一种碱基编辑工具及其用途的制作方法

本发明涉及生物
技术领域：
，特别是涉及一种碱基编辑工具及其用途。
背景技术：
：自2013年crispr/cas9被发表于应用真核生物细胞中的基因编辑。此后，基于crispr/cas9系统的基因编辑技术得到了极大的发展。该系统仅由两个部分组成：负责定位靶位点序列的引导rna(guiderna，grna)和核酸内切酶cas9蛋白。二者配合能够高效特异的靶向切割目的位点，造成dna双链断裂(doublestrainbreak,bsd)，从而允许人们利用细胞自身的非同源重组修复(non-homologousendjoining,nhej)途径产生dna片段的缺失或引起移码突变，从而造成基因的敲除。人们也可以利用细胞的同源重组修复(homologydirectedrepair,hdr)途径对靶位点进行精确的dna片段替换或者敲入。随着crispr系统的研究逐渐深入，研究人员发现基于dna双链断裂的基因编辑存在种种问题。首先，就是编辑产物的不可控性。细胞对dna上dsb位点进行的nhej途径修复产物是随机的，有时只会产生极小片段丢失且没有造成移码突变，因此虽然能够产生dsb，但依然不能保证高效的敲除效率。其次，基于hdr修复途径的编辑效率始终不高，很难在体实现高效的基因编辑效率。最后，crispr/cas9系统的脱靶效应也会导致在编辑过程中对基因组上其他位点上产生不可回复的序列改变。人类绝大多数的遗传疾病都是单碱基突变导致。因此，针对以上问题，开发能够精确编辑单碱基的技术将为能为基础研究和临床疾病治疗提供极大助益。2017年，哈佛大学davidrliu实验室在nature上报道了一种基于cas9的单碱基编辑(baseediting，be)工具。该系统利用ncas9和apobec1以及ugi融合能够高效实现胞嘧啶c到胸腺嘧啶t的定向单碱基编辑。单碱基编辑技术一经发表便得到了广泛的关注和应用，研究人员在不同的细胞系以及植物和动物中都实现了高效的编辑。随着剪辑编辑技术的广泛应用，科研人员也一直开发更精确和灵敏的脱靶检测技术对be进行更加高要求的检验。2019年，杨辉和高彩霞实验室分别独立在science上报道了cbe会产生非grna依赖的dna脱靶情况。在培养细胞系中，每个细胞内产生的随机脱靶都不同，这些脱靶位点在细胞群体中会被稀释，从而无法检测。杨辉团队开发了一种更敏感的无偏差脱靶检测方法goti来检测be3的脱靶情况，巧妙利用小鼠胚胎发育放大脱靶位点，便于检测。由于dna上的随机脱靶无法预测并无法恢复，因此这一脱靶现象也引起了大家对cbe在临床治疗应用上的担忧。同年，keithjoung实验室和杨辉实验室先后在nature上报道了cbe在转录组上的严重脱靶情况，be3能诱导原癌基因和抑癌基因等基因上的数百个基因突变，还可能引起其他严重危害健康的突变。真核细胞中rna虽然不会被遗传，但是理论上所有rna都会以表达蛋白或者直接参与细胞功能的调控，因此，脱靶突变的产生也会对细胞产生直接影响。通过对脱氨酶进行氨基酸突变，能够部分消除be在rna上的脱靶编辑。但是，这种方法无法完全保证成功，消除脱靶编辑的同时，可能也会损失靶向编辑效率。此外，针对每种脱氨酶，都需要从头进化并验证，因此该种方法的工作量也巨大。最后be3在dna上造成的随机脱靶问题仍是个问题。因此，亟待开发一个通用、方便、低成本的进化技术或者策略对降低be造成的rna和dna脱靶。技术实现要素：鉴于以上所述现有技术的缺点，本发明的目的在于提供一种碱基编辑工具及其用途，用于解决现有技术中的问题。为实现上述目的及其他相关目的，本发明一方面提供一种融合蛋白，所述融合蛋白自n端至c端依次包括第一ncas9片段、嵌合插入片段、第二ncas9片段和两个ugi片段，所述嵌合插入片段选自apobec1片段或apobec3a片段。在本发明一些实施方式中，所述第一ncas9片段的氨基酸序列包括：a)如seqidno.1所示的氨基酸序列；或，b)与seqidno.1具有80％以上序列相似性、且具有a)所限定的氨基酸序列的功能的氨基酸序列，优选为具有ncas9靶向活性；和/或，所述第二ncas9片段的氨基酸序列包括：c)如seqidno.2所示的氨基酸序列；或，d)与seqidno.2具有80％以上序列相似性、且具有e)所限定的氨基酸序列的功能的氨基酸序列，优选为具有ncas9靶向活性。在本发明一些实施方式中，所述apobec1片段的氨基酸序列包括：e)如seqidno.3所示的氨基酸序列；或，f)与seqidno.3具有80％以上序列相似性的氨基酸序列、且具有a)所限定的氨基酸序列的功能，优选为具有胞嘧啶脱氨酶活性。在本发明一些实施方式中，所述apobec3a片段的氨基酸序列包括：i)如seqidno.4所示的氨基酸序列；或，j)与seqidno.4具有80％以上序列相似性的氨基酸序列、且具有c)所限定的氨基酸序列的功能，优选为具有胞嘧啶脱氨酶活性。在本发明一些实施方式中，所述ugi片段的氨基酸序列包括：k)如seqidno.5所示的氨基酸序列；或，l)与seqidno.5具有80％以上序列相似性的氨基酸序列、且具有c)所限定的氨基酸序列的功能，优选为具有抑制尿嘧啶dna糖基化活性。在本发明一些实施方式中，所述融合蛋白还包括核定位信号片段，优选的，所述核定位信号片段的氨基酸序列包括seqidno.6所示的氨基酸序列。在本发明一些实施方式中，所述融合蛋白还包括柔性连接肽片段，优选的，所述柔性连接肽片段的氨基酸序列包括seqidno.7或seqidno.8所示的氨基酸序列。在本发明一些实施方式中，所述融合蛋白的氨基酸序列如seqidno.9～10其中之一所示。本发明另一方面提供一种分离的多核苷酸，编码上述的融合蛋白。本发明另一方面提供一种构建体，所述构建体含有上述的分离的多核苷酸。本发明另一方面提供一种表达系统，所述表达系统含有上述的构建体或基因组中整合有外源的上述的多核苷酸。在本发明一些实施方式中，所述表达系统的宿主细胞选自真核细胞或原核细胞，优选选自小鼠细胞、人细胞，更优选选自小鼠脑神经瘤细胞、人胚胎肾细胞、或人宫颈癌细胞、人结肠癌细胞、人骨肉瘤细胞，更优选选自n2a细胞、hek293ft细胞、hela细胞、hct116细胞、或u2os细胞。本发明另一方面提供上述的融合蛋白、上述的分离的多核苷酸、上述的构建体或上述的表达系统在基因编辑中的用途。在本发明一些实施方式中，所述用途具体为在真核生物的基因编辑中的用途。本发明另一方面提供一种碱基编辑体系，包括上述的融合蛋白，所述碱基编辑体系还包括sgrna。本发明另一方面提供一种基因编辑方法，包括：通过上述的融合蛋白、或上述的碱基编辑体系进行基因编辑。附图说明图1显示为本发明基于mu转座酶构建ncas9随机插入文库的示意图。图2显示为本发明筛选得到的ncas9有效插入位点和碱基编辑效率结果示意图。图3显示为本发明比较spcas9的同源物的不保守区域的结果示意图。图4显示为本发明筛选得到的ce-abe在人细胞基因组上的碱基编辑结果示意图。图5显示为本发明ce-abe在预测rna位点上脱靶编辑结果示意图。图6显示为本发明ce-abe在转录组水平产生的脱靶编辑结果示意图。图7显示为本发明ce-abe在脱靶检测样本中的靶向编辑效率结果示意图。图8显示为本发明ce-abe1048-1063和abemax在293t细胞中具有相当的编辑效率结果示意图。图9显示为本发明ce-abe1048-1063和abemax在n2a细胞中具有相当的编辑效率结果示意图。图10显示为本发明ce-be1048-1063和ancbe4max在293t细胞中具有相当的编辑效率结果示意图。图11显示为本发明ce-a3a1048-1063和be-a3a在293t细胞中具有相当的编辑效率结果示意图。图12显示为本发明ce-be和ancbe4max在293t细胞中rna上的脱靶编辑情况示意图。图13显示为本发明ce-a3a和be-a3a在293t细胞中rna上的脱靶编辑情况示意图。图14显示为本发明be4max、be-a3a、ce-be1048-1063和ce-a3a1048-1063在dna上的靶向编辑结果示意图。图15显示为本发明be4max、ce-be1048-1063(ce-be4max)在dna上的脱靶编辑结果示意图。图16显示为本发明be-a3a和ce-a3a1048-1063(ce-a3a)在dna上的脱靶编辑结果示意图。具体实施方式本发明发明人经过大量探索性研究，发现在ncas9蛋白内部的合适位置嵌合融合功能片段，能够极大地同时降低be在rna和dna上的脱靶情况，并且不影响be的靶向编辑效率，在此基础上完成了本发明。本发明第一方面提供一种融合蛋白，所述融合蛋白自n端至c端依次包括第一ncas9片段、嵌合插入片段、第二ncas9片段和两个ugi片段，所述嵌合插入片段选自apobec1片段或apobec3a片段。所述融合蛋白是将ncas9(genbank:mk048158.1)的1048thr-1063ile片段替换为嵌合插入片段，在sgrna的引导下在靶向位点做碱基编辑，能够极大地同时降低be在rna和dna上的脱靶情况，并且不影响be的靶向编辑效率。本发明所提供的融合蛋白中，所述第一ncas9片段的氨基酸序列可以包括：a)如seqidno.1所示的氨基酸序列；或，b)与seqidno.1具有80％以上序列相似性的氨基酸序列、且具有a)所限定的氨基酸序列的功能。具体的，所述b)中的氨基酸序列具体指：如seqidno.1所示的氨基酸序列经过取代、缺失或者添加一个或多个(具体可以是1-50、1-30个、1-20个、1-10个、1-5个、1-3个、1个、2个、或3个)氨基酸而得到的，或者在n-末端和/或c-末端添加一个或多个(具体可以是1-50个、1-30个、1-20个、1-10个、1-5个、1-3个、1个、2个、或3个)氨基酸而得到的，且具有氨基酸如seqidno.1所示的多肽片段的功能的多肽片段，例如，可以是ncas9第一片段与ncas9第二片段配合后依然具有ncas9的靶向活性，更具体可以是能够在合适的grna的引导下靶向dna的活性。所述b)中的氨基酸序列可与seqidno.1具有80％、85％、90％、93％、95％、97％、或99％以上的相似性。所述第一ncas9片段通常源自大肠杆菌。本申请中，序列的相似性(sequenceidentity)通常指参与对比的序列中相同的氨基酸残基的百分比，可采用本领域公知的计算软件计算两条或多条目的序列的相似性，例如，可以是来自ncbi的软件。本发明所提供的融合蛋白中，所述第二ncas9片段的氨基酸序列可以包括：c)如seqidno.2所示的氨基酸序列；或，d)与seqidno.2具有80％以上序列相似性的氨基酸序列、且具有c)所限定的氨基酸序列的功能。具体的，所述d)中的氨基酸序列具体指：如seqidno.2所示的氨基酸序列经过取代、缺失或者添加一个或多个(具体可以是1-50、1-30个、1-20个、1-10个、1-5个、1-3个、1个、2个、或3个)氨基酸而得到的，或者在n-末端和/或c-末端添加一个或多个(具体可以是1-50个、1-30个、1-20个、1-10个、1-5个、1-3个、1个、2个、或3个)氨基酸而得到的，且具有氨基酸如seqidno.2所示的多肽片段的功能的多肽片段，例如，可以是ncas9第一片段与ncas9第二片段配合后依然具有ncas9的靶向活性，更具体可以是能够在合适的grna的引导下靶向dna的活性。所述d)中的氨基酸序列可与seqidno.2具有80％、85％、90％、93％、95％、97％、或99％以上的相似性。所述第二ncas9片段通常源自大肠杆菌(streptococcuspyogenes)。本发明所提供的融合蛋白中，所述apobec1片段的氨基酸序列可以包括：e)如seqidno.3所示的氨基酸序列；或，f)与seqidno.3具有80％以上序列相似性的氨基酸序列、且具有e)所限定的氨基酸序列的功能。具体的，所述f)中的氨基酸序列具体指：如seqidno.3所示的氨基酸序列经过取代、缺失或者添加一个或多个(具体可以是1-50、1-30个、1-20个、1-10个、1-5个、1-3个、1个、2个、或3个)氨基酸而得到的，或者在n-末端和/或c-末端添加一个或多个(具体可以是1-50个、1-30个、1-20个、1-10个、1-5个、1-3个、1个、2个、或3个)氨基酸而得到的，且具有氨基酸如seqidno.3所示的多肽片段的功能的多肽片段，例如，可以是具有胞嘧啶脱氨酶活性，更具体可以是将胞嘧啶(cytosine，c)脱氨产生尿嘧啶(uracil，u)的功能。所述f)中的氨基酸序列可与seqidno.3具有80％、85％、90％、93％、95％、97％、或99％以上的相似性。所述apobec1片段通常源自大鼠(rat)。本发明所提供的融合蛋白中，所述apobec3a片段的氨基酸序列可以包括：g)如seqidno.4所示的氨基酸序列；或，h)与seqidno.4具有80％以上序列相似性的氨基酸序列、且具有g)所限定的氨基酸序列的功能。具体的，所述h)中的氨基酸序列具体指：如seqidno.4所示的氨基酸序列经过取代、缺失或者添加一个或多个(具体可以是1-50、1-30个、1-20个、1-10个、1-5个、1-3个、1个、2个、或3个)氨基酸而得到的，或者在n-末端和/或c-末端添加一个或多个(具体可以是1-50个、1-30个、1-20个、1-10个、1-5个、1-3个、1个、2个、或3个)氨基酸而得到的，且具有氨基酸如seqidno.4所示的多肽片段的功能的多肽片段，例如，可以是具有胞嘧啶脱氨酶活性，更具体可以是将胞嘧啶(cytosine，c)脱氨产生尿嘧啶(uracil，u)的功能。所述h)中的氨基酸序列可与seqidno.4具有80％、85％、90％、93％、95％、97％、或99％以上的相似性。所述apobec3a片段通常源自人。本发明所提供的融合蛋白中，可以包括两个独立的ugi片段，这两个ugi片段的氨基酸序列可以各自独立地包括：i)如seqidno.5所示的氨基酸序列；或，j)与seqidno.5具有80％以上序列相似性的氨基酸序列、且具有i)所限定的氨基酸序列的功能。具体的，所述j)中的氨基酸序列具体指：如seqidno.5所示的氨基酸序列经过取代、缺失或者添加一个或多个(具体可以是1-50、1-30个、1-20个、1-10个、1-5个、1-3个、1个、2个、或3个)氨基酸而得到的，或者在n-末端和/或c-末端添加一个或多个(具体可以是1-50个、1-30个、1-20个、1-10个、1-5个、1-3个、1个、2个、或3个)氨基酸而得到的，且具有氨基酸如seqidno.5所示的多肽片段的功能的多肽片段，例如，可以是具有抑制尿嘧啶dna糖基化反应的功能。所述j)中的氨基酸序列可与seqidno.5具有80％、85％、90％、93％、95％、97％、或99％以上的相似性。所述ugi片段通常来自枯草芽孢杆菌噬菌体(bacillussubtilisbacteriophage)。本发明所提供的融合蛋白中，所述的取代、缺失或者添加可以是保守氨基酸取代。所述“保守氨基酸取代”具体可以是指氨基酸残基被其他具有相似侧链的氨基酸残基取代的情况。具有相似侧链的氨基酸残基家族对于本领域技术人员来说应该是已知的，例如，可以是包括但不限于碱性侧链(例如赖氨酸，精氨酸，组氨酸)，酸性侧链(例如天冬氨酸，谷氨酸)，不带电荷的极性侧链(例如，甘氨酸，天冬酰胺，谷氨酰胺，丝氨酸，苏氨酸，酪氨酸，半胱氨酸)，非极性侧链(例如丙氨酸，缬氨酸，亮氨酸，异亮氨酸，脯氨酸，苯丙氨酸，甲硫氨酸，色氨酸)异亮氨酸)和芳族侧链(例如酪氨酸，苯丙氨酸，色氨酸，组氨酸)等家族。保守型氨基酸取代更具体可以包括但不限于下表中所列的具体情况，表1(氨基酸相似度矩阵)中的数字表示两个氨基酸之间的相似度，当数字大于等于0时认为是保守氨基酸取代，表2为示例性的保守氨基酸取代的方案。表1cgpsatdenqhkrvmilfyww-8-7-6-2-6-5-7-7-4-5-3-32-6-4-5-20017y0-5-5-3-3-3-4-4-2-40-4-5-2-2-1-1710f-4-5-5-3-4-3-6-5-4-5-2-5-4-10129l-6-4-3-3-2-2-4-3-3-2-2-3-32426i-2-3-2-1-10-2-2-2-2-2-2-2425m-5-3-2-2-1-1-3-20-1-20026v-2-1-1-100-2-2-2-2-2-2-24r-4-300-2-1-1-101236k-5-2-10-10001105h-3-20-1-1-111236q-5-10-10-12214n-40-1100212e-50-100034d-51-10004t-200113a-21112s0111p-3-16g-35c12表2本发明所提供的融合蛋白中，所述融合蛋白还可以包括核定位信号片段(bpnls片段)，所述核定位信号片段通常可以与入核载体相互作用，从而可以使蛋白能够被运送进细胞核。所述核定位信号片段可以位于ncas9第一段片段的n端以及两个ugi片段中第二个ugi的c端，即整个融合蛋白两端各一个bpnls片段。所述bpnls片段的氨基酸序列可以包括seqidno.6所示的氨基酸序列。本发明所提供的融合蛋白中，所述融合蛋白还可以包括柔性连接肽片段。所述柔性连接肽片段通常是成柔软线性的一类易弯折的氨基酸片段，通常可以使连接的两个蛋白之间有一定的活动空间。例如，柔性连接肽片段可以是xten肽段等。所述柔性连接肽片段(例如，xten肽段)可以位于ncas9第一个片段和嵌入片段(abobec1或apobec3a)之间，也可以位于嵌入片段(abobec1或apobec3a)和ncas9第二个片段之间。所述xten肽段的氨基酸序列可以包括seqidno.7所示的氨基酸序列。再例如，柔性连接肽片段可以是gs肽段等。所述柔性连接肽片段(例如，gs肽段)可以位于ncas9第二个片段与两个ugi片段中第一个ugi之间，也可以位于两个ugi之间。所述柔性连接肽片段的氨基酸序列可以包括seqidno.8所示的氨基酸序列。本发明所提供的融合蛋白中，所述融合蛋白自n端至c端可以依次包括bpnls肽段、第一ncas9片段、xten肽段、apobec1、xten肽段、第二ncas9片段、gs肽段和两个ugi片段。在本发明一具体实施例中，所述融合蛋白自n端至c端可以依次包括bpnls肽段、第一ncas9片段、xten肽段、apobec1、xten肽段、第二ncas9片段、gs肽段和两个ugi片段，所述融合蛋白的氨基酸序列如seqidno.9所示。本发明所提供的融合蛋白中，所述融合蛋白自n端至c端可以依次包括bpnls肽段、第一ncas9片段、xten肽段、apobec3a、xten肽段、第二ncas9片段、gs肽段和两个ugi片段。在本发明一具体实施例中，所述融合蛋白自n端至c端可以依次包括bpnls肽段、第一ncas9片段、xten肽段、apobec3a、xten肽段、第二ncas9片段、gs肽段和两个ugi片段，所述融合蛋白的氨基酸序列如seqidno.10所示。本发明第二方面提供一种分离的多核苷酸，编码本发明第一方面所提供的融合蛋白。本发明第三方面提供一种构建体，所述构建体含有本发明第二方面所提供的分离的多核苷酸。所述构建体通常可以通过将所述分离的多核苷酸插入合适的表达载体中构建获得，本领域技术人员可选择合适的表达载体，例如，所述表达载体可以是包括但不限于pcmv表达载体、psv2表达载体等。本发明第四方面提供一种表达系统，所述表达系统含有本发明第三方面所提供的构建体或基因组中整合有外源的本发明第二方面所提供的分离的多核苷酸。所述表达系统可以是宿主细胞，所述宿主细胞可以表达如上所述的融合蛋白，所述融合蛋白可以与sgrna相配合，从而可以将所述融合蛋白定位到目标区域，实现目标区域的碱基编辑。在本发明另一具体实施例中，所述宿主细胞可以是真核细胞和/或原核细胞，更具体可以是小鼠细胞、人细胞等，更具体可以是小鼠脑神经瘤细胞、人胚胎肾细胞、人宫颈癌细胞、人结肠癌细胞、人骨肉瘤细胞等，更具体可以是n2a细胞、hek293ft细胞、hela细胞、hct116细胞、或u2os细胞等。本发明第五方面提供本发明第一方面所提供的融合蛋白、或本发明第二方面所提供的分离的多核苷酸、或本发明第三方面所提供的构建体、或本发明第四方面所提供的表达系统在基因编辑中的用途，优选为真核生物的基因编辑中的用途，所述真核生物具体可以是后生动物，具体可以是包括但不限于人、小鼠等。所述用途具体可以是包括但不限于由c到t的碱基编辑等，这些碱基编辑可以应用于编辑剪接受体/供体位点来调节rna剪接，也可以用于进行模型(例如，疾病模型、细胞模型、动物模型等)的构建或人类疾病的治疗等。在本发明一具体实施例中，被编辑的对象可以是胚胎、细胞等。本发明第六方面提供一种碱基编辑体系，包括本发明第一方面所提供的融合蛋白，所述碱基编辑体系还包括sgrna。本领域技术人员可以根据基因的目标编辑区域，选择合适的靶向特异性位点的sgrna。例如，所述sgrna的序列通常可以与目标区域至少部分互补，从而可以与所述融合蛋白相配合，将所述融合蛋白定位到目标区域，实现靶点区域内的碱基编辑，例如，可以是胞嘧啶脱氨基反应，即将胞嘧啶(cytosin，c)脱氨产生胸腺嘧啶(thymine，t)。本发明第七方面提供一种碱基编辑方法，包括：通过本发明第一方面所提供的融合蛋白、或本发明第六方面所提供的碱基编辑体系进行基因编辑。例如，所述基因编辑方法可以包括：在适当条件下培养本发明第四方面所提供的表达系统，从而表达所述融合蛋白，所述融合蛋白可以在与其配合的靶向目标区域的sgrna存在的条件下，对靶标区域进行碱基编辑。提供所述sgrna存在的条件的方法对于本领域技术人员来说应该是已知的，例如，可以是在适当条件下培养能够表达所述sgrna的表达系统，所述表达系统可以是包括含有编码所述sgrna的多核苷酸的表达载体的宿主细胞、或染色体中整合有编码所述sgrna的多核苷酸的宿主细胞。在本发明一具体实施例中，所述sgrna与所述融合蛋白可以在同一宿主细胞中表达，所述宿主细胞可以为靶细胞。在本发明另一具体实施例中，所述基因编辑为体外基因编辑。本发明提供了一种新的碱基编辑工具，其通过在ncas9上可嵌合位点能够兼容多种脱氨酶的嵌入，相较于ncas9末端融合碱基编辑器，在维持特定的靶向碱基编辑效率的同时，在dna和rna上的脱靶情况都极大降低，具有更高的特异性，具有良好的产业化前景。以下通过特定的具体实例说明本发明的实施方式，本领域技术人员可由本说明书所揭露的内容轻易地了解本发明的其他优点与功效。本发明还可以通过另外不同的具体实施方式加以实施或应用，本说明书中的各项细节也可以基于不同观点与应用，在没有背离本发明的精神下进行各种修饰或改变。在进一步描述本发明具体实施方式之前，应理解，本发明的保护范围不局限于下述特定的具体实施方案；还应当理解，本发明实施例中使用的术语是为了描述特定的具体实施方案，而不是为了限制本发明的保护范围；在本发明说明书和权利要求书中，除非文中另外明确指出，单数形式“一个”、“一”和“这个”包括复数形式。当实施例给出数值范围时，应理解，除非本发明另有说明，每个数值范围的两个端点以及两个端点之间任何一个数值均可选用。除非另外定义，本发明中使用的所有技术和科学术语与本
技术领域：
技术人员通常理解的意义相同。除实施例中使用的具体方法、设备、材料外，根据本
技术领域：
的技术人员对现有技术的掌握及本发明的记载，还可以使用与本发明实施例中所述的方法、设备、材料相似或等同的现有技术的任何方法、设备和材料来实现本发明。除非另外说明，本发明中所公开的实验方法、检测方法、制备方法均采用本
技术领域：
常规的分子生物学、生物化学、染色质结构和分析、分析化学、细胞培养、重组dna技术及相关领域的常规技术。这些技术在现有文献中已有完善说明，具体可参见sambrook等molecularcloning：alaboratorymanual，secondedition，coldspringharborlaboratorypress，1989andthirdedition，2001；ausubel等，currentprotocolsinmolecularbiology，johnwiley&sons，newyork，1987andperiodicupdates；theseriesmethodsinenzymology，academicpress，sandiego；wolffe，chromatinstructureandfunction，thirdedition，academicpress，sandiego，1998；methodsinenzymology，vol.304，chromatin(p.m.wassarmananda.p.wolffe，eds.)，academicpress，sandiego，1999；和methodsinmolecularbiology，vol.119，chromatinprotocols(p.b.becker，ed.)humanapress，totowa，1999等。实施例11、基于mua转座酶的tada-tada*转座子的构建由上海铂尚生物科技有限公司合成tada-tada*转座子序列(seqidno.11)，使用诺唯赞公司的高保真酶试剂盒(vazyme,p501-d2)进行pcr扩增。正向引物：ggtctctgatccggcgcacgaa(seqidno.71)，反相引物：ggtctctgatccggcgcacgaa(seqidno.72)；扩增体系如下：表3pcr条件如下：表4pcr扩增产物经通过axypreppcrclean-up试剂盒(axygen,ap-pcr-500g)纯化回收，待用。2、sgrna的构建检测abe(adeninebaseediting)在真核细胞中靶向编辑效率所用sgrna为abe-site1。后续检测abe和ce-abe(centrallyencapsulateabe)在hek293t细胞的8个内源基因位点sgrna为site2-site9。后续检测abe和ce-abe在n2a细胞的12个内源基因位点sgrna为site10-site21。位点序列如seqidno.12～32所示。检测ce-cbe和ce-a3a所用sgrna均为靶向hek293t细胞的内源基因位点，sgrna为site22-site32。位点序列如seqidno.57～67所示。根据靶向位点序列设计20个碱基互补配对的上下游引物，加灭菌水溶解至100μm。经退火后连接到pgl3-u6-sgrna(addgene#51133)载体上，以构建靶向特异性sgrna。退火体系如下所示：表5正向引物4.5μl反向引物4.5μl10×nebbuffer21μl退火程序如下：表695℃5min95-85℃-2℃/s85-25℃-0.1℃/s4℃∞利用bsai(neb,r0535s)对pgl3-u6-sgrna(addgene#51133)质粒进行酶切以得到线性化sgrna载体。酶切体系如下所示：表7水补水至50μlpgl3-u6质粒10μg10×cutsmartbuffer5μlbsai酶5μl以上反应体系配置好后，置于37℃条件下反应5h，酶切产物用axyprepdna凝胶回收试剂盒(axygen,ap-gx-250g)做割胶回收得到线性化载体。取50ng线性化载体与3μl退火产物通过t4连接酶(neb,m0202s)连接，16℃孵育2小时后并转化涂板，经sanger测序得到正确的靶向特异性sgrna。连接体系如下：表8水补水至10μlpgl3-u6-bsai酶切线性片段20ng退货产物1μlsolutioni5μl连接产物随后进行转化，复苏30min，涂板于氨卞抗性的lb琼脂平板，37℃培养过夜。挑选单克隆进行测序验证，abesite1-site21的sgrna。3、用于mua转座酶随机插入的受体质粒构建质粒构建所用引物均由上海铂尚生物科技有限公司合成。首先以pcmv-abemax(addgene，#112095)质粒为模板，正向引物：gacaagaagtacagcatcggcc(seqidno.73)，反向引物：gctgtacttcttgtcactgctgactttccgcttcttc(seqidno.74)，得到7629bp长度的片段，pcr扩增产物经通过axypreppcrclean-up试剂盒(axygen,ap-pcr-500g)纯化回收，再使用gibsonassemblymastermix重组试剂盒(neb，e2611s)对该片段进行重组，反应体系如下：表9gibsonassemblymastermix(2x)5μl7629bppcr片段200ng无菌水补水至10μl反应液混合后置于50℃孵育1h，随后进行转化，复苏30min，涂板于氨卞抗性的lb琼脂平板，37℃培养过夜。挑选单克隆进行测序验证，得到pcmv-ncas9质粒(seqidno.33)。构建成功的质粒(seqidno.33)采用axyprepplasmidsminiprep试剂盒(axygen,ap-mn-p-250g)进行质粒小提。以seqidno.33为模板，正向引物：gaagaagcggaaagtcgacaagaagtacagcatcgg(seqidno.75)，反相引物：ctgagctagctgtcaacgagccccagctggttcttt(seqidno.76)，pcr扩增得到4507bp长度的ncas9片段；以pet30质粒为模板，正向引物：ctcactgattaagcattggtaagcgcggaacccctatttgtt(seqidno.77)，反相引物：ccgtttcatggtggcatgtatatctccttcttaaagttaaacaaaatt(seqidno.78)；pcr扩增4620bp长度的kanr片段；以pgl3-u6-sgrna质粒为模板，正向引物：gtataatactagtgctcttgcccggcgtcaatacgttttagagctagaaatagcaagtt(seqidno.79)，反相引物：gttagcagccggatcaaaaaaagcaccgactcgg(seqidno.80)，进行pcr扩增132bp长度的u6-sgrna片段，再以u6-sgrna产物为模板，正向引物：ttgacagctagctcagtcctaggtataatactagtgctcttgcc(seqidno.81)，反相引物：gttagcagccggatcaaaaaaagcaccgactcgg(seqidno.82)，pcr扩增154bp长度的j23119promoter-grna片段；以pcmv-abemax(addgene，#112095)质粒为模板，正向引物：cttttcggggaaatgtgggaaatgtgcgcggaacc(seqidno.83)，反相引物：cccggcgtcaatacgggata(seqidno.84)，pcr进行扩增，得到386bp长度的ampr-1片段以pcmv-abemax(addgene，#112095)质粒为模板，正向引物：gtattgacgccgggtaagagcaactcggtcgccgc(seqidno.85)，反向引物：ttaccaatgcttaatcagtgaggcacc(seqidno.86)，pcr进行扩增，得到620bp长度的ampr-1片段。以上pcr所用的都是诺唯赞公司的高保真酶试剂盒(vazyme,p501-d2)进行，pcr反应体系如下：表10水补加至50μl2xbuffer25μldntp1μl正向引物(10μm)2μl反向引物(10μm)2μl高保真酶1μl模板1ngpcr程序如下：表11以上所有pcr扩增产物都经axypreppcrclean-up试剂盒(axygen,ap-pcr-500g)纯化回收，再使用gibsonassemblymastermix重组试剂盒(neb，e2611s)对该片段进行重组，反应体系如下：表12gibsonassemblymastermix(2x)10μlncas9片段(4507bp)80ngkanr片段(4620bp)80ngj23119promoter-grna片段(154bp)10ngampr-1片段(386bp)20ngampr-2片段(620bp)30ng无菌水补水至20μl反应液混合后置于50℃孵育1h，随后进行转化，复苏30min，涂板于卡那抗性的lb琼脂平板，37℃培养过夜。挑选单克隆进行测序验证，得到pet-ncas9-grna-ampr(a118x)-kanr质粒(seqidno.34)。构建成功的质粒(seqidno.34)采用axyprepplasmidsminiprep试剂盒(axygen,ap-mn-p-250g)进行质粒小提。4、体外随机插入文库的构建pcr得到的tada-tada*转座子片段、pet-ncas9-grna-ampr(a118x)-kanr质粒(seqidno.34)和mua转座酶(thermofisher，f-701)在体外反应，形成质粒中随机插入tada-tada*转座子片段的插入质粒库，流程具体如图1所示。具体反应体系如下：表13tada-tada*片段250ng质粒(seqidno.34)500ngmua转座酶1μl5×reactionbufferformuatransposase4μl水补水至20μl反应液在30℃孵育1小时以实现随机插入，然后在75℃孵育10分钟以灭活mua转座酶。随后用异丙醇沉淀纯化出dna，并重悬于5μl去离子水中，然后电转到100μlbl21(de3)electro(shanghaiweidibiotechnology，ee1002)感受态细胞中。然后加入1mlsoc培养基，并将细菌在37℃下培养1小时。将上述转化后在soc培养基中复苏1小后的细菌涂布在数个含10μg/ml卡那霉素的lb琼脂板上，并在37℃下孵育16小时。然后刮下来平板上的菌落，然后采用axyprepplasmidsminiprep试剂盒(axygen,ap-mn-p-250g)进行质粒小提。提取的mua随机插入质粒库在中国北京的novogene生物信息学研究所进行测序，使用illuminahiseqxten(2×150pe)对构建的转座子文库进行测序。首先使用默认参数bwav0.7.16将所有数据读段映射到主链序列。提取断的读段，然后将其映射到插入序列。检查所有映射的读取，并将断点记录为插入位点。最终得到插入文库的随机插入情况，具体在ncas9上的插入位点以氨基酸碳基端计算(如：插入发生在第五位天冬氨酸碳基端，则该插入位点为5)，统计后发现，基于mua的随机插入文库覆盖度非常高，ncas9上99.99％的氨基酸位点都至少有一次插入发生，将插入频次和插入位点从小到大排列如下：表145、大肠杆菌中筛选功能性嵌入融合abe蛋白的表达质粒将上述转化后在soc培养基中复苏1小后的细菌涂布在数个含10μg/ml卡那霉素的lb琼脂板上，并在37℃下孵育16小时。然后刮下来平板上的菌落，并将其重悬于含有500μmiptg的100mllb中。将培养物温育10-12h以诱导ncas9表达并修复ampr(a118x)上的突变。然后将减少量的细胞(5ml，1ml，500μl，100μl)接种到含有氨苄青霉素(10μg/ml)和卡那霉素(10μg/ml)的15cmlb琼脂平板上。37℃孵育过夜后，挑选菌落并进行sanger测序，以评估ampr(a118x)上的碱基编辑并确定tada-tada*插入位点。筛选到以下插入位点，具体位置为51，62，63，249，531，584，719，768，770，776，782，790，808，819，831，832，842，893，924，1009，1010，1018，1033，1050，1051，1063，1072，1073，1090，1227，1246，1248，1253，1260，1263，1276，1290，1302，1346，tada-tada*片段插入在这些位点的碳端。经过ampr抗性筛选，以及测序分析ampr(a118x)位点的修复情况，发现以上位点插入tada-tada*能够形成有碱基编辑功能的嵌合融合版本abe，对应的插入位置和碱基编辑效率如图2所示。6、大肠杆菌中ce-abe突变效率的检测首先，我们在将上述电转随机插入文库的大肠杆菌均匀涂布于含有氨苄抗生素的琼脂糖平板上，培养箱中过夜培养。挑取阳性克隆，分别用引物(cttttcggggaaatgtgggaaatgtgcgcggaacc)(seqidno.87)和引物(cggatgcctagacaggtgttcaa)(seqidno.88)对a118x位点腺嘌呤突变效率以及对应的tada-tada*片段在ncas9上插入位置进行sanger测序分析(图2)。在筛选的文库中回收的43个插入位点中，有9个位点聚集在短片段(16-aa)内，它们位于1048thr，1050ile，1051thr，1052leu，1054asn，1056glu，1057ile，1059lys和1063ile的位置。这些位点在筛选库中的富集是特定的，因为在未筛选的库中，这些位点分别仅被插入了61、39、90、38、5、29、76、53和25次，比某些位置少得多筛选后未恢复的其他位点(例如1090pro，插入280次)。因此，16个氨基酸的片段对外源片段插入具有高度的耐受性，并且可能对于ncas9功能是非必需的。该片段在28个spcas9直系同源物中不保守(图3)。因此，在后续构建真核表达载体时，我们用tada-tada*替换了1048thr-1063ile区，以生成ce-abe1048-1063。7、比较abemax和多种ce-abe在人细胞中靶向编辑效率在原核细胞中筛选得到功能性的ce-abe后，我们进一步检测了ce-abe在hek293t细胞中检测靶向碱基编辑效率，过程如下：首先分别构建ce-abe真核表达载体：利用正向引物(agggagagccgccaccatgaaacggacagccgac)(seqidno.89)和反向引物(tcctcttcttcttgggctcgaattcgctgccgtcggc)(seqidno.90)对上述43个tada-tada*片段成功插入并具有腺嘌呤脱氨作用的编辑器进行pcr扩增，得到20个ce-abe片段。利用正向引物(ggtggcggctctccctatagtgagtc)(seqidno.91)和反向引物(cccaagaagaagaggaaagtctaacc)(seqidno.92)对pcmv-abemax进行扩增，得到片段seqidno.35。用诺唯赞高保真酶试剂盒(vazyme,p501-d2)pcr扩增片段。pcr反应体系如下所示：表15水补加至50μl2xbuffer25μldntp1μl正向引物(10μm)2μl反向引物(10μm)2μl高保真酶1μl细胞裂解液3-5μlpcr程序如下：表16pcr扩增产物经axypreppcrclean-up试剂盒(axygen,ap-pcr-500g)纯化回收，并且进行重组反应，再使用gibsonassemblymastermix重组试剂盒(neb，e2611s)对该片段进行重组，反应体系如下：表17gibsonassemblymastermix(2x)5μlce-abe的pcr片段150ngpcmv-abe的pcr片段(seqidno.35)50ng无菌水补水至10μl反应液混合后置于50℃孵育1h，随后进行转化，复苏30min，涂板于氨卞抗性的lb琼脂平板，37℃培养过夜。挑选单克隆进行测序验证，得到pcmv-ce-abe质粒(seqidno.36-55)。采用axyprepplasmidsminiprep试剂盒(axygen,ap-mn-p-250g)进行质粒小提。进行sanger测序。复苏hek293ft细胞(来自atcc)，并培养在10cm培养皿(corning,430167)中，培养基为含有10％体积比的胎牛血清(hyclone,sv30087)的dmem(hyclone,sh30243.01)。培养温度为37℃，二氧化碳浓度为5％。传代后当细胞密度为80％时，细胞分盘至12孔板。12孔板使用前用1:10稀释的多聚赖氨酸溶液(sigma，p4707-50ml)包被处理。1)接种细胞12-14h后，细胞浓度约为80％时，进行转染。每孔转染的质粒的量是ce-abe(seqidno.36-55)质粒700ng，abemax-site1(seqidno.12)的sgrna300ng将质粒混在100μl的opti-mem(gibco,11058021)培养基中。以pcmv-abemax作为阳性对照组，每孔加700ng质粒(addgene，#112095)和abemax-site1(seqidno.12)的sgrna300ng。2)另外，将3μl的lipofectamine2000转染试剂(thermo,11668019)混入100μl的opti-mem培养基，静置5分钟。3)将混有质粒的opti-mem加入混有lipofectamine2000的opti-mem，慢速吹打混匀，静置20分钟。4)将上述混和静置好的转染液分别加入培养的细胞中。5)转染6小时后用含有10％fbs的dmem换液。6)转染48小时后，去培基，用pbs清洗一次细胞，然后用te(thermofisher，r001100)将细胞消化下来，再用含有10％fbs的dmem终止消化，并离心收集细胞，最后用培养基重悬。7)重悬后的细胞进行facs(fluorescenceactivatedcellsorting)分选，收集gfp荧光强度前5％的细胞，每个样品至少收集5000个细胞。取上述收集的细胞的1/6进行直接裂解，并pcr扩增靶向位点片段，正向引物：aaagatcttcacaggctaccccc(seqidno.103)；反向引物：aatccacagcaacaccctctcc(seqidno.104)。用诺唯赞高保真酶试剂盒(vazyme,p501-d2)pcr扩增各基因组靶向位点片段。pcr反应体系如下所示：表18pcr程序如下：表19pcr扩增产物经axypreppcrclean-up试剂盒(axygen,ap-pcr-500g)纯化回收，进行sanger测序。测序情况与ce-abe对应的插入位点对应结果如图4所示。8、比较abemax和ce-abe在人细胞中的脱靶情况收集上述5％gfp阳性细胞30000个，离心后去上清，然后加入trizol(thermofisher，15596018)试剂，按照说明书进行总rna的抽提。然后取部分进行反转录，详细步骤如下：1)抽取总rna：加入1mltrizol试剂，吹打数次，使细胞均质化，吸取trizol于无核酸酶的离心管中。然后加入200μl三氯甲烷，混合均匀，4℃预冷离心机中12000rpm离心15分钟；小心吸取400μl上清于新的无核酸酶离心管中，加入400μl异丙醇，室温混匀后静置10分钟；4℃预冷离心机中12000rpm离心15分钟，弃上清；加入75％乙醇1ml，混匀后于4℃预冷离心机中12000rpm离心15分钟，弃上清，自然晾干，加入无核酸酶的水20-30μl，使用nanodrop进行rna浓度测定。2)总rna反转录成cdna，我们使用iiqrtsupermixforqpcr(+gdnawiper)试剂盒进行，首先去除总rna中的基因组dna500ng总rna2μl4×gdnawipermix(vazyme，r223-01)补水至8μl，42℃孵育5分钟。然后开始反转录反应，向上述8μl反应液中加入iiqrtsupermixiia(vazyme，r223-01)，50℃孵育20分钟，再置于85℃反应2分钟灭活反转录酶活性。即可得到cdna，进行后续检测。在之前的abemax转染细胞的rna-seq数据中分析得到三个脱靶编辑高的rna脱靶位点(chr19(14518195)，chr11(62594034)，chr16(25164711))，针对这三个位点进行引物设计，对ce-abe的cdna样本进行这三个位点的扩增，随后进行sanger测序分析，结果如图5所示。通过分析可以发现，相较于abemax，所有ce-abe在检测的3个rna脱靶位点上都有显著性的降低。这说明嵌合脱氨酶在ncas9的内部，能够有效的降低tada-tada*在部分rna位点上的脱靶编辑(图6)。随后，我们针对ce-abe1048-1063和ce-abe1072(编号后的数字为tada-tada*片段在ncas9内部的插入位点)以及abemax转染的细胞rna进行全转录组测序，所有rna样品均使用位于中国北京的novogene生物信息学研究所的illuminahiseqx10(2×150pe)进行测序，每个样品的读取深度约为2000万。通过star软件(版本2.5.1)将读段映射到人参考基因组(hg38)，使用来自gencodev30版的注释。删除重复后，通过gatkhaplotypecaller(版本4.1.2)识别变体，并用qd(质量按深度)过滤，所有变体均通过bam-readcount进行验证并量化，参数为-q20-b30。给定的编辑至少应为10倍，并且要求这些编辑至少要有99％的读数支持野生型样品中的参考等位基因。最后，仅考虑转录链中的a到g(对于abe)编辑用于下游分析。具体结果如图6，嵌合在1048thr-1063ile和1072val位置的ce-abe能够在全转录组水平极大的降低tada-tada*在rna上的脱靶编辑。同时还对abemax、ce-abe1048-1063和ce-abe1072三个编辑器靶向编辑效率进行监测。结果显示，虽然ce-abe-1072的靶向效率显著低于abemax，而ce-abe1048-1063的靶向编辑效率与abemax没有显著差异，具体结果如图7所示。9、ce-abe1048-1063在多个内源基因位点上的碱基编辑结果我们进一步在hek293t和n2a细胞中表征了ce-abe1048-1063的靶向碱基编辑效率以及编辑窗口，过程如下：复苏hek293ft和n2a细胞(来自atcc)，并分别培养在10cm培养皿(corning,430167)中，培养基为含有10％体积比的胎牛血清(hyclone,sv30087)的dmem(hyclone,sh30243.01)。培养温度为37℃，二氧化碳浓度为5％。传代后当细胞密度为80％时，细胞分盘至12孔板。12孔板使用前用1:10稀释的多聚赖氨酸溶液(sigma，p4707-50ml)包被处理。2)接种细胞12-14h后，细胞浓度约为80％时，进行转染。每孔转染的质粒的量是ce-abe1048-1063(seqidno.45)质粒700ng，对于hek293ft细胞，各个位点(seqidno.13-20)的grna质粒300ng；对于n2a细胞，各个位点(seqidno.21-32)的grna质粒300ng。将质粒混在100μl的opti-mem(gibco,11058021)培养基中。以pcmv-ancbe4max作为对照，每个孔加700ngpcmv-abe4max质粒和300ng各位点的sgrna。3)另外，将3μl的lipofectamine2000转染试剂(thermo,11668019)混入100μl的opti-mem培养基，静置5分钟。4)将混有质粒的opti-mem加入混有lipofectamine2000的opti-mem，慢速吹打混匀，静置20分钟。5)将上述混和静置好的转染液分别加入培养的细胞中。6)转染6小时后用含有10％fbs的dmem换液。转染48小时后，去培基，用pbs清洗一次细胞，然后用te(thermofisher，r001100)将细胞消化下来，再用含有10％fbs的dmem终止消化，并离心收集细胞，最后用培养基重悬。7)重悬后的细胞进行facs(fluorescenceactivatedcellsorting)分选，由于gfp信号在grna质粒上，因此，我们直接分选所有的gfp阳性细胞，每个样品至少收集5000个细胞。上述收集的细胞直接进行裂解，并pcr扩增靶向位点片段。用诺唯赞高保真酶试剂盒(vazyme,p501-d2)pcr扩增各基因组靶向位点片段。pcr反应体系如下所示：表20水补加至50μl2xbuffer25μldntp1μl正向引物(10μm)2μl反向引物(10μm)2μl高保真酶1μl细胞裂解液3-5μlpcr程序如下：表21pcr扩增产物经axypreppcrclean-up试剂盒(axygen,ap-pcr-500g)纯化回收。将具有不同条形码的pcr产物集中在一起，在中国北京的novogene生物信息研究所的illuminahiseqxten(2×150pe)平台上进行深度测序。使用adapterremoval版本2.2.2删除了配对末端读取的适配器对，并将11bp或更多碱基的配对末端读取比对合并为单个共有读取。然后使用bwa-mem算法(bwav0.7.16)将所有处理的读段映射到靶序列。对于每个位点，使用参数-q20-b30的bam读数计数来计算突变率。基于原型间隔子中至少包含1个插入或缺失的核苷酸的读数计算插入缺失。插入缺失频率计算为含插入缺失的读段数/总映射读段数。测序情况结果见附图8和9。结果表明ce-abe1048-1063在hek293t细胞的多个内源位点的靶向碱基编辑效率与abe4max相当。另外，ce-abe1048-1063编辑窗口没有发生显著改变，具体结果见附图8和9。9、ce-cbe1048-1063在多个内源基因位点上的碱基编辑结果在上述实验中已经发现将ncas9上1048thr-106ile之间的片段替换成tada-tada*的ce-abe靶向效率最高，并且脱靶效率较低。进一步将apobec1(seqidno.68)和apobec3a(seqidno.69)分别替换ncas9的1048thr-1063ile肽段，并在hek293t细胞中表征了ce-cbe1048-1063的靶向碱基编辑效率以及编辑窗口，过程如下：1)首先分别构建ce-cbe1048-1063真和ce-a3a1048-1063核表达载体：利用正向引物：catgaactttttcaagtccggatccgagaccccaggc(seqidno.93)和反向引物：tttcgccgtttgtctcgctctctggtgttgctgac(seqidno.94)对apobec1片段进行pcr扩增。利用正向引物：catgaactttttcaagtccggatccgagaccccaggc(seqidno.95)和反向引物：tttcgccgtttgtctcgctctctggtgttgctgac(seqidno.96)对apobec3a片段进行pcr扩增。利用正向引物：gagacaaacggcgaaaccggggagatc(seqidno.97)和反向引物：cttgaaaaagttcatgatgttgc(seqidno.98)以pcmv-ancbe4max为模板pcr扩增。用诺唯赞高保真酶试剂盒(vazyme,p501-d2)pcr扩增片段。pcr反应体系如下所示：表22水补加至50μl2xbuffer25μldntp1μl正向引物(10μm)2μl反向引物(10μm)2μl高保真酶1μl模板dna1μlpcr程序如下：表23pcr扩增产物经axypreppcrclean-up试剂盒(axygen,ap-pcr-500g)纯化回收，并且进行重组反应，再使用gibsonassemblymastermix重组试剂盒(neb，e2611s)对该片段进行重组，反应体系如下：表24gibsonassemblymastermix(2x)5μlapobec1和apobec3a的pcr片段150ngpcmv-ancbe4max的pcr片段50ng无菌水补水至10μl反应液混合后置于50℃孵育1h，随后进行转化，复苏30min，涂板于氨卞抗性的lb琼脂平板，37℃培养过夜。挑选单克隆进行测序验证，得到pcmv-ce-cbe1048-1063质粒(seqidno.56)和pcmv-ce-a3a1048-1063质粒(seqidno.70)。采用axyprepplasmidsminiprep试剂盒(axygen,ap-mn-p-250g)进行质粒小提。进行sanger测序。复苏hek293ft细胞(来自atcc)，并分别培养在10cm培养皿(corning,430167)中，培养基为含有10％体积比的胎牛血清(hyclone,sv30087)的dmem(hyclone,sh30243.01)。培养温度为37℃，二氧化碳浓度为5％。传代后当细胞密度为80％时，细胞分盘至12孔板。12孔板使用前用1:10稀释的多聚赖氨酸溶液(sigma，p4707-50ml)包被处理。2)接种细胞12-14h后，细胞浓度约为80％时，进行转染。每孔转染的质粒的量是ce-cbe(seqidno.56)质粒或ce-a3a(seqidno.70)700ng，各个位点的grna质粒300ng(seqidno.57-67)。将质粒混在100μl的opti-mem(gibco,11058021)培养基中。以pcmv-ancbe4max作为对照，每个孔加700ngpcmv-ancbe4max质粒和300ng各位点的sgrna。3)另外，将3μl的lipofectamine2000转染试剂(thermo,11668019)混入100μl的opti-mem培养基，静置5分钟。4)将混有质粒的opti-mem加入混有lipofectamine2000的opti-mem，慢速吹打混匀，静置20分钟。5)将上述混和静置好的转染液分别加入培养的细胞中。6)转染6小时后用含有10％fbs的dmem换液。转染48小时后，去培基，用pbs清洗一次细胞，然后用te(thermofisher，r001100)将细胞消化下来，再用含有10％fbs的dmem终止消化，并离心收集细胞，最后用培养基重悬。7)重悬后的细胞进行facs(fluorescenceactivatedcellsorting)分选，由于gfp信号在grna质粒上，因此，我们直接分选所有的gfp阳性细胞，每个样品至少收集5000个细胞。上述收集的细胞直接进行裂解，并pcr扩增靶向位点片段。用诺唯赞高保真酶试剂盒(vazyme,p501-d2)pcr扩增各基因组靶向位点片段。pcr反应体系如下所示：表25水补加至50μl2xbuffer25μldntp1μl正向引物(10μm)2μl反向引物(10μm)2μl高保真酶1μl细胞裂解液3-5μlpcr程序如下：表26pcr扩增产物经axypreppcrclean-up试剂盒(axygen,ap-pcr-500g)纯化回收。将具有不同条形码的pcr产物集中在一起，在中国北京的novogene生物信息研究所的illuminahiseqxten(2×150pe)平台上进行深度测序。使用adapterremoval版本2.2.2删除了配对末端读取的适配器对，并将11bp或更多碱基的配对末端读取比对合并为单个共有读取。然后使用bwa-mem算法(bwav0.7.16)将所有处理的读段映射到靶序列。对于每个位点，使用参数-q20-b30的bam读数计数来计算突变率。基于原型间隔子中至少包含1个插入或缺失的核苷酸的读数计算插入缺失。插入缺失频率计算为含插入缺失的读段数/总映射读段数。测序情况结果见附图10和11。结果表明ce-cbe在hek293t细胞的多个内源位点的靶向碱基编辑效率与be4max相当。另外，ce-cbe编辑窗口没有发生显著改变，具体结果见附图8和附图10和11。11、ce-cbe和ce-a3a在人细胞内rna脱靶编辑结果利用facs分选收集上述5％gfp阳性细胞30000个，离心后去上清，然后加入trizol(thermofisher，15596018)试剂，按照说明书进行总rna的抽提。然后取部分进行反转录，详细步骤如下：抽取总rna：加入1mltrizol试剂，吹打数次，使细胞均质化，吸取trizol于无核酸酶的离心管中。然后加入200μl三氯甲烷，混合均匀，4℃预冷离心机中12000rpm离心15分钟；小心吸取400μl上清于新的无核酸酶离心管中，加入400μl异丙醇，室温混匀后静置10分钟；4℃预冷离心机中12000rpm离心15分钟，弃上清；加入75％乙醇1ml，混匀后于4℃预冷离心机中12000rpm离心15分钟，弃上清，自然晾干，加入无核酸酶的水20-30μl，使用nanodrop进行rna浓度测定。随后，针对be4max、ce-cbe1048-1063、ce-cbe1072、be-a3a、ce-a3a1048-1063、ce-a3a1072进行全转录组测序，所有rna样品均使用位于中国北京的novogene生物信息学研究所的illuminahiseqx10(2×150pe)进行测序，每个样品的读取深度约为2000万。通过star软件(版本2.5.1)将读段映射到人参考基因组(hg38)，使用来自gencodev30版的注释。删除重复后，通过gatkhaplotypecaller(版本4.1.2)识别变体，并用qd(质量按深度)过滤，所有变体均通过bam-readcount进行验证并量化，参数为-q20-b30。给定的编辑至少应为10倍，并且要求这些编辑至少要有99％的读数支持野生型样品中的参考等位基因。最后，仅考虑转录链中的c到t编辑用于下游分析。附图12和13表明嵌合在1048thr-1063ile和1072val位置的ce-cbe1048-1063、ce-cbe1072、ce-cbe1048-1063和ce-cbe1072都能够在全转录组水平极大的降低apobec1和apobec3a在rna上的脱靶编辑。12、ce-cbe1048-1063和ce-a3a1048-1063在小鼠胚胎中产生的脱靶dna编辑结果对ce-cbe1048-1063和ce-a3a1048-1063进行体外转录成mrna，首先采用正向引物：atgcctgctattgtcttcccaa(seqidno.99)，和反向引物：aacgggactttccaaaatgtc(seqidno.100)，分别对ce-cbe1048-1063和ce-a3a1048-1063进行pcr扩增，得到线性化片段ce-cbe1048-1063和ce-a3a1048-1063。对于sgrna的转录，首先合成寡聚核苷酸链，退火后，连接到线性化的puc57-spsgrna质粒中，构建的puc57质粒通过sanger测序确认后使用正向引物：tctcgcgcgtttcggtgatgacgg(seqidno.101)和反向引物：aaaaaaatctcgccaacaagttgac(seqidno.102)对sgrna进行pcr扩增：具体步骤如下：表27pcr程序如下：表28无核酸酶环境下进行以下操作：首先，按照1:25的比例向pcr产物中加rnasecuretmrnaseinactivationreagent(invitrogentm，am7005)，60℃干浴处理10分钟；然后使用minelutepcrpurificationkit(qiagen，28004)试剂盒回收pcr片段。(1)cas9的体外转录按照试剂盒mmessagemmachinetmt7ultratranscriptionkit(invitrogentm，am1345)说明书进行cas9的体外转录，如下加反应液：10μlt72×ntp/arca2μl10×t7reactionbuffer600ng模板cas9pcr片段2μlt7enzymemix加nuclease-freewater至20μl反应液混合均匀后在pcr热循环仪上进行反应，热盖温度设为50℃，体系温度设为37℃；反应进行2小时后加1μlturbodnase消化模板dna，37℃反应15分钟。然后进行后续加poly-a反应，体系如下：20μl上述转录产物20μl5×e-papbuffer10μl25mmmncl210μlatpsolution36μlnuclease-freewater在加入e-pap酶之前，吸取2.5μl混合反应液用做后续凝胶电泳，然后向96μl反应液中加入4μle-pap酶，37℃反应30分钟，吸取2.5μl加尾后的反应液，与加尾前的反应液一起在0.8％琼脂糖凝胶中进行电泳，电压为180v，时间为10分钟。确认条带正常后，使用rneasyminikit(qiagen，74104)试剂盒回收cas9mrna。(2)sgrna的体外转录上述得到的纯化产物进行后续步骤。按照试剂盒megashortscripttmt7transcriptionkit(invitrogentm，am1354)说明书进行sgrna的体外转录，600ng模板dna用于反应，如下混合反应液：1μlt710×reactionbuffer1μlt7atpsolution(75mm)1μlt7ctpsolution(75mm)1μlt7gtpsolution(75mm)1μlt7utpsolution(75mm)1μlt7enzymemix2μlt7enzymemix600ng模板sgrnapcr片段加nuclease-freewater至20μl反应液混合均匀后在pcr热循环仪上进行反应，热盖温度设为50℃，体系温度设为37℃。反应进行6小时后，加1μlturbodnase对模板dna进行消化，37℃反应15分钟，然后吸取1μl混合反应液在0.8％琼脂糖凝胶中进行电泳，电压为180v，时间为10分钟。确定目的条带正常后，用megaclearkit(invitrogentm，am1908)试剂盒回收sgrna的mrna。(3)受精卵注射和胚胎移植取6-8周龄c57雌鼠，腹腔注射人绒毛膜促性腺激素hcg(宁波三生药物，b141002)，48小时后腹腔注射孕马血清促性腺激素pmsg(宁波三生药物，s141004)，然后与7-8周龄c57雄鼠进行合笼。12小时后麻醉处死小鼠，取卵细胞，待受精卵发育到2细胞时期，将细胞分开，其中一个细胞转移到另一个透明带中，不注射，直接与其余20-25枚icr小鼠受精卵一起转移到假孕icr雌性小鼠的输卵管中。分别将cbe4max/ce-cbe1048-1063/ce-a3a1048-1063(100ng/μl)与sgrna(50ng/μl)的mrna混合，然后12000rpm离心5分钟，吸取上清mrna并使用femtojet显微注射器在含有5μg/ml细胞松弛素b的hepes-czb培养基液滴中注入剩下的细胞胞质中。然后，将注射的受精卵培养至双细胞阶段，并与其余20-25枚icr小鼠受精卵一起转移到假孕icr雌性小鼠的输卵管中。13.5天后，解剖母鼠，观察小鼠眼睛颜色，选出c57小鼠胚胎，进行裂解，并提取基因组dna，进行后续检测。首先检测sgrna靶向效率，确定编辑效率，具体结果如图14所示。随后，对基因组dna分别进行wgs测序分析编辑器在dna上的脱靶情况，具体结果如图15和图16所示。可见，ce-cbe1048-1063和ce-a3a1048-1063在小鼠胚胎中具有更好的编辑效率和更低的脱靶率。综上所述，本发明有效克服了现有技术中的种种缺点而具高度产业利用价值。上述实施例仅例示性说明本发明的原理及其功效，而非用于限制本发明。任何熟悉此技术的人士皆可在不违背本发明的精神及范畴下，对上述实施例进行修饰或改变。因此，举凡所属
技术领域：
中具有通常知识者在未脱离本发明所揭示的精神与技术思想下所完成的一切等效修饰或改变，仍应由本发明的权利要求所涵盖。序列表<110>上海科技大学<120>一种碱基编辑工具及其用途<160>104<170>siposequencelisting1.0<210>1<211>1046<212>prt<213>人工序列(artificialsequence)<400>1asplyslystyrserileglyleualaileglythrasnservalgly151015trpalavalilethraspglutyrlysvalproserlyslysphelys202530valleuglyasnthrasparghisserilelyslysasnleuilegly354045alaleuleupheaspserglygluthralaglualathrargleulys505560argthralaargargargtyrthrargarglysasnargilecystyr65707580leuglngluilepheserasnglumetalalysvalaspaspserphe859095phehisargleuglugluserpheleuvalglugluasplyslyshis100105110gluarghisproilepheglyasnilevalaspgluvalalatyrhis115120125glulystyrprothriletyrhisleuarglyslysleuvalaspser130135140thrasplysalaaspleuargleuiletyrleualaleualahismet145150155160ilelyspheargglyhispheleuilegluglyaspleuasnproasp165170175asnseraspvalasplysleupheileglnleuvalglnthrtyrasn180185190glnleupheglugluasnproileasnalaserglyvalaspalalys195200205alaileleuseralaargleuserlysserargargleugluasnleu210215220ilealaglnleuproglyglulyslysasnglyleupheglyasnleu225230235240ilealaleuserleuglyleuthrproasnphelysserasnpheasp245250255leualagluaspalalysleuglnleuserlysaspthrtyraspasp260265270aspleuaspasnleuleualaglnileglyaspglntyralaaspleu275280285pheleualaalalysasnleuseraspalaileleuleuseraspile290295300leuargvalasnthrgluilethrlysalaproleuseralasermet305310315320ilelysargtyraspgluhishisglnaspleuthrleuleulysala325330335leuvalargglnglnleuproglulystyrlysgluilephepheasp340345350glnserlysasnglytyralaglytyrileaspglyglyalasergln355360365glugluphetyrlyspheilelysproileleuglulysmetaspgly370375380thrglugluleuleuvallysleuasnarggluaspleuleuarglys385390395400glnargthrpheaspasnglyserileprohisglnilehisleugly405410415gluleuhisalaileleuargargglngluaspphetyrpropheleu420425430lysaspasnargglulysileglulysileleuthrpheargilepro435440445tyrtyrvalglyproleualaargglyasnserargphealatrpmet450455460thrarglysserglugluthrilethrprotrpasnpheglugluval465470475480valasplysglyalaseralaglnserpheilegluargmetthrasn485490495pheasplysasnleuproasnglulysvalleuprolyshisserleu500505510leutyrglutyrphethrvaltyrasngluleuthrlysvallystyr515520525valthrgluglymetarglysproalapheleuserglygluglnlys530535540lysalailevalaspleuleuphelysthrasnarglysvalthrval545550555560lysglnleulysgluasptyrphelyslysileglucyspheaspser565570575valgluileserglyvalgluaspargpheasnalaserleuglythr580585590tyrhisaspleuleulysileilelysasplysasppheleuaspasn595600605glugluasngluaspileleugluaspilevalleuthrleuthrleu610615620phegluaspargglumetileglugluargleulysthrtyralahis625630635640leupheaspasplysvalmetlysglnleulysargargargtyrthr645650655glytrpglyargleuserarglysleuileasnglyileargasplys660665670glnserglylysthrileleuasppheleulysseraspglypheala675680685asnargasnphemetglnleuilehisaspaspserleuthrphelys690695700gluaspileglnlysalaglnvalserglyglnglyaspserleuhis705710715720gluhisilealaasnleualaglyserproalailelyslysglyile725730735leuglnthrvallysvalvalaspgluleuvallysvalmetglyarg740745750hislysprogluasnilevalileglumetalaarggluasnglnthr755760765thrglnlysglyglnlysasnserarggluargmetlysargileglu770775780gluglyilelysgluleuglyserglnileleulysgluhisproval785790795800gluasnthrglnleuglnasnglulysleutyrleutyrtyrleugln805810815asnglyargaspmettyrvalaspglngluleuaspileasnargleu820825830serasptyraspvalasphisilevalproglnserpheleulysasp835840845aspserileaspasnlysvalleuthrargserasplysasnarggly850855860lysseraspasnvalproserglugluvalvallyslysmetlysasn865870875880tyrtrpargglnleuleuasnalalysleuilethrglnarglysphe885890895aspasnleuthrlysalagluargglyglyleusergluleuasplys900905910alaglypheilelysargglnleuvalgluthrargglnilethrlys915920925hisvalalaglnileleuaspserargmetasnthrlystyraspglu930935940asnasplysleuilearggluvallysvalilethrleulysserlys945950955960leuvalseraspphearglysasppheglnphetyrlysvalargglu965970975ileasnasntyrhishisalahisaspalatyrleuasnalavalval980985990glythralaleuilelyslystyrprolysleugluserglupheval99510001005tyrglyasptyrlysvaltyraspvalarglysmetilealalysser101010151020gluglngluileglylysalathralalystyrphephetyrserasn1025103010351040ilemetasnphephelys1045<210>2<211>305<212>prt<213>人工序列(artificialsequence)<400>2gluthrasnglygluthrglygluilevaltrpasplysglyargasp151015phealathrvalarglysvalleusermetproglnvalasnileval202530lyslysthrgluvalglnthrglyglypheserlysgluserileleu354045prolysargasnserasplysleuilealaarglyslysasptrpasp505560prolyslystyrglyglypheaspserprothrvalalatyrserval65707580leuvalvalalalysvalglulysglylysserlyslysleulysser859095vallysgluleuleuglyilethrilemetgluargserserpheglu100105110lysasnproileasppheleuglualalysglytyrlysgluvallys115120125lysaspleuileilelysleuprolystyrserleuphegluleuglu130135140asnglyarglysargmetleualaseralaglygluleuglnlysgly145150155160asngluleualaleuproserlystyrvalasnpheleutyrleuala165170175serhistyrglulysleulysglyserprogluaspasngluglnlys180185190glnleuphevalgluglnhislyshistyrleuaspgluileileglu195200205glnileserglupheserlysargvalileleualaaspalaasnleu210215220asplysvalleuseralatyrasnlyshisargasplysproilearg225230235240gluglnalagluasnileilehisleuphethrleuthrasnleugly245250255alaproalaalaphelystyrpheaspthrthrileasparglysarg260265270tyrthrserthrlysgluvalleuaspalathrleuilehisglnser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3>人工序列(artificialsequence)<400>60acagctgcagagagccctgcagg23<210>61<211>23<212>dna<213>人工序列(artificialsequence)<400>61ttccgcctccgacctgtggctgg23<210>62<211>23<212>dna<213>人工序列(artificialsequence)<400>62ttccttcaggctctgaatcttgg23<210>63<211>23<212>dna<213>人工序列(artificialsequence)<400>63aggccgggagctggaggagctgg23<210>64<211>23<212>dna<213>人工序列(artificialsequence)<400>64agagccccccctcaaagagaggg23<210>65<211>23<212>dna<213>人工序列(artificialsequence)<400>65ggagccacaggagccgctgcagg23<210>66<211>23<212>dna<213>人工序列(artificialsequence)<400>66tactcccaggtcctcttcaaggg23<210>67<211>23<212>dna<213>人工序列(artificialsequence)<400>67ggcccagactgagcacgtgatgg23<210>68<211>678<212>dna<213>人工序列(artificialsequence)<400>68gaaaccggaccagtggcagtggacccaaccctgaggagacggattgagccccatgaattt60gaagtgttctttgacccaagggagctgaggaaggagacatgcctgctgtacgagatcaag120tggggcacaagccacaagatctggcgccacagctccaagaacaccacaaagcacgtggaa180gtgaatttcatcgagaagtttacctccgagcggcacttctgcccctctaccagctgttcc240atcacatggtttctgtcttggagcccttgcggcgagtgttccaaggccatcaccgagttc300ctgtctcagcaccctaacgtgaccctggtcatctacgtggcccggctgtatcaccacatg360gaccagcagaacaggcagggcctgcgcgatctggtgaattctggcgtgaccatccagatc420atgacagccccagagtacgactattgctggcggaacttcgtgaattatccacctggcaag480gaggcacactggccaagatacccacccctgtggatgaagctgtatgcactggagctgcac540gcaggaatcctgggcctgcctccatgtctgaatatcctgcggagaaagcagccccagctg600acatttttcaccattgctctgcagtcttgtcactatcagcggctgcctcctcatattctg660tgggctacaggcctgaag678<210>69<211>609<212>dna<213>人工序列(artificialsequence)<400>69atggaagccagcccagcatccgggcccagacacttgatggatccacacatattcacttcc60aactttaacaatggcattggaaggcataagacctacctgtgctacgaagtggagcgcctg120gacaatggcacctcggtcaagatggaccagcacaggggctttctacacaaccaggctaag180aatcttctctgtggcttttacggccgccatgcggagctgcgcttcttggacctggttcct240tctttgcagttggacccggcccagatctacagggtcacttggttcatctcctggagcccc300tgcttctcctggggctgtgccggggaagtgcgtgcgttccttcaggagaacacacacgtg360agactgcgtatcttcgctgcccgcatctttgattacgaccccctatataaggaggcactg420caaatgctgcgggatgctggggcccaagtctccatcatgacctacgatgaatttaagcac480tgctgggacacctttgtggaccaccagggatgtcccttccagccctgggatggactagat540gagcacagccaagccctgagtgggaggctgcgggccattctccagaatcagggaaacagc600ggcagcgag609<210>70<211>8855<212>dna<213>人工序列(artificialsequence)<400>70atatgccaagtacgccccctattgacgtcaatgacggtaaatggcccgcctggcattatg60cccagtacatgaccttatgggactttcctactggcagtacatctacgtattagtcatcgc120tattaccatggtgatgcggttttggcagtacatcaatgggcgtggatagcggtttgactc180acggggatttccaagtctccaccccattgacgtcaatgggagtttgttttggcaccaaaa240tcaacgggactttccaaaatgtcgtaacaactccgccccattgacgcaaatgggcggtag300gcgtgtacggtgggaggtctatataagcagagctggtttagtgaaccgtcagatccgcta360gagatccgcggccgctaatacgactcactatagggagagccgccaccatgaaacggacag420ccgacggaagcgagttcgagtcaccaaagaagaagcggaaagtcagcagtgacaagaagt480acagcatcggcctggccatcggcaccaactctgtgggctgggccgtgatcaccgacgagt540acaaggtgcccagcaagaaattcaaggtgctgggcaacaccgaccggcacagcatcaaga600agaacctgatcggagccctgctgttcgacagcggcgaaacagccgaggccacccggctga660agagaaccgccagaagaagatacaccagacggaagaaccggatctgctatctgcaagaga720tcttcagcaacgagatggccaaggtggacgacagcttcttccacagactggaagagtcct780tcctggtggaagaggataagaagcacgagcggcaccccatcttcggcaacatcgtggacg840aggtggcctaccacgagaagtaccccaccatctaccacctgagaaagaaactggtggaca900gcaccgacaaggccgacctgcggctgatctatctggccctggcccacatgatcaagttcc960ggggccacttcctgatcgagggcgacctgaaccccgacaacagcgacgtggacaagctgt1020tcatccagctggtgcagacctacaaccagctgttcgaggaaaaccccatcaacgccagcg1080gcgtggacgccaaggccatcctgtctgccagactgagcaagagcagacggctggaaaatc1140tgatcgcccagctgcccggcgagaagaagaatggcctgttcggaaacctgattgccctga1200gcctgggcctgacccccaacttcaagagcaacttcgacctggccgaggatgccaaactgc1260agctgagcaaggacacctacgacgacgacctggacaacctgctggcccagatcggcgacc1320agtacgccgacctgtttctggccgccaagaacctgtccgacgccatcctgctgagcgaca1380tcctgagagtgaacaccgagatcaccaaggcccccctgagcgcctctatgatcaagagat1440acgacgagcaccaccaggacctgaccctgctgaaagctctcgtgcggcagcagctgcctg1500agaagtacaaagagattttcttcgaccagagcaagaacggctacgccggctacattgacg1560gcggagccagccaggaagagttctacaagttcatcaagcccatcctggaaaagatggacg1620gcaccgaggaactgctcgtgaagctgaacagagaggacctgctgcggaagcagcggacct1680tcgacaacggcagcatcccccaccagatccacctgggagagctgcacgccattctgcggc1740ggcaggaagatttttacccattcctgaaggacaaccgggaaaagatcgagaagatcctga1800ccttccgcatcccctactacgtgggccctctggccaggggaaacagcagattcgcctgga1860tgaccagaaagagcgaggaaaccatcaccccctggaacttcgaggaagtggtggacaagg1920gcgcttccgcccagagcttcatcgagcggatgaccaacttcgataagaacctgcccaacg1980agaaggtgctgcccaagcacagcctgctgtacgagtacttcaccgtgtataacgagctga2040ccaaagtgaaatacgtgaccgagggaatgagaaagcccgccttcctgagcggcgagcaga2100aaaaggccatcgtggacctgctgttcaagaccaaccggaaagtgaccgtgaagcagctga2160aagaggactacttcaagaaaatcgagtgcttcgactccgtggaaatctccggcgtggaag2220atcggttcaacgcctccctgggcacataccacgatctgctgaaaattatcaaggacaagg2280acttcctggacaatgaggaaaacgaggacattctggaagatatcgtgctgaccctgacac2340tgtttgaggacagagagatgatcgaggaacggctgaaaacctatgcccacctgttcgacg2400acaaagtgatgaagcagctgaagcggcggagatacaccggctggggcaggctgagccgga2460agctgatcaacggcatccgggacaagcagtccggcaagacaatcctggatttcctgaagt2520ccgacggcttcgccaacagaaacttcatgcagctgatccacgacgacagcctgaccttta2580aagaggacatccagaaagcccaggtgtccggccagggcgatagcctgcacgagcacattg2640ccaatctggccggcagccccgccattaaga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cttggtctgacagttaccaatg7380cttaatcagtgaggcacctatctcagcgatctgtctatttcgttcatccatagttgcctg7440actccccgtcgtgtagataactacgatacgggagggcttaccatctggccccagtgctgc7500aatgataccgcgagacccacgctcaccggctccagatttatcagcaataaaccagccagc7560cggaagggccgagcgcagaagtggtcctgcaactttatccgcctccatccagtctattaa7620ttgttgccgggaagctagagtaagtagttcgccagttaatagtttgcgcaacgttgttgc7680cattgctacaggcatcgtggtgtcacgctcgtcgtttggtatggcttcattcagctccgg7740ttcccaacgatcaaggcgagttacatgatcccccatgttgtgcaaaaaagcggttagctc7800cttcggtcctccgatcgttgtcagaagtaagttggccgcagtgttatcactcatggttat7860ggcagcactgcataattctcttactgtcatgccatccgtaagatgcttttctgtgactgg7920tgagtactcaaccaagtcattctgagaatagtgtatgcggcgaccgagttgctcttgccc7980ggcgtcaatacgggataataccgcgccacatagcagaactttaaaagtgctcatcattgg8040aaaacgttcttcggggcgaaaactctcaaggatcttaccgctgttgagatccagttcgat8100gtaacccactcgtgcacccaactgatcttcagcatcttttactttcaccagcgtttctgg8160gtgagcaaaaacaggaaggcaaaatgccgcaaaaaagggaataagggcgacacggaaatg8220ttgaatactcatactcttcctttttcaatattattgaagcatttatcagggttattgtct8280catgagcggatacatatttgaatgtatttagaaaaataaacaaataggggttccgcgcac8340atttccccgaaaagtgccacctgacgtcgacggatcgggagatcgatctcccgatcccct8400agggtcgactctcagtacaatctgctctgatgccgcatagttaagccagtatctgctccc8460tgcttgtgtgttggaggtcgctgagtagtgcgcgagcaaaatttaagctacaacaaggca8520aggcttgaccgacaattgcatgaagaatctgcttagggttaggcgttttgcgctgcttcg8580cgatgtacgggccagatatacgcgttgacattgattattgactagttattaatagtaatc8640aattacggggtcattagttcatagcccatatatggagttccgcgttacataacttacggt8700aaatggcccgcctggctgaccgcccaacgacccccgcccattgacgtcaataatgacgta8760tgttcccatagtaacgccaatagggactttccattgacgtcaatgggtggagtatttacg8820gtaaactgcccacttggcagtacatcaagtgtatc8855<210>71<211>22<212>dna<213>人工序列(artificialsequence)<400>71ggtctctgatccggcgcacgaa22<210>72<211>22<212>dna<213>人工序列(artificialsequence)<400>72ggtctctgatccggcgcacgaa22<210>73<211>22<212>dna<213>人工序列(artificialsequence)<400>73gacaagaagtacagcatcggcc22<210>74<211>37<212>dna<213>人工序列(artificialsequence)<400>74gctgtacttcttgtcactgctgactttccgcttcttc37<210>75<211>36<212>dna<213>人工序列(artificialsequence)<400>75gaagaagcggaaagtcgacaagaagtacagcatcgg36<210>77<211>42<212>dna<213>人工序列(artificialsequence)<400>77ctcactgattaagcattggtaagcgcggaacccctatttgtt42<210>78<211>48<212>dna<213>人工序列(artificialsequence)<400>78ccgtttcatggtggcatgtatatctccttcttaaagttaaacaaaatt48<210>79<211>59<212>dna<213>人工序列(artificialsequence)<400>79gtataatactagtgctcttgcccggcgtcaatacgttttagagctagaaatagcaagtt59<210>80<211>34<212>dna<213>人工序列(artificialsequence)<400>80gttagcagccggatcaaaaaaagcaccgactcgg34<210>81<211>44<212>dna<213>人工序列(artificialsequence)<400>81ttgacagctagctcagtcctaggtataatactagtgctcttgcc44<210>82<211>34<212>dna<213>人工序列(artificialsequence)<400>82gttagcagccggatcaaaaaaagcaccgactcgg34<210>83<211>35<212>dna<213>人工序列(artificialsequence)<400>83cttttcggggaaatgtgggaaatgtgcgcggaacc35<210>84<211>20<212>dna<213>人工序列(artificialsequence)<400>84cccggcgtcaatacgggata20<210>85<211>35<212>dna<213>人工序列(artificialsequence)<400>85gtattgacgccgggtaagagcaactcggtcgccgc35<210>86<211>27<212>dna<213>人工序列(artificialsequence)<400>86ttaccaatgcttaatcagtgaggcacc27<210>87<211>35<212>dna<213>人工序列(artificialsequence)<400>87cttttcggggaaatgtgggaaatgtgcgcggaacc35<210>88<211>23<212>dna<213>人工序列(artificialsequence)<400>88cggatgcctagacaggtgttcaa23<210>89<211>34<212>dna<213>人工序列(artificialsequence)<400>89agggagagccgccaccatgaaacggacagccgac34<210>90<211>37<212>dna<213>人工序列(artificialsequence)<400>90tcctcttcttcttgggctcgaattcgctgccgtcggc37<210>91<211>26<212>dna<213>人工序列(artificialsequence)<400>91ggtggcggctctccctatagtgagtc26<210>92<211>26<212>dna<213>人工序列(artificialsequence)<400>92cccaagaagaagaggaaagtctaacc26<210>93<211>37<212>dna<213>人工序列(artificialsequence)<400>93catgaactttttcaagtccggatccgagaccccaggc37<210>94<211>35<212>dna<213>人工序列(artificialsequence)<400>94tttcgccgtttgtctcgctctctggtgttgctgac35<210>94<211>35<212>dna<213>人工序列(artificialsequence)<400>94tttcgccgtttgtctcgctctctggtgttgctgac35<210>95<211>37<212>dna<213>人工序列(artificialsequence)<400>95catgaactttttcaagtccggatccgagaccccaggc37<210>96<211>35<212>dna<213>人工序列(artificialsequence)<400>96tttcgccgtttgtctcgctctctggtgttgctgac35<210>97<211>27<212>dna<213>人工序列(artificialsequence)<400>97gagacaaacggcgaaaccggggagatc27<210>98<211>23<212>dna<213>人工序列(artificialsequence)<400>98cttgaaaaagttcatgatgttgc23<210>99<211>22<212>dna<213>人工序列(artificialsequence)<400>99atgcctgctattgtcttcccaa22<210>100<211>21<212>dna<213>人工序列(artificialsequence)<400>100aacgggactttccaaaatgtc21<210>101<211>24<212>dna<213>人工序列(artificialsequence)<400>101tctcgcgcgtttcggtgatgacgg24<210>102<211>25<212>dna<213>人工序列(artificialsequence)<400>102aaaaaaatctcgccaacaagttgac25<210>103<211>23<212>dna<213>人工序列(artificialsequence)<400>103aaagatcttcacaggctaccccc23<210>104<211>22<212>dna<213>人工序列(artificialsequence)<400>104aatccacagcaacaccctctcc22当前第1页12