突变。
在本发明的一个实施方式中，所述突变型TadA-8e是指TadA-8e中氨基酸位点第28、48、49、82、108位的氨基酸同时突变后的脱氨酶；
优选地，所述突变型TadA-8e是指TadA-8e中氨基酸进行以下五点突变后所得的脱氨酶：
TadA-8e中氨基酸位点第108位进行突变N108Y或N108H的同时，TadA-8e中氨基酸位点第28、48、49、82位点还进行如下突变后的脱氨酶；
TadA-8e中氨基酸位点第28位进行突变V28G，
TadA-8e中氨基酸位点第48位进行突变A48G，
TadA-8e中氨基酸位点第49位进行突变I49A或I49G，
TadA-8e中氨基酸位点第82位进行突变V82T或V82G。
所述V28G-A48G-I49G-V82T-N108H表示第28位氨基酸由缬氨酸突变为甘氨酸、第48位氨基酸由丙氨酸突变为甘氨酸、第49位氨基酸由异亮氨酸突变为甘氨酸、第82位氨基酸由缬氨酸突变为苏氨酸以及第108位氨基酸由天冬酰胺突变为组氨酸，，上文或下文中其他类似的表达方式均做与此相似的解释。
通过上述五点突变得到的碱基编辑器介导C·G--T·A突变。
在本发明的一个实施方式中，突变型TadA-8e的5’和3’分别添加不同序列的各类变体后置于Cas核酸酶的N-末端、C-末端或内部的融合位点融合形成的融合蛋白；
优选地，突变型TadA-8e的5’和3’分别添加NLS序列和linker序列后获得NLS-突变型TadA-8e-linker，NLS-突变型TadA-8e-linker置于Cas核酸酶的N-末端、C-末端或内部的融合位点融合形成的融合蛋白；
其中，NLS-突变型TadA-8e-linker是指相对于NLS-TadA-8e-linker而言，TadA-8e中氨基酸中一个或几个发生了突变，NLS-TadA-8e-linker的氨基酸序列如SEQ ID NO.4所示，NLS-突变型TadA-8e-linker的氨基酸序列为NLS-TadA-8e-linker氨基酸序列中将TadA-8e置换为突变型TadA-8e得到的氨基酸序列。NLS序列为PKKKRKV，linker序列为SGGSSGGSSGSETPGTSESATPESSGGSSGGSSR。
在本发明的一个实施方式中，所述Cas核酸酶选择为nSpCas9时，所述nSpCas9内部的融合位点包括第113、203、312、459、535、583、687、701、715、730、770、793、801、895、905、919、946、1010、1029、1047、1117、1154、1249、1282位融合位点；
优选为第1249位融合位点。
本发明的第二个目的是提供一种碱基编辑器，TadA-8e的5’和3’分别添加NLS序列和linker序列后获得NLS-TadA-8e-linker，NLS-TadA-8e-linker置于Cas核酸酶的N-末端、C-末端或内部的融合位点融合形成的融合蛋白；
NLS-TadA-8e-linker的氨基酸序列如SEQ ID NO.4所示；
所述Cas核酸酶选自Cas9、Cas12、Cas13蛋白家族，优选地，所述Cas核酸酶选自nSpCas9及其突变体、SaCas9及其突变体、Cas12a及其突变体或Cas12f及其突变体，进一步优选为nSpCas9。
在本发明的一个实施方式中，所述Cas核酸酶选择为nSpCas9时，所述nSpCas9内部的融合位点包括第113、203、312、459、535、583、687、701、715、730、770、793、801、895、905、919、946、1010、1029、1047、1117、1154、1249、1282位融合位点；
优选为第1249位融合位点。
本发明的第三个目的是提供一种碱基编辑器，为突变型TadA-X置于Cas核酸酶的N-末端、C-末端或内部的融合位点融合形成的融合蛋白；
突变型TadA-X是指TadA-X中第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺的脱氨酶；
TadA-X是指不同来源的TadA；优选地，TadA-X有多组，其突变体氨基酸序列分别如SEQ ID NO.5-SEQ ID NO.57中的任一所示；
所述Cas核酸酶选自Cas9、Cas12、Cas13蛋白家族，优选地，所述Cas核酸酶选自nSpCas9及其突变体、SaCas9及其突变体、Cas12a及其突变体或Cas12f及其突变体，进一步优选为nSpCas9。
在本发明的一个实施方式中，突变型TadA-X是指Q2LYC0、F2G306、G0JN58、B5ZCW4、Q1AS41、E4TTF3、Q57LE3、Q99W51、ecTadA中与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺的脱氨酶；
Q2LYC0、F2G306、G0JN58、B5ZCW4、Q1AS41、E4TTF3、Q57LE3、Q99W51或ecTadA突变体的氨基酸序列分别为SEQ ID NO.8、SEQ ID NO.14、SEQ ID NO.20、SEQ ID NO.22、SEQ IDNO.27、SEQ ID NO.43、SEQ ID NO.54、SEQ ID NO.55、SEQ ID NO.58。
上述得到的碱基编辑器介导A·T--G·C突变。
在本发明的一个实施方式中，突变型TadA-X是指B3PCY2、A5WFT9、C1DXN9、B2UR34、R4K908、D5W8R4、E1TAF4中与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺的脱氨酶；
B3PCY2、A5WFT9、C1DXN9、B2UR34、R4K908、D5W8R4、E1TAF4突变体的氨基酸序列分别为SEQ ID NO.15、SEQ ID NO.19、SEQ ID NO.29、SEQ ID NO.32、SEQ ID NO.44、SEQ IDNO.49、SEQ ID NO.57。
上述得到的碱基编辑器介导C·G--T·A突变。
在本发明的实施方式中，突变型TadA-X是指Q13XZ4、A8F050、E8WVH3、P44931、G0EXN4、E0TBL5、A1W869、D5X1Y1、I3YB54与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺的脱氨酶；
Q13XZ4、A8F050、E8WVH3、P44931、G0EXN4、E0TBL5、A1W869、D5X1Y1、I3YB54突变体的氨基酸序列分别为SEQ ID NO.10、SEQ ID NO.18、SEQ ID NO.25、SEQ ID NO.28、SEQ IDNO.38、SEQ ID NO.46、SEQ ID NO.48、SEQ ID NO.50、SEQ ID NO.53。
上述得到的碱基编辑器介导A·T--G·C和C·G--T·A突变。
在本发明的实施方式中，突变型TadA-X的5’和3’分别添加NLS序列和linker序列后获得NLS-突变型TadA-X-linker，NLS-突变型TadA-X-linker置于Cas核酸酶的N-末端、C-末端或内部的融合位点融合形成的融合蛋白；
其中，NLS-突变型TadA-X-linker是指相对于NLS-TadA-8e-linker而言，TadA-8e替换为突变型TadA-X，NLS-TadA-8e-linker的氨基酸序列如SEQ ID NO.4所示。
在本发明的实施方式中，所述Cas核酸酶选择为nSpCas9时，所述nSpCas9内部的融合位点包括第113、203、312、459、535、583、687、701、715、730、770、793、801、895、905、919、946、1010、1029、1047、1117、1154、1249、1282位融合位点；
优选为第1249位融合位点。
本发明的第四个目的是提供一种碱基编辑器的构建方法，其特征在于，包括以下步骤：
将TadA-8e、突变型TadA-8e或突变型TadA-X和Cas核酸酶不同位点融合，构建得到碱基编辑器；
TadA-8e的氨基酸序列如SEQ ID NO.1所示；
突变型TadA-8e是指TadA-8e中氨基酸位点为第28、30、46、48、49、82、84、108、110或111位的氨基酸中一个或几个突变后的脱氨酶；
突变型TadA-X是指TadA-X中与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺的脱氨酶；TadA-X有多组，其突变体氨基酸序列分别如SEQ ID NO.5-SEQID NO.57中的任一所示。
在本发明的一个实施方式中，还可以将TadA-8e、突变型TadA-8e或突变型TadA-X5’和3’分别添加NLS序列和linker序列。
本发明的第五个目的是提供一种碱基编辑器ABE的应用，所述碱基编辑器ABE介导A·T--G·C突变；
碱基编辑器ABE是指Q2LYC0、F2G306、G0JN58、B5ZCW4、Q1AS41、E4TTF3、Q57LE3、Q99W51、ecTadA与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺后置于nSpCas9的N-末端、第1249位融合位点融合形成的融合蛋白，其中，Q2LYC0、F2G306、G0JN58、B5ZCW4、Q1AS41、E4TTF3、Q57LE3、Q99W51或ecTadA突变体的氨基酸序列分别为SEQ IDNO.8、SEQ ID NO.14、SEQ ID NO.20、SEQ ID NO.22、SEQ ID NO.27、SEQ ID NO.43、SEQ IDNO.54、SEQ ID NO.55、SEQ ID NO.58；
碱基编辑器ABE还指TadA-8e中氨基酸进行以下单点突变或点突变组合后所得的脱氨酶置于nSpCas9的N-末端、第1249位融合位点融合形成的融合蛋白，这类融合蛋白仅具有明显的介导A·T--G·C突变能力。
本发明的第六个目的是提供一种碱基编辑器CBE的应用，所述碱基编辑器CBE介导C·G--T·A突变；
碱基编辑器CBE是指B3PCY2、A5WFT9、C1DXN9、B2UR34、R4K908、D5W8R4、E1TAF4与ecTadA第106和第108位同源的氨基酸分别突变为缬氨酸和天冬酰胺后置于nSpCas9的N-末端、第1249位融合位点融合形成的融合蛋白，其中，B3PCY2、A5WFT9、C1DXN9、B2UR34、R4K908、D5W8R4、E1TAF4突变体的氨基酸序列分别为SEQ ID NO.15、SEQ ID NO.19、SEQ IDNO.29、SEQ ID NO.32、SEQ ID NO.44、SEQ ID NO.49、SEQ ID NO.57；
碱基编辑器CBE还指TadA-8e中氨基酸进行以下突变后所得的脱氨酶置于nSpCas9的N-末端、第1249位融合位点融合形成的融合蛋白，
单点突变：N46P
双点突变：V28G-N108Y，V28G-N108C，V28G-N108H，V28G-N46L，V28G-N46C，V28G-N46P，V28W-N46L，V28W-N46C，V28W-N46P，N46P-N108Y，N46P-N108S，N46P-N108Q，N46P-N108D，N46P-N108C，N46P-N108H，N46P-F84M，N46P-I49A，N46P-I49G，N46P-A48G，N46C-N108Y，N46C-N108S，N46C-N108Q，N46C-N108M，N46C-N108C，N46C-N108H，N46L-N108Y，N46L-N108Q，N46L-N108D，N46L-N108M，N46L-N108H，N46L-F84M，N46L-I49A，N46L-I49G，N46L-V82G，I49A-N108Y
三点突变：I49A-V82T-N108Y、I49A-V82G1-N08Y、I49G-V82T-N108Y、I49G-V82G-N108Y、A48G-V82G-N108Y、A48G-I49G-N108Y、V28G-V82G-N108Y、V28G-V82G-N108H、V28G-I49A-N108Y、V28G-I49A-N108H、V28G-I49G-N108H、V28G-A48G-N108Y、V28G-A48G-N108H；
四点突变：V28G-A48G-I49A-N108Y、V28G-A48G-I49G-N108Y、V28G-A48G-V82T-N108Y、V28G-A48G-V82G-N108Y、V28G-I49A-V82T-N108Y、V28G-I49A-V82G-N108Y、V28G-I49G-V82T-N108Y、V28G-I49G-V82G-N108Y、A48G-I49A-V82T-N108Y、A48G-I49A-V82G-N108Y、A48G-I49G-V82T-N108Y、A48G-I49G-V82G-N108Y、V28G-A48G-I49A-N108H、V28G-A48G-I49G-N108H、V28G-A48G-V82T-N108H、V28G-A48G-V82G-N108H、V28G-I49A-V82T-N108H、V28G-I49A-V82G-N108H、V28G-I49G-V82T-N108H、V28G-I49G-V82G-N108H、A48G-I49A-V82T-N108H、A48G-I49A-V82G-N108H、A48G-I49G-V82T-N108H、A48G-I49G-V82G-N108H。
五点突变：V28G-A48G-I49A-V82T-N108Y、V28G-A48G-I49 G-V82T-N108Y、V28G-A48G-I49A-V82G-N108Y、V28G-A48G-I49G-V82G-N108Y、V28G-A48G-I49A-V82T-N108H、V28G-A48G-I49G-V82T-N108H、V28G-A48G-I49A-V82G-N108H、V28G-A48G-I49G-V82G-N108H。
本发明的第七个目的是提供一种碱基编辑器ACBE的应用，所述碱基编辑器ACBE介导A·T--G·C和C·G--T·A突变；
碱基编辑器ACBE是指Q13XZ4、A8F050、E8WVH3、P44931、G0EXN4、E0TBL5、A1W869、D5X1Y1、I3YB54与ecTadA第106和第108位同源的氨基酸分别突变为缬氨酸和天冬酰胺后置于nSpCas9的N-末端、第1249位融合位点融合形成的融合蛋白，其中，Q13XZ4、A8F050、E8WVH3、P44931、G0EXN4、E0TBL5、A1W869、D5X1Y1、I3YB54突变体的氨基酸序列分别为SEQID NO.10、SEQ ID NO.18、SEQ ID NO.25、SEQ ID NO.28、SEQ ID NO.38、SEQ ID NO.46、SEQID NO.48、SEQ ID NO.50、SEQ ID NO.53；
碱基编辑器ACBE还指TadA-8e中氨基酸进行点突变或点突变组合后所得的脱氨酶置于nSpCas9的N-末端、第1249位融合位点融合形成的融合蛋白，这类融合蛋白具有介导A·T--G·C和C·G--T·A突变能力。
本发明第七方面，提供突变型TadA-8e蛋白，所述突变蛋白为非天然蛋白，突变型TadA-8e蛋白指TadA-8e中氨基酸位点为第28、30、46、48、49、82、84、108、110或111位的氨基酸中一个或几个突变后的脱氨酶；
优选地，突变型TadA-8e是指TadA-8e中氨基酸进行以下单点突变后所得的脱氨酶：
V28W、V28G、N46L、N46P、N46C、A48C、A48G、A48N、A48S、I49D、I49E、I49H、I49N、I49P、I49Q、I49V、I49W、I49Y、I49R、I49A、I49T、I49G、I49L、I49S、I49M、I49C、I49F、I49K、V82G、V82A、V82T、V82C、V82S、V82D、F84M、N108H、N108M、N108C、N108D、N108F、N108Q、N108S、N108Y、V28A、V28D、V28E、V28K、V28M、V28R、V28L、V28Q、V30A、V30D、V30G、V30W、V30E、V30N、V30Q、V30Y、V30T、V30S、N46H、N46K、N46Y、N46F、N46E、N46Q、N46D、N46G、N46M、N46V、N46S、N46T、A48D、A48E、A48Y、A48W、A48F、A48Q、V82F、F84D、F84R、F84A、F84E、F84P、F84K、F84C、F84H、F84I、F84S、F84T、F84V、F84Q、F84N、F84G、F84Y、N108V、N108I、N108L、N108A、N108T、N108R、K110D、K110W、K110E、R111P、R111E、R111T、R111A、R111V、R111G、R111I、R111W、R111L、R111D；
或，突变型TadA-8e是指TadA-8e中氨基酸进行以下双点突变后所得的脱氨酶：
V28G-N108Y，V28G-N108C，V28G-N108H，V28G-N46L，V28G-N46C，V28G-N46P，V28W-N46L，V28W-N46C，V28W-N46P，N46P-N108Y，N46P-N108S，N46P-N108Q，N46P-N108D，N46P-N108C，N46P-N108H，N46P-F84M，N46P-I49A，N46P-I49G，N46P-A48G，N46C-N108Y，N46C-N108S，N46C-N108Q，N46C-N108M，N46C-N108C，N46C-N108H，N46L-N108Y，N46L-N108Q，N46L-N108D，N46L-N108M，N46L-N108H，N46L-F84M，N46L-I49A，N46L-I49G，N46L-V82G，I49A-N108Y、V28G-A48G、V28G-I49A、V28G-I49G、V28G-V82T、V28G-F84Y、V28W-N108H、N46C-V82G、N46C-V82T、N46C-F84M、N46L-V82T、N46P-F84M、N46P-N108M、A48G-N108H、I49A-N108H、I49G-N108H、I49G-N108Y、I49G-V82G、I49G-V82T、I49G-F84Y、V82G-N108Y、V82T-N108H、V82T-N108C、F84M-N108H、F84M-N108M、F84M-N108Q；
或，所述突变型TadA-8e是指TadA-8e中氨基酸进行以下三点突变后所得的脱氨酶：
I49A-V82T-N108Y、I49A-V82G1-N08Y、I49G-V82T-N108Y、I49G-V82G-N108Y、A48G-V82G-N108Y、A48G-I49G-N108Y、V28G-V82G-N108Y、V28G-V82G-N108H、V28G-I49A-N108Y、V28G-I49A-N108H、V28G-I49G-N108H、V28G-A48G-N108Y、V28G-A48G-N108H、V28G-A48G-I49G、V28G-A48G-I49A、V28G-A48G-V82G、V28G-A48G-V82T、V28G-I49G-V82G、V28G-I49G-V82T、V28G-I49G-N108Y、V28G-I49A-V82G、V28G-I49A-V82T、I49G-V82G-N108H、I49G-V82T-N108H；
或，所述突变型TadA-8e是指TadA-8e中氨基酸进行以下四点突变后所得的脱氨酶：V28G-A48G-I49A-N108Y、V28G-A48G-I49G-N108Y、V28G-A48G-V82T-N108Y、V28G-A48G-V82G-N108Y、V28G-I49A-V82T-N108Y、V28G-I49A-V82G-N108Y、V28G-I49G-V82T-N108Y、V28G-I49G-V82G-N108Y、A48G-I49A-V82T-N108Y、A48G-I49A-V82G-N108Y、A48G-I49G-V82T-N108Y、A48G-I49G-V82G-N108Y、V28G-A48G-I49A-N108H、V28G-A48G-I49G-N108H、V28G-A48G-V82T-N108H、V28G-A48G-V82G-N108H、V28G-I49A-V82T-N108H、V28G-I49A-V82G-N108H、V28G-I49G-V82T-N108H、V28G-I49G-V82G-N108H、A48G-I49A-V82T-N108H、A48G-I49A-V82G-N108H、A48G-I49G-V82T-N108H、A48G-I49G-V82G-N108H；
或，所述突变型TadA-8e是指TadA-8e中氨基酸进行以下五点突变后所得的脱氨酶：V28G-A48G-I49A-V82T-N108Y、V28G-A48G-I49 G-V82T-N108Y、V28G-A48G-I49A-V82G-N108Y、V28G-A48G-I49G-V82G-N108Y、V28G-A48G-I49A-V82T-N108H、V28G-A48G-I49G-V82T-N108H、V28G-A48G-I49A-V82G-N108H、V28G-A48G-I49G-V82G-N108H。
本发明第八方面，提供腺嘌呤脱氨酶TadA突变蛋白，所述腺嘌呤脱氨酶TadA突变蛋白为非天然蛋白，所述腺嘌呤脱氨酶TadA突变蛋白是指TadA-X中与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺的脱氨酶；
TadA-X是指不同来源的TadA；优选地，TadA-X突变体有多组，其氨基酸序列分别如SEQ ID NO.5-SEQ ID NO.57中的任一所示。
进一步地，所述的突变型TadA-8e蛋白或所述的腺嘌呤脱氨酶TadA突变蛋白为单体或二聚体。
本发明第九方面，提供一种多核苷酸，编码所述碱基编辑器中的融合蛋白。
或，多核苷酸，编码所述的突变型TadA-8e蛋白或所述的腺嘌呤脱氨酶TadA突变蛋白。
本发明第十方面，提供一种载体，含有所述的多核苷酸。
本发明第十一方面，提供一种宿主细胞，含所述碱基编辑器，或含有所述的突变型TadA-8e蛋白或所述的腺嘌呤脱氨酶TadA突变蛋白，或含有所述载体。
本发明第十二方面，提供一种试剂盒，包含用于构建所述碱基编辑器的试剂。
与现有技术相比，本发明具有以下有益效果：
(1)本发明中开发的ABE工具在介导A·T--G·C突变时有更高的特异性，其介导C·G--T·A突变的能力被明显抑制，甚至清除。
(2)本发明开发的CBE工具在介导C·G--T·A突变时有更高的精确度，且RNA和DNA水平的脱靶效应有明显降低，甚至消除。
(3)本发明中开发的ACBE工具，只需要单一的脱氨酶，其体积更小，且在RNA和DNA水平的脱靶效应有明显降低，甚至消除，具有更高的应用价值。
附图说明
图1为TadA-8e融合于nSpCas9蛋白不同位置(包括N-、C-和内部的融合位点)时所得的多种碱基编辑器ABEs在sgRNA-S1位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率示意图。
图2为NLS-TadA-8e-linker融合于nSpCas9蛋白不同位置(包括N-、C-和内部的融合位点)时所得的多种碱基编辑器ABEs在sgRNA-S1位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率示意图。
图3a为1249-NLS-TadA-8e-linker和1249-TadA-8e在sgRNA-S1位点处对第四位碱基C(C4)和第六位碱基C(C6)的编辑效率(C·G--T·A突变率)示意图。
图3b为V28、V30、N46、A48、I49、V82、F84、N108、K110、R111在TadA-8e结构(PDB：6VPC)中的空间位置示意图；图中标示的氨基酸残基位于TadA-8e的活性口袋。
图3c为增强1249-NLS-TadA-8e-linker介导C·G--T·A突变能力和对胞嘧啶(C)偏好性的单点突变。柱状图从左至右分别为1249-NLS-TadA-8e-linker突变体的胞嘧啶(C)偏好性(胞嘧啶(C)偏好性＝C·G--T·A突变率/A·T--G·C突变率)，胞嘧啶(C)相对编辑活性(胞嘧啶(C)相对编辑活性＝突变型/无突变型1249-NLS-TadA-8e-linker的C·G--T·A突变率)以及腺嘌呤(A)相对编辑活性(腺嘌呤(A)相对编辑活性＝突变型/无突变型1249-NLS-TadA-8e-linker的A·T--G·C突变率)。
图4为1249-NLS-TadA-8e-linker中TadA-8e在V28(A)和V30(B)饱和突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中深色框标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力和对胞嘧啶(C)的偏好性。
图5为1249-NLS-TadA-8e-linker中TadA-8e在N46(A)和A48(B)饱和突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中深色框标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力和对胞嘧啶(C)的偏好性。
图6为1249-NLS-TadA-8e-linker中TadA-8e在I49(A)和V82(B)饱和突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中深色框标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力和对胞嘧啶(C)的偏好性。
图7为1249-NLS-TadA-8e-linker中TadA-8e在F84(A)和N108(B)饱和突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中深色框标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力和对胞嘧啶(C)的偏好性。
图8为1249-NLS-TadA-8e-linker中TadA-8e在K110(A)和R111(B)饱和突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图。
图9为1249-NLS-TadA-8e-linker中TadA-8e在V28位点饱和突变后在sgRNA-S1位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率(热图)示意图。
图10为1249-NLS-TadA-8e-linker中TadA-8e在V30位点饱和突变后在sgRNA-S1位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率(热图)示意图。
图11为1249-NLS-TadA-8e-linker中TadA-8e在N46位点饱和突变后在sgRNA-S1位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率(热图)示意图。
图12为1249-NLS-TadA-8e-linker中TadA-8e在A48位点饱和突变后在sgRNA-S1位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率(热图)示意图。
图13为1249-NLS-TadA-8e-linker中TadA-8e在I49位点饱和突变后在sgRNA-S1位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率(热图)示意图。
图14为1249-NLS-TadA-8e-linker中TadA-8e在V82位点饱和突变后在sgRNA-S1位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率(热图)示意图。
图15为1249-NLS-TadA-8e-linker中TadA-8e在F84位点饱和突变后在sgRNA-S1位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率(热图)示意图。
图16为1249-NLS-TadA-8e-linker中TadA-8e在N108位点饱和突变后在sgRNA-S1位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率(热图)示意图。
图17为1249-NLS-TadA-8e-linker中TadA-8e在K110位点饱和突变后在sgRNA-S1位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率(热图)示意图。
图18为1249-NLS-TadA-8e-linker中TadA-8e在R111位点饱和突变后在sgRNA-S1位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率(热图)示意图。
图19为1249-NLS-TadA-8e-linker中TadA-8e引入V28G-X(a)和V28W-X(b)双点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；所述V28G-X表示在第28位氨基酸由缬氨酸突变为甘氨酸的基础上其它氨基酸位点还存在突变X。
图中深色框标示的点突变型1249-NLS-TadA-8e-linker几乎无A·T--G·C突变能力，但其C·G--T·A突变能力仍不弱于野生型1249-NLS-TadA-8e-linker。
图20为1249-NLS-TadA-8e-linker中TadA-8e引入N46P-X(a)、N46C-X(b)和N46L-X(c)双点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中深色框标示的点突变型1249-NLS-TadA-8e-linker几乎无A·T--G·C突变能力，但其C·G--T·A突变能力仍不弱于野生型1249-NLS-TadA-8e-linker。
图21为1249-NLS-TadA-8e-linker中TadA-8e引入A48G-X(a)、I49G-X(b)和I49A-X(c)双点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中深色框标示的点突变型1249-NLS-TadA-8e-linker几乎无A·T--G·C突变能力，但其C·G--T·A突变能力仍不弱于野生型1249-NLS-TadA-8e-linker。
图22为1249-NLS-TadA-8e-linker中TadA-8e引入V82G-X(a)、V82T-X(b)和F84M/Y-X(c)双点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中深色框标示的点突变型1249-NLS-TadA-8e-linker几乎无A·T--G·C突变能力，但其C·G--T·A突变能力仍不弱于野生型1249-NLS-TadA-8e-linker。
图23为1249-NLS-TadA-8e-linker中TadA-8e引入N108Y-X(a)、N108H-X(b)和N108M/C/D/Q/S-X(C)双点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中深色框标示的点突变型1249-NLS-TadA-8e-linker几乎无A·T--G·C突变能力，但其C·G--T·A突变能力仍不弱于野生型1249-NLS-TadA-8e-linker。
图24为1249-NLS-TadA-8e-linker中TadA-8e引入N108M-X(a)、N108C-X(b)、N108D-X(c)、N108S-X(d)和N108Q-X(e)双点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中深色框标示的点突变型1249-NLS-TadA-8e-linker几乎无A·T--G·C突变能力，但其C·G--T·A突变能力仍不弱于野生型1249-NLS-TadA-8e-linker。
图25与单点突变相比，能进一步增强1249-NLS-TadA-8e-linker对胞嘧啶(C)的编辑能力和偏好性的双点突变汇总。
图26为1249-NLS-TadA-8e-linker中TadA-8e引入各类三点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图。图中深色框标示的点突变型1249-NLS-TadA-8e-linker的A·T--G·C突变能力显著降低，但其C·G--T·A突变能力仍不弱于野生型1249-NLS-TadA-8e-linker。
图27为1249-NLS-TadA-8e-linker中TadA-8e引入V28G-X-Y(A)和A48G-X-Y(B)三点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；所述V28G-X-Y表示在第28位氨基酸由缬氨酸突变为甘氨酸的基础上其它两处氨基酸位点还存在突变X和突变Y；
图中红色箭头标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力，其中深红色箭头为优选点突变。X和Y指代其它位点的氨基酸突变。
图28为1249-NLS-TadA-8e-linker中TadA-8e引入I49A-X-Y(A)和I49G-X-Y(B)三点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中红色箭头标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力，其中深红色箭头为优选点突变。X和Y指代其它位点的氨基酸突变。
图29为1249-NLS-TadA-8e-linker中TadA-8e引入V82G-X-Y(A)和V82T-X-Y(B)三点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中红色箭头标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力，其中深红色箭头为优选点突变。X和Y指代其它位点的氨基酸突变。
图30为1249-NLS-TadA-8e-linker中TadA-8e引入N108H-X-Y(A)和N108Y-X-Y(B)三点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中红色箭头标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力，其中深红色箭头为优选点突变。X和Y指代其它位点的氨基酸突变。
图31为1249-NLS-TadA-8e-linker中TadA-8e引入V28G-X-Y-Z(A)和A48G-X-Y-Z(B)四点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
示意的，所述V28G-X-Y-Z表示在第28位氨基酸由缬氨酸突变为甘氨酸的基础上还存在突变X、突变Y和突变Z；
图中红色箭头标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力，其中深红色箭头为优选点突变。X、Y和Z指代其它位点的氨基酸突变。
图32为1249-NLS-TadA-8e-linker中TadA-8e引入I49A-X-Y-Z(A)和I49G-X-Y-Z(B)四点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中红色箭头标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力，其中深红色箭头为优选点突变。X、Y和Z指代其它位点的氨基酸突变。
图33为1249-NLS-TadA-8e-linker中TadA-8e引入V82G-X-Y-Z(A)和V82T-X-Y-Z(B)四点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中红色箭头标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力，其中深红色箭头为优选点突变。X、Y和Z指代其它位点的氨基酸突变。
图34为1249-NLS-TadA-8e-linker中TadA-8e引入N108H-X-Y-Z(A)和N108Y-X-Y-Z(B)四点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中红色箭头标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力，其中深红色箭头为优选点突变。X、Y和Z指代其它位点的氨基酸突变。
图35为1249-NLS-TadA-8e-linker中TadA-8e引入V28G-X-Y-Z-U(A)、A48G-X-Y-Z-U(B)、I49A-X-Y-Z-U(C)和I49G-X-Y-Z-U(D)五点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
示意的，所述V28G-X-Y-Z-U表示在第28位氨基酸由缬氨酸突变为甘氨酸的基础上还存在突变X、突变Y、突变Z和突变U；
图中红色箭头标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力，其中深红色箭头为优选点突变。X、Y、Z和U指代其它位点的氨基酸突变。
图36为1249-NLS-TadA-8e-linker中TadA-8e引入V82G-X-Y-Z-U(A)、V82T-X-Y-Z-U(B)、N108H-X-Y-Z-U(C)和N108Y-X-Y-Z-U(D)四点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图；
图中红色箭头标示的点突变能明显增强1249-NLS-TadA-8e-linker的C·G--T·A突变能力，其中深红色箭头为优选点突变。X、Y、Z和U指代其它位点的氨基酸突变。
图37为1249-NLS-TadA-8e-linker中TadA-8e引入各类四点和五点突变后在sgRNA-S1位点处介导A·T--G·C突变能力和C·G--T·A突变能力改变示意图。图中深色框标示的点突变型1249-NLS-TadA-8e-linker的A·T--G·C突变能力显著降低，但其C·G--T·A突变能力仍不弱于野生型1249-NLS-TadA-8e-linker。
图38为不同类型的TadA融合于nSpCas9蛋白1249位点时所得的多种碱基编辑器ABEs在sgRNA-1、sgRNA-18、sgRNA-PT14、sgRNA-HPE6和sgRNA-S5位点处对碱基A(A·T--G·C突变)和C(C·G--T·A突变率)的编辑效率示意图(实验结果为两次独立生物重复实验的平均值)。
图39为不同类型的碱基编辑器在RNA水平和DNA水平(非Cas依赖型)的脱靶效应示意图。
a.各碱基编辑器在RNA水平的A-I脱靶数目。
b.各碱基编辑器在RNA水平的C-U脱靶数目。
c.各碱基编辑器在在DNA水平的A-G脱靶效应(非Cas9依赖型)。共四种脱靶位点R-loop 1,2,3,4用于检测。
d.各碱基编辑器在在DNA水平的C-T脱靶效应(非Cas9依赖型)。共四种脱靶位点R-loop 1,2,3,4用于检测。
*指P<0.05；**指P<0.01；***指指P<0.001。(结果来自三次独立生物重复实验)。
图40为优选碱基编辑器介导C·G--T·A突变和A·T--G·C突变的活性窗口及介导C·G--T·A突变时对碱基的偏好性示意图；
图中为1249-NLS-TadA-8e-linker介导C·G--T·A突变和A·T--G·C突变的活性窗口(a)及介导C·G--T·A突变时对碱基的偏好性(b)；1249-NLS-TadA-8e-linker(V28G)介导C·G--T·A突变和A·T--G·C突变的活性窗口(c)及介导C·G--T·A突变时对碱基的偏好性(d)；1249-NLS-TadA-8e-linker(N108Y)介导C·G--T·A突变和A·T--G·C突变的活性窗口(e)及介导C·G--T·A突变时对碱基的偏好性(f)。
图41为优选碱基编辑器介导C·G--T·A突变和A·T--G·C突变的活性窗口及介导C·G--T·A突变时对碱基的偏好性示意图；
图中为1249-NLS-TadA-8e-linker(N46L)介导C·G--T·A突变和A·T--G·C突变的活性窗口(a)及介导C·G--T·A突变时对碱基的偏好性(b)；1249-NLS-TadA-8e-linker(N46C)介导C·G--T·A突变和A·T--G·C突变的活性窗口(c)及介导C·G--T·A突变时对碱基的偏好性(d)；1249-NLS-TadA-8e-linker(N46P)介导C·G--T·A突变和A·T--G·C突变的活性窗口(e)及介导C·G--T·A突变时对碱基的偏好性(f)。
图42为优选碱基编辑器介导C·G--T·A突变和A·T--G·C突变的活性窗口及介导C·G--T·A突变时对碱基的偏好性示意图；
图中为1249-NLS-TadA-8e-linker(V28G-N46L)介导C·G--T·A突变和A·T--G·C突变的活性窗口(a)及介导C·G--T·A突变时对碱基的偏好性(b)；1249-NLS-TadA-8e-linker(V28G-N46C)介导C·G--T·A突变和A·T--G·C突变的活性窗口(c)及介导C·G--T·A突变时对碱基的偏好性(d)；1249-NLS-TadA-8e-linker(V28G-N46P)介导C·G--T·A突变和A·T--G·C突变的活性窗口(e)及介导C·G--T·A突变时对碱基的偏好性(f)。
图43为优选碱基编辑器介导C·G--T·A突变和A·T--G·C突变的活性窗口及介导C·G--T·A突变时对碱基的偏好性示意图；
图中为1249-NLS-TadA-8e-linker(N46C-N108Y)介导C·G--T·A突变和A·T--G·C突变的活性窗口(a)及介导C·G--T·A突变时对碱基的偏好性(b)；1249-NLS-TadA-8e-linker(N46P-N108S)介导C·G--T·A突变和A·T--G·C突变的活性窗口(c)及介导C·G--T·A突变时对碱基的偏好性(d)。
图44为优选碱基编辑器介导C·G--T·A突变和A·T--G·C突变的活性窗口及介导C·G--T·A突变时对碱基的偏好性示意图；
图中为1249-NLS-TadA-8e-linker(F84M-N108Y)介导C·G--T·A突变和A·T--G·C突变的活性窗口(a)及介导C·G--T·A突变时对碱基的偏好性(b)；1249-NLS-TadA-8e-linker(F84M-N108C)介导C·G--T·A突变和A·T--G·C突变的活性窗口(c)及介导C·G--T·A突变时对碱基的偏好性(d)。
图45为优选碱基编辑器介导C·G--T·A突变和A·T--G·C突变的活性窗口及介导C·G--T·A突变时对碱基的偏好性示意图；
图中为1249-NLS-Q13XZ4(VN)-linker介导C·G--T·A突变和A·T--G·C突变的活性窗口(a)及介导C·G--T·A突变时对碱基的偏好性(b)；1249-NLS-B3PCY2(VN)-linker介导C·G--T·A突变和A·T--G·C突变的活性窗口(c)及介导C·G--T·A突变时对碱基的偏好性(d)；1249-NLS-E8WVH3(VN)-linker介导C·G--T·A突变和A·T--G·C突变的活性窗口(e)及介导C·G--T·A突变时对碱基的偏好性(f)。
图46为优选碱基编辑器介导C·G--T·A突变和A·T--G·C突变的活性窗口示意图；
图中为1249-NLS-B5ZCW4(VN)-linker介导C·G--T·A突变和A·T--G·C突变的活性窗口(a)；1249-NLS-Q57LE3(VN)-linker介导C·G--T·A突变和A·T--G·C突变的活性窗口(b)；1249-NLS-Q99W51(VN)-linker介导C·G--T·A突变和A·T--G·C突变的活性窗口(c)。
具体实施方式
下面结合附图和具体实施例对本发明进行详细说明。
实施例1
TadA-8e为基础的碱基编辑器ABE构建
已知ecTadA*可放置于nSpCas9(D10A)的N-末端、C-末端或是内部的融合位点用于功能性ABE的构建(Nat Commun,2020.11(1):5827)。本实施例中，为获得编辑活性更高和编辑窗口更多样化的ABE工具，根据已有研究(Nat Commun,2020.11(1):5827)，将TadA-8e(SEQ ID NO.1)分别置于SV40NLS-linker-nSpCas9-nucleoplasmin NLS(SEQ.ID NO.2，序列中2-17位为SV40NLS-linker序列，18-1384位对应nSpCas9(D10A)的2-1368位序列，1385-1400位为nucleoplasmin NLS序列)的N-末端、C末端(C末端有额外的link序列(SEQ IDNO.3))和多个内部融合的位点(表1)，构建出多种ABE工具表达载体ABE-Internal-8e(通过2A-EGFP元件同时表达EGFP)。此外，考虑到蛋白接头(linker)或能改善内部融合后融合蛋白的构象，申请人在TadA-8e的5’和3’分别添加NLS序列和linker序列获得NLS-TadA-8e-linker(SEQ ID NO.4)，并分别置于nSpCas9的N-末端和多个内部融合的位点，构建出多种ABE工具表达载体ABE-Internal-8e(通过2A-EGFP元件同时表达EGFP)。为评估ABE-Internal-8e的编辑活性，将不同的ABE-Internal-8e与单向导RNA(sgRNA)(sgRNA同时表达尿嘧啶糖基化酶抑制因子UGI和2A-mCherry)共转染至培养的293T细胞。细胞培养72小时后，通过FACS收集同时表达EGFP和mCherry的双阳性细胞。本实施例中使用的sgRNA为S1(序列为GAACACAAAGCATAGACTGC)。
表1 nspCas9融合位点
收集的双阳性细胞抽提基因组，使用可特异性扩增sgRNA-S1位点信息的引物进行定向PCR。PCR产物通过sanger测序验证不同ABE-Internal-8e在S1位点的编辑效率。结果显示，与TadA-8e相比，NLS-TadA-8e-linker能增强多种ABE-Internal-8e的编辑活性，并在一定程度上拓展编辑窗口(图1-2)。此外，结果显示，在nSpCas9第1249位融合位点构建的1249-TadA-8e和1249-NLS-TadA-8e-linker具有一定的C·G--T·A突变活性，其中，1249-NLS-TadA-8e-linker在C4位点的C·G--T·A突变活性高于1249-TadA-8e(图3a)。
实施例2
对1249-NLS-TadA-8e-linker工具中的TadA-8e进行点突变筛选
本实施例中，根据TadA-8e的结构信息(PDB：6VPC)，选取TadA-8e中位于酶活性口袋中与反应底物邻近的数个氨基酸(第28、30、46、48、49、82、84、108、110或111位的氨基酸)中任一个进行饱和突变(图3b)，分析不同点突变对1249-NLS-TadA-8e-linker的碱基编辑活性的影响。
为评估不同点突变对1249-NLS-TadA-8e-linker的碱基编辑活性的影响，将1249-NLS-TadA-8e-linker突变体(通过2A-EGFP元件同时表达EGFP)与单向导RNA(sgRNA)(sgRNA同时表达尿嘧啶糖基化酶抑制因子UGI和2A-mCherry)共转染至培养的293T细胞。细胞培养72小时后，通过FACS收集同时表达EGFP和mCherry的双阳性细胞(方案1，FACS)或直接收取细胞(方案2，NO.FACS)。本实施例中使用的sgRNA为S1(序列为GAACACAAAGCATAGACTGC)。
收集的细胞抽提基因组，使用可特异性扩增sgRNA-S1位点信息的引物进行定向PCR。PCR产物通过sanger测序验证1249-NLS-TadA-8e-linker各突变体的碱基编辑活性改变。
本实施例中，对TadA-8e单点突变后的碱基编辑器的A·T--G·C突变和C·G--T·A突变的能力进行分析。
分析指标为：(1)点突变后，第三位腺嘌呤(A3)、第四位胞嘧啶(C4)、第五位腺嘌呤(A5)、第六位胞嘧啶(C6)、第七位腺嘌呤(A7)的编辑效率变化(PAM为21-23位点)(相对效率＝点突变组/无突变组，相对效率>1表明突变型1249-NLS-TadA-8e-linker的A·T--G·C突变或C·G--T·A突变的能力增强)；
(2)根据1249-NLS-TadA-8e-linker对S1位点中各碱基的编辑情况可知，其对A3、A5、A7、A8和A9的编辑效率最高且各位点效率接近，因此选取A5作为1249-NLS-TadA-8e-linker介导A·T--G·C突变能力的代表。点突变后，C4突变效率与A5突变效率的比值变化(C4/A5)，以及C6突变效率与A5突变效率的比值变化(C6/A5)可反映1249-NLS-TadA-8e-linker对胞嘧啶(C)和腺嘌呤(A)两种碱基偏好性的改变。C4/A5的相对效率＝(点突变组C4/A5)/(无突变组C4/A5)，C6/A5的相对效率＝(点突变组C6/A5)/(无突变组C6/A5)，相对效率>1表明突变型1249-NLS-TadA-8e-linker对胞嘧啶(C)的编辑能力和偏好性增强。
结合以上两种指标，申请人对可促进1249-NLS-TadA-8e-linker介导C·G--T·A突变能力和增加胞嘧啶(C)偏好性的点突变进行筛选。结果显示，V28W、V28G、N46L、N46P、N46C、A48C、A48G、A48N、A48S、I49D、I49E、I49H、I49N、I49P、I49Q、I49V、I49W、I49Y、I49R、I49A、I49T、I49G、I49L、I49S、I49M、I49C、I49F、I49K、V82G、V82A、V82T、V82C、V82S、V82D、F84M、N108H、N108M、N108C、N108D、N108F、N108Q、N108S、N108Y点突变能增强1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力及对胞嘧啶(C)的编辑偏好性(图3c)。其余点突变对1249-NLS-TadA-8e-linker功能的影响可参见图4-图8。
此外，对点突变后1249-NLS-TadA-8e-linker编辑S1位点各碱基的效率(方案1，FACS分选部分)进行分析后发现(效率热图)，V28位点的不同点突变中，V28D、V28E、V28K能有效提高1249-NLS-TadA-8e-linker的碱基编辑特异性和精确度，V28D、V28E突变在A5位点(PAM位点编号为21-23)具有明显的编辑活性(A·T--G·C突变)；V28K突变则在A3-A7具有明显的编辑活性，其中在A5-A7具有最高的编辑效率(A·T--G·C突变)。此外，V28M、V28R、V28D、V28Q、V28K、V28L、V28E突变均可显著抑制1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(<10％的编辑效率)。V28W、V28A、V28G则能明显提高1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(提升超过20％)，其中V28G还能同时抑制1249-NLS-TadA-8e-linker介导A·T--G·C突变的能力并导致其活性窗口变窄(图9)。
V30位点的不同点突变中，V30D、V30G、V30W、V30E、V30N、V30Q、V30Y、V30T、V30S均能明显提高1249-NLS-TadA-8e-linker的碱基编辑特异性和精确度。V30D、V30W、V30E的核心活性窗口为A5；V30N、V30Q、V30Y的活性窗口为A5-A7；V30T、V30S的活性窗口为A3-A7。此外，V30A、V30D、V30T、V30G、V30P、V30W、V30E、V30K、V30N、V30Q、V30R、V30S、V30Y能显著抑制1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(<10％的编辑效率)，其中V30G、V30P、V30K、V30R还同时抑制了1249-NLS-TadA-8e-linker介导A·T--G·C突变的能力(图10)。
N46位点的所有19种点突变均导致1249-NLS-TadA-8e-linker活性窗口的缩小。其中N46H、N46K、N46Y、N46F只在A5有较低的编辑活性(～30％)。N46G、N46M的活性窗口为A5-A7，其中A5为核心活性窗口(A5编辑效率～60％，A7约～20％)。N46E、N46Q、N46D的活性窗口为A3-A7。N46H、N46K、N46Y、N46F、N46G、N46M、N46E、N46Q、N46D能显著抑制1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(<10％的编辑效率)。有趣的是，N46P在S1位点仅能介导C4-C6位点的C·G--T·A突变。N46A、N46V的活性窗口位于C4-A5，其中N46A活性较低(<20％)；N46L的活性窗口位于C4-A5-C6；N46C的活性窗口位于C4-A5-C6-A7。N46A、N46V、N46L、N46C突变后的1249-NLS-TadA-8e-linker具有更窄的活性窗口，并具备介导C·G--T·A突变和A·T--G·C突变的两种能力。N46A、N46R、N46W点突变后，1249-NLS-TadA-8e-linker基本不具备任何编辑活性。N46S、N46T具有更窄的活性窗口，具备介导C·G--T·A突变和A·T--G·C突变的两种能力，但A·T--G·C突变的能力显著强于C·G--T·A突变的能力(图11)。
A48位点的不同点突变中，A48D、A48E能明显提高1249-NLS-TadA-8e-linker的碱基编辑特异性和精确度,活性窗口主要位于A3-A7。A48Y、A48W、A48E、A48D、A48F、A48Q能显著抑制1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(<10％的编辑效率)。A48G、A48S、A48N能明显提高1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(提升超过20％)。A48L点突变导致1249-NLS-TadA-8e-linker基本不具备任何编辑活性(图12)。
I49位点的不同点突变中，所有19种点突变均能在一定程度上提高1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力，且大部分点突变能明显提高1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(提升超过20％)。(图13)
V82位点的不同点突变中，V82F可显著抑制1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(<10％的编辑效率)。V82K、V82W、V82R、V82Y导致1249-NLS-TadA-8e-linker基本不具备任何编辑活性。V82A、V82T、V82C、V82S、V82D能明显提高1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(提升超过20％)(图14)。
F84位点的不同点突变中，F84D、F84R能明显提高1249-NLS-TadA-8e-linker的碱基编辑特异性和精确度,活性窗口主要位于A5；F84A、F84E、F84P能明显提高1249-NLS-TadA-8e-linker的碱基编辑特异性和精确度，活性窗口主要位于A3-A7。此外，F84K、F84C、F84H、F84I、F84S、F84T、F84V、F84Q、F84N、F84G能导致其活性窗口变窄。F84Y、F84E、F84R可显著抑制1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(<10％的编辑效率)。F84M能明显提高1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(提升超过20％)(图15)。
N108位点的不同点突变中，除N108V外，均能导致1249-NLS-TadA-8e-linker的活性窗口明显变窄。N108I、N108L、N108A、N108T能明显提高1249-NLS-TadA-8e-linker的碱基编辑特异性和精确度,活性窗口主要位于A5，N108R能明显提高1249-NLS-TadA-8e-linker的碱基编辑特异性和精确度,活性窗口主要位于A3-A7。N108I、N108L、N108R、N108A、N108T可显著抑制1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(<10％的编辑效率)。N108H、N108M、N108C、N108D、N108F、N108Q、N108S、N108Y能明显提高1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(提升超过20％)(图16)。
K110位点的不同点突变中，K110D、K110W能明显提高1249-NLS-TadA-8e-linker的碱基编辑特异性和精确度,活性窗口主要位于A2-A8。K110E、K110W可显著抑制1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(<10％的编辑效率)(图17)。
R111位点的不同点突变中，R111P、R111E、R111T能明显提高1249-NLS-TadA-8e-linker的碱基编辑特异性和精确度，活性窗口主要位于A5。R111A、R111V、R111G、R111I、R111W能明显提高1249-NLS-TadA-8e-linker的碱基编辑特异性和精确度,活性窗口主要位于A3-A7。R111L能明显提高1249-NLS-TadA-8e-linker的碱基编辑特异性和精确度,活性窗口主要位于A2-A7。R111P、R111D、R111L、R111E、R111I、R111T、R111W可显著抑制1249-NLS-TadA-8e-linker介导C·G--T·A突变的能力(<10％的编辑效率)(图18)。
实施例3
对1249-NLS-TadA-8e-linker工具中的TadA-8e进行点突变组合筛选(双点突变)
本实施例中，根据TadA-8e的结构信息(PDB：6VPC)，选取能增强TadA-8e介导C·G--T·A突变能力的关键点突变V28G、V28W(优选V28G)，N46P、N46C、N46L，A48G，I49A、I49G，V82T，F84M，N108H、N108M、N108C、N108D、N108Q、N108S、N108Y(优选N108H、N108Y)，以及能降低RNA脱靶风险的V82G点突变，通过构建双点突变型1249-NLS-TadA-8e-linker，分析不同双点突变组合对1249-NLS-TadA-8e-linker碱基编辑活性的影响。
为评估不同点突变对1249-NLS-TadA-8e-linker的碱基编辑活性的影响，将1249-NLS-TadA-8e-linker突变体(通过2A-EGFP元件同时表达EGFP)与单向导RNA(sgRNA)(sgRNA同时表达尿嘧啶糖基化酶抑制因子UGI和2A-mCherry)共转染至培养的293T细胞。细胞培养72小时后，细胞培养72小时后，通过FACS收集同时表达EGFP和mCherry的双阳性细胞(方案1)，或直接收集所有细胞(方案2)。本实施例中使用的sgRNA为S1(序列为GAACACAAAGCATAGACTGC)。
收集好的细胞抽提基因组，使用可特异性扩增sgRNA-S1位点信息的引物进行定向PCR。PCR产物通过sanger测序验证1249-NLS-TadA-8e-linker各突变体的活性变化。
分析指标为：(1)双点突变后，第四位胞嘧啶(C4)、第五位腺嘌呤(A5)、第六位胞嘧啶(C6)的编辑效率变化(PAM为21-23位点)(相对效率＝双点突变组/无突变组，双点突变相对效率>单点突变相对效率时表明，与单点突变相比，双点突变后1249-NLS-TadA-8e-linker的A·T--G·C突变或C·G--T·A突变的能力增强)；
(2)双点突变后，C4/A5的相对效率＝(双点突变组C4/A5)/(无突变组C4/A5)，C6/A5的相对效率＝(双点突变组C6/A5)/(无突变组C6/A5)，双点突变相对效率>单点突变相对效率时表明，与单点突变相比，双点突变后1249-NLS-TadA-8e-linker对胞嘧啶(C)的编辑能力和偏好性增强。
综合以上两种指标，申请人对可促进1249-NLS-TadA-8e-linker介导C·G--T·A突变能力的双点突变进行筛选(图19-24)。其中双点突变V28G-N108Y，V28G-N108C，V28G-N108H，V28G-N46L，V28G-N46C，V28G-N46P，V28W-N46L，V28W-N46C，V28W-N46P，N46P-N108Y，N46P-N108S，N46P-N108Q，N46P-N108D，N46P-N108C，N46P-N108H，N46P-F84M，N46P-I49A，N46P-I49G，N46P-A48G，N46C-N108Y，N46C-N108S，N46C-N108Q，N46C-N108M，N46C-N108C，N46C-N108H，N46L-N108Y，N46L-N108Q，N46L-N108D，N46L-N108M，N46L-N108H，N46L-F84M，N46L-I49A，N46L-I49G，N46L-V82G，I49A-N108Y能几乎完全抑制1249-NLS-TadA-8e-linker的A·T--G·C突变能力，同时保持不弱于1249-NLS-TadA-8e-linker的C·G--T·A突变能力。此外，与相应的单点突变相比，V28G-N46C、V28G-N46P、V28G-A48G、V28G-I49A、V28G-I49G、V28G-V82T、V28G-F84Y、V28W-N108H、V28W-N46P、V28W-N46C、N46C-V82G、N46C-V82T、N46C-F84M、N46L-A48G、N46L-V82T、N46L-F84M、N46P-F84M、N46P-N108H、N46P-N108M、N46P-N108C、N46P-N108Q、N46P-N108S、A48G-N108H、I49A-N108H、I49G-N108H、I49G-N108Y、I49G-V82G、I49G-V82T、I49G-F84Y、V82G-N108Y、V82T-N108H、V82T-N108C、F84M-N108H、F84M-N108M、F84M-N108Q双点突变能进一步增强1249-NLS-TadA-8e-linker对胞嘧啶(C)的编辑能力和偏好性(图25)。
实施例4
对1249-NLS-TadA-8e-linker工具中的TadA-8e进行点突变组合筛选(三点突变)
本实施例中，根据TadA-8e的结构信息(PDB：6VPC)，以及单点突变和三点突变的结果，选取能增强TadA-8e介导C·G--T·A突变能力的关键点突变V28G，A48G，I49A、I49G，V82T，N108H、N108Y，以及能降低RNA脱靶风险的V82G点突变，通过构建三点突变型1249-NLS-TadA-8e-linker，分析不同三点突变组合对1249-NLS-TadA-8e-linker的碱基编辑活性的影响。
为评估不同点突变对1249-NLS-TadA-8e-linker的碱基编辑活性的影响，将1249-NLS-TadA-8e-linker突变体(通过2A-EGFP元件同时表达EGFP)与单向导RNA(sgRNA)(sgRNA同时表达尿嘧啶糖基化酶抑制因子UGI和2A-mCherry)共转染至培养的293T细胞。细胞培养72小时后，细胞培养72小时后，通过FACS收集同时表达EGFP和mCherry的双阳性细胞(方案1)，或直接收集所有细胞(方案2)。本实施例中使用的sgRNA为S1(序列为GAACACAAAGCATAGACTGC)。
收集好的细胞抽提基因组，使用可特异性扩增sgRNA-S1位点信息的引物进行定向PCR。PCR产物通过sanger测序验证1249-NLS-TadA-8e-linker各突变体的活性变化。
分析指标为：(1)三点突变后，第四位胞嘧啶(C4)、第五位腺嘌呤(A5)、第六位胞嘧啶(C6)的编辑效率变化(PAM为21-23位点)(相对效率＝三点突变组/无突变组，相对效率>单点突变组表明，与单点突变相比，三点突变后1249-NLS-TadA-8e-linker的A·T--G·C突变或C·G--T·A突变的能力增强)；
(2)三点突变后，C4/A5的相对效率＝(三点突变组C4/A5)/(无突变组C4/A5)，C6/A5的相对效率＝(三点突变组C6/A5)/(无突变组C6/A5)，相对效率>单点突变组表明，与单点突变相比，三点突变后1249-NLS-TadA-8e-linker的C·G--T·A突变的能力增强。
综合以上两种指标，我们发现，显著抑制1249-NLS-TadA-8e-linker的A·T--G·C突变能力，同时保持或增强C·G--T·A突变能力的三点突变均包含有N108Y或N108H点突变。这类三点突变有I49A-V82T-N108Y、I49A-V82G1-N08Y、I49G-V82T-N108Y、I49G-V82G-N108Y、A48G-V82G-N108Y、A48G-I49G-N108Y、V28G-V82G-N108Y、V28G-V82G-N108H、V28G-I49A-N108Y、V28G-I49A-N108H、V28G-I49G-N108H、V28G-A48G-N108Y、V28G-A48G-N108H(图26)。
分析指标为：(1)三点突变后，第四位胞嘧啶(C4)、第五位腺嘌呤(A5)、第六位胞嘧啶(C6)的编辑效率变化(PAM为21-23位点)(三/单相对效率＝三点突变组/对应的单点突变组，相对效率>1表明，与单点突变相比，三点突变后1249-NLS-TadA-8e-linker的A·T--G·C突变或C·G--T·A突变的能力增强)；
(2)三点突变后，C4/A5的三/单相对效率＝(三点突变组C4/A5)/(对应的单点突变组C4/A5)，C6/A5的三/单相对效率＝(三点突变组C6/A5)/(对应的单点突变组C6/A5)，相对效率>1表明，与单点突变相比，三点突变后1249-NLS-TadA-8e-linker的C·G--T·A突变的能力增强。
结合以上两种指标，申请人对可促进1249-NLS-TadA-8e-linker介导C·G--T·A突变能力的三点突变进行筛选(图27-30)。结果显示，V28G-A48G-I49G、V28G-A48G-I49A、V28G-A48G-V82G、V28G-A48G-V82T、V28G-A48G-N108H、V28G-A48G-N108Y、V28G-I49G-V82G、V28G-I49G-V82T、V28G-I49G-N108H、V28G-I49G-N108Y、V28G-I49A-V82G、V28G-I49A-V82T、V28G-I49A-N108H、V28G-I49A-N108Y、I49G-V82G-N108H、I49G-V82G-N108Y、I49G-V82T-N108H、I49G-V82T-N108Y三点突变能进一步增强1249-NLS-TadA-8e-linker介导C·G--T·A突变能力。
实施例5
对1249-NLS-TadA-8e-linker工具中的TadA-8e进行点突变组合筛选(四点突变)
本实施例中，根据TadA-8e的结构信息(PDB：6VPC)，以及单点突变、二点突变和三点突变的结果，选取能增强TadA-8e介导C·G--T·A突变能力的关键点突变V28G，A48G，I49A、I49G，V82T，N108H、N108Y，以及能降低RNA脱靶风险的V82G点突变，通过构建1249-NLS-TadA-8e-linker的四点突变表达载体，分析不同四点突变组合对1249-NLS-TadA-8e-linker的碱基编辑活性的影响。
为评估不同点突变对1249-NLS-TadA-8e-linker的碱基编辑活性的影响，将1249-NLS-TadA-8e-linker突变体(通过2A-EGFP元件同时表达EGFP)与单向导RNA(sgRNA)(sgRNA同时表达尿嘧啶糖基化酶抑制因子UGI和2A-mCherry)共转染至培养的293T细胞。细胞培养72小时后，细胞培养72小时后，通过FACS收集同时表达EGFP和mCherry的双阳性细胞(方案1)，或直接收集所有细胞(方案2)。本实施例中使用的sgRNA为S1(序列为GAACACAAAGCATAGACTGC)。
收集好的细胞抽提基因组，使用可特异性扩增sgRNA-S1位点信息的引物进行定向PCR。PCR产物通过sanger测序验证1249-NLS-TadA-8e-linker各突变体的活性变化。
分析指标为：(1)四点突变后，第四位胞嘧啶(C4)、第五位腺嘌呤(A5)、第六位胞嘧啶(C6)的编辑效率变化(PAM为21-23位点)(四/单相对效率＝四点突变组/对应的单点突变组，相对效率>1表明，与单点突变相比，四点突变后1249-NLS-TadA-8e-linker的A·T--G·C突变或C·G--T·A突变的能力增强)；
(2)四点突变后，C4/A5的四/单相对效率＝(四点突变组C4/A5)/(对应的单点突变组C4/A5)，C6/A5的四/单相对效率＝(四点突变组C6/A5)/(对应的单点突变组C6/A5)，相对效率>1表明，与单点突变相比，四点突变后1249-NLS-TadA-8e-linker的C·G--T·A突变的能力增强。
结合以上两种指标，申请人对可促进1249-NLS-TadA-8e-linker介导C·G--T·A突变能力的四点突变进行筛选(图31-34)。结果显示，V28G-I49G-V82T-N108Y、A48G-I49A-V82T-N108H、A48G-I49G-V82G-N108H四点突变能进一步增强1249-NLS-TadA-8e-linker介导C·G--T·A突变能力。
实施例6
对1249-NLS-TadA-8e-linker工具中的TadA-8e进行点突变组合筛选(五点突变)
本实施例中，根据TadA-8e的结构信息(PDB：6VPC)，以及单点突变二点突变、三点突变和四点突变的结果，选取能增强TadA-8e介导C·G--T·A突变能力的关键点突变V28G，A48G，I49A、I49G，V82T，N108H、N108Y，以及能降低RNA脱靶风险的V82G点突变，通过构建1249-NLS-TadA-8e-linker的五点突变表达载体，分析不同五点突变组合对1249-NLS-TadA-8e-linker的碱基编辑活性的影响。
为评估不同点突变对1249-NLS-TadA-8e-linker的碱基编辑活性的影响，将1249-NLS-TadA-8e-linker突变体(通过2A-EGFP元件同时表达EGFP)与单向导RNA(sgRNA)(sgRNA同时表达尿嘧啶糖基化酶抑制因子UGI和2A-mCherry)共转染至培养的293T细胞。细胞培养72小时后，细胞培养72小时后，通过FACS收集同时表达EGFP和mCherry的双阳性细胞(方案1)，或直接收集所有细胞(方案2)。本实施例中使用的sgRNA为S1(序列为GAACACAAAGCATAGACTGC)。
收集好的细胞抽提基因组，使用可特异性扩增sgRNA-S1位点信息的引物进行定向PCR。PCR产物通过sanger测序验证1249-NLS-TadA-8e-linker各突变体的活性变化。
分析指标为：(1)五点突变后，第四位胞嘧啶(C4)、第五位腺嘌呤(A5)、第六位胞嘧啶(C6)的编辑效率变化(PAM为21-23位点)(五/单相对效率＝五点突变组/对应的单点突变组，相对效率>1表明，与单点突变相比，五点突变后1249-NLS-TadA-8e-linker的A·T--G·C突变或C·G--T·A突变的能力增强)；
(2)五点突变后，C4/A5的五/单相对效率＝(五点突变组C4/A5)/(对应的单点突变组C4/A5)，C6/A5的五/单相对效率＝(五点突变组C6/A5)/(对应的单点突变组C6/A5)，相对效率>1表明，与单点突变相比，五点突变后1249-NLS-TadA-8e-linker的C·G--T·A突变的能力增强。
结合以上两种指标，申请人对可促进1249-NLS-TadA-8e-linker介导C·G--T·A突变能力的五点突变进行筛选(图35-36)。结果显示，V28G-A48G-I49G-V82T-N108H、V28G-A48G-I49G-V82T-N108Y、V28G-A48G-I49A-V82T-N108H、V28G-A48G-I49A-V82T-N108Y五点突变能进一步增强1249-NLS-TadA-8e-linker介导C·G--T·A突变能力。
此外，我们对四点突变和五点突变的数据做了进一步分析，分别计算两种指标：
(1)多点突变后，第四位胞嘧啶(C4)、第五位腺嘌呤(A5)、第六位胞嘧啶(C6)的编辑效率变化(PAM为21-23位点)(相对效率＝多点突变组/无点突变组)。
(2)多点突变后，C4/A5的相对效率＝(多点突变组C4/A5)/(无点突变组C4/A5)，C6/A5的相对效率＝(多点突变组C6/A5)/(无点突变组C6/A5)。
综合以上两种指标，我们发现，显著抑制1249-NLS-TadA-8e-linker的A·T--G·C突变能力，同时保持或增强C·G--T·A突变能力的四/五点突变均包含有N108Y或N108H点突变。这表明N108位点的点突变对改变碱基编辑器编辑胞嘧啶(C)及腺嘌呤(A)的偏好性非常关键，N108Y/H点突变能明显增强1249-NLS-TadA-8e-linker编辑胞嘧啶(C)的偏好性，除A48G-I49A-V82T-N108H外，所有包含N108Y/H的四点突变和五点突变(V28G、A48G、I49A/G、V82T/G和N108H/Y的各类四点突变和五点突变组合)均可显著抑制1249-NLS-TadA-8e-linker的A·T--G·C突变能力。(图37)。
实施例7
自然界中其它类型TadA的筛选和碱基编辑器的构建
本实施例中，在自然界中筛选数十种不同类型的TadA，将其与ecTadA第106和第108对应的氨基酸位点分别突变为缬氨酸和天冬酰胺(TadA-X,SEQ ID NO.5-SEQ IDNO.57)。并以A106V和D108N位点突变的野生型ecTadA(SEQ ID NO.58)为对照。之后将1249-NLS-TadA-8e-linker中的TadA-8e替换为突变型TadA-X，构建1249-TadA-X表达载体，分析1249-TadA-X的碱基编辑能力和特性。
为评估1249-TadA-X的碱基编辑能力，将1249-TadA-X(通过2A-EGFP元件同时表达EGFP)与单向导RNA(sgRNA)(sgRNA同时表达尿嘧啶糖基化酶抑制因子UGI和2A-mCherry)共转染至培养的293T细胞。细胞培养72小时后，细胞培养72小时后，通过FACS收集同时表达EGFP和mCherry的双阳性细胞(方案1)。本实施例中方案1使用的sgRNA为S1(序列为GAACACAAAGCATAGACTGC)、S18(ACACACACACTTAGAATCTG)、PT14(序列为TCCAAACCCATATATACAGC)、HPE6(序列为GTATAATCCAAAGATGGTCA)、S5(GATGAGATAATGATGAGTCA)和PT14(TCCAAACCCATATATACAGC)，结果显示，1249-NLS-Q2LYC0(VN)-linker，1249-NLS-Q13XZ4(VN)-linker，1249-NLS-F2G306(VN)-linker，1249-NLS-B3PCY2(VN)-linker，1249-NLS-A8F050(N)-linker，1249-NLS-A5WFT9(VN)-linker，1249-NLS-G0JN58(VN)-linker，1249-NLS-B5ZCW4(VN)-linker，1249-NLS-E8WVH3(VN)-linker，1249-NLS-Q1AS41(VN)-linker，1249-NLS-P44931(VN)-linker，1249-NLS-C1DXN9(VN)-linker，1249-NLS-B2UR34(VN)-linker，1249-NLS-G0EXN4(VN)-linker，1249-NLS-E4TTF3(VN)-linker，1249-NLS-R4K908(VN)-linker，1249-NLS-E0TBL5(VN)-linker，1249-NLS-A1W869(VN)-linker，1249-NLS-D5W8R4(VN)-linker，1249-NLS-D5X1Y1(VN)-linker，1249-NLS-I3YB54(VN)-linker，1249-NLS-Q57LE3(VN)-linker，1249-NLS-Q99W51(VN)-linker，1249-NLS-E1TAF4(VN)-linker，1249-NLS-ecTadA(VN)-linker在至少一个位点处有明显的碱基编辑活性(>10％)。就碱基编辑的特异性而言，1249-NLS-B3PCY2(VN)-linker，1249-NLS-A5WFT9(VN)-linker，1249-NLS-C1DXN9(VN)-linker，1249-NLS-B2UR34(VN)-linker，1249-NLS-R4K908(VN)-linker，1249-NLS-D5W8R4(VN)-linker，1249-NLS-E1TAF4(VN)-linker主要介导C·G--T·A突变，主要用作CBE工具；1249-NLS-Q13XZ4(VN)-linker，1249-NLS-A8F050(N)-linker，1249-NLS-E8WVH3(VN)-linker，1249-NLS-P44931(VN)-linker，1249-NLS-G0EXN4(VN)-linker，1249-NLS-E0TBL5(VN)-linker，1249-NLS-A1W869(VN)-linker，1249-NLS-D5X1Y1(VN)-linker，1249-NLS-I3YB54(VN)-linker介导A·T--G·C和C·G--T·A突变的能力相当，具备做ACBE的潜力，1249-NLS-Q2LYC0(VN)-linker，1249-NLS-F2G306(VN)-linker，1249-NLS-G0JN58(VN)-linker，1249-NLS-B5ZCW4(VN)-linker，1249-NLS-Q1AS41(VN)-linker，1249-NLS-E4TTF3(VN)-linker，1249-NLS-Q57LE3(VN)-linker，1249-NLS-Q99W51(VN)-linker，1249-NLS-ecTadA(VN)-linker主要用作ABE工具(图38)。
实施例8
优选工具在RNA水平的脱靶效应分析
本实施例中，优选1249-NLS-TadA-8e-linker、1249-NLS-TadA-8e-linker(V28G)、1249-NLS-TadA-8e-linker(N46L)、1249-NLS-TadA-8e-linker(N46C)、1249-NLS-TadA-8e-linker(N46P)、1249-NLS-TadA-8e-linker(N108Y)、、1249-NLS-TadA-8e-linker(V28G-N46L)、1249-NLS-TadA-8e-linker(V28G-N46P)、1249-NLS-TadA-8e-linker(V28G-N46C)、1249-NLS-TadA-8e-linker(N46C-N108Y)、1249-NLS-TadA-8e-linker(N46P-N108S)、1249-NLS-TadA-8e-linker(F84M-N108Y)、1249-NLS-TadA-8e-linker(F84M-N108C)、1249-NLS-Q13XZ4(VN)-linker、1249-NLS-B3PCY2(VN)-linker、1249-NLS-B5ZCW4(VN)-linker、1249-NLS-E8WVH3(VN)-linker、1249-NLS-Q57LE3(VN)-linker、1249-NLS-Q99W51(VN)-linker、1249-NLS-ecTadA(VN)-linker以分析其在RNA水平的脱靶效应。其中1249-NLS-TadA-8e-linker(8E)为阳性对照；nCas9-D10A为阴性对照。
为评估各优选工具在RNA水平的脱靶效应，将各优选表达载体(通过2A-EGFP元件同时表达EGFP)与单向导RNA(sgRNA)(sgRNA同时表达尿嘧啶糖基化酶抑制因子UGI和2A-puromycin)共转染至培养的293T细胞。转染后24小时加入嘌呤霉素筛选阳性细胞。嘌呤霉素处理48小时后，收集细胞提取总RNA进行RNA-seq实验。本方案使用的sgRNA为sg-RNF(序列为GTCATCTTAGTCATTACCTG)。
RNA编辑是指在mRNA水平上改变遗传信息的过程。具体来说，它指的是mRNA分子中核苷酸的缺失、插入或化学修饰。这种修饰影响基因的表达，不同氨基酸的产生和开放阅读框的形成。在哺乳动物细胞中，主要存在两类RNA编辑，分别为A-I编辑和C-U编辑。这种RNA编辑是由RNA依赖的腺嘌呤脱氨酶介导的。
我们首先对A-I编辑结果进行分析，发现TadA-8e*点突变可显著降低1249-NLS-TadA-8e-linker在RNA水平的脱靶效应。点突变后，RNA水平的脱靶效应与本底(N.C.，仅转染sgRNA)及阴性对照组(Cas9-D10A+sgRNA)类似。此外，其它新筛选的功能性TadA为基础的碱基编辑器中，1249-NLS-B5ZCW4(VN)-linker和1249-NLS-B3PCY2(VN)-linker在RNA水平的脱靶效应与本底及阴性对照相似，1249-NLS-Q13XZ4(VN)-linker、1249-NLS-ecTadA(VN)-linker、1249-NLS-Q57LE3(VN)-linker和1249-NLS-E8WVH3(VN)-linker在RNA水平的脱靶效应比本底及阴性对照略高，但明显低于1249-NLS-TadA-8e-linker。1249-NLS-Q99W51(VN)-linker在RNA水平的脱靶效应明显高于1249-NLS-TadA-8e-linker(图39a)。
RNA水平的C-U编辑结果显示，各样品组与本底(N.C.，仅转染sgRNA)及阴性对照组(Cas9-D10A+sgRNA)类似，没有明显的增加(图39b)。
实施例9
部分工具在DNA水平的脱靶效应分析(非Cas依赖型脱靶效应)
本实施例中，优选1249-NLS-TadA-8e-linker(V28G)、1249-NLS-TadA-8e-linker(N46L)、1249-NLS-TadA-8e-linker(N46C)、1249-NLS-TadA-8e-linker(N46P)、1249-NLS-TadA-8e-linker(N108Y)、1249-NLS-TadA-8e-linker(V28G-N46L)、1249-NLS-TadA-8e-linker(V28G-N46P)、1249-NLS-TadA-8e-linker(V28G-N46C)、1249-NLS-TadA-8e-linker(N46C-N108Y)、1249-NLS-TadA-8e-linker(N46P-N108S)、1249-NLS-TadA-8e-linker(F84M-N108Y)、1249-NLS-TadA-8e-linker(F84M-N108C)、1249-NLS-Q13XZ4(VN)-linker、1249-NLS-B3PCY2(VN)-linker、1249-NLS-B5ZCW4(VN)-linker、1249-NLS-E8WVH3(VN)-linker、1249-NLS-Q57LE3(VN)-linker、1249-NLS-Q99W51(VN)-linker以分析其在DNA水平的脱靶效应(非Cas依赖型脱靶效应)。另选择1249-NLS-TadA-8e-linker(8e)为对照。
DNA水平的脱靶效应(非Cas依赖型脱靶效应)通过R-Loop实验进行分析[8]。其基本原理是，dSaCas9与靶向基因组特定位点(位点X)的sasgRNA(sasgRNA-X)在细胞中共表达，会在位点X处诱导形成单链DNA(ssDNA)区域；此时，细胞内共表达的碱基编辑器(ABE或CBE)因其自身对ssDNA的亲和力，便可能结合ssDNA并介导C·G--T·A(CBE)或A·T--G·C(ABE)突变。
人源HEK293T细胞系复苏培养后以30％-50％密度铺12孔平板，24小时后采用脂质体共转染碱基编辑器的各优选表达载体(通过2A-EGFP元件同时表达EGFP)和失活型SaCas9(dSaCas9)-sasgRNA共表达载体(同时表达红色荧光蛋白)，培养72小时后胰酶消化收取细胞，流式细胞分选双阳性细胞，通过裂解液裂解细胞提取基因组，之后针对sasgRNA位点设计引物进行定向扩增，扩增产物经纯化后进行深度测序，进行数据分析了解sasgRNA位点处的A-G和C-T突变频率。研究中共选取四种不同的sasgRNA脱靶位点(sasgRNA-1～6)用于在DNA水平的脱靶效应分析(非Cas依赖型脱靶效应)。
A-G突变效率结果显示，1249-NLS-TadA-8e-linker(8e)在sasgRNA-1，2,4,6四个位点处的碱基编辑效率均<8％(该编辑效率为脱靶位点所有A-G突变频率的总和)，表明其DNA水平的脱靶效应(非Cas依赖型脱靶效应)很低。而与1249-NLS-TadA-8e-linker(8e)相比，1249-NLS-TadA-8e-linker(V28G)、1249-NLS-TadA-8e-linker(N46L)、1249-NLS-TadA-8e-linker(N46C)、1249-NLS-TadA-8e-linker(N46P)、1249-NLS-TadA-8e-linker(N108Y)、1249-NLS-TadA-8e-linker(V28G-N46L)、1249-NLS-TadA-8e-linker(V28G-N46P)、1249-NLS-TadA-8e-linker(V28G-N46C)、1249-NLS-TadA-8e-linker(N46C-N108Y)、1249-NLS-TadA-8e-linker(N46P-N108S)、1249-NLS-TadA-8e-linker(F84M-N108Y)、1249-NLS-TadA-8e-linker(F84M-N108C)、1249-NLS-Q13XZ4(VN)-linker、1249-NLS-B3PCY2(VN)-linker、1249-NLS-B5ZCW4(VN)-linker、1249-NLS-E8WVH3(VN)-linker、1249-NLS-Q57LE3(VN)-linker、1249-NLS-Q99W51(VN)-linker等点突变能显著降低sasgRNA-1,2，4四个位点的DNA水平脱靶效应(非Cas依赖型脱靶效应)；在sasgRNA-6位点处，以上碱基编辑器的DNA水平脱靶效应(非Cas依赖型脱靶效应)也低于8e，其在1,2,4,6四种脱靶位点的A-G碱基编辑效率均<4％(图39c)。
C-T突变效率分析显示，除V28G-N46C在sasgRNA-6位点处的突变效率接近4％外，其余碱基编辑器在四种位点的C-T突变效率均<2％(图39d)。
DNA脱靶效应分析结果表明，点突变和新的TadA脱氨基酶构建的碱基编辑器在非Cas依赖型脱靶效应上处于较低的水平，安全性有明显提高。
实施例10优选碱基编辑器的活性窗口和碱基偏好性
本实施例中，优选1249-NLS-TadA-8e-linker、1249-NLS-TadA-8e-linker(V28G)、1249-NLS-TadA-8e-linker(N46L)、1249-NLS-TadA-8e-linker(N46C)、1249-NLS-TadA-8e-linker(N46P)、1249-NLS-TadA-8e-linker(N108Y)、1249-NLS-TadA-8e-linker(V28G-N46L)、1249-NLS-TadA-8e-linker(V28G-N46P)、1249-NLS-TadA-8e-linker(V28G-N46C)、1249-NLS-TadA-8e-linker(N46C-N108Y)、1249-NLS-TadA-8e-linker(N46P-N108S)、1249-NLS-TadA-8e-linker(F84M-N108Y)、1249-NLS-TadA-8e-linker(F84M-N108C)、1249-NLS-Q13XZ4(VN)-linker、1249-NLS-B3PCY2(VN)-linker、1249-NLS-B5ZCW4(VN)-linker、1249-NLS-E8WVH3(VN)-linker、1249-NLS-Q57LE3(VN)-linker、1249-NLS-Q99W51(VN)-linker分析其活性窗口和碱基偏好性，另选择1249-NLS-TadA-8e-linker(8e)为对照。
为系统分析碱基编辑器的活性窗口和碱基偏好性，选择12种sgRNA(S18序列：ACACACACACTTAGAATCTG；HEK4序列：GGCACTGCGGCTGGAGGTGG；ZSCAN2序列：GTGCGGCAAGAGCTTCAGCC；AAVS1序列：GACCCTCAGCCGTGCTGCTC；FAN3序列：CGCCGTCTCCAAGGTGAAAG；EMX序列：GAGTCCGAGCAGAAGAAGAA；MSKC序列：CGTCGCCGATCTTCACAGGG；PT14序列：TCCAAACCCATATATACAGC；RNF序列：GTCATCTTAGTCATTACCTG；S1序列：GAACACAAAGCATAGACTGC；sgB序列：AGAGCCCCCCCTCAAAGAGA；UBE3A序列：GTACAGTTAGTACTCAGCAG)开展相关研究。将1249-NLS-TadA-8e-linker突变体(通过2A-EGFP元件同时表达EGFP)与单向导RNA(sgRNA)(sgRNA同时表达尿嘧啶糖基化酶抑制因子UGI和2A-mCherry)共转染至培养的293T细胞。细胞培养72小时后，细胞培养72小时后，通过FACS收集同时表达EGFP和mCherry的双阳性细胞(方案1)。
收集好的细胞抽提基因组，使用可特异性扩增目标sgRNA位点信息的引物进行定向PCR。PCR产物通过扩增子测序，以分析优选碱基编辑器的活性窗口和碱基偏好性(n＝3次生物学重复)。
结果显示，1249-NLS-TadA-8e-linker具有介导C·G--T·A突变(CBE活性)和A·T--G·C突变(ABE活性)的能力，但其ABE活性显著高于其CBE活性。其中1249-NLS-TadA-8e-linker的ABE活性窗口为A2-A14(PAM为21-23)，CBE活性窗口为(C4-C7)(图40a)。1249-NLS-TadA-8e-linker的CBE活性呈现出碱基偏好性，其对不同碱基位点的偏好性为CA≈CT>CG≈CC，AC≈TC>GC≈CC(图40b)。
TadA-8e不同点突变对其碱基编辑活性和碱基偏好性都有影响。单点突变而言，其中V28G点突变后，1249-NLS-TadA-8e-linker(V28G)的CBE活性高于ABE活性，CBE活性窗口为(C4-C7)，ABE活性窗口为A5(图40c)。其对不同碱基位点的偏好性为CA≈CT>CG≈CC，AC≈TC>CC>GC(图40d)。1249-NLS-TadA-8e-linker(N108Y)的CBE活性与ABE活性类似，其中CBE活性窗口为(C4-C7)，ABE活性窗口为(A5-A7)(图40e)。其对不同碱基位点的偏好性为CT≈CA≈CC>CG，TC>AC>CC≈GC(图40f)。1249-NLS-TadA-8e-linker(N46L)几乎无ABE活性，其CBE活性窗口为(C4-C8)(图41a)。其对不同碱基位点的偏好性为CG>CA≈CC≈CT，CC≈TC>GC≈AC(图41b)。、1249-NLS-TadA-8e-linker(N46C)的CBE活性与ABE活性类似，其中CBE活性窗口为(C4-C8)，ABE活性窗口为(A5-A7)(图41c)。其对不同碱基位点的偏好性为CT≈CA≈CC>CG，CC≈TC>GC≈AC(图41d)。1249-NLS-TadA-8e-linker(N46P)仅有CBE活性，活性窗口为(C4-C8)(图41e)。其对不同碱基位点的偏好性为CC≈TC>GC≈AC(图41f)。
双点突变而言，1249-NLS-TadA-8e-linker(V28G-N46L)的ABE活性很低，CBE活性窗口为(C4-C8)(图42a)，其对不同碱基位点的偏好性为CG≈CT≈CA>CC，TC≈CC>GC≈AC(图42b)。1249-NLS-TadA-8e-linker(V28G-N46C)的ABE活性很低，CBE活性窗口为(C4-C9)(图42c)，其对不同碱基位点的偏好性为TC≈CC≈AC>GC(图42d)。1249-NLS-TadA-8e-linker(V28G-N46P)的ABE活性很低，CBE活性窗口为(C4-C8)(图42e)，其对不同碱基位点的偏好性为CT≈CG≈CA>CC，TC≈CC>GC≈AC(图42f)。1249-NLS-TadA-8e-linker(N46C-N108Y)的ABE活性和CBE活性均非常低，ABE活性窗口为A5，CBE活性窗口为C4-C6(图43a)。其对不同碱基位点的偏好性为CA≈CT>CC≈CG，AC≈TC>CC≈GC(图43b)。1249-NLS-TadA-8e-linker(N46P-N108S)的CBE活性显著高于ABE活性，ABE活性窗口为A5，CBE活性窗口为C4-C9(图43c)，其并无明显的碱基偏好性(图43d)。1249-NLS-TadA-8e-linker(F84M-N108Y)的ABE活性和CBE活性相似，ABE的活性窗口为A5-A7，CBE的活性窗口为C4-C7(图44a)，其对不同碱基位点的偏好性为CT≈CA≈CC>CG，TC≈AC>CC≈GC(图44b)。1249-NLS-TadA-8e-linker(F84M-N108C)的ABE活性显著高于CBE，ABE的活性窗口为A5-A7，CBE的活性窗口为C4-C7(图44c)。其对不同碱基位点的偏好性为CA≈CT>CC≈CG，AC≈TC>CC≈GC(图44d)。
总结可知，TadA-8e不同点突变会显著改变1249-NLS-TadA-8e-linker的碱基编辑活性和偏好性，且不同点突变对ABE和CBE活性的影响有明显差别。
TadA同源蛋白来源的碱基编辑器而言，1249-NLS-Q13XZ4(VN)-linker的CBE活性显著高于ABE活性，CBE活性窗口为C4-6，ABE活性窗口为A5，A7(图45a)。其对不同碱基位点的偏好性为CA≈CT>CC≈CG，AC≈TC>CC≈GC(图45b)。1249-NLS-B3PCY2(VN)-linker的ABE活性很低，CBE活性窗口为C4-7(图45c)。其对不同碱基位点的偏好性为，在CT/AC位点具有高编辑活性，在CG/GC位点编辑活性很低(图45d)。1249-NLS-E8WVH3(VN)-linker具有相似的ABE和CBE活性，CBE活性窗口为C4-6，ABE活性窗口为A3-7(图45e)。其对不同碱基位点的偏好性为CA≈CT>CC>CG，AC>TC≈CC≈GC(图45f)。1249-NLS-B5ZCW4(VN)-linker的CBE活性很低，ABE活性窗口为A5-7(图46a)、1249-NLS-Q57LE3(VN)-linker的CBE活性很低，ABE活性窗口为A5,(图46b)，1249-NLS-Q99W51(VN)-linker的CBE和ABE活性均很低，CBE活性窗口为C5，ABE活性窗口为A3，A5(图46c)
上述的对实施例的描述是为便于该技术领域的普通技术人员能理解和使用发明。熟悉本领域技术的人员显然可以容易地对这些实施例做出各种修改，并把在此说明的一般原理应用到其他实施例中而不必经过创造性的劳动。因此，本发明不限于上述实施例，本领域技术人员根据本发明的揭示，不脱离本发明范畴所做出的改进和修改都应该在本发明的保护范围之内。
序列表
<110> 复旦大学
<120> 碱基编辑器及其构建方法与应用
<160> 58
<170> SIPOSequenceListing 1.0
<210> 1
<211> 167
<212> PRT
<213> 大肠杆菌(Escherichia coli)
<400> 1
Met Ser Glu Val Glu Phe Ser His Glu Tyr Trp Met Arg His Ala Leu
1 5 10 15
Thr Leu Ala Lys Arg Ala Arg Asp Glu Arg Glu Val Pro Val Gly Ala
20 25 30
Val Leu Val Leu Asn Asn Arg Val Ile Gly Glu Gly Trp Asn Arg Ala
35 40 45
Ile Gly Leu His Asp Pro Thr Ala His Ala Glu Ile Met Ala Leu Arg
50 55 60
Gln Gly Gly Leu Val Met Gln Asn Tyr Arg Leu Ile Asp Ala Thr Leu
65 70 75 80
Tyr Val Thr Phe Glu Pro Cys Val Met Cys Ala Gly Ala Met Ile His
85 90 95
Ser Arg Ile Gly Arg Val Val Phe Gly Val Arg Asn Ser Lys Arg Gly
100 105 110
Ala Ala Gly Ser Leu Met Asn Val Leu Asn Tyr Pro Gly Met Asn His
115 120 125
Arg Val Glu Ile Thr Glu Gly Ile Leu Ala Asp Glu Cys Ala Ala Leu
130 135 140
Leu Cys Asp Phe Tyr Arg Met Pro Arg Gln Val Phe Asn Ala Gln Lys
145 150 155 160
Lys Ala Gln Ser Ser Ile Asn
165
<210> 2
<211> 1400
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 2
Met Ala Pro Lys Lys Lys Arg Lys Val Gly Ile His Gly Val Pro Ala
1 5 10 15
Ala Asp Lys Lys Tyr Ser Ile Gly Leu Ala Ile Gly Thr Asn Ser Val
20 25 30
Gly Trp Ala Val Ile Thr Asp Glu Tyr Lys Val Pro Ser Lys Lys Phe
35 40 45
Lys Val Leu Gly Asn Thr Asp Arg His Ser Ile Lys Lys Asn Leu Ile
50 55 60
Gly Ala Leu Leu Phe Asp Ser Gly Glu Thr Ala Glu Ala Thr Arg Leu
65 70 75 80
Lys Arg Thr Ala Arg Arg Arg Tyr Thr Arg Arg Lys Asn Arg Ile Cys
85 90 95
Tyr Leu Gln Glu Ile Phe Ser Asn Glu Met Ala Lys Val Asp Asp Ser
100 105 110
Phe Phe His Arg Leu Glu Glu Ser Phe Leu Val Glu Glu Asp Lys Lys
115 120 125
His Glu Arg His Pro Ile Phe Gly Asn Ile Val Asp Glu Val Ala Tyr
130 135 140
His Glu Lys Tyr Pro Thr Ile Tyr His Leu Arg Lys Lys Leu Val Asp
145 150 155 160
Ser Thr Asp Lys Ala Asp Leu Arg Leu Ile Tyr Leu Ala Leu Ala His
165 170 175
Met Ile Lys Phe Arg Gly His Phe Leu Ile Glu Gly Asp Leu Asn Pro
180 185 190
Asp Asn Ser Asp Val Asp Lys Leu Phe Ile Gln Leu Val Gln Thr Tyr
195 200 205
Asn Gln Leu Phe Glu Glu Asn Pro Ile Asn Ala Ser Gly Val Asp Ala
210 215 220
Lys Ala Ile Leu Ser Ala Arg Leu Ser Lys Ser Arg Arg Leu Glu Asn
225 230 235 240
Leu Ile Ala Gln Leu Pro Gly Glu Lys Lys Asn Gly Leu Phe Gly Asn
245 250 255
Leu Ile Ala Leu Ser Leu Gly Leu Thr Pro Asn Phe Lys Ser Asn Phe
260 265 270
Asp Leu Ala Glu Asp Ala Lys Leu Gln Leu Ser Lys Asp Thr Tyr Asp
275 280 285
Asp Asp Leu Asp Asn Leu Leu Ala Gln Ile Gly Asp Gln Tyr Ala Asp
290 295 300
Leu Phe Leu Ala Ala Lys Asn Leu Ser Asp Ala Ile Leu Leu Ser Asp
305 310 315 320
Ile Leu Arg Val Asn Thr Glu Ile Thr Lys Ala Pro Leu Ser Ala Ser
325 330 335
Met Ile Lys Arg Tyr Asp Glu His His Gln Asp Leu Thr Leu Leu Lys
340 345 350
Ala Leu Val Arg Gln Gln Leu Pro Glu Lys Tyr Lys Glu Ile Phe Phe
355 360 365
Asp Gln Ser Lys Asn Gly Tyr Ala Gly Tyr Ile Asp Gly Gly Ala Ser
370 375 380
Gln Glu Glu Phe Tyr Lys Phe Ile Lys Pro Ile Leu Glu Lys Met Asp
385 390 395 400
Gly Thr Glu Glu Leu Leu Val Lys Leu Asn Arg Glu Asp Leu Leu Arg
405 410 415
Lys Gln Arg Thr Phe Asp Asn Gly Ser Ile Pro His Gln Ile His Leu
420 425 430
Gly Glu Leu His Ala Ile Leu Arg Arg Gln Glu Asp Phe Tyr Pro Phe
435 440 445
Leu Lys Asp Asn Arg Glu Lys Ile Glu Lys Ile Leu Thr Phe Arg Ile
450 455 460
Pro Tyr Tyr Val Gly Pro Leu Ala Arg Gly Asn Ser Arg Phe Ala Trp
465 470 475 480
Met Thr Arg Lys Ser Glu Glu Thr Ile Thr Pro Trp Asn Phe Glu Glu
485 490 495
Val Val Asp Lys Gly Ala Ser Ala Gln Ser Phe Ile Glu Arg Met Thr
500 505 510
Asn Phe Asp Lys Asn Leu Pro Asn Glu Lys Val Leu Pro Lys His Ser
515 520 525
Leu Leu Tyr Glu Tyr Phe Thr Val Tyr Asn Glu Leu Thr Lys Val Lys
530 535 540
Tyr Val Thr Glu Gly Met Arg Lys Pro Ala Phe Leu Ser Gly Glu Gln
545 550 555 560
Lys Lys Ala Ile Val Asp Leu Leu Phe Lys Thr Asn Arg Lys Val Thr
565 570 575
Val Lys Gln Leu Lys Glu Asp Tyr Phe Lys Lys Ile Glu Cys Phe Asp
580 585 590
Ser Val Glu Ile Ser Gly Val Glu Asp Arg Phe Asn Ala Ser Leu Gly
595 600 605
Thr Tyr His Asp Leu Leu Lys Ile Ile Lys Asp Lys Asp Phe Leu Asp
610 615 620
Asn Glu Glu Asn Glu Asp Ile Leu Glu Asp Ile Val Leu Thr Leu Thr
625 630 635 640
Leu Phe Glu Asp Arg Glu Met Ile Glu Glu Arg Leu Lys Thr Tyr Ala
645 650 655
His Leu Phe Asp Asp Lys Val Met Lys Gln Leu Lys Arg Arg Arg Tyr
660 665 670
Thr Gly Trp Gly Arg Leu Ser Arg Lys Leu Ile Asn Gly Ile Arg Asp
675 680 685
Lys Gln Ser Gly Lys Thr Ile Leu Asp Phe Leu Lys Ser Asp Gly Phe
690 695 700
Ala Asn Arg Asn Phe Met Gln Leu Ile His Asp Asp Ser Leu Thr Phe
705 710 715 720
Lys Glu Asp Ile Gln Lys Ala Gln Val Ser Gly Gln Gly Asp Ser Leu
725 730 735
His Glu His Ile Ala Asn Leu Ala Gly Ser Pro Ala Ile Lys Lys Gly
740 745 750
Ile Leu Gln Thr Val Lys Val Val Asp Glu Leu Val Lys Val Met Gly
755 760 765
Arg His Lys Pro Glu Asn Ile Val Ile Glu Met Ala Arg Glu Asn Gln
770 775 780
Thr Thr Gln Lys Gly Gln Lys Asn Ser Arg Glu Arg Met Lys Arg Ile
785 790 795 800
Glu Glu Gly Ile Lys Glu Leu Gly Ser Gln Ile Leu Lys Glu His Pro
805 810 815
Val Glu Asn Thr Gln Leu Gln Asn Glu Lys Leu Tyr Leu Tyr Tyr Leu
820 825 830
Gln Asn Gly Arg Asp Met Tyr Val Asp Gln Glu Leu Asp Ile Asn Arg
835 840 845
Leu Ser Asp Tyr Asp Val Asp His Ile Val Pro Gln Ser Phe Leu Lys
850 855 860
Asp Asp Ser Ile Asp Asn Lys Val Leu Thr Arg Ser Asp Lys Asn Arg
865 870 875 880
Gly Lys Ser Asp Asn Val Pro Ser Glu Glu Val Val Lys Lys Met Lys
885 890 895
Asn Tyr Trp Arg Gln Leu Leu Asn Ala Lys Leu Ile Thr Gln Arg Lys
900 905 910
Phe Asp Asn Leu Thr Lys Ala Glu Arg Gly Gly Leu Ser Glu Leu Asp
915 920 925
Lys Ala Gly Phe Ile Lys Arg Gln Leu Val Glu Thr Arg Gln Ile Thr
930 935 940
Lys His Val Ala Gln Ile Leu Asp Ser Arg Met Asn Thr Lys Tyr Asp
945 950 955 960
Glu Asn Asp Lys Leu Ile Arg Glu Val Lys Val Ile Thr Leu Lys Ser
965 970 975
Lys Leu Val Ser Asp Phe Arg Lys Asp Phe Gln Phe Tyr Lys Val Arg
980 985 990
Glu Ile Asn Asn Tyr His His Ala His Asp Ala Tyr Leu Asn Ala Val
995 1000 1005
Val Gly Thr Ala Leu Ile Lys Lys Tyr Pro Lys Leu Glu Ser Glu Phe
1010 1015 1020
Val Tyr Gly Asp Tyr Lys Val Tyr Asp Val Arg Lys Met Ile Ala Lys
1025 1030 1035 1040
Ser Glu Gln Glu Ile Gly Lys Ala Thr Ala Lys Tyr Phe Phe Tyr Ser
1045 1050 1055
Asn Ile Met Asn Phe Phe Lys Thr Glu Ile Thr Leu Ala Asn Gly Glu
1060 1065 1070
Ile Arg Lys Arg Pro Leu Ile Glu Thr Asn Gly Glu Thr Gly Glu Ile
1075 1080 1085
Val Trp Asp Lys Gly Arg Asp Phe Ala Thr Val Arg Lys Val Leu Ser
1090 1095 1100
Met Pro Gln Val Asn Ile Val Lys Lys Thr Glu Val Gln Thr Gly Gly
1105 1110 1115 1120
Phe Ser Lys Glu Ser Ile Leu Pro Lys Arg Asn Ser Asp Lys Leu Ile
1125 1130 1135
Ala Arg Lys Lys Asp Trp Asp Pro Lys Lys Tyr Gly Gly Phe Asp Ser
1140 1145 1150
Pro Thr Val Ala Tyr Ser Val Leu Val Val Ala Lys Val Glu Lys Gly
1155 1160 1165
Lys Ser Lys Lys Leu Lys Ser Val Lys Glu Leu Leu Gly Ile Thr Ile
1170 1175 1180
Met Glu Arg Ser Ser Phe Glu Lys Asn Pro Ile Asp Phe Leu Glu Ala
1185 1190 1195 1200
Lys Gly Tyr Lys Glu Val Lys Lys Asp Leu Ile Ile Lys Leu Pro Lys
1205 1210 1215
Tyr Ser Leu Phe Glu Leu Glu Asn Gly Arg Lys Arg Met Leu Ala Ser
1220 1225 1230
Ala Gly Glu Leu Gln Lys Gly Asn Glu Leu Ala Leu Pro Ser Lys Tyr
1235 1240 1245
Val Asn Phe Leu Tyr Leu Ala Ser His Tyr Glu Lys Leu Lys Gly Ser
1250 1255 1260
Pro Glu Asp Asn Glu Gln Lys Gln Leu Phe Val Glu Gln His Lys His
1265 1270 1275 1280
Tyr Leu Asp Glu Ile Ile Glu Gln Ile Ser Glu Phe Ser Lys Arg Val
1285 1290 1295
Ile Leu Ala Asp Ala Asn Leu Asp Lys Val Leu Ser Ala Tyr Asn Lys
1300 1305 1310
His Arg Asp Lys Pro Ile Arg Glu Gln Ala Glu Asn Ile Ile His Leu
1315 1320 1325
Phe Thr Leu Thr Asn Leu Gly Ala Pro Ala Ala Phe Lys Tyr Phe Asp
1330 1335 1340
Thr Thr Ile Asp Arg Lys Arg Tyr Thr Ser Thr Lys Glu Val Leu Asp
1345 1350 1355 1360
Ala Thr Leu Ile His Gln Ser Ile Thr Gly Leu Tyr Glu Thr Arg Ile
1365 1370 1375
Asp Leu Ser Gln Leu Gly Gly Asp Lys Arg Pro Ala Ala Thr Lys Lys
1380 1385 1390
Ala Gly Gln Ala Lys Lys Lys Lys
1395 1400
<210> 3
<211> 95
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 3
Glu Leu Gly Gly Gly Gly Thr Gly Gly Gly Gly Ser Ala Glu Tyr Val
1 5 10 15
Arg Ala Leu Phe Asp Phe Asn Gly Asn Asp Glu Glu Asp Leu Pro Phe
20 25 30
Lys Lys Gly Asp Ile Leu Arg Ile Arg Asp Lys Pro Glu Glu Gln Trp
35 40 45
Trp Asn Ala Glu Asp Ser Glu Gly Lys Arg Gly Met Ile Leu Val Pro
50 55 60
Tyr Val Glu Lys Tyr Ser Gly Asp Tyr Lys Asp His Asp Gly Asp Tyr
65 70 75 80
Lys Asp His Asp Ile Asp Tyr Lys Asp Asp Asp Asp Lys Glu Phe
85 90 95
<210> 4
<211> 211
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 4
Thr Ser Gly Ser Pro Lys Lys Lys Arg Lys Val Ser Glu Val Glu Phe
1 5 10 15
Ser His Glu Tyr Trp Met Arg His Ala Leu Thr Leu Ala Lys Arg Ala
20 25 30
Arg Asp Glu Arg Glu Val Pro Val Gly Ala Val Leu Val Leu Asn Asn
35 40 45
Arg Val Ile Gly Glu Gly Trp Asn Arg Ala Ile Gly Leu His Asp Pro
50 55 60
Thr Ala His Ala Glu Ile Met Ala Leu Arg Gln Gly Gly Leu Val Met
65 70 75 80
Gln Asn Tyr Arg Leu Ile Asp Ala Thr Leu Tyr Val Thr Phe Glu Pro
85 90 95
Cys Val Met Cys Ala Gly Ala Met Ile His Ser Arg Ile Gly Arg Val
100 105 110
Val Phe Gly Val Arg Asn Ser Lys Arg Gly Ala Ala Gly Ser Leu Met
115 120 125
Asn Val Leu Asn Tyr Pro Gly Met Asn His Arg Val Glu Ile Thr Glu
130 135 140
Gly Ile Leu Ala Asp Glu Cys Ala Ala Leu Leu Cys Asp Phe Tyr Arg
145 150 155 160
Met Pro Arg Gln Val Phe Asn Ala Gln Lys Lys Ala Gln Ser Ser Ile
165 170 175
Asn Ser Gly Gly Ser Ser Gly Gly Ser Ser Gly Ser Glu Thr Pro Gly
180 185 190
Thr Ser Glu Ser Ala Thr Pro Glu Ser Ser Gly Gly Ser Ser Gly Gly
195 200 205
Ser Ser Arg
210
<210> 5
<211> 169
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 5
Glu His Thr Val Lys Ile Thr Gly Glu His Lys Asp Leu Tyr Arg Gly
1 5 10 15
Ser Phe Arg Asp Thr Pro Asp Glu Lys Ile Met Arg Lys Ala Leu Cys
20 25 30
Leu Ala His Ala Ala Gly Glu Lys Gly Glu Ile Pro Val Gly Ala Val
35 40 45
Ile Ala Asp Val Asn Gly Lys Thr Ile Ala Glu Gly Phe Asn Lys Arg
50 55 60
Glu Glu Ser Lys Asp Pro Thr Asp His Ala Glu Ile Ile Ala Ile Arg
65 70 75 80
Lys Ala Ser Gln Met Leu Lys Asp Trp Arg Leu Ser Gly Leu Thr Leu
85 90 95
Phe Val Thr Met Glu Pro Cys Thr Met Cys Ala Gly Ala Ile Val Thr
100 105 110
Ser Arg Ile Ser Arg Val Val Phe Gly Val Phe Asn Asn Lys Thr Gly
115 120 125
Ser Val Gly Ser Arg Ile Asp Ile Leu Arg Asp Glu Ile Leu Asn Thr
130 135 140
Asn Ile Glu Val Val Ser Gly Val Leu Ile Ser Glu Cys Glu Gly Val
145 150 155 160
Leu Ala Lys Phe Phe Ala Asp Leu Arg
165
<210> 6
<211> 243
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 6
Thr Leu Thr Ser Pro Pro Leu Pro Pro Ala Ala Ile Ala Trp Leu Ala
1 5 10 15
Glu Leu Asp Ile His Thr Arg Gln Gln Leu His Ala Arg Gly Val Val
20 25 30
Thr Ala Phe Leu Gln Leu Lys Ala Ala Gly His Thr Ala Thr Arg Arg
35 40 45
Leu Leu Phe Ala Leu Glu Ala Ala Ala Arg Gly Ile His Trp Arg Gln
50 55 60
Leu Glu Glu Glu Asp Lys Ala Leu Leu Leu Arg Arg Leu Ser Asp His
65 70 75 80
Pro Pro Val Ala Leu Pro Pro Asp Ala Asp Asp Arg Leu Arg Tyr Met
85 90 95
Arg Gln Ala Arg Asp Leu Ala Gln Gln Ala Ala Ala Glu Gly Glu Val
100 105 110
Pro Val Gly Ala Leu Val Val Lys Asp Gly Glu Ile Ile Gly Arg Gly
115 120 125
Tyr Asn Gln Pro Ile Gly Arg His Asp Pro Ser Ala His Ala Glu Met
130 135 140
Gln Ala Leu Arg Asp Ala Ala Ala Arg Leu Arg Asn Tyr Arg Leu Asp
145 150 155 160
Gly Cys Asp Leu Tyr Val Thr Leu Glu Pro Cys Pro Met Cys Ser Gly
165 170 175
Ala Ile Leu His Ala Arg Val Ala Arg Val Ile Tyr Gly Val Ala Asn
180 185 190
Ala Lys Thr Gly Ala Ala Gly Ser Thr Val Asp Leu Phe Ala Asp Pro
195 200 205
Arg Leu Asn His His Ala Ala Val Phe Gly Gly Val Glu Ala Glu Ala
210 215 220
Cys Ala Ala Gln Leu Ser Ala Phe Phe Arg Gln Arg Arg Gln Ser Ala
225 230 235 240
Gly Glu Asp
<210> 7
<211> 195
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 7
Val Val Ile Leu Ile Glu Thr Ser His Ser Phe Gln Ala Phe Pro Ala
1 5 10 15
Arg His Pro His Pro Ser Glu Ala Gly Thr Ala Pro Thr Gly Thr Met
20 25 30
Gly Ala Val Pro Ile Pro Thr Ala Gln Tyr Glu Pro Trp Met Arg Arg
35 40 45
Ala Leu Lys Leu Ala Thr Thr Leu Pro Asp Pro Gly Ala Asp Asp Pro
50 55 60
Pro Val Gly Ala Val Ile Tyr Gly Pro Asp Gly Thr Glu Ile Ala Ala
65 70 75 80
Ala His His Asp Arg Glu Arg Thr Ala Asp Pro Thr Ala Tyr Ala Glu
85 90 95
Ile Leu Ala Leu Arg Gln Ala Ala Gln Ala Leu Gly Thr Trp Arg Leu
100 105 110
Thr Asp Cys Thr Leu Val Thr Thr Leu Glu Pro Gly Thr Met Ser Ala
115 120 125
Gly Ala Ile Val Leu Ala Arg Ile Pro Arg Leu Ile Ile Gly Val Trp
130 135 140
Asn Lys Tyr Asn Gly Ala Val Cys Ser Leu Trp Asp Val Val Arg Asp
145 150 155 160
Arg Arg Leu Asn His Phe Val Glu Val Ile Pro Asp Val Leu Lys Asp
165 170 175
Glu Cys Asp Ala Leu Leu Asp Ser Tyr Leu Asp Ser Pro Ser Asn Leu
180 185 190
Ala Arg Gly
195
<210> 8
<211> 186
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 8
Asp Thr Glu Asn Gly Ser Ala Glu Ser Asp Arg Asn Pro Gly Phe Pro
1 5 10 15
Ser Gly Ala Ser Glu Thr Glu Arg Asp His Asp Glu Arg Trp Met Arg
20 25 30
Leu Ala Leu Glu Glu Ala Arg Leu Ala Ala Ser Glu Gly Glu Val Pro
35 40 45
Ile Gly Ala Val Ile Val Arg Glu Asn Glu Val Ile Ala Arg Ser His
50 55 60
Asn Met Pro Val Asp Arg His Asp Pro Thr Ala His Ala Glu Ile Leu
65 70 75 80
Ala Ile Arg Glu Ala Ala Glu Lys Met Lys Asn Tyr Arg Leu Thr Gly
85 90 95
Met Thr Leu Tyr Val Thr Leu Glu Pro Cys Ile Met Cys Ala Gly Ala
100 105 110
Ile Leu Gln Ala Arg Leu Lys Arg Leu Val Tyr Gly Val Gly Asn Pro
115 120 125
Lys Gly Gly Ala Val Asp Ser Leu Tyr Arg Leu Leu Gln Asp Ser Arg
130 135 140
Leu Asn His Phe Val Glu Val Thr Gly Gly Val Leu Gln Ala Ser Cys
145 150 155 160
Ala Glu Ile Leu Ser Gly Phe Phe Arg Glu Lys Arg Val Thr Ser Pro
165 170 175
Leu Leu Lys Ser Glu Gly Gln Val Lys Ser
180 185
<210> 9
<211> 202
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 9
Gly Ser Glu His Ser Pro Leu Trp Trp His Ile Lys Val Val Ala His
1 5 10 15
Gln Thr Met Arg Ser Ala Lys Ser Leu Thr Met Gln Ser Pro Ile Glu
20 25 30
Leu Ser Pro Gln Lys Met Gln Ala Trp Met Gly Ile Leu Ile Glu Arg
35 40 45
Ala Arg Arg Phe Gly Glu Arg Gly Glu Val Pro Val Ser Ala Val Val
50 55 60
Leu Asp His His Gly Arg Cys Ile Gly His Gly Ile Asn Gln Arg Glu
65 70 75 80
Leu His His Asp Pro Leu Gly His Ala Glu Leu Met Ala Val Arg Gln
85 90 95
Ala Cys Arg Leu Arg Gly Asp Trp Arg Leu Asn Asp Cys Thr Leu Leu
100 105 110
Val Thr Leu Glu Pro Cys Pro Met Cys Ala Gly Ala Leu Val Gln Ala
115 120 125
Arg Val Gly Gln Val Ile Phe Ala Val Thr Asn Pro Lys Arg Gly Ala
130 135 140
Met Gly Ser Thr Ile Asn Leu Ala Thr His Ile Ser Ala His His Arg
145 150 155 160
Met Thr Val Ile Gly Gly Val Leu Gly Glu Glu Ala Lys Glu Met Leu
165 170 175
Ser Ser Trp Phe Lys Gln Arg Arg Arg Arg Ser Asp Gly Thr Gly Ala
180 185 190
Ala Pro Phe Gln Thr Gly Ser Thr Thr Cys
195 200
<210> 10
<211> 194
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 10
Glu Ala Ala Ala Gly Ser Val Pro Gly Asp Ser Pro Ala Ser Ser Ala
1 5 10 15
Val Leu Glu Arg Asp Arg Arg Phe Met Ala Leu Ala Gln Ala Ala Ala
20 25 30
Glu Glu Ala Arg Ala Ile Gly Glu Val Pro Val Gly Ala Val Leu Val
35 40 45
Arg Gly Asp Glu Val Ile Ala Lys Gly Phe Asn His Pro Ile Gly Ala
50 55 60
His Asp Pro Ser Ala His Ala Glu Met Val Ala Leu Arg Ala Ala Ala
65 70 75 80
Gln Ala Val Glu Asn Tyr Arg Leu Pro Gly Cys Glu Leu Tyr Val Thr
85 90 95
Leu Glu Pro Cys Leu Met Cys Ala Gly Ala Ile Met His Ala Arg Ile
100 105 110
Ala Arg Val Val Phe Gly Val Arg Asn Pro Lys Thr Gly Ala Cys Gly
115 120 125
Ser Val Val Asp Ala Phe Ala Asn Pro Gln Leu Asn His His Thr Thr
130 135 140
Val Thr Gly Gly Val Leu Glu Ser Glu Cys Gly Ala Ala Leu Lys Ala
145 150 155 160
Phe Phe Ala Glu Arg Arg Arg Ala Ser Arg Glu Ala Arg Val Ser Ala
165 170 175
Arg Ala Glu Ala Gly Ala Asp Ala Gly Ala Asp Ala Gly Pro Ala Lys
180 185 190
Ala Pro
<210> 11
<211> 238
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 11
Leu Thr Thr Pro Pro Leu Ala Pro Lys Thr Val Ala Ala Leu His Arg
1 5 10 15
Leu Gly Ile Arg Thr Leu Glu Glu Leu Arg Gln Thr Gly Ala Val Gln
20 25 30
Thr Phe Leu Leu Leu Lys Ala Ser Gly Leu Thr Leu Thr Lys Ser Thr
35 40 45
Leu Trp Gln Leu Glu Ser Leu Leu Asp Gly Thr Pro Pro Gln Glu Met
50 55 60
Ser Gln Ala His Lys Ala Arg Leu Leu Ala Glu Leu Lys Asn His Pro
65 70 75 80
Pro Val Ala Ala Phe Pro Pro Gln Gly Glu Met Ala His Phe Met Gly
85 90 95
Leu Ala Leu Glu Gln Ala Lys Leu Ser Ala Leu Met Gly Glu Ile Pro
100 105 110
Val Gly Ala Val Ile Val Ser Asp Gly Lys Ile Ile Ala Ser Ala His
115 120 125
Asn Thr Cys Ile Ala Asp Cys Asn Val Ser Arg His Ala Glu Ile Asn
130 135 140
Ala Leu Ala Gln Ala Gly Ser Lys Ile Gln Asn Tyr Arg Leu Asp Gly
145 150 155 160
Cys Asp Ile Tyr Ile Thr Leu Glu Pro Cys Ala Met Cys Ala Ser Ala
165 170 175
Leu Ile Gln Ala Arg Val Lys Arg Val Ile Tyr Gly Val Gly Asn Pro
180 185 190
Lys Thr Gly Ala Ala Gly Ser Ile Val Asn Leu Phe Ala Asp Lys Arg
195 200 205
Leu Asn Thr His Thr Ala Ile Arg Gly Gly Ile Leu Gln Glu Glu Cys
210 215 220
Arg Ala Val Leu Ser Arg Phe Phe Gln Asn Lys Arg Lys Gly
225 230 235
<210> 12
<211> 150
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 12
Leu Glu Glu Ala Ile Leu Arg Thr Ala Leu Ala Ala Ala Tyr Ser Glu
1 5 10 15
Ala Glu Leu Ala Phe Ala Glu Asn Glu Val Pro Val Gly Ala Val Val
20 25 30
Val Trp Asn Asn Glu Ile Ile Ala Arg Asp His Asn Arg Ile Val Gln
35 40 45
Ser Ala Asn Ala Leu Arg His Ala Glu Leu Ser Val Leu Glu Arg Ala
50 55 60
Ala Gly Val Val Lys Asn Glu Arg Leu Ile Asp Cys His Leu Val Thr
65 70 75 80
Thr Leu Glu Pro Cys Met Met Cys Ser Gly Ala Ile Val His Ser Arg
85 90 95
Val Ala Ser Val His Phe Leu Val Glu Asn Glu Arg Ala Pro Gly Leu
100 105 110
Lys Gln Leu Leu Ala Leu Pro Asn Val Asn His Lys Pro Gln Leu Tyr
115 120 125
Arg His Arg Ser Glu Asp Trp Asp Ser Ala Ala Leu Leu Arg Arg Phe
130 135 140
Phe Gln Met Arg Arg Gln
145 150
<210> 13
<211> 143
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 13
Asp Lys Lys Phe Met Glu Leu Ala Met Ser Val Ala Arg Gln Ser Asp
1 5 10 15
Cys Glu Val Pro Val Gly Ala Val Leu Val Arg Gln Gly Ala Val Ile
20 25 30
Ala Ser Ser Gly Asn Arg Val Phe Arg Asp Ser Asp Pro Thr Ala His
35 40 45
Ala Glu Met Phe Val Ile Arg Glu Ala Ile Gln Arg Leu Gly Val Lys
50 55 60
Phe Leu Val Asp Phe Glu Leu Tyr Val Thr Leu Glu Pro Cys Pro Met
65 70 75 80
Cys Met Tyr Ala Met Ser Leu Ser Arg Val Gly Thr Leu Cys Phe Ala
85 90 95
Val Tyr Asn Glu Lys Arg Gly Ala Ile Ser Asn Gly Cys Leu Gly Thr
100 105 110
Gly Thr Tyr Phe His Val Pro Ile Ile Tyr Glu Gly Leu Met Glu Asp
115 120 125
Glu Ala Lys Glu Leu Leu Ser Ser Phe Phe Arg Ile Leu Arg Asn
130 135 140
<210> 14
<211> 222
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 14
Ser Asp Glu Asn Asn Asn Pro Ser Pro Met Pro Ser Glu Leu Asn His
1 5 10 15
Gln Gln Ser Val Arg Arg Asp Arg Ser Gln Ser Gly Ser Asp Val Ala
20 25 30
Asn Ile Lys Ser Leu Ser His Met His Glu Ser Asp Val Ser Asn Val
35 40 45
Ala Val Ser Asp Glu Glu Leu Glu Gln His Ile Arg Trp Met Lys His
50 55 60
Ala Leu Ala Leu Ala Asp Ser Ala Glu Ser Ile Gly Glu Val Pro Val
65 70 75 80
Gly Ala Cys Val Val Leu Asn Gly Glu Leu Ile Gly Glu Gly Tyr Asn
85 90 95
Thr Pro Ile Ser Asp His Asp Pro Ser Ala His Ala Glu Leu Arg Ala
100 105 110
Val Lys Gln Ala Ala Ser Lys Val Gln Asn Tyr Arg Leu Ile Asp Ala
115 120 125
Thr Leu Tyr Val Thr Leu Glu Pro Cys Ser Met Cys Ala Gly Met Leu
130 135 140
Val His Ala Arg Val Lys Arg Val Val Phe Gly Val Lys Asn Ala Lys
145 150 155 160
Thr Gly Ala Ala Gly Ser Val Met Asn Leu Leu Gln His Pro Ala Leu
165 170 175
Asn His Gln Val Asp Ile Val Ser Gly Val Leu Ala Ser Thr Cys Ala
180 185 190
Asn Lys Leu Ser Asp Phe Phe Arg Lys Arg Arg Glu Glu Ile Lys Ala
195 200 205
Ala Lys Lys Ala Lys Lys Arg Leu Glu Gly Lys Met Pro Asp
210 215 220
<210> 15
<211> 200
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 15
Val Pro Leu Ser Asp Arg Gln Asp Asp Ser Tyr Trp Met Arg Arg Ala
1 5 10 15
Ile Ala Leu Ala Ser Gln Gly Glu Ala Leu Gly Glu Val Pro Val Gly
20 25 30
Ala Val Ile Val Arg Glu Gly Ile Ala Ile Gly Glu Gly Phe Asn Gln
35 40 45
Pro Ile Thr Ser Arg Asp Pro Ser Ala His Ala Glu Ile Val Ala Leu
50 55 60
Arg Gln Ala Ala Ala His Leu Gln Asn Tyr Arg Leu Pro Gly Ala Thr
65 70 75 80
Leu Tyr Val Thr Leu Glu Pro Cys Thr Met Cys Val Gly Ala Leu Val
85 90 95
His Ala Arg Ile Thr Arg Leu Val Tyr Gly Val Ala Asn Pro Lys Ala
100 105 110
Gly Ala Val Thr Ser Arg Ala Arg Leu Leu Asp Ala Asp Tyr Val Asn
115 120 125
His Arg Val Ser Tyr Glu Gly Gly Leu Met Ala Ala Glu Cys Gln His
130 135 140
Gln Leu Ser His Phe Phe Gln Gln Arg Arg Leu Gln His Lys Gln Ala
145 150 155 160
Arg Gly Ala Gly Thr Thr Asn Gln Gly Ile Ile Ala Val Glu Gln Ser
165 170 175
Phe Asp Glu Lys Gln Leu Glu Asp Asp Gly Leu Pro Gly Ile Gly Glu
180 185 190
Phe Gly Lys Asn Thr Pro Leu Lys
195 200
<210> 16
<211> 161
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 16
His Asp Ser Asp Lys Tyr Phe Met Lys Cys Ala Ile Phe Leu Ala Lys
1 5 10 15
Ile Ser Glu Met Ile Gly Glu Val Pro Val Gly Ala Val Leu Val Phe
20 25 30
Asn Asn Thr Ile Ile Gly Lys Gly Leu Asn Ser Ser Ile Leu Asn His
35 40 45
Asp Pro Thr Ala His Ala Glu Ile Lys Ala Leu Arg Asn Gly Ala Lys
50 55 60
Phe Leu Lys Asn Tyr Arg Leu Leu His Thr Thr Leu Tyr Val Thr Leu
65 70 75 80
Glu Pro Cys Ile Met Cys Tyr Gly Ala Ile Ile His Ser Arg Ile Ser
85 90 95
Arg Leu Val Phe Gly Val Lys Tyr Lys Asn Leu Gln Lys Tyr Ile Cys
100 105 110
Cys Lys Asn His Phe Phe Ile Asn Lys Asn Phe Arg Lys Ile Ser Ile
115 120 125
Thr Gln Glu Val Leu Glu Ser Glu Cys Ser Asn Leu Leu Ser Ser Phe
130 135 140
Phe Lys Arg Lys Arg Lys Ile Ala Thr Lys Tyr Phe Asn Asn Asn Ile
145 150 155 160
Ile
<210> 17
<211> 197
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 17
Trp Leu Arg Ser Trp Ala Ser Thr Arg Ser Ser Pro Thr Cys Trp Thr
1 5 10 15
Pro Thr Cys Leu Thr Thr Arg Thr Met Thr Thr Glu Pro Gly Ser Pro
20 25 30
Ala Gln Pro Asp Arg Leu Ile Gly Gly Gly Leu Pro Arg Gln Gly Gly
35 40 45
His Val Ala Asp Glu Glu Leu Met Arg Gln Ala Ile Asp Val Ala Cys
50 55 60
Ser Thr Pro Pro Gly Asp Val Pro Val Gly Ala Leu Val Val Asp Ala
65 70 75 80
Asp Gly Thr Val Leu Ala Thr Ala Thr Asn Arg Arg Glu Gln Asp Ala
85 90 95
Asp Pro Leu Ala His Ala Glu Val Leu Ala Leu Arg Gln Ala Val Arg
100 105 110
Ala Leu Gly Asp Ser Trp Arg Leu Glu Arg Cys Thr Leu Val Val Thr
115 120 125
Leu Glu Pro Cys Ala Met Cys Ala Gly Ala Ala Leu Gly Ala Arg Val
130 135 140
Gly Arg Ile Val Phe Gly Val Tyr Asn Pro Arg Thr Gly Ala Cys Gly
145 150 155 160
Ser Val Trp Asp Leu Pro Arg Glu Ala Leu Leu His Lys Ala Glu Val
165 170 175
Arg Gly Gly Val Leu Gln Pro Glu Cys Glu Gln Leu Leu Arg Gly Phe
180 185 190
Phe Glu Lys Leu Arg
195
<210> 18
<211> 199
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 18
Asn Leu Leu Leu Asp Lys Glu Thr Ser Phe Asn Asn Phe Phe Met Gln
1 5 10 15
Gln Ala Leu Lys Gln Ala Arg Ile Ala Phe Asp Lys Asn Glu Val Pro
20 25 30
Val Gly Ala Val Ile Val Asp Arg Leu Asn Gln Lys Ile Ile Ala Ser
35 40 45
Ser Tyr Asn Asn Thr Glu Glu Lys Asn Asn Ala Leu Tyr His Ala Glu
50 55 60
Ile Ile Thr Ile Asn Glu Ala Cys Asn Ile Ile Ser Cys Lys Asn Leu
65 70 75 80
Asn Asp Tyr Asp Ile Tyr Val Thr Leu Glu Pro Cys Ala Met Cys Ala
85 90 95
Ala Ala Ile Ser His Ser Arg Leu Lys Arg Leu Phe Tyr Gly Val Ser
100 105 110
Asn Pro Lys His Gly Ala Val Glu Ser Asn Leu Arg Tyr Phe Asn Ser
115 120 125
Ser Ala Cys Phe His Arg Pro Glu Ile Tyr Ser Gly Ile Leu Ala Glu
130 135 140
Asp Ser Gly Leu Leu Met Lys Glu Phe Phe Lys Lys Ile Arg Thr Thr
145 150 155 160
Ile Pro Asn His Gly Arg Ala Leu Leu Arg Asn Ser Asn Asp Val Ile
165 170 175
Pro Val Lys Val Gly Asp Cys Cys Met Ala Arg Lys Ile Ala Gln Met
180 185 190
Ser Phe Gln Arg Lys Cys Glu
195
<210> 19
<211> 189
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 19
Thr Val Thr Ala Val Asn Leu Leu Thr Pro His Ser Ala Leu Leu Ser
1 5 10 15
Glu Ala Tyr Pro Gly Asp Leu Thr Ala Ala Asn Tyr Trp Leu Leu Ser
20 25 30
Asp Val Gly Phe Met Arg Arg Ala Leu Ala Leu Ala Gln Gln Gly Ala
35 40 45
Ser Gln Glu Glu Val Pro Val Gly Ala Val Leu Val Cys Asp Asn Lys
50 55 60
Ile Ile Gly Glu Gly Phe Asn Gln Pro Ile Thr Thr Ser Asp Pro Thr
65 70 75 80
Ala His Ala Glu Val Val Ala Leu Arg Cys Ala Cys Gln Thr Leu Gln
85 90 95
Asn Tyr Arg Leu Pro Pro Asn Thr Thr Leu Tyr Val Thr Leu Glu Pro
100 105 110
Cys Thr Met Cys Leu Gly Ala Leu Ile His Ala Arg Leu Ala Arg Leu
115 120 125
Val Phe Ala Val Phe Asn Pro Arg Ala Gly Met Val Gly Ser Gln Leu
130 135 140
Asn Leu Thr Glu Met Asp Phe Tyr Asn His Lys Met Gln Ala Tyr Ser
145 150 155 160
Gly Leu Leu Gln Gln Gln Ser Gln Thr Gln Leu Arg Ser Phe Phe Arg
165 170 175
Ala Arg Arg Gly Lys Lys Lys Pro Gln Lys Gln Ser Asp
180 185
<210> 20
<211> 187
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 20
Ile Leu Thr Glu Ala Asp Gly His Cys Arg Asp Leu Ala Trp Met Thr
1 5 10 15
Leu Ala Leu Asp Tyr Ala Ala Arg Gly Ala Val Gln Gly Glu Val Pro
20 25 30
Val Gly Ala Val Leu Leu Gly Ala Asp Gly His Leu Leu Ala Ala Ala
35 40 45
His Asn Thr Pro Val Gln Ser His Asp Pro Ser Ala His Ala Glu Met
50 55 60
Leu Val Leu Arg Gln Ala Ala Leu Val Leu Gln Asn Tyr Arg Leu Thr
65 70 75 80
Gly Ala Thr Leu Tyr Val Thr Leu Glu Pro Cys Val Met Cys Ile Gly
85 90 95
Ala Met Leu His Ala Arg Val Ala Arg Leu Val Tyr Ala Val Ser Asn
100 105 110
Pro Lys Thr Gly Ala Val Glu Ser Leu Tyr His Leu Leu Glu Asp Asp
115 120 125
Arg Phe Asn His Arg Ile Val Thr Glu Gly Gly Val Leu Ala Ala Pro
130 135 140
Ser Ala Thr Leu Leu Arg Asn Phe Phe Arg Ala Arg Arg Arg Gly Lys
145 150 155 160
Gly Ala Gly Glu Ala Asp Val Ala Thr Ala Glu Thr Gly Val Glu Thr
165 170 175
Gly Gln Arg Ser Gly Asp Ala Ala Leu Gln Ser
180 185
<210> 21
<211> 172
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 21
Gly Lys Glu Met Thr Ala Gln Gly Gly Asp Glu Arg Leu Ser Arg Asp
1 5 10 15
Tyr Tyr Phe Met Gly Leu Ala Leu Glu Lys Ala Lys Glu Ala Phe Asp
20 25 30
Asn Gly Glu Val Pro Val Gly Ala Val Ile Val Arg Gly Glu Glu Ile
35 40 45
Leu Gly Phe Gly Arg Asn Arg Val Glu Arg His Arg Asp Val Thr Ala
50 55 60
His Ala Glu Met Glu Ala Ile Arg Gln Ser Gln Gln Arg Val Gly Asp
65 70 75 80
Trp Arg Leu Asp Ser Thr Thr Leu Tyr Val Thr Lys Glu Pro Cys Leu
85 90 95
Met Cys Trp Gly Ala Val Phe Leu Ser Arg Ile Glu Arg Val Val Phe
100 105 110
Gly Val Ser Asn Pro Lys Gln Ala Asp Phe Cys Cys Ile Lys Asp Phe
115 120 125
Phe Thr Ala Arg Lys Lys Pro Glu Ile Leu Pro Gly Val Arg Ser Gln
130 135 140
Glu Ser Leu Glu Leu Met His Gln Phe Phe Phe Ile Leu Arg Asn Lys
145 150 155 160
Lys Arg Glu Leu Pro Arg Cys Asn Ile Gly Ile Ser
165 170
<210> 22
<211> 177
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 22
Thr Asn Gly Ala Ala Arg Gly Gln Ala Gly Asp Gly Pro Met Ala Leu
1 5 10 15
Ala Leu Ala Glu Ala Arg Ala Ala Ala Ala Arg Gly Glu Val Pro Val
20 25 30
Gly Ala Val Leu Leu Gly Pro Asp Gly Thr Ile Leu Ala Arg Ala Gly
35 40 45
Asn Gln Val Glu Ala Gly His Asp Ala Ala Ala His Ala Glu Met Leu
50 55 60
Val Met Arg Ala Ala Ala Arg Ile Leu Gly Ser Pro Arg Leu Thr Gly
65 70 75 80
Cys Thr Leu Val Val Thr Leu Glu Pro Cys Pro Met Cys Ala Ala Ala
85 90 95
Ser Val His Phe Arg Val Gly Arg Ile Val Phe Gly Val Tyr Asn Pro
100 105 110
Lys Gly Gly Gly Val Glu His Gly Pro Arg Leu Leu Ala Arg Pro Asp
115 120 125
Cys Leu His Arg Pro Glu Trp Val Gly Gly Val Arg Glu Arg Glu Ala
130 135 140
Ala Ala Leu Leu Arg Asp Phe Phe Ala Ala Leu Arg Gly Arg Pro Glu
145 150 155 160
Gly Gly Pro Pro Glu Gly Asp Gly Lys Gly Asp Leu Arg Glu Ala Ser
165 170 175
Ser
<210> 23
<211> 173
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 23
Val His Arg Ile Ser Tyr Gln Gln His Cys Glu Trp Met Glu Gln Ala
1 5 10 15
Ile Ala Leu Ala Glu Gln Ala Gly Ala Ala Gly Glu Val Pro Val Gly
20 25 30
Ala Ile Val Val Ser Pro Gln Gly Glu Leu Leu Gly Thr Gly Glu Asn
35 40 45
Arg Arg Glu Arg Asp Gly Asp Pro Thr Ala His Ala Glu Ile Leu Ala
50 55 60
Leu Arg Ala Ala Gly Gln Lys Leu Gln Thr Trp Tyr Leu Lys Asn Cys
65 70 75 80
Thr Leu Tyr Val Asn Leu Glu Pro Cys Pro Met Cys Ala Gly Ala Ile
85 90 95
Leu Gln Ala Arg Ile Gly Leu Leu Val Tyr Gly Val Asp Asn Ala Lys
100 105 110
Thr Gly Ala Ile Arg Thr Val Leu Asn Leu Pro Asp Ser Arg Cys Ser
115 120 125
Phe His Arg Leu Thr Thr Leu Gly Gly Ile Leu Glu Thr Thr Cys Arg
130 135 140
Gln Gln Leu Gln Thr Trp Phe Ala Arg Lys Arg Gly Asp Ser Leu Ser
145 150 155 160
Pro Ala Leu Val Asp Arg Asp Pro Lys Leu Pro Leu Val
165 170
<210> 24
<211> 629
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 24
Asp Phe Gly Asp Phe Asn Val Tyr His Ile Phe Thr Met Gly Phe Ala
1 5 10 15
Gly Ala Glu Glu Tyr Gln Glu Gln Cys Gly Gln Thr Lys His Arg Leu
20 25 30
Trp Lys Ile Glu Asn Leu Leu Glu Tyr Leu Lys Lys Leu Gly Ile Asn
35 40 45
Thr Ile Leu Leu Gly Pro Leu Phe Ser Ser Val Ser His Gly Tyr Asp
50 55 60
Thr Thr Asp Tyr Tyr Thr Val Asp Arg Arg Leu Gly Thr Asn Glu Asp
65 70 75 80
Leu Ala Ala Leu Met Asn Lys Leu His Asp Ser Gly Phe Lys Val Val
85 90 95
Leu Asp Cys Val Phe Asn His Val Gly Arg Asp Phe Phe Ala Phe Gln
100 105 110
Asp Met Lys Gln Asn Arg Glu Asn Ser Arg Tyr Lys Asp Trp Phe Leu
115 120 125
Gly Val Asn Phe Trp Gly Asn Asn Ser Phe Asn Asp Gly Leu Ser Tyr
130 135 140
Glu Asn Trp Ala Gly His Asp Asn Leu Val Lys Leu Asn Leu Tyr Asn
145 150 155 160
Pro Glu Val Lys Asp Tyr Leu Lys Glu Val Met Arg Phe Trp Ile Glu
165 170 175
Thr Phe His Ile Asp Gly Val Arg Met Asp Ala Ala Asn Val Met Asn
180 185 190
Thr Glu Phe Leu Arg Glu Leu Ser Asp Phe Ser Lys Asn Leu Lys Gln
195 200 205
Asp Phe Trp Phe Val Gly Glu Ser Val His Gly Asp Tyr Asn Val Leu
210 215 220
Val Gln Glu Gly His Leu Asp Ser Val Thr Asn Tyr Glu Asp Tyr Lys
225 230 235 240
Gly Leu Tyr Ser Ser Leu Asn Thr Lys Asn Tyr Phe Glu Ile Ser Phe
245 250 255
Ser Leu Asn Arg Leu Phe Gly Glu Tyr Gly Ile Tyr Lys Ser Phe Tyr
260 265 270
Thr Phe Asn Phe Val Asp Asn His Asp Val Asp Arg Val Ala Ser Thr
275 280 285
Leu Thr Arg Glu Ala Trp Leu Tyr Pro Leu Tyr Leu Met Leu Tyr Thr
290 295 300
Met Pro Gly Ile Pro Ser Ile Tyr Tyr Gly Ser Glu Gln Gly Ala Lys
305 310 315 320
Gly Val Lys Gly Asn Gly Thr Asp Ala Pro Leu Arg Pro Asn Tyr Glu
325 330 335
Ser Met His Phe Asp Glu His Cys Asp Leu Tyr Gln Lys Ile Cys Arg
340 345 350
Met Ala Ala Val Arg Arg Ala Ser Arg Ala Ile Arg Tyr Gly Asp Tyr
355 360 365
Lys Glu Leu Phe Val Lys Ser Gln Gln Met Gly Phe Met Arg Arg Phe
370 375 380
Glu Asn Glu Gln Val Tyr Ala Leu Phe Asn Ser Glu Glu Asn Ser Val
385 390 395 400
Arg Val Glu His His Asp Leu Arg Gly Lys Phe Arg Asp Leu Tyr Asn
405 410 415
Asp Glu Ile Val Glu Cys Asp Gly Ser Val Glu Ile Pro Gly Asp Ser
420 425 430
Gly Arg Val Leu Ala Ala Glu Ser Cys Gly Leu Val Val Glu Glu Ile
435 440 445
Thr Glu Gln Pro Ser Ser Ser Gln Ala Glu Glu Gln Ser Ala Gly Glu
450 455 460
Ile Lys Ser Arg Glu Glu Lys Cys Asp Gln Thr Leu Glu Ile Pro Ile
465 470 475 480
Gly Met Thr Met Gln Asp Ile Met Ser Lys Ala Ile Glu Glu Ala Glu
485 490 495
Arg Gly Ile Glu Glu Gly Glu Val Pro Val Gly Ala Val Ile Leu His
500 505 510
Asn Gly Glu Ile Ile Ala Ala Ala His Asn Gln Lys Glu Thr Leu Gln
515 520 525
Asp Pro Thr Ala His Ala Glu Met Leu Val Ile Arg Glu Ala Ser Lys
530 535 540
Lys Leu Gly Arg Trp Arg Leu Asp Asp Cys Glu Leu Tyr Val Thr Ala
545 550 555 560
Glu Pro Cys Pro Met Cys Met Gly Ala Val Ile Gln Ser Arg Ile Arg
565 570 575
Lys Leu Val Tyr Gly Thr Trp Glu Thr Arg Phe Gly Gly Val Glu Thr
580 585 590
Thr Ala Glu Leu Gly Lys His Pro Met Leu Ser Asn Ala Thr Glu Ile
595 600 605
Tyr Ser Gly Ile Cys Glu Glu Lys Cys Gln Glu Leu Leu Glu Arg Phe
610 615 620
Phe Gln Asn Asn Arg
625
<210> 25
<211> 291
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 25
Thr Pro Asp Glu Gln Phe Leu Arg Glu Ala Ile Ala Glu Ala Arg Ala
1 5 10 15
Ala Glu Gln Ala Gly Glu Val Pro Val Gly Ala Val Leu Val Leu Asn
20 25 30
Asn Glu Ile Ile Ala Arg Gly Arg Asn Arg Val Ile Leu Asp Ser Asp
35 40 45
Pro Thr Ala His Ala Glu Ile Val Ala Leu Arg Glu Ala Gly Arg Ile
50 55 60
Leu Gly Asn Tyr Arg Leu Glu Asn Cys Asp Leu Tyr Thr Thr Leu Glu
65 70 75 80
Pro Cys Ala Met Cys Ala Gly Ala Ile Leu His Ala Arg Ile Arg Arg
85 90 95
Leu Ile Tyr Ala Val Ala Asn Pro Lys Ala Gly Ala Cys Gly Ser Ala
100 105 110
Leu Asp Val Met Asn His Pro Arg Leu Asn His Arg Met Glu Val Ala
115 120 125
Val Gly Leu Leu Ala Glu Glu Cys Gly Glu Met Leu Thr Ser Phe Phe
130 135 140
Arg Thr Arg Arg Leu Lys Asn Lys Glu Asn Ala Ala Ser Ala Ile Gly
145 150 155 160
Ile Glu Ala Leu Met Thr Thr Val Lys Lys Ser Ala Ala Pro Lys Lys
165 170 175
Ser Ala Ala Lys Thr Thr Ala Lys Lys Thr Ala Ala Lys Pro Pro His
180 185 190
Lys Trp Ser Ala Lys Val Thr Thr Asp Ser Thr His Pro Asp Glu Gly
195 200 205
Leu Phe Asn Glu Asp Ala Gln Thr Ile Ala Lys Lys Leu Ala Ser Lys
210 215 220
Lys Val Ser Pro Lys Gly Pro Ala Ser Gly Met Gln Met Leu Asn Phe
225 230 235 240
Tyr Ile Asn Arg Ala Gly Lys Asn Leu Pro Lys Ala Arg Gln Ala Glu
245 250 255
Leu Glu Lys Ala Lys Asp Ile Leu Ser Gln Ile Ile Ala Asp Ala Lys
260 265 270
Pro Lys Ala Pro Ala Lys Lys Ala Gly Arg Lys Thr Pro Ala Lys Lys
275 280 285
Thr Ala Asn
290
<210> 26
<211> 180
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 26
Thr Asn Asp Cys Pro Phe Asp Lys Ile Tyr Pro Ser Glu Leu Asn Arg
1 5 10 15
Asn Glu Asp Tyr Phe Met Ala His Ala Phe Asn Glu Ala Ile Glu Ala
20 25 30
Trp Lys Lys Asp Glu Val Pro Ile Gly Ala Val Ile Glu Tyr Glu Gly
35 40 45
Arg Ile Ile Ala Ser Ala His Asn Gln Ser Arg Ser Thr Asn Asp Pro
50 55 60
Thr Ala His Ala Glu Ile Leu Ala Ile Ser Gln Ala Ala Asn Thr Leu
65 70 75 80
Gly Asp Trp Arg Leu Asn Glu Cys Arg Leu Tyr Val Thr Lys Glu Pro
85 90 95
Cys Pro Met Cys Ser Gly Ala Leu Val Ile Ala Arg Ile Gly Lys Val
100 105 110
Tyr Tyr Gly Val Pro Asn Glu Lys Met Gly Cys Val Gly Gly Ala Val
115 120 125
Asp Leu Gly Ala Leu Pro Arg Ser Asn His His Phe Glu Ser Ile Gly
130 135 140
Gly Val Met Glu Asp Ala Asn His Ala Ile Leu Lys Ala Phe Phe Glu
145 150 155 160
Lys Lys Arg Ala Ile Asn Ala Ala Lys Lys Glu Ala Ala Arg Leu Glu
165 170 175
Asp Pro Lys Glu
180
<210> 27
<211> 166
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 27
Ala Val His Glu Pro Gly Ala Gln Asn Pro Pro Cys Ala Gly Arg Arg
1 5 10 15
Asp Leu Arg Met Met Arg Leu Ala Leu Arg Ala Ala Arg Glu Ala Ala
20 25 30
Gly Arg Gly Glu Val Pro Val Gly Ala Val Val Ala Arg Gly Glu Glu
35 40 45
Val Leu Ala Leu Ala Ser Asn Glu Arg Glu Ala Thr Gly Asp Pro Thr
50 55 60
Ala His Ala Glu Leu Leu Ala Ile Arg Arg Ala Ala Ala Ala Leu Gly
65 70 75 80
Gly Trp Arg Leu Thr Gly Cys Thr Leu Tyr Ser Thr Leu Glu Pro Cys
85 90 95
Pro Met Cys Ala Gly Ala Ala Trp Ala Ala Arg Leu Ser Arg Ile Val
100 105 110
Tyr Ala Val Pro Asn Pro Lys Ala Gly Tyr Ala Gly Thr Leu His Asn
115 120 125
Thr Pro Ser Asp Arg Arg Leu Asn His Thr Ala Ser Val Leu Gly Gly
130 135 140
Leu Leu Ala Gln Glu Ser Ala Thr Leu Leu Arg Asp Phe Phe Arg Glu
145 150 155 160
Arg Arg Leu Lys Arg Ala
165
<210> 28
<211> 172
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 28
Asp Ala Ala Lys Val Arg Ser Glu Phe Asp Glu Lys Met Met Arg Tyr
1 5 10 15
Ala Leu Glu Leu Ala Asp Lys Ala Glu Ala Leu Gly Glu Ile Pro Val
20 25 30
Gly Ala Val Leu Val Asp Asp Ala Arg Asn Ile Ile Gly Glu Gly Trp
35 40 45
Asn Leu Ser Ile Val Gln Ser Asp Pro Thr Ala His Ala Glu Ile Ile
50 55 60
Ala Leu Arg Asn Gly Ala Lys Asn Ile Gln Asn Tyr Arg Leu Leu Asn
65 70 75 80
Ser Thr Leu Tyr Val Thr Leu Glu Pro Cys Thr Met Cys Ala Gly Ala
85 90 95
Ile Leu His Ser Arg Ile Lys Arg Leu Val Phe Gly Val Ser Asn Tyr
100 105 110
Lys Thr Gly Ala Ile Gly Ser Arg Phe His Phe Phe Asp Asp Tyr Lys
115 120 125
Met Asn His Thr Leu Glu Val Thr Ser Gly Val Leu Ala Glu Glu Cys
130 135 140
Ser Gln Lys Leu Ser Thr Phe Phe Gln Lys Arg Arg Glu Glu Lys Lys
145 150 155 160
Ile Glu Lys Ala Leu Leu Lys Ser Leu Ser Asp Lys
165 170
<210> 29
<211> 156
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 29
Lys Glu Ile Asn Thr Lys Phe Ile Glu Leu Ala Ile Lys Glu Ala Glu
1 5 10 15
Lys Ala Leu Lys Lys Gly Glu Val Pro Val Gly Ala Val Leu Val Lys
20 25 30
Asp Asp Lys Val Val Ser Lys Gly Tyr Asn Leu Arg Ile Ser Lys Lys
35 40 45
Asn Ala Leu Tyr His Ala Glu Ile Val Ala Ile Glu Arg Ala Cys Lys
50 55 60
Lys Leu Lys Ser Trp Arg Leu Asp Asp Thr Val Leu Tyr Thr Thr Leu
65 70 75 80
Glu Pro Cys Leu Met Cys Ala Gly Ala Ile Met Gln Ala Arg Ile Lys
85 90 95
Lys Val Val Phe Leu Val Lys Asn Glu Lys Gly Gly Ala Val Leu Ser
100 105 110
Asn Tyr Thr Val Phe Asp Asp Lys Lys Leu Pro Phe Lys Val Glu Tyr
115 120 125
Ser Tyr Ile Pro Asp Glu Arg Ala Glu Lys Leu Leu Lys Asp Phe Phe
130 135 140
Lys Ile Leu Arg Glu Ala Lys His His Asp Lys Val
145 150 155
<210> 30
<211> 157
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 30
Glu Asn His Glu Pro Phe Leu Arg Glu Ala Ile Ala Leu Ser Arg Ser
1 5 10 15
Ala Ile Glu Gln Gly Gly Gly Glu Pro Phe Gly Ser Val Leu Val Lys
20 25 30
Asn Gly Glu Val Ile Leu Arg Ala Glu Asn Ser Val Phe Ser Gly His
35 40 45
Asp Met Thr Asn His Ala Glu Met Asn Leu Val Lys Leu Ala Ala Gln
50 55 60
His Tyr Asp Thr Ala Phe Leu Ala Asp Cys Thr Leu Tyr Thr Ser Thr
65 70 75 80
Glu Pro Cys Ala Met Cys Ser Gly Ala Ile Tyr Trp Ser Gly Ile Gly
85 90 95
His Met Val Phe Ala Cys Ser Glu Thr Arg Leu Gly Glu Ile Ala Gly
100 105 110
Ile Gly Leu Asn Val Pro Ser Arg Pro Val Leu Gln Thr Gly Ala Arg
115 120 125
Leu Val Thr Val Val Gly Pro Thr Ser Leu Glu Asp Glu Ala Ala Glu
130 135 140
Val His Gln Glu Phe Trp Pro Lys His Leu Gly Lys Ala
145 150 155
<210> 31
<211> 173
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 31
Pro Asp Ala Pro Arg Asp Leu Gln Tyr Met Glu Arg Ala Leu Gln Leu
1 5 10 15
Ala Ala Tyr Ala Gly Glu Met Gly Glu Val Pro Ile Gly Ala Val Val
20 25 30
Val Ala Gly Gly Lys Ile Leu Gly Glu Gly Cys Asn Leu Arg Glu Met
35 40 45
Trp Asn Asp Pro Thr Ala His Ala Glu Met Val Ala Leu Arg Gln Ala
50 55 60
Ala Gln Ala Val Gly Gly Trp Arg Leu Ser Gly Thr Thr Leu Tyr Val
65 70 75 80
Thr Leu Glu Pro Cys Ala Met Cys Val Gly Ala Ala Val Leu Ala Arg
85 90 95
Val Glu Arg Val Val Phe Gly Val Pro Asn Pro Lys Ala Gly Ala Cys
100 105 110
Gly Ser Val Leu Gln Val Ala Asp Ala Pro Leu Asn His Arg Leu Lys
115 120 125
Val Glu Gly Gly Val Leu Gln Gln Ser Cys Ser His Leu Leu Lys Gln
130 135 140
Phe Phe Arg Glu Leu Arg Met Arg Lys Arg Pro Gln Ala Gly Arg Ser
145 150 155 160
Ser Val Leu Ser Phe Asp Asp Pro Ser Arg Arg Asp Val
165 170
<210> 32
<211> 180
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 32
Ile Thr Ser Val Met Met Glu Met Pro Gly Ser Asp Glu Trp Phe Met
1 5 10 15
Arg Gln Ala Met Lys Glu Ser Arg Lys Ala Leu Val Lys Gly Glu Val
20 25 30
Pro Val Gly Ala Ile Val Val Lys Asp Gly Arg Val Ile Gly Arg Gly
35 40 45
Trp Asn Gln Val Glu Thr Leu Lys Asp Ala Thr Ala His Ala Glu Met
50 55 60
Ile Ala Leu Thr Ala Ala Gln Glu Ala Leu Gly Asp Trp Arg Leu Glu
65 70 75 80
Gly Cys Thr Leu Tyr Val Thr Lys Glu Pro Cys Pro Met Cys Ala Gly
85 90 95
Ala Ile Val His Cys Arg Pro Asp Arg Val Val Phe Gly Val Pro Asn
100 105 110
Ala Lys Thr Gly Ala Ala Gly Gly Trp Ile Asn Leu Leu Asp Ser Asn
115 120 125
Pro Pro Leu Asn His Lys Cys Glu Val Arg Pro Gly Val Leu Gly Asp
130 135 140
Glu Cys Leu Leu His Leu Gln Glu Phe Phe Arg Ala Ala Arg Leu Ala
145 150 155 160
Ala Lys Glu Arg Lys Lys Leu Ser Ala Arg Leu Ser Ser Pro Pro Asp
165 170 175
Glu Asn Gly Glu
180
<210> 33
<211> 160
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 33
Lys Tyr Glu Lys Asp Lys Asn Trp Met Lys Ile Ala Leu Lys Tyr Ala
1 5 10 15
Tyr Tyr Ala Lys Glu Lys Gly Glu Ile Pro Ile Gly Ala Ile Leu Val
20 25 30
Phe Lys Glu Arg Ile Ile Gly Ile Gly Trp Asn Ser Ser Ile Ser Lys
35 40 45
Asn Asp Pro Thr Ala His Ala Glu Ile Ile Ala Leu Arg Gly Ala Gly
50 55 60
Lys Lys Ile Lys Asn Tyr Arg Leu Leu Asn Thr Thr Leu Tyr Val Thr
65 70 75 80
Leu Gln Pro Cys Ile Met Cys Cys Gly Ala Ile Ile Gln Ser Arg Ile
85 90 95
Lys Arg Leu Val Phe Gly Val Asn Cys Asn Ser Ser Asp His Arg Phe
100 105 110
Ser Leu Lys Asn Leu Phe Cys Asp Pro Gln Lys Asp Tyr Lys Leu Asp
115 120 125
Ile Lys Lys Asn Val Met Gln Arg Glu Cys Ser Asp Ile Leu Ile Asn
130 135 140
Phe Phe Gln Lys Lys Arg Lys Asn Lys Ile His Ile Cys Lys Lys Ile
145 150 155 160
<210> 34
<211> 365
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 34
Arg Val Pro Val Pro Gly Thr Arg Ala Val Ala Lys Gln Thr Ser Thr
1 5 10 15
Gly Pro Thr Pro Val Ala Met Glu Asp Asn Gln Ala Ser Ala Gln Leu
20 25 30
Arg Ile Val Pro Ala Thr Ala Ala Asp Ile Asp Ala Leu Ser Ala Leu
35 40 45
Leu Val Glu Thr Val Ala Ser Asn Gly Ser Val Gly Phe Met His Pro
50 55 60
Leu Pro Met Glu Gln Ala Arg Ala Phe Trp Thr Ala Ala Leu Glu Glu
65 70 75 80
Ala Gly Gln Gly Arg Arg Val Val Leu Cys Ala Trp Arg Asp Gly Glu
85 90 95
Leu Leu Ala Thr Gly Ser Leu Ala Leu Val Gln Ala Pro Ser Gln Pro
100 105 110
His Arg Ala Glu Leu Thr Lys Val Met Thr Ala Pro Ala Ala Arg Gly
115 120 125
Ala Gly Val Gly Ala Ile Leu Val Arg Ala Leu Glu Gln Val Ala Arg
130 135 140
Glu Arg Gly Arg Arg Leu Leu Thr Leu Asp Ala Ser Leu Val Glu Gly
145 150 155 160
Ala Thr Ser Phe Tyr Arg Arg Leu Gly Tyr His Thr Thr Gly Asp Ile
165 170 175
Pro Glu Tyr Ala Tyr Lys Pro Leu Gly Gly Leu Gly Ala Thr Arg Ile
180 185 190
Met Trp Lys Leu Leu Asp Ala Arg Glu Pro Met Arg Val Thr Ala Val
195 200 205
Asp Arg Ala Ile Asp Glu Arg His Met Cys His Ala Leu Asp Leu Ala
210 215 220
Glu Arg Ala Glu Lys Glu Phe Asp Glu Ile Pro Val Gly Ala Val Val
225 230 235 240
Val Ser Ala Ala Gly Glu Val Ile Gly Glu Gly Trp Asn Arg Asn Ile
245 250 255
Leu Asp His Asp Pro Ser Ala His Ala Glu Ile Met Ala Met Arg Gln
260 265 270
Ala Gly Gln Ala Leu Ser Asn His Arg Leu Val Gly Ala Thr Leu Tyr
275 280 285
Val Thr Leu Glu Pro Cys Ala Met Cys Ala Met Ala Leu Val His Ala
290 295 300
Arg Ile Ala Arg Val Val Tyr Gly Val Ala Asn Pro Lys Thr Gly Ala
305 310 315 320
Cys Gly Ser Val Phe Asp Val Ile Gly Asp Pro Arg His Asn His Arg
325 330 335
Val Gln Val Gln Gly Gly Val Leu Gly Glu Glu Ala Gly Arg Arg Leu
340 345 350
Thr Asn Tyr Phe Arg Ala Lys Arg Gly Lys Pro Leu Val
355 360 365
<210> 35
<211> 168
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 35
Arg Leu Ser Asp Tyr Asp Tyr Leu Ala Leu Ala Phe Glu Glu Ala Asn
1 5 10 15
Gln Ala Phe Glu Glu Gly Thr Tyr Pro Ile Gly Ala Val Ile Val Asp
20 25 30
Asp Asp Gly Asn Ile Val Ser Lys Gly Arg Asn Arg Val Phe Thr Glu
35 40 45
Ser Asp Cys Thr Ala His Ala Glu Val Asp Ala Ile Arg Lys Ala Gly
50 55 60
His Lys Leu Leu Asp Ile Pro Asn Lys Arg Phe Val Lys Asn Asn Leu
65 70 75 80
Thr Leu Tyr Thr Thr Cys Glu Pro Cys Pro Met Cys Thr Cys Thr Ile
85 90 95
Leu Leu Ser Gly Ile Lys Arg Val Val Trp Ala Val Asp Asn Asp Glu
100 105 110
Tyr Gly Gly Ile Arg Lys Phe Lys Glu Gly Pro His Phe Ile Asn Leu
115 120 125
Phe Asp Thr Ile Thr Tyr Thr Ala Ser Pro Tyr Asp Asp Leu Glu Asn
130 135 140
Lys Gln Arg Asp Leu Leu Ala Arg Trp Asn Ile Gly Arg Gly Ile Leu
145 150 155 160
Asn Thr Glu Trp Glu Lys Pro Lys
165
<210> 36
<211> 239
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 36
Leu Phe Gly Met Ile Cys Cys Ser Trp Ile Leu Ser Ala Val Ile Leu
1 5 10 15
Ser Gln Phe Asp Tyr Phe Asp Ser Ile Val Ser Ile Asn Ile Glu Val
20 25 30
Ser Leu Glu Arg Gly Phe Trp Ile Ser Ala Met Phe Phe Gly Arg Cys
35 40 45
Ala Ser Leu Trp Leu Tyr Leu Ala Ala Phe Leu Phe Gln Leu Pro Leu
50 55 60
Lys Asn Pro Gly Asp Ser Phe Ala Tyr Thr Met Ile Asp Thr Asp Tyr
65 70 75 80
Met Phe Glu Ala Leu Ala Gln Ala Ala Leu Ala Glu Ala Ala Gly Glu
85 90 95
Val Pro Val Gly Ala Val Val Val Lys Asp Gly Gln Ile Ile Gly Arg
100 105 110
Gly Tyr Asn Ala Pro Ile Ser Leu His Asp Pro Ser Ala His Ala Glu
115 120 125
Met Gln Ala Leu Arg Ala Ala Ala Gln Tyr Leu Gly Asn Tyr Arg Leu
130 135 140
Val Gly Cys Glu Leu Phe Val Thr Leu Glu Pro Cys Val Met Cys Ala
145 150 155 160
Gly Ala Ile Met His Ala Arg Ile Arg Arg Leu Val Tyr Gly Val Ser
165 170 175
Asn Phe Lys Thr Gly Val Cys Gly Ser Leu Leu Asp Leu Phe Ala Glu
180 185 190
Gln Arg Leu Asn His His Thr Glu Val Ala Gly Gly Val Leu Ala Glu
195 200 205
Ala Cys Gly Ala Thr Leu Ser Arg Phe Phe Ser Leu Arg Arg Glu Gln
210 215 220
Lys Lys Ile Asn Glu Asn Lys His Ile His Gln Asn Ser Ala Ala
225 230 235
<210> 37
<211> 168
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 37
Thr Asn Ile Ile Thr Arg Gly Thr Pro Pro Ala Asn Pro Pro Gln Gly
1 5 10 15
Gln Thr Trp Arg Ser Met Met Asp Thr Ala Ile Arg Glu Ala Phe Lys
20 25 30
Ala Arg Arg His Glu Glu Val Pro Ile Gly Ala Ala Leu Phe Thr Ala
35 40 45
Glu Gly Glu Leu Leu Ala Thr Gly Asn Asn Thr Pro Leu Thr Gln Asn
50 55 60
Asp Pro Thr Gly His Ala Glu Val Asn Cys Ile Arg Asn Ala Cys Lys
65 70 75 80
Asn Leu Asp Asn Tyr Arg Leu Pro Arg Gly Thr Ile Leu Val Val Thr
85 90 95
Leu Glu Pro Cys Ile Met Cys Leu Gly Ala Ile Ile His Ala Arg Val
100 105 110
Gly Gly Val Val Phe Gly Val Pro Asn Pro Lys Ala Gly Ala Val Val
115 120 125
Ser Asn Leu Glu Gly Thr Asp Leu Ser Phe Ala Asn His Lys Phe Trp
130 135 140
Thr Ile Gly Gly Val Cys Glu Asn Glu Cys Lys Glu Ile Leu Gln Ser
145 150 155 160
Phe Phe Leu His Lys Arg Lys Lys
165
<210> 38
<211> 199
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 38
Thr Glu Thr Pro Pro Leu Pro Leu Arg Ala Leu Arg Ala Leu Pro Glu
1 5 10 15
Asp Pro Val Glu Ala Ala Glu Arg Asp Ala Arg Tyr Met Arg Ala Ala
20 25 30
Leu Glu Glu Ala Arg Leu Ala Glu Ala Ala Gly Glu Val Pro Val Gly
35 40 45
Ala Val Val Val Trp Asn Asp Thr Ile Ile Ala Arg Gly His Asn Leu
50 55 60
Pro Ile Arg Ser Val Asp Pro Ser Ala His Ala Glu Met Gln Ala Leu
65 70 75 80
Arg Ala Ala Ala Gln Val Ile Gly Asn Tyr Arg Met Pro Glu Cys Glu
85 90 95
Leu Tyr Val Thr Leu Glu Pro Cys Val Met Cys Ser Gly Ala Ile Leu
100 105 110
His Ala Arg Leu Arg His Val Val Phe Gly Val Thr Asn Pro Lys Thr
115 120 125
Gly Ala Ala Gly Ser Val Leu Asn Leu Phe Glu Gln Ala Gln Leu Asn
130 135 140
His Gln Thr Thr Ile Ala Gly Gly Val Leu Ala Asp Thr Cys Gly Gln
145 150 155 160
Met Leu Lys Asp Phe Phe Gly Ala Arg Arg Arg Ala Gln Lys Ala Ala
165 170 175
Gln Lys Ala Ala Gln Gln Asp Ala Ala Ala Pro Val Thr Asp Pro Asp
180 185 190
Lys Asn Asn Asp Ile Glu Ala
195
<210> 39
<211> 170
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 39
Ile Ile Gln Lys Thr Val Phe Asp Asp Lys Lys Ile Gln Lys Asn Leu
1 5 10 15
Leu Lys Ile Asp Lys Glu Lys Gly Asp Lys Leu Met Tyr Ile Tyr Asn
20 25 30
Val Met Lys Thr Leu Ile Glu Tyr Ala Ser Arg Ser Asn Asp Ile Pro
35 40 45
Val Ala Ala Ala Val Val Lys Asn Gly Lys Ile Ile Ser Ile Lys Arg
50 55 60
Asn Asp Ser Lys Lys Ala Ile Tyr His Ala Glu Ile Leu Ala Ile Ile
65 70 75 80
Asp Ala Thr Ser Lys Leu Ser Thr Lys Asp Leu Arg Ser Cys Glu Met
85 90 95
Phe Val Thr Lys Glu Pro Cys Pro Met Cys Met Ser Ala Ile Val Leu
100 105 110
Ser Lys Val Lys Arg Leu Tyr Phe Gly Val Arg Asn Phe Lys Met Gly
115 120 125
Ala Ala Glu Ser Cys Phe Asn Leu Ser Gln Asn Pro Phe Leu Asn His
130 135 140
Lys Val Glu Val Ile Gly Gly Ile Cys Glu Asp Glu Cys Arg Leu Leu
145 150 155 160
Leu Lys Arg Phe Phe Glu Glu Lys Arg Arg
165 170
<210> 40
<211> 206
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 40
Leu Gln Pro His Arg Pro Lys Glu Pro Ala Thr Ala Thr Arg Ala Ala
1 5 10 15
Ser Gly Ser Pro Arg Gly Pro Glu Ala Ser Gln Asp Glu Ala Leu Ala
20 25 30
Ala Pro Ala Ala Arg Pro Ala Pro Asp Asp Pro Asp Val Leu Tyr Met
35 40 45
Ala Glu Ala Leu Ala Glu Ala Arg Arg Ala Leu Ala Leu Gly Glu Val
50 55 60
Pro Val Gly Ala Val Ala Val His Asp Gly Arg Ile Ile Ala Arg Gly
65 70 75 80
His Asn Leu Arg Glu Arg Leu Gly Asp Pro Thr Ala His Ala Glu Ile
85 90 95
Leu Val Leu Arg Glu Ala Ala Ala Arg Leu Gly Gly Trp Arg Leu Glu
100 105 110
Gly Val Thr Leu Tyr Val Thr Leu Glu Pro Cys Pro Met Cys Ala Gly
115 120 125
Ala Ile Val Leu Ala Arg Val Pro Arg Leu Val Tyr Gly Val Pro Asn
130 135 140
Pro Lys Ala Gly Ala Ala Gly Ser Leu Met Asn Leu Val Gln His Asp
145 150 155 160
Lys Leu Asn His Arg Val Glu Leu Arg Ala Gly Val Leu Ala Glu Ala
165 170 175
Ser Ala Ala Leu Leu Arg Gly Phe Phe Arg Gln Leu Arg Gly Ala Gly
180 185 190
Asp Pro Ala Pro Ala Pro Gly Pro Ala Glu Thr Gly Asp Pro
195 200 205
<210> 41
<211> 164
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 41
Phe Leu Trp Gln Leu Asp Cys Phe Ala Lys Gly Gly Ser His Ile Lys
1 5 10 15
Lys Asn Leu Gln Glu Met Met Glu Gln Ala Ile Ile Ile Ala Lys Gln
20 25 30
Gly Glu Ala Lys Gly Glu Val Pro Ile Gly Ala Leu Val Val Cys Gln
35 40 45
Gly Glu Val Ile Ala Arg Ala His Asn Gln Lys Glu Thr Leu Ala Asp
50 55 60
Pro Thr Ala His Ala Glu Met Leu Val Ile Arg Glu Ala Ala Lys Lys
65 70 75 80
Leu Gly Arg Trp Arg Leu Asp Asp Cys Gln Leu Phe Val Thr Ala Glu
85 90 95
Pro Cys Val Met Cys Met Gly Ala Ile Ile Gln Ala Arg Ile Pro Phe
100 105 110
Leu Val Tyr Gly Val Thr Asn Lys Lys Phe Gly Gly Val Glu Ser Thr
115 120 125
Ala Tyr Leu Cys Gln His Pro Met Leu Pro Lys Gln Met Glu Ile Tyr
130 135 140
Ala Gly Ile Cys Glu Gln Gln Cys Glu Ala Leu Leu Lys Asn Phe Phe
145 150 155 160
Ser Ser Lys Arg
<210> 42
<211> 162
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 42
Cys Asp His Val Val His Glu Gln Glu Lys Gln Ile Met Arg Gln Ala
1 5 10 15
Ile Glu Phe Ser Arg Glu Lys Met Ile Ala Gly Phe Gly Gly Pro Phe
20 25 30
Gly Ala Val Ile Ser Arg Asn Gly Glu Val Ile Ala Thr Gly Phe Asn
35 40 45
Gln Val Thr Ser Ala Asn Asp Pro Thr Ala His Ala Glu Val Ser Ala
50 55 60
Ile Arg Ala Ala Cys Gln Val Leu Asn Thr Phe Asp Leu Ser Gly Cys
65 70 75 80
Glu Ile Tyr Thr Ser Cys Glu Pro Cys Pro Met Cys Leu Ser Ala Ile
85 90 95
Tyr Trp Ala Arg Leu Asp Arg Ile Tyr Tyr Ala Asn Ser Arg Gln Asp
100 105 110
Ala Ala Asp Ile Gly Phe Asp Asp Ala Phe Leu Tyr Glu Glu Ile Ala
115 120 125
Lys Asp Ile Pro Asp Arg Gln Ile Pro Thr Ile Arg Leu Leu Glu Lys
130 135 140
Glu Ala Ile Leu Pro Phe Asn Glu Trp Arg Ala Lys Pro Asp Lys Ile
145 150 155 160
Ala Tyr
<210> 43
<211> 151
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 43
Glu Leu Thr Val Asn Ser Asp Glu His Phe Met Arg Glu Ala Leu Arg
1 5 10 15
Gln Ala Glu Ile Ala Tyr Glu Glu Gly Glu Ile Pro Val Gly Ala Val
20 25 30
Val Val Cys Gln Lys Lys Ile Ile Ala Lys Ala Tyr Asn Gln Thr Glu
35 40 45
Arg Leu Asn Asp Val Thr Ala His Ala Glu Met Leu Ala Ile Thr Ser
50 55 60
Ala Ala Asn His Leu Gly Gly Lys Tyr Leu Thr Asp Cys Thr Leu Tyr
65 70 75 80
Val Ser Leu Glu Pro Cys Gly Met Cys Ala Gly Ala Leu Asn Trp Ser
85 90 95
Gln Ile Ser Glu Ile Val Tyr Ala Val Ala Asn Glu Lys Arg Gly Phe
100 105 110
Thr Lys Ile Asn Pro Asn Met Ile His Pro Lys Thr Lys Val Lys Ser
115 120 125
Gly Leu Met Ala Ala Glu Ser Lys Lys Leu Ile Asp Asp Phe Phe Ala
130 135 140
Lys Met Arg Asn Lys Lys Asn
145 150
<210> 44
<211> 281
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 44
Pro Arg Pro Ser Tyr Gly Val Ser Gly Thr Thr Gly Ala Glu Gly Ala
1 5 10 15
Ala Thr Arg Ala Ala Gly Ala Asp Arg Gly Gly Asn Asp Phe Leu Thr
20 25 30
Ser Gly Met Val Trp Ala Cys Val Asp Trp Ile Val Tyr Ile Ser Tyr
35 40 45
Asn Ile Ser Trp Ala Gln Gly Cys Gly Leu Gly Glu Phe Val Gly Ser
50 55 60
Arg Leu Gly Ala Asn Ile Ser Asn Cys Thr Gln Lys Arg His Leu Gly
65 70 75 80
Ala Ile Pro Lys Asn Asp Lys Ala Ser Asp Thr Lys Gly Leu Gln Gly
85 90 95
Tyr Ala Asn Tyr Lys Lys Gly Val Phe His Ile Pro Lys Pro Arg Ala
100 105 110
Gly Gly Ser Ser Pro Ser Arg Arg Thr Ile Ile Phe Lys Gly Glu Pro
115 120 125
Leu Leu Asn Gln Ala Gly Tyr Met Arg Glu Ala Leu Ala Glu Ala Glu
130 135 140
Lys Ala Tyr Ala Leu Gly Glu Val Pro Val Gly Ala Val Val Val Leu
145 150 155 160
Asp Asp Lys Ile Ile Gly Arg Gly His Asn Leu Arg Glu Thr Leu Leu
165 170 175
Asp Ser Thr Ala His Ala Glu Ile Ile Ala Leu Arg Gln Ala Ala Arg
180 185 190
His Ile Gly Asp Trp Arg Leu Asn Gly Ser Val Leu Tyr Val Thr Leu
195 200 205
Glu Pro Cys Pro Met Cys Ala Gly Ala Met Ile Gln Phe Arg Val Gln
210 215 220
Thr Leu Val Tyr Gly Val Gln Asn Pro Lys Ala Gly Ala Val Asp Ser
225 230 235 240
Ile Met Asp Val Val Arg Glu Pro Arg Phe Asn His Gln Val Ala Val
245 250 255
Val Ser Gly Val Leu Glu Asp Glu Cys Arg Ala Ile Leu Lys Arg Phe
260 265 270
Phe Lys Glu Leu Arg Gly Asn Lys Lys
275 280
<210> 45
<211> 226
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 45
Asn Val Pro Lys Glu Arg Val Lys Gln Asn Met Cys Met Glu Glu His
1 5 10 15
Lys Thr Gln Glu Glu Pro Val Arg Glu Glu Leu Arg Lys Lys Asp Thr
20 25 30
Gly Glu Glu Glu Ala Lys Lys Glu Thr Arg Glu Glu Lys Lys Glu Glu
35 40 45
Glu Ser Gln Val Pro Ala Phe Ser Arg Lys Asp Ile Lys Tyr Met Lys
50 55 60
Glu Ala Ile Arg Gln Ala Lys Lys Ala Lys Asp Leu Gly Glu Val Pro
65 70 75 80
Ile Gly Cys Val Ile Val Tyr Glu Asp Arg Ile Ile Gly Arg Gly Tyr
85 90 95
Asn Arg Arg Asn Thr Asp Arg Ser Thr Leu Ala His Ala Glu Leu Ala
100 105 110
Ala Ile Arg Lys Ala Cys Lys Lys Met Gly Asp Trp Arg Leu Glu Gly
115 120 125
Cys Thr Met Tyr Val Thr Leu Glu Pro Cys Gln Met Cys Ser Gly Ala
130 135 140
Leu Val Gln Ser Arg Ile Ser Lys Val Val Ile Gly Val Met Asn Pro
145 150 155 160
Lys Ala Gly Cys Ala Gly Ser Val Leu Asn Leu Leu Gln Met Lys Ala
165 170 175
Phe Asn His Gln Val Asp Val Glu Tyr Gly Leu Cys Lys Glu Met Cys
180 185 190
Ala Glu Ile Leu Arg Asp Phe Phe Arg Asp Leu Arg Arg Lys Asn Lys
195 200 205
Glu Ile Arg Leu Gln Glu Arg Lys Gln Lys Glu Ile Gln Ser Gln Glu
210 215 220
Leu Gly
225
<210> 46
<211> 171
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 46
Asp Arg Pro Asp Leu Ser Ala Ser Asp Ala Asp Leu Glu Arg Trp Met
1 5 10 15
Asp His Ala Leu Gly Leu Ala Ser Met Ala Ala Arg Asn Gly Glu Val
20 25 30
Pro Val Gly Ala Val Leu Leu Ser Ala Thr Gly Gln Leu Ile Ala Glu
35 40 45
Ala Val Asn Thr Pro Ile Ala Gln Cys Asp Pro Thr Ala His Ala Glu
50 55 60
Leu Ala Val Leu Arg Lys Gly Ala Leu Ala Thr Gly Asn Tyr Arg Leu
65 70 75 80
Thr Gly Thr Thr Leu Leu Val Thr Leu Glu Pro Cys Ala Met Cys Ala
85 90 95
Gly Ala Ile Cys His Ala Arg Ile Gly His Leu Val Tyr Gly Val Asp
100 105 110
Asn Pro Lys Gly Gly Ala Val Arg His Gly Ala Thr Leu Phe Asp Gln
115 120 125
Val Thr Thr His His Arg Pro Arg Val Thr Ala Gly Ile Arg Ala Asp
130 135 140
Glu Ser Ala Ala Leu Leu Arg Ser Phe Phe Ala Glu Arg Arg Ala Met
145 150 155 160
Thr Arg Pro Arg Arg His Gly Glu Ser Gly Pro
165 170
<210> 47
<211> 170
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 47
Ile Ile Glu Gln Lys Pro Ser Lys Asn Gln Leu Lys Lys Asp Phe Phe
1 5 10 15
Trp Met Asn Lys Ser Leu Asp Leu Ala Lys Lys Ser Glu Ile Leu Gly
20 25 30
Glu Ile Pro Ile Gly Ala Val Leu Ile Tyr Gln Asp Asn Ile Val Ala
35 40 45
Ser Ser Gly Asn Glu Val Ile Leu Arg Asn Asp Pro Ser Ala His Ala
50 55 60
Glu Ile Ile Val Ile Arg Glu Gly Ala Lys His Phe Lys Asn Tyr Arg
65 70 75 80
Leu Lys Glu Asn Leu Thr Ile Tyr Val Thr Ile Phe Pro Cys Ile Met
85 90 95
Cys Met Gly Ala Ile Phe Gln Ser Arg Ile Ser Arg Val Val Tyr Gly
100 105 110
Ser Glu Asn Tyr Lys Asn Asn Leu Ser Phe Phe Leu Phe Glu Lys Asn
115 120 125
Asn Thr Asn Ile Ser Met Asn Tyr Gly His Ile Thr Val Tyr Gly Gly
130 135 140
Ile Leu Ser Arg Ser Cys Ser Ser Ile Leu Lys Asn Phe Phe Lys Lys
145 150 155 160
Arg Arg Asn Leu Ile Gln Pro Lys Lys Phe
165 170
<210> 48
<211> 360
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 48
Ser Glu Ser Asp Asp Ala His Trp Met Arg Glu Ala Ile Ala Gln Ala
1 5 10 15
Arg Ala Ala Gln Gln Ala Gly Glu Val Pro Val Gly Ala Val Leu Val
20 25 30
Arg Asp Gly Gln Val Ile Ala Thr Gly Arg Asn Ala Pro Ile Ala Gly
35 40 45
His Asp Pro Thr Ala His Ala Glu Met Ala Ala Leu Arg Ala Gly Ala
50 55 60
Ala Gln Leu Ser Asn Tyr Arg Leu Asp Gly Cys Thr Leu Tyr Val Thr
65 70 75 80
Leu Glu Pro Cys Ala Met Cys Ser Gly Ala Met Leu His Ala Arg Leu
85 90 95
Pro Arg Val Val Tyr Gly Val Ala Asn Ala Lys Thr Gly Ala Ala Gly
100 105 110
Ser Val Val Asn Leu Phe Ala Glu Pro Arg Leu Asn His Gln Thr Ala
115 120 125
Val Gln Gly Gly Val Leu Ala Asp Glu Cys Gly Ala Leu Leu Ser Asp
130 135 140
Phe Phe Arg Asp Arg Arg Arg Ala Gln Arg Ala Gln Gln Leu Ala Ala
145 150 155 160
His Pro Leu Arg Gln Asp Ala Leu Arg Thr Pro Asp Ala Ala Phe Ala
165 170 175
His Leu Asp Gly Pro Ala Pro His Tyr Val Ser Asp Leu Pro Ala Leu
180 185 190
Ala Gly Leu Arg Leu His Tyr Leu Asp Glu Gly Pro Arg Asp Ala Ala
195 200 205
Arg Thr Trp Leu Leu Leu His Ala Ser Pro Gly Trp Ser His Gly Trp
210 215 220
Gln Pro Met Leu Ala Ala Phe Ala Gln Ala Gly His Arg Val Val Ala
225 230 235 240
Pro Asp Leu Ile Gly Phe Gly Arg Ser Asp Lys Pro Lys Lys Glu Ala
245 250 255
Ala His Arg Leu Asp Trp His Arg Gln Val Leu Ala Glu Leu Val Gln
260 265 270
Arg Leu Ala Leu Arg Gly Val Val Gln Val Ala Gln Pro Gly Ala Pro
275 280 285
Leu Leu Pro Asp Ala Val Leu Ala Pro Leu Val Arg Val Leu His Val
290 295 300
Arg Gly Gly Ala Pro Cys Asp Ala Ala Pro Phe Pro Asp Ala Gly His
305 310 315 320
Arg Ala Gly Pro Arg Ala Phe Ala Arg Phe Ala Asp Ala Asp Ala Ala
325 330 335
Pro Leu Leu Thr Val Pro Asp Ala Ser Ala Ala Ser Ala Leu Arg Ala
340 345 350
Met Glu Tyr Phe Thr Arg Tyr Ala
355 360
<210> 49
<211> 248
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 49
Asn Glu Pro Leu Ala Cys Thr Ala Ser Ile Gly Ser Ser Ala Gln Thr
1 5 10 15
Glu Asp Ser Glu Lys Ser Val Asp Ala Ser Ile Gly Ala Val Ser Ala
20 25 30
Pro Thr Ala Asp His Pro Val Ala Gly Pro Gly Val Pro Pro Gly Asn
35 40 45
Ser Glu Ala Pro Ile Ile Ser Glu Arg Asp Leu Arg Phe Met Ala Leu
50 55 60
Ala Gln Ala Ala Ala Glu Glu Ala Arg Ala Ala Gly Glu Val Pro Val
65 70 75 80
Gly Ala Val Leu Val Arg Gly Asp Glu Val Ile Ala Thr Gly Phe Asn
85 90 95
His Pro Ile Gly Ala His Asp Pro Ser Ala His Ala Glu Met Val Ala
100 105 110
Leu Arg Ala Ala Ala Gln Ser Leu Glu Asn Tyr Arg Leu Pro Gly Cys
115 120 125
Glu Leu Tyr Val Thr Leu Glu Pro Cys Leu Met Cys Ala Gly Ala Ile
130 135 140
Met His Ala Arg Ile Ala Arg Val Val Phe Gly Val Arg Asn Pro Lys
145 150 155 160
Thr Gly Ala Cys Gly Ser Val Val Asp Ala Phe Ala Asn Pro Arg Leu
165 170 175
Asn His His Thr Thr Val Ser Gly Gly Val Leu Glu Ala Glu Cys Gly
180 185 190
Ala Ala Leu Arg Ser Phe Phe Ala Glu Arg Arg Arg Ala Ser Arg Ala
195 200 205
Ala Arg Ala Ala Ala Arg Gly Glu Ala Thr Lys Ala Val Glu Val Ala
210 215 220
Glu Gly Ser Val Gly Thr Ala Thr Asp Ser Ala Thr Gln Pro Asn Ser
225 230 235 240
Glu Pro Pro Pro Thr Thr Pro Leu
245
<210> 50
<211> 186
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 50
Thr Pro Thr Phe Ala Pro Asp Leu Glu Ala Asp Arg Ala Phe Met Arg
1 5 10 15
Leu Ala Leu Asp Gln Ala Gln Asn Ala Trp Leu Leu Gly Glu Val Pro
20 25 30
Val Gly Ala Val Ile Val Lys Asp Gly Lys Val Ile Ala Thr Gly Tyr
35 40 45
Asn Arg Pro Ile Gly Asp His Asp Pro Thr Ala His Ala Glu Ile Val
50 55 60
Ala Leu Arg Gln Ala Ala His Leu Leu Gly Asn Tyr Arg Leu Pro Glu
65 70 75 80
Cys Thr Leu Tyr Val Thr Leu Glu Pro Cys Ala Met Cys Ala Met Ala
85 90 95
Leu Leu His Ala Arg Phe Ala Arg Val Val Phe Gly Val Arg Asn Pro
100 105 110
Lys Thr Gly Ala Ala Gly Ser Val Val Asp Leu Phe Ala Glu Pro Arg
115 120 125
Leu Asn His His Cys Asp Ile Thr Pro Glu Ile Glu Gln Glu Ala Cys
130 135 140
Ser Ala Leu Leu Gln Asp Phe Phe Arg Ala Arg Arg Lys Glu Gln Lys
145 150 155 160
Arg Pro Glu Glu Ser Val His Asp Ala Cys Leu Ser Gly Ile Glu Val
165 170 175
Thr Glu Leu Asp Ile Glu Pro Pro Leu Ser
180 185
<210> 51
<211> 233
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 51
Ala Lys Asp Phe Ser Ile Phe Asp Gln Lys Met Met Gln Leu Ala Leu
1 5 10 15
Glu Glu Ala Lys Leu Ala Phe Asn Lys Glu Glu Val Pro Val Gly Ala
20 25 30
Val Leu Thr Leu Gly Asp Gln Val Ile Ala Arg Ala His Asn Thr Met
35 40 45
Glu Thr Lys Gly Asp Pro Thr Ala His Ala Glu Ile Leu Cys Ile Arg
50 55 60
Lys Gly Ala Glu Ala Ile Gly Asp Trp Arg Leu Leu Gly Ala Thr Leu
65 70 75 80
Tyr Thr Thr Leu Glu Pro Cys Val Met Cys Gly Gly Ala Met Leu Leu
85 90 95
Ala Arg Ile Thr Arg Val Val Trp Gly Val Pro Asn Ile Arg His Gly
100 105 110
Ser Asn Gly Ser Trp Ile Asp Leu Phe Glu Lys Lys His Pro Thr His
115 120 125
Gln Leu Glu Ile Ala Gly Gly Leu Tyr Glu Glu Glu Ala Ala Asn Leu
130 135 140
Met Arg Leu Phe Phe Lys Asp Lys Gly Gln Asn Met Lys Lys Thr Asn
145 150 155 160
Phe Leu Leu Val Glu Leu Phe Asp Glu Leu Ile Ala Phe Gln Glu Thr
165 170 175
Lys Leu Leu Asn Tyr Ala Ala Glu Ile Ile Pro Asn Val Thr Ser Asp
180 185 190
Asp Ile Leu Gln Pro Asn Asp Tyr Asp Glu Leu Glu Asn His Pro Phe
195 200 205
Phe Arg Tyr Glu Glu Gly Val Leu Lys Gly Ile Gln Thr Ala Lys Met
210 215 220
Ala Val Leu Ala Lys Leu Arg Glu Asp
225 230
<210> 52
<211> 517
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 52
Ala Val Gly Val Ser Gly Gly Ala Asp Ser Val Ala Leu Leu Arg Ala
1 5 10 15
Leu Val Leu Val Gly Ala Arg Pro Val Ala Ala His Leu Asp His Ala
20 25 30
Leu Arg Pro Glu Ser Ala Gln Asp Ala Ala Trp Val Gly Ala Leu Ala
35 40 45
Ala Asp Leu Gly Val Pro Phe Gly Thr Ala Arg Val Asp Val Gly Ala
50 55 60
Val Ala Gly Arg Arg Gly Trp Asn Val Glu Glu Ala Ala Arg Arg Val
65 70 75 80
Arg Tyr Asp Phe Leu Ala Arg Thr Ala Arg Ala His Gly Ala Gly Leu
85 90 95
Val Leu Thr Ala His Thr Arg Arg Asp Gln Ala Glu Thr Val Leu Trp
100 105 110
Gly Leu Leu Arg Gly Glu Ala Thr Leu Arg Gly Ile Ala Pro Val Trp
115 120 125
Gly Arg Val Arg Arg Pro Trp Leu Asp Ala Pro Arg Ala Asp Leu Glu
130 135 140
Thr Phe Leu Arg Ala Leu Gly Gln Asp Trp Arg Glu Asp Pro Thr Asn
145 150 155 160
Ala Asp Pro Ala Tyr Thr Arg Ala Trp Leu Arg Cys Glu Val Met Pro
165 170 175
Val Leu Arg Ala Arg Phe Pro Ala Val Glu Glu Ala Leu Ala His Phe
180 185 190
Gly Arg Leu Asn Ala Glu Asp Asp Asp Ala Leu Ser Ala Gln Ala Ala
195 200 205
Ala Leu Ser Ala His Ala Pro Leu Glu Arg Gln Ser Pro Ala Val Leu
210 215 220
Arg Arg His Val Ala Ala Glu Leu Arg Ala Ala Gly Leu Glu Tyr His
225 230 235 240
Gly Leu His Val Gly Ala Leu Ala Glu Gly Leu Arg Ala Gly Thr Thr
245 250 255
Arg His Leu Thr Leu Pro Gly Pro Arg Glu Val Thr Val Thr Gly Gly
260 265 270
Arg Leu His Leu Gly Pro Leu Ala Trp Pro Gly Pro Asp Phe Ala Leu
275 280 285
Pro Ser Gly Trp Thr Arg Arg Thr Arg Gln Pro Gly Asp Arg Leu Arg
290 295 300
Leu Pro Gly Gly Thr Arg Lys Leu Ser Asp Val Leu Thr Asp Ala Lys
305 310 315 320
Val Pro Arg Ala Glu Arg Asp Arg Val Pro Leu Ala Val Asp Ala Gly
325 330 335
Gly Ala Val Gln Trp Val Gly Leu Thr Ser Pro Leu Trp Ala Ala Gly
340 345 350
Ala Arg Glu Ala Ala Gly Ala Val Pro Asp Pro Leu His Ala Ala Met
355 360 365
Gly Glu Ala Leu Ala Leu Ala His Glu Ala Ala Gly Ala Gly Glu Val
370 375 380
Pro Val Gly Ala Val Val Leu Asp Ala Ala Gly Gly Val Val Gly Arg
385 390 395 400
Gly Arg Asn Thr Ser Arg Glu His Gly Asp Met Thr Arg His Ala Glu
405 410 415
Leu Ala Ala Leu Arg Glu Ala Ser Glu Ala Val Gly Ala Tyr Leu Gly
420 425 430
Asp Cys Thr Leu Val Val Thr Leu Glu Pro Cys Pro Met Cys Leu Gly
435 440 445
Ala Ala Ile Glu Ala Arg Val Gly His Val Val Tyr Gly Val Ala Asn
450 455 460
Pro Lys Ala Gly Ala Leu Gly Gly Val Ala Asp Val Leu Ala Ala His
465 470 475 480
Trp Gly His Ala Pro Arg Val Thr Pro Gly Val Arg Ala Ala Glu Ala
485 490 495
Gly Ala Leu Leu Arg Arg Ser Phe Gln Ala Ile Arg Gly Gln Gly Arg
500 505 510
Gly Glu Asp Gly Ala
515
<210> 53
<211> 290
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 53
Asp Val Val Asp Ser Arg Ser Asp Gln Asp Trp Met Arg Leu Ala Leu
1 5 10 15
Thr Leu Ala Gln Arg Ala Ala Ala Ala Gly Glu Val Pro Val Gly Ala
20 25 30
Leu Leu Val Arg Asp Gly Val Ile Ile Gly Glu Gly Trp Asn Arg Pro
35 40 45
Ile Lys Thr His Asp Ala Ser Ala His Ala Glu Ile Gln Ala Leu Arg
50 55 60
Asp Ala Gly Gln Arg Val Gly Asn Tyr Arg Leu Pro Gly Thr Thr Leu
65 70 75 80
Tyr Val Thr Leu Glu Pro Cys Val Met Cys Ala Gly Ala Ile Val His
85 90 95
Ala Arg Val Gly Gln Val Ile Tyr Gly Val Pro Asn Pro Lys Ala Gly
100 105 110
Ala Cys Gly Ser Val Phe Asp Leu Leu Pro Ser Asp Ala Arg Phe Asn
115 120 125
His Arg Thr Asp Cys Lys Gly Gly Val Leu Thr Asp Asp Cys Ala Glu
130 135 140
Thr Leu Arg Ala Phe Phe Arg Ala Arg Arg Ser Thr Ala Phe Lys Ala
145 150 155 160
Ala Thr Ser Arg Pro Glu Thr Ser Ile Met Ser Ile Asp Asn Val Tyr
165 170 175
Ser Asp Asp Ala Ile Arg Asn Arg Leu Ala Thr Glu Leu Pro Arg Trp
180 185 190
Ser Leu Asp Ala Gly Cys Leu Gln Arg Thr Tyr Arg Thr Ser Gly Trp
195 200 205
Lys Gly Thr Leu Met Val Ile Asn Thr Val Gly His Leu Ala Glu Ala
210 215 220
Ala Trp His His Pro Asp Leu Thr Ala Ser Tyr Asp Arg Val Thr Val
225 230 235 240
Ser Leu Thr Thr His Ser Ala Lys Gly Val Thr Asp Lys Asp Phe Glu
245 250 255
Leu Ala Asn Arg Ile Glu Glu Thr Val Met Trp Arg Pro Ala Ser Gly
260 265 270
Ser Ala Leu Glu Gly Thr Pro Asp Asp Val Arg Tyr Ala Tyr Ile Gln
275 280 285
Tyr Asp
290
<210> 54
<211> 182
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 54
Pro Pro Ala Phe Ile Thr Gly Val Thr Ser Leu Ser Asp Val Glu Leu
1 5 10 15
Asp His Glu Tyr Trp Met Arg His Ala Leu Thr Leu Ala Lys Arg Ala
20 25 30
Trp Asp Glu Arg Glu Val Pro Val Gly Ala Val Leu Val His Asn His
35 40 45
Arg Val Ile Gly Glu Gly Trp Asn Arg Pro Ile Gly Arg His Asp Pro
50 55 60
Thr Ala His Ala Glu Ile Met Ala Leu Arg Gln Gly Gly Leu Val Leu
65 70 75 80
Gln Asn Tyr Arg Leu Leu Asp Thr Thr Leu Tyr Val Thr Phe Glu Pro
85 90 95
Cys Val Met Cys Ala Gly Ala Met Val His Ser Arg Ile Gly Arg Val
100 105 110
Val Phe Gly Val Arg Asn Ala Lys Thr Gly Ala Ala Gly Ser Leu Ile
115 120 125
Asp Val Leu His His Pro Gly Met Asn His Arg Val Glu Ile Ile Glu
130 135 140
Gly Val Leu Arg Asp Glu Cys Ala Thr Leu Leu Ser Asp Phe Phe Arg
145 150 155 160
Met Arg Arg Gln Glu Ile Lys Ala Leu Lys Lys Ala Asp Arg Ala Glu
165 170 175
Gly Ala Gly Pro Ala Val
180
<210> 55
<211> 155
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 55
Thr Asn Asp Ile Tyr Phe Met Thr Leu Ala Ile Glu Glu Ala Lys Lys
1 5 10 15
Ala Ala Gln Leu Gly Glu Val Pro Ile Gly Ala Ile Ile Thr Lys Asp
20 25 30
Asp Glu Val Ile Ala Arg Ala His Asn Leu Arg Glu Thr Leu Gln Gln
35 40 45
Pro Thr Ala His Ala Glu His Ile Ala Ile Glu Arg Ala Ala Lys Val
50 55 60
Leu Gly Ser Trp Arg Leu Glu Gly Cys Thr Leu Tyr Val Thr Leu Glu
65 70 75 80
Pro Cys Val Met Cys Ala Gly Thr Ile Val Met Ser Arg Ile Pro Arg
85 90 95
Val Val Tyr Gly Val Asp Asn Pro Lys Gly Gly Cys Ser Gly Ser Leu
100 105 110
Met Asn Leu Leu Gln Gln Ser Asn Phe Asn His Arg Ala Ile Val Asp
115 120 125
Lys Gly Val Leu Lys Glu Ala Cys Ser Thr Leu Leu Thr Thr Phe Phe
130 135 140
Lys Asn Leu Arg Ala Asn Lys Lys Ser Thr Asn
145 150 155
<210> 56
<211> 178
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 56
Glu Leu Gly Glu Ser Phe Leu Met Pro Tyr Ser Leu Glu Glu Gln Thr
1 5 10 15
Tyr Phe Met Gln Glu Ala Leu Lys Glu Ser Glu Lys Ser Leu Gln Lys
20 25 30
Ala Glu Ile Pro Ile Gly Cys Val Ile Val Lys Asp Gly Glu Ile Ile
35 40 45
Gly Arg Gly His Asn Ala Arg Glu Glu Ser Asn Gln Ala Ile Met His
50 55 60
Ala Glu Met Met Ala Ile Asn Glu Ala Asn Ala His Glu Gly Asn Trp
65 70 75 80
Arg Leu Leu Asp Thr Thr Leu Phe Val Thr Ile Glu Pro Cys Val Met
85 90 95
Cys Ser Gly Ala Ile Gly Leu Ala Arg Ile Pro His Val Ile Tyr Gly
100 105 110
Val Ser Asn Gln Lys Phe Gly Gly Val Asp Ser Leu Tyr Gln Ile Leu
115 120 125
Thr Asp Glu Arg Leu Asn His Arg Val Gln Val Glu Arg Gly Leu Leu
130 135 140
Ala Ala Asp Cys Ala Asn Ile Met Gln Thr Phe Phe Arg Gln Gly Arg
145 150 155 160
Glu Arg Lys Lys Ile Ala Lys His Leu Ile Lys Glu Gln Ser Asp Pro
165 170 175
Phe Asp
<210> 57
<211> 349
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 57
Ser Val Pro Ala Asn Pro Cys Gly Ala Pro Leu Ala Asp Ala Asp Arg
1 5 10 15
Leu Glu Ala Gly Lys Ala Ser Ala Ala Pro Leu Glu Gly Ala Ser Gly
20 25 30
Val Asp Leu Ile Gly Gly Asp Ala Ala Cys Gly Thr Ala Gly Ala Gly
35 40 45
Phe Ala Ala Thr Thr Gln Ala Ala Arg Thr Ser Lys Val Pro Ala Ala
50 55 60
Thr Ala Ser Asp Pro Ala Ala Gln Ala Ser Val Ala His Thr Ala Ser
65 70 75 80
Ala Pro Thr Thr Ala Glu His Thr Ala Thr Gly Arg Thr Pro Ala Ala
85 90 95
His Ala Tyr Ala Thr Gln Thr Pro Ala His His Thr Pro Ala Glu Pro
100 105 110
Thr Pro Ala Glu Pro Thr Pro Ala Gln Pro Thr Pro Ala Gln Pro Thr
115 120 125
Pro Ala Gln Pro Thr Pro Ala Gln Pro Thr Pro Ala Gln Pro Thr Pro
130 135 140
Ala Glu Pro Thr Pro Ala Glu Pro Thr Pro Ala Glu Pro Thr Pro Ala
145 150 155 160
Glu Pro Thr Pro Ala Glu Pro Thr Pro Thr Thr Pro Ala Pro Thr Val
165 170 175
Ser Glu Arg Asp Leu Arg Phe Met Ala Leu Ala Gln Ala Ala Ala Glu
180 185 190
Glu Ala Arg Ala Ala Gly Glu Val Pro Val Gly Ala Val Leu Val Arg
195 200 205
Gly Asp Glu Val Ile Ala Lys Gly Phe Asn His Pro Ile Ser Ala His
210 215 220
Asp Pro Ser Ala His Ala Glu Met Val Ala Leu Arg Ala Ala Ala Gln
225 230 235 240
Val Val Glu Asn Tyr Arg Leu Pro Gly Cys Glu Leu Tyr Val Thr Leu
245 250 255
Glu Pro Cys Leu Met Cys Ala Gly Ala Ile Met His Ala Arg Ile Ala
260 265 270
Arg Val Val Phe Gly Val Arg Asn Pro Lys Thr Gly Ala Cys Gly Ser
275 280 285
Val Val Asp Ala Phe Ala Asn Pro Gln Leu Asn His His Thr Glu Val
290 295 300
Thr Gly Gly Val Leu Glu Asn Glu Cys Gly Ala Ala Leu Lys Ser Phe
305 310 315 320
Phe Ala Glu Arg Arg Arg Ala Ser Arg Asp Ala Arg Ala Ala Ala Arg
325 330 335
Ala Gly Asn Gly Ala Gln Ala Glu Pro Asn Glu Pro Arg
340 345
<210> 58
<211> 165
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 58
Glu Val Glu Phe Ser His Glu Tyr Trp Met Arg His Ala Leu Thr Leu
1 5 10 15
Ala Lys Arg Ala Trp Asp Glu Arg Glu Val Pro Val Gly Ala Val Leu
20 25 30
Val His Asn Asn Arg Val Ile Gly Glu Gly Trp Asn Arg Pro Ile Gly
35 40 45
Arg His Asp Pro Thr Ala His Ala Glu Ile Met Ala Leu Arg Gln Gly
50 55 60
Gly Leu Val Met Gln Asn Tyr Arg Leu Ile Asp Ala Thr Leu Tyr Val
65 70 75 80
Thr Leu Glu Pro Cys Val Met Cys Ala Gly Ala Met Ile His Ser Arg
85 90 95
Ile Gly Arg Val Val Phe Gly Val Arg Asn Ala Lys Thr Gly Ala Ala
100 105 110
Gly Ser Leu Met Asp Val Leu His His Pro Gly Met Asn His Arg Val
115 120 125
Glu Ile Thr Glu Gly Ile Leu Ala Asp Glu Cys Ala Ala Leu Leu Ser
130 135 140
Asp Phe Phe Arg Met Arg Arg Gln Glu Ile Lys Ala Gln Lys Lys Ala
145 150 155 160
Gln Ser Ser Thr Asp
165
Claims (30)
1.碱基编辑器，其特征在于，为突变型TadA-8e置于Cas核酸酶的N-末端、C-末端或内部的融合位点融合形成的融合蛋白；
突变型TadA-8e是指TadA-8e中氨基酸位点为第28、30、46、48、49、82、84、108、110或111位的氨基酸中一个或几个突变后的脱氨酶；
TadA-8e的氨基酸序列如SEQ ID NO.1所示。
2.根据权利要求1所述的碱基编辑器，其特征在于，所述Cas核酸酶选自Cas9、Cas12、Cas13蛋白家族；
优选地，所述Cas核酸酶选自nSpCas9及其突变体、SaCas9及其突变体、Cas12a及其突变体或Cas12f及其突变体，优选为nSpCas9。
3.根据权利要求1所述的碱基编辑器，其特征在于，突变型TadA-8e是指TadA-8e中氨基酸位点为第28、30、46、48、49、82、84、108、110或111位的氨基酸中一个突变为不同于所在位点自身原来氨基酸的另外19种氨基酸中的一种以后所得的脱氨酶；
优选地，突变型TadA-8e是指TadA-8e中氨基酸进行以下单点突变后所得的脱氨酶：V28W、V28G、N46L、N46P、N46C、A48C、A48G、A48N、A48S、I49D、I49E、I49H、I49N、I49P、I49Q、I49V、I49W、I49Y、I49R、I49A、I49T、I49G、I49L、I49S、I49M、I49C、I49F、I49K、V82G、V82A、V82T、V82C、V82S、V82D、F84M、N108H、N108M、N108C、N108D、N108F、N108Q、N108S、N108Y、V28A、V28D、V28E、V28K、V28M、V28R、V28L、V28Q、V30A、V30D、V30G、V30W、V30E、V30N、V30Q、V30Y、V30T、V30S、N46H、N46K、N46Y、N46F、N46E、N46Q、N46D、N46G、N46M、N46V、N46S、N46T、A48D、A48E、A48Y、A48W、A48F、A48Q、V82F、F84D、F84R、F84A、F84E、F84P、F84K、F84C、F84H、F84I、F84S、F84T、F84V、F84Q、F84N、F84G、F84Y、N108V、N108I、N108L、N108A、N108T、N108R、K110D、K110W、K110E、R111P、R111E、R111T、R111A、R111V、R111G、R111I、R111W、R111L、R111D。
4.根据权利要求1所述的碱基编辑器，其特征在于，突变型TadA-8e是指TadA-8e中氨基酸进行以下双点突变后所得的脱氨酶：
(1)氨基酸位点第28位的氨基酸突变的同时，第46、48、49、82、84、108、110或111位的氨基酸中一个突变后的脱氨酶，或，
(2)氨基酸位点第46位的氨基酸突变的同时，第28、82、84、108位的氨基酸中一个突变后的脱氨酶，或，
(3)氨基酸位点第48、108位的氨基酸同时突变后的脱氨酶，或，
(4)氨基酸位点第49位的氨基酸突变的同时，第82、84、108位的氨基酸中一个突变后的脱氨酶，或，
(5)氨基酸位点第108位的氨基酸突变的同时，第82或84位的氨基酸中一个突变后的脱氨酶；
优选地，突变型TadA-8e是指TadA-8e中氨基酸进行以下双点突变后所得的脱氨酶：
V28G-N108Y，V28G-N108C，V28G-N108H，V28G-N46L，V28G-N46C，V28G-N46P，V28W-N46L，V28W-N46C，V28W-N46P，N46P-N108Y，N46P-N108S，N46P-N108Q，N46P-N108D，N46P-N108C，N46P-N108H，N46P-F84M，N46P-I49A，N46P-I49G，N46P-A48G，N46C-N108Y，N46C-N108S，N46C-N108Q，N46C-N108M，N46C-N108C，N46C-N108H，N46L-N108Y，N46L-N108Q，N46L-N108D，N46L-N108M，N46L-N108H，N46L-F84M，N46L-I49A，N46L-I49G，N46L-V82G，I49A-N108Y、V28G-A48G、V28G-I49A、V28G-I49G、V28G-V82T、V28G-F84Y、V28W-N108H、N46C-V82G、N46C-V82T、N46C-F84M、N46L-V82T、N46P-F84M、N46P-N108M、A48G-N108H、I49A-N108H、I49G-N108H、I49G-N108Y、I49G-V82G、I49G-V82T、I49G-F84Y、V82G-N108Y、V82T-N108H、V82T-N108C、F84M-N108H、F84M-N108M、F84M-N108Q。
5.根据权利要求1所述的碱基编辑器，其特征在于，所述突变型TadA-8e是指TadA-8e中氨基酸进行以下三点突变后所得的脱氨酶：
(1)氨基酸位点第28、48、49位的氨基酸同时突变后的脱氨酶，或，
(2)氨基酸位点第28、48、82位的氨基酸同时突变后的脱氨酶，或，
(3)氨基酸位点第28、48、108位的氨基酸同时突变后的脱氨酶，或，
(4)氨基酸位点第28、49、82位的氨基酸同时突变后的脱氨酶，或，
(5)氨基酸位点第28、49、108位的氨基酸同时突变后的脱氨酶，或，
(6)氨基酸位点第49、82、108位的氨基酸同时突变后的脱氨酶；
优选地，所述突变型TadA-8e是指TadA-8e中氨基酸进行以下三点突变后所得的脱氨酶：
I49A-V82T-N108Y、I49A-V82G1-N08Y、I49G-V82T-N108Y、I49G-V82G-N108Y、A48G-V82G-N108Y、A48G-I49G-N108Y、V28G-V82G-N108Y、V28G-V82G-N108H、V28G-I49A-N108Y、V28G-I49A-N108H、V28G-I49G-N108H、V28G-A48G-N108Y、V28G-A48G-N108H、V28G-A48G-I49G、V28G-A48G-I49A、V28G-A48G-V82G、V28G-A48G-V82T、V28G-I49G-V82G、V28G-I49G-V82T、V28G-I49G-N108Y、V28G-I49A-V82G、V28G-I49A-V82T、I49G-V82G-N108H、I49G-V82T-N108H。
6.根据权利要求1所述的碱基编辑器，其特征在于，所述突变型TadA-8e是指TadA-8e中氨基酸位点第108位的氨基酸突变的同时，TadA-8e中氨基酸位点第28、48、49、82中还有三个位点突变后的脱氨酶；
优选地，所述突变型TadA-8e是指TadA-8e中氨基酸位点第108位进行突变N108Y或N108H的同时，TadA-8e中氨基酸位点第28、48、49、82中还有三个位点进行如下突变后的脱氨酶；
TadA-8e中氨基酸位点第28位进行突变V28G，
TadA-8e中氨基酸位点第48位进行突变A48G，
TadA-8e中氨基酸位点第49位进行突变I49A或I49G，
TadA-8e中氨基酸位点第82位进行突变V82T或V82G。
7.根据权利要求1所述的碱基编辑器，其特征在于，所述突变型TadA-8e是指TadA-8e中氨基酸位点第28、48、49、82、108位的氨基酸同时突变后的脱氨酶；
优选地，所述突变型TadA-8e是指TadA-8e中氨基酸进行以下五点突变后所得的脱氨酶：
TadA-8e中氨基酸位点第108位进行突变N108Y或N108H的同时，TadA-8e中氨基酸位点第28、48、49、82位点还进行如下突变后的脱氨酶；
TadA-8e中氨基酸位点第28位进行突变V28G，
TadA-8e中氨基酸位点第48位进行突变A48G，
TadA-8e中氨基酸位点第49位进行突变I49A或I49G，
TadA-8e中氨基酸位点第82位进行突变V82T或V82G。
8.根据权利要求1或2或3或4或5或6或7所述的碱基编辑器，其特征在于，突变型TadA-8e的5’和3’分别添加不同序列的各类变体后置于Cas核酸酶的N-末端、C-末端或内部的融合位点融合形成的融合蛋白；
优选地，突变型TadA-8e的5’和3’分别添加NLS序列和linker序列后获得NLS-突变型TadA-8e-linker，NLS-突变型TadA-8e-linker置于Cas核酸酶的N-末端、C-末端或内部的融合位点融合形成的融合蛋白；
其中，NLS-突变型TadA-8e-linker是指相对于NLS-TadA-8e-linker而言，TadA-8e中氨基酸中一个或几个发生了突变，NLS-TadA-8e-linker的氨基酸序列如SEQ ID NO.4所示，NLS-突变型TadA-8e-linker的氨基酸序列为NLS-TadA-8e-linker氨基酸序列中将TadA-8e置换为突变型TadA-8e得到的氨基酸序列。
9.根据权利要求1或2或3或4或5或6或7所述的碱基编辑器，其特征在于，所述Cas核酸酶选择为nSpCas9时，内部的融合位点包括第113、203、312、459、535、583、687、701、715、730、770、793、801、895、905、919、946、1010、1029、1047、1117、1154、1249、1282位融合位点；
优选为第1249位融合位点。
10.碱基编辑器，其特征在于，TadA-8e的5’和3’分别添加NLS序列和linker序列后获得NLS-TadA-8e-linker，NLS-TadA-8e-linker置于Cas核酸酶的N-末端、C-末端或内部的融合位点融合形成的融合蛋白；
NLS-TadA-8e-linker的氨基酸序列如SEQ ID NO.4所示；
所述Cas核酸酶选自Cas9、Cas12、Cas13蛋白家族，优选地，所述Cas核酸酶选自nSpCas9及其突变体、SaCas9及其突变体、Cas12a及其突变体或Cas12f及其突变体，进一步优选为nSpCas9。
11.根据权利要求10所述的碱基编辑器，其特征在于，所述Cas核酸酶选择为nSpCas9时，所述nSpCas9内部的融合位点包括第113、203、312、459、535、583、687、701、715、730、770、793、801、895、905、919、946、1010、1029、1047、1117、1154、1249、1282位融合位点；
优选为第1249位融合位点。
12.碱基编辑器，其特征在于，为突变型TadA-X置于Cas核酸酶的N-末端、C-末端或内部的融合位点融合形成的融合蛋白；
突变型TadA-X是指TadA-X中与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺的脱氨酶；
TadA-X是指不同来源的TadA；
优选地，TadA-X有多组，其突变体氨基酸序列分别如SEQ ID NO.5-SEQ ID NO.57中的任一所示；
所述Cas核酸酶选自Cas9、Cas12、Cas13蛋白家族，优选地，所述Cas核酸酶选自nSpCas9及其突变体、SaCas9及其突变体、Cas12a及其突变体或Cas12f及其突变体，进一步优选为nSpCas9。
13.根据权利要求12所述的碱基编辑器，其特征在于，突变型TadA-X是指Q2LYC0、F2G306、G0JN58、B5ZCW4、Q1AS41、E4TTF3、Q57LE3、Q99W51、ecTadA中与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺的脱氨酶；
Q2LYC0、F2G306、G0JN58、B5ZCW4、Q1AS41、E4TTF3、Q57LE3、Q99W51或ecTadA突变体的氨基酸序列分别为SEQ ID NO.8、SEQ ID NO.14、SEQ ID NO.20、SEQ ID NO.22、SEQ IDNO.27、SEQ ID NO.43、SEQ ID NO.54、SEQ ID NO.55、SEQ ID NO.58。
14.根据权利要求12所述的碱基编辑器，其特征在于，突变型TadA-X是指B3PCY2、A5WFT9、C1DXN9、B2UR34、R4K908、D5W8R4、E1TAF4中与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺的脱氨酶；
B3PCY2、A5WFT9、C1DXN9、B2UR34、R4K908、D5W8R4、E1TAF4突变体的氨基酸序列分别为SEQ ID NO.15、SEQ ID NO.19、SEQ ID NO.29、SEQ ID NO.32、SEQ ID NO.44、SEQ IDNO.49、SEQ ID NO.57。
15.根据权利要求12所述的碱基编辑器，其特征在于，突变型TadA-X是指Q13XZ4、A8F050、E8WVH3、P44931、G0EXN4、E0TBL5、A1W869、D5X1Y1、I3YB54与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺的脱氨酶；
Q13XZ4、A8F050、E8WVH3、P44931、G0EXN4、E0TBL5、A1W869、D5X1Y1、I3YB54突变体的氨基酸序列分别为SEQ ID NO.10、SEQ ID NO.18、SEQ ID NO.25、SEQ ID NO.28、SEQ IDNO.38、SEQ ID NO.46、SEQ ID NO.48、SEQ ID NO.50、SEQ ID NO.53。
16.根据权利要求12或13或14或15所述的碱基编辑器，其特征在于，突变型TadA-X的5’和3’分别添加NLS序列和linker序列后获得NLS-突变型TadA-X-linker，NLS-突变型TadA-X-linker置于Cas核酸酶的N-末端、C-末端或内部的融合位点融合形成的融合蛋白；
其中，NLS-突变型TadA-X-linker是指相对于NLS-TadA-8e-linker而言，TadA-8e替换为突变型TadA-X，NLS-TadA-8e-linker的氨基酸序列如SEQ ID NO.4所示。
17.根据权利要求12或13或14或15所述的碱基编辑器，其特征在于，所述Cas核酸酶选择为nSpCas9时，所述nSpCas9内部的融合位点包括第113、203、312、459、535、583、687、701、715、730、770、793、801、895、905、919、946、1010、1029、1047、1117、1154、1249、1282位融合位点；
优选为第1249位融合位点。
18.碱基编辑器的构建方法，其特征在于，包括以下步骤：
将TadA-8e、突变型TadA-8e或突变型TadA-X和Cas核酸酶不同位点融合，构建得到碱基编辑器；
TadA-8e的氨基酸序列如SEQ ID NO.1所示；
突变型TadA-8e是指TadA-8e中氨基酸位点为第28、30、46、48、49、82、84、108、110或111位的氨基酸中一个或几个突变后的脱氨酶；
突变型TadA-X是指TadA-X中与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺的脱氨酶；TadA-X有多组，其突变体氨基酸序列分别如SEQ ID NO.5-SEQ IDNO.57中的任一所示。
19.根据权利要求18所述的碱基编辑器的构建方法，其特征在于，还可以将TadA-8e、突变型TadA-8e或突变型TadA-X 5’和3’分别添加NLS序列和linker序列。
20.碱基编辑器ABE的应用，其特征在于，所述碱基编辑器ABE介导A·T--G·C突变；
碱基编辑器ABE是指Q2LYC0、F2G306、G0JN58、B5ZCW4、Q1AS41、E4TTF3、Q57LE3、Q99W51、ecTadA与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺后置于nSpCas9的N-末端、第1249位融合位点融合形成的融合蛋白，其中，Q2LYC0、F2G306、G0JN58、B5ZCW4、Q1AS41、E4TTF3、Q57LE3、Q99W51或ecTadA突变体的氨基酸序列分别为SEQ ID NO.8、SEQ IDNO.14、SEQ ID NO.20、SEQ ID NO.22、SEQ ID NO.27、SEQ ID NO.43、SEQ ID NO.54、SEQ IDNO.55、SEQ ID NO.58；
碱基编辑器ABE还指TadA-8e中氨基酸进行以下单点突变或点突变组合后所得的脱氨酶置于nSpCas9的N-末端、第1249位融合位点融合形成的融合蛋白，这类融合蛋白仅具有明显的介导A·T--G·C突变能力。
21.碱基编辑器CBE的应用，其特征在于，所述碱基编辑器CBE介导C·G--T·A突变；
碱基编辑器CBE是指B3PCY2、A5WFT9、C1DXN9、B2UR34、R4K908、D5W8R4、E1TAF4与ecTadA第106和第108位同源的氨基酸分别突变为缬氨酸和天冬酰胺后置于nSpCas9的N-末端、第1249位融合位点融合形成的融合蛋白，其中，B3PCY2、A5WFT9、C1DXN9、B2UR34、R4K908、D5W8R4、E1TAF4突变体的氨基酸序列分别为SEQ ID NO.15、SEQ ID NO.19、SEQ ID NO.29、SEQ ID NO.32、SEQ ID NO.44、SEQ ID NO.49、SEQ ID NO.57；
碱基编辑器CBE还指TadA-8e中氨基酸进行以下突变后所得的脱氨酶置于nSpCas9的N-末端、第1249位融合位点融合形成的融合蛋白，
单点突变：N46P
双点突变：V28G-N108Y，V28G-N108C，V28G-N108H，V28G-N46L，V28G-N46C，V28G-N46P，V28W-N46L，V28W-N46C，V28W-N46P，N46P-N108Y，N46P-N108S，N46P-N108Q，N46P-N108D，N46P-N108C，N46P-N108H，N46P-F84M，N46P-I49A，N46P-I49G，N46P-A48G，N46C-N108Y，N46C-N108S，N46C-N108Q，N46C-N108M，N46C-N108C，N46C-N108H，N46L-N108Y，N46L-N108Q，N46L-N108D，N46L-N108M，N46L-N108H，N46L-F84M，N46L-I49A，N46L-I49G，N46L-V82G，I49A-N108Y
三点突变：I49A-V82T-N108Y、I49A-V82G1-N08Y、I49G-V82T-N108Y、I49G-V82G-N108Y、A48G-V82G-N108Y、A48G-I49G-N108Y、V28G-V82G-N108Y、V28G-V82G-N108H、V28G-I49A-N108Y、V28G-I49A-N108H、V28G-I49G-N108H、V28G-A48G-N108Y、V28G-A48G-N108H；
四点突变：V28G-A48G-I49A-N108Y、V28G-A48G-I49G-N108Y、V28G-A48G-V82T-N108Y、V28G-A48G-V82G-N108Y、V28G-I49A-V82T-N108Y、V28G-I49A-V82G-N108Y、V28G-I49G-V82T-N108Y、V28G-I49G-V82G-N108Y、A48G-I49A-V82T-N108Y、A48G-I49A-V82G-N108Y、A48G-I49G-V82T-N108Y、A48G-I49G-V82G-N108Y、V28G-A48G-I49A-N108H、V28G-A48G-I49G-N108H、V28G-A48G-V82T-N108H、V28G-A48G-V82G-N108H、V28G-I49A-V82T-N108H、V28G-I49A-V82G-N108H、V28G-I49G-V82T-N108H、V28G-I49G-V82G-N108H、A48G-I49A-V82T-N108H、A48G-I49A-V82G-N108H、A48G-I49G-V82T-N108H、A48G-I49G-V82G-N108H。
五点突变：V28G-A48G-I49A-V82T-N108Y、V28G-A48G-I49 G-V82T-N108Y、V28G-A48G-I49A-V82G-N108Y、V28G-A48G-I49G-V82G-N108Y、V28G-A48G-I49A-V82T-N108H、V28G-A48G-I49G-V82T-N108H、V28G-A48G-I49A-V82G-N108H、V28G-A48G-I49G-V82G-N108H。
22.碱基编辑器ACBE的应用，其特征在于，所述碱基编辑器ACBE介导A·T--G·C和C·G--T·A突变；
碱基编辑器ACBE是指Q13XZ4、A8F050、E8WVH3、P44931、G0EXN4、E0TBL5、A1W869、D5X1Y1、I3YB54与ecTadA第106和第108位同源的氨基酸分别突变为缬氨酸和天冬酰胺后置于nSpCas9的N-末端、第1249位融合位点融合形成的融合蛋白，其中，Q13XZ4、A8F050、E8WVH3、P44931、G0EXN4、E0TBL5、A1W869、D5X1Y1、I3YB54突变体的氨基酸序列分别为SEQID NO.10、SEQ ID NO.18、SEQ ID NO.25、SEQ ID NO.28、SEQ ID NO.38、SEQ ID NO.46、SEQID NO.48、SEQ ID NO.50、SEQ ID NO.53；
碱基编辑器ACBE还指TadA-8e中氨基酸进行点突变或点突变组合后所得的脱氨酶置于nSpCas9的N-末端、第1249位融合位点融合形成的融合蛋白，这类融合蛋白具有介导A·T--G·C和C·G--T·A突变能力。
23.突变型TadA-8e蛋白，其特征在于，所述突变蛋白为非天然蛋白，突变型TadA-8e蛋白指TadA-8e中氨基酸位点为第28、30、46、48、49、82、84、108、110或111位的氨基酸中一个或几个突变后的脱氨酶；
优选地，突变型TadA-8e是指TadA-8e中氨基酸进行以下单点突变后所得的脱氨酶：
V28W、V28G、N46L、N46P、N46C、A48C、A48G、A48N、A48S、I49D、I49E、I49H、I49N、I49P、I49Q、I49V、I49W、I49Y、I49R、I49A、I49T、I49G、I49L、I49S、I49M、I49C、I49F、I49K、V82G、V82A、V82T、V82C、V82S、V82D、F84M、N108H、N108M、N108C、N108D、N108F、N108Q、N108S、N108Y、V28A、V28D、V28E、V28K、V28M、V28R、V28L、V28Q、V30A、V30D、V30G、V30W、V30E、V30N、V30Q、V30Y、V30T、V30S、N46H、N46K、N46Y、N46F、N46E、N46Q、N46D、N46G、N46M、N46V、N46S、N46T、A48D、A48E、A48Y、A48W、A48F、A48Q、V82F、F84D、F84R、F84A、F84E、F84P、F84K、F84C、F84H、F84I、F84S、F84T、F84V、F84Q、F84N、F84G、F84Y、N108V、N108I、N108L、N108A、N108T、N108R、K110D、K110W、K110E、R111P、R111E、R111T、R111A、R111V、R111G、R111I、R111W、R111L、R111D；
或，突变型TadA-8e是指TadA-8e中氨基酸进行以下双点突变后所得的脱氨酶：
V28G-N108Y，V28G-N108C，V28G-N108H，V28G-N46L，V28G-N46C，V28G-N46P，V28W-N46L，V28W-N46C，V28W-N46P，N46P-N108Y，N46P-N108S，N46P-N108Q，N46P-N108D，N46P-N108C，N46P-N108H，N46P-F84M，N46P-I49A，N46P-I49G，N46P-A48G，N46C-N108Y，N46C-N108S，N46C-N108Q，N46C-N108M，N46C-N108C，N46C-N108H，N46L-N108Y，N46L-N108Q，N46L-N108D，N46L-N108M，N46L-N108H，N46L-F84M，N46L-I49A，N46L-I49G，N46L-V82G，I49A-N108Y、V28G-A48G、V28G-I49A、V28G-I49G、V28G-V82T、V28G-F84Y、V28W-N108H、N46C-V82G、N46C-V82T、N46C-F84M、N46L-V82T、N46P-F84M、N46P-N108M、A48G-N108H、I49A-N108H、I49G-N108H、I49G-N108Y、I49G-V82G、I49G-V82T、I49G-F84Y、V82G-N108Y、V82T-N108H、V82T-N108C、F84M-N108H、F84M-N108M、F84M-N108Q；
或，所述突变型TadA-8e是指TadA-8e中氨基酸进行以下三点突变后所得的脱氨酶：
I49A-V82T-N108Y、I49A-V82G1-N08Y、I49G-V82T-N108Y、I49G-V82G-N108Y、A48G-V82G-N108Y、A48G-I49G-N108Y、V28G-V82G-N108Y、V28G-V82G-N108H、V28G-I49A-N108Y、V28G-I49A-N108H、V28G-I49G-N108H、V28G-A48G-N108Y、V28G-A48G-N108H、V28G-A48G-I49G、V28G-A48G-I49A、V28G-A48G-V82G、V28G-A48G-V82T、V28G-I49G-V82G、V28G-I49G-V82T、V28G-I49G-N108Y、V28G-I49A-V82G、V28G-I49A-V82T、I49G-V82G-N108H、I49G-V82T-N108H；
或，所述突变型TadA-8e是指TadA-8e中氨基酸进行以下四点突变后所得的脱氨酶：V28G-A48G-I49A-N108Y、V28G-A48G-I49G-N108Y、V28G-A48G-V82T-N108Y、V28G-A48G-V82G-N108Y、V28G-I49A-V82T-N108Y、V28G-I49A-V82G-N108Y、V28G-I49G-V82T-N108Y、V28G-I49G-V82G-N108Y、A48G-I49A-V82T-N108Y、A48G-I49A-V82G-N108Y、A48G-I49G-V82T-N108Y、A48G-I49G-V82G-N108Y、V28G-A48G-I49A-N108H、V28G-A48G-I49G-N108H、V28G-A48G-V82T-N108H、V28G-A48G-V82G-N108H、V28G-I49A-V82T-N108H、V28G-I49A-V82G-N108H、V28G-I49G-V82T-N108H、V28G-I49G-V82G-N108H、A48G-I49A-V82T-N108H、A48G-I49A-V82G-N108H、A48G-I49G-V82T-N108H、A48G-I49G-V82G-N108H；
或，所述突变型TadA-8e是指TadA-8e中氨基酸进行以下五点突变后所得的脱氨酶：V28G-A48G-I49A-V82T-N108Y、V28G-A48G-I49 G-V82T-N108Y、V28G-A48G-I49A-V82G-N108Y、V28G-A48G-I49G-V82G-N108Y、V28G-A48G-I49A-V82T-N108H、V28G-A48G-I49G-V82T-N108H、V28G-A48G-I49A-V82G-N108H、V28G-A48G-I49G-V82G-N108H。
24.腺嘌呤脱氨酶TadA突变蛋白，其特征在于，所述腺嘌呤脱氨酶TadA突变蛋白为非天然蛋白，所述腺嘌呤脱氨酶TadA突变蛋白是指TadA-X中与ecTadA第106和第108同源的氨基酸分别突变为缬氨酸和天冬酰胺的脱氨酶；
TadA-X是指不同来源的TadA；优选地，TadA-X有多组，其突变体氨基酸序列分别如SEQID NO.5-SEQ ID NO.57中的任一所示。
25.根据权利要求23所述的突变型TadA-8e蛋白或根据权利要求24所述的腺嘌呤脱氨酶TadA突变蛋白，其特征在于，突变蛋白为单体或二聚体。
26.多核苷酸，其特征在于，编码权利要求1-17中任一项所述碱基编辑器中的融合蛋白。
27.多核苷酸，其特征在于，编码权利要求23、24或25任一项所述蛋白。
28.载体，其特征在于，含有权利要求26或27所述的多核苷酸。
29.宿主细胞，其特征在于，含有权利要求1-17中任一项所述碱基编辑器，或含有权利要求23、24或25任一项所述蛋白，或含有权利要求28所述载体。
30.一种试剂盒，其特征在于，包含用于构建权利要求1-17中任一项所述碱基编辑器的试剂。
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