本发明属于基因工程领域，更具体地说涉及crispr/cas9靶向敲除人肠癌细胞cnr1基因及其特异性的grna。
背景技术：
：内源性大麻素系统(endocannabmd，ec)是近年来研究较为深入的一个脂质信号和神经调节系统，广泛参与了体内一些生理或病理过程的调节。主要由内源性大麻素、大麻素受体、内源性大麻素的合成酶及降解酶等组成。近年已发现有两种大麻素受体，cnr1受体和cnr2受体，二者均属g蛋白偶联受体。已有大量的研究表明，cnr1是介导内源性大麻素发挥作用的关键因子。cnr1受体包含473个氨基酸，主要位于脑、脊髓与外周神经系统中，脑中的cnr1受体主要分布于基底神经节(黑质、苍白球、外侧纹状体)、海马ca锥体细胞层、小脑和大脑皮质中。推测cnr1受体的这种分布可能与大麻类物质对认知、记忆、运动控制的调节有关；下丘脑的cnr1受体表达区域与食欲调节密切相关。cnr1受体也在外周组织表达，如眼、胃肠道、胎盘和前列腺等；其在胃肠道的分布可能与胃肠蠕动和食管下括约肌功能调节相关，并成为在外周组织发挥作用的基础。cnr1可以激活多重细胞内信号传导通路，调节能量的动态平衡，与肥胖症和糖尿病发生密切相关。肠癌是指发生于肠道的癌病类疾病，是胃肠道中常见的恶性肿瘤，发病率仅次于胃癌和食管癌。目前，研究人员在肠癌的发生机制方面做了大量研究，发现了众多参与肠癌进展的途径和机制，如原癌基因激活、抑癌基因失活(点突变、重排、缺失)及染色体异常等，但目前肠癌发生与进展中所涉及的分子机制仍有许多尚未明确，需要进一步研究。crispr/cas系统是最近几年兴起用于靶向基因特定dna修饰的重要工具，具有快速、高效及特异性靶向等优势，已被应用于多种模式生物，包括人、小鼠、大鼠、斑马鱼、秀丽隐杆线虫、植物及细菌。crisprrna(crrna)与转录激活crrna退火形成的复合物能特异识别基因组序列，引导cas9核酸内切酶在目的片段生成dna双链断裂。这个识别复合体可以通过融合crrna与tracrrna序列形成sgrna进行简化。基因组的靶序列中有长约20bp的片段与crrna或sgrna互补配对；靶序列末端的三核苷酸区域pam(5’-ngg-3’)为spcas9识别位点，是实现剪切功能的关键。sgrna能否做到特异性、精确靶向目标基因是crispr/cas9特异性敲除目标基因的先决条件，脱靶和错误靶向都会影响crispr/cas9对目标基因的特异性敲除，因此，能够设计、制备出精确性和特异性靶向目标基因的sgrna成为crispr/cas9基因敲除的关键技术。本发明利用crispr/cas9慢病毒系统，获得敲除cnr1基因的人肠癌细胞，为研究肠癌细胞中cnr1的作用机制提供有力工具。参考文献：1)bensaidm,gary-bobom,esclangona,maffrandjp,lefurg,oury-donatfandsoubriep,2003.thecannabinoidcb1receptorantagonistsr141716increasesacrp30mrnaexpressioninadiposetissueofobesefa/faratsandinculturedadipocytecells.molecularpharmacology,63(4):908～9142)benzinoum,chevrejc,wardkj,lecoeurc,dinac,lobbenss,durande,delplanquej,horberff,heudeb,balkaub,borch-johnsenk,jorgensent,hansent,pederseno,meyredandfroguelp,2008.endocannabinoidreceptor1genevariationsincreaseriskforobesityandmodulatebodymassindexineuropeanpopulations.humanmoleculargenetics,17(13):1916～19213)blazquezc,chiarlonea,sagredoo,aguadot,pazosmr,resele,palazuelosj,julienb,salazarm,bornerc,benitoc,carrascoc,diez-zaeram,paolettip,diaz-hernandezm,ruizc,sendtnerm,lucasjj,deyebenesjg,marsicanog,monoryk,lutzb,romeroj,alberchj,giness,krausj,fernandez-ruizj,galve-roperhiandguzmanm,2011.lossofstriataltype1cannabinoidreceptorsisakeypathogenicfactorinhuntington'sdisease.brain,134(pt1):119～136技术实现要素：本发明的首要目的在于克服现有技术的缺点与不足，提供一种靶向敲除cnr1基因的sgrna序列。本发明的另一目的在于提供所述一种靶向敲除cnr1基因的crispr/cas9慢病毒系统的应用。本发明的再一目的在于提供靶向敲除cnr1基因的人肠癌细胞ht-29细胞的应用。本发明的目的通过下述技术方案实现：一种靶向敲除cnr1基因的sgrna序列，选自dna序列如下的cnr1sgrna：cnr1sgrna的序列如下：cnr1sgrnaoligo1：5’-caccgttatgaagtcgatcctaga-3’；cnr1sgrnaoligo2：5’-aaactctaggatcgacttcataac-3’；一种靶向敲除cnr1基因的crispr/cas9慢病毒系统，含有上述靶向敲除cnr1基因的sgrna的dna序列。所述的靶向敲除cnr1基因的crispr/cas9慢病毒系统的构建，包括如下步骤：(1)使用bsmbi酶切crispr/cas9慢病毒载体lenti-crisprv2，得到酶切后的crispr/cas9慢病毒载体；(2)将上述靶向敲除cnr1基因的sgrna的dna序列磷酸化后与酶切后的crispr/cas9慢病毒载体连接，得到靶向敲除cnr1基因的crispr/cas9慢病毒系统。步骤(2)中所述的dna序列是将寡核苷酸链1(oligo1)和寡核苷酸链2(oligo2)退火得到双链序列。所述的靶向敲除cnr1基因的crispr/cas9慢病毒系统在制备敲除cnr1基因的细胞株中的应用。一种敲除cnr1基因的细胞株，是将所述的靶向敲除cnr1基因的crispr/cas9慢病毒系统转染目的细胞株得到的。所述的敲除cnr1基因的细胞株，具体是通过如下步骤构建得到：(1)将所述的靶向敲除cnr1基因的crispr/cas9慢病毒系统通过包装细胞进行包装，得到慢病毒颗粒；(2)将慢病毒颗粒感染目的细胞株；(3)将感染后的细胞铺单克隆，得到敲除cnr1基因的细胞株。所述的目的细胞株优选为肿瘤细胞株。所述的肿瘤细胞株优选为肠癌细胞株。所述的肠癌细胞株优选为人结肠癌永生化细胞ht-29。所述的敲除cnr1基因的细胞株，是ht-29细胞中的cnr1基因缺失或插入核苷酸得到的细胞株，为如下的细胞株中的任一株：细胞株2#：sgrna的切割位点位于在野生型cnr1基因上的sgrnapam区前10个碱基处，sgrna序列gttatgaagtcgatcctaga缺失5个碱基，突变为gttatgaagtctaga，导致cnr1基因的第二号外显子缺失5个碱基；细胞株11#：是sgrna的切割位点位于在野生型cnr1基因上的sgrnapam区前3个碱基处，导致cnr1基因的第二号外显子缺失25个碱基。cnr1基因的具体靶向敲除位点与野生型cnr1基因序列的对比如图6和图7。本发明相对于现有技术具有如下的优点及效果：本发明提供cnr1基因的sgrna能特异靶向cnr1基因，将其构建入crispr/cas9慢病毒系统，该系统能特异靶向敲除cnr1基因，得到敲除cnr1基因的细胞株，从而有利于研究细胞株中的cnr1的作用机制。附图说明图1为本发明实施例中使用的载体质粒lenti-crisprv2的质粒图谱；图2为本发明实施例中使用的表达载体质粒lenti-crisprv2-hcnr1sg的质粒图谱；图3为本发明实施例中使用的包装质粒plp-vsvg的质粒图谱；图4为本发明实施例中使用的包装质粒pspax2的质粒图谱；图5为ht-29细胞病毒感染含有cnr1sgrna的活性鉴定测序图；图6为敲除cnr1基因的单克隆ht-29细胞株2#中cnr1基因的测序图；图7为敲除cnr1基因的单克隆ht-29细胞株11#中cnr1基因的测序图。具体实施方式下面结合实施例及附图对本发明作进一步详细的描述，但本发明的实施方式不限于此。实施例11.使用crispr/cas9技术构建敲除cnr1质粒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cnr1的第二外显子上设计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，设计1对crispr寡核苷酸链，见表1。表1cnr1靶向位点及sgrna寡核苷酸序列1.2载体构建1.2.1使用bsmbi酶切2μglenti-crisprv2质粒(购于addgene公司)，2h，37℃,酶切体系如下：2μg(2μl)lenti-crisprv21μlbsmbi(neb)5μl10xcutsmart42μlddh2o50μl合计1.2.2使用捷瑞基因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sgrnaoligos，酶连体系如下：1μloligo1(100μm)1μloligo2(100μm)1μl10xt4链接缓冲液(neb)6.5μlddh2o0.5μlt4pnk(neb)10μl合计pcr仪器退火程序为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oligo双链和酶切后的lenti-crisprv2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lenti-crisprv2-hcnr1sg。1.5敲除效率验证。1)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1×105/孔接种到24孔板培养。待细胞融合度达到70％～80％时更换成opti-mem培养基，1小时后将相应的crispr/cas9敲除质粒0.8μg经lipo2000试剂，转染到293t细胞中，转染48小时后，荧光显微镜下观察转染效果，消化收集细胞，采用基因组dna提取试剂盒提取细胞基因组dna。2)以各组细胞基因组dna为模板，利用表2的引物(seqidno.4～7)pcr扩增靶向序列。用cnr1-4-1和cnr1-4-2进行pcr，得到1034bp的片段a，以a为模板，用cnr1-4-3和cnr1-4-4进行pcr，得到383bp的片段b，该片段电泳后胶回收用于测序，测序引物为cnr1-4-3。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。表2引物序列表引物名称引物序列cnr1-4-1catgtgacacatcagtgctctccnr1-4-2tggaagtcaatgaagctgtagaccnr1-4-3cccaagcttgacctcctactgttcagagacagcnr1-4-4ccggaattccgcagtcatcttctcttgga2.包装慢病毒100mm皿种入4x106个293tn细胞，培养24h(37℃,5％co2)后，更换新鲜培养基，转染包装细胞：a.混匀以下3个质粒：包装质粒pspax212μg；辅助质粒plp-vsvg10μg；表达质粒lenti-crisprv2-hcnr1sg22μg；加入cacl2250μl，加入ddh2o至总体积为500μl，37℃放置20min；b.2xbes500μl，37℃放置20min；c.把a中500μl三质粒混合液逐滴加入b中，并轻轻混匀；d.小心的将c中混合液加入种有293tn的100mm皿中；培养12h后移除感染液，更换10ml新鲜培养基生产病毒液；培养48h后，收获病毒液至50ml离心管中，并重新向100mm皿中补加10ml培养基再次生产病毒；培养24h后，再次收获病毒液，室温离心，1000rpm，10min；使用0.22μm的滤器过滤病毒上清液；将病毒上清液装入超离管中，4℃，100000g，2h；弃去上清液后用opti-mem重悬过夜；再次使用0.22μm的滤器过滤重悬的病毒；分装后，-80℃保存。3.病毒感染1)将要感染的ht-29细胞(购于美国atcc细胞库)接种到六孔板中，过夜，待细胞融合度约为50％时，移去培养基，更换2ml新鲜培养基；2)每孔加入20μl病毒液，再加入12μl聚凝胺polybrene，至终浓度为6μg/ml；3)37℃培养12h后，移除培养液，更换新鲜培养液后继续培养48h；4.筛选稳定细胞株感染结束48h后，向每孔加入嘌呤霉素(2.0μg/ml)，隔天换液，并保持培养基的嘌呤霉素浓度恒定，筛选阳性克隆细胞。所得细胞株命名为ht-29-cnr1。并使用有限稀释法挑选ht-29-cnr1单克隆敲除细胞株。5.稳定细胞株鉴定及基因打靶方式鉴定以各组敲除cnr1的单克隆细胞株基因组dna为模板，分别设计针对cnr1第二外显子的引物，序列如seqidno.4～7进行两轮巢式pcr扩增，对获得的第二轮pcr产物进行测序，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cnr1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cnr1基因敲除。结果显示，鉴定出sgrna是有活性的(图5)，并且找到2种类型的具有cnr1基因缺失或插入突变的稳定细胞株(图6和图7)。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>crispr/cas9靶向敲除人肠癌细胞cnr1基因及其特异性的sgrna<160>11<170>siposequencelisting1.0<210>1<211>20<212>dna<213>人工序列(artificialsynthesis)<400>1gttatgaagtcgatcctaga20<210>2<211>24<212>dna<213>人工序列(artificialsynthesis)<400>2caccgttatgaagtcgatcctaga24<210>3<211>24<212>dna<213>人工序列(artificialsynthesis)<400>3aaactctaggatcgacttcataac24<210>4<211>22<212>dna<213>人工序列(artificialsynthesis)<400>4catgtgacacatcagtgctctc22<210>5<211>23<212>dna<213>人工序列(artificialsynthesis)<400>5tggaagtcaatgaagctgtagac23<210>6<211>32<212>dna<213>人工序列(artificialsynthesis)<400>6cccaagcttgacctcctactgttcagagacag32<210>7<211>29<212>dna<213>人工序列(artificialsynthesis)<400>7ccggaattccgcagtcatcttctcttgga29<210>8<211>14873<212>dna<213>人工序列(artificialsynthesis)<400>8tgatgcggttttggcagtacatcaatgggcgtggatagcggtttgactcacggggatttc60caagtctccaccccattgacgtcaatgggagtttgttttggcaccaaaatcaacgggact120ttccaaaatgtcgtaacaactccgccccattgacgcaaatgggcggtaggcgtgtacggt180gggaggtctatataagcagcgcgttttgcctgtactgggtctctctggttagaccagatc240tgagcctgggagctctctggctaactagggaacccactgcttaagcctcaataaagcttg300ccttgagtgcttcaagtagtgtgtgcccgtctgttgtgtgactctggtaactagagatcc360ctcagacccttttagtcagtgtggaaaatctctagcagtggcgcccgaacagggacttga420aagcgaaagggaaaccagaggagctctctcgacgcaggactcggcttgctgaagcgcgca480cggcaagaggcgaggggcggcgactggtgagtacgccaaaaattttgactagcggaggct540agaaggagagagatgggtgcgagagcgtcagtattaagcgggggagaattagatcgcgat600gggaaaaaattcggttaaggccagggggaaagaaaaaatataaattaaaacatatagtat660gggcaagcagggagctagaacgattcgcagttaatcctggcctgttagaaacatcagaag720gctgtagacaaatactgggacagctacaaccatcccttcagacaggatcagaagaactta780gatcattatataatacagtagcaaccctctattgtgtgcatcaaaggatagagataaaag840acaccaaggaagctttagacaagatagaggaagagcaaaacaaaagtaagaccaccgcac900agcaagcggccgctgatcttcagacctggaggaggagatatgagggacaattggagaagt960gaattatataaatataaagtagtaaaaattgaaccattaggagtagcacccaccaaggca1020aagagaagagtggtgcagagagaaaaaagagcagtgggaataggagctttgttccttggg1080ttcttgggagcagcaggaagcactatgggcgcagcgtcaatgacgctgacggtacaggcc1140agacaattattgtctggtatagtgcagcagcagaacaatttgctgagggctattgaggcg1200caacagcatctgttgcaactcacagtctggggcatcaagcagctccaggcaagaatcctg1260gctgtggaaagatacctaaaggatcaacagctcctggggatttggggttgctctggaaaa1320ctcatttgcaccactgctgtgccttggaatgctagttggagtaataaatctctggaacag1380atttggaatcacacgacctggatggagtgggacagagaaattaacaattacacaagctta1440atacactccttaattgaagaatcgcaaaaccagcaagaaaagaatgaacaagaattattg1500gaattagataaatgggcaagtttgtggaattggtttaacataacaaattggctgtggtat1560ataaaattattcataatgatagtaggaggcttggtaggtttaagaatagtttttgctgta1620ctttctatagtgaatagagttaggcagggatattcaccattatcgtttcagacccacctc1680ccaaccccgaggggacccgacaggcccgaaggaatagaagaagaaggtggagagagagac1740agagacagatccattcgattagtgaacggatcggcactgcgtgcgccaattctgcagaca1800aatggcagtattcatccacaattttaaaagaaaaggggggattggggggtacagtgcagg1860ggaaagaatagtagacataatagcaacagacatacaaactaaagaattacaaaaacaaat1920tacaaaaattcaaaattttcgggtttattacagggacagcagagatccagtttggttaat1980taaggtaccgagggcctatttcccatgattccttcatatttgcatatacgatacaaggct2040gttagagagataattagaattaatttgactgtaaacacaaagatattagtacaaaatacg2100tgacgtagaaagtaataatttcttgggtagtttgcagttttaaaattatgttttaaaatg2160gactatcatatgcttaccgtaacttgaaagtatttcgatttcttggctttatatatcttg2220tggaaaggacgaaacaccggagacggttgtaaatgagcacacaaaatacacatgctaaaa2280tattatattctatgacctttataaaatcaaccaaaatcttctttttaataactttagtat2340caataattagaatttttatgttcctttttgcaaacttttaataaaaatgagcaaaataaa2400aaaacgctagttttagtaactcgcgttgttttcttcacctttaataatagctactccacc2460acttgttcctaagcggtcagctcctgcttcaatcattttttgagcatcttcaaatgttct2520aactccaccagctgctttaactaaagcattgtctttaacaactgacttcattagtttaac2580atcttcaaatgttgcacctgattttgaaaatcctgttgatgttttaacaaattctaatcc2640agcttcaacagctatttcacaagctttcatgatttcttcttttgttaataaacaattttc2700cataatacatttaacaacatgtgatccagctgctttttttacagctttcatgtcttctaa2760aactaattcataatttttgtcttttaatgcaccaatatttaataccatatcaatttctgt2820tgcaccatctttaattgcttcagaaacttcgaatgcttttgtagctgttgtgcatgcacc2880tagaggaaaacctacaacatttgttattcctacatttgtgccttttaataattctttaca2940atagcttgttcaatatgaattaacacaaactgttgcaaaatcaaattcaattgcttcatc3000acataattgtttaatttcagcttt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