CN115247186A - 一种构建af双基因突变的动脉粥样硬化模型猪核移植供体细胞的基因编辑系统及其应用 - Google Patents
一种构建af双基因突变的动脉粥样硬化模型猪核移植供体细胞的基因编辑系统及其应用 Download PDFInfo
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Abstract
本发明公开了一种构建AF双基因突变的动脉粥样硬化模型猪核移植供体细胞的基因编辑系统及其应用。本发明提供了的包括质粒pKG‑U6gRNA(APOE‑E2‑gRNA2)、靶序列结合区如SEQ ID NO：25中第3‑22位核苷酸所示的FBN1‑gRNA4、靶序列结合区如SEQ ID NO：26中第3‑22位核苷酸所示的FBN1‑gRNA6和SEQ ID NO：27所示的FBN1‑mutant‑ss163。所述试剂盒的用途：制备重组细胞；制备动脉粥样硬化模型猪；制备动脉粥样硬化细胞模型或动脉粥样硬化组织模型或动脉粥样硬化器官模型。质粒pKG‑U6gRNA(APOE‑E2‑gRNA2)，其转录得到靶序列结合区如SEQ ID NO：11中第1‑20位核苷酸所示的sgRNAAPOE‑E2‑gRNA2。本发明为通过体细胞核移植动物克隆技术培育动脉粥样硬化疾病模型猪奠定了基础。
Description
技术领域
本发明属于生物技术领域，具体属于基因编辑技术领域，更具体涉及一种构建AF双基因突变的动脉粥样硬化模型猪核移植供体细胞的基因编辑系统及其应用。
背景技术
心血管疾病是目前我国居民所有疾病中的致死首因，据报道，心血管疾病死亡占居民疾病死亡构成的40％以上，远高于癌症及其他疾病。动脉粥样硬化(Atherosclerosis)是冠心病、脑梗死、外周血管病的主要原因。脂质代谢异常和炎性反应为动脉粥样硬化的主要病变基础。
载脂蛋白E(APOE)是一个主要的乳糜微粒的辅基蛋白，可以与肝脏细胞或者周围细胞上的受体结合。APOE基因的缺陷可能导致由于乳糜微粒和极低密度脂蛋白无法被正常清除而引发血清胆固醇和甘油三酯升高。原纤蛋白1(Fibrillin-1)是微纤维的主要结构成分，可为弹性蛋白的沉积和交联提供支架，由FBN1基因编码。在小鼠模型研究中发现该蛋白C1039G突变会导致动脉硬化增加，进而加快动脉粥样硬化疾病的发展进程，经序列比对发现突变区域在物种间高度保守。APOE和FBN1基因的异常已被证实与人动脉粥样硬化的发生发展密切相关，因此，迫切需要开发出基于APOE和FBN1联合突变导致的动脉粥样硬化动物模型以尽快解开动脉粥样硬化疾病进展机制及疾病进程谜团，并为进一步的治疗奠定基础。
目前，已建立了APOE敲除联合FBN1点突变的小鼠模型，虽可准确模拟人类动脉粥样硬化疾病前期高度不稳定的斑块的发展进程，但是小鼠不论从体型、器官大小、生理、病理等方面都与人相差巨大，不能真实地模拟人类正常的生理、病理状态。而猪作为大动物，是人类长期以来主要的肉食供应动物，其体型大小和生理功能与人类近似，易于大规模繁殖饲养，而且在伦理道德及动物保护等方面要求较低，是理想的人类疾病模型动物。
基因编辑是近年来不断取得重大发展的一种生物技术，其包括从基于同源重组的基因编辑到基于核酸酶的ZFN、TALEN、CRISPR/Cas9等编辑技术，其中CRISPR/Cas9技术是当前最先进的基因编辑技术。目前，基因编辑技术被越来越多地应用到动物模型的制作上。
同源重组(HDR)是通过序列同源性交换DNA序列信息：即修复模板中包含所需插入片段，修复模板的两端则是与插入位点附近具有序列同源性的重组臂。过去通常使用双链DNA(dsDNA)作为修复模板，但最近的研究揭示了单链寡核苷酸脱氧核苷酸(ssODN)作为HDR供体模板的优越性。首先，ssODN作为供体模板比dsDNA模板的插入位点特异性高，dsDNA模板容易产生随机插入。其次，ssODN对同源重组臂的长度要求比dsDNA模板更短，单侧30-60个碱基的重组臂设计可以获得高效且稳定的HDR，相比类似的dsDNA模板，其提供的插入效率更高。第三，dsDNA容易被NHEJ修复途径合并，从而导致同源臂的复制或者dsDNA模板的部分整合，而ssODN就不易产生这种现象。另外，dsDNAs对培养的细胞是有害的，线型或者质粒dsDNAs的转染效率较低，并使细胞产生不良反应，而ssODN模板在这些方面就更有优势。
发明内容
本发明的目的是提供一种构建AF双基因突变的动脉粥样硬化模型猪核移植供体细胞的基因编辑系统及其应用。
本发明提供了一种试剂盒，包括质粒pKG-U6gRNA(APOE-E2-gRNA2)、FBN1-gRNA4、FBN1-gRNA6和FBN1-mutant-ss163。
所述试剂盒还包括表达Cas9蛋白的质粒。
所述试剂盒还包括NCN蛋白。
所述试剂盒还包括PRONCN蛋白。
所述试剂盒还包括质粒pKG-GE4。
所述试剂盒还包括猪细胞。
所述试剂盒的用途为如下(a)或(b)或(c)：(a)制备重组细胞；(b)制备动脉粥样硬化模型猪；(c)制备动脉粥样硬化细胞模型或动脉粥样硬化组织模型或动脉粥样硬化器官模型。
本发明还提供了一种制备重组细胞的方法，包括如下步骤：将质粒pKG-U6gRNA(APOE-E2-gRNA2)、表达Cas9蛋白的质粒、FBN1-gRNA4、FBN1-gRNA6、FBN1-mutant-ss163和NCN蛋白共转染猪细胞，得到APOE基因发生突变且FBN1基因发生目标突变的重组细胞；
FBN1基因发生目标突变指的是：用SEQ ID NO：27所示的DNA分子取代猪细胞的染色体DNA中SEQ ID NO：28所示的DNA分子，得到重组细胞。
质粒pKG-U6gRNA(APOE-E2-gRNA2)、表达Cas9蛋白的质粒、FBN1-gRNA4、FBN1-gRNA6、FBN1-mutant-ss163和NCN蛋白的配比依次为：0.8-1.0μg质粒pKG-U6gRNA(APOE-E2-gRNA2)：0.9-1.2μg表达Cas9蛋白的质粒：0.8-1.2μg FBN1-gRNA4：0.8-1.2μg FBN1-gRNA6：1.8-2.2μg FBN1-mutant-ss163：3-5μg NCN蛋白。
质粒pKG-U6gRNA(APOE-E2-gRNA2)、表达Cas9蛋白的质粒、FBN1-gRNA4、FBN1-gRNA6、FBN1-mutant-ss163和NCN蛋白的配比依次为：0.92μg质粒pKG-U6gRNA(APOE-E2-gRNA2)：1.08μg表达Cas9蛋白的质粒：1μg FBN1-gRNA4：1μg FBN1-gRNA6：2μg FBN1-mutant-ss163：4μg NCN蛋白。
猪细胞、质粒pKG-U6gRNA(APOE-E2-gRNA2)、表达Cas9蛋白的质粒、FBN1-gRNA4、FBN1-gRNA6、FBN1-mutant-ss163和NCN蛋白的配比依次为：20万个猪细胞：0.8-1.0μg质粒pKG-U6gRNA(APOE-E2-gRNA2)：0.9-1.2μg表达Cas9蛋白的质粒：0.8-1.2μg FBN1-gRNA4：0.8-1.2μg FBN1-gRNA6：1.8-2.2μg FBN1-mutant-ss163：3-5μg NCN蛋白。
猪细胞、质粒pKG-U6gRNA(APOE-E2-gRNA2)、表达Cas9蛋白的质粒、FBN1-gRNA4、FBN1-gRNA6、FBN1-mutant-ss163和NCN蛋白的配比依次为：20万个猪细胞：0.92μg质粒pKG-U6gRNA(APOE-E2-gRNA2)：1.08μg表达Cas9蛋白的质粒：1μg FBN1-gRNA4：1μgFBN1-gRNA6：2μg FBN1-mutant-ss163：4μg NCN蛋白。
质粒pKG-U6gRNA(APOE-E2-gRNA2)，其转录得到sgRNAAPOE-E2-gRNA2；sgRNAAPOE-E2-gRNA2，为sgRNA，其靶序列结合区如SEQ ID NO：11中第1-20位核苷酸所示。
具体的，所述sgRNAAPOE-E2-gRNA2如SEQ ID NO：11所示。
FBN1-gRNA4，为sgRNA，其靶序列结合区如SEQ ID NO：25中第3-22位核苷酸所示。
具体的，所述FBN1-gRNA4如SEQ ID NO：25所示。
具体的，所述FBN1-gRNA4如SEQ ID NO：19所示。
FBN1-gRNA6，为sgRNA，其靶序列结合区如SEQ ID NO：26中第3-22位核苷酸所示。
具体的，所述FBN1-gRNA6如SEQ ID NO：26所示。
具体的，所述FBN1-gRNA6如SEQ ID NO：21所示。
FBN1-mutant-ss163为SEQ ID NO：27所示的单链DNA分子。
所述NCN蛋白为Cas9蛋白或具有Cas9蛋白的融合蛋白。
具体的，所述NCN蛋白如SEQ ID NO：3所示。
以上任一所述猪细胞为猪成纤维细胞。
以上任一所述猪细胞为猪原代成纤维细胞。
所述NCN蛋白的制备方法包括如下步骤：
(1)将质粒pKG-GE4导入大肠杆菌BL21(DE3)，得到重组菌；
(2)采用液体培养基30℃培养所述重组菌，然后加入IPTG并进行25℃诱导培养，然后收集菌体；
(3)将收集的菌体进行菌体破碎，收集粗蛋白溶液；
(4)采用亲和层析从所述粗蛋白溶液中纯化具有His6标签的融合蛋白；
(5)采用具有His6标签的肠激酶酶切具有His6标签的融合蛋白，然后采用Ni-NTA树脂去除具有His6标签的蛋白，得到纯化的NCN蛋白；
质粒pKG-GE4中具有SEQ ID NO：1中第5209-9852位核苷酸所示的融合基因。
所述NCN蛋白的制备方法具体包括如下步骤：
(1)将质粒pKG-GE4导入大肠杆菌BL21(DE3)，得到重组菌。
(2)将步骤(1)得到的重组菌接种至含氨苄青霉素的液体LB培养基，振荡培养；
(3)将步骤(2)得到的菌液接种至液体LB培养基，30℃、230rpm振荡培养至OD600nm值＝1.0，然后加入IPTG并使其在体系中的浓度为0.5mM，然后25℃、230rpm振荡培养12小时，然后离心收集菌体；
(4)取步骤(3)得到的菌体，用PBS缓冲液洗涤；
(5)取步骤(4)得到的菌体，加入粗提缓冲液并悬浮菌体，然后进行菌体破碎，然后离心收集上清液，采用0.22μm孔径滤膜过滤，收集滤液；
(6)采用亲和层析从步骤(5)得到的滤液中纯化具有His6标签的融合蛋白(SEQ IDNO：2所示的融合蛋白)；
(7)取步骤(6)收集的过柱后溶液，使用超滤管浓缩，然后用25mM Tris-HCl(pH8.0)稀释；
(8)将具有His6标签的重组牛肠激酶加入到步骤(7)得到的溶液中，酶切；
(9)将完成步骤(8)的溶液与Ni-NTA树脂混匀，孵育，然后离心收集上清液；
(10)取步骤(9)得到的上清液，使用超滤管浓缩，然后加入酶贮存液中，即为NCN蛋白溶液。
采用亲和层析从步骤(5)得到的滤液中纯化具有His6标签的融合蛋白的具体方法如下：
首先采用5个柱体积的平衡液平衡Ni-NTA琼脂糖柱(流速为1ml/min)；然后上样50ml步骤(5)得到的滤液(流速为0.5-1ml/min)；然后用5个柱体积的平衡液洗涤柱子(流速为1ml/min)；然后用5个柱体积的缓冲液洗涤柱子(流速为1ml/min)，以去除杂蛋白；然后用10个柱体积的洗脱液以0.5-1ml/min的流速洗脱，收集过柱后溶液(90-100ml)。
所述PRONCN蛋白自上游至下游依次包括如下元件：信号肽、分子伴侣蛋白、蛋白标签、蛋白酶酶切位点、核定位信号、Cas9蛋白、核定位信号。
所述信号肽的功能为促进蛋白分泌表达。所述信号肽可选自大肠杆菌碱性磷酸酶(phoA)信号肽、金黄色葡萄球菌蛋白A信号肽、大肠杆菌外膜蛋白(ompa)信号肽或任何其他原核基因的信号肽，优选为碱性磷酸酶信号肽(phoA signal peptide)。碱性磷酸酶信号肽用来引导目的蛋白分泌表达至细菌周质腔中，从而与细菌胞内蛋白分离，且分泌到细菌周质腔中的目的蛋白为可溶性表达，可被细菌周质腔中的信号肽酶裂解。
所述分子伴侣蛋白的功能为增加蛋白的可溶性。所述分子伴侣可为任何帮助形成二硫键的蛋白，优选为硫氧还原蛋白(TrxA蛋白)。硫氧还原蛋白，其能作为分子伴侣帮助所共表达的目的蛋白(例如Cas9蛋白)形成二硫键，提高蛋白的稳定性、折叠的正确性，增加目的蛋白的溶解性及活性。
所述蛋白标签的功能为用于蛋白纯化。所述标签可为His标签(His-Tag，His6蛋白标签)、GST标签、Flag标签、HA标签、c-Myc标签或其他任何蛋白标签，进一步优选为His标签。His标签能与Ni柱结合，可以通过一步法Ni柱亲和层析纯化目的蛋白，可极大地简化目的蛋白的纯化流程。
所述蛋白酶酶切位点的功能为纯化后用于切除非功能区段，以释放天然形式Cas9蛋白。所述蛋白酶可选自肠激酶(Enterokinase)、因子Xa(Factor Xa)、凝血酶(Thrombin)、TEV蛋白酶(TEV protease)、HRV 3C蛋白酶(HRV 3C protease)、WELQut蛋白酶或任何其他内切蛋白酶，进一步优选为肠激酶。EK为肠激酶酶切位点，便于使用肠激酶切除所融合的TrxA-His区段，得到天然形式的Cas9蛋白。本申请使用带His标签的商品肠激酶酶切融合蛋白后，可通过一次亲和层析除去TrxA-His区段及带His标签的肠激酶，得到天然形式的Cas9蛋白，避免了多次纯化透析对目的蛋白的伤害和损耗。
所述核定位信号可为任何核定位信号，优选为SV40核定位信号和/或nucleoplasmin核定位信号。NLS为核定位信号，在Cas9的N端及C端分别设计了一个NLS位点，使Cas9能更有效地进入细胞核进行基因编辑。
所述Cas9蛋白可为saCas9或spCas9,优选为spCas9蛋白。
PRONCN蛋白具体如SEQ ID NO：2所示。
所述质粒pKG-GE4自上游至下游依次包括如下元件：启动子、操纵子、核糖体结合位点、PRONCN蛋白的编码基因、终止子。
所述启动子具体可为T7启动子。T7启动子为原核表达强启动子，能高效驱动外源基因的表达。
所述操纵子具体可为Lac操纵子。Lac操纵子为乳糖诱导表达的调控元件，可在细菌生长至一定数量后，再用IPTG在低温下诱导目的蛋白的表达，可避免目的蛋白过早表达对宿主菌生长的影响，低温下诱导表达也显著提高所表达的目的蛋白的可溶性。
所述核糖体结合位点是蛋白翻译时的核糖体结合位点，对蛋白质的翻译是必要的。
所述终止子具体可为T7终止子。T7终止子可在目的基因的末端有效终止基因转录，避免目的基因之外的其他下游序列得到转录和翻译。
对于spCas9蛋白的密码子，本申请对其密码子进行了优化，使之完全适应本申请所选用的大肠杆菌高效表达菌株E.coli BL21(DE3)的密码子偏好，从而提高Cas9蛋白的表达水平。
T7启动子如SEQ ID NO：1中第5121-5139位核苷酸所示。
Lac操纵子如SEQ ID NO：1中第5140-5164位核苷酸所示。
核糖体结合位点如SEQ ID NO：1中第5178-5201位核苷酸所示。
碱性磷酸酶信号肽的编码序列如SEQ ID NO：1中第5209-5271位核苷酸所示。
TrxA蛋白的编码序列如SEQ ID NO：1中第5272-5598位核苷酸所示。
His-Tag的编码序列如SEQ ID NO：1中第5620-5637位核苷酸所示。
肠激酶酶切位点的编码序列如SEQ ID NO：1中第5638-5652位核苷酸所示。
核定位信号的编码序列如SEQ ID NO：1中第5656-5670位核苷酸所示。
spCas9蛋白的编码序列如SEQ ID NO：1中第5701-9801位核苷酸所示。
核定位信号的编码序列如SEQ ID NO：1中第9802-9849位核苷酸所示。
T7终止子如SEQ ID NO：1中第9902-9949位核苷酸。
质粒pKG-GE4中具有SEQ ID NO：1中第5121-9949位核苷酸所示的DNA分子。
具体的，质粒pKG-GE4如SEQ ID NO：1所示。
表达Cas9蛋白的质粒具体可为质粒pKG-GE3。
质粒pKG-GE3中，具有特异融合基因；所述特异融合基因编码特异融合蛋白；
所述特异融合蛋白自N端至C端依次包括如下元件：两个核定位信号(NLS)、Cas9蛋白、两个核定位信号、自剪切多肽P2A、荧光报告蛋白、自裂解多肽T2A、抗性筛选标记蛋白；
质粒pKG-GE3中，由EF1a启动子启动所述特异融合基因的表达；
质粒pKG-GE3中，所述特异融合基因下游具有WPRE序列元件、3’LTR序列元件和bGHpoly(A)signal序列元件。
质粒pKG-GE3中，依次具有如下元件：CMV增强子、EF1a启动子、所述特异融合基因、WPRE序列元件、3’LTR序列元件、bGH poly(A)signal序列元件。
所述特异融合蛋白中，Cas9蛋白上游的两个核定位信号为SV40核定位信号，Cas9蛋白下游的两个核定位信号为nucleoplasmin核定位信号。
所述特异融合蛋白中，荧光报告蛋白具体可为EGFP蛋白。
所述特异融合蛋白中，抗性筛选标记蛋白具体可为Puromycin抗性蛋白。
自剪切多肽P2A的氨基酸序列为“ATNFSLLKQAGDVEENPGP”(发生自剪切的断裂位置为C端开始第一个氨基酸残基和第二个氨基酸残基之间)。
自裂解多肽T2A的氨基酸序列为“EGRGSLLTCGDVEENPGP”(发生自裂解的断裂位置为C端开始第一个氨基酸残基和第二个氨基酸残基之间)。
特异融合基因具体如SEQ ID NO：29中第911-6706位核苷酸所示。
CMV增强子如SEQ ID NO：29中第395-680位核苷酸所示。
EF1a启动子如SEQ ID NO：29中第682-890位核苷酸所示。
WPRE序列元件如SEQ ID NO：29中第6722-7310位核苷酸所示。
3’LTR序列元件如SEQ ID NO：29中第7382-7615位核苷酸所示。
bGH poly(A)signal序列元件如SEQ ID NO：29中第7647-7871位核苷酸所示。
质粒pKG-GE3具体如SEQ ID NO：29所示。
质粒pKG-U6gRNA中，具有SEQ ID NO：30中第2280-2637位核苷酸所示的DNA分子。
质粒pKG-U6gRNA具体如SEQ ID NO：30所示。
具体来说，所述质粒pKG-U6gRNA(APOE-E2-gRNA2)是借助限制性内切酶BbsI将sgRNAAPOE-E2-gRNA2的靶序列结合区的编码序列插入pKG-U6gRNA载体得到的。
本发明还保护以上任一所述方法制备得到的重组细胞。
本发明还保护所述重组细胞在制备动脉粥样硬化模型猪中的应用。
将所述重组细胞作为核移植供体细胞进行体细胞克隆，可以得到克隆猪，即为动脉粥样硬化模型猪。
本发明还保护利用所述重组细胞制备的模型猪的猪组织，即动脉粥样硬化组织模型。
本发明还保护利用所述重组细胞制备的模型猪的猪器官，即动脉粥样硬化器官模型。
本发明还保护利用所述重组细胞制备的模型猪的猪细胞，即动脉粥样硬化细胞模型。
本发明还保护所述重组细胞、所述动脉粥样硬化组织模型、所述动脉粥样硬化器官模型、所述动脉粥样硬化细胞模型或者所述动脉粥样硬化模型猪的应用，为如下(d1)或(d2)或(d3)或(d4)：
(d1)筛选治疗动脉粥样硬化的药物；
(d2)进行动脉粥样硬化药物的药效评价；
(d3)进行动脉粥样硬化的基因治疗和/或细胞治疗的疗效评价；
(d4)研究动脉粥样硬化的发病机制。
以上任一所述猪具体可为从江香猪。
猪APOE基因信息：编码Apolipoprotein E蛋白；位于6号染色体；GeneID为397576，Sus scrofa。猪APOE基因编码的蛋白质如SEQ ID NO：8所示。猪APOE基因具有SEQ ID NO：9所示的DNA区段。
猪FBN1基因信息：编码原纤蛋白1(Fibrillin-1)；位于猪1号染色体；GeneID为414836，Sus scrofa。猪FBN1基因编码的蛋白质如SEQ ID NO：14所示。猪FBN1基因具有SEQID NO：15所示的DNA区段。
大小鼠等啮齿类动物不论从体型、器官大小、生理、病理等方面都与人相差巨大，无法真实地模拟人类正常的生理、病理状态。研究表明，95％以上在大小鼠中验证有效的药物在人类临床试验中是无效的。就大动物而言，灵长类是与人亲缘关系最近的动物，但其体型小、性成熟晚(6-7岁开始交配)，且为单胎动物，群体扩繁速度极慢，饲养成本很高。另外，灵长类动物克隆效率低、难度大、成本高。而猪作为模型动物就没有上述缺点，猪是除灵长类外与人亲缘关系最近的动物，其体型、体重、器官大小等与人相近，在解剖学、生理学、免疫学、营养代谢、疾病发病机制等方面与人类极为相似。同时，猪的性成熟早(4-6个月)，繁殖力高，一胎多仔，在2-3年内即可形成一个较大群体。另外，猪的克隆技术非常成熟，克隆及饲养成本也较灵长类低得多。因此猪是非常适合作为人类疾病模型的动物。
本发明采用CRISPR/Cas9技术联合ssODN同源重组技术进行了FBN1基因的点突变基因编辑，同时对APOE基因进行了编辑，获得了APOE基因敲除和FBN1基因精确点突变的单细胞克隆，为后期通过体细胞核移植动物克隆技术培育动脉粥样硬化疾病模型猪奠定了基础。该模型猪预期可模拟动脉粥样硬化疾病的人类疾病进程特征，将为研究动脉粥样硬化的发病机制、疾病进程及药物研发提供有力的实验工具。
附图说明
图1为质粒pET-32a的结构示意图。
图2为质粒pKG-GE4的结构示意图。
图3为实施例3中gRNA与NCN蛋白用量配比优化的电泳图。
图4为实施例3中NCN蛋白与商品Cas9蛋白的基因编辑效率比较的电泳图。
图5为实施例4中分别以18只猪的基因组DNA为模板采用APOE-E2-F和APOE-E2-R组成的引物对进行PCR扩增的电泳图。
图6为实施例4中不同靶点的测序峰图。
图7为实施例5中用命名为1的猪的耳组织提取基因组作为模板采用不同引物对进行PCR扩增的电泳图。
图8为实施例5中分别以18只猪的基因组DNA为模板采用FBN1-JDF419和FBN1-JDR673组成的引物对进行PCR扩增的电泳图。
图9为实施例5中不同靶点的编辑效率比较的电泳图。
图10是实施例6中编号为2的单细胞克隆的反向测序与APOE基因靶标位点野生型序列的比对结果。
图11是实施例6中编号为3的单细胞克隆的反向测序与APOE基因靶标位点野生型序列的比对结果。
图12是实施例6中编号为10的单细胞克隆的反向测序与APOE基因靶标位点野生型序列的比对结果。
图13是实施例6中编号为8的单细胞克隆的正向和反向测序同时与APOE基因靶标位点野生型序列的比对结果。
图14是实施例6中编号为3的单细胞克隆的反向测序与FBN1基因靶标位点野生型序列的比对结果。
图15是实施例6中编号为1的单细胞克隆的反向测序与FBN1基因靶标位点野生型序列的比对结果。
图16是实施例6中编号为2的单细胞克隆的反向测序与FBN1基因靶标位点野生型序列的比对结果。
图17是实施例6中编号为11的单细胞克隆的反向测序与FBN1基因靶标位点野生型序列的比对结果。
图18是实施例6中编号为10的单细胞克隆的反向测序与FBN1基因靶标位点野生型序列的比对结果。
图19是实施例6中编号为15的单细胞克隆的反向测序与FBN1基因靶标位点野生型序列的比对结果。
具体实施方式
下面结合具体实施方式对本发明进行进一步的详细描述，给出的实施例仅为了阐明本发明，而不是为了限制本发明的范围。以下提供的实施例可作为本技术领域普通技术人员进行进一步改进的指南，并不以任何方式构成对本发明的限制。
下述实施例中的实验方法，如无特殊说明，均为常规方法，按照本领域内的文献所描述的技术或条件或者按照产品说明书进行。下述实施例中所用的材料、试剂等，如无特殊说明，均可从商业途径得到。实施例中构建的重组质粒，均已进行测序验证。商品Cas9-A蛋白为市售的效果好的Cas9蛋白。商品Cas9-B蛋白为市售的效果好的Cas9蛋白。完全培养液(％为体积比)：15％胎牛血清(Gibco)+83％DMEM培养基(Gibco)+1％Penicillin-Streptomycin(Gibco)+1％HEPES(Solarbio)。细胞培养条件：37℃，5％CO2、5％O2的恒温培养箱。
实施例中采用的猪原代成纤维细胞均是用初生从江香猪耳组织制备得到的。制备猪原代成纤维细胞的方法：①取猪耳组织0.5g，去除毛发及骨组织，然后用75％酒精浸泡30-40s，然后用含5％(体积比)Penicillin-Streptomycin(Gibco)的PBS缓冲液洗涤5次，然后用PBS缓冲液洗涤一次；②用剪刀将组织剪碎，采用5mL 0.1％胶原酶溶液(Sigma)，37℃消化1h，然后500g离心5min，弃上清；③将沉淀用1mL完全培养液重悬，然后铺入含10mL完全培养液并已用0.2％明胶(VWR)封盘的直径为10cm的细胞培养皿中，培养至细胞长满皿底60％左右；④完成步骤③后，采用胰蛋白酶消化并收集细胞，然后重悬于完全培养液。用于进行后续电转实验。
实施例1、质粒的构建
一、质粒pKG-GE3的构建
质粒pKG-GE3，为环形质粒，如SEQ ID NO：29所示。SEQ ID NO：29中，第395-680位核苷酸组成CMV增强子，第682-890位核苷酸组成EF1a启动子，第986-1006位核苷酸编码核定位信号(NLS)，第1016-1036位核苷酸编码核定位信号(NLS)，第1037-5161位核苷酸编码Cas9蛋白，第5162-5209位核苷酸编码核定位信号(NLS)，第5219-5266位核苷酸编码核定位信号(NLS)，第5276-5332位核苷酸编码自剪切多肽P2A(自剪切多肽P2A的氨基酸序列为“ATNFSLLKQAGDVEENPGP”，发生自剪切的断裂位置为C端开始第一个氨基酸残基和第二个氨基酸残基之间)，第5333-6046位核苷酸编码EGFP蛋白，第6056-6109位核苷酸编码自裂解多肽T2A(自裂解多肽T2A的氨基酸序列为“EGRGSLLTCGDVEENPGP”，发生自裂解的断裂位置为C端开始第一个氨基酸残基和第二个氨基酸残基之间)，第6110-6703位核苷酸编码Puromycin蛋白(简称Puro蛋白)，第6722-7310位核苷酸组成WPRE序列元件，第7382-7615位核苷酸组成3’LTR序列元件，第7647-7871位核苷酸组成bGH poly(A)signal序列元件。SEQID NO：29中，第911-6706位核苷酸形成融合基因，表达融合蛋白。由于自剪切多肽P2A和自裂解多肽T2A的存在，融合蛋白自发形成如下三个蛋白：具有Cas9蛋白的蛋白、具有EGFP蛋白的蛋白和具有Puro蛋白的蛋白。
二、质粒pKG-U6gRNA的构建
pKG-U6gRNA载体即质粒pKG-U6gRNA，为环形质粒，如SEQ ID NO：30所示。SEQ IDNO：30中，第2280-2539位核苷酸组成hU6启动子，第2558-2637位核苷酸用于转录形成gRNA骨架。使用时，将20bp左右的DNA分子(用于转录形成gRNA的靶序列结合区)插入质粒pKG-U6gRNA，形成重组质粒，在细胞中重组质粒转录得到gRNA。
三、原核Cas9高效表达载体的构建
质粒pET-32a的结构示意图见图1。
质粒pKG-GE4是以质粒pET-32a为出发质粒进行改造得到的。质粒pET32a-T7lac-phoA:SP-TrxA-His-EK-NLS-spCas9-NLS-T7ter(简称质粒pKG-GE4)，如SEQ ID NO：1所示，为环形质粒，结构示意图见图2。
SEQ ID NO：1中，第5121-5139位核苷酸组成T7启动子，第5140-5164位核苷酸编码Lac操纵子(lac operator)，第5178-5201位核苷酸组成核糖体结合位点(RBS),第5209-5271位核苷酸编码碱性磷酸酶信号肽(phoA signal peptide),第5272-5598位核苷酸编码TrxA蛋白，第5620-5637位核苷酸编码His-Tag，第5638-5652位核苷酸编码肠激酶酶切位点(EK酶切位点)，第5656-5670位核苷酸编码核定位信号，第5701-9801位核苷酸编码spCas9蛋白，第9802-9849位核苷酸编码核定位信号，第9902-9949位核苷酸组成T7终止子。编码spCas9蛋白的核苷酸已进行针对大肠杆菌BL21(DE3)菌株的密码子优化。
质粒pKG-GE4的主要改造如下：①保留了TrxA蛋白的编码区域，TrxA蛋白可以帮助所表达的目的蛋白形成二硫键、增加目的蛋白的溶解性及活性；在TrxA蛋白的编码区域之前加入碱性磷酸酶信号肽的编码序列，碱性磷酸酶信号肽可以引导所表达的目的蛋白分泌至细菌的膜周质腔中并可被原核周质信号肽酶酶切；②在TrxA蛋白的编码序列之后增加His-Tag的编码序列，His-Tag可用于所表达的目的蛋白的富集；③在His-Tag的编码序列下游增加肠激酶酶切位点DDDDK(Asp-Asp-Asp-Asp-Lys)的编码序列，纯化出的蛋白将在肠激酶作用下去除His-Tag和上游所融合的TrxA蛋白；④插入密码子优化后的适宜大肠杆菌BL21(DE3)菌株表达的Cas9基因，同时在该基因的上游和下游均增加核定位信号编码序列，增加后期纯化出的Cas9蛋白的核定位能力。
质粒pKG-GE4中的融合基因如SEQ ID NO：1中第5209-9852位核苷酸所示，编码SEQID NO：2所示的融合蛋白(融合蛋白TrxA-His-EK-NLS-spCas9-NLS，简称为PRONCN蛋白)。由于碱性磷酸酶信号肽以及肠激酶酶切位点的存在，融合蛋白被肠激酶酶切后形成SEQ IDNO：3所示的蛋白质，将SEQ ID NO：3所示的蛋白质命名为NCN蛋白。
实施例2、NCN蛋白的制备和纯化
一、诱导表达
1、将质粒pKG-GE4导入大肠杆菌BL21(DE3)，得到重组菌。
2、将步骤1得到的重组菌接种至含100μg/ml氨苄青霉素的液体LB培养基，37℃、200rpm振荡培养过夜。
3、将步骤2得到的菌液接种至液体LB培养基，30℃、230rpm振荡培养至OD600nm值＝1.0，然后加入异丙基硫代半乳糖苷(IPTG)并使其在体系中的浓度为0.5mM，然后25℃、230rpm振荡培养12小时，然后4℃、10000g离心15分钟，收集菌体。
4、取步骤3得到的菌体，用PBS缓冲液洗涤。
二、融合蛋白TrxA-His-EK-NLS-spCas9-NLS的纯化
1、取步骤一得到的菌体，加入粗提缓冲液并悬浮菌体，然后采用均质机进行菌体破碎(1000par循环三次)，然后4℃、15000g离心30min，收集上清液，上清液采用0.22μm孔径滤膜过滤，收集滤液。本步骤中，每g湿重的菌体配比10ml粗提缓冲液。
粗提缓冲液：含20mM Tris-HCl(pH8.0)、0.5M NaCl、5mM Imidazole、1mM PMSF，余量为ddH2O。
2、采用亲和层析纯化融合蛋白。
首先采用5个柱体积的平衡液平衡Ni-NTA琼脂糖柱(流速为1ml/min)；然后上样50ml步骤1得到的滤液(流速为0.5-1ml/min)；然后用5个柱体积的平衡液洗涤柱子(流速为1ml/min)；然后用5个柱体积的缓冲液洗涤柱子(流速为1ml/min)，以去除杂蛋白；然后用10个柱体积的洗脱液以0.5-1ml/min的流速洗脱，收集过柱后溶液(90-100ml)。
Ni-NTA琼脂糖柱：金斯瑞，L00250/L00250-C，填料为10ml。
平衡液：含20mM Tris-HCl(pH 8.0)、0.5M NaCl、5mM Imidazole，余量为ddH2O。
缓冲液：含20mM Tris-HCl(pH 8.0)、0.5M NaCl、50mM Imidazole，余量为ddH2O。
洗脱液：含20mM Tris-HCl(pH 8.0)、0.5M NaCl、500mM Imidazole，余量为ddH2O。
三、融合蛋白TrxA-His-EK-NLS-spCas9-NLS的酶切与NCN蛋白的纯化
1、取15ml步骤二收集的过柱后溶液，使用Amicon超滤管(Sigma,UFC9100，容量为15ml)将其浓缩至200μl，然后用25mM Tris-HCl(pH8.0)稀释至1ml。采用6个超滤管，共得到6ml。
2、将商品来源的具有His6标签的重组牛肠激酶(生工生物，C620031，重组牛肠激酶轻链，带His6标签，Recombinant Bovine Enterokinase Light Chain,His)加入到步骤1得到的溶液(约6ml)中，25℃酶切16小时。每50μg蛋白量配比加入2个单位的肠激酶。
3、取完成步骤2的溶液(约6ml)，与480μl Ni-NTA树脂(金斯瑞，L00250/L00250-C)混匀，在室温下旋转混匀15min，然后7000g离心3min，收集上清液(4-5.5ml)。
4、取步骤3得到的上清液，使用Amicon超滤管(Sigma,UFC9100，容量为15ml)将其浓缩至200μl，然后加入酶贮存液中，调整蛋白浓度为5mg/ml，即为NCN蛋白溶液。
经测序，NCN蛋白溶液中的蛋白质，N端15个氨基酸残基如SEQ ID NO：3第1至15位所示，即NCN蛋白。
用于后续实施例的NCN蛋白均由NCN蛋白溶液提供。
酶贮存液(pH7.4)：含10mM Tris,300mM NaCl,0.1mM EDTA,1mM DTT,50％(体积比)甘油，余量为ddH2O。
实施例3、NCN蛋白的性能
选择靶向TTN基因的2个gRNA靶点如下：
TTN-gRNA1：AGAGCACAGTCAGCCTGGCG；
TTN-gRNA2：CTTCCAGAATTGGATCTCCG。
用于鉴定包含TTN基因中gRNA的靶点片段的引物如下：
TTN-F55：TACGGAATTGGGGAGCCAGCGGA；
TTN-R560：CAAAGTTAACTCTCTGTGTCT。
一、制备gRNA
1、制备TTN-T7-gRNA1转录模板和TTN-T7-gRNA2转录模板
TTN-T7-gRNA1转录模板为双链DNA分子，如SEQ ID NO：4所示。
TTN-T7-gRNA2转录模板为双链DNA分子，如SEQ ID NO：5所示。
2、体外转录得到gRNA
取TTN-T7-gRNA1转录模板，采用Transcript Aid T7 High Yield TranscriptionKit(Fermentas，K0441)进行体外转录，然后用MEGA clearTM Transcription Clean-Up Kit(Thermo，AM1908)进行回收纯化，得到TTN-gRNA1。TTN-gRNA1为单链RNA，如SEQ ID NO：6所示。
取TTN-T7-gRNA2转录模板，采用Transcript Aid T7 High Yield TranscriptionKit(Fermentas，K0441)进行体外转录，然后用MEGA clearTM Transcription Clean-Up Kit(Thermo，AM1908)进行回收纯化，得到TTN-gRNA2。TTN-gRNA2为单链RNA，如SEQ ID NO：7所示。
二、gRNA与NCN蛋白用量配比优化
1、共转染猪原代成纤维细胞
第一组：将TTN-gRNA1、TTN-gRNA2和NCN蛋白共转染猪原代成纤维细胞。配比：约10万个猪原代成纤维细胞：0.5μg TTN-gRNA1：0.5μg TTN-gRNA2：4μg NCN蛋白。
第二组：将TTN-gRNA1、TTN-gRNA2和NCN蛋白共转染猪原代成纤维细胞。配比：约10万个猪原代成纤维细胞：0.75μg TTN-gRNA1：0.75μg TTN-gRNA2：4μg NCN蛋白。
第三组：将TTN-gRNA1、TTN-gRNA2和NCN蛋白共转染猪原代成纤维细胞。配比：约10万个猪原代成纤维细胞：1μg TTN-gRNA1：1μg TTN-gRNA2：4μg NCN蛋白。
第四组：将TTN-gRNA1、TTN-gRNA2和NCN蛋白共转染猪原代成纤维细胞。配比：约10万个猪原代成纤维细胞：1.25μg TTN-gRNA1：1.25μg TTN-gRNA2：4μg NCN蛋白。
第五组：将TTN-gRNA1和TTN-gRNA2共转染猪原代成纤维细胞。配比：约10万个猪原代成纤维细胞：1μg TTN-gRNA1：1μg TTN-gRNA2。
共转染采用电击转染的方式，采用哺乳动物核转染试剂盒(Neon kit,Thermofisher)与Neon TM transfection system电转仪(参数设置为：1450V、10ms、3pulse)。
2、完成步骤1后，采用完全培养液培养12-18小时，然后更换新的完全培养液进行培养。电转后培养总时间为48小时。
3、完成步骤2后，采用胰蛋白酶消化并收集细胞，提取基因组DNA，采用TTN-F55和TTN-R560组成的引物对进行PCR扩增，然后进行1％琼脂糖凝胶电泳。
电泳图见图3。505bp条带为野生型条带(WT)，254bp左右(野生型条带505bp理论缺失251bp)为缺失突变条带(MT)。
基因缺失突变效率＝(MT灰度/MT条带bp数)/(WT灰度/WT条带bp数+MT灰度/MT条带bp数)×100％。第一组基因缺失突变效率为19.9％，第二组基因缺失突变效率为39.9％，第三组基因缺失突变效率为79.9％，第四组基因缺失突变效率为44.3％。第五组未发生突变。
结果表明，当两个gRNA与NCN蛋白的质量配比为1：1：4，实际用量为1μg：1μg：4μg时基因编辑效率最高。因此，确定两个gRNA与NCN蛋白的最适用量为1μg：1μg：4μg。
三、NCN蛋白与商品Cas9蛋白的基因编辑效率比较
1、共转染猪原代成纤维细胞
Cas9-A组：将TTN-gRNA1、TTN-gRNA2和商品Cas9-A蛋白共转染猪原代成纤维细胞。配比：约10万个猪原代成纤维细胞：1μg TTN-gRNA1：1μg TTN-gRNA2：4μg Cas9-A蛋白。
pKG-GE4组：将TTN-gRNA1、TTN-gRNA2和NCN蛋白共转染猪原代成纤维细胞。配比：约10万个猪原代成纤维细胞：1μg TTN-gRNA1：1μg TTN-gRNA2：4μg NCN蛋白。
Cas9-B组：将TTN-gRNA1、TTN-gRNA2和商品Cas9-B蛋白共转染猪原代成纤维细胞。配比：约10万个猪原代成纤维细胞：1μg TTN-gRNA1：1μg TTN-gRNA2：4μg Cas9-B蛋白。
Control组：将TTN-gRNA1、TTN-gRNA2共转染猪原代成纤维细胞。配比：约10万个猪原代成纤维细胞：1μg TTN-gRNA1：1μg TTN-gRNA2。
共转染采用电击转染的方式，采用哺乳动物核转染试剂盒(Neon kit,Thermofisher)与Neon TM transfection system电转仪(参数设置为：1450V、10ms、3pulse)。
2、完成步骤1后，采用完全培养液培养12-18小时，然后更换新的完全培养液进行培养。电转后培养总时间为48小时。
3、完成步骤2后，采用胰蛋白酶消化并收集细胞，提取基因组DNA，采用TTN-F55和TTN-R560组成的引物对进行PCR扩增，然后进行1％琼脂糖凝胶电泳。
电泳图见图4。采用商品Cas9-A蛋白的基因缺失突变效率为28.5％，采用NCN蛋白的基因缺失突变效率为85.6％，采用商品Cas9-B蛋白的基因缺失突变效率为16.6％。
结果表明，与采用商品的Cas9蛋白相比，采用本发明制备的NCN蛋白使得基因编辑效率显著提高。
实施例4、APOE基因高效gRNA靶点的筛选
猪APOE基因信息：编码Apolipoprotein E蛋白；位于6号染色体；GeneID为397576，Sus scrofa。猪APOE基因编码的蛋白质如SEQ ID NO：8所示。基因组DNA中，猪APOE基因具有3个外显子。猪APOE基因部分序列(含第2外显子及其上下游各350bp)如SEQ ID NO：9所示。
一、APOE基因预设点突变位点及邻近基因组序列保守性分析
18只初生从江香猪，其中雌性10只(分别命名为1、2、3、4、5、6、7、8、9、10)、雄性8只(分别命名为A、B、C、D、E、F、G、H)。
APOE-E2-F：ACCTGATGGCTGTGAACTGG；
APOE-E2-R：GGCGACAAGGACAGAAGGAA。
分别以18只猪的基因组DNA为模板，采用APOE-E2-F和APOE-E2-R组成的引物对进行PCR扩增，然后进行1％琼脂糖凝胶电泳。电泳图见图5。回收PCR扩增产物并进行测序，将测序结果与公共数据库中的APOE基因序列进行比对分析。选择18只猪中共有的保守区进行gRNA靶点的设计。
二、筛选靶点
通过筛选NGG(避开可能的突变位点)初步筛选到若干靶点，经过预实验进一步从中筛选到4个靶点。
4个靶点分别如下：
APOE-E2-gRNA1：GTAATCCCAGAAGCGGCCCA；
APOE-E2-gRNA2：TGTGGTGGGAGGAGCCCAAG；
APOE-E2-gRNA3：CCTGTCTGACCAAGTGCAGG；
APOE-E2-gRNA4：CACCACACGTGCACCTCCGG。
三、制备重组质粒
取质粒pKG-U6gRNA，用限制性内切酶BbsI进行酶切，回收载体骨架(约3kb的线性大片段)。
分别合成APOE-E2-gRNA1-S和APOE-E2-gRNA1-A，然后混合并进行退火，得到具有粘性末端的双链DNA分子。将具有粘性末端的双链DNA分子和载体骨架连接，得到质粒pKG-U6gRNA(APOE-E2-gRNA1)。质粒pKG-U6gRNA(APOE-E2-gRNA1)表达SEQ ID NO：10所示的sgRNAAPOE-E2-gRNA1。
sgRNAAPOE-E2-gRNA1(SEQ ID NO：10)：
GUAAUCCCAGAAGCGGCCCAguuuuagagcuagaaauagcaaguuaaaauaaggcuaguccguuaucaacuugaaaaaguggcaccgagucggugcuuuu。
分别合成APOE-E2-gRNA2-S和APOE-E2-gRNA2-A，然后混合并进行退火，得到具有粘性末端的双链DNA分子。将具有粘性末端的双链DNA分子和载体骨架连接，得到质粒pKG-U6gRNA(APOE-E2-gRNA2)。质粒pKG-U6gRNA(APOE-E2-gRNA2)表达SEQ ID NO：11所示的sgRNAAPOE-E2-gRNA2。
sgRNAAPOE-E2-gRNA2(SEQ ID NO：11)：
UGUGGUGGGAGGAGCCCAAGguuuuagagcuagaaauagcaaguuaaaauaaggcuaguccguuaucaacuugaaaaaguggcaccgagucggugcuuuu。
分别合成APOE-E2-gRNA3-S和APOE-E2-gRNA3-A，然后混合并进行退火，得到具有粘性末端的双链DNA分子。将具有粘性末端的双链DNA分子和载体骨架连接，得到质粒pKG-U6gRNA(APOE-E2-gRNA3)。质粒pKG-U6gRNA(APOE-E2-gRNA3)表达SEQ ID NO：12所示的sgRNAAPOE-E2-gRNA3。
sgRNAAPOE-E2-gRNA3(SEQ ID NO：12)：
CCUGUCUGACCAAGUGCAGGguuuuagagcuagaaauagcaaguuaaaauaaggcuaguccguuaucaacuugaaaaaguggcaccgagucggugcuuuu。
分别合成APOE-E2-gRNA4-S和APOE-E2-gRNA4-A，然后混合并进行退火，得到具有粘性末端的双链DNA分子。将具有粘性末端的双链DNA分子和载体骨架连接，得到质粒pKG-U6gRNA(APOE-E2-gRNA4)。质粒pKG-U6gRNA(APOE-E2-gRNA4)表达SEQ ID NO：13所示的sgRNAAPOE-E2-gRNA4。
sgRNAAPOE-E2-gRNA4(SEQ ID NO：13)：
CACCACACGUGCACCUCCGGguuuuagagcuagaaauagcaaguuaaaauaaggcuaguccguuaucaacuugaaaaaguggcaccgagucggugcuuuu。
APOE-E2-gRNA1-S：caccGTAATCCCAGAAGCGGCCCA；
APOE-E2-gRNA1-A：aaacTGGGCCGCTTCTGGGATTAC；
APOE-E2-gRNA2-S：caccgTGTGGTGGGAGGAGCCCAAG；
APOE-E2-gRNA2-A：aaacCTTGGGCTCCTCCCACCACAc；
APOE-E2-gRNA3-S：caccgCCTGTCTGACCAAGTGCAGG；
APOE-E2-gRNA3-A：aaacCCTGCACTTGGTCAGACAGGc；
APOE-E2-gRNA4-S：caccgCACCACACGTGCACCTCCGG；
APOE-E2-gRNA4-A：aaacCCGGAGGTGCACGTGTGGTGc。
APOE-E2-gRNA1-S、APOE-E2-gRNA1-A、APOE-E2-gRNA2-S、APOE-E2-gRNA2-A、APOE-E2-gRNA3-S、APOE-E2-gRNA3-A、APOE-E2-gRNA4-S和APOE-E2-gRNA4-A均为单链DNA分子。
四、不同靶点的编辑效率比较
1、共转染
第一组：将质粒pKG-U6gRNA(APOE-E2-gRNA1)、质粒pKG-GE3共转染猪原代成纤维细胞。配比：约20万个猪原代成纤维细胞：0.92μg质粒pKG-U6gRNA(APOE-E2-gRNA1)：1.08μg质粒pKG-GE3。
第二组：将质粒pKG-U6gRNA(APOE-E2-gRNA2)、质粒pKG-GE3共转染猪原代成纤维细胞。配比：约20万个猪原代成纤维细胞：0.92μg质粒pKG-U6gRNA(APOE-E2-gRNA2)：1.08μg质粒pKG-GE3。
第三组：将质粒pKG-U6gRNA(APOE-E2-gRNA3)、质粒pKG-GE3共转染猪原代成纤维细胞。配比：约20万个猪原代成纤维细胞：0.92μg质粒pKG-U6gRNA(APOE-E2-gRNA3)：1.08μg质粒pKG-GE3。
第四组：将质粒pKG-U6gRNA(APOE-E2-gRNA4)、质粒pKG-GE3共转染猪原代成纤维细胞。配比：约20万个猪原代成纤维细胞：0.92μg质粒pKG-U6gRNA(APOE-E2-gRNA4)：1.08μg质粒pKG-GE3。
第五组：猪原代成纤维细胞，同等电转参数不加质粒进行电转操作。
共转染采用电击转染的方式，采用哺乳动物核转染试剂盒(Neon kit,Thermofisher)与Neon TM transfection system电转仪(参数设置为：1450V、10ms、3pulse)。
2、完成步骤1后，采用完全培养液培养12-18小时，然后更换新的完全培养液进行培养。电转后培养总时间为48小时。
3、完成步骤2后，采用胰蛋白酶消化并收集细胞，裂解细胞，提取基因组DNA，采用APOE-E2-F和APOE-E2-R组成的引物对进行PCR扩增，然后进行1％琼脂糖凝胶电泳。
将目的产物切胶回收后送测序公司进行测序，测序峰图见图6。将测序结果利用网页版Synthego ICE工具分析测序峰图得出不同靶点的基因编辑效率。第一组至第四组的基因编辑效率依次为19％、61％、12％、8％。第五组未发生基因编辑。结果表明，APOE-E2-gRNA2编辑效率较高。
实施例5、FBN1基因高效gRNA靶点的筛选
猪FBN1基因信息：编码原纤蛋白1(Fibrillin-1)；位于猪1号染色体；GeneID为414836，Sus scrofa。猪FBN1基因编码的蛋白质如SEQ ID NO：14所示。基因组DNA中，猪FBN1基因具有65个外显子。猪FBN1基因部分序列(含第25外显子和第26外显子)如SEQ ID NO：15所示。FBN1基因C1039G突变会导致动脉硬化增加。
一、FBN1基因预设点突变位点及邻近基因组序列保守性分析
18只初生从江香猪，其中雌性10只(分别命名为1、2、3、4、5、6、7、8、9、10)、雄性8只(分别命名为A、B、C、D、E、F、G、H)。
FBN1-JDF412：CCTGGGTTTAGAATGGGGTGT；
FBN1-JDR692：ACGCTTTCCTTCTTGAGGCA；
FBN1-JDF419：TTAGAATGGGGTGTTGAGCCC；
FBN1-JDR673：AGTGGGTTAAAGGCGAGGTG。
用命名为1的猪的耳组织提取基因组作为模板，采用不同引物对进行PCR扩增，然后进行1％琼脂糖凝胶电泳。电泳图见图7。图7中：组1：FBN1-JDF412/FBN1-JDR692；组2：FBN1-JDF412/FBN1-JDR673；组3：FBN1-JDF419/FBN1-JDR692；组4：FBN1-JDF419/FBN1-JDR673。结果表明，优选采用FBN1-JDF419和FBN1-JDR673组成的引物对进行目的片段扩增。
分别以18只猪的基因组DNA为模板，采用FBN1-JDF419和FBN1-JDR673组成的引物对进行PCR扩增，然后进行1％琼脂糖凝胶电泳。电泳图见图8。回收PCR扩增产物并进行测序，将测序结果与公共数据库中的FBN1基因序列进行比对分析。选择18只猪中共有的保守区进行gRNA靶点的设计。
二、筛选靶点
通过筛选NGG(避开可能的突变位点)初步筛选到若干靶点，经过预实验进一步从中筛选到7个靶点。
7个靶点分别如下：
FBN1-E25-gRNA1：ATCCCCAACCTCTGTACCCA；
FBN1-E25-gRNA2：AGTGTTCCTGCACTTGCCGT；
FBN1-E25-gRNA3：TAGTGTTCCTGCACTTGCCG；
FBN1-E25-gRNA4：TGCACTTGCCGTGGGTACAG；
FBN1-E25-gRNA5：GGCAAGTGCAGGAACACTAT；
FBN1-E25-gRNA6：CTTCAACAAAAATTGAATGT；
FBN1-E25-gRNA7：TCTTCAACAAAAATTGAATG。
三、制备重组质粒
取质粒pKG-U6gRNA，用限制性内切酶BbsI进行酶切，回收载体骨架(约3kb的线性大片段)。
分别合成FBN1-E25-gRNA1-S和FBN1-E25-gRNA1-A，然后混合并进行退火，得到具有粘性末端的双链DNA分子。将具有粘性末端的双链DNA分子和载体骨架连接，得到质粒pKG-U6gRNA(FBN1-E25-gRNA1)。质粒pKG-U6gRNA(FBN1-E25-gRNA1)表达SEQ ID NO：16所示的sgRNAFBN1-E25-gRNA1。
sgRNAFBN1-E25-gRNA1(SEQ ID NO：16)：
AUCCCCAACCUCUGUACCCAguuuuagagcuagaaauagcaaguuaaaauaaggcuaguccguuaucaacuugaaaaaguggcaccgagucggugcuuuu。
分别合成FBN1-E25-gRNA2-S和FBN1-E25-gRNA2-A，然后混合并进行退火，得到具有粘性末端的双链DNA分子。将具有粘性末端的双链DNA分子和载体骨架连接，得到质粒pKG-U6gRNA(FBN1-E25-gRNA2)。质粒pKG-U6gRNA(FBN1-E25-gRNA2)表达SEQ ID NO：17所示的sgRNAFBN1-E25-gRNA2。
sgRNAFBN1-E25-gRNA2(SEQ ID NO：17)：
AGUGUUCCUGCACUUGCCGUguuuuagagcuagaaauagcaaguuaaaauaaggcuaguccguuaucaacuugaaaaaguggcaccgagucggugcuuuu。
分别合成FBN1-E25-gRNA3-S和FBN1-E25-gRNA3-A，然后混合并进行退火，得到具有粘性末端的双链DNA分子。将具有粘性末端的双链DNA分子和载体骨架连接，得到质粒pKG-U6gRNA(FBN1-E25-gRNA3)。质粒pKG-U6gRNA(FBN1-E25-gRNA3)表达SEQ ID NO：18所示的sgRNAFBN1-E25-gRNA3。
sgRNAFBN1-E25-gRNA3(SEQ ID NO：18)：
UAGUGUUCCUGCACUUGCCGguuuuagagcuagaaauagcaaguuaaaauaaggcuaguccguuaucaacuugaaaaaguggcaccgagucggugcuuuu。
分别合成FBN1-E25-gRNA4-S和FBN1-E25-gRNA4-A，然后混合并进行退火，得到具有粘性末端的双链DNA分子。将具有粘性末端的双链DNA分子和载体骨架连接，得到质粒pKG-U6gRNA(FBN1-E25-gRNA4)。质粒pKG-U6gRNA(FBN1-E25-gRNA4)表达SEQ ID NO：19所示的sgRNAFBN1-E25-gRNA4。
sgRNAFBN1-E25-gRNA4(SEQ ID NO：19)：
UGCACUUGCCGUGGGUACAGguuuuagagcuagaaauagcaaguuaaaauaaggcuaguccguuaucaacuugaaaaaguggcaccgagucggugcuuuu。
分别合成FBN1-E25-gRNA5-S和FBN1-E25-gRNA5-A，然后混合并进行退火，得到具有粘性末端的双链DNA分子。将具有粘性末端的双链DNA分子和载体骨架连接，得到质粒pKG-U6gRNA(FBN1-E25-gRNA5)。质粒pKG-U6gRNA(FBN1-E25-gRNA5)表达SEQ ID NO：20所示的sgRNAFBN1-E25-gRNA5。
sgRNAFBN1-E25-gRNA5(SEQ ID NO：20)：
GGCAAGUGCAGGAACACUAUguuuuagagcuagaaauagcaaguuaaaauaaggcuaguccguuaucaacuugaaaaaguggcaccgagucggugcuuuu。
分别合成FBN1-E25-gRNA6-S和FBN1-E25-gRNA6-A，然后混合并进行退火，得到具有粘性末端的双链DNA分子。将具有粘性末端的双链DNA分子和载体骨架连接，得到质粒pKG-U6gRNA(FBN1-E25-gRNA6)。质粒pKG-U6gRNA(FBN1-E25-gRNA6)表达SEQ ID NO：21所示的sgRNAFBN1-E25-gRNA6。
sgRNAFBN1-E25-gRNA6(SEQ ID NO：21)：
CUUCAACAAAAAUUGAAUGUguuuuagagcuagaaauagcaaguuaaaauaaggcuaguccguuaucaacuugaaaaaguggcaccgagucggugcuuuu。
分别合成FBN1-E25-gRNA7-S和FBN1-E25-gRNA7-A，然后混合并进行退火，得到具有粘性末端的双链DNA分子。将具有粘性末端的双链DNA分子和载体骨架连接，得到质粒pKG-U6gRNA(FBN1-E25-gRNA7)。质粒pKG-U6gRNA(FBN1-E25-gRNA7)表达SEQ ID NO：22所示的sgRNAFBN1-E25-gRNA7。
sgRNAFBN1-E25-gRNA7(SEQ ID NO：22)：
UCUUCAACAAAAAUUGAAUGguuuuagagcuagaaauagcaaguuaaaauaaggcuaguccguuaucaacuugaaaaaguggcaccgagucggugcuuuu。
FBN1-E25-gRNA1-S：caccgATCCCCAACCTCTGTACCCA；
FBN1-E25-gRNA1-A：aaacTGGGTACAGAGGTTGGGGATc；
FBN1-E25-gRNA2-S：caccgAGTGTTCCTGCACTTGCCGT；
FBN1-E25-gRNA2-A：aaacACGGCAAGTGCAGGAACACTc；
FBN1-E25-gRNA3-S：caccgTAGTGTTCCTGCACTTGCCG；
FBN1-E25-gRNA3-A：aaacCGGCAAGTGCAGGAACACTAc；
FBN1-E25-gRNA4-S：caccgTGCACTTGCCGTGGGTACAG；
FBN1-E25-gRNA4-A：aaacCTGTACCCACGGCAAGTGCAc；
FBN1-E25-gRNA5-S：caccGGCAAGTGCAGGAACACTAT；
FBN1-E25-gRNA5-A：aaacATAGTGTTCCTGCACTTGCC；
FBN1-E25-gRNA6-S：caccgCTTCAACAAAAATTGAATGT；
FBN1-E25-gRNA6-A：aaacACATTCAATTTTTGTTGAAGc；
FBN1-E25-gRNA7-S：caccgTCTTCAACAAAAATTGAATG；
FBN1-E25-gRNA7-A：aaacCATTCAATTTTTGTTGAAGAc。
FBN1-E25-gRNA1-S、FBN1-E25-gRNA1-A、FBN1-E25-gRNA2-S、FBN1-E25-gRNA2-A、FBN1-E25-gRNA3-S、FBN1-E25-gRNA3-A、FBN1-E25-gRNA4-S、FBN1-E25-gRNA4-A、FBN1-E25-gRNA5-S、FBN1-E25-gRNA5-A、FBN1-E25-gRNA6-S、FBN1-E25-gRNA6-A、FBN1-E25-gRNA7-S、FBN1-E25-gRNA7-A均为单链DNA分子。
四、不同靶点的编辑效率比较
1、共转染
第一组：将质粒pKG-U6gRNA(FBN1-E25-gRNA1)、质粒pKG-GE3共转染猪原代成纤维细胞。配比：约20万个猪原代成纤维细胞：0.92μg质粒pKG-U6gRNA(FBN1-E25-gRNA1)：1.08μg质粒pKG-GE3。
第二组：将质粒pKG-U6gRNA(FBN1-E25-gRNA2)、质粒pKG-GE3共转染猪原代成纤维细胞。配比：约20万个猪原代成纤维细胞：0.92μg质粒pKG-U6gRNA(FBN1-E25-gRNA2)：1.08μg质粒pKG-GE3。
第三组：将质粒pKG-U6gRNA(FBN1-E25-gRNA3)、质粒pKG-GE3共转染猪原代成纤维细胞。配比：约20万个猪原代成纤维细胞：0.92μg质粒pKG-U6gRNA(FBN1-E25-gRNA3)：1.08μg质粒pKG-GE3。
第四组：将质粒pKG-U6gRNA(FBN1-E25-gRNA4)、质粒pKG-GE3共转染猪原代成纤维细胞。配比：约20万个猪原代成纤维细胞：0.92μg质粒pKG-U6gRNA(FBN1-E25-gRNA4)：1.08μg质粒pKG-GE3。
第五组：将质粒pKG-U6gRNA(FBN1-E25-gRNA5)、质粒pKG-GE3共转染猪原代成纤维细胞。配比：约20万个猪原代成纤维细胞：0.92μg质粒pKG-U6gRNA(FBN1-E25-gRNA5)：1.08μg质粒pKG-GE3。
第六组：将质粒pKG-U6gRNA(FBN1-E25-gRNA6)、质粒pKG-GE3共转染猪原代成纤维细胞。配比：约20万个猪原代成纤维细胞：0.92μg质粒pKG-U6gRNA(FBN1-E25-gRNA6)：1.08μg质粒pKG-GE3。
第七组：将质粒pKG-U6gRNA(FBN1-E25-gRNA7)、质粒pKG-GE3共转染猪原代成纤维细胞。配比：约20万个猪原代成纤维细胞：0.92μg质粒pKG-U6gRNA(FBN1-E25-gRNA7)：1.08μg质粒pKG-GE3。
第八组：猪原代成纤维细胞，同等电转参数不加质粒进行电转操作。
共转染采用电击转染的方式，采用哺乳动物核转染试剂盒(Neon kit,Thermofisher)与Neon TM transfection system电转仪(参数设置为：1450V、10ms、3pulse)。
2、完成步骤1后，采用完全培养液培养12-18小时，然后更换新的完全培养液进行培养。电转后培养总时间为48小时。
3、完成步骤2后，采用胰蛋白酶消化并收集细胞，裂解细胞，提取基因组DNA，采用FBN1-JDF419和FBN1-JDR673组成的引物对进行PCR扩增，然后进行1％琼脂糖凝胶电泳。电泳图见图9。
将目的产物切胶回收后送测序公司进行测序。将测序结果利用网页版SynthegoICE工具分析测序峰图得出不同靶点的基因编辑效率。第一组至第七组的基因编辑效率依次为25％、20％、13％、45％、12％、52％、9％。第八组未发生基因编辑。结果表明，FBN1-E25-gRNA4和FBN1-E25-gRNA6编辑效率较高。
实施例6、制备APOE和FBN1基因同时编辑的单细胞克隆
选用实施例4中筛到的一个针对APOE基因的高效gRNA靶点(APOE-E2-gRNA2)，该靶点通过将质粒pKG-U6gRNA(APOE-E2-gRNA2)和质粒pKG-GE3共转染细胞进行基因编辑；选用实施例5中筛到的两个针对FBN1基因的高效gRNA靶点(FBN1-E25-gRNA4和FBN1-E25-gRNA6)，通过将相应的两个sgRNA、NCN蛋白及FBN1-mutant-ss163 donor DNA共转染细胞进行定点突变(FBN1基因C1039G突变)。
一、制备gRNA
1、制备FBN1-T7-gRNA4转录模板和FBN1-T7-gRNA6转录模板。
FBN1-T7-gRNA4转录模板为双链DNA分子，如SEQ ID NO：23所示。
FBN1-T7-gRNA6转录模板为双链DNA分子，如SEQ ID NO：24所示。
2、体外转录得到gRNA
取FBN1-T7-gRNA4转录模板，采用Transcript Aid T7 High YieldTranscription Kit(Fermentas，K0441)进行体外转录，然后用MEGA clearTMTranscription Clean-Up Kit(Thermo，AM1908)进行回收纯化，得到FBN1-gRNA4。FBN1-gRNA4为单链RNA，如SEQ ID NO：25所示。
SEQ ID NO：25：
GGUGCACUUGCCGUGGGUACAGGUUUUAGAGCUAGAAAUAGCAAGUUAAAAUAAGGCUAGUCCGUUAUCAACUUGAAAAAGUGGCACCGAGUCGGUGCUUUU。
取FBN1-T7-gRNA6转录模板，采用Transcript Aid T7 High YieldTranscription Kit(Fermentas，K0441)进行体外转录，然后用MEGA clearTMTranscription Clean-Up Kit(Thermo，AM1908)进行回收纯化，得到FBN1-gRNA6。FBN1-gRNA6为单链RNA，如SEQ ID NO：26所示。
SEQ ID NO：26：
GGCUUCAACAAAAAUUGAAUGUGUUUUAGAGCUAGAAAUAGCAAGUUAAAAUAAGGCUAGUCCGUUAUCAACUUGAAAAAGUGGCACCGAGUCGGUGCUUUU。
二、合成含有FBN1突变位点的单链Donor DNA
合成对应C1039G突变的单链DNA作为Donor DNA，该单链DNA除靶位点突变外还含有FBN1-E25-gRNA4和FBN1-E25-gRNA6靶点的PAM序列同义突变。该单链Donor DNA命名为FBN1-mutant-ss163。
FBN1-mutant-ss163如SEQ ID NO：27所示。
SEQ ID NO：27：
ggtttagaatggggtgttgagccctcctcactgatcactgatggacttttAccacattcaatttttgttgaagacatcaacgagtgcaagatgatccccaaTctcGgtacccacggcaagtgcaggaacactatcggcagcttcaagtgcagatgtgacagtg。
三、转染猪原代成纤维细胞
1、将pKG-U6gRNA(APOE-E2-gRNA2)质粒、质粒pKG-GE3、FBN1-gRNA4、FBN1-gRNA6、FBN1-mutant-ss163和NCN蛋白共转染猪原代成纤维细胞。配比：约20万个猪原代成纤维细胞：0.92μg质粒pKG-U6gRNA(APOE-E2-gRNA2)：1.08μg质粒pKG-GE3：1μg FBN1-gRNA4：1μgFBN1-gRNA6：2μg FBN1-mutant-ss163：4μg NCN蛋白。共转染采用电击转染的方式，采用哺乳动物核转染试剂盒(Neon kit,Thermofisher)与Neon TM transfection system电转仪(参数设置为：1450V、10ms、3pulse)。
2、完成步骤1后，采用完全培养液培养16-18小时，然后更换新的完全培养液进行培养。电转后培养总时间为48小时。
3、完成步骤2后，采用胰蛋白酶消化并收集细胞，然后用完全培养液洗涤，然后用完全培养液重悬，然后分别挑取各个单细胞转移到96孔板中(每个孔1个细胞，每个孔中装有100μl完全培养液)，培养2周(每2-3天更换新的完全培养液)。
4、完成步骤3后，采用胰蛋白酶消化并收集细胞(每孔得到的细胞，约2/3接种到装有完全培养液的6孔板中，剩余的1/3收集在1.5mL离心管中)。
5、取步骤4的6孔板，培养直至细胞长至80％汇合度，采用胰蛋白酶消化并收集细胞，使用细胞冻存液(90％完全培养基+10％DMSO，体积比)将细胞冻存。
6、取步骤4的离心管，取细胞，进行细胞裂解并提取基因组DNA，分别采用APOE-E2-F和APOE-E2-R组成的引物对和FBN1-JDF419和FBN1-JDR673组成的引物对进行PCR扩增，将猪原代成纤维细胞作为野生型对照(WT)。
7、回收PCR扩增产物并测序以确认APOE基因的编辑情况和FBN1基因的点突变情况。
猪原代成纤维细胞的测序结果只有一种，其基因型为纯合野生型。如果某一单细胞克隆的测序结果有两种，一种与猪原代成纤维细胞的测序结果一致，另一种与猪原代成纤维细胞的测序结果相比发生了突变(突变包括一个或多个核苷酸的缺失、插入或替换)，该单细胞克隆的基因型为杂合型；如果某一单细胞克隆的测序结果为两种，均与猪原代成纤维细胞的测序结果相比发生了突变(突变包括一个或多个核苷酸的缺失、插入或替换)，该单细胞克隆的基因型为双等位基因不同突变型；如果某一单细胞克隆的测序结果为一种，且与猪原代成纤维细胞的测序结果相比发生了突变(突变包括一个或多个核苷酸的缺失、插入或替换)，该单细胞克隆的基因型为双等位基因相同突变型；如果某一单细胞克隆的测序结果为一种，且与猪原代成纤维细胞的测序结果一致，该单细胞克隆的基因型为纯合野生型。
表1和表2中编号相同的单细胞克隆为同一单细胞克隆。
根据测序比对确认APOE基因的编辑结果见表1。编号为2、7、17、19、22、24、28、30、33、36、40的单细胞克隆的基因型为纯合野生型。编号为1、3、9、11、14、16、18、23、25、27、29、35、37的单细胞克隆的基因型为杂合型。编号为5、10、26、31、39的单细胞克隆的基因型为双等位基因相同突变型。编号为4、6、8、12、13、15、20、21、32、34、38的单细胞克隆的基因型为双等位基因不同突变型。得到APOE基因编辑单细胞克隆的比率为72.5％。
示例性的测序比对结果见图10至图13。图10是编号为2的单细胞克隆的反向测序与野生型序列的比对结果，判定为纯合野生型。图11是编号为3的单细胞克隆的反向测序与野生型序列的比对结果，判定为杂合型。图12是编号为10的单细胞克隆的正向测序与野生型序列的比对结果，判定为双等位基因相同突变型。图13是编号为8的单细胞克隆的正向和反向测序同时与野生型序列的比对结果，判定为双等位基因不同突变型。
表1 APOE基因编辑的基因型鉴定结果
根据测序比对确认FBN1基因点突变单细胞克隆的基因型结果见表2。编号为3、8、17、21、33、38的单细胞克隆的基因型为纯合野生型。编号为1、5、6、10、12、14、18、24、26、30、34、35的单细胞克隆的基因型为杂合型。编号为2、7、9、13、20、22、23、25、27、28、31、36、37、39的单细胞克隆的基因型为双等位基因不同突变型。编号为4、11、15、16、19、29、32、40的单细胞克隆的基因型为双等位基因相同突变型。编号为10、14、26、30的单细胞克隆为靶标位点突变的杂合型(即两条同源染色体中的一条完成了单链Donor DNA的替换)。编号为22、25、28、37的单细胞克隆为靶位点突变的双等位基因不同突变型(即两条同源染色体中的一条完成了单链Donor DNA的替换)。编号为15的单细胞克隆为靶位点突变的双等位基因相同突变型(即两条同源染色体均完成了单链Donor DNA的替换)。得到FBN1基因编辑单细胞克隆的比率为85％，得到靶位点突变的单细胞克隆的比率为22.5％。
示例性的测序比对结果见图14至图19。图14是编号为3单细胞克隆的反向测序与靶标位点野生型序列的比对结果，判定为纯合野生型。图15是编号为1的单细胞克隆的反向测序与靶标位点野生型序列的比对结果，判定为杂合型。图16是编号为2的单细胞克隆的反向测序与靶标位点野生型序列的比对结果，判定为双等位基因不同突变型。图17是编号为11的单细胞克隆的反向测序与靶标位点野生型序列的比对结果，判定为双等位基因相同突变型。图18是编号为10的单细胞克隆的反向测序与靶标位点野生型序列的比对结果，判定为具有靶标位点点突变的杂合型。图19是编号为15的单细胞克隆的反向测序与靶标位点野生型序列的比对结果，判定为具有靶标位点点突变的双等位基因相同突变型。
表2 FBN1基因点突变单细胞克隆的基因型结果
注：靶位点突变指的是完成了单链Donor DNA的替换；即用SEQ ID NO：27所示的DNA分子取代了染色体DNA中SEQ ID NO：28所示的DNA分子。
APOE基因为双等位基因突变型(包括双等位基因相同突变型或双等位基因不同突变型)且FBN1基因为靶位点突变杂合型(靶位点突变的双等位基因不同突变型、靶位点突变的杂合型)的单细胞克隆为目标单细胞克隆。目标单细胞克隆可作为核移植供体细胞进行体细胞克隆，可以得到克隆猪，即为动脉粥样硬化疾病模型猪。
编号为10、26的单细胞克隆为目标单细胞克隆。目标单细胞克隆可作为核移植供体细胞进行体细胞克隆，可以得到克隆猪，即为动脉粥样硬化疾病模型猪。
以上对本发明进行了详述。对于本领域技术人员来说，在不脱离本发明的宗旨和范围，以及无需进行不必要的实验情况下，可在等同参数、浓度和条件下，在较宽范围内实施本发明。虽然本发明给出了特殊的实施例，应该理解为，可以对本发明作进一步的改进。总之，按本发明的原理，本申请欲包括任何变更、用途或对本发明的改进，包括脱离了本申请中已公开范围，而用本领域已知的常规技术进行的改变。按以下附带的权利要求的范围，可以进行一些基本特征的应用。
序列表
<110> 南京启真基因工程有限公司
<120> 一种构建AF双基因突变的动脉粥样硬化模型猪核移植供体细胞的基因编辑系统及其应用
<130> GNCYX211723
<160> 30
<170> SIPOSequenceListing 1.0
<210> 1
<211> 9974
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 1
tggcgaatgg gacgcgccct gtagcggcgc attaagcgcg gcgggtgtgg tggttacgcg 60
cagcgtgacc gctacacttg ccagcgccct agcgcccgct cctttcgctt tcttcccttc 120
ctttctcgcc acgttcgccg gctttccccg tcaagctcta aatcgggggc tccctttagg 180
gttccgattt agtgctttac ggcacctcga ccccaaaaaa cttgattagg gtgatggttc 240
acgtagtggg ccatcgccct gatagacggt ttttcgccct ttgacgttgg agtccacgtt 300
ctttaatagt ggactcttgt tccaaactgg aacaacactc aaccctatct cggtctattc 360
ttttgattta taagggattt tgccgatttc ggcctattgg ttaaaaaatg agctgattta 420
acaaaaattt aacgcgaatt ttaacaaaat attaacgttt acaatttcag gtggcacttt 480
tcggggaaat gtgcgcggaa cccctatttg tttatttttc taaatacatt caaatatgta 540
tccgctcatg agacaataac cctgataaat gcttcaataa tattgaaaaa ggaagagtat 600
gagtattcaa catttccgtg tcgcccttat tccctttttt gcggcatttt gccttcctgt 660
ttttgctcac ccagaaacgc tggtgaaagt aaaagatgct gaagatcagt tgggtgcacg 720
agtgggttac atcgaactgg atctcaacag cggtaagatc cttgagagtt ttcgccccga 780
agaacgtttt ccaatgatga gcacttttaa agttctgcta tgtggcgcgg tattatcccg 840
tattgacgcc gggcaagagc aactcggtcg ccgcatacac tattctcaga atgacttggt 900
tgagtactca ccagtcacag aaaagcatct tacggatggc atgacagtaa gagaattatg 960
cagtgctgcc ataaccatga gtgataacac tgcggccaac ttacttctga caacgatcgg 1020
aggaccgaag gagctaaccg cttttttgca caacatgggg gatcatgtaa ctcgccttga 1080
tcgttgggaa ccggagctga atgaagccat accaaacgac gagcgtgaca ccacgatgcc 1140
tgcagcaatg gcaacaacgt tgcgcaaact attaactggc gaactactta ctctagcttc 1200
ccggcaacaa ttaatagact ggatggaggc ggataaagtt gcaggaccac ttctgcgctc 1260
ggcccttccg gctggctggt ttattgctga taaatctgga gccggtgagc gtgggtctcg 1320
cggtatcatt gcagcactgg ggccagatgg taagccctcc cgtatcgtag ttatctacac 1380
gacggggagt caggcaacta tggatgaacg aaatagacag atcgctgaga taggtgcctc 1440
actgattaag cattggtaac tgtcagacca agtttactca tatatacttt agattgattt 1500
aaaacttcat ttttaattta aaaggatcta ggtgaagatc ctttttgata atctcatgac 1560
caaaatccct taacgtgagt tttcgttcca ctgagcgtca gaccccgtag aaaagatcaa 1620
aggatcttct tgagatcctt tttttctgcg cgtaatctgc tgcttgcaaa caaaaaaacc 1680
accgctacca gcggtggttt gtttgccgga tcaagagcta ccaactcttt ttccgaaggt 1740
aactggcttc agcagagcgc agataccaaa tactgtcctt ctagtgtagc cgtagttagg 1800
ccaccacttc aagaactctg tagcaccgcc tacatacctc gctctgctaa tcctgttacc 1860
agtggctgct gccagtggcg ataagtcgtg tcttaccggg ttggactcaa gacgatagtt 1920
accggataag gcgcagcggt cgggctgaac ggggggttcg tgcacacagc ccagcttgga 1980
gcgaacgacc tacaccgaac tgagatacct acagcgtgag ctatgagaaa gcgccacgct 2040
tcccgaaggg agaaaggcgg acaggtatcc ggtaagcggc agggtcggaa caggagagcg 2100
cacgagggag cttccagggg gaaacgcctg gtatctttat agtcctgtcg ggtttcgcca 2160
cctctgactt gagcgtcgat ttttgtgatg ctcgtcaggg gggcggagcc tatggaaaaa 2220
cgccagcaac gcggcctttt tacggttcct ggccttttgc tggccttttg ctcacatgtt 2280
ctttcctgcg ttatcccctg attctgtgga taaccgtatt accgcctttg agtgagctga 2340
taccgctcgc cgcagccgaa cgaccgagcg cagcgagtca gtgagcgagg aagcggaaga 2400
gcgcctgatg cggtattttc tccttacgca tctgtgcggt atttcacacc gcatatatgg 2460
tgcactctca gtacaatctg ctctgatgcc gcatagttaa gccagtatac actccgctat 2520
cgctacgtga ctgggtcatg gctgcgcccc gacacccgcc aacacccgct gacgcgccct 2580
gacgggcttg tctgctcccg gcatccgctt acagacaagc tgtgaccgtc tccgggagct 2640
gcatgtgtca gaggttttca ccgtcatcac cgaaacgcgc gaggcagctg cggtaaagct 2700
catcagcgtg gtcgtgaagc gattcacaga tgtctgcctg ttcatccgcg tccagctcgt 2760
tgagtttctc cagaagcgtt aatgtctggc ttctgataaa gcgggccatg ttaagggcgg 2820
ttttttcctg tttggtcact gatgcctccg tgtaaggggg atttctgttc atgggggtaa 2880
tgataccgat gaaacgagag aggatgctca cgatacgggt tactgatgat gaacatgccc 2940
ggttactgga acgttgtgag ggtaaacaac tggcggtatg gatgcggcgg gaccagagaa 3000
aaatcactca gggtcaatgc cagcgcttcg ttaatacaga tgtaggtgtt ccacagggta 3060
gccagcagca tcctgcgatg cagatccgga acataatggt gcagggcgct gacttccgcg 3120
tttccagact ttacgaaaca cggaaaccga agaccattca tgttgttgct caggtcgcag 3180
acgttttgca gcagcagtcg cttcacgttc gctcgcgtat cggtgattca ttctgctaac 3240
cagtaaggca accccgccag cctagccggg tcctcaacga caggagcacg atcatgcgca 3300
cccgtggggc cgccatgccg gcgataatgg cctgcttctc gccgaaacgt ttggtggcgg 3360
gaccagtgac gaaggcttga gcgagggcgt gcaagattcc gaataccgca agcgacaggc 3420
cgatcatcgt cgcgctccag cgaaagcggt cctcgccgaa aatgacccag agcgctgccg 3480
gcacctgtcc tacgagttgc atgataaaga agacagtcat aagtgcggcg acgatagtca 3540
tgccccgcgc ccaccggaag gagctgactg ggttgaaggc tctcaagggc atcggtcgag 3600
atcccggtgc ctaatgagtg agctaactta cattaattgc gttgcgctca ctgcccgctt 3660
tccagtcggg aaacctgtcg tgccagctgc attaatgaat cggccaacgc gcggggagag 3720
gcggtttgcg tattgggcgc cagggtggtt tttcttttca ccagtgagac gggcaacagc 3780
tgattgccct tcaccgcctg gccctgagag agttgcagca agcggtccac gctggtttgc 3840
cccagcaggc gaaaatcctg tttgatggtg gttaacggcg ggatataaca tgagctgtct 3900
tcggtatcgt cgtatcccac taccgagatg tccgcaccaa cgcgcagccc ggactcggta 3960
atggcgcgca ttgcgcccag cgccatctga tcgttggcaa ccagcatcgc agtgggaacg 4020
atgccctcat tcagcatttg catggtttgt tgaaaaccgg acatggcact ccagtcgcct 4080
tcccgttccg ctatcggctg aatttgattg cgagtgagat atttatgcca gccagccaga 4140
cgcagacgcg ccgagacaga acttaatggg cccgctaaca gcgcgatttg ctggtgaccc 4200
aatgcgacca gatgctccac gcccagtcgc gtaccgtctt catgggagaa aataatactg 4260
ttgatgggtg tctggtcaga gacatcaaga aataacgccg gaacattagt gcaggcagct 4320
tccacagcaa tggcatcctg gtcatccagc ggatagttaa tgatcagccc actgacgcgt 4380
tgcgcgagaa gattgtgcac cgccgcttta caggcttcga cgccgcttcg ttctaccatc 4440
gacaccacca cgctggcacc cagttgatcg gcgcgagatt taatcgccgc gacaatttgc 4500
gacggcgcgt gcagggccag actggaggtg gcaacgccaa tcagcaacga ctgtttgccc 4560
gccagttgtt gtgccacgcg gttgggaatg taattcagct ccgccatcgc cgcttccact 4620
ttttcccgcg ttttcgcaga aacgtggctg gcctggttca ccacgcggga aacggtctga 4680
taagagacac cggcatactc tgcgacatcg tataacgtta ctggtttcac attcaccacc 4740
ctgaattgac tctcttccgg gcgctatcat gccataccgc gaaaggtttt gcgccattcg 4800
atggtgtccg ggatctcgac gctctccctt atgcgactcc tgcattagga agcagcccag 4860
tagtaggttg aggccgttga gcaccgccgc cgcaaggaat ggtgcatgca aggagatggc 4920
gcccaacagt cccccggcca cggggcctgc caccataccc acgccgaaac aagcgctcat 4980
gagcccgaag tggcgagccc gatcttcccc atcggtgatg tcggcgatat aggcgccagc 5040
aaccgcacct gtggcgccgg tgatgccggc cacgatgcgt ccggcgtaga ggatcgagat 5100
cgatctcgat cccgcgaaat taatacgact cactataggg gaattgtgag cggataacaa 5160
ttcccctcta gaaataattt tgtttaactt taagaaggag atatacatat gaaacaaagc 5220
actattgcac tggcactctt accgttactg tttacccctg tgacaaaagc catgagcgat 5280
aaaattattc acctgactga cgacagtttt gacacggatg tactcaaagc ggacggggcg 5340
atcctcgtcg atttctgggc agagtggtgc ggtccgtgca aaatgatcgc cccgattctg 5400
gatgaaatcg ctgacgaata tcagggcaaa ctgaccgttg caaaactgaa catcgatcaa 5460
aaccctggca ctgcgccgaa atatggcatc cgtggtatcc cgactctgct gctgttcaaa 5520
aacggtgaag tggcggcaac caaagtgggt gcactgtcta aaggtcagtt gaaagagttc 5580
ctcgacgcta acctggccgg ttctggttct ggccatatgc accatcatca tcatcatgac 5640
gatgacgata agatgcccaa aaagaaacga aaggtgggta tccacggagt cccagcagcc 5700
gacaaaaaat atagcatcgg cctggacatc ggtaccaaca gcgttggctg ggcagtgatc 5760
actgatgaat acaaagttcc atccaaaaaa tttaaagtac tgggcaacac cgaccgtcac 5820
tctatcaaaa aaaacctgat tggtgctctg ctgtttgaca gcggcgaaac tgctgaggct 5880
acccgtctga aacgtacggc tcgccgtcgc tacactcgtc gtaaaaaccg catctgttat 5940
ctgcaggaaa ttttctctaa cgaaatggca aaagttgatg atagcttctt tcatcgtctg 6000
gaagagagct tcctggtgga agaagataaa aaacacgaac gtcacccgat tttcggtaac 6060
attgtggatg aggttgccta ccacgagaaa tatccgacca tctaccatct gcgtaaaaaa 6120
ctggttgata gcactgacaa agcggatctg cgtctgatct acctggctct ggcacacatg 6180
atcaaattcc gtggtcactt cctgatcgaa ggtgatctga accctgataa ctccgacgtg 6240
gacaaactgt tcattcagct ggttcagacc tataaccagc tgttcgaaga aaacccgatc 6300
aacgcgtccg gtgtagacgc taaggcaatt ctgtctgcgc gtctgtctaa gtctcgtcgt 6360
ctggaaaacc tgattgcgca actgccaggt gaaaagaaaa acggcctgtt cggcaatctg 6420
atcgccctgt ccctgggtct gactccgaac tttaaatcca actttgacct ggcggaagat 6480
gccaagctgc agctgagcaa agatacctat gacgatgacc tggataacct gctggcacag 6540
atcggtgatc agtatgccga tctgttcctg gccgcgaaaa acctgtctga tgcgattctg 6600
ctgtctgata tcctgcgcgt taacactgaa attactaaag cgccgctgag cgcatccatg 6660
attaaacgtt acgatgaaca ccaccaggat ctgaccctgc tgaaagcgct ggtgcgtcag 6720
cagctgccgg aaaaatacaa ggagatcttc ttcgaccaga gcaaaaacgg ttacgcgggc 6780
tacattgatg gtggtgcatc tcaggaggaa ttctacaaat tcattaaacc gatcctggaa 6840
aaaatggatg gtactgaaga gctgctggtt aaactgaatc gtgaagatct gctgcgcaaa 6900
cagcgtacct tcgataacgg ttccatcccg catcagattc atctgggcga actgcacgct 6960
atcctgcgcc gtcaggaaga cttttatccg ttcctgaaag acaaccgtga gaaaattgaa 7020
aaaatcctga ccttccgtat tccgtactat gtaggtccgc tggcgcgtgg taactcccgt 7080
ttcgcttgga tgacccgcaa aagcgaagaa accatcaccc cgtggaattt cgaagaagtc 7140
gttgacaaag gcgcgtccgc gcagtctttc atcgaacgca tgacgaactt cgacaaaaac 7200
ctgccgaacg agaaagtgct gccgaaacac tctctgctgt acgagtactt cactgtgtac 7260
aacgaactga ccaaagtgaa atacgtcacc gaaggtatgc gtaaaccggc attcctgtcc 7320
ggtgagcaaa aaaaagcaat cgtggatctg ctgttcaaaa ccaaccgtaa agtaaccgtg 7380
aaacagctga aggaagacta tttcaagaaa atcgaatgtt ttgattctgt tgaaatctcc 7440
ggcgtggaag atcgcttcaa tgcgtccctg ggtacgtatc acgacctgct gaaaattatc 7500
aaagacaaag attttctgga caacgaggaa aacgaagaca tcctggagga tattgtactg 7560
accctgaccc tgttcgaaga ccgtgagatg atcgaagaac gcctgaaaac ctacgcccac 7620
ctgttcgatg acaaggtaat gaagcagctg aaacgtcgtc gttataccgg ctggggtcgt 7680
ctgtcccgta aactgatcaa tggcatccgt gataaacagt ctggcaaaac catcctggac 7740
ttcctgaaat ccgacggttt cgcgaatcgt aacttcatgc aactgattca tgacgattct 7800
ctgactttca aagaagacat ccagaaagca caggtttccg gccagggtga ctctctgcac 7860
gagcacattg ccaatctggc tggttctccg gctattaaaa agggtattct gcagactgtg 7920
aaagtagttg atgagctggt caaagtaatg ggccgtcaca agccggaaaa cattgtgatc 7980
gaaatggcac gtgaaaacca gacgacccag aaaggtcaga aaaactctcg tgaacgcatg 8040
aaacgtatcg aagaaggcat caaagaactg ggctctcaga tcctgaagga acaccctgta 8100
gaaaataccc agctgcagaa cgaaaagctg tatctgtatt acctgcagaa cggccgcgat 8160
atgtatgtgg accaggaact ggatatcaac cgcctgtccg attacgatgt agatcacatc 8220
gtgccgcaaa gcttcctgaa agacgacagc attgacaaca aagtactgac ccgttctgat 8280
aagaaccgtg gcaaatccga taacgtcccg tctgaagaag ttgttaaaaa aatgaaaaac 8340
tattggcgtc agctgctgaa cgcgaaactg atcacccagc gtaagttcga caatctgact 8400
aaagctgagc gcggtggtct gtccgaactg gataaagcgg gttttatcaa acgccagctg 8460
gttgaaaccc gtcagatcac gaagcacgtt gcgcagattc tggactctcg tatgaacacc 8520
aaatacgacg aaaacgacaa actgatccgc gaggttaagg ttatcaccct gaaaagcaaa 8580
ctggtatccg attttcgtaa agactttcag ttctacaaag tgcgcgaaat taacaactat 8640
caccacgctc acgatgcata tctgaatgca gttgttggca cggcgctgat caaaaagtat 8700
ccgaaactgg aatctgaatt cgtatacggc gattacaaag tgtatgacgt tcgtaagatg 8760
atcgcaaaat ccgagcagga aattggtaag gcgacggcga aatacttctt ttattccaat 8820
attatgaact ttttcaaaac cgaaatcacc ctggcgaatg gtgaaattcg taaacgcccg 8880
ctgatcgaaa ccaacggtga aactggtgaa atcgtttggg acaaaggccg cgacttcgcg 8940
accgtgcgta aagttctgtc tatgccgcaa gtgaacatcg tcaagaagac cgaagtacaa 9000
accggcggtt ttagcaaaga gagcattctg ccaaaacgta actccgacaa actgatcgcg 9060
cgcaagaaag actgggatcc gaaaaaatac ggtggtttcg attctccaac cgttgcttat 9120
tccgttctgg tggtagccaa agttgagaaa ggtaaaagca aaaaactgaa atccgtaaag 9180
gaactgctgg gtattactat catggagcgt agctccttcg aaaaaaaccc gatcgatttt 9240
ctggaagcga aaggctataa agaagtcaaa aaggacctga tcatcaaact gccaaaatac 9300
agcctgttcg agctggaaaa cggccgtaaa cgtatgctgg catctgcggg cgaactgcag 9360
aaaggcaacg agctggctct gccgtccaaa tacgtgaact ttctgtacct ggcctctcac 9420
tacgaaaaac tgaaaggttc cccggaagac aacgaacaga aacagctgtt cgtagagcag 9480
cacaaacact acctggacga gatcatcgaa cagatttctg aattttctaa acgtgtgatt 9540
ctggctgatg cgaatctgga taaagttctg tctgcctata acaagcatcg tgacaaaccg 9600
atccgcgaac aggctgagaa catcatccac ctgttcactc tgactaacct gggcgcgcca 9660
gcggctttca agtactttga taccaccatt gaccgcaagc gttacacctc cactaaagaa 9720
gtgctggacg cgactctgat ccaccagtcc atcaccggtc tgtacgagac ccgtatcgat 9780
ctgagccagc tgggcggtga caaaaggccg gcggccacga aaaaggccgg ccaggcaaaa 9840
aagaaaaagt gacaaagccc gaaaggaagc tgagttggct gctgccaccg ctgagcaata 9900
actagcataa ccccttgggg cctctaaacg ggtcttgagg ggttttttgc tgaaaggagg 9960
aactatatcc ggat 9974
<210> 2
<211> 1547
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 2
Met Lys Gln Ser Thr Ile Ala Leu Ala Leu Leu Pro Leu Leu Phe Thr
1 5 10 15
Pro Val Thr Lys Ala Met Ser Asp Lys Ile Ile His Leu Thr Asp Asp
20 25 30
Ser Phe Asp Thr Asp Val Leu Lys Ala Asp Gly Ala Ile Leu Val Asp
35 40 45
Phe Trp Ala Glu Trp Cys Gly Pro Cys Lys Met Ile Ala Pro Ile Leu
50 55 60
Asp Glu Ile Ala Asp Glu Tyr Gln Gly Lys Leu Thr Val Ala Lys Leu
65 70 75 80
Asn Ile Asp Gln Asn Pro Gly Thr Ala Pro Lys Tyr Gly Ile Arg Gly
85 90 95
Ile Pro Thr Leu Leu Leu Phe Lys Asn Gly Glu Val Ala Ala Thr Lys
100 105 110
Val Gly Ala Leu Ser Lys Gly Gln Leu Lys Glu Phe Leu Asp Ala Asn
115 120 125
Leu Ala Gly Ser Gly Ser Gly His Met His His His His His His Asp
130 135 140
Asp Asp Asp Lys Met Pro Lys Lys Lys Arg Lys Val Gly Ile His Gly
145 150 155 160
Val Pro Ala Ala Asp Lys Lys Tyr Ser Ile Gly Leu Asp Ile Gly Thr
165 170 175
Asn Ser Val Gly Trp Ala Val Ile Thr Asp Glu Tyr Lys Val Pro Ser
180 185 190
Lys Lys Phe Lys Val Leu Gly Asn Thr Asp Arg His Ser Ile Lys Lys
195 200 205
Asn Leu Ile Gly Ala Leu Leu Phe Asp Ser Gly Glu Thr Ala Glu Ala
210 215 220
Thr Arg Leu Lys Arg Thr Ala Arg Arg Arg Tyr Thr Arg Arg Lys Asn
225 230 235 240
Arg Ile Cys Tyr Leu Gln Glu Ile Phe Ser Asn Glu Met Ala Lys Val
245 250 255
Asp Asp Ser Phe Phe His Arg Leu Glu Glu Ser Phe Leu Val Glu Glu
260 265 270
Asp Lys Lys His Glu Arg His Pro Ile Phe Gly Asn Ile Val Asp Glu
275 280 285
Val Ala Tyr His Glu Lys Tyr Pro Thr Ile Tyr His Leu Arg Lys Lys
290 295 300
Leu Val Asp Ser Thr Asp Lys Ala Asp Leu Arg Leu Ile Tyr Leu Ala
305 310 315 320
Leu Ala His Met Ile Lys Phe Arg Gly His Phe Leu Ile Glu Gly Asp
325 330 335
Leu Asn Pro Asp Asn Ser Asp Val Asp Lys Leu Phe Ile Gln Leu Val
340 345 350
Gln Thr Tyr Asn Gln Leu Phe Glu Glu Asn Pro Ile Asn Ala Ser Gly
355 360 365
Val Asp Ala Lys Ala Ile Leu Ser Ala Arg Leu Ser Lys Ser Arg Arg
370 375 380
Leu Glu Asn Leu Ile Ala Gln Leu Pro Gly Glu Lys Lys Asn Gly Leu
385 390 395 400
Phe Gly Asn Leu Ile Ala Leu Ser Leu Gly Leu Thr Pro Asn Phe Lys
405 410 415
Ser Asn Phe Asp Leu Ala Glu Asp Ala Lys Leu Gln Leu Ser Lys Asp
420 425 430
Thr Tyr Asp Asp Asp Leu Asp Asn Leu Leu Ala Gln Ile Gly Asp Gln
435 440 445
Tyr Ala Asp Leu Phe Leu Ala Ala Lys Asn Leu Ser Asp Ala Ile Leu
450 455 460
Leu Ser Asp Ile Leu Arg Val Asn Thr Glu Ile Thr Lys Ala Pro Leu
465 470 475 480
Ser Ala Ser Met Ile Lys Arg Tyr Asp Glu His His Gln Asp Leu Thr
485 490 495
Leu Leu Lys Ala Leu Val Arg Gln Gln Leu Pro Glu Lys Tyr Lys Glu
500 505 510
Ile Phe Phe Asp Gln Ser Lys Asn Gly Tyr Ala Gly Tyr Ile Asp Gly
515 520 525
Gly Ala Ser Gln Glu Glu Phe Tyr Lys Phe Ile Lys Pro Ile Leu Glu
530 535 540
Lys Met Asp Gly Thr Glu Glu Leu Leu Val Lys Leu Asn Arg Glu Asp
545 550 555 560
Leu Leu Arg Lys Gln Arg Thr Phe Asp Asn Gly Ser Ile Pro His Gln
565 570 575
Ile His Leu Gly Glu Leu His Ala Ile Leu Arg Arg Gln Glu Asp Phe
580 585 590
Tyr Pro Phe Leu Lys Asp Asn Arg Glu Lys Ile Glu Lys Ile Leu Thr
595 600 605
Phe Arg Ile Pro Tyr Tyr Val Gly Pro Leu Ala Arg Gly Asn Ser Arg
610 615 620
Phe Ala Trp Met Thr Arg Lys Ser Glu Glu Thr Ile Thr Pro Trp Asn
625 630 635 640
Phe Glu Glu Val Val Asp Lys Gly Ala Ser Ala Gln Ser Phe Ile Glu
645 650 655
Arg Met Thr Asn Phe Asp Lys Asn Leu Pro Asn Glu Lys Val Leu Pro
660 665 670
Lys His Ser Leu Leu Tyr Glu Tyr Phe Thr Val Tyr Asn Glu Leu Thr
675 680 685
Lys Val Lys Tyr Val Thr Glu Gly Met Arg Lys Pro Ala Phe Leu Ser
690 695 700
Gly Glu Gln Lys Lys Ala Ile Val Asp Leu Leu Phe Lys Thr Asn Arg
705 710 715 720
Lys Val Thr Val Lys Gln Leu Lys Glu Asp Tyr Phe Lys Lys Ile Glu
725 730 735
Cys Phe Asp Ser Val Glu Ile Ser Gly Val Glu Asp Arg Phe Asn Ala
740 745 750
Ser Leu Gly Thr Tyr His Asp Leu Leu Lys Ile Ile Lys Asp Lys Asp
755 760 765
Phe Leu Asp Asn Glu Glu Asn Glu Asp Ile Leu Glu Asp Ile Val Leu
770 775 780
Thr Leu Thr Leu Phe Glu Asp Arg Glu Met Ile Glu Glu Arg Leu Lys
785 790 795 800
Thr Tyr Ala His Leu Phe Asp Asp Lys Val Met Lys Gln Leu Lys Arg
805 810 815
Arg Arg Tyr Thr Gly Trp Gly Arg Leu Ser Arg Lys Leu Ile Asn Gly
820 825 830
Ile Arg Asp Lys Gln Ser Gly Lys Thr Ile Leu Asp Phe Leu Lys Ser
835 840 845
Asp Gly Phe Ala Asn Arg Asn Phe Met Gln Leu Ile His Asp Asp Ser
850 855 860
Leu Thr Phe Lys Glu Asp Ile Gln Lys Ala Gln Val Ser Gly Gln Gly
865 870 875 880
Asp Ser Leu His Glu His Ile Ala Asn Leu Ala Gly Ser Pro Ala Ile
885 890 895
Lys Lys Gly Ile Leu Gln Thr Val Lys Val Val Asp Glu Leu Val Lys
900 905 910
Val Met Gly Arg His Lys Pro Glu Asn Ile Val Ile Glu Met Ala Arg
915 920 925
Glu Asn Gln Thr Thr Gln Lys Gly Gln Lys Asn Ser Arg Glu Arg Met
930 935 940
Lys Arg Ile Glu Glu Gly Ile Lys Glu Leu Gly Ser Gln Ile Leu Lys
945 950 955 960
Glu His Pro Val Glu Asn Thr Gln Leu Gln Asn Glu Lys Leu Tyr Leu
965 970 975
Tyr Tyr Leu Gln Asn Gly Arg Asp Met Tyr Val Asp Gln Glu Leu Asp
980 985 990
Ile Asn Arg Leu Ser Asp Tyr Asp Val Asp His Ile Val Pro Gln Ser
995 1000 1005
Phe Leu Lys Asp Asp Ser Ile Asp Asn Lys Val Leu Thr Arg Ser Asp
1010 1015 1020
Lys Asn Arg Gly Lys Ser Asp Asn Val Pro Ser Glu Glu Val Val Lys
1025 1030 1035 1040
Lys Met Lys Asn Tyr Trp Arg Gln Leu Leu Asn Ala Lys Leu Ile Thr
1045 1050 1055
Gln Arg Lys Phe Asp Asn Leu Thr Lys Ala Glu Arg Gly Gly Leu Ser
1060 1065 1070
Glu Leu Asp Lys Ala Gly Phe Ile Lys Arg Gln Leu Val Glu Thr Arg
1075 1080 1085
Gln Ile Thr Lys His Val Ala Gln Ile Leu Asp Ser Arg Met Asn Thr
1090 1095 1100
Lys Tyr Asp Glu Asn Asp Lys Leu Ile Arg Glu Val Lys Val Ile Thr
1105 1110 1115 1120
Leu Lys Ser Lys Leu Val Ser Asp Phe Arg Lys Asp Phe Gln Phe Tyr
1125 1130 1135
Lys Val Arg Glu Ile Asn Asn Tyr His His Ala His Asp Ala Tyr Leu
1140 1145 1150
Asn Ala Val Val Gly Thr Ala Leu Ile Lys Lys Tyr Pro Lys Leu Glu
1155 1160 1165
Ser Glu Phe Val Tyr Gly Asp Tyr Lys Val Tyr Asp Val Arg Lys Met
1170 1175 1180
Ile Ala Lys Ser Glu Gln Glu Ile Gly Lys Ala Thr Ala Lys Tyr Phe
1185 1190 1195 1200
Phe Tyr Ser Asn Ile Met Asn Phe Phe Lys Thr Glu Ile Thr Leu Ala
1205 1210 1215
Asn Gly Glu Ile Arg Lys Arg Pro Leu Ile Glu Thr Asn Gly Glu Thr
1220 1225 1230
Gly Glu Ile Val Trp Asp Lys Gly Arg Asp Phe Ala Thr Val Arg Lys
1235 1240 1245
Val Leu Ser Met Pro Gln Val Asn Ile Val Lys Lys Thr Glu Val Gln
1250 1255 1260
Thr Gly Gly Phe Ser Lys Glu Ser Ile Leu Pro Lys Arg Asn Ser Asp
1265 1270 1275 1280
Lys Leu Ile Ala Arg Lys Lys Asp Trp Asp Pro Lys Lys Tyr Gly Gly
1285 1290 1295
Phe Asp Ser Pro Thr Val Ala Tyr Ser Val Leu Val Val Ala Lys Val
1300 1305 1310
Glu Lys Gly Lys Ser Lys Lys Leu Lys Ser Val Lys Glu Leu Leu Gly
1315 1320 1325
Ile Thr Ile Met Glu Arg Ser Ser Phe Glu Lys Asn Pro Ile Asp Phe
1330 1335 1340
Leu Glu Ala Lys Gly Tyr Lys Glu Val Lys Lys Asp Leu Ile Ile Lys
1345 1350 1355 1360
Leu Pro Lys Tyr Ser Leu Phe Glu Leu Glu Asn Gly Arg Lys Arg Met
1365 1370 1375
Leu Ala Ser Ala Gly Glu Leu Gln Lys Gly Asn Glu Leu Ala Leu Pro
1380 1385 1390
Ser Lys Tyr Val Asn Phe Leu Tyr Leu Ala Ser His Tyr Glu Lys Leu
1395 1400 1405
Lys Gly Ser Pro Glu Asp Asn Glu Gln Lys Gln Leu Phe Val Glu Gln
1410 1415 1420
His Lys His Tyr Leu Asp Glu Ile Ile Glu Gln Ile Ser Glu Phe Ser
1425 1430 1435 1440
Lys Arg Val Ile Leu Ala Asp Ala Asn Leu Asp Lys Val Leu Ser Ala
1445 1450 1455
Tyr Asn Lys His Arg Asp Lys Pro Ile Arg Glu Gln Ala Glu Asn Ile
1460 1465 1470
Ile His Leu Phe Thr Leu Thr Asn Leu Gly Ala Pro Ala Ala Phe Lys
1475 1480 1485
Tyr Phe Asp Thr Thr Ile Asp Arg Lys Arg Tyr Thr Ser Thr Lys Glu
1490 1495 1500
Val Leu Asp Ala Thr Leu Ile His Gln Ser Ile Thr Gly Leu Tyr Glu
1505 1510 1515 1520
Thr Arg Ile Asp Leu Ser Gln Leu Gly Gly Asp Lys Arg Pro Ala Ala
1525 1530 1535
Thr Lys Lys Ala Gly Gln Ala Lys Lys Lys Lys
1540 1545
<210> 3
<211> 1399
<212> PRT
<213> 人工序列(Artificial Sequence)
<400> 3
Met Pro Lys Lys Lys Arg Lys Val Gly Ile His Gly Val Pro Ala Ala
1 5 10 15
Asp Lys Lys Tyr Ser Ile Gly Leu Asp Ile Gly Thr Asn Ser Val Gly
20 25 30
Trp Ala Val Ile Thr Asp Glu Tyr Lys Val Pro Ser Lys Lys Phe Lys
35 40 45
Val Leu Gly Asn Thr Asp Arg His Ser Ile Lys Lys Asn Leu Ile Gly
50 55 60
Ala Leu Leu Phe Asp Ser Gly Glu Thr Ala Glu Ala Thr Arg Leu Lys
65 70 75 80
Arg Thr Ala Arg Arg Arg Tyr Thr Arg Arg Lys Asn Arg Ile Cys Tyr
85 90 95
Leu Gln Glu Ile Phe Ser Asn Glu Met Ala Lys Val Asp Asp Ser Phe
100 105 110
Phe His Arg Leu Glu Glu Ser Phe Leu Val Glu Glu Asp Lys Lys His
115 120 125
Glu Arg His Pro Ile Phe Gly Asn Ile Val Asp Glu Val Ala Tyr His
130 135 140
Glu Lys Tyr Pro Thr Ile Tyr His Leu Arg Lys Lys Leu Val Asp Ser
145 150 155 160
Thr Asp Lys Ala Asp Leu Arg Leu Ile Tyr Leu Ala Leu Ala His Met
165 170 175
Ile Lys Phe Arg Gly His Phe Leu Ile Glu Gly Asp Leu Asn Pro Asp
180 185 190
Asn Ser Asp Val Asp Lys Leu Phe Ile Gln Leu Val Gln Thr Tyr Asn
195 200 205
Gln Leu Phe Glu Glu Asn Pro Ile Asn Ala Ser Gly Val Asp Ala Lys
210 215 220
Ala Ile Leu Ser Ala Arg Leu Ser Lys Ser Arg Arg Leu Glu Asn Leu
225 230 235 240
Ile Ala Gln Leu Pro Gly Glu Lys Lys Asn Gly Leu Phe Gly Asn Leu
245 250 255
Ile Ala Leu Ser Leu Gly Leu Thr Pro Asn Phe Lys Ser Asn Phe Asp
260 265 270
Leu Ala Glu Asp Ala Lys Leu Gln Leu Ser Lys Asp Thr Tyr Asp Asp
275 280 285
Asp Leu Asp Asn Leu Leu Ala Gln Ile Gly Asp Gln Tyr Ala Asp Leu
290 295 300
Phe Leu Ala Ala Lys Asn Leu Ser Asp Ala Ile Leu Leu Ser Asp Ile
305 310 315 320
Leu Arg Val Asn Thr Glu Ile Thr Lys Ala Pro Leu Ser Ala Ser Met
325 330 335
Ile Lys Arg Tyr Asp Glu His His Gln Asp Leu Thr Leu Leu Lys Ala
340 345 350
Leu Val Arg Gln Gln Leu Pro Glu Lys Tyr Lys Glu Ile Phe Phe Asp
355 360 365
Gln Ser Lys Asn Gly Tyr Ala Gly Tyr Ile Asp Gly Gly Ala Ser Gln
370 375 380
Glu Glu Phe Tyr Lys Phe Ile Lys Pro Ile Leu Glu Lys Met Asp Gly
385 390 395 400
Thr Glu Glu Leu Leu Val Lys Leu Asn Arg Glu Asp Leu Leu Arg Lys
405 410 415
Gln Arg Thr Phe Asp Asn Gly Ser Ile Pro His Gln Ile His Leu Gly
420 425 430
Glu Leu His Ala Ile Leu Arg Arg Gln Glu Asp Phe Tyr Pro Phe Leu
435 440 445
Lys Asp Asn Arg Glu Lys Ile Glu Lys Ile Leu Thr Phe Arg Ile Pro
450 455 460
Tyr Tyr Val Gly Pro Leu Ala Arg Gly Asn Ser Arg Phe Ala Trp Met
465 470 475 480
Thr Arg Lys Ser Glu Glu Thr Ile Thr Pro Trp Asn Phe Glu Glu Val
485 490 495
Val Asp Lys Gly Ala Ser Ala Gln Ser Phe Ile Glu Arg Met Thr Asn
500 505 510
Phe Asp Lys Asn Leu Pro Asn Glu Lys Val Leu Pro Lys His Ser Leu
515 520 525
Leu Tyr Glu Tyr Phe Thr Val Tyr Asn Glu Leu Thr Lys Val Lys Tyr
530 535 540
Val Thr Glu Gly Met Arg Lys Pro Ala Phe Leu Ser Gly Glu Gln Lys
545 550 555 560
Lys Ala Ile Val Asp Leu Leu Phe Lys Thr Asn Arg Lys Val Thr Val
565 570 575
Lys Gln Leu Lys Glu Asp Tyr Phe Lys Lys Ile Glu Cys Phe Asp Ser
580 585 590
Val Glu Ile Ser Gly Val Glu Asp Arg Phe Asn Ala Ser Leu Gly Thr
595 600 605
Tyr His Asp Leu Leu Lys Ile Ile Lys Asp Lys Asp Phe Leu Asp Asn
610 615 620
Glu Glu Asn Glu Asp Ile Leu Glu Asp Ile Val Leu Thr Leu Thr Leu
625 630 635 640
Phe Glu Asp Arg Glu Met Ile Glu Glu Arg Leu Lys Thr Tyr Ala His
645 650 655
Leu Phe Asp Asp Lys Val Met Lys Gln Leu Lys Arg Arg Arg Tyr Thr
660 665 670
Gly Trp Gly Arg Leu Ser Arg Lys Leu Ile Asn Gly Ile Arg Asp Lys
675 680 685
Gln Ser Gly Lys Thr Ile Leu Asp Phe Leu Lys Ser Asp Gly Phe Ala
690 695 700
Asn Arg Asn Phe Met Gln Leu Ile His Asp Asp Ser Leu Thr Phe Lys
705 710 715 720
Glu Asp Ile Gln Lys Ala Gln Val Ser Gly Gln Gly Asp Ser Leu His
725 730 735
Glu His Ile Ala Asn Leu Ala Gly Ser Pro Ala Ile Lys Lys Gly Ile
740 745 750
Leu Gln Thr Val Lys Val Val Asp Glu Leu Val Lys Val Met Gly Arg
755 760 765
His Lys Pro Glu Asn Ile Val Ile Glu Met Ala Arg Glu Asn Gln Thr
770 775 780
Thr Gln Lys Gly Gln Lys Asn Ser Arg Glu Arg Met Lys Arg Ile Glu
785 790 795 800
Glu Gly Ile Lys Glu Leu Gly Ser Gln Ile Leu Lys Glu His Pro Val
805 810 815
Glu Asn Thr Gln Leu Gln Asn Glu Lys Leu Tyr Leu Tyr Tyr Leu Gln
820 825 830
Asn Gly Arg Asp Met Tyr Val Asp Gln Glu Leu Asp Ile Asn Arg Leu
835 840 845
Ser Asp Tyr Asp Val Asp His Ile Val Pro Gln Ser Phe Leu Lys Asp
850 855 860
Asp Ser Ile Asp Asn Lys Val Leu Thr Arg Ser Asp Lys Asn Arg Gly
865 870 875 880
Lys Ser Asp Asn Val Pro Ser Glu Glu Val Val Lys Lys Met Lys Asn
885 890 895
Tyr Trp Arg Gln Leu Leu Asn Ala Lys Leu Ile Thr Gln Arg Lys Phe
900 905 910
Asp Asn Leu Thr Lys Ala Glu Arg Gly Gly Leu Ser Glu Leu Asp Lys
915 920 925
Ala Gly Phe Ile Lys Arg Gln Leu Val Glu Thr Arg Gln Ile Thr Lys
930 935 940
His Val Ala Gln Ile Leu Asp Ser Arg Met Asn Thr Lys Tyr Asp Glu
945 950 955 960
Asn Asp Lys Leu Ile Arg Glu Val Lys Val Ile Thr Leu Lys Ser Lys
965 970 975
Leu Val Ser Asp Phe Arg Lys Asp Phe Gln Phe Tyr Lys Val Arg Glu
980 985 990
Ile Asn Asn Tyr His His Ala His Asp Ala Tyr Leu Asn Ala Val Val
995 1000 1005
Gly Thr Ala Leu Ile Lys Lys Tyr Pro Lys Leu Glu Ser Glu Phe Val
1010 1015 1020
Tyr Gly Asp Tyr Lys Val Tyr Asp Val Arg Lys Met Ile Ala Lys Ser
1025 1030 1035 1040
Glu Gln Glu Ile Gly Lys Ala Thr Ala Lys Tyr Phe Phe Tyr Ser Asn
1045 1050 1055
Ile Met Asn Phe Phe Lys Thr Glu Ile Thr Leu Ala Asn Gly Glu Ile
1060 1065 1070
Arg Lys Arg Pro Leu Ile Glu Thr Asn Gly Glu Thr Gly Glu Ile Val
1075 1080 1085
Trp Asp Lys Gly Arg Asp Phe Ala Thr Val Arg Lys Val Leu Ser Met
1090 1095 1100
Pro Gln Val Asn Ile Val Lys Lys Thr Glu Val Gln Thr Gly Gly Phe
1105 1110 1115 1120
Ser Lys Glu Ser Ile Leu Pro Lys Arg Asn Ser Asp Lys Leu Ile Ala
1125 1130 1135
Arg Lys Lys Asp Trp Asp Pro Lys Lys Tyr Gly Gly Phe Asp Ser Pro
1140 1145 1150
Thr Val Ala Tyr Ser Val Leu Val Val Ala Lys Val Glu Lys Gly Lys
1155 1160 1165
Ser Lys Lys Leu Lys Ser Val Lys Glu Leu Leu Gly Ile Thr Ile Met
1170 1175 1180
Glu Arg Ser Ser Phe Glu Lys Asn Pro Ile Asp Phe Leu Glu Ala Lys
1185 1190 1195 1200
Gly Tyr Lys Glu Val Lys Lys Asp Leu Ile Ile Lys Leu Pro Lys Tyr
1205 1210 1215
Ser Leu Phe Glu Leu Glu Asn Gly Arg Lys Arg Met Leu Ala Ser Ala
1220 1225 1230
Gly Glu Leu Gln Lys Gly Asn Glu Leu Ala Leu Pro Ser Lys Tyr Val
1235 1240 1245
Asn Phe Leu Tyr Leu Ala Ser His Tyr Glu Lys Leu Lys Gly Ser Pro
1250 1255 1260
Glu Asp Asn Glu Gln Lys Gln Leu Phe Val Glu Gln His Lys His Tyr
1265 1270 1275 1280
Leu Asp Glu Ile Ile Glu Gln Ile Ser Glu Phe Ser Lys Arg Val Ile
1285 1290 1295
Leu Ala Asp Ala Asn Leu Asp Lys Val Leu Ser Ala Tyr Asn Lys His
1300 1305 1310
Arg Asp Lys Pro Ile Arg Glu Gln Ala Glu Asn Ile Ile His Leu Phe
1315 1320 1325
Thr Leu Thr Asn Leu Gly Ala Pro Ala Ala Phe Lys Tyr Phe Asp Thr
1330 1335 1340
Thr Ile Asp Arg Lys Arg Tyr Thr Ser Thr Lys Glu Val Leu Asp Ala
1345 1350 1355 1360
Thr Leu Ile His Gln Ser Ile Thr Gly Leu Tyr Glu Thr Arg Ile Asp
1365 1370 1375
Leu Ser Gln Leu Gly Gly Asp Lys Arg Pro Ala Ala Thr Lys Lys Ala
1380 1385 1390
Gly Gln Ala Lys Lys Lys Lys
1395
<210> 4
<211> 225
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 4
ggcttgtcgg actcttcgct attacgccag ctggcgaagg gggatgtgct gcaaggcgat 60
taagttgggt aacgccaggg ttttcccagt cacgacgtta ggaaattaat acgactcact 120
ataggagagc acagtcagcc tggcggtttt agagctagaa atagcaagtt aaaataaggc 180
tagtccgtta tcaacttgaa aaagtggcac cgagtcggtg ctttt 225
<210> 5
<211> 225
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
ggcttgtcgg actcttcgct attacgccag ctggcgaagg gggatgtgct gcaaggcgat 60
taagttgggt aacgccaggg ttttcccagt cacgacgtta ggaaattaat acgactcact 120
ataggcttcc agaattggat ctccggtttt agagctagaa atagcaagtt aaaataaggc 180
tagtccgtta tcaacttgaa aaagtggcac cgagtcggtg ctttt 225
<210> 6
<211> 102
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 6
ggagagcaca gucagccugg cgguuuuaga gcuagaaaua gcaaguuaaa auaaggcuag 60
uccguuauca acuugaaaaa guggcaccga gucggugcuu uu 102
<210> 7
<211> 102
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 7
ggcuuccaga auuggaucuc cgguuuuaga gcuagaaaua gcaaguuaaa auaaggcuag 60
uccguuauca acuugaaaaa guggcaccga gucggugcuu uu 102
<210> 8
<211> 317
<212> PRT
<213> Sus scrofa
<400> 8
Met Arg Val Leu Trp Val Ala Leu Val Val Thr Leu Leu Ala Gly Cys
1 5 10 15
Arg Thr Glu Asp Glu Pro Gly Pro Pro Pro Glu Val His Val Trp Trp
20 25 30
Glu Glu Pro Lys Trp Gln Gly Ser Gln Pro Trp Glu Gln Ala Leu Gly
35 40 45
Arg Phe Trp Asp Tyr Leu Arg Trp Val Gln Ser Leu Ser Asp Gln Val
50 55 60
Gln Glu Glu Leu Leu Ser Thr Lys Val Thr Gln Glu Leu Thr Glu Leu
65 70 75 80
Ile Glu Glu Ser Met Lys Glu Val Lys Ala Tyr Arg Glu Glu Leu Glu
85 90 95
Ala Gln Leu Gly Pro Val Thr Gln Glu Thr Gln Ala Arg Leu Ser Lys
100 105 110
Glu Leu Gln Ala Ala Gln Ala Arg Val Gly Ala Asp Met Glu Asp Val
115 120 125
Arg Asn Arg Leu Val Leu Tyr Arg Ser Glu Val His Asn Met Leu Gly
130 135 140
Gln Thr Thr Glu Glu Leu Arg Ser Arg Leu Ala Ser His Leu Arg Asn
145 150 155 160
Val Arg Lys Arg Leu Val Arg Asp Thr Glu Asp Leu Gln Lys Arg Leu
165 170 175
Ala Val Tyr Gln Ala Gly Leu Arg Glu Gly Ala Glu Arg Ser Val Ser
180 185 190
Ala Leu Arg Glu Arg Leu Gly Pro Leu Val Glu Gln Gly Arg Leu Arg
195 200 205
Ala Ala Thr Leu Ser Thr Arg Ala Gly Gln Pro Leu Arg Glu Arg Ala
210 215 220
Glu Ala Trp Gly Gln Lys Leu Arg Gly Arg Leu Glu Glu Met Gly Ser
225 230 235 240
Arg Thr Arg Asp Arg Leu Asp Glu Met Arg Asp Glu Leu Glu Glu Val
245 250 255
Arg Thr Lys Val Glu Glu Gln Gly Ser Gln Leu Arg Leu Gln Ala Glu
260 265 270
Ala Phe Gln Ala Arg Leu Lys Gly Trp Phe Glu Pro Leu Val Glu Asp
275 280 285
Met Arg Arg Gln Trp Ala Gly Leu Val Glu Arg Met Gln Ser Ala Val
290 295 300
Ser Ile Ser Ser Ser Thr Ser Ala Pro Ser Asp Asn Gln
305 310 315
<210> 9
<211> 890
<212> DNA
<213> Sus scrofa
<400> 9
ggacccctcg cctgggaact tccacgtatg ccactggtgc agccctaaaa gacaaacaaa 60
caaaaacgaa agaaagagaa aagaaaggaa agggggcttc tgtttctaat gcgttgttgc 120
ctggcagggc gtgagcatta gatacgtgtc agctgtgact agcgtgcacg gagcacacaa 180
tccatgcttg tccagtaatt agacaggctg ggtgtccttc cacccctccc tgcccaccag 240
tgctctagag aagcccaccc accggggctg ggggagcacc tgctctgtac caggtaccgt 300
gtgctgggag ggggcagagg acctgatggc tgtgaactgg ctcggtgcag gatgccggac 360
agaggacgag ccggggccgc cgccggaggt gcacgtgtgg tgggaggagc ccaagtggca 420
gggcagccag ccctgggagc aggccctggg ccgcttctgg gattacctgc gctgggtgca 480
gtccctgtct gaccaagtgc aggaggagct gctcagcacc aaggtcaccc aggaactgac 540
gtaagtgccc acccgactcc cgccgcgcgc gcgcgcgcgc gcgcgcgcct gaccctcctg 600
gcgaaccgtg tgttctggac cctcaggctc cacccgtccg ggtttccttc tgtccttgtc 660
gccaactctt gggggtctgg gtctctgttt cttttttttc cttcttcctt ttttgggggg 720
agtttacttt ttcttttttc tttcatttga cttcatgtct tgctttcttt ccatcttgag 780
ctcctgcctt cgcctgtctc tgggtcagtc ttgccgtcct tgctgtctct gaatctctgg 840
cacgtcctgg ccatcgccag ctcaggagcc ctccttctcc ccctccccgc 890
<210> 10
<211> 100
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 10
guaaucccag aagcggccca guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu 100
<210> 11
<211> 100
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 11
ugugguggga ggagcccaag guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu 100
<210> 12
<211> 100
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 12
ccugucugac caagugcagg guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu 100
<210> 13
<211> 100
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 13
caccacacgu gcaccuccgg guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu 100
<210> 14
<211> 2871
<212> PRT
<213> Sus scrofa
<400> 14
Met Arg Arg Gly Arg Leu Leu Glu Val Ala Leu Gly Phe Thr Val Leu
1 5 10 15
Leu Ala Ser Tyr Thr Ser His Arg Ala Glu Ala Asn Leu Glu Ala Gly
20 25 30
Asn Gly Lys Glu Thr Arg Ala Ser Arg Ala Lys Arg Arg Gly Gly Gly
35 40 45
Gly His Asp Ala Leu Lys Gly Pro Asn Val Cys Gly Ser Arg Tyr Asn
50 55 60
Ala Tyr Cys Cys Pro Gly Trp Lys Thr Leu Pro Gly Gly Asn Gln Cys
65 70 75 80
Ile Val Pro Ile Cys Arg His Ser Cys Gly Asp Gly Phe Cys Ser Arg
85 90 95
Pro Asn Met Cys Thr Cys Pro Ser Gly Gln Ile Ala Pro Ser Cys Gly
100 105 110
Ser Arg Ser Ile Gln His Cys Asn Ile Arg Cys Met Asn Gly Gly Ser
115 120 125
Cys Ser Asp Asp His Cys Leu Cys Gln Lys Gly Tyr Ile Gly Thr His
130 135 140
Cys Gly Gln Pro Val Cys Glu Ser Gly Cys Leu Asn Gly Gly Arg Cys
145 150 155 160
Val Ala Pro Asn Arg Cys Ala Cys Thr Tyr Gly Phe Thr Gly Pro Gln
165 170 175
Cys Glu Arg Asp Tyr Arg Thr Gly Pro Cys Phe Thr Val Val Ser Asn
180 185 190
Gln Met Cys Gln Gly Gln Leu Ser Gly Ile Val Cys Thr Lys Thr Leu
195 200 205
Cys Cys Ala Thr Val Gly Arg Ala Trp Gly His Pro Cys Glu Met Cys
210 215 220
Pro Ala Gln Pro His Pro Cys Arg Arg Gly Phe Ile Pro Asn Ile Arg
225 230 235 240
Thr Gly Ala Cys Gln Asp Val Asp Glu Cys Gln Ala Ile Pro Gly Leu
245 250 255
Cys Gln Gly Gly Asn Cys Ile Asn Thr Val Gly Ser Phe Glu Cys Lys
260 265 270
Cys Pro Ala Gly His Lys Phe Asn Glu Val Ser Gln Lys Cys Glu Asp
275 280 285
Ile Asp Glu Cys Ser Thr Ile Pro Gly Ile Cys Asp Gly Gly Glu Cys
290 295 300
Thr Asn Thr Val Ser Ser Tyr Phe Cys Lys Cys Pro Pro Gly Phe Tyr
305 310 315 320
Thr Ser Pro Asp Gly Thr Arg Cys Ile Asp Val Arg Pro Gly Tyr Cys
325 330 335
Tyr Thr Ala Leu Thr Asn Gly Arg Cys Ser Asn Gln Leu Pro Gln Ser
340 345 350
Ile Thr Lys Met Gln Cys Cys Cys Asp Val Gly Arg Cys Trp Ser Pro
355 360 365
Gly Val Thr Val Thr Pro Glu Met Cys Pro Ile Arg Ala Thr Glu Asp
370 375 380
Phe Asn Lys Leu Cys Ser Val Pro Met Val Val Pro Glu Arg Pro Gly
385 390 395 400
Tyr Pro Ser Pro Pro Leu Gly Pro Ile Pro Pro Val His Pro Val Pro
405 410 415
Pro Gly Phe Pro Pro Gly Pro Gln Ile Pro Val Pro Arg Pro Pro Val
420 425 430
Glu Tyr Pro Tyr Pro Ser Arg Glu Pro Pro Arg Val Leu Pro Val Asn
435 440 445
Val Thr Asp Tyr Cys Gln Leu Phe Arg Tyr Leu Cys His Asn Gly Arg
450 455 460
Cys Ile Pro Thr Pro Gly Ser Tyr Arg Cys Glu Cys Asn Lys Gly Phe
465 470 475 480
Gln Leu Asp Leu Arg Gly Glu Cys Ile Asp Val Asp Glu Cys Glu Lys
485 490 495
Asn Pro Cys Ala Gly Gly Glu Cys Ile Asn Asn Gln Gly Ser Tyr Thr
500 505 510
Cys Gln Cys Arg Pro Gly Tyr Gln Ser Thr Leu Thr Arg Thr Glu Cys
515 520 525
Arg Asp Ile Asp Glu Cys Leu Gln Asn Gly Arg Ile Cys Asn Asn Gly
530 535 540
Arg Cys Ile Asn Thr Asp Gly Ser Phe His Cys Val Cys Asn Ala Gly
545 550 555 560
Phe His Val Thr Arg Asp Gly Lys Asn Cys Glu Asp Met Asp Glu Cys
565 570 575
Ser Ile Arg Asn Met Cys Leu Asn Gly Met Cys Ile Asn Glu Asp Gly
580 585 590
Ser Phe Lys Cys Ile Cys Lys Pro Gly Phe Gln Leu Ala Ser Asp Gly
595 600 605
Arg Tyr Cys Lys Asp Ile Asn Glu Cys Glu Thr Ser Gly Ile Cys Met
610 615 620
Asn Gly Arg Cys Val Asn Thr Asp Gly Ser Tyr Arg Cys Glu Cys Phe
625 630 635 640
Pro Gly Leu Ala Val Gly Leu Asp Gly Arg Val Cys Val Asp Thr His
645 650 655
Met Arg Ser Thr Cys Tyr Gly Gly Tyr Lys Arg Gly Gln Cys Val Lys
660 665 670
Pro Leu Phe Gly Ala Val Thr Lys Ser Glu Cys Cys Cys Ala Ser Thr
675 680 685
Glu Tyr Ala Phe Gly Glu Pro Cys Gln Pro Cys Pro Ser Gln Asn Ser
690 695 700
Ala Glu Tyr Gln Ala Leu Cys Ser Ser Gly Pro Gly Met Thr Ser Ala
705 710 715 720
Gly Ser Asp Ile Asn Glu Cys Ala Leu Asp Pro Asp Ile Cys Pro Asn
725 730 735
Gly Ile Cys Glu Asn Leu Arg Gly Thr Tyr Lys Cys Ile Cys Asn Ser
740 745 750
Gly Tyr Glu Val Asp Ser Thr Gly Lys Asn Cys Val Asp Ile Asn Glu
755 760 765
Cys Val Leu Asn Ser Leu Leu Cys Asp Asn Gly Gln Cys Arg Asn Thr
770 775 780
Pro Gly Ser Phe Val Cys Thr Cys Pro Lys Gly Phe Ile Tyr Lys Pro
785 790 795 800
Asp Leu Lys Thr Cys Glu Asp Ile Asp Glu Cys Glu Ser Ser Pro Cys
805 810 815
Ile Asn Gly Val Cys Lys Asn Ser Pro Gly Ser Phe Ile Cys Glu Cys
820 825 830
Ser Ser Glu Ser Thr Leu Asp Pro Thr Lys Thr Ile Cys Ile Glu Thr
835 840 845
Ile Lys Gly Thr Cys Trp Gln Thr Ile Ile Asp Gly Arg Cys Glu Ile
850 855 860
Asn Ile Asn Gly Ala Thr Leu Lys Ser Gln Cys Cys Ser Ser Leu Gly
865 870 875 880
Ala Ala Trp Gly Ser Pro Cys Thr Pro Cys Gln Val Asp Pro Ile Cys
885 890 895
Gly Lys Gly Tyr Ser Arg Ile Lys Gly Thr Gln Cys Glu Asp Ile Asp
900 905 910
Glu Cys Glu Val Phe Pro Gly Val Cys Lys Asn Gly Leu Cys Val Asn
915 920 925
Ser Lys Gly Ser Phe Lys Cys Gln Cys Pro Asn Gly Met Thr Leu Asp
930 935 940
Ala Thr Gly Arg Ile Cys Leu Asp Ile Arg Leu Glu Thr Cys Phe Leu
945 950 955 960
Arg Tyr Glu Asp Glu Glu Cys Thr Leu Pro Val Val Gly Arg His Arg
965 970 975
Met Asp Ala Cys Cys Cys Ser Val Gly Ala Ala Trp Gly Thr Glu Glu
980 985 990
Cys Glu Glu Cys Pro Pro Arg Asn Thr Pro Glu Tyr Glu Glu Leu Cys
995 1000 1005
Pro Arg Gly Pro Gly Phe Ala Thr Lys Glu Ile Thr Asn Gly Lys Pro
1010 1015 1020
Phe Phe Lys Asp Ile Asn Glu Cys Lys Met Ile Pro Asn Leu Cys Thr
1025 1030 1035 1040
His Gly Lys Cys Arg Asn Thr Ile Gly Ser Phe Lys Cys Arg Cys Asp
1045 1050 1055
Ser Gly Phe Ala Leu Asp Ser Glu Glu Arg Asn Cys Thr Asp Ile Asp
1060 1065 1070
Glu Cys Arg Ile Ser Pro Asp Leu Cys Gly Arg Gly Gln Cys Val Asn
1075 1080 1085
Thr Pro Gly Asp Phe Glu Cys Lys Cys Asp Glu Gly Tyr Glu Ser Gly
1090 1095 1100
Phe Met Met Met Lys Asn Cys Met Asp Ile Asp Glu Cys Gln Arg Asp
1105 1110 1115 1120
Pro Leu Leu Cys Arg Gly Gly Val Cys Leu Asn Thr Glu Gly Ser Tyr
1125 1130 1135
Arg Cys Glu Cys Pro Ser Gly His Gln Met Ser Pro Asn Ile Ser Ala
1140 1145 1150
Cys Ile Asp Ile Asn Glu Cys Glu Leu Ser Ala His Leu Cys Pro His
1155 1160 1165
Gly Arg Cys Val Asn Leu Ile Gly Lys Tyr Gln Arg Ala Arg Asn Pro
1170 1175 1180
Gly Tyr His Ser Thr Pro Asp Arg Leu Phe Cys Val Asp Ile Asp Glu
1185 1190 1195 1200
Cys Ser Ile Met Asn Gly Gly Cys Glu Thr Phe Cys Thr Asn Ser Glu
1205 1210 1215
Gly Ser Tyr Glu Cys Ser Cys Gln Pro Gly Phe Ala Leu Met Pro Asp
1220 1225 1230
Gln Arg Ser Cys Thr Asp Ile Asp Glu Cys Glu Asp Asn Pro Asn Ile
1235 1240 1245
Cys Asp Gly Gly Gln Cys Thr Asn Ile Pro Gly Glu Tyr Arg Cys Leu
1250 1255 1260
Cys Tyr Asp Gly Phe Met Ala Ser Glu Asp Met Lys Thr Cys Val Asp
1265 1270 1275 1280
Val Asn Glu Cys Asp Leu Asn Pro Asn Ile Cys Leu Ser Gly Thr Cys
1285 1290 1295
Glu Asn Thr Lys Gly Ser Phe Ile Cys His Cys Asp Met Gly Tyr Ser
1300 1305 1310
Gly Lys Lys Gly Lys Thr Gly Cys Thr Asp Ile Asn Glu Cys Glu Ile
1315 1320 1325
Gly Ala His Asn Cys Asp Arg His Ala Val Cys Thr Asn Thr Ala Gly
1330 1335 1340
Ser Phe Asn Cys Ser Cys Ser Pro Gly Trp Ile Gly Asp Gly Ile Lys
1345 1350 1355 1360
Cys Thr Asp Leu Asp Glu Cys Ser Asn Gly Thr His Met Cys Ser Gln
1365 1370 1375
His Ala Asp Cys Lys Asn Thr Met Gly Ser Tyr Arg Cys Leu Cys Lys
1380 1385 1390
Glu Gly Tyr Thr Gly Asp Gly Phe Thr Cys Ala Asp Leu Asp Glu Cys
1395 1400 1405
Ser Glu Asn Val Lys Leu Cys Gly Asn Val Gln Cys Leu Tyr Ala Pro
1410 1415 1420
Gly Gly Tyr His Cys Glu Tyr Asp Met Gly Phe Val Pro Ser Ala Asp
1425 1430 1435 1440
Arg Lys Ser Cys Val Asp Ser Asp Glu Cys Ser Leu Pro Asn Ile Cys
1445 1450 1455
Val Phe Gly Thr Cys His Asn Leu Pro Gly Leu Phe Arg Cys Glu Cys
1460 1465 1470
Glu Ile Gly Tyr Glu Leu Asp Arg Ser Gly Gly Asn Cys Thr Asp Val
1475 1480 1485
Asn Glu Cys Leu Glu Pro Pro Thr Cys Ile Ser Gly Asn Cys Val Asn
1490 1495 1500
Thr Pro Gly Ser Tyr Thr Cys Val Cys Pro Pro Asp Phe Glu Leu Asn
1505 1510 1515 1520
Pro Thr Arg Val Gly Cys Val Asp Thr Arg Ser Gly Asn Cys Tyr Leu
1525 1530 1535
Asp Val Arg Pro Arg Gly Asp Asn Gly Asp Thr Ala Cys Ser Asn Glu
1540 1545 1550
Ile Gly Val Gly Val Ser Lys Ala Ser Cys Cys Cys Ser Leu Gly Lys
1555 1560 1565
Ala Trp Gly Thr Pro Cys Glu Gln Cys Pro Pro Val Asn Thr Ser Glu
1570 1575 1580
Tyr Lys Ile Leu Cys Pro Gly Gly Glu Gly Phe Arg Pro Asn Pro Ile
1585 1590 1595 1600
Thr Val Ile Leu Glu Asp Ile Asp Glu Cys Gln Glu Leu Pro Gly Leu
1605 1610 1615
Cys Gln Gly Gly Lys Cys Ile Asn Thr Phe Gly Ser Phe Gln Cys Arg
1620 1625 1630
Cys Pro Thr Gly Tyr Tyr Leu Asn Glu Asp Thr Arg Val Cys Asp Asp
1635 1640 1645
Val Asn Glu Cys Glu Thr Pro Gly Ile Cys Gly Pro Gly Thr Cys Tyr
1650 1655 1660
Asn Thr Val Gly Asn Tyr Thr Cys Ile Cys Pro Pro Asp Tyr Met Gln
1665 1670 1675 1680
Val Asn Gly Gly Asn Asn Cys Met Asp Met Arg Arg Ser Leu Cys Tyr
1685 1690 1695
Arg Asn Tyr Tyr Ala Asp Asn Gln Thr Cys Asp Gly Glu Leu Leu Phe
1700 1705 1710
Asn Met Thr Lys Lys Met Cys Cys Cys Ser Tyr Asn Ile Gly Arg Ala
1715 1720 1725
Trp Asn Lys Pro Cys Glu Gln Cys Pro Ile Pro Ser Thr Asp Glu Phe
1730 1735 1740
Ala Thr Leu Cys Gly Ser Gln Arg Pro Gly Phe Val Ile Asp Ile Tyr
1745 1750 1755 1760
Thr Gly Leu Pro Val Asp Ile Asp Glu Cys Arg Glu Ile Pro Gly Val
1765 1770 1775
Cys Glu Asn Gly Val Cys Ile Asn Met Val Gly Ser Phe Arg Cys Glu
1780 1785 1790
Cys Pro Val Gly Phe Phe Tyr Asn Asp Lys Leu Leu Val Cys Glu Asp
1795 1800 1805
Ile Asp Glu Cys Gln Asn Gly Pro Val Cys Gln Arg Asn Ala Glu Cys
1810 1815 1820
Ile Asn Thr Ala Gly Ser Tyr Arg Cys Asp Cys Lys Pro Gly Tyr Arg
1825 1830 1835 1840
Phe Thr Ser Thr Gly Gln Cys Asn Asp Arg Asn Glu Cys Gln Glu Ile
1845 1850 1855
Pro Asn Ile Cys Ser His Gly Gln Cys Ile Asp Thr Val Gly Ser Phe
1860 1865 1870
Tyr Cys Leu Cys His Thr Gly Phe Lys Thr Asn Ala Asp Gln Thr Met
1875 1880 1885
Cys Leu Asp Ile Asn Glu Cys Glu Arg Asp Ala Cys Gly Asn Gly Thr
1890 1895 1900
Cys Arg Asn Thr Ile Gly Ser Phe Asn Cys Arg Cys Asn His Gly Phe
1905 1910 1915 1920
Ile Leu Ser His Asn Asn Asp Cys Ile Asp Val Asp Glu Cys Ala Thr
1925 1930 1935
Gly Asn Gly Asn Leu Cys Arg Asn Gly Gln Cys Ile Asn Thr Val Gly
1940 1945 1950
Ser Phe Gln Cys Gln Cys Asn Glu Gly Tyr Glu Val Ala Pro Asp Gly
1955 1960 1965
Arg Thr Cys Val Asp Ile Asn Glu Cys Leu Leu Glu Pro Gly Lys Cys
1970 1975 1980
Ala Pro Gly Thr Cys Gln Asn Leu Asp Gly Ser Tyr Arg Cys Ile Cys
1985 1990 1995 2000
Pro Pro Gly Tyr Ser Leu Gln Asn Asp Lys Cys Glu Asp Ile Asp Glu
2005 2010 2015
Cys Val Glu Glu Pro Glu Ile Cys Ala Leu Gly Thr Cys Ser Asn Thr
2020 2025 2030
Glu Gly Ser Phe Lys Cys Leu Cys Pro Asp Gly Phe Ser Leu Ser Ser
2035 2040 2045
Thr Gly Arg Arg Cys Gln Asp Leu Arg Met Ser Tyr Cys Tyr Ala Lys
2050 2055 2060
Phe Glu Gly Gly Lys Cys Ser Ser Pro Lys Ser Arg Asn His Ser Lys
2065 2070 2075 2080
Gln Glu Cys Cys Cys Ala Leu Lys Gly Glu Gly Trp Gly Asp Pro Cys
2085 2090 2095
Glu Leu Cys Pro Thr Glu Pro Asp Glu Ala Phe Arg Gln Ile Cys Pro
2100 2105 2110
Tyr Gly Ser Gly Ile Ile Val Gly Pro Asp Asp Ser Ala Val Asp Met
2115 2120 2125
Asp Glu Cys Lys Glu Pro Asp Val Cys Lys His Gly Gln Cys Ile Asn
2130 2135 2140
Thr Asp Gly Ser Tyr Arg Cys Glu Cys Pro Phe Gly Tyr Ile Leu Glu
2145 2150 2155 2160
Gly Asn Glu Cys Val Asp Thr Asp Glu Cys Ser Val Gly Asn Pro Cys
2165 2170 2175
Gly Asn Gly Thr Cys Lys Asn Val Ile Gly Gly Phe Glu Cys Thr Cys
2180 2185 2190
Glu Glu Gly Phe Glu Pro Gly Pro Met Met Thr Cys Glu Asp Ile Asn
2195 2200 2205
Glu Cys Ala Gln Asn Pro Leu Leu Cys Ala Phe Arg Cys Val Asn Thr
2210 2215 2220
Tyr Gly Ser Tyr Glu Cys Lys Cys Pro Thr Gly Tyr Val Leu Arg Glu
2225 2230 2235 2240
Asp Arg Arg Met Cys Lys Asp Glu Asp Glu Cys Glu Glu Gly Lys His
2245 2250 2255
Asp Cys Ala Glu Lys Gln Met Glu Cys Lys Asn Leu Ile Gly Met Tyr
2260 2265 2270
Ile Cys Ile Cys Gly Pro Gly Tyr Gln Arg Arg Pro Asp Gly Glu Gly
2275 2280 2285
Cys Val Asp Glu Asn Glu Cys Gln Thr Lys Pro Gly Ile Cys Glu Asn
2290 2295 2300
Gly Arg Cys Leu Asn Thr Arg Gly Ser Tyr Thr Cys Glu Cys Asn Asp
2305 2310 2315 2320
Gly Phe Thr Ala Ser Pro Thr Gln Asp Glu Cys Leu Asp Asn Arg Glu
2325 2330 2335
Gly Tyr Cys Phe Thr Glu Val Leu Gln Asn Met Cys Gln Ile Gly Ser
2340 2345 2350
Ser Asn Arg Asn Pro Val Thr Lys Ser Glu Cys Cys Cys Asp Gly Gly
2355 2360 2365
Arg Gly Trp Gly Pro His Cys Glu Ile Cys Pro Phe Gln Gly Thr Val
2370 2375 2380
Ala Phe Lys Lys Leu Cys Pro His Gly Arg Gly Phe Met Thr Asn Gly
2385 2390 2395 2400
Ala Asp Ile Asp Glu Cys Lys Val Ile His Asp Val Cys Arg Asn Gly
2405 2410 2415
Glu Cys Ile Asn Asp Arg Gly Ser Tyr His Cys Ile Cys Lys Thr Gly
2420 2425 2430
Tyr Thr Pro Asp Ile Thr Gly Thr Ala Cys Val Asp Leu Asn Glu Cys
2435 2440 2445
Asn Gln Ala Pro Lys Pro Cys Asn Phe Ile Cys Lys Asn Thr Glu Gly
2450 2455 2460
Ser Tyr Gln Cys Ser Cys Pro Lys Gly Tyr Ile Leu Gln Glu Asp Gly
2465 2470 2475 2480
Arg Ser Cys Lys Asp Leu Asp Glu Cys Ala Thr Lys Gln His Asn Cys
2485 2490 2495
Gln Phe Leu Cys Val Asn Thr Ile Gly Ser Phe Ala Cys Lys Cys Pro
2500 2505 2510
Pro Gly Phe Thr Gln His His Thr Ala Cys Ile Asp Asn Asn Glu Cys
2515 2520 2525
Thr Ser Asp Ile Asn Leu Cys Gly Ala Lys Gly Ile Cys Gln Asn Thr
2530 2535 2540
Pro Gly Ser Phe Thr Cys Glu Cys Gln Arg Gly Phe Ser Leu Asp Gln
2545 2550 2555 2560
Ser Gly Ala Ser Cys Glu Asp Val Asp Glu Cys Glu Gly Asn His Arg
2565 2570 2575
Cys Gln His Gly Cys Gln Asn Ile Ile Gly Gly Tyr Arg Cys Ser Cys
2580 2585 2590
Pro Gln Gly Tyr Leu Gln His Tyr Gln Trp Asn Gln Cys Val Asp Glu
2595 2600 2605
Asn Glu Cys Leu Ser Ala His Ile Cys Gly Gly Ala Ser Cys His Asn
2610 2615 2620
Thr Leu Gly Ser Tyr Lys Cys Met Cys Pro Ala Gly Phe Gln Tyr Glu
2625 2630 2635 2640
Gln Phe Ser Gly Gly Cys Gln Asp Ile Asn Glu Cys Gly Ser Ser Gln
2645 2650 2655
Ala Pro Cys Ser Tyr Gly Cys Ser Asn Thr Glu Gly Gly Tyr Leu Cys
2660 2665 2670
Gly Cys Pro Pro Gly Tyr Phe Arg Ile Gly Gln Gly His Cys Val Ser
2675 2680 2685
Gly Met Gly Met Gly Arg Gly Ser Pro Glu Pro Pro Ala Ser Gly Glu
2690 2695 2700
Met Asp Asp Asn Ser Leu Ser Pro Glu Ala Cys Tyr Glu Cys Lys Ile
2705 2710 2715 2720
Asn Gly Tyr Pro Lys Arg Gly Arg Lys Arg Arg Ser Thr Asn Glu Thr
2725 2730 2735
Asp Ala Phe Asn Ile Glu Asp Gln Pro Glu Thr Glu Ser Asn Val Ser
2740 2745 2750
Leu Ala Ser Trp Asp Val Glu Lys Thr Ala Val Phe Ala Phe Asn Ile
2755 2760 2765
Ser His Ile Ser Asn Lys Val Arg Ile Leu Glu Leu Leu Pro Ala Leu
2770 2775 2780
Thr Thr Leu Thr Asn His Asn Arg Tyr Leu Ile Glu Ser Gly Asn Glu
2785 2790 2795 2800
Asn Gly Phe Phe Lys Ile Asn Gln Lys Glu Gly Ile Ser Tyr Leu His
2805 2810 2815
Phe Thr Lys Lys Lys Pro Val Ala Gly Thr Tyr Ser Leu Gln Ile Ser
2820 2825 2830
Ser Thr Pro Leu Tyr Lys Lys Lys Glu Leu Asn Gln Leu Glu Asp Lys
2835 2840 2845
Tyr Asp Lys Asp Tyr Leu Ser Gly Glu Leu Gly Asp Asn Leu Lys Met
2850 2855 2860
Lys Ile Gln Ile Leu Leu His
2865 2870
<210> 15
<211> 926
<212> DNA
<213> Sus scrofa
<400> 15
taaatgaata tctcattcca ctacatactc aattggttgc tctgaggcaa tctatttaaa 60
aatcttctac atgctttaga tacagaagct aatggtttag aatttttatt cttacaagca 120
attacttcag tgtttagtgt gtataaggta gtgccctggg ctttagaagt ccaagtaaac 180
acactcttta cttctttaat acgcctttct aaaatgtaga cttttaaaag tctagaactg 240
tttacggaat cctatttctc acggatcatc acatatcttt aaaactgagt atgaccaaga 300
ccttaaatca agatcctcta attcctgggt ttagaatggg gtgttgagcc ctcctcactg 360
atcactgatg gacttttccc acattcaatt tttgttgaag acatcaacga gtgcaagatg 420
atccccaacc tctgtaccca cggcaagtgc aggaacacta tcggcagctt caagtgcaga 480
tgtgacagtg gctttgctct ggactctgaa gaaaggaact gtacaggtca gtaaggggct 540
tagcccagag ggacatcccc tgggacacct cgcctttaac ccactgcctc aagaaggaaa 600
gcgttcctgc tgctcatttg ctgcccttgc tgctcctgtg cagatattga tgaatgccgc 660
atttctcctg acctctgtgg ccgaggccag tgtgtgaaca cccctgggga cttcgaatgc 720
aagtgtgatg aaggctatga aagtggattc atgatgatga agaactgcat gggtaagtgg 780
gtgagccttt gatcatacaa attctaagaa gctttggtga ctcatctcct gtgttcccag 840
ttcttcctgc tcccctaaag ccctggtctc ctgaccagtg agtcagcaat tggtcagttg 900
cagagagaga ctgacatgtg gaagga 926
<210> 16
<211> 100
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 16
auccccaacc ucuguaccca guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu 100
<210> 17
<211> 100
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 17
aguguuccug cacuugccgu guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu 100
<210> 18
<211> 100
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 18
uaguguuccu gcacuugccg guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu 100
<210> 19
<211> 100
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 19
ugcacuugcc guggguacag guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu 100
<210> 20
<211> 100
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 20
ggcaagugca ggaacacuau guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu 100
<210> 21
<211> 100
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 21
cuucaacaaa aauugaaugu guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu 100
<210> 22
<211> 100
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 22
ucuucaacaa aaauugaaug guuuuagagc uagaaauagc aaguuaaaau aaggcuaguc 60
cguuaucaac uugaaaaagu ggcaccgagu cggugcuuuu 100
<210> 23
<211> 225
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 23
ggcttgtcgg actcttcgct attacgccag ctggcgaagg gggatgtgct gcaaggcgat 60
taagttgggt aacgccaggg ttttcccagt cacgacgtta ggaaattaat acgactcact 120
ataggtgcac ttgccgtggg tacaggtttt agagctagaa atagcaagtt aaaataaggc 180
tagtccgtta tcaacttgaa aaagtggcac cgagtcggtg ctttt 225
<210> 24
<211> 225
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 24
ggcttgtcgg actcttcgct attacgccag ctggcgaagg gggatgtgct gcaaggcgat 60
taagttgggt aacgccaggg ttttcccagt cacgacgtta ggaaattaat acgactcact 120
ataggcttca acaaaaattg aatgtgtttt agagctagaa atagcaagtt aaaataaggc 180
tagtccgtta tcaacttgaa aaagtggcac cgagtcggtg ctttt 225
<210> 25
<211> 102
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 25
ggugcacuug ccguggguac agguuuuaga gcuagaaaua gcaaguuaaa auaaggcuag 60
uccguuauca acuugaaaaa guggcaccga gucggugcuu uu 102
<210> 26
<211> 102
<212> RNA
<213> 人工序列(Artificial Sequence)
<400> 26
ggcuucaaca aaaauugaau guguuuuaga gcuagaaaua gcaaguuaaa auaaggcuag 60
uccguuauca acuugaaaaa guggcaccga gucggugcuu uu 102
<210> 27
<211> 163
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 27
ggtttagaat ggggtgttga gccctcctca ctgatcactg atggactttt accacattca 60
atttttgttg aagacatcaa cgagtgcaag atgatcccca atctcggtac ccacggcaag 120
tgcaggaaca ctatcggcag cttcaagtgc agatgtgaca gtg 163
<210> 28
<211> 163
<212> DNA
<213> Sus scrofa
<400> 28
ggtttagaat ggggtgttga gccctcctca ctgatcactg atggactttt cccacattca 60
atttttgttg aagacatcaa cgagtgcaag atgatcccca acctctgtac ccacggcaag 120
tgcaggaaca ctatcggcag cttcaagtgc agatgtgaca gtg 163
<210> 29
<211> 10476
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 29
gagggcctat ttcccatgat tccttcatat ttgcatatac gatacaaggc tgttagagag 60
ataattggaa ttaatttgac tgtaaacaca aagatattag tacaaaatac gtgacgtaga 120
aagtaataat ttcttgggta gtttgcagtt ttaaaattat gttttaaaat ggactatcat 180
atgcttaccg taacttgaaa gtatttcgat ttcttggctt tatatatctt gtggaaagga 240
cgaaacaccg ggtcttcgag aagacctgtt ttagagctag aaatagcaag ttaaaataag 300
gctagtccgt tatcaacttg aaaaagtggc accgagtcgg tgcttttttc tagcgcgtgc 360
gccaattctg cagacaaatg gctctagagg tacccgttac ataacttacg gtaaatggcc 420
cgcctggctg accgcccaac gacccccgcc cattgacgtc aatagtaacg ccaataggga 480
ctttccattg acgtcaatgg gtggagtatt tacggtaaac tgcccacttg gcagtacatc 540
aagtgtatca tatgccaagt acgcccccta ttgacgtcaa tgacggtaaa tggcccgcct 600
ggcattgtgc ccagtacatg accttatggg actttcctac ttggcagtac atctacgtat 660
tagtcatcgc tattaccatg ggggcagagc gcacatcgcc cacagtcccc gagaagttgg 720
ggggaggggt cggcaattga tccggtgcct agagaaggtg gcgcggggta aactgggaaa 780
gtgatgtcgt gtactggctc cgcctttttc ccgagggtgg gggagaaccg tatataagtg 840
cagtagtcgc cgtgaacgtt ctttttcgca acgggtttgc cgccagaaca caggttggac 900
cggtgccacc atggactata aggaccacga cggagactac aaggatcatg atattgatta 960
caaagacgat gacgataaga tggcccccaa aaagaaacga aaggtgggtg ggtccccaaa 1020
gaagaagcgg aaggtcggta tccacggagt cccagcagcc gacaagaagt acagcatcgg 1080
cctggacatc ggcaccaact ctgtgggctg ggccgtgatc accgacgagt acaaggtgcc 1140
cagcaagaaa ttcaaggtgc tgggcaacac cgaccggcac agcatcaaga agaacctgat 1200
cggagccctg ctgttcgaca gcggcgaaac agccgaggcc acccggctga agagaaccgc 1260
cagaagaaga tacaccagac ggaagaaccg gatctgctat ctgcaagaga tcttcagcaa 1320
cgagatggcc aaggtggacg acagcttctt ccacagactg gaagagtcct tcctggtgga 1380
agaggataag aagcacgagc ggcaccccat cttcggcaac atcgtggacg aggtggccta 1440
ccacgagaag taccccacca tctaccacct gagaaagaaa ctggtggaca gcaccgacaa 1500
ggccgacctg cggctgatct atctggccct ggcccacatg atcaagttcc ggggccactt 1560
cctgatcgag ggcgacctga accccgacaa cagcgacgtg gacaagctgt tcatccagct 1620
ggtgcagacc tacaaccagc tgttcgagga aaaccccatc aacgccagcg gcgtggacgc 1680
caaggccatc ctgtctgcca gactgagcaa gagcagacgg ctggaaaatc tgatcgccca 1740
gctgcccggc gagaagaaga atggcctgtt cggaaacctg attgccctga gcctgggcct 1800
gacccccaac ttcaagagca acttcgacct ggccgaggat gccaaactgc agctgagcaa 1860
ggacacctac gacgacgacc tggacaacct gctggcccag atcggcgacc agtacgccga 1920
cctgtttctg gccgccaaga acctgtccga cgccatcctg ctgagcgaca tcctgagagt 1980
gaacaccgag atcaccaagg cccccctgag cgcctctatg atcaagagat acgacgagca 2040
ccaccaggac ctgaccctgc tgaaagctct cgtgcggcag cagctgcctg agaagtacaa 2100
agagattttc ttcgaccaga gcaagaacgg ctacgccggc tacattgacg gcggagccag 2160
ccaggaagag ttctacaagt tcatcaagcc catcctggaa aagatggacg gcaccgagga 2220
actgctcgtg aagctgaaca gagaggacct gctgcggaag cagcggacct tcgacaacgg 2280
cagcatcccc caccagatcc acctgggaga gctgcacgcc attctgcggc ggcaggaaga 2340
tttttaccca ttcctgaagg acaaccggga aaagatcgag aagatcctga ccttccgcat 2400
cccctactac gtgggccctc tggccagggg aaacagcaga ttcgcctgga tgaccagaaa 2460
gagcgaggaa accatcaccc cctggaactt cgaggaagtg gtggacaagg gcgcttccgc 2520
ccagagcttc atcgagcgga tgaccaactt cgataagaac ctgcccaacg agaaggtgct 2580
gcccaagcac agcctgctgt acgagtactt caccgtgtat aacgagctga ccaaagtgaa 2640
atacgtgacc gagggaatga gaaagcccgc cttcctgagc ggcgagcaga aaaaggccat 2700
cgtggacctg ctgttcaaga ccaaccggaa agtgaccgtg aagcagctga aagaggacta 2760
cttcaagaaa atcgagtgct tcgactccgt ggaaatctcc ggcgtggaag atcggttcaa 2820
cgcctccctg ggcacatacc acgatctgct gaaaattatc aaggacaagg acttcctgga 2880
caatgaggaa aacgaggaca ttctggaaga tatcgtgctg accctgacac tgtttgagga 2940
cagagagatg atcgaggaac ggctgaaaac ctatgcccac ctgttcgacg acaaagtgat 3000
gaagcagctg aagcggcgga gatacaccgg ctggggcagg ctgagccgga agctgatcaa 3060
cggcatccgg gacaagcagt ccggcaagac aatcctggat ttcctgaagt ccgacggctt 3120
cgccaacaga aacttcatgc agctgatcca cgacgacagc ctgaccttta aagaggacat 3180
ccagaaagcc caggtgtccg gccagggcga tagcctgcac gagcacattg ccaatctggc 3240
cggcagcccc gccattaaga agggcatcct gcagacagtg aaggtggtgg acgagctcgt 3300
gaaagtgatg ggccggcaca agcccgagaa catcgtgatc gaaatggcca gagagaacca 3360
gaccacccag aagggacaga agaacagccg cgagagaatg aagcggatcg aagagggcat 3420
caaagagctg ggcagccaga tcctgaaaga acaccccgtg gaaaacaccc agctgcagaa 3480
cgagaagctg tacctgtact acctgcagaa tgggcgggat atgtacgtgg accaggaact 3540
ggacatcaac cggctgtccg actacgatgt ggaccatatc gtgcctcaga gctttctgaa 3600
ggacgactcc atcgacaaca aggtgctgac cagaagcgac aagaaccggg gcaagagcga 3660
caacgtgccc tccgaagagg tcgtgaagaa gatgaagaac tactggcggc agctgctgaa 3720
cgccaagctg attacccaga gaaagttcga caatctgacc aaggccgaga gaggcggcct 3780
gagcgaactg gataaggccg gcttcatcaa gagacagctg gtggaaaccc ggcagatcac 3840
aaagcacgtg gcacagatcc tggactcccg gatgaacact aagtacgacg agaatgacaa 3900
gctgatccgg gaagtgaaag tgatcaccct gaagtccaag ctggtgtccg atttccggaa 3960
ggatttccag ttttacaaag tgcgcgagat caacaactac caccacgccc acgacgccta 4020
cctgaacgcc gtcgtgggaa ccgccctgat caaaaagtac cctaagctgg aaagcgagtt 4080
cgtgtacggc gactacaagg tgtacgacgt gcggaagatg atcgccaaga gcgagcagga 4140
aatcggcaag gctaccgcca agtacttctt ctacagcaac atcatgaact ttttcaagac 4200
cgagattacc ctggccaacg gcgagatccg gaagcggcct ctgatcgaga caaacggcga 4260
aaccggggag atcgtgtggg ataagggccg ggattttgcc accgtgcgga aagtgctgag 4320
catgccccaa gtgaatatcg tgaaaaagac cgaggtgcag acaggcggct tcagcaaaga 4380
gtctatcctg cccaagagga acagcgataa gctgatcgcc agaaagaagg actgggaccc 4440
taagaagtac ggcggcttcg acagccccac cgtggcctat tctgtgctgg tggtggccaa 4500
agtggaaaag ggcaagtcca agaaactgaa gagtgtgaaa gagctgctgg ggatcaccat 4560
catggaaaga agcagcttcg agaagaatcc catcgacttt ctggaagcca agggctacaa 4620
agaagtgaaa aaggacctga tcatcaagct gcctaagtac tccctgttcg agctggaaaa 4680
cggccggaag agaatgctgg cctctgccgg cgaactgcag aagggaaacg aactggccct 4740
gccctccaaa tatgtgaact tcctgtacct ggccagccac tatgagaagc tgaagggctc 4800
ccccgaggat aatgagcaga aacagctgtt tgtggaacag cacaagcact acctggacga 4860
gatcatcgag cagatcagcg agttctccaa gagagtgatc ctggccgacg ctaatctgga 4920
caaagtgctg tccgcctaca acaagcaccg ggataagccc atcagagagc aggccgagaa 4980
tatcatccac ctgtttaccc tgaccaatct gggagcccct gccgccttca agtactttga 5040
caccaccatc gaccggaaga ggtacaccag caccaaagag gtgctggacg ccaccctgat 5100
ccaccagagc atcaccggcc tgtacgagac acggatcgac ctgtctcagc tgggaggcga 5160
caaaaggccg gcggccacga aaaaggccgg ccaggcaaaa aagaaaaagg gcggctccaa 5220
gcggcctgcc gcgacgaaga aagcgggaca ggccaagaaa aagaaaggat ccggcgcaac 5280
aaacttctct ctgctgaaac aagccggaga tgtcgaagag aatcctggac cggtgagcaa 5340
gggcgaggag ctgttcaccg gggtggtgcc catcctggtc gagctggacg gcgacgtaaa 5400
cggccacaag ttcagcgtgt ccggcgaggg cgagggcgat gccacctacg gcaagctgac 5460
cctgaagttc atctgcacca ccggcaagct gcccgtgccc tggcccaccc tcgtgaccac 5520
cctgacctac ggcgtgcagt gcttcagccg ctaccccgac cacatgaagc agcacgactt 5580
cttcaagtcc gccatgcccg aaggctacgt ccaggagcgc accatcttct tcaaggacga 5640
cggcaactac aagacccgcg ccgaggtgaa gttcgagggc gacaccctgg tgaaccgcat 5700
cgagctgaag ggcatcgact tcaaggagga cggcaacatc ctggggcaca agctggagta 5760
caactacaac agccacaacg tctatatcat ggccgacaag cagaagaacg gcatcaaggt 5820
gaacttcaag atccgccaca acatcgagga cggcagcgtg cagctcgccg accactacca 5880
gcagaacacc cccatcggcg acggccccgt gctgctgccc gacaaccact acctgagcac 5940
ccagtccgcc ctgagcaaag accccaacga gaagcgcgat cacatggtcc tgctggagtt 6000
cgtgaccgcc gccgggatca ctctcggcat ggacgagctg tacaagggct ccggcgaggg 6060
caggggaagt cttctaacat gcggggacgt ggaggaaaat cccggcccaa ccgagtacaa 6120
gcccacggtg cgcctcgcca cccgcgacga cgtccccagg gccgtacgca ccctcgccgc 6180
cgcgttcgcc gactaccccg ccacgcgcca caccgtcgat ccggaccgcc acatcgagcg 6240
ggtcaccgag ctgcaagaac tcttcctcac gcgcgtcggg ctcgacatcg gcaaggtgtg 6300
ggtcgcggac gacggcgccg cggtggcggt ctggaccacg ccggagagcg tcgaagcggg 6360
ggcggtgttc gccgagatcg gcccgcgcat ggccgagttg agcggttccc ggctggccgc 6420
gcagcaacag atggaaggcc tcctggcgcc gcaccggccc aaggagcccg cgtggttcct 6480
ggccaccgtc ggagtctcgc ccgaccacca gggcaagggt ctgggcagcg ccgtcgtgct 6540
ccccggagtg gaggcggccg agcgcgccgg ggtgcccgcc ttcctggaga cctccgcgcc 6600
ccgcaacctc cccttctacg agcggctcgg cttcaccgtc accgccgacg tcgaggtgcc 6660
cgaaggaccg cgcacctggt gcatgacccg caagcccggt gcctgaacgc gttaagtcga 6720
caatcaacct ctggattaca aaatttgtga aagattgact ggtattctta actatgttgc 6780
tccttttacg ctatgtggat acgctgcttt aatgcctttg tatcatgcta ttgcttcccg 6840
tatggctttc attttctcct ccttgtataa atcctggttg ctgtctcttt atgaggagtt 6900
gtggcccgtt gtcaggcaac gtggcgtggt gtgcactgtg tttgctgacg caacccccac 6960
tggttggggc attgccacca cctgtcagct cctttccggg actttcgctt tccccctccc 7020
tattgccacg gcggaactca tcgccgcctg ccttgcccgc tgctggacag gggctcggct 7080
gttgggcact gacaattccg tggtgttgtc ggggaaatca tcgtcctttc cttggctgct 7140
cgcctgtgtt gccacctgga ttctgcgcgg gacgtccttc tgctacgtcc cttcggccct 7200
caatccagcg gaccttcctt cccgcggcct gctgccggct ctgcggcctc ttccgcgtct 7260
tcgccttcgc cctcagacga gtcggatctc cctttgggcc gcctccccgc gtcgacttta 7320
agaccaatga cttacaaggc agctgtagat cttagccact ttttaaaaga aaagggggga 7380
ctggaagggc taattcactc ccaacgaaga caagatctgc tttttgcttg tactgggtct 7440
ctctggttag accagatctg agcctgggag ctctctggct aactagggaa cccactgctt 7500
aagcctcaat aaagcttgcc ttgagtgctt caagtagtgt gtgcccgtct gttgtgtgac 7560
tctggtaact agagatccct cagacccttt tagtcagtgt ggaaaatctc tagcagggcc 7620
cgtttaaacc cgctgatcag cctcgactgt gccttctagt tgccagccat ctgttgtttg 7680
cccctccccc gtgccttcct tgaccctgga aggtgccact cccactgtcc tttcctaata 7740
aaatgaggaa attgcatcgc attgtctgag taggtgtcat tctattctgg ggggtggggt 7800
ggggcaggac agcaaggggg aggattggga agacaatagc aggcatgctg gggatgcggt 7860
gggctctatg gcctgcaggg gcgcctgatg cggtattttc tccttacgca tctgtgcggt 7920
atttcacacc gcatacgtca aagcaaccat agtacgcgcc ctgtagcggc gcattaagcg 7980
cggcgggtgt ggtggttacg cgcagcgtga ccgctacact tgccagcgcc ttagcgcccg 8040
ctcctttcgc tttcttccct tcctttctcg ccacgttcgc cggctttccc cgtcaagctc 8100
taaatcgggg gctcccttta gggttccgat ttagtgcttt acggcacctc gaccccaaaa 8160
aacttgattt gggtgatggt tcacgtagtg ggccatcgcc ctgatagacg gtttttcgcc 8220
ctttgacgtt ggagtccacg ttctttaata gtggactctt gttccaaact ggaacaacac 8280
tcaactctat ctcgggctat tcttttgatt tataagggat tttgccgatt tcggtctatt 8340
ggttaaaaaa tgagctgatt taacaaaaat ttaacgcgaa ttttaacaaa atattaacgt 8400
ttacaatttt atggtgcact ctcagtacaa tctgctctga tgccgcatag ttaagccagc 8460
cccgacaccc gccaacaccc gctgacgcgc cctgacgggc ttgtctgctc ccggcatccg 8520
cttacagaca agctgtgacc gtctccggga gctgcatgtg tcagaggttt tcaccgtcat 8580
caccgaaacg cgcgagacga aagggcctcg tgatacgcct atttttatag gttaatgtca 8640
tgataataat ggtttcttag acgtcaggtg gcacttttcg gggaaatgtg cgcggaaccc 8700
ctatttgttt atttttctaa atacattcaa atatgtatcc gctcatgaga caataaccct 8760
gataaatgct tcaataatat tgaaaaagga agagtatgag tattcaacat ttccgtgtcg 8820
cccttattcc cttttttgcg gcattttgcc ttcctgtttt tgctcaccca gaaacgctgg 8880
tgaaagtaaa agatgctgaa gatcagttgg gtgcacgagt gggttacatc gaactggatc 8940
tcaacagcgg taagatcctt gagagttttc gccccgaaga acgttttcca atgatgagca 9000
cttttaaagt tctgctatgt ggcgcggtat tatcccgtat tgacgccggg caagagcaac 9060
tcggtcgccg catacactat tctcagaatg acttggttga gtactcacca gtcacagaaa 9120
agcatcttac ggatggcatg acagtaagag aattatgcag tgctgccata accatgagtg 9180
ataacactgc ggccaactta cttctgacaa cgatcggagg accgaaggag ctaaccgctt 9240
ttttgcacaa catgggggat catgtaactc gccttgatcg ttgggaaccg gagctgaatg 9300
aagccatacc aaacgacgag cgtgacacca cgatgcctgt agcaatggca acaacgttgc 9360
gcaaactatt aactggcgaa ctacttactc tagcttcccg gcaacaatta atagactgga 9420
tggaggcgga taaagttgca ggaccacttc tgcgctcggc ccttccggct ggctggttta 9480
ttgctgataa atctggagcc ggtgagcgtg gaagccgcgg tatcattgca gcactggggc 9540
cagatggtaa gccctcccgt atcgtagtta tctacacgac ggggagtcag gcaactatgg 9600
atgaacgaaa tagacagatc gctgagatag gtgcctcact gattaagcat tggtaactgt 9660
cagaccaagt ttactcatat atactttaga ttgatttaaa acttcatttt taatttaaaa 9720
ggatctaggt gaagatcctt tttgataatc tcatgaccaa aatcccttaa cgtgagtttt 9780
cgttccactg agcgtcagac cccgtagaaa agatcaaagg atcttcttga gatccttttt 9840
ttctgcgcgt aatctgctgc ttgcaaacaa aaaaaccacc gctaccagcg gtggtttgtt 9900
tgccggatca agagctacca actctttttc cgaaggtaac tggcttcagc agagcgcaga 9960
taccaaatac tgttcttcta gtgtagccgt agttaggcca ccacttcaag aactctgtag 10020
caccgcctac atacctcgct ctgctaatcc tgttaccagt ggctgctgcc agtggcgata 10080
agtcgtgtct taccgggttg gactcaagac gatagttacc ggataaggcg cagcggtcgg 10140
gctgaacggg gggttcgtgc acacagccca gcttggagcg aacgacctac accgaactga 10200
gatacctaca gcgtgagcta tgagaaagcg ccacgcttcc cgaagggaga aaggcggaca 10260
ggtatccggt aagcggcagg gtcggaacag gagagcgcac gagggagctt ccagggggaa 10320
acgcctggta tctttatagt cctgtcgggt ttcgccacct ctgacttgag cgtcgatttt 10380
tgtgatgctc gtcagggggg cggagcctat ggaaaaacgc cagcaacgcg gcctttttac 10440
ggttcctggc cttttgctgg ccttttgctc acatgt 10476
<210> 30
<211> 3120
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 30
gacgaaaggg cctcgtgata cgcctatttt tataggttaa tgtcatgata ataatggttt 60
cttagacgtc aggtggcact tttcggggaa atgtgcgcgg aacccctatt tgtttatttt 120
tctaaataca ttcaaatatg tatccgctca tgagacaata accctgataa atgcttcaat 180
aatattgaaa aaggaagagt atgagtattc aacatttccg tgtcgccctt attccctttt 240
ttgcggcatt ttgccttcct gtttttgctc acccagaaac gctggtgaaa gtaaaagatg 300
ctgaagatca gttgggtgca cgagtgggtt acatcgaact ggatctcaac agcggtaaga 360
tccttgagag ttttcgcccc gaagaacgtt ttccaatgat gagcactttt aaagttctgc 420
tatgtggcgc ggtattatcc cgtattgacg ccgggcaaga gcaactcggt cgccgcatac 480
actattctca gaatgacttg gttgagtact caccagtcac agaaaagcat cttacggatg 540
gcatgacagt aagagaatta tgcagtgctg ccataaccat gagtgataac actgcggcca 600
acttacttct gacaacgatc ggaggaccga aggagctaac cgcttttttg cacaacatgg 660
gggatcatgt aactcgcctt gatcgttggg aaccggagct gaatgaagcc ataccaaacg 720
acgagcgtga caccacgatg cctgtagcaa tggcaacaac gttgcgcaaa ctattaactg 780
gcgaactact tactctagct tcccggcaac aattaataga ctggatggag gcggataaag 840
ttgcaggacc acttctgcgc tcggcccttc cggctggctg gtttattgct gataaatctg 900
gagccggtga gcgtgggtct cgcggtatca ttgcagcact ggggccagat ggtaagccct 960
cccgtatcgt agttatctac acgacgggga gtcaggcaac tatggatgaa cgaaatagac 1020
agatcgctga gataggtgcc tcactgatta agcattggta actgtcagac caagtttact 1080
catatatact ttagattgat ttaaaacttc atttttaatt taaaaggatc taggtgaaga 1140
tcctttttga taatctcatg accaaaatcc cttaacgtga gttttcgttc cactgagcgt 1200
cagaccccgt agaaaagatc aaaggatctt cttgagatcc tttttttctg cgcgtaatct 1260
gctgcttgca aacaaaaaaa ccaccgctac cagcggtggt ttgtttgccg gatcaagagc 1320
taccaactct ttttccgaag gtaactggct tcagcagagc gcagatacca aatactgttc 1380
ttctagtgta gccgtagtta ggccaccact tcaagaactc tgtagcaccg cctacatacc 1440
tcgctctgct aatcctgtta ccagtggctg ctgccagtgg cgataagtcg tgtcttaccg 1500
ggttggactc aagacgatag ttaccggata aggcgcagcg gtcgggctga acggggggtt 1560
cgtgcacaca gcccagcttg gagcgaacga cctacaccga actgagatac ctacagcgtg 1620
agctatgaga aagcgccacg cttcccgaag ggagaaaggc ggacaggtat ccggtaagcg 1680
gcagggtcgg aacaggagag cgcacgaggg agcttccagg gggaaacgcc tggtatcttt 1740
atagtcctgt cgggtttcgc cacctctgac ttgagcgtcg atttttgtga tgctcgtcag 1800
gggggcggag cctatggaaa aacgccagca acgcggcctt tttacggttc ctggcctttt 1860
gctggccttt tgctcacatg ttctttcctg cgttatcccc tgattctgtg gataaccgta 1920
ttaccgcctt tgagtgagct gataccgctc gccgcagccg aacgaccgag cgcagcgagt 1980
cagtgagcga ggaagcggaa gagcgcccaa tacgcaaacc gcctctcccc gcgcgttggc 2040
cgattcatta atgcagctgg cacgacaggt ttcccgactg gaaagcgggc agtgagcgca 2100
acgcaattaa tgtgagttag ctcactcatt aggcacccca ggctttacac tttatgcttc 2160
cggctcgtat gttgtgtgga attgtgagcg gataacaatt tcacacagga aacagctatg 2220
accatgatta cgccaagctt gcatgcaggc ctctgcagtc gacgggcccg ggatccgatg 2280
ataaacatgt gagggcctat ttcccatgat tccttcatat ttgcatatac gatacaaggc 2340
tgttagagag ataattggaa ttaatttgac tgtaaacaca aagatattag tacaaaatac 2400
gtgacgtaga aagtaataat ttcttgggta gtttgcagtt ttaaaattat gttttaaaat 2460
ggactatcat atgcttaccg taacttgaaa gtatttcgat ttcttggctt tatatatctt 2520
gtggaaagga cgaaacaccg ggtcttcgag aagacctgtt ttagagctag aaatagcaag 2580
ttaaaataag gctagtccgt tatcaacttg aaaaagtggc accgagtcgg tgcttttttc 2640
tagcgcgtgc gccaattctg cagacaaatg gctctagagg tacccataga tctagatgca 2700
ttcgcgaggt accgagctcg aattcactgg ccgtcgtttt acaacgtcgt gactgggaaa 2760
accctggcgt tacccaactt aatcgccttg cagcacatcc ccctttcgcc agctggcgta 2820
atagcgaaga ggcccgcacc gatcgccctt cccaacagtt gcgcagcctg aatggcgaat 2880
ggcgcctgat gcggtatttt ctccttacgc atctgtgcgg tatttcacac cgcatatggt 2940
gcactctcag tacaatctgc tctgatgccg catagttaag ccagccccga cacccgccaa 3000
cacccgctga cgcgccctga cgggcttgtc tgctcccggc atccgcttac agacaagctg 3060
tgaccgtctc cgggagctgc atgtgtcaga ggttttcacc gtcatcaccg aaacgcgcga 3120
Claims (13)
1.一种试剂盒，包括质粒pKG-U6gRNA(APOE-E2-gRNA2)、FBN1-gRNA4、FBN1-gRNA6和FBN1-mutant-ss163；
质粒pKG-U6gRNA(APOE-E2-gRNA2)，其转录得到sgRNAAPOE-E2-gRNA2；sgRNAAPOE-E2-gRNA2，为sgRNA，其靶序列结合区如SEQ ID NO：11中第1-20位核苷酸所示；
FBN1-gRNA4，为sgRNA，其靶序列结合区如SEQ ID NO：25中第3-22位核苷酸所示；
FBN1-gRNA6，为sgRNA，其靶序列结合区如SEQ ID NO：26中第3-22位核苷酸所示；
FBN1-mutant-ss163为SEQ ID NO：27所示的单链DNA分子；
所述试剂盒的用途为如下(a)或(b)或(c)：(a)制备重组细胞；(b)制备动脉粥样硬化模型猪；(c)制备动脉粥样硬化细胞模型或动脉粥样硬化组织模型或动脉粥样硬化器官模型。
2.如权利要求1所述的试剂盒，其特征在于：所述试剂盒还包括表达Cas9蛋白的质粒。
3.如权利要求1或2所述的试剂盒，其特征在于：所述试剂盒还包括NCN蛋白；所述NCN蛋白为Cas9蛋白或具有Cas9蛋白的融合蛋白。
4.如权利要求1或2所述的试剂盒，其特征在于：所述试剂盒还包括PRONCN蛋白；所述PRONCN蛋白自上游至下游依次包括如下元件：信号肽、分子伴侣蛋白、蛋白标签、蛋白酶酶切位点、核定位信号、Cas9蛋白、核定位信号。
5.如权利要求1或2所述的试剂盒，其特征在于：所述试剂盒还包括质粒pKG-GE4；
所述质粒pKG-GE4自上游至下游依次包括如下元件：启动子、操纵子、核糖体结合位点、PRONCN蛋白的编码基因、终止子；所述PRONCN蛋白自上游至下游依次包括如下元件：信号肽、分子伴侣蛋白、蛋白标签、蛋白酶酶切位点、核定位信号、Cas9蛋白、核定位信号。
6.如权利要求1至5中任一所述的试剂盒，其特征在于：所述试剂盒还包括猪细胞。
7.一种制备重组细胞的方法，包括如下步骤：将质粒pKG-U6gRNA(APOE-E2-gRNA2)、表达Cas9蛋白的质粒、FBN1-gRNA4、FBN1-gRNA6、FBN1-mutant-ss163和NCN蛋白共转染猪细胞，得到APOE基因发生突变且FBN1基因发生目标突变的重组细胞；FBN1基因发生目标突变指的是：用SEQ ID NO：27所示的DNA分子取代猪细胞的染色体DNA中SEQ ID NO：28所示的DNA分子，得到重组细胞；
质粒pKG-U6gRNA(APOE-E2-gRNA2)，其转录得到sgRNAAPOE-E2-gRNA2；sgRNAAPOE-E2-gRNA2，为sgRNA，其靶序列结合区如SEQ ID NO：11中第1-20位核苷酸所示；
FBN1-gRNA4，为sgRNA，其靶序列结合区如SEQ ID NO：25中第3-22位核苷酸所示；
FBN1-gRNA6，为sgRNA，其靶序列结合区如SEQ ID NO：26中第3-22位核苷酸所示；
FBN1-mutant-ss163为SEQ ID NO：27所示的单链DNA分子；
所述NCN蛋白为Cas9蛋白或具有Cas9蛋白的融合蛋白。
8.如权利要求3所述的试剂盒或如权利要求7所述的方法，其特征在于：所述NCN蛋白如SEQ ID NO：3所示。
9.如权利要求8所述的试剂盒或方法，其特征在于：
所述NCN蛋白的制备方法包括如下步骤：
(1)将质粒pKG-GE4导入大肠杆菌BL21(DE3)，得到重组菌；
(2)采用液体培养基30℃培养所述重组菌，然后加入IPTG并25℃诱导培养，然后收集菌体；
(3)将收集的菌体进行菌体破碎，收集粗蛋白溶液；
(4)采用亲和层析从所述粗蛋白溶液中纯化具有His6标签的融合蛋白；
(5)采用具有His6标签的肠激酶酶切具有His6标签的融合蛋白，然后采用Ni-NTA树脂去除具有His6标签的蛋白，得到纯化的NCN蛋白；
质粒pKG-GE4中具有SEQ ID NO：1中第5209-9852位核苷酸所示的融合基因。
10.权利要求7至9中任一所述方法制备得到的重组细胞。
11.权利要求10所述重组细胞在制备动脉粥样硬化模型猪中的应用。
12.利用权利要求10所述重组细胞所制备的动脉粥样硬化模型猪的猪组织、猪器官或猪细胞。
13.权利要求10所述重组细胞、权利要求12所述猪组织、权利要求12所述猪器官、权利要求12所述猪细胞或者利用权利要求10所述重组细胞所制备的动脉粥样硬化模型猪的应用，为如下(d1)或(d2)或(d3)或(d4)：
(d1)筛选治疗动脉粥样硬化的药物；
(d2)进行动脉粥样硬化药物的药效评价；
(d3)进行动脉粥样硬化的基因治疗和/或细胞治疗的疗效评价；
(d4)研究动脉粥样硬化的发病机制。
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