特异性靶向YBX1基因的sgRNAs及其应用的制作方法

本发明属于基因工程领域，更具体地说，涉及特异性靶向ybx1基因的sgrna及利用该sgrna并基于crispr-cas9技术改造293t细胞提高腺相关病毒产毒效率的方法。
背景技术：
：腺相关病毒(adeno-associatedviralvector，aav)属于微小病毒科(parvovirus)，为无包膜的单链线状dna病毒，具有宿主范围广、安全性高、免疫原性低、表达稳定和物理性质稳定等优点，已被广泛地应用于基础研究和临床试验中，并且腺相关病毒载体已成为世界上最常用的基因治疗载体之一。然而，较低滴度aav的生产仍然是限制该技术进一步推广应用的一项挑战。293t细胞是hek-293(人胚肾细胞)细胞株插入了sv40t-antigen的温度敏感基因形成的高转染效率衍生株。ybx1是一个dna和rna结合蛋白，参与了几乎所有的dna和mrna依赖的过程。ybx1对单链dna(ssdna)比双链dna有更高的亲和力(hasegawaetal.,1991；izumietal.,2001)，尤其是对单链dna基序gggg(tt)有最大的结合偏好(zasedatelevaetal，2002)。aav2单链dnaitr区域内126-146核苷酸(包含位于137-142的gggg(tt)片段)，以及紧跟的125个核苷酸长发夹结构，对aav基因组dna的包装是必须的，也因此被作为aav的包装信号，aav衣壳蛋白的n端与itr区域内126-146核苷酸序列的结合，致使aav单链dna包入aav衣壳内(wangandsrivastava,1997,xiaoetal.,1997)。因此，yb1和aav衣壳竞争性与itr126-146核苷酸序列结合，影响aav基因组的包装效率。最近的报告显示，通过引入shrna序列下调yb1来建立的239t细胞系，可以使aav2和aav8的物理滴度分别增加了45和9倍，aav2感染基因组的滴度增加7倍。同时发现，yb1基因敲除促进了aav2rep的表达和载体dna的产生，减少了aav2产物中空颗粒的数量。因此，敲除ybx1有助于提高293t细胞生产aav的效率。技术实现要素：为了更好的提高293t细胞产aav的效率，本研究采用crispr/cas9系统，通过在ybx1基因第二外显子上下游设计全新的sgrna片段，并通过cre-loxp介导的同源重组，删除ybx1的第二外显子，从而特异性的靶向敲除ybx1基因。本发明的首要目的在于提供用于靶向剪切ybx1基因的sgrnas、loxp供体(donor)质粒、载体、试剂盒、crispr-cas9系统等。本发明的另一目的在利用上述sgrnas、loxp供体(donor)质粒、载体、试剂盒、crispr-cas9系统等靶向剪切ybx1基因，并提高293t细胞生产aav的产毒效率。本发明的目的通过下述技术方案实现：在第一个方面，本发明提供了一对特异性靶向ybx1基因第一内含子的sgrna1和第三内含子的sgrna2，所述sgrna1的序列如seqidno.1所示：tttccaaatccgcccggctt；所述sgrna2的序列如seqidno.2所示：cctgctctgtcggcttctcg。所述sgrna1和sgrna2在ybx1基因上的靶序列是唯一的。应该理解，在保持功能不变的情况下，与上述序列(及本发明中的其他序列)具有至少91％、至少92％、至少93％、至少94％、至少95％、至少96％、至少97％、至少98％、至少99％序列同一性的序列也在本发明的保护范围之内。所述的序列的差异由碱基的取代、缺失或添加所致，本领域技术人员有能力，例如通过非保守碱基的替换，获得功能相似的序列，这样的序列在本发明的保护范围之内。在本发明的一些实施方式中，sgrna1和sgrna2的引物包括：针对ybx1基因的靶点位于第一外显子和第三内含子上的引物对：ybx1sgrna1正向寡核苷酸链：如seqidno.3所示：5’-accgtttccaaatccgcccggctt-3’，ybx1sgrna1反向寡核苷酸链：如seqidno.4所示：5’-aaacaagccgggcggatttggaaa-3’；ybx1sgrna2正向寡核苷酸链：如seqidno.5所示：5’-accgcctgctctgtcggcttctcg-3’，ybx1sgrna2反向寡核苷酸链：如seqidno.6所示：5’-aaaccgagaagccgacagagcagg-3’。在第二个方面，本发明提供了一对载体，分别含有如上所述的sgrna1和sgrna2。在本发明的一些实施方式中，所述载体是慢病毒载体。应该理解，本发明不限于慢病毒载体，在本发明的教导之下，本领域技术人员能够采用其他合适的载体，并且在本发明的保护范围之内。在本发明的一些实施方式中，含有sgrna1的慢病毒载体是靶向剪切ybx1基因第一内含子的crispr-cas9重组表达慢病毒载体plenti-u6-ybx1spgrna1-cmv-puro-p2a-3flag-spcas9，其含有特异性靶向ybx1基因第一内含子的sgrna1和cas9蛋白的重组表达载体，带有puro(嘌呤霉素)筛选标记。应该理解，在不改变功能的情况下，可以对其中一个或多个成分进行替换，例如可以选择除puro之外的其他合适的筛选标记，而不超出本发明的保护范围。作为示例，如下制备含有sgrna1的慢病毒载体：(1)提供sgrna1，所述sgrna1在ybx1基因上的靶序列符合5’-n(19)g的序列排列规则，所述sgrna1在ybx1基因上的靶序列位于基因的第一内含子，所述sgrna1在ybx1基因上的靶序列是唯一的，并且所述sgrna1在ybx1上的靶位点序列如序列表seqidno.1序列所示，在所述sgrna1在ybx1上的靶位点序列的5’-端加上accg序列合成得到正向寡核苷酸即forwardoligo1；获得sgrna1在ybx1上的靶位点序列的互补链，并且在互补链的5’-端加上aaac序列得到反向寡核苷酸即reverseoligo1；将合成的1对互补的sgrna1寡聚核苷酸的forwardoligo1和reverseoligo1成对变性、退火，退火之后形成可以连入包含u6启动子的慢病毒表达载体的双链sgrna1寡聚核苷酸；(2)线性化序列如序列表seqidno.7所示的plenti-u6-spgrnav2.0-cmv-puro-p2a-3flag-spcas9质粒；将退火的双链sgrna1寡聚核苷酸与线性化的载体plenti-u6-spgrnav2.0-cmv-puro-p2a-3flag-spcas9连接获得携带含相应靶序列的sgrna1寡聚核苷酸的表达载体plenti-u6-ybx1spgrna1-cmv-puro-p2a-3flag-spcas9质粒，转化感受态细菌并涂amp+平板，挑取单克隆并用通用引物u6通过测序鉴定出阳性克隆，并对所述阳性克隆摇菌、提取质粒。在本发明的一些实施方式中，含有sgrna2的慢病毒载体是靶向剪切ybx1基因第三内含子的crispr-cas9重组表达慢病毒载体plenti-u6-ybx1spgrna2-cmv-blasticidin(杀稻瘟菌素)-p2a-3flag-spcas9，其含有特异性靶向ybx1基因第三内含子的sgrna2和cas9蛋白的重组表达载体，带有杀稻瘟菌素筛选标记。应该理解，在不改变功能的情况下，可以对其中一个或多个成分进行替换，例如可以选择除杀稻瘟菌素之外的其他合适的筛选标记，而不超出本发明的保护范围。作为示例，如下制备含有sgrna2的慢病毒载体：(1)提供sgrna2，所述sgrna2在ybx1基因上的靶序列符合5’-n(19)g的序列排列规则，所述sgrna2在ybx1基因上的靶序列位于基因的第三内含子，所述sgrna2在ybx1基因上的靶序列是唯一的，并且所述sgrna2在ybx1上的靶位点序列如序列表seqidno.2序列所示，在所述sgrna2在ybx1上的靶位点序列的5’-端加上accg序列合成得到正向寡核苷酸即forwardoligo2；获得sgrna2在ybx1上的靶位点序列的互补链，并且在互补链的5’-端加上aaac序列得到反向寡核苷酸即reverseoligo2；将合成的1对互补的sgrna2寡聚核苷酸的forwardoligo2和reverseoligo2成对变性、退火，退火之后形成可以连入包含u6启动子的慢病毒表达载体的双链sgrna2寡聚核苷酸；(2)线性化序列如序列表seqidno.8所示的plenti-u6-spgrnav2.0-cmv-blasticidin-p2a-3flag-spcas9质粒；将退火的双链sgrna2寡聚核苷酸与线性化的载体plenti-u6-spgrnav2.0-cmv-blasticidin-p2a-3flag-spcas9连接获得携带含相应靶序列的sgrna2寡聚核苷酸的表达载体plenti-u6-ybx1spgrna2-cmv-blasticidin-p2a-3flag-spcas9质粒，转化感受态细菌并涂amp+平板，挑取单克隆并用通用引物u6通过测序鉴定出阳性克隆，并对所述阳性克隆摇菌、提取质粒。在本发明的一些实施方式中，含有sgrna1的crispr-cas9重组表达载体的序列如序列表中seqidno.10所示。在本发明的一些实施方式中，含有sgrna1的crispr-cas9重组表达载体的骨架载体的序列如序列表中seqidno.7所示。在本发明的一些实施方式中，含有sgrna2的crispr-cas9重组表达载体的序列如序列表中seqidno.11所示。在本发明的一些实施方式中，含有sgrna2的crispr-cas9重组表达载体的骨架载体的序列如序列表中seqidno.8所示。在第三个方面，本发明提供了试剂盒，包含上面所述的载体。在本发明的一些实施方式中，试剂盒还包括loxp供体载体质粒，所述loxp供体载体质粒是一种靶向ybx1基因的loxp重组表达腺相关病毒供体载体paav-ybx1供体优选地，loxp前端加sgrna1-cas9剪切位点上游800bp的同源臂，loxp后端加sgrna2-cas9剪切位点下游800bp的同源臂，loxp中间添加从sgrna1-cas9剪切位点到sgrna2-cas9剪切位点之间含有第二外显子的切除片段。在本发明的一些实施方式中，所述loxp供体载体质粒的序列如序列表中seqidno.12所示。在本发明的一些实施方式中，所述loxp供体载体质粒的骨架载体的序列如序列表中seqidno.9所示。在第四个方面，本发明提供了一种特异性靶向ybx1基因第一内含子的sgrna1和第三内含子的sgrna2的crispr-cas9系统，其含有前面所述的一对特异性靶向ybx1基因第一内含子的sgrna1和第三内含子的sgrna2。在第五个方面，本发明提供了前面所述的sgrna1和sgrna2、载体或crispr-cas9系统在特异性靶向删除ybx1基因中的用途。在第六个方面，本发明提供了前面所述的sgrna1和sgrna2、载体或crispr-cas9系统在制备高产腺相关病毒的细胞中的用途。所述的细胞优选为人胚肾细胞株，所述人胚肾细胞优选为人胚肾细胞株293t。然而，应该理解，在本发明的教导之下，本领域技术人员能够采用其他合适的细胞，并且在本发明的保护范围之内。在第七个方面，本发明提供了一种基于crispr-cas9删除ybx1基因的方法，所述方法包括如下步骤：(1)分别构建本发明的ybx1基因的sgrna1、sgrna2到慢病毒载体；(2)构建包含第二外显子切除片段的loxp供体载体；(3)将前面三种载体转染293t细胞；(4)药筛后转入cre质粒，通过同源重组删除ybx1基因。在第八个方面，本发明提供了一种基于crispr-cas9获得高产腺相关病毒的293t细胞株的方法，所述方法包括如下步骤：(1)分别构建本发明的ybx1基因的sgrna1、sgrna2到慢病毒载体；(2)构建包含第二外显子切除片段的loxp供体载体；(3)将三种载体转染293t细胞；(4)进行药物筛选，转入cre质粒，得到高产腺相关病毒的293t细胞株。在第九个方面，本发明提供了一种高产腺相关病毒的293t细胞株，该细胞株由上面所述的方法制备而得，其ybx1基因已被删除。应该理解，本发明并不限于293t细胞株，任何利用本发明方法删除ybx1基因的细胞株都在本发明的保护范围之内。在第十个方面，本发明提供了一种基于crispr-cas9提高293t细胞产aav的方法，所述方法包括如下步骤：(1)分别构建ybx1基因的sgrna1、sgrna2到慢病毒载体；(2)构建包含第二外显子切除片段的loxp供体载体质粒；(3)将三种载体质粒转染293t细胞；(4)药筛后转入cre质粒，得到ybx1基因敲除细胞株。在本发明的实施方式中，所述的靶向剪切ybx1基因的慢病毒载体的构建，包括如下步骤：(1)使用bsmbi酶切慢病毒载体plenti-u6-spgrnav2.0-cmv-puro-p2a-3flagspcas9和plenti-u6-spgrnav2.0-cmv-杀稻瘟菌素-p2a-3flag-spcas9得到酶切后的plenti-u6-spgrnav2.0-cmv-puro-p2a-3flag-spcas9和plenti-u6-spgrnav2.0-cmv-杀稻瘟菌素-p2a-3flag-spcas9慢病毒载体；(2)将上述靶向剪切ybx1基因的sgrna1的dna序列磷酸化后与酶切后的plenti-u6-spgrnav2.0-cmv-puro-p2a-3flag-spcas9慢病毒载体连接，得到靶向剪切ybx1基因的plenti-u6-ybx1spgrna1-cmv-puro-p2a-3flag-spcas9慢病毒载体；将上述靶向剪切ybx1基因的sgrna2的dna序列磷酸化后与酶切后的plenti-u6-spgrnav2.0-cmv-杀稻瘟菌素-p2a-3flag-spcas9慢病毒载体连接，得到靶向剪切ybx1基因的plenti-u6-ybx1spgrna2-cmv-杀稻瘟菌素-p2a-3flag-spcas9慢病毒载体。步骤(2)中所述的dna序列是将正向寡核苷酸链(forwardoligo)和反向寡核苷酸链(reverseoligo)变性、退火之后形成可以连入u6真核表达载体的双链。所述的loxp供体质粒为包含第二外显子切除片段的loxp供体载体质粒paav-ybx1供体，其构建包括如下步骤：loxp前端加sgrna1-cas9剪切位点上游800bp的同源臂，loxp后端加sgrna2-cas9剪切位点下游800bp的同源臂，loxp中间添加从sgrna1-cas9剪切位点到sgrna2-cas9剪切位点之间含有第二外显子的切除片段。在本发明的实施方式中，所述的敲除ybx1基因的细胞株，具体是通过如下步骤构建得到：(1)将所述的靶向剪切ybx1基因的慢病毒载体质粒及loxp供体质粒转染目的细胞；(2)转染48小时后，用2ug/ml嘌呤霉素(puromycin)和5ug/ml杀稻瘟菌素(blasticidin)筛选一周后，活下来的混合细胞扩大培养，再用cre过表达质粒转染混合细胞，所得细胞即为靶向敲除ybx1的混合细胞株，进一步培养得到混合克隆；(3)铺单克隆细胞，并进一步放大培养；收集单克隆细胞，以其基因组dna为模板扩增包含所述靶序列的基因片段，ta克隆测序确认ybx1基因已经被敲除并获得基因敲除的细胞。在本发明的一些实施方式中，目的细胞为293t细胞，是293t细胞中的ybx1基因第二外显子缺失得到的细胞株，经sgrna1和sgrna2的切割后，由donor质粒提供片段同源重组，再经cre剪切，导致ybx1基因的第二号外显子缺失；经过比对，蛋白翻译提前终止。本发明相对于现有技术具有如下的显著优点及效果：本发明提供ybx1基因的sgrnas能有效靶向ybx1基因的第一和第三内含子，将其构建入慢病毒载体，能靶向剪切ybx1基因，通过cre-loxp同源重组，能简单快速得到靶向敲除ybx1基因的细胞株，从而有利于提高293t细胞产aav的产毒效率，所获得的ybx1的293t敲除细胞株具有产毒能力明显高于未敲除细胞的优势。本发明具有位点可控、操作步骤简单、sgrna靶向性好，且对ybx1基因切割效率高，为基础研究和临床应用提供更好的细胞工具。本发明提供的ybx1基因的sgrnas是本申请发明人精心设计的，具有新颖性和创造性。附图说明图1为本发明实施例中使用的载体质粒plenti-u6-spgrnav2.0-cmv-嘌呤霉素(puro)-p2a-3flag-spcas9的质粒图谱。图2为本发明实施例中使用的载体质粒plenti-u6-spgrnav2.0-cmv-杀稻瘟菌素(blasticidin)-p2a-3flag-spcas9的质粒图谱。图3为本发明实施例中使用的载体质粒paav-ybx1供体的质粒图谱。图4为本发明实施例中293t细胞转染含有ybx1-sgrna的活性鉴定测序图。图5为本发明实施例中鉴定到的切割活性较高的一个单克隆细胞株与对照组细胞293t的westernblot检测图。图6为本发明实施例中单克隆细胞株与对照空细胞株(293t)的包装aav的质粒图谱图7为本发明实施例中单克隆细胞株与对照空细胞株(293t)的包装aav上清感染293t的对比图。具体实施方式下面结合附图和实施例对本发明的技术方案进行详细描述，但并不因此将本发明限制在所述的实施例范围之中。下列实施例中未注明具体条件的实验方法，按照常规方法和条件，或按照商品说明书选择。本发明所用试剂和原料均市售可得。本发明首先设计获得特异性靶向ybx1基因第一内含子的sgrna1、第三内含子的sgrna2，并分别构建ybx1基因的sgrna1、sgrna2到慢病毒载体；其次构建包含第二外显子切除片段的loxp供体载体质粒；然后将三种载体质粒转染293t细胞，药筛后转入cre质粒，通过挑取单克隆，得到ybx1基因敲除细胞株。详细过程如下描述：实施例11.靶向剪切ybx1慢病毒质粒构建1.1sgrna寡核苷酸链合成使用crispr在线设计工具(http://crispr.mit.edu/)，根据评分系统，在靠近ybx1的二号外显子的上游和下游内含子中各设计1个20bp的sgrna，并通过blast验证无非特异性基因。编码链模板5′端添加accg，非编码链模板3′端添加aaac，与bsmbi酶切后形成的粘性末端互补，各设计1对crispr寡核苷酸链，见表1。表1.ybx1靶向位点及sgrnas寡核苷酸序列sgrna名称sgrna寡核苷酸序列ybx1sgrna1tttccaaatccgcccggcttybx1sgrna1oligo15’-accgtttccaaatccgcccggctt-3’ybx1sgrna1oligo25’-aaacaagccgggcggatttggaaa-3’ybx1sgrna2cctgctctgtcggcttctcgybx1sgrna2oligo15’-accgcctgctctgtcggcttctcg-3’ybx1sgrna2oligo25’-aaaccgagaagccgacagagcagg-3’1.2载体构建1.2.1使用bsmbi分别酶切2μgplenti-u6-spgrnav2.0-cmv-puro-p2a-3flag-spcas9质粒和plenti-u6-spgrnav2.0-cmv-杀稻瘟菌素-p2a-3flag-spcas9质粒，2h，37℃，酶切体系：1.2.2使用genry胶回收试剂盒纯化酶切质粒产物，按说明书进行操作。1.2.3磷酸化并退火sgrna，酶连体系：pcr仪器退火程序：37℃30min，95℃维持5min，每分钟降低5℃至25℃，4℃维持。1.2.4将退火形成的sgrna1双链和酶切后的plenti-u6-spgrnav2.0-cmv-purop2a-3flag-spcas9载体直接连接；将退火形成的sgrna2双链和酶切后的plenti-u6-spgrnav2.0-cmv-杀稻瘟菌素-p2a-3flag-spcas9载体直接连接。室温下，10min。1.2.5将连接后的质粒转化至感受态细胞dh5α中，均匀涂至lb固体培养基平板中，置于37℃培养箱中培养12-16小时，单个菌落即可出现。1.3挑取单个菌落扩大培养并质粒小提。1.4测序鉴定质粒构建成功，并命名为plenti-u6-ybx1spgrna1-cmvpuro-p2a-3flag-spcas9和plenti-u6-ybx1spgrna2-cmv-杀稻瘟菌素-p2a-3flag-spcas9。2.loxp供体载体构建2.1选取sgrna1-cas9剪切位点上游800bp和sgrna2-cas9剪切位点下游800bp做为同源臂，上游同源臂3’端连接loxp序列，下游同源臂5’端连接loxp序列，两个loxp序列中间连接sgrna1-cas9剪切位点和sgrna2-cas9剪切位点的中间序列(包含完整的第二外显子)，合成寡核苷酸片段。2.2将2.1步合成的寡核苷酸连接到paav-供体载体中，转化至感受态细胞dh5α中，涂板，挑单克隆，扩增并小抽质粒。2.3测序鉴定质粒构建成功，并命名为paav-ybx1-供体。3.筛选稳定细胞株用含10％胎牛血清的高糖dmem培养基于5％co2，37℃恒温培养293t细胞(购于美国atcc细胞库)。取对数期细胞以2×105/孔接种到12孔板培养。待细胞融合度达到60％～70％时更换成opti-mem培养基，1小时后将敲除两种sgrna剪切质粒以及loxp供体质粒各2μg经lipo2000试剂，转染到293t细胞中，转染48小时后，向每孔加入嘌呤霉素(2μg/ml)和杀稻瘟菌素(5μg/ml)，隔天换液，并保持培养基的嘌呤霉素和杀稻瘟菌素浓度恒定，筛选阳性克隆细胞。进一步扩大阳性克隆细胞，转染cre质粒2μg，将所得细胞株命名为293t-ybx1。转染48h后，使用有限稀释法挑选293t-ybx1单克隆敲除细胞株。4.稳定细胞株鉴定及基因打靶方式鉴定以各组敲除ybx1的单克隆细胞株基因组dna为模板，设计针对ybx1第一外显子和第三内含子的引物，序列如表2，进行两轮巢式pcr扩增。用h9271jd-4-1和h9271jd-4-2进行pcr，得到1481bp的片段a，以a为模板，用h9271jd-4-3和h9271jd-4-4进行pcr，得到1373的片段b，该片段电泳后，胶回收用于测序，测序引物为h9271jd-4-3和h9271jd-4-4，表2为引物序列。如测序结果打靶位点附近出现双峰的情况，则可能为打靶成功。如测序结果ybx1基因靶位点附近发生非三倍数的碱基插入或碱基缺失，导致阅读框移码突变，则可判断为ybx1基因敲除。鉴定出sgrna是有活性的(图3)，并且找到1株具有ybx1基因插入突变的单克隆细胞株，经wb检测ybx1敲除，蛋白不表达(图4)。表2.测序引物名称及序列引物名称引物序列h9271jd-4-1gacagtaccactggccagtgaach9271jd-4-2ctgttaaggaatggctcattcach9271jd-4-3attctcgctagttcgatcggtagh9271jd-4-4aattgctttgtactgtgacgaagc实施例21.使用ybx1敲除细胞株进行aav包装和感染测试1.1ybx1敲除细胞株和对照空细胞株(293t)分别铺于6孔板1个孔。1.2第二天细胞汇合度达80％的时候安排转染；转染时配制dna和转染试剂：6孔板质粒时每孔比例为质粒paav-gfp(图6)：phelper：paav-rc：转染试剂(lipofectmine2000)＝1ug：1ug：1ug：6ul；将稀释好dna及转染试剂，常温孵育5min；将稀释好的dna和转染试剂混匀，常温孵育20min；将孔板中的培养基弃上清，把残液吸干净，加入新鲜不含双抗的培养基；将混好的质粒及转染试剂滴加于孔中；转染6小时后，更换新鲜完全培养基；转染48小时后，显微镜下观察，x号克隆ybx1敲除株转染效率明显高于未进行敲除的空细胞，收集上清。测滴度，结果如下：。1.3将293t铺于24孔板，待第二天细胞汇合度达到70％的时候进行病毒感染。将对照组(293t)和目的组(ybx1敲除株)包装得到的aav上清，取相同体积(500ul)的病毒上清分别加入孔中；感染24小时后，更换新鲜完全培养基；感染48小时后，显微镜下观察，目的组(ybx1敲除株)的感染效率明显高于对照组(293t)(图7)，说明，ybx1敲除株能明显提高aav的产量。参考文献：hasegawa,s.l.,doetsch,p.w.,hamilton,k.k.,etal.(1991).dnabindingpropertiesofyb-1anddbpa:bindingtodouble-stranded,single-stranded,andabasicsitecontainingdnas.nucleicacidsres19,4915-4920.izumi,h.,imamura,t.,nagatani,g.,etal.(2001).yboxbindingprotein-1bindspreferentiallytosingle-strandednucleicacidsandexhibits3’–>5’exonucleaseactivity.nucleicacidsres29,1200-1207.wang,x.s.,andsrivasatava,a.(1997).anovelterminalresolution-likesiteintheadeno-associatedvirustype2genome.jvirol71,1140–1146.xiao,x.,xiao,w.,li,j.,andsamulski,r.j.(1997).anovel165-base-pairterminalrepeatsequenceisthesolecisrequirementfortheadeno-associatedviruslifecycle.jvirol71,941–948.satkunanathan,s.,wheeler,j.,thorpe,r.,andzhaoy.(2014).establishmentofanovelcelllinefortheenhancedproductionofrecombinantadeno-associatedvirusvectorsforgenetherapy.humangenetherapy25,929-941.zasedateleva,o.a.,krylov,a.s.,prokopenko,d.v.,etal.(2002).specificityofmammaliany-boxbindingproteinp50ininteractionwithssanddsdnaanalyzedwithgenericoligonucleotidemicrochip.jmolbiol324,73–87.上述实施例为本发明较佳的实施方式，但本发明的实施方式并不受上述实施例的限制，其他的任何未背离本发明的精神实质与原理下所作的改变、修饰、替代、组合、简化，均应为等效的置换方式，都包含在本发明的保护范围之内。序列表<110>和元生物技术(上海)股份有限公司<120>特异性靶向ybx1基因的sgrnas及其应用<160>12<170>siposequencelisting1.0<210>1<211>20<212>dna<213>人工序列(artificialsequence)<400>1tttccaaatccgcccggctt20<210>2<211>20<212>dna<213>人工序列(artificialsequence)<400>2cctgctctgtcggcttctcg20<210>3<211>24<212>dna<213>人工序列(artificialsequence)<400>3accgtttccaaatccgcccggctt24<210>4<211>24<212>dna<213>人工序列(artificialsequence)<400>4aaacaagccgggcggatttggaaa24<210>5<211>24<212>dna<213>人工序列(artificialsequence)<400>5accgcctgctctgtcggcttctcg24<210>6<211>24<212>dna<213>人工序列(artificialsequence)<400>6aaaccgagaagccgacagagcagg24<210>7<211>12584<212>dna<213>人工序列(artificialsequence)<400>7tggaagggctaattcactcccaaagaagacaagatatccttgatctgtggatctaccaca60cacaaggctacttccctgattagcagaactacacaccagggccaggggtcagatatccac120tgacctttggatggtgctacaagctagtaccagttgagccagataaggtagaagaggcca180ataaaggagagaacaccagcttgttacaccctgtgagcctgcatgggatggatgacccgg240agagagaagtgttagagtggaggtttgacagccgcctagcatttcatcacgtggcccgag300agctgcatccggagtacttcaagaactgctgatatcgagcttgctacaagggactttccg360ctggggactttccagggaggcgtggcctgggcgggactggggagtggcgagccctcagat420cctgcatataagcagctgctttttgcctgtactgggtctctctggttagaccagatctga480gcctgggagctctctggctaactagggaacccactgcttaagcctcaataaagcttgcct540tgagtgcttcaagtagtgtgtgcccgtctgttgtgtgactctggtaactagagatccctc600agacccttttagtcagtgtggaaaatctctagcagtggcgcccgaacagggacttgaaag660cgaaagggaaaccagaggagctctctcgacgcaggactcggcttgctgaagcgcgcacgg720caagaggcgaggggcggcgactggtgagtacgccaaaaattttgactagcggaggctaga780aggagagagatgggtgcgagagcgtcagtattaagcgggggagaattagatcgcgatggg840aaaaaattcggttaaggccagggggaaagaaaaaatataaattaaaacatatagtatggg900caagcagggagctagaacgattcgcagttaatcctggcctgttagaaacatcagaaggct960gtagacaaatactgggacagctacaaccatcccttcagacaggatcagaagaacttagat1020cattatataatacagtagcaaccctctattgtgtgcatcaaaggatagagataaaagaca1080ccaaggaagctttagacaagatagaggaagagcaaaacaaaagtaagaccaccgcacagc1140aagcggccggccgctgatcttcagacctggaggaggagatatgagggacaattggagaag1200tgaattatataaatataaagtagtaaaaattgaaccattaggagtagcacccaccaaggc1260aaagagaagagtggtgcagagagaaaaaagagcagtgggaataggagctttgttccttgg1320gttcttgggagcagcaggaagcactatgggcgcagcgtcaatgacgctgacggtacaggc1380cagacaattattgtctggtatagtgcagcagcagaacaatttgctgagggctattgaggc1440gcaacagcatctgttgcaactcacagtctggggcatcaagcagctccaggcaagaatcct1500ggctgtggaaagatacctaaaggatcaacagctcctggggatttggggttgctctggaaa1560actcatttgcaccactgctgtgccttggaatgctagttggagtaataaatctctggaaca1620gatttggaatcacacgacctggatggagtgggacagagaaattaacaattacacaagctt1680aatacactccttaattgaagaatcgcaaaaccagcaagaaaagaatgaacaagaattatt1740ggaattagataaatgggcaagtttgtggaattggtttaacataacaaattggctgtggta1800tataaaattattcataatgatagtaggaggcttggtaggtttaagaatagtttttgctgt1860actttctatagtgaatagagttaggcagggatattcaccattatcgtttcagacccacct1920cccaaccccgaggggacccgacaggcccgaaggaatagaagaagaaggtggagagagaga1980cagagacagatccattcgattagtgaacggatctcgacggtatcgcctttaaaagaaaag2040gggggattggggggtacagtgcaggggaaagaatagtagacataatagcaacagacatac2100aaactaaagaactacaaaaacaaattacaaaaattcaaaattttcgggtttattacaggg2160acagcagagatccagtttatcgatacgcgtgcggccgcccccttcaccgagggcctattt2220cccatgattccttcatatttgcatatacgatacaaggctgttagagagataattggaatt2280aatttgactgtaaacacaaagatattagtacaaaatacgtgacgtagaaagtaataattt2340cttgggtagtttgcagttttaaaattatgttttaaaatggactatcatatgcttaccgta2400acttgaaagtatttcgatttcttggctttatatatcttgtggaaaggacgaaacaccggg2460agacgatgcagtttaaggtttacacctataaaagagagagccgttatcgtctgtttgtgg2520atgtacagagtgatattattgacacgcccgggcgacggatggtgatccccctggccagtg2580cacgtctgctgtcagataaagtctcccgtgaactttacccggtggtgcatatcggggatg2640aaagctggcgcatgatgaccaccgatatggccagtgtgccggtctccgttatcggggaag2700aagtggctgatctcagccaccgcgaaaatgacatcaaaaacgccattaacctgatgttct2760ggggaatataacgtctcagtttcagagctatgctggaaacagcatagcaagttgaaataa28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