一种可实现靶基因两种剂量表达的基因修饰模型构建方法与流程

本发明属于生物
技术领域：
，具体涉及一种可实现靶基因两种剂量表达的基因修饰模型的构建方法。
背景技术：
：利用基因过表达制备细胞或者动物模型，是研究基因功能的常用技术，广泛应用于生命科学和医学基础研究以及临床前转化研究。常用的转基因动物模型为研究早期发育、疾病的发病过程和临床前药物评价提供了强有力的工具。研究发现，在这些模型中基因的表达剂量和表型密切相关，如：fus基因发现与肌萎缩性侧索硬化症（als）和额颞叶变性（ftld）有关，在过表达野生型人fus基因的转基因纯合子小鼠中（fus蛋白表达量是对照组的1.9倍），早期出现震颤，随后后肢逐渐瘫痪，12周的时候小鼠死亡；然而在过表达野生型人fus基因的转基因杂合子小鼠中（fus蛋白表达量是对照组的1.4倍），没有出现运动障碍的表型或者病理病变。这说明人fus基因过表达导致进行性神经元病变是剂量依赖的(jacquelinec.mitchelletal.,overexpressionofhumanwild-typefuscausesprogressivemotorneurondegenerationinanage-anddose-dependentfashion.actaneuropathol(2013)125:273–288,doi10.1007/s00401-012-1043-z）。同样的情况也发生在人源tlr8转基因小鼠模型中(cristianaguiducci,et.al,rnarecognitionbyhumantlr8canleadtoautoimmuneinflammation,j.exp.med.2013vol.210no.132903-2919),人源tlr8的表达会导致小鼠自身免疫性疾病的发生，小鼠的存活率及表型和人tlr8的表达量呈明显的负相关。由此可见，不同剂量的基因表达水平，可以使基因展现出不同的生物学功能；不同剂量的表达模型可以为我们理解基因功能提供更全面的线索。而我们目前常用的不同表达剂量模型，通常有：1）杂合子和纯合子动物模型间的基因表达剂量差异，但两种基因型间的基因表达水平差异有限；2）通过传统的随机转基因方式，因为插入位点和基因拷贝数不同，可以获得不同表达水平的动物模型，但因为常规的转基因模型需要复杂的建系过程，从多个品系中筛选出可传代，目的基因表达的动物模型，增加了时间和成本。鉴于此，通过一种小鼠模型，高效获得两种或者多种不同表达剂量的模型，对于基因生命科学、医学研究具有重要意义，亦有需求，特提出本发明。技术实现要素：本发明的目的在于提供一种可实现靶基因高、低两种剂量表达的基因修饰模型构建方法以及用于实施该方法的核酸序列、载体和宿主，利用该方法制备的细胞和动物模型中，可以实现敲入基因分别由内源启动子驱动表达，实现较低水平的转基因表达或由强启动子（如cag启动子）来驱动目的基因高表达。根据本发明的一个方面，提供一种用于基因重组表达的核酸序列，其包含：外源强启动子序列和至少一个控制启动子序列功能的调控元件。优选地，所述在所述外源强自动子上游和下游各有一个控制启动子序列功能的调控元件，其能够调控外源强启动子和/或所述核酸序列插入位置的内源启动子的功能。优选地，所述调控序列为终止元件，其两端具有重组酶识别元件。本发明还提供了包含上述核酸序列的载体、宿主，以及构建所述载体和宿主的方法，所述方法包括讲所述核酸序列引入所述载体以及将所述核酸序列或载体引入所述宿主。进一步地，本发明还提供了利用所述核酸序列、载体或宿主表达外源基因的方法。敲入基因可以根据需求实现不同强度的过表达。具体的方法为：如图1所示，通过定点敲入的方式，将rna剪接受体序列（sa）-重组酶1识别元件-终止元件1-cag强启动子-重组酶1识别元件-重组酶2识别元件-终止元件2-重组酶2识别元件-靶基因-polya片段插入基因编码外显子之前的内含子中。正常情况下，因为转录终止元件1和2的存在，所插入位置基因内源启动子和cag强启动子的转录都被终止，目的基因不表达；当通过重组酶2作用后，stop2元件将被剔除掉，目的基因为cag强启动子所驱动，实现目的基因高表达；在此基础上，可通过重组酶1进一步作用，将stop1元件和cag启动子剔除掉，目的基因受插入位置基因内源启动子驱动表达，实现一个较低剂量表达。上述名词中，cag启动子为巨细胞病毒（cmv）增强子元件、鸡β-肌动蛋白基因的启动子和兔β-珠蛋白基因的剪接受体组合而成的一个人工启动子，具有较高的转录活性，强启动子也可以选择其他启动子，优选cag启动子；重组酶识别元件为各种重组酶所识别的dna序列，常用的有loxp，frt,rox序列等，重组酶1识别元件可以为loxp，frt,rox序列的任意一种，优选非loxp序列；重组酶2识别元件可以为loxp，frt,rox序列的任意一种，优选loxp序列；重组酶1识别元件和重组酶2识别元件需为不同种类的识别元件，对应的重组酶应为两种不同的重组酶；终止元件1为转录终止元件，发挥终止内源基因转录的作用，避免对外源强启动子表达有影响；终止元件2为转录或者翻译终止元件，用于终止强启动子的转录或者终止强启动子转录出mrna的翻译，避免外源基因、蛋白表达。polya为polyadenylation缩写，用于终止转录。本发明可以通过构建一个模型，实现靶基因的两种剂量表达，为观察不同靶基因表达剂量对基因功能的影响研究提供了简便的方法。附图说明为了更清楚的说明本发明实施例或现有技术中的技术方案，下面将对实施例中所需使用的附图做简单的介绍，应当理解，以下附图仅示出了本发明的某些实施例，因此不应被看作是对范围的限定，对于本领域普通技术人员来讲，在不付出创造性劳动的前提下，还可以根据这些附图获得其他相关的附图。图1是本
发明内容的示意图。图2tdtomato-luciferase条件性高低剂量过表达小鼠胚胎干细胞模型构建。图a为模型构建验证示意图；图b为tdtomato-luciferase酶切验证结果。载体经ecori限制性内切酶，酶切后理论条带大小为9831bp、5617bp、3969bp、3640bp、932bp，结果符合预期（m为1kbdnamarker）；图c为同源重组es细胞鉴定电泳结果，上图为5’同源臂鉴定结果，下图为3’同源臂鉴定结果（m为1kbdnamarker，图上方编号为es细胞克隆编号）。图3tdtomato-luciferase阳性es细胞克隆目的基因高低剂量表达验证。图a为cre作用后阳性es细胞克隆pcr鉴定电泳结果；图b为cre/flp两种重组酶作用后阳性es细胞克隆pcr鉴定电泳结果；图c为野生型es细胞克隆、tdtomato-luciferase阳性es细胞克隆、cre作用后阳性es细胞克隆、cre/flp两种重组酶作用后阳性es细胞克隆egfp和tdtomato荧光表达检测结果；图d为野生型es细胞克隆、tdtomato-luciferase阳性es细胞克隆、cre作用后阳性es细胞克隆、cre/flp两种重组酶作用后阳性es细胞克隆luciferase荧光素酶活性表达检测结果。图4egfp高、低剂量过表达小鼠模型构建及验证。图a为模型构建验证示意图；图b为同源重组f1代小鼠鉴定电泳结果，左图为5’同源臂鉴定结果，右图为3’同源臂鉴定结果（m为1kbdnamarker，图上方数字为f1代小鼠编号）；图c-f图为分别检测野生型小鼠（wt）、cre重组酶去stop2小鼠（cag-egfp）、cre/flp重组酶去stop1和stop2小鼠（r26-egfp）的脑（c）、肾脏（d）、肝脏（e）、肺（f）四个组织egfp荧光表达结果，左图为荧光显微镜拍照结果，右图为左图荧光结果的定量分析结果。具体实施方式实施例一tdtomato-luciferase条件性高低剂量过表达小鼠胚胎干细胞模型为验证本发明在细胞水平实现高低剂量过表达调控的可行性，发明人通过同源重组方式将sa-frt-egfp-stop1-cag-frt-loxp-stop2-loxp-tdtomato-ires-luciferase-wpre-polya-pgk-neo-polya元件插入在小鼠rosa26基因的内含子1中，策略示意图2所示。利用该策略，发明人可通过检测tdtomato红色荧光和luciferase荧光素酶催化底物发光的活性来反映是否可以实现目的基因tdtomato和luciferase高低剂量表达。同时该策略引入了一个egfp绿色荧光蛋白，便于验证片段插入后rosa26启动子的活性。该模型的构建和验证过程如下：1）同源重组载体构建通过全基因合成获得同源重组左臂(seqidno:1)，同源重组右臂(seqidno:2)，sa-frt-egfp-stop1(sa、frt、egfp、stop1序列依次为seqidno:3-6)，frt-loxp-stop2-loxp(loxp、stop2序列依次为seqidno:7-8)，tdtomato-ires-luciferase-wpre-polya(tdtomato、ires、luciferase、wpre-polya序列依次为seqidno:9-12),pgk-neo-polya(seqidno:13)片段，通过pcr扩增的方式以plvct-ttr-krab质粒为模板扩增获得cag启动子(seqidno:14)，然后通过in-fusion的方法，将同源重组左臂、sa-frt-egfp-stop1、cag启动子、frt-loxp-stop2-loxp、tdtomato-ires-luciferase-wpre-polya、pgk-neo-polya和同源重组右臂片段按序插入到pbr322骨架载体（seqidno:15）中，构建获得tdtomato-luciferase同源重组载体(seqidno:16)，该载体经酶切（ecori酶切结果如图2b所示）、测序确认正确后进行后续的es细胞转染。2)tdtomato-luciferase阳性es细胞克隆获得验证正确的重组载体经大肠杆菌转化扩增，抽提纯化后，将重组打靶载体利用电转的方式转染小鼠胚胎干细胞（es细胞）。利用neomycine抗生素进行筛选，共计获得96个抗性克隆，通过pcr技术对抗性克隆进行鉴定，共计获得10个同源重组阳性克隆，阳性克隆两个同源臂重组的pcr鉴定结果如图2c所示（上图为5’臂同源重组阳性克隆鉴定结果，下图为3’臂同源重组阳性克隆鉴定结果，编号为克隆编号，m为1kbdnamarker），5’臂同源重组阳性克隆可扩增出3.4kb大小的阳性条带，阴性克隆只能扩增出6.8kb的阴性条带；3’臂同源重组阳性克隆应扩增出7.7kb片段，阴性克隆应无产物。pcr信息和反应条件如下：引物序列信息：引物名称序列信息（5’->3'）5'臂鉴定正向引物gccgggcctcgtcgtct(seqidno:17)5'臂鉴定反向引物tttttgggggtgatggtggtc(seqidno:18)3’臂鉴定正向引物tcgcagcctaccgtggtgttcgtt(seqidno:19)3’臂鉴定反向引物ggtgttaattggcagtctatgagc(seqidno:20)pcr反应体系：pcr反应组成体积(µl)ddh2o31pcrbuffer102.5mmdntp4正向引物1反向引物1dnapolymerase2genomicdna1总计50pcr反应程序：步骤温度（℃）时间备注1982min29815sec36815sec4684min重复步骤2-4共34个循环5681min61210min　3)tdtomato-luciferasees细胞克隆cre重组酶、cre/flp重组酶处理tdtomato-luciferase阳性es细胞克隆分别转染cre重组酶表达质粒、cre/flp重组酶表达质粒,转染72小时后，进行单克隆培养，分别获得剔除stop2元件、剔除stop1和stop2元件的单克隆。tdtomato-luciferase阳性es细胞克隆，转染cre质粒，获得剔除stop2元件，由cag启动子驱动tdtomato-luciferase表达的es细胞克隆，pcr鉴定方法如下所示：引物序列信息如下表，pcr反应体系和反应程序除退火温度为63℃外，其他同上引物名称序列信息（5’->3'）鉴定正向引物cgccgccgcaccccctcta(seqidno:21)鉴定反向引物ggcaccagcagcgcactttgaatc(seqidno:22)pcr鉴定阳性克隆应可扩增出3.4kb大小的条带，电泳结果如图3a所示，经pcr产物测序确认d7和d11为去除stop2的阳性克隆。tdtomato-luciferase阳性es细胞克隆，转染cre/flp质粒，获得剔除stop1和stop2元件，由rosa26基因启动子驱动tdtomato-luciferase表达的es细胞克隆，pcr鉴定方法如下所示：引物序列信息如下表，pcr反应体系和反应程序同上引物名称序列信息（5’->3'）鉴定正向引物tccgcccaccgccccacact(seqidno:23)鉴定反向引物ggcaccagcagcgcactttgaatc(seqidno:24)pcr鉴定阳性克隆应可扩增出7.3kb大小的条带，电泳结果如图3a所示，经pcr产物测序确认g4和g6为去除stop1和stop2的阳性克隆。4）重组酶作用前后目的基因tdtomato和luciferase表达量检测。①tdtomato荧光表达检测：分别对c57bl/6野生型细胞、tdtomato-luciferase打靶阳性es细胞、转染cre质粒去stop2的es细胞和转染cre+flp质粒去stop1及stop2的es细胞，观察每组细胞的荧光表达情况。如图3c所示，c57bl/6野生型细胞作为对照组没有检测到任何荧光表达；tdtomato-luciferase打靶阳性es细胞只能检测到rosa26内源启动子驱动的绿色荧光表达，无tdtomato荧光表达；tdtomato-luciferase打靶阳性转染cre质粒es细胞可以检测到rosa26内源启动子驱动的绿色荧光和cag强启动子驱动的红色荧光表达；tdtomato-luciferase打靶阳性转染cre+flp质粒的es细胞只能检测到rosa26基因启动子驱动的tdtomato红色荧光的表达，而且红色荧光的表达强度明显弱于转染cre质粒es细胞组，这是因为在转染cre质粒es细胞组中红色荧光是受cag启动子驱动表达，在转染cre+flp质粒es细胞中红色荧光是受rosa26基因内源性启动子驱动表达。这说明本发明的基因双表达系统可以通过重组酶作用实现诱导不同强度的靶基因过表达。②体外luciferase荧光素酶活性检测:在上述tdtomato荧光检测部分，发明人通过荧光对目的基因表达进行了比较分析，为进一步说明本应用的可行性，发明人通过检测另一个目的基因luciferase催化底物产生荧光的能力，从功能层面上也对目的基因表达进行了检测。分别对c57bl/6野生型细胞、tdtomato-luciferase打靶阳性es细胞、转染cre质粒去stop2的es细胞和转染cre+flp质粒去stop1及stop2的es细胞进行培养，培养至80%密度时，去培养基，pbs洗两次后，分别加入500ml裂解液（promega），4℃，12000rpm，离心15分钟，取上清即为蛋白样品。使用luciferaseassaysystem试剂盒（promega），用化学发光检测仪luminometertd-20/20(lumatlb9507)检测荧光强度。细胞蛋白浓度采用bca法测定（pierce）,定量后将测得的荧光数值用蛋白浓度来标准化，得到每毫克蛋白的荧光值。如图3d所示，c57bl/6野生型es细胞和tdtomato-luciferase打靶阳性es细胞不能检测到luciferase表达，而转染cre质粒后luciferase受cag启动子驱动表达强度是转染cre/flp质粒后luciferase受rosa26基因内源性启动子驱动表达强度的3倍左右。实施例二egfp高、低剂量过表达小鼠模型为验证本发明在动物体内实现高低剂量目的基因过表达调控的可行性，发明人通过同源重组方式将sa-frt-stop1-cag-frt-loxp-stop2-loxp-egfp-wpre-polya-pgk-neo-polya元件插入在小鼠rosa26基因的内含子1中，制备策略如示意图4a，具体构建及检测过程描述如下：1）同源重组载体构建利用实施例一中构建获得的tdtomato-luciferase同源重组载体(seqidno:16)为模板，pcr扩增获得同源重组左臂(seqidno:1)，同源重组右臂(seqidno:2)、sa-frt-stop1、cag、frt-loxp-stop2-loxp、egfp-wpre-polya等载体元件，然后通过in-fusion的方法，将各元件插入到pbr322骨架载体（seqidno:15）中，构建获得egfp同源重组载体，载体经酶切、测序确认后进行后续的受精卵注射。2）egfp基因敲入小鼠获得egfp同源重组载体质粒与cas9、grna混合后进行小鼠受精卵显微注射，注射后的受精卵经培养箱短暂培养后，移植至受体母鼠的输卵管，获得基因修饰小鼠f0代小鼠，同源重组阳性的f0代小鼠通过与野生型小鼠交配，繁育获得f1代小鼠。通过pcr方法对f1代小鼠是否为正确发生同源重组的阳性小鼠进行鉴定，发生5’臂同源重组阳性的小鼠应可扩增出3.4kb的pcr产物，阴性小鼠只能扩增出6.8kb的pcr产物；发生3’臂同源重组阳性的小鼠应可扩增出4.1kb的pcr产物，阴性小鼠无pcr产物扩增出。部分小鼠的鉴定结果如图4b所示，1-4号小鼠为5’和3’臂同源重组阳性的小鼠（该小鼠简称：egfp小鼠）；该小鼠通过和dppa3-cre小鼠（一种在胚胎早期及生殖细胞系中高表达cre的小鼠品系，可用于小鼠整体全身性的去除loxp之间的元件）交配，获得去掉stop2元件的egfp小鼠,在该小鼠中egfp表达受cag启动子驱动（简称:cag-egfp）；将cag-egfp小鼠与cag-flp转基因小鼠交配，获得同时去掉stop1和stop2元件的egfp小鼠,在该小鼠中egfp表达受rosa26内源启动子驱动（简称:r26-egfp）。3）egfp小鼠、r26-egfp小鼠、cag-egfp小鼠体内各组织egfp荧光表达检测分别取egfp小鼠、r26-egfp小鼠、cag-egfp小鼠的脑（图4c）、肾脏(图4d)、肝脏(图4e)和肺(图4f)组织进行egfp荧光检测,并对荧光结果进行定量比较分析（图4c-f,左图为荧光照片结果，右图为荧光定量分析）。结果显示：egfp小鼠各组织中无egfp荧光检测到，r26-egfp小鼠和cag-egfp小鼠4个组织均可以检测到egfp荧光的表达，cag-egfp小鼠各组织中荧光显著高于r26-egfp小鼠，4个组织中cag-egfp小鼠较r26-egfp小鼠荧光强度的比值略有不同，cag-egfp小鼠组织荧光约是r26-egfp小鼠的3-4倍。该结果说明本发明的两种剂量表达系统可以在体内通过重组酶作用实现诱导不同强度的靶基因过表达。综上所述，给予两个实施例的结果，发明人通过在rosa26基因位点分别插入不同的目的基因，借助重组酶系统，分别在细胞水平和小鼠体内水平证明本发明“一种可实现靶基因两种剂量表达的基因修饰模型构建方法”，可以通过构建一个模型，实现靶基因的两种剂量表达，为观察不同靶基因表达剂量对基因功能的影响研究提供了简便的方法。以上所述，仅为本发明的具体实施方式，但本发明的保护范围并不局限于此，任何熟悉本
技术领域：
的技术人员在本发明揭露的技术范围内，可轻易想到变化或替换，都应涵盖在本发明的保护范围之内。因此，本发明的保护范围应以所述权利要求的保护范围为准。序列表<110>上海南方模式生物科技股份有限公司上海砥石生物科技有限公司广东南模生物科技有限公司<120>一种可实现靶基因两种剂量表达的基因修饰模型构建方法<160>24<170>siposequencelisting1.0<210>1<211>3300<212>dna<213>人工序列(artificialsequence)<400>1gccagcgggggcggcgaggaggcgctcccaggttccggccctcccctcggccccgcgccg60cagagtctggccgcgcgcccctgcgcaacgtggcaggaagcgcgcgctgggggcggggac120gggcagtagggctgagcggctgcggggcgggtgcaagcacgtttccgacttgagttgcct180caagaggggcgtgctgagccagacctccatcgcgcactccggggagtggagggaaggagc240gagggctcagttgggctgttttggaggcaggaagcacttgctctcccaaagtcgctctga300gttgttatcagtaagggagctgcagtggagtaggcggggagaaggccgcacccttctccg360gaggggggaggggagtgttgcaatacctttctgggagttctctgctgcctcctggcttct420gaggaccgccctgggcctgggagaatcccttccccctcttccctcgtgatctgcaactcc480agtctttctagaagatgggcgggagtcttctgggcaggcttaaaggctaacctggtgtgt540gggcgttgtcctgcaggggaattgaacaggtgtaaaattggagggacaagacttcccaca600gattttcggttttgtcgggaagttttttaataggggcaaataaggaaaatgggaggatag660gtagtcatctggggttttatgcagcaaaactacaggttattattgcttgtgatccgcctc720ggagtattttccatcgaggtagattaaagacatgctcacccgagttttatactctcctgc780ttgagatccttactacagtatgaaattacagtgtcgcgagttagactatgtaagcagaat840tttaatcatttttaaagagcccagtacttcatatccatttctcccgctccttctgcagcc900ttatcaaaaggtattttagaacactcattttagccccattttcatttattatactggctt960atccaacccctagacagagcattggcattttccctttcctgatcttagaagtctgatgac1020tcatgaaaccagacagattagttacatacaccacaaatcgaggctgtagctggggcctca1080acactgcagttcttttataactccttagtacactttttgttgatcctttgccttgatcct1140taattttcagtgtctatcacctctcccgtcaggtggtgttccacatttgggcctattctc1200agtccagggagttttacaacaatagatgtattgagaatccaacctaaagcttaactttcc1260actcccatgaatgcctctctcctttttctccatttataaactgagctattaaccattaat1320ggtttccaggtggatgtctcctcccccaatattacctgatgtatcttacatattgccagg1380ctgatattttaagacattaaaaggtatatttcattattgagccacatggtattgattact1440gcttactaaaattttgtcattgtacacatctgtaaaaggtggttccttttggaatgcaaa1500gttcaggtgtttgttgtctttcctgacctaaggtcttgtgagcttgtattttttctattt1560aagcagtgctttctcttggactggcttgactcatggcattctacacgttattgctggtct1620aaatgtgattttgccaagcttcttcaggacctataattttgcttgacttgtagccaaaca1680caagtaaaatgattaagcaacaaatgtatttgtgaagcttggtttttaggttgttgtgtt1740gtgtgtgcttgtgctctataataatactatccaggggctggagaggtggctcggagttca1800agagcacagactgctcttccagaagtcctgagttcaattcccagcaaccacatggtggct1860cacaaccatctgtaatgggatctgatgccctcttctggtgtgtctgaagaccacaagtgt1920attcacattaaataaataaatcctccttcttcttcttttttttttttttaaagagaatac1980tgtctccagtagaatttactgaagtaatgaaatactttgtgtttgttccaatatggtagc2040caataatcaaattactctttaagcactggaaatgttaccaaggaactaatttttatttga2100agtgtaactgtggacagaggagccataactgcagacttgtgggatacagaagaccaatgc2160agactttaatgtcttttctcttacactaagcaataaagaaataaaaattgaacttctagt2220atcctatttgtttaaactgctagctttacttaacttttgtgcttcatctatacaaagctg2280aaagctaagtctgcagccattactaaacatgaaagcaagtaatgataattttggatttca2340aaaatgtagggccagagtttagccagccagtggtggtgcttgcctttatgcctttaatcc2400cagcactctggaggcagagacaggcagatctctgagtttgagcccagcctggtctacaca2460tcaagttctatctaggatagccaggaatacacacagaaaccctgttggggaggggggctc2520tgagatttcataaaattataattgaagcattccctaatgagccactatggatgtggctaa2580atccgtctacctttctgatgagatttgggtattattttttctgtctctgctgttggttgg2640gtcttttgacactgtgggctttctttaaagcctccttcctgccatgtggtctcttgtttg2700ctactaacttcccatggcttaaatggcatggctttttgccttctaagggcagctgctgag2760atttgcagcctgatttccagggtggggttgggaaatctttcaaacactaaaattgtcctt2820taattttttttttaaaaaatgggttatataataaacctcataaaatagttatgaggagtg2880aggtggactaatattaaatgagtccctcccctataaaagagctattaaggctttttgtct2940tatacttaactttttttttaaatgtggtatctttagaaccaagggtcttagagttttagt3000atacagaaactgttgcatcgcttaatcagattttctagtttcaaatccagagaatccaaa3060ttcttcacagccaaagtcaaattaagaatttctgacttttaatgttaatttgcttactgt3120gaatataaaaatgatagcttttcctgaggcagggtctcactatgtatctctgcctgatct3180gcaacaagatatgtagactaaagttctgcctgcttttgtctcctgaatactaaggttaaa3240atgtagtaatacttttggaacttgcaggtcagattcttttataggggacacactaaggga3300<210>2<211>3300<212>dna<213>人工序列(artificialsequence)<400>2gcttgggtgatagttggtaaaatgtgtttcaagtgatgaaaacttgaattattatcaccg60caacctactttttaaaaaaaaaagccaggcctgttagagcatgcttaagggatccctagg120acttgctgagcacacaagagtagttacttggcaggctcctggtgagagcatatttcaaaa180aacaaggcagacaaccaagaaactacagttaaggttacctgtctttaaaccatctgcata240tacacagggatattaaaatattccaaataatatttcattcaagttttcccccatcaaatt300gggacatggatttctccggtgaataggcagagttggaaactaaacaaatgttggttttgt360gatttgtgaaattgttttcaagtgatagttaaagcccatgagatacagaacaaagctgct420atttcgaggtctcttggtttatactcagaagcacttctttgggtttccctgcactatcct480gatcatgtgctaggcctaccttaggctgattgttgttcaaataaacttaagtttcctgtc540aggtgatgtcatatgatttcatatatcaaggcaaaacatgttatatatgttaaacatttg600tacttaatgtgaaagttaggtctttgtgggtttgatttttaattttcaaaacctgagcta660aataagtcatttttacatgtcttacatttggtggaattgtataattgtggtttgcaggca720agactctctgacctagtaaccctacctatagagcactttgctgggtcacaagtctaggag780tcaagcatttcaccttgaagttgagacgttttgttagtgtatactagtttatatgttgga840ggacatgtttatccagaagatattcaggactatttttgactgggctaaggaattgattct900gattagcactgttagtgagcattgagtggcctttaggcttgaattggagtcacttgtata960tctcaaataatgctggccttttttaaaaagcccttgttctttatcaccctgttttctaca1020taatttttgttcaaagaaatacttgtttggatctccttttgacaacaatagcatgttttc1080aagccatattttttttcctttttttttttttttttggtttttcgagacagggtttctctg1140tatagccctggctgtcctggaactcactttgtagaccaggctggcctcgaactcagaaat1200ccgcctgcctctgcctcctgagtgccgggattaaaggcgtgcaccaccacgcctggctaa1260gttggatattttgttatataactataaccaatactaactccactgggtggatttttaatt1320cagtcagtagtcttaagtggtctttattggcccttcattaaaatctactgttcactctaa1380cagaggctgttggtactagtggcacttaagcaacttcctacggatatactagcagattaa1440gggtcagggatagaaactagtctagcgttttgtatacctaccagctttatactaccttgt1500tctgatagaaatatttcaggacatctagagtgtactataaggttgatggtaagcttataa1560ggaacttgaaagtggagtaactactccatttctctgaggggagaattaaaatttttgacc1620aagtgttgttgagccactgagaatggtctcagaacataacttcttaaggaaccttcccag1680attgccctcaacactgcaccacatttggtcctgcttgaacattgccatggctcttaaagt1740cttaattaagaatattaattgtgtaattattgtttttcctcctttagatcattccttgag1800gacaggacagtgcttgtttaaggctatatttctgctgtctgagcagcaacaggtcttcga1860gatcaacatgatgttcataatcccaagatgttgccatttatgttctcagaagcaagcaga1920ggcatgatggtcagtgacagtaatgtcactgtgttaaatgttgctatgcagtttggattt1980ttctaatgtagtgtaggtagaacatatgtgttctgtatgaattaaactcttaagttacac2040cttgtataatccatgcaatgtgttatgcaattaccattttaagtattgtagctttctttg2100tatgtgaggataaaggtgtttgtcataaaatgttttgaacatttccccaaagttccaaat2160tataaaaccacaacgttagaacttatttatgaacaatggttgtagtttcatgcttttaaa2220atgcttaattattcaattaacaccgtttgtgttataatatatataaaactgacatgtaga2280agtgtttgtccagaacatttcttaaatgtatactgtctttagagagtttaatatagcatg2340tcttttgcaacatactaacttttgtgttggtgcgagcaatattgtgtagtcattttgaaa2400ggagtcatttcaatgagtgtcagattgttttgaatgttattgaacattttaaatgcagac2460ttgttcgtgttttagaaagcaaaactgtcagaagctttgaactagaaattaaaaagctga2520agtatttcagaagggaaataagctacttgctgtattagttgaaggaaagtgtaatagctt2580agaaaatttaaaaccatatagttgtcattgctgaatatctggcagatgaaaagaaatact2640cagtggttcttttgagcaatataacagcttgttatattaaaaattttccccacagatata2700aactctaatctataactcataaatgttacaaatggatgaagcttacaaatgtggcttgac2760ttgtcactgtgcttgttttagttatgtgaaagtttggcaataaacctatgtcctaaatag2820tcaaactgtggaatgactttttaatctattggtttgtctagaacagttatgttgccattt2880gccctaatggtgaaagaaaaagtggggagtgccttggcactgttcatttgtggtgtgaac2940caaagaggggggcatgcacttacacttcaaacatccttttgaaagactgacaagtttggg3000tcttcacagttggaattgggcatcccttttgtcagggagggagggagggagggaggctgg3060cttgttatgctgacaagtgtgattaaattcaaactttgaggtaagttggaggaacttgta3120cattgttaggagtgtgacaatttggactcttaatgatttggtcatacaaaatgaacctag3180accaacttctggaagatgtatataataactccatgttacattgatttcacctgactaata3240cttatcccttatcaattaaatacagaagatgccagccatctgggccttttaacccagaaa3300<210>3<211>182<212>dna<213>人工序列(artificialsequence)<400>3ttaattaagggatctgtagggcgcagtagtccagggtttccttgatgatgtcatacttat60cctgtcccttttttttccacagctcgcggttgaggacaaactcttcgcggtctttccagt120ggggatcgacggtatcgtagagtcgaggccgctctagaactagtggatccggaaccctta180at182<210>4<211>34<212>dna<213>人工序列(artificialsequence)<400>4gaagttcctattctctagaaagtataggaacttc34<210>5<211>720<212>dna<213>人工序列(artificialsequence)<400>5atggtgagcaagggcgaggagctgttcaccggggtggtgcccatcctggtcgagctggac60ggcgacgtaaacggccacaagttcagcgtgtccggcgagggcgagggcgatgccacctac120ggcaagctgaccctgaagttcatctgcaccaccggcaagctgcccgtgccctggcccacc180ctcgtgaccaccctgacctacggcgtgcagtgcttcagccgctaccccgaccacatgaag240cagcacgacttcttcaagtccgccatgcccgaaggctacgtccaggagcgcaccatcttc300ttcaaggacgacggcaactacaagacccgcgccgaggtgaagttcgagggcgacaccctg360gtgaaccgcatcgagctgaagggcatcgacttcaaggaggacggcaacatcctggggcac420aagctggagtacaactacaacagccacaacgtctatatcatggccgacaagcagaagaac480ggcatcaaggtgaacttcaagatccgccacaacatcgaggacggcagcgtgcagctcgcc540gaccactaccagcagaacacccccatcggcgacggccccgtgctgctgcccgacaaccac600tacctgagcacccagtccgccctgagcaaagaccccaacgagaagcgcgatcacatggtc660ctgctggagttcgtgaccgccgccgggatcactctcggcatggacgagctgtacaagtaa720<210>6<211>555<212>dna<213>人工序列(artificialsequence)<400>6tcgcgatgaataaatgaaagcttgcagatctgcgactctagaggatctgcgactctagag60gatcataatcagccataccacatttgtagaggttttacttgctttaaaaaacctcccaca120cctccccctgaacctgaaacataaaatgaatgcaattgttgttgttaacttgtttattgc180agcttataatggttacaaataaagcaatagcatcacaaatttcacaaataaagcattttt240ttcactgcattctagttgtggtttgtccaaactcatcaatgtatcttatcatgtctggat300ctgcgactctagaggatcataatcagccataccacatttgtagaggttttacttgcttta360aaaaacctcccacacctccccctgaacctgaaacataaaatgaatgcaattgttgttgtt420aacttgtttattgcagcttataatggttacaaataaagcaatagcatcacaaatttcaca480aataaagcatttttttcactgcattctagttgtggtttgtccaaactcatcaatgtatct540tatcatgtctggatc555<210>7<211>34<212>dna<213>人工序列(artificialsequence)<400>7ataacttcgtatagcatacattatacgaagttat34<210>8<211>1731<212>dna<213>人工序列(artificialsequence)<400>8cctcagcaccatggctagcggcagcctcggagtttgaatagatagaataaaatatcttta60ttttcattccatctgtgtgttggttttttgtgtgagatctacgggtggcatccctgtgac120ccctccccagtgcctctcctggccctggaagttgccactccagtgcccaccagccttgtc180ctaataaaattaagttgcatcattttgtctgactaggtgtccttctataatattatgggg240tggaggggggtggtatggagcaaggggcaagttgggaagacaacctgtagggcctgcggg300gtctattgggaaccaagctggagtgcagtggcacaatcttggctcactgcaatctccgcc360tcctgggttcaagcgattctcctgcctcaggctcccgagttgttgggattccaggcatgc420atgaccaggctcagctaatttttgtttttttggtagagacggggtttcaccatattggcc480aggctggtctccaactcctaatctcaggtgatctacccaccttggcctcccaaattgctg540ggattacaggcgtgaaccactgctcccttccctgtccttctgcctcagctaattgagtag600gggggaggctaactgaaacacggaaggagacaataccggaaggaacccgcgctatgacgg660caataaaaagacagaataaaacgcacgggtgttgggtcgtttgttcataaacgcggggtt720cggtcccagggctggcactctgtcgataccccaccgagaccccattggggccaatacgcc780cgcgtttcttccttttccccaccccaccccccaagttcgggtgaaggcccagggctcgca840gccaacgtcggggcggcaggccgtggaattcgtaaatgaattttctgtatgaggtcgcga900tgaataaatgaaagcttgcagatctgcgactctagaggatctgcgactctagaggatcat960aatcagccataccacatttgtagaggttttacttgctttaaaaaacctcccacacctccc1020cctgaacctgaaacataaaatgaatgcaattgttgttgttaacttgtttattgcagctta1080taatggttacaaataaagcaatagcatcacaaatttcacaaataaagcatttttttcact1140gcattctagttgtggtttgtccaaactcatcaatgtatcttatcatgtctggatctgcga1200ctctagaggatcataatcagccataccacatttgtagaggttttacttgctttaaaaaac1260ctcccacacctccccctgaacctgaaacataaaatgaatgcaattgttgttgttaacttg1320tttattgcagcttataatggttacaaataaagcaatagcatcacaaatttcacaaataaa1380gcatttttttcactgcattctagttgtggtttgtccaaactcatcaatgtatcttatcat1440gtctggatctgcgactctagaggatcataatcagccataccacatttgtagaggttttac1500ttgctttaaaaaacctcccacacctccccctgaacctgaaacataaaatgaatgcaattg1560ttgttgttaacttgtttattgcagcttataatggttacaaataaagcaatagcatcacaa1620atttcacaaataaagcatttttttcactgcattctagttgtggtttgtccaaactcatca1680atgtatcttatcatgtctggatccccatcaagctgatccggaacccttaat1731<210>9<211>1431<212>dna<213>人工序列(artificialsequence)<400>9atggtgagcaagggcgaggaggtcatcaaagagttcatgcgcttcaaggtgcgcatggag60ggctccatgaacggccacgagttcgagatcgagggcgagggcgagggccgcccctacgag120ggcacccagaccgccaagctgaaggtgaccaagggcggccccctgcccttcgcctgggac180atcctgtccccccagttcatgtacggctccaaggcgtacgtgaagcaccccgccgacatc240cccgattacaagaagctgtccttccccgagggcttcaagtgggagcgcgtgatgaacttc300gaggacggcggtctggtgaccgtgacccaggactcctccctgcaggacggcacgctgatc360tacaaggtgaagatgcgcggcaccaacttcccccccgacggccccgtaatgcagaagaag420accatgggctgggaggcctccaccgagcgcctgtacccccgcgacggcgtgctgaagggc480gagatccaccaggccctgaagctgaaggacggcggccactacctggtggagttcaagacc540atctacatggccaagaagcccgtgcaactgcccggctactactacgtggacaccaagctg600gacatcacctcccacaacgaggactacaccatcgtggaacagtacgagcgctccgagggc660cgccaccacctgttcctggggcatggcaccggcagcaccggcagcggcagctccggcacc720gcctcctccgaggacaacaacatggccgtcatcaaagagttcatgcgcttcaaggtgcgc780atggagggctccatgaacggccacgagttcgagatcgagggcgagggcgagggccgcccc840tacgagggcacccagaccgccaagctgaaggtgaccaagggcggccccctgcccttcgcc900tgggacatcctgtccccccagttcatgtacggctccaaggcgtacgtgaagcaccccgcc960gacatccccgattacaagaagctgtccttccccgagggcttcaagtgggagcgcgtgatg1020aacttcgaggacggcggtctggtgaccgtgacccaggactcctccctgcaggacggcacg1080ctgatctacaaggtgaagatgcgcggcaccaacttcccccccgacggccccgtaatgcag1140aagaagaccatgggctgggaggcctccaccgagcgcctgtacccccgcgacggcgtgctg1200aagggcgagatccaccaggccctgaagctgaaggacggcggccactacctggtggagttc1260aagaccatctacatggccaagaagcccgtgcaactgcccggctactactacgtggacacc1320aagctggacatcacctcccacaacgaggactacaccatcgtggaacagtacgagcgctcc1380gagggccgccaccacctgttcctgtacggcatggacgagctgtacaagtaa1431<210>10<211>585<212>dna<213>人工序列(artificialsequence)<400>10gcccctctccctcccccccccctaacgttactggccgaagccgcttggaataaggccggt60gtgcgtttgtctatatgttattttccaccatattgccgtcttttggcaatgtgagggccc120ggaaacctggccctgtcttcttgacgagcattcctaggggtctttcccctctcgccaaag180gaatgcaaggtctgttgaatgtcgtgaaggaagcagttcctctggaagcttcttgaagac240aaacaacgtctgtagcgaccctttgcaggcagcggaaccccccacctggcgacaggtgcc300tctgcggccaaaagccacgtgtataagatacacctgcaaaggcggcacaaccccagtgcc360acgttgtgagttggatagttgtggaaagagtcaaatggctctcctcaagcgtattcaaca420aggggctgaaggatgcccagaaggtaccccattgtatgggatctgatctggggcctcggt480gcacatgctttacatgtgtttagtcgaggttaaaaaaacgtctaggccccccgaaccacg540gggacgtggttttcctttgaaaaacacgatgataatatggccaca585<210>11<211>1658<212>dna<213>人工序列(artificialsequence)<400>11atggaagacgccaaaaacataaagaaaggcccggcgccattctatccgctggaagatgga60accgctggagagcaactgcataaggctatgaagagatacgccctggttcctggaacaatt120gcttttacagatgcacatatcgaggtggacatcacttacgctgagtacttcgaaatgtcc180gttcggttggcagaagctatgaaacgatatgggctgaatacaaatcacagaatcgtcgta240tgcagtgaaaactctcttcaattctttatgccggtgttgggcgcgttatttatcggagtt300gcagttgcgcccgcgaacgacatttataatgaacgtgaattgctcaacagtatgggcatt360tcgcagcctaccgtggtgttcgtttccaaaaaggggttgcaaaaaattttgaacgtgcaa420aaaaagctcccaatcatccaaaaaattattatcatggattctaaaacggattaccaggga480tttcagtcgatgtacacgttcgtcacatctcatctacctcccggttttaatgaatacgat540tttgtgccagagtccttcgatagggacaagacaattgcactgatcatgaactcctctgga600tctactggtctgcctaaaggtgtcgctctgcctcatagaactgcctgcgtgagattctcg660catgccagagatcctatttttggcaatcaaatcattccggatactgcgattttaagtgtt720gttccattccatcacggttttggaatgtttactacactcggatatttgatatgtggattt780cgagtcgtcttaatgtatagatttgaagaagagctgtttctgaggagccttcaggattac840aagattcaaagtgcgctgctggtgccaaccctattctccttcttcgccaaaagcactctg900attgacaaatacgatttatctaatttacacgaaattgcttctggtggcgctcccctctct960aaggaagtcggggaagcggttgccaagaggttccatctgccaggtatcaggcaaggatat1020gggctcactgagactacatcagctattctgattacacccgagggggatgataaaccgggc1080gcggtcggtaaagttgttccattttttgaagcgaaggttgtggatctggataccgggaaa1140acgctgggcgttaatcaaagaggcgaactgtgtgtgagaggtcctatgattatgtccggt1200tatgtaaacaatccggaagcgaccaacgccttgattgacaaggatggatggctacattct1260ggagacatagcttactgggacgaagacgaacacttcttcatcgttgaccgcctgaagtct1320ctgattaagtacaaaggctatcaggtggctcccgctgaattggaatccatcttgctccaa1380caccccaacatcttcgacgcaggtgtcgcaggtcttcccgacgatgacgccggtgaactt1440cccgccgccgttgttgttttggagcacggaaagacgatgacggaaaaagagatcgtggat1500tacgtcgccagtcaagtaacaaccgcgaaaaagttgcgcggaggagttgtgtttgtggac1560gaagtaccgaaaggtcttaccggaaaactcgacgcaagaaaaatcagagagatcctcata1620aaggccaagaagggcggaaagatcgccgtgtaactcga1658<210>12<211>828<212>dna<213>人工序列(artificialsequence)<400>12tcaacctctggattacaaaatttgtgaaagattgactggtattcttaactatgttgctcc60ttttacgctatgtggatacgctgctttaatgcctttgtatcatgctattgcttcccgtat120ggctttcattttctcctccttgtataaatcctggttgctgtctctttatgaggagttgtg180gcccgttgtcaggcaacgtggcgtggtgtgcactgtgtttgctgacgcaacccccactgg240ttggggcattgccaccacctgtcagctcctttccgggactttcgctttccccctccctat300tgccacggcggaactcatcgccgcctgccttgcccgctgctggacaggggctcggctgtt360gggcactgacaattccgtggtgttgtcggggaaatcatcgtcctttccttggctgctcgc420ctgtgttgccacctggattctgcgcgggacgtccttctgctacgtcccttcggccctcaa480tccagcggaccttccttcccgcggcctgctgccggctctgcggcctcttccgcgtcttcg540ccttcgccctcagacgagtcggatctccctttgggccgcctccccgcatcgataccgtcg600atcctgtgccttctagttgccagccatctgttgtttgcccctcccccgtgccttccttga660ccctggaaggtgccactcccactgtcctttcctaataaaatgaggaaattgcatcgcatt720gtctgagtaggtgtcattctattctggggggtggggtggggcaggacagcaagggggagg780attgggaagacaatagcaggcatgctggggatgcggtgggctctatgg828<210>13<211>1638<212>dna<213>人工序列(artificialsequence)<400>13ggtctgaagaggagtttacgtccagccaagctagcttggctgcaggtcgtcgaaattcta60ccgggtaggggaggcgcttttcccaaggcagtctggagcatgcgctttagcagccccgct120gggcacttggcgctacacaagtggcctctggcctcgcacacattccacatccaccggtag180gcgccaaccggctccgttctttggtggccccttcgcgccaccttctactcctcccctagt240caggaagttcccccccgccccgcagctcgcgtcgtgcaggacgtgacaaatggaagtagc300acgtctcactagtctcgtgcagatggacagcaccgctgagcaatggaagcgggtaggcct360ttggggcagcggccaatagcagctttgctccttcgctttctgggctcagaggctgggaag420gggtgggtccgggggcgggctcaggggcgggctcaggggcggggcgggcgcccgaaggtc480ctccggaggcccggcattctgcacgcttcaaaagcgcacgtctgccgcgctgttctcctc540ttcctcatctccgggcctttcgacctgcagcctgttgacaattaatcatcggcatagtat600atcggcatagtataatacgacaaggtgaggaactaaaccatgggatcggccattgaacaa660gatggattgcacgcaggttctccggccgcttgggtggagaggctattcggctatgactgg720gcacaacagacaatcggctgctctgatgccgccgtgttccggctgtcagcgcaggggcgc780ccggttctttttgtcaagaccgacctgtccggtgccctgaatgaactgcaggacgaggca840gcgcggctatcgtggctggccacgacgggcgttccttgcgcagctgtgctcgacgttgtc900actgaagcgggaagggactggctgctattgggcgaagtgccggggcaggatctcctgtca960tctcaccttgctcctgccgagaaagtatccatcatggctgatgcaatgcggcggctgcat1020acgcttgatccggctacctgcccattcgaccaccaagcgaaacatcgcatcgagcgagca1080cgtactcggatggaagccggtcttgtcgatcaggatgatctggacgaagagcatcagggg1140ctcgcgccagccgaactgttcgccaggctcaaggcgcgcatgcccgacggcgatgatctc1200gtcgtgacccatggcgatgcctgcttgccgaatatcatggtggaaaatggccgcttttct1260ggattcatcgactgtggccggctgggtgtggcggaccgctatcaggacatagcgttggct1320acccgtgatattgctgaagagcttggcggcgaatgggctgaccgcttcctcgtgctttac1380ggtatcgccgctcccgattcgcagcgcatcgccttctatcgccttcttgacgagttcttc1440tgaggggatcaattctctagagctcgctgatcagcctcgactgtgccttctagttgccag1500ccatctgttgtttgcccctcccccgtgccttccttgaccctggaaggtgccactcccact1560gtcctttcctaataaaatgaggaaattgcatcgcattgtctgagtaggtgtcattctatt1620ctggggggtggggtgggg1638<210>14<211>1598<212>dna<213>人工序列(artificialsequence)<400>14acataacttacggtaaatggcccgcctggctgaccgcccaacgacccccgcccattgacg60tcaataatgacgtatgttcccatagtaacgccaatagggactttccattgacgtcaatgg120gtggagtatttacggtaaactgcccacttggcagtacatcaagtgtatcatatgccaagt180acgccccctattgacgtcaatgacggtaaatggcccgcctggcattatgcccagtacatg240accttatgggactttcctacttggcagtacatctacgtattagtcatcgctattaccatg300gtcgaggtgagccccacgttctgcttcactctccccatctcccccccctccccaccccca360attttgtatttatttattttttaattattttgtgcagcgatgggggcggggggggggggg420gggcgcgcgccaggcggggcggggcggggcgaggggcggggcggggcgaggcggagaggt480gcggcggcagccaatcagagcggcgcgctccgaaagtttccttttatggcgaggcggcgg540cggcggcggccctataaaaagcgaagcgcgcggcgggcggggagtcgctgcgacgctgcc600ttcgccccgtgccccgctccgccgccgcctcgcgccgcccgccccggctctgactgaccg660cgttactcccacaggtgagcgggcgggacggcccttctcctccgggctgtaattagcgct720tggtttaatgacggcttgtttcttttctgtggctgcgtgaaagccttgaggggctccggg780agggccctttgtgcggggggagcggctcggggggtgcgtgcgtgtgtgtgtgcgtgggga840gcgccgcgtgcggctccgcgctgcccggcggctgtgagcgctgcgggcgcggcgcggggc900tttgtgcgctccgcagtgtgcgcgaggggagcgcggccgggggcggtgccccgcggtgcg960gggggggctgcgaggggaacaaaggctgcgtgcggggtgtgtgcgtgggggggtgagcag1020ggggtgtgggcgcgtcggtcgggctgcaaccccccctgcacccccctccccgagttgctg1080agcacggcccggcttcgggtgcggggctccgtacggggcgtggcgcggggctcgccgtgc1140cgggcggggggtggcggcaggtgggggtgccgggcggggcggggccgcctcgggccgggg1200agggctcgggggaggggcgcggcggcccccggagcgccggcggctgtcgaggcgcggcga1260gccgcagccattgccttttatggtaatcgtgcgagagggcgcagggacttcctttgtccc1320aaatctgtgcggagccgaaatctgggaggcgccgccgcaccccctctagcgggcgcgggg1380cgaagcggtgcggcgccggcaggaaggaaatgggcggggagggccttcgtgcgtcgccgc1440gccgccgtccccttctccctctccagcctcggggctgtccgcggggggacggctgccttc1500gggggggacggggcagggcggggttcggcttctggcgtgtgaccggcggctctagagcct1560ctgctaaccatgttcatgccttcttctttttcctacag1598<210>15<211>5869<212>dna<213>人工序列(artificialsequence)<400>15gatcctctagagtcgagcagtgtggttttcaagaggaagcaaaaagcctctccacccagg60cctggaatgtttccacccaatgtcgagcagtgtggttttgcaagaggaagcaaaaagcct120ctccacccaggcctggaatgtttccacccaatgtcgagcaaaccccgcccagcgtcttgt180cattggcgaattcgaacacgcagatgcagtcggggcggcgcggtcccaggtccacttcgc240atattaaggtgacgcgtgtggcctcgaacaccgagcgaccctgcagcgacccgcttaaca300gcgtcaacagcgtgccgcaga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