CN111499758A - 筛选人钾离子通道调节剂的方法 - Google Patents
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Abstract
本发明涉及筛选人钾离子通道调节剂的方法。本发明中，确定了一系列离子通道蛋白的融合蛋白，适用于在体高通量筛选调控人的KCNQ2、KV1.2和BK钾通道功能的物质，如小分子化合物等药物。本发明还提供了编码所述融合蛋白的多核苷酸序列，相关的转基因线虫，以及相关的筛选方法和应用。为筛选药物治疗离子通道疾病提供了新的途径。
Description
技术领域
本发明属于生物技术领域，更具体地，本发明涉及一种筛选人钾离子通道调节剂的方法。
背景技术
癫痫是一类由于大脑神经元突发性异常放电，导致短暂的大脑功能障碍的一种慢性疾病。由于神经元异常放电的起始部位和传递方式不同，癫痫发作的临床表现复杂多样，可出现抽搐、肌肉痉挛和意识丧失等症状。癫痫是最常见的神经性疾病之一，在不同年龄段的病人都有发生，但主要在小孩和老年人群中高发；它的发病率大约是1％，全球有6500多万人患有癫痫疾病，带来严重的家庭和社会医疗负担。
根据降低大脑神经元兴奋性来减缓癫痫疾病的理论，临床上通过增加GABA能抑制性神经元活性的药物或抑制兴奋性神经元活性的药物如电压门控钠、钙通道的抑制剂等来治疗癫痫疾病。目前，临床上尽管有20多种抗癫痫药物，但是仍然有大约1/3的病人对目前的抗癫痫药物有抵抗作用，而且大约有40％病人在服用抗癫痫药物时会产生副反应{Alves，M.等(2018)，Front Pharmacol 9:193}。因而需要筛选特异的作用于致病离子通道并恢复其功能的化合物来精准化治疗癫痫疾病。
几十年来的研究表明大脑内部兴奋性和抑制性输入的不平衡是导致癫痫发生的主要原因。离子通道是神经元兴奋的分子基础，因而通道功能异常会导致癫痫的发生；这些通道功能的改变会增加兴奋性神经元的活性或降低抑制性神经元的活性。
线虫中表达许多离子通道并控制很多重要的生理功能，这些离子通道包括：电压门控的钾通道，钙离子激活的钾通道，电压门控的钙通道，环核苷酸门控离子通道和瞬时受体电位通道等。而且线虫和人的同源离子通在氨基酸序列方面具有保守性，尤其体现在通道的跨膜区域；而且这些离子通道的生物学合成调控机制在线虫和人中同样非常保守(Chen et al.,2015；Li et al.,2017；Petersen et al.,2004)。秀丽隐杆线虫(Caenorhabditis elegans)是生物学研究中的模式生物。之前，人们用线虫基因的启动子驱动人的离子通道(hERG)直接表达在线虫中，转基因线虫并不会产生行为改变，进一步研究发现全长的人离子通道在线虫细胞中并不能成功运输到细胞质膜，滞留在细胞质中。
在本发明人在先的研究工作中，制备了用于筛选hERG钾离子通道激动剂和毒性检测的转基因线虫模型，利用的是ERG家族钾离子通道UNC-103以及人hERG通道，选择两者的适当区域进行融合，获得融合蛋白。但是，尽管这一融合蛋白能够被应用于筛选hERG钾离子通道激动剂等，本领域仍然需要进一步探寻合适的靶蛋白，开发针对其它疾病的筛选模型。
发明内容
本发明的目的在于提供筛选人钾离子通道调节剂的方法。
在本发明的第一方面，提供钾离子通道蛋白的融合蛋白，其具有N端区、跨膜区以及C端区，包括：(a)线虫KQT-1(较佳地为其b亚型)的N端区、人KCNQ2(较佳地为其d亚型)的S1-S6跨膜区和线虫KQT-1的C端区；或(b)线虫SHK-1的N端区、人KV1.2的S1-S6跨膜区和C端区；或(c)人BK的N端区和跨膜区S0-S6、线虫SLO-1的C端区。
在一个优选例中，所述人KCNQ2蛋白的氨基酸序列如SEQ ID NO:1所示；所述线虫KQT-1蛋白的氨基酸序列如SEQ ID NO:2所示。
在另一优选例中，所述人KV1.2蛋白的氨基酸序列如SEQ ID NO:7所示；所述线虫SHK-1蛋白的氨基酸序列如SEQ ID NO:8所示。
在另一优选例中，所述人BK蛋白的氨基酸序列可如SEQ ID NO:13所示；所述线虫SLO-1蛋白的氨基酸序列可如SEQ ID NO:14所示。
在另一优选例中，(a)中：所述线虫KQT-1的N端区包含其N端第1～145位氨基酸残基中长120～145个氨基酸残基的序列(如为其N端第2、4、6、8、10、12位到第122、124、126、128、130、132、134、136、138、140、142、144位氨基酸残基的序列)；较佳地为包含N端第1～130位氨基酸残基或由第1～130位氨基酸残基组成的序列；所述人KCNQ2的S1-S6跨膜区包含自其第66～75氨基酸残基中的任一位起至第347～360位氨基酸中的任一位氨基酸残基的序列(如起始于第68、70、72、74位，终止于第348、350、352、354、356、358位)；较佳地为包含第75～347位氨基酸残基或由第75～347位氨基酸残基组成的序列；所述线虫KQT-1的C端区包含其C端第379～739位氨基酸残基中长330～361个氨基酸残基的序列；较佳地为其C端第379～406氨基酸残基中的任一位起至第730～739位氨基酸中的任一位氨基酸残基的序列(如起始于第381、385、389、393、396、399、402位，终止于第732、734、736、738位)；更佳地为至少包含C端第405～739位氨基酸残基或由第405～739位氨基酸残基组成的序列。
在另一优选例中，所述融合蛋白为线虫KQT-1第1～130位氨基酸残基、人KCNQ2第75～347位氨基酸残基与KQT-1蛋白C端第405～739氨基酸残基组成的融合蛋白。
在另一优选例中，(a)中所述的跨膜区序列中存在通道功能获得性突变，其中所述突变包括(但不限于)：在人KCNQ2蛋白第201位上的R201C突变。
在另一优选例中，所述的融合蛋白(KCNQ2chimera)的氨基酸序列为SEQ ID NO:3所示；其cDNA序列为SEQ ID NO:5所示；或，含有功能获得性突变的融合蛋白(KCNQ2chimera /R201C)的氨基酸序列为SEQ ID NO:4所示，其cDNA序列为SEQ ID NO:6所示。
在另一优选例中，(b)中：所述线虫SHK-1的N端区包含其N端第1～260位氨基酸残基之间，长240～260个氨基酸残基的序列(如为SHK-1的N端第2、3、4、5、6、7、8、9、10、11位到第245、246、247、248、249、250、251、252、253、254或255位氨基酸残基的序列)，较佳地为包含第1～245位氨基酸残基，或由第1～245位氨基酸残基组成的序列；所述人KV1.2的S1-S6跨膜区和C端区包含其第135～449位氨基酸残基之间，长为295～315个氨基酸残基的片段；较佳地为其第135～150氨基酸残基中的任一位起到第435～449位氨基酸中的任一位氨基酸残基的序列(如起始于第136、138、140、142、144、146、148位，终止于第436、438、440、442、444、446、448位)；更佳地为至少包含其第146～435位氨基酸残基或由其第146～435位氨基酸残基组成。
在另一优选例中，所述融合蛋白为线虫SHK-1的N端区第1～245位氨基酸残基，人KV1.2第146～449位氨基酸残基组成的融合蛋白。
在另一优选例中，(b)中所述的跨膜区序列中存在通道功能获得性突变，其中所述突变包括(但不限于)：在人KV1.2蛋白第297位上的R297Q突变。
在另一优选例中，所述融合蛋白(KV1.2chimera)的氨基酸序列为SEQ ID NO:9；其cDNA序列为SEQ ID NO:11所示；或，含有功能获得性突变的融合蛋白(KV1.2chimera/R297Q)的氨基酸序列为SEQ ID NO:10所示；其cDNA序列为SEQ ID NO:12所示。
在另一优选例中，(c)中：所述人BK的N端区和跨膜区S0-S6包含其第1～604位氨基酸残基之间，长570～604个氨基酸残基的片段；较佳地为其第1～20位氨基酸残基中的任一位起到第581～604位氨基酸中的任一位氨基酸残基的序列(如起始于第2、4、6、8、10、12、14、16、18位，终止于第582、584、586、588、590、592、594、596、598、600、602位)；更佳地为至少包含其第1～581位氨基酸残基或由其第1～581位氨基酸残基组成；所述线虫SLO-1的C端区包含其第488～1140位氨基酸残基之间，长595～653个氨基酸残基的片段；较佳地为其第488～546氨基酸残基中的任一位起到第1120～1140位氨基酸中的任一位氨基酸残基的序列(如起始于第490、494、498、502、506、510、514、518、522、526、530、534、538、542位，终止于第1122、1124、1126、1128、1130、1132、1134、1136、1138位)；更佳地为至少包含其第546～1140位氨基酸残基或由其第546～1140位氨基酸残基组成。
在另一优选例中，所述融合蛋白为人BK的N端区和跨膜区S0-S6第1～581位氨基酸残基，线虫SLO-1的C端区第546～1140位氨基酸残基组成的融合蛋白。
在另一优选例中，所述的跨膜区序列中存在通道功能获得性突变，其中所述突变包括(但不限于)：在人BK蛋白第434位上的D434G突变。
在另一优选例中，(c)中所述融合蛋白(BKchimera)的氨基酸序列为SEQ ID NO:15所示，其cDNA序列为SEQ ID NO:17所示；或，含有功能获得性突变的融合蛋白(BKchimera/D434G)的氨基酸序列为SEQ ID NO:16所示，其cDNA序列为SEQ ID NO:18所示。
在另一优选例中，所述的融合蛋白还含有可被检测的蛋白质或酶(包括报告基因编码的蛋白，如荧光蛋白)。
在本发明的另一方面，提供分离的多核苷酸序列，所述多核苷酸序列选自：(a)编码前面任一项所述的融合蛋白的多核苷酸序列；和(b)与(a)的多核苷酸序列互补的序列。
在本发明的另一方面，提供一种表达构建物或表达载体，其包含所述的多核苷酸序列。
在本发明的另一方面，提供基因工程细胞或转基因线虫，其包含所述的多核苷酸序列或转入了所述的表达载体，优选地，所述转基因线虫具有以下一项或多项特征：(i)所述多核苷酸序列整合入所述转基因线虫的基因组中；(ii)所述转基因线虫表达权利要求4、8或12所述的融合蛋白；和(iii)所述转基因线虫为acs-20基因缺陷型线虫。
在另一优选例中，所述的基因工程细胞或转基因线虫中，ifd-2和/或c15c7.5基因或其编码的蛋白被下调；较佳地，所述ifd-2和/或c15c7.5基因或其编码的蛋白藉由RNAi干扰、CRISPR/Cas9基因编辑、基因敲除、抗体靶向结合的方式被下调。
在本发明的另一方面，提供前面任一项所述的融合蛋白、所述的多核苷酸、所述的表达构建物或表达载体或所述的基因工程细胞或转基因线虫的用途，用于筛选人KCNQ2、KV1.2或BK通道的抑制剂或激活剂，或用于制备筛选人KCNQ2、KV1.2或BK通道的抑制剂或激活剂的筛选模型。
在本发明的另一方面，提供一种筛选人钾离子通道抑制剂和激活剂的方法，其中所述钾离子通道为KCNQ2、KV1.2或BK通道，该方法包括：所述筛选人KCNQ2通道抑制剂和激活剂的方法包括：使待筛选的化合物与表达前面中(a)所述的融合蛋白的转基因线虫共同孵育，并观察、比较共同孵育前和共同孵育后线虫的行为表现；与共同孵育前相比，共同孵育后使共同孵育前的线虫的行为缺陷(如发育迟缓和咽喉肌肉跳动频率降低)得以改善或恢复的化合物鉴别为KCNQ2通道的抑制剂；共同孵育后使共同孵育前的线虫的行为缺陷加重的化合物鉴别为KCNQ2通道的激活剂；或
所述筛选人KV1.2通道抑制剂和激活剂的方法包括：使待筛选的化合物与表达前面(2)所述的融合蛋白的转基因线虫共同孵育，并观察、比较共同孵育前和共同孵育后线虫的行为表现；与共同孵育前相比，共同孵育后使共同孵育前的线虫的行为缺陷(如运动和/或产卵障碍)得以改善或恢复的化合物鉴别为KV1.2通道的抑制剂；共同孵育后使共同孵育前的线虫的行为缺陷加重的化合物鉴别为KV1.2通道的激活剂；或
所述筛选人BK通道抑制剂和激活剂的方法包括：使待筛选的化合物与表达前面(3)所述的融合蛋白的转基因线虫共同孵育，并观察、比较共同孵育前和共同孵育后线虫的行为表现；与共同孵育前相比，共同孵育后使共同孵育前的线虫的行为缺陷(如运动和/或产卵障碍)得以改善或恢复的化合物鉴别为BK通道的抑制剂；共同孵育后使共同孵育前的线虫的行为缺陷加重的化合物鉴别为BK通道的激活剂；
优选的，上述方法使用前面所述的转基因线虫进行筛选。
在另一优选例中，所述的候选物质包括(但不限于)：小分子化合物、针对人KCNQ2、KV1.2或BK通道设计的干扰分子、核酸抑制物、结合分子(如抗体或配体)。
在另一优选例中，所述的转基因线虫中，ifd-2和/或c15c7.5基因被下调；较佳地，藉由RNAi干扰、CRISPR/Cas9基因编辑、基因敲除、抗体靶向结合的方式下调ifd-2和/或c15c7.5基因。
在另一优选例中，在含诱导剂(如IPTG)和抗生素(如羧苄青霉素)的线虫培养基(如NGM)中接种所述靶向ifd-2和c15c7.5基因的RNAi菌，培养后(培养数天后，如1～4天，进一步如2、3天)将待筛选的化合物加入，之后(较佳地数小时后，如2～6小时，进一步如2、3、4小时)接种所述的转基因线虫(较佳地为同步化的线虫)，优选为L3-L4时期的线虫，观察(如在24-48小时后)转基因线虫行为的变化。
在本发明的另一方面，提供用于筛选人KCNQ2、KV1.2或BK通道的抑制剂或激活剂的试剂盒，其中含有：前面任一项所述的融合蛋白、所述的多核苷酸、所述的表达构建物或表达载体或所述的基因工程细胞或转基因线虫。
本发明的其它方面由于本文的公开内容，对本领域的技术人员而言是显而易见的。
附图说明
图1、人KCNQ2通道蛋白与线虫KQT-1通道蛋白氨基酸序列比对图。通道的跨膜区S1-S6用下划线标记出来，其中三角形指示通道功能增强型点突变R201C的位置。
图2、人KV1.2通道蛋白与线虫SHK-1通道蛋白氨基酸序列比对图。通道的跨膜区S1-S6用下划线标记出来，其中三角形指示通道功能增强型点突变R297Q的位置。
图3、人BK通道蛋白与线虫SLO-1通道蛋白氨基酸序列比对图。通道的跨膜区S0-S6用灰色下划线标记出来，其中三角形指示通道功能增强型点突变D434G的位置。灰色带点划线标记RCK1/2结构域；RCK1(or 2)，regulator of K+conductance domain 1(or 2)，钾电导调节结构域1或2。
图4、构建通道疾病线虫模型筛选通道化合物的策略图。
A，构建线虫-人嵌合离子通道的示意图。红色圈表示通道功能增强型点突变；黑色圈表示通道功能缺失型点突变。
B，利用转基因线虫筛选通道化合物的策略示意图。由于离子通道过表达会改变细胞的兴奋性，表达功能增强型通道的转基因线虫会产生表型缺陷，当用通道的抑制剂处理这种转基因线虫会降低通道功能、从而减缓线虫的表型缺陷。表达功能缺失型通道的转基因线虫由于过表达通道电流很小，因而不会产生表型缺陷，通道的激活剂或能特异性的修复突变通道功能缺陷的矫正剂能够提高功能缺失型通道的功能、改变线虫细胞的兴奋性，从而使转基因线虫产生新的表型缺陷。
图5、离子通道病线虫模型具有明显表型缺陷。A，构建线虫-人嵌合离子通道BKchimera/D434G，Kv1.2chimera/R297Q和KCNQ2chimera/R201C的示意图。B，三种转基因线虫的显微镜拍摄图像，标尺为200μM。C，转基因线虫的运动行为缺陷分析。D，转基因线虫的产卵行为缺陷分析。E，KCNQ2chimera/R201C转基因线虫的咽喉肌肉跳动行为分析。结果以平均值±标准误(Mean±SEM)展示，显著性以***P<0.01表示。图E用Student’s T test分析，图C和D用one-way ANOVA Dunnett’s test分析显著性。
图6、已知的通道阻断剂减缓通道疾病线虫模型的表型缺陷。acs-20；BKChimera /D434G，acs-20；Kv1.2Chimera/R297Q和acs-20；KCNQ2Chimera/R201C突变线虫在DMSO或已知通道阻断剂处理后的显微图像。白色箭头指示线虫卵，图像标尺为400μm。
具体实施方式
本发明人经过深入的研究，确定了一系列离子通道蛋白的融合蛋白，适用于在体高通量筛选调控人的KCNQ2、KV1.2和BK钾通道功能的物质(如小分子化合物等药物)。本发明还提供了编码所述融合蛋白的多核苷酸序列，相关的转基因线虫，以及相关的筛选方法和应用。为筛选药物治疗离子通道疾病提供了新的途径。
在本发明中，构建了离子通道疾病线虫模型，其可被应用于筛选调控KCNQ2、KV1.2和BK钾通道功能的化合物。将带有功能增强型点突变的线虫-人嵌合离子通道KCNQ2chimera /R201C，Kv1.2chimera/R297Q和BKchimera/D434G表达在线虫中。表达存在通道功能获得性突变的BKchimera/D434G和Kv1.2chimera/R297Q通道转基因线虫呈现出非常明显的运动和产卵障碍；通过抑制剂降低BK或KV1.2通道功能可以缓解这些转基因线虫的上述行为缺陷，而通道的激活剂能够进一步加剧线虫的表型缺陷。表达存在通道功能获得性突变的KCNQ2chimera/R201C通道的线虫呈现出非常明显的发育迟缓和线虫咽喉肌肉跳动频率减慢的表型缺陷；通过抑制剂降低KCNQ2通道功能可以缓解该转基因线虫的行为缺陷，通道的激活剂能够进一步加剧线虫的表型缺陷。因此，这是一种新的筛选调控KCNQ2、BK和KV1.2钾离子通道功能小分子化合物的方法。
如本发明所用，所述线虫可以是来自广杆线虫属的线虫，优选为秀丽隐杆线虫(Caenorhabditis elegans)。本发明所构建的转基因线虫能表达线虫-人嵌合的钾通道KCNQ2chimera/R201C，Kv1.2chimera/R297Q和BKchimera/D434G的融合蛋白。
“片段”在本文中指全长序列的一部分连续的序列。除非另外说明，所述的“N端区”是指自蛋白跨膜区之前的序列区段，该区域不包括蛋白的跨膜区。所述的“C端区”是指自蛋白跨膜区之后的序列区段，该区域不包括蛋白的跨膜区。
KCNQ是一类电压门控的钾通道，有KCNQ1-5五种亚型，其中KCNQ2和KCNQ3形成M型钾通道，它决定神经元的兴奋性阈值、放电特性等，是神经元兴奋性的重要调节器。KCNQ2或KCNQ3基因上的功能缺失突变会导致良性家族性新生儿婴儿惊厥疾病的发生{Biervert，C.等，Science 279(5349):403-406；Jentsch，T.J.(2000)，Nat Rev Neurosci 1(1):21-30.}(Biervert et al.,1998；Jentsch,2000)(Biervert et al.,1998；Jentsch,2000)(Biervert et al.,1998；Jentsch,2000)(Biervert et al.,1998；Jentsch,2000)。在优选的实施方案中，所述人KCNQ2蛋白的氨基酸序列如SEQ ID NO:1所示；相应地，用于与之融合的所述线虫KQT-1蛋白的氨基酸序列如SEQ ID NO:2所示。
KV1.1和KV1.2通道控制神经元的兴奋性，KV1.2通道上存在与癫痫性脑病、智力障碍等疾病相关的通道功能缺失或增强型突变。大电导的钙离子激活钾通道(BK)可以被电压和胞内钙离子激活，能够耦合细胞内钙信号和细胞膜去极化过程，是控制神经元兴奋性的重要分子。在优选的实施方案中，所述人KV1.2蛋白的氨基酸序列如SEQ ID NO:7所示；相应地，用于与之融合的所述线虫SHK-1蛋白的氨基酸序列如SEQ ID NO:8所示。
BK通道的功能缺失突变会导致神经元过度兴奋，进而造成颞叶癫痫以及全身性强直阵挛发作；另有研究报道BK通道上的功能增强突变(如D434G)也会导致癫痫疾病发生{Du，W.等，Nat Genet 37(7):733-738.}。癫痫发作本身也会增加BK通道电流，进一步加重疾病症状，抑制BK通道则可阻止全身强直性阵挛癫痫发作{N’Gouemo，P.(2011)."Targeting BK(big potassium)channels in epilepsy."Expert Opin Ther Targets 15(11):1283-1295.}。在优选的实施方案中，所述人BK蛋白的氨基酸序列可如SEQ ID NO:13所示；相应地，用于与之融合的所述线虫SLO-1蛋白的氨基酸序列可如SEQ ID NO:14所示。
上述的人KCNQ2蛋白、线虫KQT-1蛋白、人KV1.2蛋白、线虫SHK-1蛋白、人BK蛋白、线虫SLO-1蛋白或其片段(例如N端区片段，跨膜区片段，C端区片段)，除了本发明所附序列表提供的具体序列以外，还可以是与所述蛋白或其片段相同功能的序列变异形式(变体)。这些变异形式包括但并不限于：若干个(通常为1-50个，较佳地1-30个，更佳地1-20个，最佳地1-10个，还更佳如1-8个、1-5个)氨基酸的缺失、插入和/或取代，以及在C末端和/或N末端添加或缺失一个或数个(通常为20个以内，较佳地为10个以内，更佳地为5个以内)氨基酸。例如，在本领域中，用性能相近或相似的氨基酸进行取代时，通常不会改变蛋白质的功能。又比如，在C末端和/或N末端添加或缺失一个或数个氨基酸通常也不会改变蛋白质的功能。本发明也包含与上述蛋白的序列有80％以上(较佳地85％，更佳地90％以上，更佳95％以上，如98％，99％)同源性且保留有所述蛋白功能的衍生的蛋白。但是，所述的变异形式(变体)及其片段、衍生物和类似物的氨基酸序列中，对应于本发明中所主张的功能获得性突变位点，需为本发明所指定的突变而不能作随意的改变，也即在人KCNQ2蛋白第201位上突变为C，在人KV1.2蛋白第297位上突变为Q，在人BK蛋白第434位上突变为G。
作为本发明的优选方式，建立了KCNQ2chimera/R201C融合蛋白，包括作为该融合蛋白N端的来自线虫KQT-1钾通道N端第1～145位氨基酸残基之间、长120～145个氨基酸残基的片段，至少包含人KCNQ2跨膜区S1～S6的片段，以及作为该融合蛋白C端的来自线虫KQT-1钾通道C端第379～739位氨基酸残基之间、长330～361个氨基酸残基的片段。
在进一步的优选方式中，所述线虫KQT-1蛋白N端长120～145个氨基酸残基的片段由KQT-1蛋白N端第2、4、6、8、10、12位到第120、121、122、123、124、125、126、127、128、129、130、131、132、133、134、135、136、137、138、139、140、141、142、143、144或145位氨基酸残基组成，优选为第1位到第125、126、127、128、129、130、131、132、133、134、135、136、137、138、139或140位氨基酸残基，更优选至少包含第1～130位氨基酸残基，或由第1～130位氨基酸残基组成。
在进一步的优选方式中，所述线虫KQT-1通道蛋白C端第379～739位氨基酸残基之间长330～361个氨基酸残基的片段可以是例如KQT-1蛋白第379～406位氨基酸残基中的任一位起到第730～739位氨基酸中的任一位所形成的片段，如第379～730位氨基酸残基，第379～739位氨基酸残基，第395～735位氨基酸残基，第405～730位氨基酸残基，或第405～739位氨基酸残基；优选地，所述片段至少含有第405～739位氨基酸残基。
本文所述的人KCNQ2通道蛋白的片段至少包含KCNQ2的S1～S6跨膜区。所述KCNQ2的含S1～S6跨膜区片段至少包括KCNQ2第66～360位氨基酸残基，优选至少包括KCNQ2第75～347位氨基酸残基片段，或由第75～347位氨基酸残基组成。
在某些实施方案中，KCNQ2的片段是第66～360位氨基酸之间长至少250～295个氨基酸残基的片段，例如可以是第66～78位氨基酸残基中的任一位起到第345～360位氨基酸中的任一位所形成的片段，如第66～345位氨基酸残基，第66～360位氨基酸残基，第78～360位氨基酸残基，第75～347位氨基酸残基等；优选地，所述片段至少含第75～347位氨基酸残基。
在优选的实施方案中，该KCNQ2或其含跨膜氨基酸序列的片段中存在通道功能获得性的突变。这些突变包括但不限于在201位上的突变。优选的突变为取代突变，例如R201C，这些通道功能获得性突变能使离子通道的激活曲线向更负的电压偏移，使通道能在较低的膜电位条件下开放，可造成动物行为异常，有利于筛选调控KCNQ2离子通道功能的化合物。
在某些实施方案中，本文的融合蛋白由KQT-1蛋白N端第1～130位氨基酸残基、人KCNQ2第75～347位氨基酸残基、和KQT-1蛋白C端第405～739位氨基酸残基组成。除此之外，应理解的是，虽然本文没有一一列出，但本文所述的任意KQT-1蛋白N端片段可与本文所述的任意KCNQ2片段及任意KQT-1蛋白C端片段组合，以形成本发明的融合蛋白。
适用于本文的人KCNQ2的氨基酸序列可如SEQ ID NO:1所示；适用于本文的KQT-1的氨基酸序列可如SEQ ID NO:2所示。文中所述的KCNQ2和KQT-1全长序列或其片段的氨基酸位置/编号均以SEQ ID NO:1和2的氨基酸位置/编号为基准。
作为本发明的优选方式，建立了KV1.2chimera/R297Q融合蛋白，包括作为该融合蛋白N端的来自线虫SHK-1钾通道N端第1～260位氨基酸残基之间、长240～260个氨基酸残基的片段，至少包含人KV1.2通道蛋白跨膜区S1～S6和C端的片段。
在优选的实施方案中，所述线虫SHK-1蛋白N端长240～260个氨基酸残基的片段由SHK-1蛋白N端第2、3、4、5、6、7、8、9、10、11位到第240、241、242、243、244、245、246、247、248、249、250、251、252、253、254、255、256、257、258、259或260位氨基酸残基组成，优选N端第1、2、3、4、5、6、7、8、9、10、11位到第245、246、247、248、249、250、251、252、253、254或255位氨基酸残基，更优选至少包含第1～245位氨基酸残基，或由第1～245位氨基酸残基组成。
本文所述的人KV1.2通道蛋白片段至少包含KV1.2的S1～S6跨膜区和C端。所述KV1.2蛋白的含S1～S6跨膜区和C端片段至少包括KV1.2第135～449位氨基酸残基，优选至少包括KCNQ2第146～435位氨基酸残基，或由第146～449位氨基酸残基组成。
在某些实施方案中，KCNQ2的片段是第135～449位氨基酸之间长至少290～315个氨基酸残基的片段，例如可以是第135～150位氨基酸残基中的任一位起到第435～449位氨基酸中的任一位所形成的片段，如第135～449位氨基酸残基，第135～435位氨基酸残基，第140～440位氨基酸残基，第145～449位氨基酸残基等；优选地，所述片段至少含第146～435位氨基酸残基。
在优选的实施方案中，该KV1.2通道或其含跨膜氨基酸序列的片段中存在通道功能获得性的突变。这些突变包括但不限于在297位上的突变。优选的突变为取代突变，例如R297Q，这些通道功能获得性突变能使离子通道的激活曲线向更负的电压偏移，使通道能在较低的膜电位条件下开放，可造成动物行为异常，有利于筛选调控KV1.2离子通道功能的化合物。
在某些实施方案中，本文的融合蛋白由线虫SHK-1通道蛋白N端第1～245位氨基酸残基、人KV1.2通道蛋白第146～449位氨基酸残基组成。除此之外，应理解的是，虽然本文没有一一列出，但本文所述的任意SHK-1蛋白N端片段可与本文所述的任意KV1.2通道蛋白包含跨膜区和C段的片段组合，以形成本发明的融合蛋白。
适用于本文的人KV1.2通道蛋白的氨基酸序列可如SEQ ID NO:7所示；适用于本文的线虫SHK-1通道蛋白的氨基酸序列可如SEQ ID NO:8所示。文中所述的KV1.2和SHK-1全长序列或其片段的氨基酸位置/编号均以SEQ ID NO:7和8的氨基酸位置/编号为基准。
作为本发明的优选方式，建立了BKchimera/D434G融合蛋白，包括作为该融合蛋白N端和跨膜区S0-S6的来自人BK钾通道蛋白第1～604位氨基酸残基之间、长570～604个氨基酸残基的片段，和至少包含线虫SLO-1通道蛋白C端第488～1140位氨基酸残基之间、长595～653个氨基酸残基的片段。
所述人BK钾通道蛋白N端和S0-S6跨膜区长570～604个氨基酸残基的片段由SHK-1蛋白N端第12、4、6、8、10、12、14、16、18位到第570、571、572、573、574、575、576、577、578、579、580、581、582、583、584、585、586、587、588、589、590、591、592、593、594、595、596、597、598、599、600、601、602、603或604位氨基酸残基组成，优选第1位到第575、576、577、578、579、580、581、582、583、584、585、586、587、588、589、590、591、592、593、594或595位氨基酸残基，更优选至少包含第1～581位氨基酸残基，或由第1～581位氨基酸残基组成。
本文所述的线虫SLO-1通道蛋白C端片段至少包含SLO-1第448～1140位氨基酸残基，优选至少包括KCNQ2第546～1120位氨基酸残基，或由第546～1120位氨基酸残基组成。
所述线虫SLO-1通道蛋白C端第448～1140位氨基酸残基之间长595～653个氨基酸残基的片段，例如可以是SLO-1蛋白第488～546位氨基酸残基中的任一位起到第1120～1140位氨基酸中的任一位所形成的片段，如第488～1120位氨基酸残基，第488～1140位氨基酸残基，第546～1120位氨基酸残基，第540～1140位氨基酸残基，或第546～1140位氨基酸残基；优选地，所述片段至少含有第546～1120位氨基酸残基。
在优选的实施方案中，该BK通道或其含跨膜氨基酸序列的片段中存在通道功能获得性的突变。这些突变包括但不限于在434位上的突变。优选的突变为取代突变，例如D434G，这些通道功能获得性突变能使离子通道的激活曲线向更负的电压偏移，使通道能在较低的膜电位条件下开放，可造成动物行为异常，有利于筛选调控BK钾离子通道功能的化合物。
在某些实施方案中，本文的融合蛋白由人BK通道蛋白N端和S0-S6跨膜区第1～581位氨基酸残基、和线虫SLO-1通道蛋白第546～1140位氨基酸残基组成。除此之外，应理解的是，虽然本文没有一一列出，但本文所述的任意人BK蛋白N端和S0-S6跨膜区片段可与本文所述的任意线虫SLO-1通道蛋白C段的片段组合，以形成本发明的融合蛋白。
适用于本文的人BK通道蛋白的氨基酸序列可如SEQ ID NO:13所示；适用于本文的线虫SLO-1通道蛋白的氨基酸序列可如SEQ ID NO:14所示。文中所述的BK和SLO-1全长序列或其片段的氨基酸位置/编号均以SEQ ID NO:13和14的氨基酸位置/编号为基准。
本领域技术人员公知，在基因克隆操作中常常需要设计合适的酶切位点，这势必在所表达的蛋白末端引入了一个或多个不相干的氨基酸残基，但这并不影响目的蛋白的活性。为了构建融合蛋白，需要在目的蛋白N端或C端加上或是促进蛋白分泌表达的信号肽、或是由利于重组蛋白纯化的如谷胱甘肽S-转移酶(GST)、6His或Flag等标签、或是指示蛋白表达和定位的荧光蛋白如绿色荧光蛋白、黄色荧光蛋白、和红色荧光蛋白等。应理解，这些氨基酸序列的存在不会影响到所需目的蛋白的活性。这些蛋白质或多肽可经由接头序列与目的蛋白中的C端或N端片段连接。接头序列可以是本领域常用的接头序列，例如含GS的接头序列，只要其不影响到融合蛋白整体的生物学功能即可。
在某些实施方案中，本发明KCNQ2chimera/R201C,Kv1.2chimera/R297Q和BKchimera/D434G融合蛋白中所述线虫通道蛋白N端和C端片段、以及人通道跨膜区片段中可存在一个或多个(例如25个以内、20个以内、15个以内、10个以内、5个以内)氨基酸突变(包括取代、缺失和/或插入数个氨基酸)。氨基酸突变优选的是保守性突变，即用性能相近或相似的氨基酸进行保守性取代的突变，通常不会改变蛋白质或多肽的功能。“性能相近或相似的氨基酸”包括多种具有相似侧链的氨基酸残基的家族，例如具有碱性侧链的氨基酸家族(包含赖氨酸、精氨酸、组氨酸)、具有酸性侧链的氨基酸家族(包含天冬氨酸、谷氨酸)、具有不带电荷的极性侧链的氨基酸家族(包含甘氨酸、天冬酰胺、谷氨酰胺、丝氨酸、苏氨酸、酪氨酸、半胱氨酸)、具有非极性侧链的氨基酸家族(包含丙氨酸、缬氨酸、亮氨酸、异亮氨酸、脯氨酸、苯丙氨酸、甲硫氨酸、色氨酸)、具有β-分支侧链的氨基酸家族(包含苏氨酸、缬氨酸、异亮氨酸)和具有芳香侧链的氨基酸家族(包含酪氨酸、苯丙氨酸、色氨酸、组氨酸)。因此，优选的突变所得的各片段或融合蛋白本身仍然保留本文所述通道蛋白的生物学活性。例如，突变所得的KQT-1蛋白N端片段和C端片段保留野生型KQT-1的N和C端在线虫细胞中正常上膜所具有的生物学活性；突变所得的KCNQ2的片段保留了S1～S6的生物学功能，或保留了与本文所述的KCNQ2R201C突变体相同的生物学功能(通道功能获得性)。
本文包括所述融合蛋白的编码序列。例如，SEQ ID NO:5显示了由线虫KQT-1蛋白N端第1～130位氨基酸残基、人KCNQ2第75～347位氨基酸残基、以及KQT-1蛋白C端第405～739氨基酸残基组成的KCNQ2chimera融合蛋白的编码cDNA序列；SEQ ID NO:6是存在R201C突变的前述融合蛋白的编码cDNA序列。SEQ ID NO:11显示了由SHK-1蛋白N端第1～245位氨基酸残基、人KV1.2蛋白第146～449位氨基酸残基片段组成的KV1.2chimera融合蛋白的编码cDNA序列；SEQ ID NO:12是存在R297Q突变的前述融合蛋白的编码cDNA序列。SEQ ID NO:17显示了由人BK通道蛋白N端和S1～S6跨膜区第1～581位氨基酸残基和线虫SLO-1蛋白C端第546～1140氨基酸残基片段组成的BKchimera融合蛋白的编码cDNA序列；SEQ ID NO:18是存在D434G突变的前述融合蛋白的编码cDNA序列。本文包括SEQ ID NO:5、6、11、12、17和18的简并异构体，即编码相同的氨基酸序列但核苷酸序列不同的序列。本文也包括本发明编码序列的互补序列。如前述所述，当本文的融合蛋白中还包括用于检测目的的蛋白质或酶时，本文的编码序列优选还包括编码该蛋白质或酶的报告基因。
为获得本文中所述的线虫-人嵌合通道蛋白的编码序列，先通过PCR扩增各cDNA片段、再用DNA重组法将这些片段按正确顺序连接在一起构建到表达载体中。根据线虫或人的通道蛋白的开放阅读框序列设计引物用来PCR扩增各cDNA片段，用市售的cDNA库或按本领域技术人员已知的常规方法所制备的cDNA库作为PCR扩增模板。
本文构建克隆用的载体为pPD95.75表达载体(序列如SEQ ID NO:19所示)。用线虫kqt-1基因的启动子驱动表达KCNQ2chimera/R201C融合蛋白；用线虫shk-1基因的启动子驱动表达KV1.2chimera/R297Q融合蛋白；用线虫slo-1基因的启动子驱动表达BKchimera/D434G融合蛋白。同样可采用本领域技术人员熟知的在线虫中常用的表达载体构建包含所述融合蛋白和相应启动子序列的克隆载体。克隆载体的构建方法包括体外重组DNA技术、DNA合成技术、体内重组技术等。
在本文中，通过显微注射的方式将本文所述表达融合通道蛋白的载体注射到线虫性腺中，处于多合胞体阶段的卵细胞在成熟过程中会把表达载体包裹进入卵细胞中，进而获得相应的转基因线虫。然后通过本领域周知的整合方法使表达载体整合入线虫染色体中获得稳定表达融合蛋白的转基因线虫，这类整合方法包括但不限于UV/TMP方法。优选的，获得稳定表达本文所述融合蛋白的转基因整合线虫后，通过与野生型N2线虫回交数4～6次，去除UV/TMP整合过程中产生的背景基因突变。此外，由于线虫具有较厚的保护性体壁，同时线虫肠道又可以主动排除外源有毒物质，这使得大多数化合物较难进入线虫体内到达作用的靶点位置。因此，进一步优选的是，本文的线虫是acs-20基因缺陷型的突变体，例如其acs-20基因被敲除，该线虫体壁被破坏，允许更多的小分子化合物进入线虫体内。例如，通过将稳定表达本文所述融合蛋白的转基因整合线虫在和野生型N2线虫回交后获得的线虫与acs-20基因缺陷型线虫(如本领域周知的acs-20(tm3232))杂交，可获得本文的表达融合通道蛋白的acs-20基因缺陷型线虫；即acs-20；BKChimera/D434G，acs-20；Kv1.2Chimera/R297Q和acs-20；KCNQ2Chimera/R201C三种转基因线虫，待用于通道化合物的线虫筛选。
本发明中，线虫的行为缺陷通常指运动、产卵障碍，以及生长迟缓等。线虫的运动行为通过Olympus显微镜录像系统(cellSense Standard)记录线虫在线虫培养基NGM平板(无OP50大肠杆菌)上的自由运动，之后用wrMtrack软件分析线虫的运动速度。刚成年的野生型N2线虫的平均运动速度约为0.18mm/s，当转基因线虫的平均运动速度低于0.10mm/s时，可认为所述转基因线虫存在运动障碍。更优选地，运动缺陷的转基因线虫的平均运动速度(单位为mm/s)低于0.09、0.08、0.07、0.06、0.05、0.04、0.03、0.02、0.01或更低。在产卵能力方面，L4晚期的野生型N2线虫在30小时内平均可产约60枚卵。因此，当L4晚期的转基因线虫在30小时内产卵数量低于35枚、优选低于30枚、更优选低于25枚、更优选低于20枚、更优选低于15枚、更优选低于10枚、更优选低于5枚时，可认为该转基因线虫存在产卵障碍。在20℃培养条件下野生型N2线虫的生长周期平均为3.5天，即从产下的线虫卵发育到可以可以产卵的成年线虫时期的阶段为3.5天；转基因线虫的生长周期长于4天、更优选的为5天或更多，可认为该转基因线虫存在发育迟缓缺陷。存在上述运动和/或产卵等缺陷的转基因线虫可用于通道化合物筛选。
可利用本发明的转基因线虫筛选KCNQ2、KV1.2和BK钾通道的抑制剂或激活剂。例如，表达存在通道功能获得性突变的融合蛋白的acs-20；BKChimera/D434G或acs-20；Kv1.2Chimera/R297Q线虫存在运动和产卵的行为缺陷，在与待筛选的物质接触后，若该转基因线虫的行为缺陷(如运动和/或产卵障碍)得以改善或恢复，则表明该待筛选物质为BK或KV1.2钾通道的抑制剂。产卵行为便于观察统计，因而化合物筛选主要采用产卵行为指标。所述“改善”指在转基因线虫在30小时内的产卵数量多于未用待筛选物质处理的转基因线虫在30小时内的产卵数量(优选至少多2枚，更优选至少至少多5枚，更优选至少多10枚)。
通常，筛选可在含有线虫培养基NGM的48孔板中进行，在培养基中可加入诱导剂(如1mM IPTG)和抗生素(如羧苄青霉素，25ug/ml)用于诱导线虫用RNAi菌表达双链dsRNA敲低基因的表达。在NGM平板中接种靶向ifd-2和c15c7.5基因的RNAi菌(RNAi菌来源于SourceBioScience公司的C.elegans RNAi Library，构建ifd-2RNAi质粒的正向引物为AATGGTTGATTTCGTTTCGC(SEQ ID NO:46)，反向引物为TGCAGTCGCTTTTCACTTTG(SEQ ID NO:47)；构建c15c7.5RNAi质粒的正向引物为GATCCCTTCCAAGAACAAGAGTT(SEQ ID NO:48)，反向引物为ATCGTCTTTTTCTCCATTTCCTC(SEQ ID NO:49))，它们可以降低线虫肠道排除外源药物能力，增加小分子化合物在线虫体内的累积。接种RNAi菌数天(如1～3天)后将孵育待测混合物加入上述培养基中，再过数小时(如2～8小时)后接种所述的转基因线虫，优选为L3-L4时期的线虫，在24-36小时后观察转基因线虫行为(如产卵和发育迟缓)的变化，确定化合物是否是KCNQ2、KV1.2和BK钾通道的抑制剂。
下面结合具体实施例，进一步阐述本发明。应理解，这些实施例仅用于说明本发明而不用于限制本发明的范围。下列实施例中未注明具体条件的实验方法，通常按照常规条件如J.萨姆布鲁克等编著，分子克隆实验指南，第三版，科学出版社，2002中所述的条件，或按照制造厂商所建议的条件。
方法
线虫品系
野生型线虫品系N2获自美国线虫遗传中心，突变体线虫acs-20(tm3232)获自日本线虫资源库。线虫饲养于含有OP50大肠杆菌的NGM平板上，于20℃恒温培养箱中培养。
质粒构建
用于转基因的质粒KCNQ2chimera/R201C经过PCR和重组法构建而成。首先从线虫或人的cDNA文库中分别PCR扩增出以下cDNA片段：①线虫KQT-1钾通道(b亚型)N端第1-130位置氨基酸对应的cDNA片段、②人KCNQ2钾通道(d亚型)S1-S6跨膜区第75-347位置氨基酸对应的cDNA片段、和③线虫KQT-1钾通道(b亚型)C端第405-739位置氨基酸对应的cDNA片段。通过DNA重组法将这三个片段按正确的顺序(按照5’至3’端①-②-③)插入pPD95.75表达载体(Addgene)的多克隆位点中，使其携带编码融合蛋白KCNQ2chimera的完整cDNA片段。接着以线虫基因组为模板PCR扩增约4Kb的kqt-1基因的启动子序列，通过酶切连接的方法将启动子序列插入含有KCNQ2chimera cDNA的pPD95.75载体中，获得Pkqt-1::KCNQ2chimera::gfp质粒。最后，通过点突变实验在融合蛋白引入一个R201C氨基酸替换的突变质粒，即Pkqt-1::KCNQ2chimera/R201C::gfp表达质粒。
用于转基因的质粒KV1.2chimera/R297Q经过PCR和重组法构建而成。首先从线虫或人的cDNA文库中分别PCR扩增出以下cDNA片段：①线虫SHK-1钾通道(b亚型)N端第1-245位置氨基酸对应的cDNA片段、②人KV1.2钾通道S1-S6跨膜区和C端第146-449位置氨基酸对应的cDNA片段。通过DNA重组法将上述片段按正确的顺序(按照5’至3’端①-②)插入pPD95.75表达载体的多克隆位点中，使其携带编码融合蛋白KV1.2chimera的完整cDNA片段。接着以线虫基因组为模板PCR扩增约4.1Kb的shk-1基因的启动子序列，通过酶切连接的方法将启动子序列插入含有KV1.2chimera cDNA的pPD95.75载体中，获得Pshk-1::Kv1.2chimera::gfp质粒。最后通过点突变实验在融合蛋白引入一个R297Q氨基酸替换的突变质粒，即Pshk-1::Kv1.2chimera /R297Q::gfp表达质粒。
用于转基因的质粒BKchimera/D434G经过PCR和重组法构建而成。首先从线虫或人的cDNA文库中分别PCR扩增出以下cDNA片段：①人BK钾通道(b亚型)N端和S0-S6跨膜区第1-581位置氨基酸对应的cDNA片段、②线虫SLO-1钾通道(a亚型)C端第546-1140位置氨基酸对应的cDNA片段。通过DNA重组法将这上述片段按正确的顺序(按照5’至3’端①-②)插入pPD95.75表达载体的多克隆位点中，使其携带编码融合蛋白BKchimera的完整cDNA片段。接着以线虫基因组为模板PCR扩增约2Kb的slo-1基因的启动子序列，通过酶切连接的方法将启动子序列插入含有BKchimera cDNA的pPD95.75载体中，获得Pslo-1::BKchimera::gfp质粒。最后通过点突变实验在融合蛋白引入一个D434G氨基酸替换的突变质粒，即Pslo-1::BKchimera /D434G::gfp表达质粒。
上述所有质粒都经过测序反应验证核苷酸序列的正确性。上述构建过程使用到的引物序列如下所示：
kqt-1启动子引物序列：
正向：GCATGCCTGCAGGCCGAAGGAATTATCACAGCA(SEQ ID NO:20)
反向：CCCCCCGGGATCTTCAAAATTGAAACTTTGACTG(SEQ ID NO:21)
KQT-1通道N端(第1-130位氨基酸)引物序列：
正向：TTCAATTTTGAAGATCCCGGGATGTCTCAAGCTTCACAAG(SEQ ID NO:22)
反向：CAGCTTGCGAAATCGTTGATCAGTACG(SEQ ID NO:23)
KCNQ2通道跨膜区S1-S6(第75-347位氨基酸)引物序列：
正向：AACGATTTCGCAAGCTGCAGAATTTCC(SEQ ID NO:24)
反向：GCAGCAGAGTAGAATCTCCAGGCCGACTG(SEQ ID NO:25)
KQT-1通道C端(第405-739位氨基酸)引物序列：
正向：GAGATTCTACTCTGCTGCACCGGAATCCAC(SEQ ID NO:26)
反向：CTCATTTTTTCTACCGGTACCCTACAACTTGAAGTACCGGGAC C(SEQ ID NO:27)
KCNQ2通道功能增强型R201C点突变引物序列：
正向：GCTCCGGAGCCTGTGCTTCCTGCAGATTC(SEQ ID NO:28)
反向：GAATCTGCAGGAAGCACAGGCTCCGGAGC(SEQ ID NO:29)
shk-1启动子引物序列：
正向：CATGCATGCACAGCGTTCTCACCTAAAAGGCC(SEQ ID NO:30)
反向：CGGGATCCCGGGTCGTAATGACCGCCTG(SEQ ID NO:31)
SHK-1通道N端(第1-245位氨基酸)引物序列：
正向：CAGGTCGACTCTAGAGGATCCATGCGATTCGGTGGTCAAC(SEQ ID NO:32)
反向：CTTGTCTCTGGGTTTTATTATTTGGCATCATCAC(SEQ ID NO:33)
KV1.2通道跨膜区S1-S6和C端(第146-449位氨基酸)引物序列：
正向：AATAAAACCCAGAGACAAGTGTGGCTTC(SEQ ID NO:34)
反向：CTCATTTTTTCTACCGGTACCCTGACATCAGTTAACATTTTGG(SEQ ID NO:35)
KV1.2通道功能增强型R297Q点突变引物序列：
正向：CCTCCGTGTCATCCAGTTGGTAAGAGTC(SEQ ID NO:36)
反向：GACTCTTACCAACTGGATGACACGGAGG(SEQ ID NO:37)
slo-1启动子引物序列：
正向：CGGGATCCAGCGCAGACAGCTAGAGTGTCTG(SEQ ID NO:38)
反向：CGGGATCCCGTGTCGACGAAGGGTCCAC(SEQ ID NO:39)
BK通道N端和S0-S6跨膜区(第1-581位氨基酸)引物序列：
正向：CAGGTCGACTCTAGAGGATCCATGGCAAATGGTGGCGG(SEQ ID NO:40)
反向：GACGTTTTGAATGACCTCATGGAGAAGAGG(SEQ ID NO:41)
SLO-1通道C端(第546-1140位氨基酸)引物序列：
正向：GGTCATTCAAAACGTCACAAACCACACCG(SEQ ID NO:42)
反向：ATTCTACGAATGACCGGTACCCTAAAAGTGTCGTTTGCCCG(SEQ ID NO:43)
BK通道功能增强型D434G点突变引物序列：
正向：GCACAAGGACCGGGGTGACGTCAATGTGG(SEQ ID NO:44)
反向：CCACATTGACGTCACCCCGGTCCTTGTGC(SEQ ID NO:45)
转基因线虫构建
将Pkqt-1::KCNQ2chimera/R201C::gfp和Pkqt-1::mCherry质粒(Pkqt-1启动子后加上报告基因mcherry)显微注射入野生型N2线虫中获得KCNQ2chimera/R201C转基因线虫。
将Pshk-1::Kv1.2chimera/R297Q::gfp和Pshk-1::mCherry质粒(Pshk-1启动子后加上报告基因mcherry)显微注射入野生型N2线虫中获得Kv1.2chimera/R297Q转基因线虫。
将Pslo-1::BKchimera/D434G::gfp和Pslo-1::mCherry质粒(Pslo-1启动子后加上报告基因mcherry)显微注射入野生型N2线虫中获得BKchimera/D434G转基因线虫。
红色mCherry荧光蛋白在线虫细胞质中表达，可用于线虫荧光挑选和标记线虫细胞质部分来反应融合gfp荧光蛋白的通道在线虫中运输状况。瞬时传代的线虫用标准的UV/TMP方法(Jiang et al.(2014)The Journal of Neuroscience 34:3947-3958)进行整合使表达质粒插入线虫染色体中，获得稳定表达的转基因线虫；之后再与野生型N2线虫反复回交4～6次以去除背景基因突变，接着与acs-20(tm3232)突变体线虫杂交获得可以进行化合物筛选的acs-20；BKChimera/D434G，acs-20；Kv1.2Chimera/R297Q和acs-20；KCNQ2Chimera/R201C转基因线虫。
线虫行为检测
线虫行为学实验所用线虫的时期一致，即经同步化培养的L4晚期或刚成年雌雄同体线虫。线虫行为学实验都是采用盲做的方式，线虫培养和实验均在20℃进行，每种行为学实验至少重复3次。线虫产卵行为实验具体如下：将L4晚期雌雄同体线虫每条挑到单独的含OP50大肠杆菌的NGM平板上培养，待培养30小时后将线虫挑出来，计数NGM平板上小虫子和卵的数目，即线虫在该时间段内的产卵数目。线虫运动行为实验具体如下：提前24小时准备实验用NGM平板(35mm平板加2ml NGM)，室温放置晾干，将约25只刚成年的雌雄同体线虫挑到一个NGM平板上(注意不要附带OP50到检测用NGM平板)，恢复1min后，用Olympus显微镜录像系统(cellSense Standard)记录线虫的自由运动1min，之后用wrMtrack软件分析线虫的运动速度。线虫咽喉肌肉跳动行为实验具体如下：先将线虫从培养箱中取出，在20℃室温静置10min再检测行为，在解剖显微镜下计数正在进食线虫的咽喉肌肉在10s内跳动的次数。
小分子化合物处理转基因线虫
化合物处理转基因线虫实验在48孔板进行，每个孔中加入500μl NGM培养基(含有1mM IPTG，异丙基硫代半乳糖苷；25μg/ml羧苄青霉素)，室温放置2天晾干后接种靶向ifd-2和c15c7.5基因的RNAi菌种于NGM培养基上，2天后把溶解在DMSO中的小分子化合物(终浓度达到20μM)加入到RNAi菌上，4小时后将同步化得到的L3～L4时期的acs-20；BKChimera/D434G或acs-20；Kv1.2Chimera/R297Q或acs-20；KCNQ2Chimera/R201C转基因线虫接种到已加化合物分子的48孔板NGM上，每个孔线虫约20只。36小时后于显微镜下观察线虫表型来初步确定目标化合物。
实施例1、构建离子通道病线虫模型
线虫表达许多离子通道，它们控制许多重要的生理功能，如线虫的运动、产卵和机械感受等。线虫的离子通道和人的同源通道在蛋白序列上保守性很高，尤其体现在通道的跨膜区域(图1，2，3)。
离子通道的N端和C端结构能够与胞内的许多蛋白发生相互作用，进而调控通道的表达和运输。为了能够在线虫中成功表达人类离子通道，本发明人构建了线虫-人嵌合的离子通道，如图4A的策略，即：构建一个包含线虫离子通道N端和C端部分以及人类离子通道跨膜区的嵌合的离子通道。由于通道化合物(阻断剂和激活剂)通常直接作用于通道的跨膜区，因此本发明人推测这种嵌合的离子通道既能够在线虫中成功表达，又能最大可能地保留人类离子通道的药物学调控特性。
接着，在这种嵌合的离子通道上引入一个功能增强型点突变(命名为嵌合通道1，图4A)，当把这种离子通道表达在线虫中会明显改变线虫细胞如神经元和肌肉细胞等的兴奋性，进而导致转基因线虫产生表型缺陷。当用通道的抑制剂处理降低通道功能时会减缓线虫的表型缺陷，因此本发明人可以利用这种转基因线虫筛选离子通道的抑制剂(图4B)。
为了能够筛选离子通道的激活剂或功能矫正剂，本发明人在嵌合通道1上再引入一个功能缺失型点突变(命名为嵌合通道2，图4A)，它会严重降低通道电流或完全丧失，因而表达嵌合通道2的转基因线虫并不会出现表型缺陷。当用化合物库处理这种转基因线虫时，离子通道的激活剂或功能矫正剂能够修复突变通道的功能缺陷使嵌合通道2重新发挥相应的通道功能，进而诱导转基因线虫产生表型缺陷；因此可以用线虫模型筛选人类离子通道的激活剂或功能矫正剂(图4B)。
采用上述策略，本发明人成功构建了3种通道(人KCNQ2钾通道、人Kv1.2钾通道、人BK钾通道)疾病的线虫模型。
(1)人的KCNQ2通道
人的KCNQ2通道在线虫的同源物是KQT-1钾通道，它调节线虫咽喉肌肉的节律跳动和进食行为(Wei，A.D.，A.Butler，et al.(2005)."KCNQ-like potassium channels inCaenorhabditis elegans.Conserved properties and modulation."J Biol Chem 280(22):21337-21345)。通过把线虫KQT-1钾通道的N端、人KCNQ2通道的S1-S6跨膜区、和KQT-1通道的C端组合在一起构建嵌合的KCNQ2chimera通道(图5A)，并引入一个与早发性癫痫脑病相关的R201C功能增强型点突变(Miceli，F.，M.V.Soldovieri，et al.(2015)."Early-onset epileptic encephalopathy caused by gain-of-function mutations in thevoltage sensor of Kv7.2and Kv7.3potassium channel subunits."J Neurosci 35(9):3782-3793)；进而得到由kqt-1基因启动子驱动表达这种嵌合离子通道的KCNQ2chimera/R201C转基因线虫。与野生型N2线虫相比，KCNQ2chimera/R201C线虫的咽喉肌肉跳动速率明显减慢，影响线虫的进食行为进而导致转基因线虫发育迟缓(图5B，E)。
(2)大电导钙激活的钾通道(BK)
大电导钙激活的钾通道(BK)调节神经递质的释放和神经元的兴奋性，它的功能异常会导致许多精神类疾病的发生，如研究报道BK通道D434G功能增强型点突变被报道与全身性癫痫与发作性运动障碍有关(Du，W.，J.F.Bautista，et al.(2005)."Calcium-sensitive potassium channelopathy in human epilepsy and paroxysmal movementdisorder."Nat Genet 37(7):733-738)。利用slo-1基因的启动子驱动嵌合的BKchimera/D434G通道在线虫中表达，构建BKchimera/D434G转基因线虫，它们出现运动和产卵行为障碍(图5B，C，D)。
(3)人Kv1.2钾通道
同样，在嵌合的KV1.2chimera通道中引入一个与癫痫疾病相关的R297Q功能增强型点突变(Syrbe，S.et al.De novo loss-or gain-of-function mutations in KCNA2causeepileptic encephalopathy.Nat Genet 47，393-399)，进而用shk-1基因启动子驱动表达在N2线虫中构建KV1.2chimera/R297Q转基因线虫(图5B，C，D)。由于线虫的shk-1基因表达在与运动和产卵行为相关的神经元和肌肉细胞中，KV1.2chimera/R297Q转基因线虫也出现运动和产卵的行为缺陷。
总之，本发明人成功构建了多种表达人类致病离子通道的线虫模型，它们具有明显的表型缺陷，这为在通道疾病线虫模型中筛选通道化合物提供了便利的检测指标。
实施例2、通道抑制剂减缓通道疾病线虫模型的表型缺陷
在构建好通道疾病线虫模型后，本发明人利用已知的通道阻断剂处理，来验证这些模型能否减缓线虫的表型缺陷。
线虫的体壁较厚，同时线虫肠道又可以主动排除外源有毒物质，这使得大多数化合物较难进入线虫体内。acs-20基因编码线虫体壁的一个脂肪转运蛋白(Kage-Nakadai，E.et al.(2010).Two very long chain fatty acid acyl-CoA synthetase genes，acs-20and acs-22，have roles in the cuticle surface barrier in Caenorhabditiselegans.PLoS One 5，e8857)，其基因敲除线虫acs-20(tm3232)体壁保护功能的缺失，允许更多小分子化合物进入线虫体内。因此，本发明人把构建好的转基因线虫与acs-20(tm3232)杂交得到acs-20；BKChimera/D434G、acs-20；Kv1.2Chimera/R297Q和acs-20；KCNQ2Chimera /R201C化合物筛选用线虫。
用已知的BK通道阻断剂PenitremA和Paxilline处理acs-20；BKChimera/D434G线虫，其运动和产卵行为缺陷可以得到明显改善(图6)。
同样，电压门控钾通道的抑制剂Tetraethylammonium(TEA)和AM92016处理，可以减缓acs-20；Kv1.2Chimera/R297Q线虫的产卵缺陷。
KCNQ2钾通道的阻断剂ML252处理，可以改善acs-20；KCNQ2Chimera/R201C线虫的发育迟缓缺陷。
这些结果说明，小分子化合物可以有效地进入线虫体内发挥作用，同时也表明本发明所构建的转基因线虫表型缺陷确实是融合的KCNQ2Chimera/R201C、Kv1.2Chimera/R297Q和BKChimera/D434G通道蛋白成功表达并发挥功能导致的。因此，可以使用这些构建的通道疾病线虫模型进行表型筛选寻找调控KCNQ2、KV1.2和BK钾通道功能的小分子化合物。
在本发明提及的所有文献都在本申请中引用作为参考，就如同每一篇文献被单独引用作为参考那样。此外应理解，在阅读了本发明的上述讲授内容之后，本领域技术人员可以对本发明作各种改动或修改，这些等价形式同样落于本申请所附权利要求书所限定的范围。
参考文献：
Biervert,C.,Schroeder,B.C.,Kubisch,C.,Berkovic,S.F.,Propping,P.,Jentsch,T.J.,and Steinlein,O.K.(1998).A potassium channel mutation inneonatal human epilepsy.Science 279,403-406.
Chen,X.,Ruan,M.Y.,and Cai,S.Q.(2015).KChIP-like auxiliary subunits ofKv4channels regulate excitability of muscle cells and control male turningbehavior during mating in Caenorhabditis elegans.J Neurosci 35,1880-1891.
Jentsch,T.J.(2000).Neuronal KCNQ potassium channels:physiology androle in disease.Nat Rev Neurosci 1,21-30.
Li,K.,Jiang,Q.,Bai,X.,Yang,Y.F.,Ruan,M.Y.,and Cai,S.Q.(2017).Tetrameric Assembly of K(+)Channels Requires ER-Located ChaperoneProteins.Mol Cell 65,52-65.
Petersen,C.I.,McFarland,T.R.,Stepanovic,S.Z.,Yang,P.,Reiner,D.J.,Hayashi,K.,George,A.L.,Roden,D.M.,Thomas,J.H.,and Jr,B.(2004).In vivoidentification of genes that modify ether-a-go-go-related gene activity inCaenorhabditis elegans may also affect human cardiac arrhythmia.P Natl AcadSci USA 101,11773-11778.
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gtctatgccc acagcaagga gctggtcact gcctggtaca tcggcttcct ttgtctcatc 900
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aggcagaagc actttgagaa gaggcggaac ccggcagcag gcctgatcca gtcggcctgg 1200
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caatcaagga tgttggcacc atcaaatgct catctagacg aagaagaagc cgtcggatat 1620
caaccacaaa caattgaaga gtttacccct gctttgaaaa attgtgttcg agcgatcaga 1680
agaatacaac tcttggtggc tcggaagaaa ttcaaggagg cattaaaacc ctatgacgtc 1740
aaagatgtga ttgaacaata ttcagctggt catgttgatt tgcaaagtcg agtgaagaca 1800
gttcaagcga aactagattt tatatgcggg aaaaatattg agaaaattga gccaaaaatt 1860
tcaatgttca ctagaattgc aactttggaa acaactgttg gtaaaatgga caaaaagctc 1920
gacttaatgg ttgaaatgct aatgggaagg caagcatccc agcgtgtctt ctcacagaac 1980
acatcgccac gtggagagtt ttctgaacca actagtgcga gacaagattt gacaaggtcg 2040
cggcggtcta tggtcagcac tgacatggaa atgtatacag caagatcaca ttcgcctggt 2100
tatcatggag atgcccgacc aattattgct caaatagatg cagatgatga cgatgaagat 2160
gagaatgttt tcgacgattc tactccattg aacaatggtc ccggtacttc aagttgt 2217
<210> 6
<211> 2217
<212> DNA
<213> 人工序列(Artificial Sequence)
<220>
<221> misc_feature
<222> (1)..(2217)
<223> KCNQ2chimera/R201C cDNA序列
<400> 6
atgtctcaag cttcacaaga agacaatgac ggtggagatt tattgtcccc agagtcacca 60
atgactccag acggaatgat attggcacga atgccttggc acccaggtct cattggaaat 120
tggaacacag tgaaatcagt gagctgcagt atgctcaatg gagacgagac tgatcaagca 180
gctccaagtg atgagcagca ggaagctggc tcgtcttcag caatcggtca agaatctcga 240
aaaacagttg tatttcaaga acccgatatt ggattcccga gtgaacacga tcaactgaca 300
actctacacg attctgagga aggaaaccga aaaatgtcat tagttggaaa accgttaacc 360
tataaaaact atcgtactga tcaacgattt cgcaagctgc agaatttcct ctacaacgtg 420
ctggagcggc cgcgcggctg ggcgttcatc taccacgcct acgtgttcct cctggttttc 480
tcctgcctcg tgctgtctgt gttttccacc atcaaggagt atgagaagag ctcggagggg 540
gccctctaca tcctggaaat cgtgactatc gtggtgtttg gcgtggagta cttcgtgcgg 600
atctgggccg caggctgctg ctgccggtac cgtggctgga gggggcggct caagtttgcc 660
cggaaaccgt tctgtgtgat tgacatcatg gtgctcatcg cctccattgc ggtgctggcc 720
gccggctccc agggcaacgt ctttgccaca tctgcgctcc ggagcctgtg cttcctgcag 780
attctgcgga tgatccgcat ggaccggcgg ggaggcacct ggaagctgct gggctctgtg 840
gtctatgccc acagcaagga gctggtcact gcctggtaca tcggcttcct ttgtctcatc 900
ctggcctcgt tcctggtgta cttggcagag aagggggaga acgaccactt tgacacctac 960
gcggatgcac tctggtgggg cctgatcacg ctgaccacca ttggctacgg ggacaagtac 1020
ccccagacct ggaacggcag gctccttgcg gcaaccttca ccctcatcgg tgtctccttc 1080
ttcgcgctgc ctgcaggcat cttggggtct gggtttgccc tgaaggttca ggagcagcac 1140
aggcagaagc actttgagaa gaggcggaac ccggcagcag gcctgatcca gtcggcctgg 1200
agattctact ctgctgcacc ggaatccacc tcacttgcaa cttggaaaat acacttagcc 1260
agagagcttc caccaatcgt caaacttact ccaaacggaa gtaacaatgc tacaggtctt 1320
ataaatcgat tacgtcaaag tacaaaacga acaccaaatt tgaataacca aaatctagcc 1380
gtcaattcac aagcaacttc aaaaaacctt tccgtcccac gagtgcacga caccataagt 1440
ctggtctcaa cgtcagatat atctgaaatt gagcagctag gagctcttgg attctctttg 1500
ggttggaagt caaagtccaa atatggagga tcaaagaaag ctacagatga ttcagttttg 1560
caatcaagga tgttggcacc atcaaatgct catctagacg aagaagaagc cgtcggatat 1620
caaccacaaa caattgaaga gtttacccct gctttgaaaa attgtgttcg agcgatcaga 1680
agaatacaac tcttggtggc tcggaagaaa ttcaaggagg cattaaaacc ctatgacgtc 1740
aaagatgtga ttgaacaata ttcagctggt catgttgatt tgcaaagtcg agtgaagaca 1800
gttcaagcga aactagattt tatatgcggg aaaaatattg agaaaattga gccaaaaatt 1860
tcaatgttca ctagaattgc aactttggaa acaactgttg gtaaaatgga caaaaagctc 1920
gacttaatgg ttgaaatgct aatgggaagg caagcatccc agcgtgtctt ctcacagaac 1980
acatcgccac gtggagagtt ttctgaacca actagtgcga gacaagattt gacaaggtcg 2040
cggcggtcta tggtcagcac tgacatggaa atgtatacag caagatcaca ttcgcctggt 2100
tatcatggag atgcccgacc aattattgct caaatagatg cagatgatga cgatgaagat 2160
gagaatgttt tcgacgattc tactccattg aacaatggtc ccggtacttc aagttgt 2217
<210> 7
<211> 499
<212> PRT
<213> 智人(Homo sapiens)
<400> 7
Met Thr Val Ala Thr Gly Asp Pro Ala Asp Glu Ala Ala Ala Leu Pro
1 5 10 15
Gly His Pro Gln Asp Thr Tyr Asp Pro Glu Ala Asp His Glu Cys Cys
20 25 30
Glu Arg Val Val Ile Asn Ile Ser Gly Leu Arg Phe Glu Thr Gln Leu
35 40 45
Lys Thr Leu Ala Gln Phe Pro Glu Thr Leu Leu Gly Asp Pro Lys Lys
50 55 60
Arg Met Arg Tyr Phe Asp Pro Leu Arg Asn Glu Tyr Phe Phe Asp Arg
65 70 75 80
Asn Arg Pro Ser Phe Asp Ala Ile Leu Tyr Tyr Tyr Gln Ser Gly Gly
85 90 95
Arg Leu Arg Arg Pro Val Asn Val Pro Leu Asp Ile Phe Ser Glu Glu
100 105 110
Ile Arg Phe Tyr Glu Leu Gly Glu Glu Ala Met Glu Met Phe Arg Glu
115 120 125
Asp Glu Gly Tyr Ile Lys Glu Glu Glu Arg Pro Leu Pro Glu Asn Glu
130 135 140
Phe Gln Arg Gln Val Trp Leu Leu Phe Glu Tyr Pro Glu Ser Ser Gly
145 150 155 160
Pro Ala Arg Ile Ile Ala Ile Val Ser Val Met Val Ile Leu Ile Ser
165 170 175
Ile Val Ser Phe Cys Leu Glu Thr Leu Pro Ile Phe Arg Asp Glu Asn
180 185 190
Glu Asp Met His Gly Ser Gly Val Thr Phe His Thr Tyr Ser Asn Ser
195 200 205
Thr Ile Gly Tyr Gln Gln Ser Thr Ser Phe Thr Asp Pro Phe Phe Ile
210 215 220
Val Glu Thr Leu Cys Ile Ile Trp Phe Ser Phe Glu Phe Leu Val Arg
225 230 235 240
Phe Phe Ala Cys Pro Ser Lys Ala Gly Phe Phe Thr Asn Ile Met Asn
245 250 255
Ile Ile Asp Ile Val Ala Ile Ile Pro Tyr Phe Ile Thr Leu Gly Thr
260 265 270
Glu Leu Ala Glu Lys Pro Glu Asp Ala Gln Gln Gly Gln Gln Ala Met
275 280 285
Ser Leu Ala Ile Leu Arg Val Ile Arg Leu Val Arg Val Phe Arg Ile
290 295 300
Phe Lys Leu Ser Arg His Ser Lys Gly Leu Gln Ile Leu Gly Gln Thr
305 310 315 320
Leu Lys Ala Ser Met Arg Glu Leu Gly Leu Leu Ile Phe Phe Leu Phe
325 330 335
Ile Gly Val Ile Leu Phe Ser Ser Ala Val Tyr Phe Ala Glu Ala Asp
340 345 350
Glu Arg Glu Ser Gln Phe Pro Ser Ile Pro Asp Ala Phe Trp Trp Ala
355 360 365
Val Val Ser Met Thr Thr Val Gly Tyr Gly Asp Met Val Pro Thr Thr
370 375 380
Ile Gly Gly Lys Ile Val Gly Ser Leu Cys Ala Ile Ala Gly Val Leu
385 390 395 400
Thr Ile Ala Leu Pro Val Pro Val Ile Val Ser Asn Phe Asn Tyr Phe
405 410 415
Tyr His Arg Glu Thr Glu Gly Glu Glu Gln Ala Gln Tyr Leu Gln Val
420 425 430
Thr Ser Cys Pro Lys Ile Pro Ser Ser Pro Asp Leu Lys Lys Ser Arg
435 440 445
Ser Ala Ser Thr Ile Ser Lys Ser Asp Tyr Met Glu Ile Gln Glu Gly
450 455 460
Val Asn Asn Ser Asn Glu Asp Phe Arg Glu Glu Asn Leu Lys Thr Ala
465 470 475 480
Asn Cys Thr Leu Ala Asn Thr Asn Tyr Val Asn Ile Thr Lys Met Leu
485 490 495
Thr Asp Val
<210> 8
<211> 524
<212> PRT
<213> 线虫(Caenorhabditis)
<400> 8
Met Arg Phe Gly Gly Gln Arg Arg Cys Ile Glu Ser Ser Asp Asp Asn
1 5 10 15
Asp Ser Ile Asp Ser Met Ser Ile Gln Gln Lys Asn Lys Glu Lys Glu
20 25 30
Glu Trp Lys Trp Ala Gln Ser Leu Ala Arg Arg Leu Glu Gln Val Leu
35 40 45
Asn Val His Pro Glu Lys Tyr Gln Asn Gly Cys Ser Ser Gln Arg Pro
50 55 60
Ser Arg Gln Pro Ser Pro Val Glu Gln Leu Ala Pro Ile Arg Ser His
65 70 75 80
Ser Ala Asp Phe Gly Arg Ala Ala Glu Met Ala Ser Gly Asn Ser Glu
85 90 95
Glu Gly Lys Gly Met Leu Leu Arg Asn Gly Asp Asp Arg Ile Arg Leu
100 105 110
Pro Ser Pro Gln Arg Gly Thr Pro Asp Thr Ser Ser Thr Gln Gly His
115 120 125
Pro Tyr Thr Asp Gln Ile Val Thr Ile Asn Val Ser Gly Met Arg Phe
130 135 140
Gln Thr Phe Glu Ser Thr Leu Ser Arg Tyr Pro Asn Ser Leu Leu Gly
145 150 155 160
Asp Arg Asn Lys Arg Gln His Phe Phe Val Ser Asp Thr Asn Glu Phe
165 170 175
Phe Phe Asp Arg His Arg Thr Thr Phe Glu Ser Ile Leu Tyr Ile Tyr
180 185 190
Gln Ser Gly Gly Arg Val Lys Arg Pro Glu Ile Val Pro Ile Asp Ile
195 200 205
Phe Leu Lys Glu Met Arg Phe Phe Gln Met Gly Asp Asp Leu Leu Glu
210 215 220
Glu Phe Trp Ile Ala Glu Gly Tyr Glu Lys Pro Lys Glu Val Met Met
225 230 235 240
Pro Asn Asn Lys Thr Gln Arg Lys Ile Trp Glu Leu Met Glu Tyr Pro
245 250 255
Asp Ser Ser Leu Ser Ala Arg Ile Ile Ala Phe Ile Ser Ile Ala Val
260 265 270
Ile Ala Leu Ser Ile Ile Ser Phe Cys Trp Glu Thr Val Pro Ser Asp
275 280 285
Ile Glu Glu Lys Pro Ile Asn Asn Ser Ala Thr Ala Glu Leu Leu Asp
290 295 300
Glu Met Asp Glu Lys His Tyr Ser Pro Phe Phe Trp Ile Glu Leu Met
305 310 315 320
Cys Ile Leu Trp Phe Thr Ile Glu Leu Ile Leu Arg Phe Ile Ser Cys
325 330 335
Pro Cys Lys Val Thr Phe Ala Thr Ser Val Leu Asn Ile Ile Asp Phe
340 345 350
Val Ala Ile Ala Pro Phe Phe Val Asn Phe Phe Phe Ala Asp Thr Ser
355 360 365
Lys Ser Asn Ser Ser Met Ser Phe Ala Val Leu Arg Val Leu Arg Leu
370 375 380
Val Arg Val Phe Arg Val Phe Lys Leu Ser Arg His Ser Val Gly Leu
385 390 395 400
Gln Ile Leu Gly Lys Thr Phe Arg Ser Ser Val Gln Glu Phe Cys Leu
405 410 415
Leu Ile Phe Phe Met Ala Ile Ala Leu Val Leu Phe Ala Ser Gly Met
420 425 430
Tyr Phe Ala Glu Gln Gly Glu Pro Asn Ser Lys Phe Thr Ser Ile Pro
435 440 445
Ala Ser Phe Trp Phe Val Leu Val Thr Met Thr Thr Val Gly Tyr Gly
450 455 460
Asp Leu Val Pro Leu Ser Pro Phe Gly Lys Val Val Gly Gly Met Cys
465 470 475 480
Ala Met Ile Gly Val Leu Thr Leu Ala Leu Pro Val Pro Ile Ile Val
485 490 495
Ala Asn Phe Lys His Phe Tyr Arg Gln Glu Asn Arg Leu Ala Ser Met
500 505 510
Lys Ser Lys Gly Asp Asp Ala Asp Asp Asp Ile Ala
515 520
<210> 9
<211> 599
<212> PRT
<213> 人工序列(Artificial Sequence)
<220>
<221> PEPTIDE
<222> (1)..(599)
<223> KV1.2chimera蛋白氨基酸序列
<400> 9
Met Arg Phe Gly Gly Gln Arg Arg Cys Ile Glu Ser Ser Asp Asp Asn
1 5 10 15
Asp Ser Ile Asp Ser Met Ser Ile Gln Gln Lys Asn Lys Glu Lys Glu
20 25 30
Glu Trp Lys Trp Ala Gln Ser Leu Ala Arg Arg Leu Glu Gln Val Leu
35 40 45
Asn Val His Pro Glu Lys Tyr Gln Asn Gly Cys Ser Ser Gln Arg Pro
50 55 60
Ser Arg Gln Pro Ser Pro Val Glu Gln Leu Ala Pro Ile Arg Ser His
65 70 75 80
Ser Ala Asp Phe Gly Arg Ala Ala Glu Met Ala Ser Gly Asn Ser Glu
85 90 95
Glu Gly Lys Gly Met Leu Leu Arg Asn Gly Asp Asp Arg Ile Arg Leu
100 105 110
Pro Ser Pro Gln Arg Gly Thr Pro Asp Thr Ser Ser Thr Gln Gly His
115 120 125
Pro Tyr Thr Asp Gln Ile Val Thr Ile Asn Val Ser Gly Met Arg Phe
130 135 140
Gln Thr Phe Glu Ser Thr Leu Ser Arg Tyr Pro Asn Ser Leu Leu Gly
145 150 155 160
Asp Arg Asn Lys Arg Gln His Phe Phe Val Ser Asp Thr Asn Glu Phe
165 170 175
Phe Phe Asp Arg His Arg Thr Thr Phe Glu Ser Ile Leu Tyr Ile Tyr
180 185 190
Gln Ser Gly Gly Arg Val Lys Arg Pro Glu Ile Val Pro Ile Asp Ile
195 200 205
Phe Leu Lys Glu Met Arg Phe Phe Gln Met Gly Asp Asp Leu Leu Glu
210 215 220
Glu Phe Trp Ile Ala Glu Gly Tyr Glu Lys Pro Lys Glu Val Met Met
225 230 235 240
Pro Asn Asn Lys Thr Gln Arg Gln Val Trp Leu Leu Phe Glu Tyr Pro
245 250 255
Glu Ser Ser Gly Pro Ala Arg Ile Ile Ala Ile Val Ser Val Met Val
260 265 270
Ile Leu Ile Ser Ile Val Ser Phe Cys Leu Glu Thr Leu Pro Ile Phe
275 280 285
Arg Asp Glu Asn Glu Asp Met His Gly Ser Gly Val Thr Phe His Thr
290 295 300
Tyr Ser Asn Ser Thr Ile Gly Tyr Gln Gln Ser Thr Ser Phe Thr Asp
305 310 315 320
Pro Phe Phe Ile Val Glu Thr Leu Cys Ile Ile Trp Phe Ser Phe Glu
325 330 335
Phe Leu Val Arg Phe Phe Ala Cys Pro Ser Lys Ala Gly Phe Phe Thr
340 345 350
Asn Ile Met Asn Ile Ile Asp Ile Val Ala Ile Ile Pro Tyr Phe Ile
355 360 365
Thr Leu Gly Thr Glu Leu Ala Glu Lys Pro Glu Asp Ala Gln Gln Gly
370 375 380
Gln Gln Ala Met Ser Leu Ala Ile Leu Arg Val Ile Arg Leu Val Arg
385 390 395 400
Val Phe Arg Ile Phe Lys Leu Ser Arg His Ser Lys Gly Leu Gln Ile
405 410 415
Leu Gly Gln Thr Leu Lys Ala Ser Met Arg Glu Leu Gly Leu Leu Ile
420 425 430
Phe Phe Leu Phe Ile Gly Val Ile Leu Phe Ser Ser Ala Val Tyr Phe
435 440 445
Ala Glu Ala Asp Glu Arg Glu Ser Gln Phe Pro Ser Ile Pro Asp Ala
450 455 460
Phe Trp Trp Ala Val Val Ser Met Thr Thr Val Gly Tyr Gly Asp Met
465 470 475 480
Val Pro Thr Thr Ile Gly Gly Lys Ile Val Gly Ser Leu Cys Ala Ile
485 490 495
Ala Gly Val Leu Thr Ile Ala Leu Pro Val Pro Val Ile Val Ser Asn
500 505 510
Phe Asn Tyr Phe Tyr His Arg Glu Thr Glu Gly Glu Glu Gln Ala Gln
515 520 525
Tyr Leu Gln Val Thr Ser Cys Pro Lys Ile Pro Ser Ser Pro Asp Leu
530 535 540
Lys Lys Ser Arg Ser Ala Ser Thr Ile Ser Lys Ser Asp Tyr Met Glu
545 550 555 560
Ile Gln Glu Gly Val Asn Asn Ser Asn Glu Asp Phe Arg Glu Glu Asn
565 570 575
Leu Lys Thr Ala Asn Cys Thr Leu Ala Asn Thr Asn Tyr Val Asn Ile
580 585 590
Thr Lys Met Leu Thr Asp Val
595
<210> 10
<211> 599
<212> PRT
<213> 人工序列(Artificial Sequence)
<220>
<221> PEPTIDE
<222> (1)..(599)
<223> KV1.2chimera/R297Q蛋白氨基酸序列
<400> 10
Met Arg Phe Gly Gly Gln Arg Arg Cys Ile Glu Ser Ser Asp Asp Asn
1 5 10 15
Asp Ser Ile Asp Ser Met Ser Ile Gln Gln Lys Asn Lys Glu Lys Glu
20 25 30
Glu Trp Lys Trp Ala Gln Ser Leu Ala Arg Arg Leu Glu Gln Val Leu
35 40 45
Asn Val His Pro Glu Lys Tyr Gln Asn Gly Cys Ser Ser Gln Arg Pro
50 55 60
Ser Arg Gln Pro Ser Pro Val Glu Gln Leu Ala Pro Ile Arg Ser His
65 70 75 80
Ser Ala Asp Phe Gly Arg Ala Ala Glu Met Ala Ser Gly Asn Ser Glu
85 90 95
Glu Gly Lys Gly Met Leu Leu Arg Asn Gly Asp Asp Arg Ile Arg Leu
100 105 110
Pro Ser Pro Gln Arg Gly Thr Pro Asp Thr Ser Ser Thr Gln Gly His
115 120 125
Pro Tyr Thr Asp Gln Ile Val Thr Ile Asn Val Ser Gly Met Arg Phe
130 135 140
Gln Thr Phe Glu Ser Thr Leu Ser Arg Tyr Pro Asn Ser Leu Leu Gly
145 150 155 160
Asp Arg Asn Lys Arg Gln His Phe Phe Val Ser Asp Thr Asn Glu Phe
165 170 175
Phe Phe Asp Arg His Arg Thr Thr Phe Glu Ser Ile Leu Tyr Ile Tyr
180 185 190
Gln Ser Gly Gly Arg Val Lys Arg Pro Glu Ile Val Pro Ile Asp Ile
195 200 205
Phe Leu Lys Glu Met Arg Phe Phe Gln Met Gly Asp Asp Leu Leu Glu
210 215 220
Glu Phe Trp Ile Ala Glu Gly Tyr Glu Lys Pro Lys Glu Val Met Met
225 230 235 240
Pro Asn Asn Lys Thr Gln Arg Gln Val Trp Leu Leu Phe Glu Tyr Pro
245 250 255
Glu Ser Ser Gly Pro Ala Arg Ile Ile Ala Ile Val Ser Val Met Val
260 265 270
Ile Leu Ile Ser Ile Val Ser Phe Cys Leu Glu Thr Leu Pro Ile Phe
275 280 285
Arg Asp Glu Asn Glu Asp Met His Gly Ser Gly Val Thr Phe His Thr
290 295 300
Tyr Ser Asn Ser Thr Ile Gly Tyr Gln Gln Ser Thr Ser Phe Thr Asp
305 310 315 320
Pro Phe Phe Ile Val Glu Thr Leu Cys Ile Ile Trp Phe Ser Phe Glu
325 330 335
Phe Leu Val Arg Phe Phe Ala Cys Pro Ser Lys Ala Gly Phe Phe Thr
340 345 350
Asn Ile Met Asn Ile Ile Asp Ile Val Ala Ile Ile Pro Tyr Phe Ile
355 360 365
Thr Leu Gly Thr Glu Leu Ala Glu Lys Pro Glu Asp Ala Gln Gln Gly
370 375 380
Gln Gln Ala Met Ser Leu Ala Ile Leu Arg Val Ile Gln Leu Val Arg
385 390 395 400
Val Phe Arg Ile Phe Lys Leu Ser Arg His Ser Lys Gly Leu Gln Ile
405 410 415
Leu Gly Gln Thr Leu Lys Ala Ser Met Arg Glu Leu Gly Leu Leu Ile
420 425 430
Phe Phe Leu Phe Ile Gly Val Ile Leu Phe Ser Ser Ala Val Tyr Phe
435 440 445
Ala Glu Ala Asp Glu Arg Glu Ser Gln Phe Pro Ser Ile Pro Asp Ala
450 455 460
Phe Trp Trp Ala Val Val Ser Met Thr Thr Val Gly Tyr Gly Asp Met
465 470 475 480
Val Pro Thr Thr Ile Gly Gly Lys Ile Val Gly Ser Leu Cys Ala Ile
485 490 495
Ala Gly Val Leu Thr Ile Ala Leu Pro Val Pro Val Ile Val Ser Asn
500 505 510
Phe Asn Tyr Phe Tyr His Arg Glu Thr Glu Gly Glu Glu Gln Ala Gln
515 520 525
Tyr Leu Gln Val Thr Ser Cys Pro Lys Ile Pro Ser Ser Pro Asp Leu
530 535 540
Lys Lys Ser Arg Ser Ala Ser Thr Ile Ser Lys Ser Asp Tyr Met Glu
545 550 555 560
Ile Gln Glu Gly Val Asn Asn Ser Asn Glu Asp Phe Arg Glu Glu Asn
565 570 575
Leu Lys Thr Ala Asn Cys Thr Leu Ala Asn Thr Asn Tyr Val Asn Ile
580 585 590
Thr Lys Met Leu Thr Asp Val
595
<210> 11
<211> 1797
<212> DNA
<213> 人工序列(Artificial Sequence)
<220>
<221> misc_feature
<222> (1)..(1797)
<223> KV1.2chimera cDNA序列
<400> 11
atgcgattcg gtggtcaacg acgctgcatc gaaagcagcg atgacaacga ttcaattgat 60
tcgatgtcaa ttcaacaaaa aaataaggaa aaagaagagt ggaaatgggc acagtcacta 120
gcaagacgtc tggaacaagt actaaacgtc catcctgaaa aatatcaaaa tggatgcagc 180
agtcaacgtc cgagtcgtca gccatcaccg gttgagcaac ttgccccgat tcggagtcat 240
agtgccgatt ttggcagagc cgctgaaatg gcatctggca acagtgaaga aggaaaagga 300
atgctattga ggaacgggga cgaccggatt cggctcccaa gtccgcaacg tggaactcct 360
gatacttcga gcacgcaggg acatccttac acggatcaaa ttgtcacaat aaatgtgtcc 420
gggatgcggt ttcaaacttt tgagagcaca ttgtcaaggt acccaaactc gttgctaggt 480
gatcggaata aacggcaaca ctttttcgtg tcggatacta acgaattttt ctttgatcgg 540
cacaggacca cttttgaatc aattctctac atatatcaaa gtggcggaag agtgaaacga 600
cctgaaattg tgccgattga catatttttg aaagaaatgc gatttttcca aatgggtgat 660
gatcttctgg aagaattctg gatagctgag ggttatgaaa aaccaaaaga agtgatgatg 720
ccaaataata aaacccagag acaagtgtgg cttctctttg aatacccaga gagctcaggg 780
cctgccagga ttatagctat tgtgtctgtc atggtgattc tgatctcaat tgtcagcttc 840
tgtctggaaa cattgcccat cttccgggat gagaatgaag acatgcatgg tagtggggtg 900
accttccaca cctattccaa cagcaccatc gggtaccagc agtccacttc cttcacagac 960
cctttcttca ttgtagagac actctgcatc atctggttct cctttgaatt cttggtgagg 1020
ttctttgcct gtcccagcaa agccggcttc ttcaccaaca tcatgaacat cattgacatt 1080
gtggccatca tcccctactt catcaccctg gggacagagt tggctgagaa gccagaggac 1140
gctcagcaag gccagcaggc catgtcactg gccatcctcc gtgtcatccg gttggtaaga 1200
gtctttagga ttttcaagtt gtccagacac tccaaaggtc tccagattct aggtcagacc 1260
ctcaaagcca gcatgagaga attgggcctc ctgatattct ttctcttcat aggggtcatc 1320
cttttctcta gtgctgtgta ttttgcagag gccgatgagc gagagtccca gttccccagc 1380
atcccagatg ccttctggtg ggcagtcgtc tccatgacaa ctgtaggcta tggagacatg 1440
gttccgacta ccattggggg aaagatagtg ggttccctat gtgcgattgc aggtgtgtta 1500
actattgcct taccggtccc tgtcattgtg tccaatttca actacttcta ccaccgggag 1560
acagagggag aggaacaggc ccaatacttg caagtgacaa gctgtccaaa gatcccatcc 1620
tcccctgacc taaagaaaag tagaagtgcc tctaccatta gtaagtctga ttacatggag 1680
atccaggagg gtgtaaataa cagtaatgag gactttagag aggaaaactt gaaaacagcc 1740
aactgtacct tggctaacac aaactatgtg aatattacca aaatgttaac tgatgtc 1797
<210> 12
<211> 1797
<212> DNA
<213> 人工序列(Artificial Sequence)
<220>
<221> misc_feature
<222> (1)..(1797)
<223> KV1.2chimera/R297Q cDNA序列
<400> 12
atgcgattcg gtggtcaacg acgctgcatc gaaagcagcg atgacaacga ttcaattgat 60
tcgatgtcaa ttcaacaaaa aaataaggaa aaagaagagt ggaaatgggc acagtcacta 120
gcaagacgtc tggaacaagt actaaacgtc catcctgaaa aatatcaaaa tggatgcagc 180
agtcaacgtc cgagtcgtca gccatcaccg gttgagcaac ttgccccgat tcggagtcat 240
agtgccgatt ttggcagagc cgctgaaatg gcatctggca acagtgaaga aggaaaagga 300
atgctattga ggaacgggga cgaccggatt cggctcccaa gtccgcaacg tggaactcct 360
gatacttcga gcacgcaggg acatccttac acggatcaaa ttgtcacaat aaatgtgtcc 420
gggatgcggt ttcaaacttt tgagagcaca ttgtcaaggt acccaaactc gttgctaggt 480
gatcggaata aacggcaaca ctttttcgtg tcggatacta acgaattttt ctttgatcgg 540
cacaggacca cttttgaatc aattctctac atatatcaaa gtggcggaag agtgaaacga 600
cctgaaattg tgccgattga catatttttg aaagaaatgc gatttttcca aatgggtgat 660
gatcttctgg aagaattctg gatagctgag ggttatgaaa aaccaaaaga agtgatgatg 720
ccaaataata aaacccagag acaagtgtgg cttctctttg aatacccaga gagctcaggg 780
cctgccagga ttatagctat tgtgtctgtc atggtgattc tgatctcaat tgtcagcttc 840
tgtctggaaa cattgcccat cttccgggat gagaatgaag acatgcatgg tagtggggtg 900
accttccaca cctattccaa cagcaccatc gggtaccagc agtccacttc cttcacagac 960
cctttcttca ttgtagagac actctgcatc atctggttct cctttgaatt cttggtgagg 1020
ttctttgcct gtcccagcaa agccggcttc ttcaccaaca tcatgaacat cattgacatt 1080
gtggccatca tcccctactt catcaccctg gggacagagt tggctgagaa gccagaggac 1140
gctcagcaag gccagcaggc catgtcactg gccatcctcc gtgtcatcca gttggtaaga 1200
gtctttagga ttttcaagtt gtccagacac tccaaaggtc tccagattct aggtcagacc 1260
ctcaaagcca gcatgagaga attgggcctc ctgatattct ttctcttcat aggggtcatc 1320
cttttctcta gtgctgtgta ttttgcagag gccgatgagc gagagtccca gttccccagc 1380
atcccagatg ccttctggtg ggcagtcgtc tccatgacaa ctgtaggcta tggagacatg 1440
gttccgacta ccattggggg aaagatagtg ggttccctat gtgcgattgc aggtgtgtta 1500
actattgcct taccggtccc tgtcattgtg tccaatttca actacttcta ccaccgggag 1560
acagagggag aggaacaggc ccaatacttg caagtgacaa gctgtccaaa gatcccatcc 1620
tcccctgacc taaagaaaag tagaagtgcc tctaccatta gtaagtctga ttacatggag 1680
atccaggagg gtgtaaataa cagtaatgag gactttagag aggaaaactt gaaaacagcc 1740
aactgtacct tggctaacac aaactatgtg aatattacca aaatgttaac tgatgtc 1797
<210> 13
<211> 1178
<212> PRT
<213> 智人(Homo sapiens)
<400> 13
Met Ala Asn Gly Gly Gly Gly Gly Gly Gly Ser Ser Gly Gly Gly Gly
1 5 10 15
Gly Gly Gly Gly Ser Ser Leu Arg Met Ser Ser Asn Ile His Ala Asn
20 25 30
His Leu Ser Leu Asp Ala Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser
35 40 45
Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser Val His Glu Pro
50 55 60
Lys Met Asp Ala Leu Ile Ile Pro Val Thr Met Glu Val Pro Cys Asp
65 70 75 80
Ser Arg Gly Gln Arg Met Trp Trp Ala Phe Leu Ala Ser Ser Met Val
85 90 95
Thr Phe Phe Gly Gly Leu Phe Ile Ile Leu Leu Trp Arg Thr Leu Lys
100 105 110
Tyr Leu Trp Thr Val Cys Cys His Cys Gly Gly Lys Thr Lys Glu Ala
115 120 125
Gln Lys Ile Asn Asn Gly Ser Ser Gln Ala Asp Gly Thr Leu Lys Pro
130 135 140
Val Asp Glu Lys Glu Glu Ala Val Ala Ala Glu Val Gly Trp Met Thr
145 150 155 160
Ser Val Lys Asp Trp Ala Gly Val Met Ile Ser Ala Gln Thr Leu Thr
165 170 175
Gly Arg Val Leu Val Val Leu Val Phe Ala Leu Ser Ile Gly Ala Leu
180 185 190
Val Ile Tyr Phe Ile Asp Ser Ser Asn Pro Ile Glu Ser Cys Gln Asn
195 200 205
Phe Tyr Lys Asp Phe Thr Leu Gln Ile Asp Met Ala Phe Asn Val Phe
210 215 220
Phe Leu Leu Tyr Phe Gly Leu Arg Phe Ile Ala Ala Asn Asp Lys Leu
225 230 235 240
Trp Phe Trp Leu Glu Val Asn Ser Val Val Asp Phe Phe Thr Val Pro
245 250 255
Pro Val Phe Val Ser Val Tyr Leu Asn Arg Ser Trp Leu Gly Leu Arg
260 265 270
Phe Leu Arg Ala Leu Arg Leu Ile Gln Phe Ser Glu Ile Leu Gln Phe
275 280 285
Leu Asn Ile Leu Lys Thr Ser Asn Ser Ile Lys Leu Val Asn Leu Leu
290 295 300
Ser Ile Phe Ile Ser Thr Trp Leu Thr Ala Ala Gly Phe Ile His Leu
305 310 315 320
Val Glu Asn Ser Gly Asp Pro Trp Glu Asn Phe Gln Asn Asn Gln Ala
325 330 335
Leu Thr Tyr Trp Glu Cys Val Tyr Leu Leu Met Val Thr Met Ser Thr
340 345 350
Val Gly Tyr Gly Asp Val Tyr Ala Lys Thr Thr Leu Gly Arg Leu Phe
355 360 365
Met Val Phe Phe Ile Leu Gly Gly Leu Ala Met Phe Ala Ser Tyr Val
370 375 380
Pro Glu Ile Ile Glu Leu Ile Gly Asn Arg Lys Lys Tyr Gly Gly Ser
385 390 395 400
Tyr Ser Ala Val Ser Gly Arg Lys His Ile Val Val Cys Gly His Ile
405 410 415
Thr Leu Glu Ser Val Ser Asn Phe Leu Lys Asp Phe Leu His Lys Asp
420 425 430
Arg Asp Asp Val Asn Val Glu Ile Val Phe Leu His Asn Ile Ser Pro
435 440 445
Asn Leu Glu Leu Glu Ala Leu Phe Lys Arg His Phe Thr Gln Val Glu
450 455 460
Phe Tyr Gln Gly Ser Val Leu Asn Pro His Asp Leu Ala Arg Val Lys
465 470 475 480
Ile Glu Ser Ala Asp Ala Cys Leu Ile Leu Ala Asn Lys Tyr Cys Ala
485 490 495
Asp Pro Asp Ala Glu Asp Ala Ser Asn Ile Met Arg Val Ile Ser Ile
500 505 510
Lys Asn Tyr His Pro Lys Ile Arg Ile Ile Thr Gln Met Leu Gln Tyr
515 520 525
His Asn Lys Ala His Leu Leu Asn Ile Pro Ser Trp Asn Trp Lys Glu
530 535 540
Gly Asp Asp Ala Ile Cys Leu Ala Glu Leu Lys Leu Gly Phe Ile Ala
545 550 555 560
Gln Ser Cys Leu Ala Gln Gly Leu Ser Thr Met Leu Ala Asn Leu Phe
565 570 575
Ser Met Arg Ser Phe Ile Lys Ile Glu Glu Asp Thr Trp Gln Lys Tyr
580 585 590
Tyr Leu Glu Gly Val Ser Asn Glu Met Tyr Thr Glu Tyr Leu Ser Ser
595 600 605
Ala Phe Val Gly Leu Ser Phe Pro Thr Val Cys Glu Leu Cys Phe Val
610 615 620
Lys Leu Lys Leu Leu Met Ile Ala Ile Glu Tyr Lys Ser Ala Asn Arg
625 630 635 640
Glu Ser Arg Ile Leu Ile Asn Pro Gly Asn His Leu Lys Ile Gln Glu
645 650 655
Gly Thr Leu Gly Phe Phe Ile Ala Ser Asp Ala Lys Glu Val Lys Arg
660 665 670
Ala Phe Phe Tyr Cys Lys Ala Cys His Asp Asp Ile Thr Asp Pro Lys
675 680 685
Arg Ile Lys Lys Cys Gly Cys Lys Arg Leu Glu Asp Glu Gln Pro Ser
690 695 700
Thr Leu Ser Pro Lys Lys Lys Gln Arg Asn Gly Gly Met Arg Asn Ser
705 710 715 720
Pro Asn Thr Ser Pro Lys Leu Met Arg His Asp Pro Leu Leu Ile Pro
725 730 735
Gly Asn Asp Gln Ile Asp Asn Met Asp Ser Asn Val Lys Lys Tyr Asp
740 745 750
Ser Thr Gly Met Phe His Trp Cys Ala Pro Lys Glu Ile Glu Lys Val
755 760 765
Ile Leu Thr Arg Ser Glu Ala Ala Met Thr Val Leu Ser Gly His Val
770 775 780
Val Val Cys Ile Phe Gly Asp Val Ser Ser Ala Leu Ile Gly Leu Arg
785 790 795 800
Asn Leu Val Met Pro Leu Arg Ala Ser Asn Phe His Tyr His Glu Leu
805 810 815
Lys His Ile Val Phe Val Gly Ser Ile Glu Tyr Leu Lys Arg Glu Trp
820 825 830
Glu Thr Leu His Asn Phe Pro Lys Val Ser Ile Leu Pro Gly Thr Pro
835 840 845
Leu Ser Arg Ala Asp Leu Arg Ala Val Asn Ile Asn Leu Cys Asp Met
850 855 860
Cys Val Ile Leu Ser Ala Asn Gln Asn Asn Ile Asp Asp Thr Ser Leu
865 870 875 880
Gln Asp Lys Glu Cys Ile Leu Ala Ser Leu Asn Ile Lys Ser Met Gln
885 890 895
Phe Asp Asp Ser Ile Gly Val Leu Gln Ala Asn Ser Gln Gly Phe Thr
900 905 910
Pro Pro Gly Met Asp Arg Ser Ser Pro Asp Asn Ser Pro Val His Gly
915 920 925
Met Leu Arg Gln Pro Ser Ile Thr Thr Gly Val Asn Ile Pro Ile Ile
930 935 940
Thr Glu Leu Val Asn Asp Thr Asn Val Gln Phe Leu Asp Gln Asp Asp
945 950 955 960
Asp Asp Asp Pro Asp Thr Glu Leu Tyr Leu Thr Gln Pro Phe Ala Cys
965 970 975
Gly Thr Ala Phe Ala Val Ser Val Leu Asp Ser Leu Met Ser Ala Thr
980 985 990
Tyr Phe Asn Asp Asn Ile Leu Thr Leu Ile Arg Thr Leu Val Thr Gly
995 1000 1005
Gly Ala Thr Pro Glu Leu Glu Ala Leu Ile Ala Glu Glu Asn Ala Leu
1010 1015 1020
Arg Gly Gly Tyr Ser Thr Pro Gln Thr Leu Ala Asn Arg Asp Arg Cys
1025 1030 1035 1040
Arg Val Ala Gln Leu Ala Leu Leu Asp Gly Pro Phe Ala Asp Leu Gly
1045 1050 1055
Asp Gly Gly Cys Tyr Gly Asp Leu Phe Cys Lys Ala Leu Lys Thr Tyr
1060 1065 1070
Asn Met Leu Cys Phe Gly Ile Tyr Arg Leu Arg Asp Ala His Leu Ser
1075 1080 1085
Thr Pro Ser Gln Cys Thr Lys Arg Tyr Val Ile Thr Asn Pro Pro Tyr
1090 1095 1100
Glu Phe Glu Leu Val Pro Thr Asp Leu Ile Phe Cys Leu Met Gln Phe
1105 1110 1115 1120
Asp His Asn Ala Gly Gln Ser Arg Ala Ser Leu Ser His Ser Ser His
1125 1130 1135
Ser Ser Gln Ser Ser Ser Lys Lys Ser Ser Ser Val His Ser Ile Pro
1140 1145 1150
Ser Thr Ala Asn Arg Gln Asn Arg Pro Lys Ser Arg Glu Ser Arg Asp
1155 1160 1165
Lys Gln Lys Tyr Val Gln Glu Glu Arg Leu
1170 1175
<210> 14
<211> 1140
<212> PRT
<213> 线虫(Caenorhabditis)
<400> 14
Met Gly Glu Ile Tyr Ser Pro Ser Gln Ser Lys Gly Phe Asn Gln Pro
1 5 10 15
Tyr Gly Tyr Pro Met Asn Cys Asn Leu Ser Arg Val Phe Met Glu Met
20 25 30
Thr Glu Glu Asp Arg Lys Cys Leu Glu Glu Arg Lys Tyr Trp Cys Phe
35 40 45
Leu Leu Ser Ser Ile Thr Thr Phe Cys Ala Ser Met Ile Leu Val Val
50 55 60
Ile Trp Arg Val Val Thr His Leu Cys Cys Gln Arg Arg Glu Lys Glu
65 70 75 80
Phe Val Glu Pro Ile Pro Ala Pro Glu Ala Val Gln Ile Asn Met Asn
85 90 95
Gly Ser Lys His Ala Pro Ser Glu Thr Asp Pro Phe Leu Lys Gln Gln
100 105 110
Glu Glu Lys His Leu Gly Trp Met Thr Glu Ala Lys Asp Trp Ala Gly
115 120 125
Glu Leu Ile Ser Gly Gln Ser Leu Thr Gly Arg Phe Leu Val Leu Leu
130 135 140
Val Phe Ile Leu Ser Ile Gly Ser Leu Ile Ile Tyr Phe Tyr Asp Ala
145 150 155 160
Ser Phe Gln Asn Phe Gln Val Glu Thr Cys Ile Pro Trp Gln Asp Ser
165 170 175
Pro Ser Gln Gln Ile Asp Leu Gly Phe Asn Ile Phe Phe Leu Val Tyr
180 185 190
Phe Phe Ile Arg Phe Ile Ala Ala Ser Asp Lys Val Trp Phe Leu Leu
195 200 205
Glu Met Tyr Ser Trp Ile Asp Phe Phe Thr Ile Pro Pro Ser Phe Val
210 215 220
Ala Ile Tyr Leu Gln Arg Asn Trp Leu Gly Phe Arg Phe Leu Arg Ala
225 230 235 240
Leu Arg Leu Met Thr Val Pro Asp Ile Leu Gln Tyr Leu Asn Ile Leu
245 250 255
Lys Thr Ser Ser Ser Ile Arg Leu Thr Gln Leu Val Thr Ile Phe Val
260 265 270
Ala Val Cys Leu Thr Gly Ala Gly Leu Val His Leu Leu Glu Asn Ser
275 280 285
Gly Asp Phe Phe Lys Gly Phe Ile Asn Pro His Arg Ile Thr Tyr Ala
290 295 300
Asp Ser Val Tyr Phe Val Leu Val Thr Met Ser Thr Val Gly Tyr Gly
305 310 315 320
Asp Ile Tyr Cys Thr Thr Leu Cys Gly Arg Leu Phe Met Ile Phe Phe
325 330 335
Ile Leu Phe Gly Leu Ala Met Phe Ala Ser Tyr Val Pro Glu Ile Ala
340 345 350
Asp Leu Ile Gly Asn Arg Gln Lys Tyr Gly Gly Glu Tyr Lys Gly Glu
355 360 365
His Gly Lys Lys His Ile Val Val Cys Gly His Ile Thr Tyr Asp Ser
370 375 380
Val Ser His Phe Leu Gln Asp Phe Leu His Glu Asp Arg Asp Asp Val
385 390 395 400
Asp Val Glu Val Val Phe Leu His Arg Val Val Pro Asp Leu Glu Leu
405 410 415
Glu Gly Leu Phe Lys Arg His Phe Thr Lys Val Glu Phe Phe Thr Gly
420 425 430
Thr Val Met Asp Ser Leu Asp Leu Ser Arg Val Lys Ile Gly Asp Ala
435 440 445
Asp Ala Cys Leu Val Leu Ala Asn Lys Tyr Ser Thr Asn Pro Asp Ala
450 455 460
Glu Asp Ala Ala Asn Ile Met Arg Val Ile Ser Ile Lys Asn Tyr Ser
465 470 475 480
Ser Asp Ile Arg Val Ile Val Gln Leu Met Gln Tyr His Asn Lys Ala
485 490 495
Tyr Leu Leu Asn Ile Pro Ser Trp Asp Trp Lys Arg Gly Asp Asp Val
500 505 510
Ile Cys Leu Ala Glu Leu Lys Leu Gly Phe Ile Ala Gln Ser Cys Leu
515 520 525
Ala Pro Gly Phe Ser Thr Met Met Ala Asn Leu Phe Ala Met Arg Ser
530 535 540
Phe Lys Thr Ser Gln Thr Thr Pro Asp Trp Leu Asn Leu Tyr Leu Cys
545 550 555 560
Gly Ala Gly Met Glu Met Tyr Thr Asp Thr Leu Ser His Ser Phe Val
565 570 575
Gly Met Thr Phe Pro Glu Ala Val Asp Leu Leu Phe Asn Arg Leu Gly
580 585 590
Leu Leu Leu Leu Ala Ile Glu Leu Lys Asp Glu Glu Asn Lys Glu Cys
595 600 605
Asn Ile Ala Ile Asn Pro Gly Pro His Ile Val Ile Gln Pro Gln Thr
610 615 620
Gln Gly Phe Phe Ile Ala Gln Ser Ala Asp Glu Val Lys Arg Ala Phe
625 630 635 640
Phe Trp Cys Lys Gln Cys His Asp Asp Ile Lys Asp Val Ser Leu Ile
645 650 655
Lys Lys Cys Lys Cys Lys Asn Leu Ala Leu Phe Arg Arg Asn Thr Lys
660 665 670
His Ser Thr Ala Ala Arg Ala Arg Ala Thr Asp Val Leu Gln Gln Phe
675 680 685
Gln Pro Gln Ala Pro Ala Gly Pro Met Gly His Leu Gly Gln Gln Val
690 695 700
Gln Leu Arg Met Ile Asn Gln Gln Ser Ser Thr Ser Asp Thr His Leu
705 710 715 720
Asn Thr Lys Ser Leu Arg Phe Ala Tyr Glu Ile Lys Lys Leu Met Pro
725 730 735
Ser Ser Gly Gly Arg Arg Asn Ser Met Ser Ile Pro Pro Asp Gly Arg
740 745 750
Gly Val Asp Phe Ser Lys Asp Phe Glu Gln Gln Phe Gln Asp Met Lys
755 760 765
Tyr Asp Ser Thr Gly Met Phe His Trp Cys Pro Ser Arg Asn Leu Glu
770 775 780
Asp Cys Val Leu Glu Arg His Gln Ala Ala Met Thr Val Leu Asn Gly
785 790 795 800
His Val Val Val Cys Leu Phe Ala Asp Gln Asp Ser Pro Leu Ile Gly
805 810 815
Leu Arg Asn Phe Ile Met Pro Leu Arg Ser Ser Asn Phe His Tyr His
820 825 830
Glu Leu Lys His Val Val Ile Val Gly Asp Leu Glu Tyr Leu Arg Lys
835 840 845
Glu Trp Lys Thr Leu Tyr Asn Leu Pro Lys Ile Ser Ile Leu Asn Gly
850 855 860
Ser Pro Leu Ser Arg Ala Asp Leu Arg Ala Val Asn Ile Asn Leu Cys
865 870 875 880
Asp Met Cys Val Ile Ile Ser Ala Arg Val Pro Asn Thr Glu Asp Thr
885 890 895
Thr Leu Ala Asp Lys Glu Ala Ile Leu Ala Ser Leu Asn Ile Lys Ala
900 905 910
Met Gln Phe Asp Asp Thr Leu Gly Phe Phe Pro Met Arg His Gln Thr
915 920 925
Gly Asp Arg Ser Pro Leu Gly Ser Pro Ile Ser Met Gln Lys Lys Gly
930 935 940
Ala Lys Phe Gly Thr Asn Val Pro Met Ile Thr Glu Leu Val Asn Asp
945 950 955 960
Ser Asn Val Gln Phe Leu Asp Gln Asp Asp Asp Asp Asp Pro Asp Thr
965 970 975
Glu Leu Tyr Leu Thr Gln Pro Phe Ala Cys Gly Thr Ala Phe Ala Ile
980 985 990
Ser Val Leu Asp Ser Leu Met Ser Thr Thr Tyr Phe Asn Asp Ser Ala
995 1000 1005
Leu Thr Leu Ile Arg Thr Leu Val Thr Gly Gly Ala Thr Pro Glu Leu
1010 1015 1020
Glu Leu Ile Leu Ala Glu Gly Ala Gly Leu Arg Gly Gly Tyr Ser Thr
1025 1030 1035 1040
Pro Glu Thr Leu Ser Asn Arg Asp Arg Cys Arg Ile Ala Gln Ile Ser
1045 1050 1055
Leu Gln Asp Asn Pro Tyr Asp Gly Val Val His Asn Thr Thr Tyr Gly
1060 1065 1070
Ala Met Phe Thr Ile Ala Leu Arg Arg Tyr Gly Gln Leu Cys Ile Gly
1075 1080 1085
Leu Tyr Arg Leu His Asp Gln Asp Asn Pro Asp Ser Met Lys Arg Tyr
1090 1095 1100
Val Ile Thr Asn Pro Pro Ala Glu Leu Arg Ile Lys Asn Thr Asp Tyr
1105 1110 1115 1120
Val Tyr Val Leu Glu Gln Phe Asp Pro Gly Leu Glu Tyr Glu Pro Gly
1125 1130 1135
Lys Arg His Phe
1140
<210> 15
<211> 1174
<212> PRT
<213> 人工序列(Artificial Sequence)
<220>
<221> PEPTIDE
<222> (1)..(1174)
<223> BKchimera蛋白氨基酸序列
<400> 15
Met Ala Asn Gly Gly Gly Gly Gly Gly Gly Ser Ser Gly Gly Gly Gly
1 5 10 15
Gly Gly Gly Gly Ser Ser Leu Arg Met Ser Ser Asn Ile His Ala Asn
20 25 30
His Leu Ser Leu Asp Ala Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser
35 40 45
Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser Val His Glu Pro
50 55 60
Lys Met Asp Ala Leu Ile Ile Pro Val Thr Met Glu Val Pro Cys Asp
65 70 75 80
Ser Arg Gly Gln Arg Met Trp Trp Ala Phe Leu Ala Ser Ser Met Val
85 90 95
Thr Phe Phe Gly Gly Leu Phe Ile Ile Leu Leu Trp Arg Thr Leu Lys
100 105 110
Tyr Leu Trp Thr Val Cys Cys His Cys Gly Gly Lys Thr Lys Glu Ala
115 120 125
Gln Lys Ile Asn Asn Gly Ser Ser Gln Ala Asp Gly Thr Leu Lys Pro
130 135 140
Val Asp Glu Lys Glu Glu Ala Val Ala Ala Glu Val Gly Trp Met Thr
145 150 155 160
Ser Val Lys Asp Trp Ala Gly Val Met Ile Ser Ala Gln Thr Leu Thr
165 170 175
Gly Arg Val Leu Val Val Leu Val Phe Ala Leu Ser Ile Gly Ala Leu
180 185 190
Val Ile Tyr Phe Ile Asp Ser Ser Asn Pro Ile Glu Ser Cys Gln Asn
195 200 205
Phe Tyr Lys Asp Phe Thr Leu Gln Ile Asp Met Ala Phe Asn Val Phe
210 215 220
Phe Leu Leu Tyr Phe Gly Leu Arg Phe Ile Ala Ala Asn Asp Lys Leu
225 230 235 240
Trp Phe Trp Leu Glu Val Asn Ser Val Val Asp Phe Phe Thr Val Pro
245 250 255
Pro Val Phe Val Ser Val Tyr Leu Asn Arg Ser Trp Leu Gly Leu Arg
260 265 270
Phe Leu Arg Ala Leu Arg Leu Ile Gln Phe Ser Glu Ile Leu Gln Phe
275 280 285
Leu Asn Ile Leu Lys Thr Ser Asn Ser Ile Lys Leu Val Asn Leu Leu
290 295 300
Ser Ile Phe Ile Ser Thr Trp Leu Thr Ala Ala Gly Phe Ile His Leu
305 310 315 320
Val Glu Asn Ser Gly Asp Pro Trp Glu Asn Phe Gln Asn Asn Gln Ala
325 330 335
Leu Thr Tyr Trp Glu Cys Val Tyr Leu Leu Met Val Thr Met Ser Thr
340 345 350
Val Gly Tyr Gly Asp Val Tyr Ala Lys Thr Thr Leu Gly Arg Leu Phe
355 360 365
Met Val Phe Phe Ile Leu Gly Gly Leu Ala Met Phe Ala Ser Tyr Val
370 375 380
Pro Glu Ile Ile Glu Leu Ile Gly Asn Arg Lys Lys Tyr Gly Gly Ser
385 390 395 400
Tyr Ser Ala Val Ser Gly Arg Lys His Ile Val Val Cys Gly His Ile
405 410 415
Thr Leu Glu Ser Val Ser Asn Phe Leu Lys Asp Phe Leu His Lys Asp
420 425 430
Arg Asp Asp Val Asn Val Glu Ile Val Phe Leu His Asn Ile Ser Pro
435 440 445
Asn Leu Glu Leu Glu Ala Leu Phe Lys Arg His Phe Thr Gln Val Glu
450 455 460
Phe Tyr Gln Gly Ser Val Leu Asn Pro His Asp Leu Ala Arg Val Lys
465 470 475 480
Ile Glu Ser Ala Asp Ala Cys Leu Ile Leu Ala Asn Lys Tyr Cys Ala
485 490 495
Asp Pro Asp Ala Glu Asp Ala Ser Asn Ile Met Arg Val Ile Ser Ile
500 505 510
Lys Asn Tyr His Pro Lys Ile Arg Ile Ile Thr Gln Met Leu Gln Tyr
515 520 525
His Asn Lys Ala His Leu Leu Asn Ile Pro Ser Trp Asn Trp Lys Glu
530 535 540
Gly Asp Asp Ala Ile Cys Leu Ala Glu Leu Lys Leu Gly Phe Ile Ala
545 550 555 560
Gln Ser Cys Leu Ala Gln Gly Leu Ser Thr Met Leu Ala Asn Leu Phe
565 570 575
Ser Met Arg Ser Phe Lys Thr Ser Gln Thr Thr Pro Asp Trp Leu Asn
580 585 590
Leu Tyr Leu Cys Gly Ala Gly Met Glu Met Tyr Thr Asp Thr Leu Ser
595 600 605
His Ser Phe Val Gly Met Thr Phe Pro Glu Ala Val Asp Leu Leu Phe
610 615 620
Asn Arg Leu Gly Leu Leu Leu Leu Ala Ile Glu Leu Lys Asp Glu Glu
625 630 635 640
Asn Lys Glu Cys Asn Ile Ala Ile Asn Pro Gly Pro His Ile Val Ile
645 650 655
Gln Pro Gln Thr Gln Gly Phe Phe Ile Ala Gln Ser Ala Asp Glu Val
660 665 670
Lys Arg Ala Phe Phe Trp Cys Lys Gln Cys His Asp Asp Ile Lys Asp
675 680 685
Val Ser Leu Ile Lys Lys Cys Lys Cys Lys Asn Leu Ala Leu Phe Arg
690 695 700
Arg Asn Thr Lys His Ser Thr Ala Ala Arg Ala Arg Ala Thr Asp Val
705 710 715 720
Leu Gln Gln Phe Gln Pro Gln Ala Pro Ala Gly Pro Met Gly His Leu
725 730 735
Gly Gln Gln Val Gln Met Ile Asn Gln Gln Ser Ser Thr Ser Asp Thr
740 745 750
His Leu Asn Thr Lys Ser Leu Arg Phe Ala Tyr Glu Ile Lys Lys Leu
755 760 765
Met Pro Ser Ser Gly Gly Arg Arg Asn Ser Met Ser Ile Pro Pro Asp
770 775 780
Gly Arg Gly Val Asp Phe Ser Lys Asp Phe Glu Gln Gln Phe Gln Asp
785 790 795 800
Met Lys Tyr Asp Ser Thr Gly Met Phe His Trp Cys Pro Ser Arg Asn
805 810 815
Leu Glu Asp Cys Val Leu Glu Arg His Gln Ala Ala Met Thr Val Leu
820 825 830
Asn Gly His Val Val Val Cys Leu Phe Ala Asp Gln Asp Ser Pro Leu
835 840 845
Ile Gly Leu Arg Asn Phe Ile Met Pro Leu Arg Ser Ser Asn Phe His
850 855 860
Tyr His Glu Leu Lys His Val Val Ile Val Gly Asp Leu Glu Tyr Leu
865 870 875 880
Arg Lys Glu Trp Lys Thr Leu Tyr Asn Leu Pro Lys Ile Ser Ile Leu
885 890 895
Asn Gly Ser Pro Leu Ser Arg Ala Asp Leu Arg Ala Val Asn Ile Asn
900 905 910
Leu Cys Asp Met Cys Val Ile Ile Ser Ala Arg Val Pro Asn Thr Glu
915 920 925
Asp Thr Thr Leu Ala Asp Lys Glu Ala Ile Leu Ala Ser Leu Asn Ile
930 935 940
Lys Ala Met Gln Phe Asp Asp Thr Leu Gly Phe Phe Pro Met Arg His
945 950 955 960
Gln Thr Gly Asp Arg Ser Pro Leu Gly Ser Pro Ile Ser Met Gln Lys
965 970 975
Lys Gly Ala Lys Phe Gly Thr Asn Val Pro Met Ile Thr Glu Leu Val
980 985 990
Asn Asp Ser Asn Val Gln Phe Leu Asp Gln Asp Asp Asp Asp Asp Pro
995 1000 1005
Asp Thr Glu Leu Tyr Leu Thr Gln Pro Phe Ala Cys Gly Thr Ala Phe
1010 1015 1020
Ala Ile Ser Val Leu Asp Ser Leu Met Ser Thr Thr Tyr Phe Asn Asp
1025 1030 1035 1040
Ser Ala Leu Thr Leu Ile Arg Thr Leu Val Thr Gly Gly Ala Thr Pro
1045 1050 1055
Glu Leu Glu Leu Ile Leu Ala Glu Gly Ala Gly Leu Arg Gly Gly Tyr
1060 1065 1070
Ser Thr Pro Glu Thr Leu Ser Asn Arg Asp Arg Cys Arg Ile Ala Gln
1075 1080 1085
Ile Ser Leu Gln Asp Asn Pro Tyr Asp Gly Val Val His Asn Thr Thr
1090 1095 1100
Tyr Gly Ala Met Phe Thr Ile Ala Leu Arg Arg Tyr Gly Gln Leu Cys
1105 1110 1115 1120
Ile Gly Leu Tyr Arg Leu His Asp Gln Asp Asn Pro Asp Ser Met Lys
1125 1130 1135
Arg Tyr Val Ile Thr Asn Pro Pro Ala Glu Leu Arg Ile Lys Asn Thr
1140 1145 1150
Asp Tyr Val Tyr Val Leu Glu Gln Phe Asp Pro Gly Leu Glu Tyr Glu
1155 1160 1165
Pro Gly Lys Arg His Phe
1170
<210> 16
<211> 1174
<212> PRT
<213> 人工序列(Artificial Sequence)
<220>
<221> PEPTIDE
<222> (1)..(1174)
<223> BKchimera/D434G蛋白氨基酸序列
<400> 16
Met Ala Asn Gly Gly Gly Gly Gly Gly Gly Ser Ser Gly Gly Gly Gly
1 5 10 15
Gly Gly Gly Gly Ser Ser Leu Arg Met Ser Ser Asn Ile His Ala Asn
20 25 30
His Leu Ser Leu Asp Ala Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser
35 40 45
Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser Ser Val His Glu Pro
50 55 60
Lys Met Asp Ala Leu Ile Ile Pro Val Thr Met Glu Val Pro Cys Asp
65 70 75 80
Ser Arg Gly Gln Arg Met Trp Trp Ala Phe Leu Ala Ser Ser Met Val
85 90 95
Thr Phe Phe Gly Gly Leu Phe Ile Ile Leu Leu Trp Arg Thr Leu Lys
100 105 110
Tyr Leu Trp Thr Val Cys Cys His Cys Gly Gly Lys Thr Lys Glu Ala
115 120 125
Gln Lys Ile Asn Asn Gly Ser Ser Gln Ala Asp Gly Thr Leu Lys Pro
130 135 140
Val Asp Glu Lys Glu Glu Ala Val Ala Ala Glu Val Gly Trp Met Thr
145 150 155 160
Ser Val Lys Asp Trp Ala Gly Val Met Ile Ser Ala Gln Thr Leu Thr
165 170 175
Gly Arg Val Leu Val Val Leu Val Phe Ala Leu Ser Ile Gly Ala Leu
180 185 190
Val Ile Tyr Phe Ile Asp Ser Ser Asn Pro Ile Glu Ser Cys Gln Asn
195 200 205
Phe Tyr Lys Asp Phe Thr Leu Gln Ile Asp Met Ala Phe Asn Val Phe
210 215 220
Phe Leu Leu Tyr Phe Gly Leu Arg Phe Ile Ala Ala Asn Asp Lys Leu
225 230 235 240
Trp Phe Trp Leu Glu Val Asn Ser Val Val Asp Phe Phe Thr Val Pro
245 250 255
Pro Val Phe Val Ser Val Tyr Leu Asn Arg Ser Trp Leu Gly Leu Arg
260 265 270
Phe Leu Arg Ala Leu Arg Leu Ile Gln Phe Ser Glu Ile Leu Gln Phe
275 280 285
Leu Asn Ile Leu Lys Thr Ser Asn Ser Ile Lys Leu Val Asn Leu Leu
290 295 300
Ser Ile Phe Ile Ser Thr Trp Leu Thr Ala Ala Gly Phe Ile His Leu
305 310 315 320
Val Glu Asn Ser Gly Asp Pro Trp Glu Asn Phe Gln Asn Asn Gln Ala
325 330 335
Leu Thr Tyr Trp Glu Cys Val Tyr Leu Leu Met Val Thr Met Ser Thr
340 345 350
Val Gly Tyr Gly Asp Val Tyr Ala Lys Thr Thr Leu Gly Arg Leu Phe
355 360 365
Met Val Phe Phe Ile Leu Gly Gly Leu Ala Met Phe Ala Ser Tyr Val
370 375 380
Pro Glu Ile Ile Glu Leu Ile Gly Asn Arg Lys Lys Tyr Gly Gly Ser
385 390 395 400
Tyr Ser Ala Val Ser Gly Arg Lys His Ile Val Val Cys Gly His Ile
405 410 415
Thr Leu Glu Ser Val Ser Asn Phe Leu Lys Asp Phe Leu His Lys Asp
420 425 430
Arg Gly Asp Val Asn Val Glu Ile Val Phe Leu His Asn Ile Ser Pro
435 440 445
Asn Leu Glu Leu Glu Ala Leu Phe Lys Arg His Phe Thr Gln Val Glu
450 455 460
Phe Tyr Gln Gly Ser Val Leu Asn Pro His Asp Leu Ala Arg Val Lys
465 470 475 480
Ile Glu Ser Ala Asp Ala Cys Leu Ile Leu Ala Asn Lys Tyr Cys Ala
485 490 495
Asp Pro Asp Ala Glu Asp Ala Ser Asn Ile Met Arg Val Ile Ser Ile
500 505 510
Lys Asn Tyr His Pro Lys Ile Arg Ile Ile Thr Gln Met Leu Gln Tyr
515 520 525
His Asn Lys Ala His Leu Leu Asn Ile Pro Ser Trp Asn Trp Lys Glu
530 535 540
Gly Asp Asp Ala Ile Cys Leu Ala Glu Leu Lys Leu Gly Phe Ile Ala
545 550 555 560
Gln Ser Cys Leu Ala Gln Gly Leu Ser Thr Met Leu Ala Asn Leu Phe
565 570 575
Ser Met Arg Ser Phe Lys Thr Ser Gln Thr Thr Pro Asp Trp Leu Asn
580 585 590
Leu Tyr Leu Cys Gly Ala Gly Met Glu Met Tyr Thr Asp Thr Leu Ser
595 600 605
His Ser Phe Val Gly Met Thr Phe Pro Glu Ala Val Asp Leu Leu Phe
610 615 620
Asn Arg Leu Gly Leu Leu Leu Leu Ala Ile Glu Leu Lys Asp Glu Glu
625 630 635 640
Asn Lys Glu Cys Asn Ile Ala Ile Asn Pro Gly Pro His Ile Val Ile
645 650 655
Gln Pro Gln Thr Gln Gly Phe Phe Ile Ala Gln Ser Ala Asp Glu Val
660 665 670
Lys Arg Ala Phe Phe Trp Cys Lys Gln Cys His Asp Asp Ile Lys Asp
675 680 685
Val Ser Leu Ile Lys Lys Cys Lys Cys Lys Asn Leu Ala Leu Phe Arg
690 695 700
Arg Asn Thr Lys His Ser Thr Ala Ala Arg Ala Arg Ala Thr Asp Val
705 710 715 720
Leu Gln Gln Phe Gln Pro Gln Ala Pro Ala Gly Pro Met Gly His Leu
725 730 735
Gly Gln Gln Val Gln Met Ile Asn Gln Gln Ser Ser Thr Ser Asp Thr
740 745 750
His Leu Asn Thr Lys Ser Leu Arg Phe Ala Tyr Glu Ile Lys Lys Leu
755 760 765
Met Pro Ser Ser Gly Gly Arg Arg Asn Ser Met Ser Ile Pro Pro Asp
770 775 780
Gly Arg Gly Val Asp Phe Ser Lys Asp Phe Glu Gln Gln Phe Gln Asp
785 790 795 800
Met Lys Tyr Asp Ser Thr Gly Met Phe His Trp Cys Pro Ser Arg Asn
805 810 815
Leu Glu Asp Cys Val Leu Glu Arg His Gln Ala Ala Met Thr Val Leu
820 825 830
Asn Gly His Val Val Val Cys Leu Phe Ala Asp Gln Asp Ser Pro Leu
835 840 845
Ile Gly Leu Arg Asn Phe Ile Met Pro Leu Arg Ser Ser Asn Phe His
850 855 860
Tyr His Glu Leu Lys His Val Val Ile Val Gly Asp Leu Glu Tyr Leu
865 870 875 880
Arg Lys Glu Trp Lys Thr Leu Tyr Asn Leu Pro Lys Ile Ser Ile Leu
885 890 895
Asn Gly Ser Pro Leu Ser Arg Ala Asp Leu Arg Ala Val Asn Ile Asn
900 905 910
Leu Cys Asp Met Cys Val Ile Ile Ser Ala Arg Val Pro Asn Thr Glu
915 920 925
Asp Thr Thr Leu Ala Asp Lys Glu Ala Ile Leu Ala Ser Leu Asn Ile
930 935 940
Lys Ala Met Gln Phe Asp Asp Thr Leu Gly Phe Phe Pro Met Arg His
945 950 955 960
Gln Thr Gly Asp Arg Ser Pro Leu Gly Ser Pro Ile Ser Met Gln Lys
965 970 975
Lys Gly Ala Lys Phe Gly Thr Asn Val Pro Met Ile Thr Glu Leu Val
980 985 990
Asn Asp Ser Asn Val Gln Phe Leu Asp Gln Asp Asp Asp Asp Asp Pro
995 1000 1005
Asp Thr Glu Leu Tyr Leu Thr Gln Pro Phe Ala Cys Gly Thr Ala Phe
1010 1015 1020
Ala Ile Ser Val Leu Asp Ser Leu Met Ser Thr Thr Tyr Phe Asn Asp
1025 1030 1035 1040
Ser Ala Leu Thr Leu Ile Arg Thr Leu Val Thr Gly Gly Ala Thr Pro
1045 1050 1055
Glu Leu Glu Leu Ile Leu Ala Glu Gly Ala Gly Leu Arg Gly Gly Tyr
1060 1065 1070
Ser Thr Pro Glu Thr Leu Ser Asn Arg Asp Arg Cys Arg Ile Ala Gln
1075 1080 1085
Ile Ser Leu Gln Asp Asn Pro Tyr Asp Gly Val Val His Asn Thr Thr
1090 1095 1100
Tyr Gly Ala Met Phe Thr Ile Ala Leu Arg Arg Tyr Gly Gln Leu Cys
1105 1110 1115 1120
Ile Gly Leu Tyr Arg Leu His Asp Gln Asp Asn Pro Asp Ser Met Lys
1125 1130 1135
Arg Tyr Val Ile Thr Asn Pro Pro Ala Glu Leu Arg Ile Lys Asn Thr
1140 1145 1150
Asp Tyr Val Tyr Val Leu Glu Gln Phe Asp Pro Gly Leu Glu Tyr Glu
1155 1160 1165
Pro Gly Lys Arg His Phe
1170
<210> 17
<211> 3522
<212> DNA
<213> 人工序列(Artificial Sequence)
<220>
<221> misc_feature
<222> (1)..(3522)
<223> BKchimera cDNA序列
<400> 17
atggcaaatg gtggcggcgg cggcggcggc agcagcggcg gcggcggcgg cggcggaggc 60
agcagtctta gaatgagtag caatatccac gcgaaccatc tcagcctaga cgcgtcctcc 120
tcctcctcct cctcctcttc ctcttcttct tcttcctcct cctcttcctc ctcgtcctcg 180
gtccacgagc ccaagatgga tgcgctcatc atcccggtga ccatggaggt gccgtgcgac 240
agccggggcc aacgcatgtg gtgggctttc ctggcctcct ccatggtgac tttcttcggg 300
ggcctcttca tcatcttgct ctggcggacg ctcaagtacc tgtggaccgt gtgctgccac 360
tgcgggggca agacgaagga ggcccagaag attaacaatg gctcaagcca ggcggatggc 420
actctcaaac cagtggatga aaaagaggag gcagtggccg ccgaggtcgg ctggatgacc 480
tccgtgaagg actgggcggg ggtgatgata tccgcccaga cactgactgg cagagtcctg 540
gttgtcttag tctttgctct cagcatcggt gcacttgtaa tatacttcat agattcatca 600
aacccaatag aatcctgcca gaatttctac aaagatttca cattacagat cgacatggct 660
ttcaacgtgt tcttccttct ctacttcggc ttgcggttta ttgcagccaa cgataaattg 720
tggttctggc tggaagtgaa ctctgtagtg gatttcttca cggtgccccc cgtgtttgtg 780
tctgtgtact taaacagaag ttggcttggt ttgagatttt taagagctct gagactgata 840
cagttttcag aaattttgca gtttctgaat attcttaaaa caagtaattc catcaagctg 900
gtgaatctgc tctccatatt tatcagcacg tggctgactg cagccgggtt catccatttg 960
gtggagaatt caggggaccc atgggaaaat ttccaaaaca accaggctct cacctactgg 1020
gaatgtgtct atttactcat ggtcacaatg tccaccgttg gttatgggga tgtttatgca 1080
aaaaccacac ttgggcgcct cttcatggtc ttcttcatcc tcgggggact ggccatgttt 1140
gccagctacg tccctgaaat catagagtta ataggaaacc gcaagaaata cgggggctcc 1200
tatagtgcgg ttagtggaag aaagcacatt gtggtctgcg gacacatcac tctggagagt 1260
gtttccaact tcctgaagga ctttctgcac aaggaccggg atgacgtcaa tgtggagatc 1320
gtttttcttc acaacatctc ccccaacctg gagcttgaag ctctgttcaa acgacatttt 1380
actcaggtgg aattttatca gggttccgtc ctcaatccac atgatcttgc aagagtcaag 1440
atagagtcag cagatgcatg cctgatcctt gccaacaagt actgcgctga cccggatgcg 1500
gaggatgcct cgaatatcat gagagtaatc tccataaaga actaccatcc gaagataaga 1560
atcatcactc aaatgctgca gtatcacaac aaggcccatc tgctaaacat cccgagctgg 1620
aattggaaag aaggtgatga cgcaatctgc ctcgcagagt tgaagttggg cttcatagcc 1680
cagagctgcc tggctcaagg cctctccacc atgcttgcca acctcttctc catgaggtca 1740
ttcaaaacgt cacaaaccac accggactgg ctgaacctgt acctttgcgg tgccggaatg 1800
gaaatgtaca ccgacacgct atcccactcg ttcgtcggaa tgacttttcc tgaagctgtc 1860
gacctattat tcaaccgcct ggggcttctt cttctggcga ttgagcttaa agatgaggaa 1920
aataaagaat gcaatattgc aatcaatcca ggcccacata ttgtcatcca gcctcaaact 1980
caaggatttt tcattgctca gagcgctgac gaggtgaagc gggcgttttt ctggtgtaag 2040
cagtgtcatg acgatattaa ggacgtttcc ttgatcaaaa aatgtaaatg caaaaatttg 2100
gccctcttcc gtcgcaacac caaacactcg acagcggccc gtgcaagagc caccgatgtg 2160
ctccagcagt tccagccaca agcgccagca ggtccgatgg gccatttggg ccaacaagtt 2220
cagatgatta atcaacagag ttcgacgagt gatactcatc tcaacacgaa atcccttcga 2280
tttgcctatg aaatcaaaaa actcatgcca tcttccggag gacgccgaaa ttcgatgagt 2340
attccgccag acggcagggg tgtcgatttt agcaaggatt ttgagcagca attccaagac 2400
atgaaatatg acagtactgg aatgttccac tggtgtccat cccgtaacct agaggactgt 2460
gttctggagc gccaccaggc cgctatgacc gtactcaacg gccacgtggt ggtttgtttg 2520
ttcgctgatc aggattctcc gctaattggg ctccggaact tcataatgcc actaagaagc 2580
tcaaacttcc attatcatga gctcaaacac gtggttattg tgggagacct ggaatatttg 2640
aggaaagagt ggaagacgct gtacaacttg ccgaaaatct cgattttgaa tggctccccg 2700
ctatctcgag cagatttacg cgccgtcaac attaacctct gcgacatgtg tgtcataatc 2760
tctgcccggg tcccaaatac cgaagatacc actttggcag ataaggaagc gattctggca 2820
tcattgaaca ttaaagctat gcaatttgat gatacccttg gctttttccc gatgaggcat 2880
caaactggcg atcggagccc gctgggctcc ccgatttcca tgcagaagaa gggcgccaag 2940
tttgggacca atgtgccgat gattacggag cttgttaacg actcaaatgt gcaattcctc 3000
gaccaggacg acgacgacga tccggacacc gagctctacc tcacgcagcc cttcgcctgc 3060
ggaactgcct tcgccatttc agtgttggat tctttgatga gtactaccta cttcaacgac 3120
tcggccctga ccctcatccg aactctggtc actggtggag ccacaccgga gcttgaactt 3180
attctggcag aaggcgccgg gctccgaggt ggctacagta ccccggagac cctctcaaac 3240
agagatcggt gccgaattgc tcagatttcg ctgcaggata acccgtacga tggagttgtg 3300
cacaacacta cctatggtgc aatgttcacg atagctctga ggaggtacgg ccaactttgt 3360
atcgggctgt ataggcttca cgaccaggat aatccggact ctatgaagcg atatgttatc 3420
actaatcctc cagcagaatt gaggattaaa aatactgatt acgtctacgt acttgaacaa 3480
ttcgatccgg gactggagta cgagccgggc aaacgacact tt 3522
<210> 18
<211> 3522
<212> DNA
<213> 人工序列(Artificial Sequence)
<220>
<221> misc_feature
<222> (1)..(3522)
<223> BKchimera/D434G cDNA序列
<400> 18
atggcaaatg gtggcggcgg cggcggcggc agcagcggcg gcggcggcgg cggcggaggc 60
agcagtctta gaatgagtag caatatccac gcgaaccatc tcagcctaga cgcgtcctcc 120
tcctcctcct cctcctcttc ctcttcttct tcttcctcct cctcttcctc ctcgtcctcg 180
gtccacgagc ccaagatgga tgcgctcatc atcccggtga ccatggaggt gccgtgcgac 240
agccggggcc aacgcatgtg gtgggctttc ctggcctcct ccatggtgac tttcttcggg 300
ggcctcttca tcatcttgct ctggcggacg ctcaagtacc tgtggaccgt gtgctgccac 360
tgcgggggca agacgaagga ggcccagaag attaacaatg gctcaagcca ggcggatggc 420
actctcaaac cagtggatga aaaagaggag gcagtggccg ccgaggtcgg ctggatgacc 480
tccgtgaagg actgggcggg ggtgatgata tccgcccaga cactgactgg cagagtcctg 540
gttgtcttag tctttgctct cagcatcggt gcacttgtaa tatacttcat agattcatca 600
aacccaatag aatcctgcca gaatttctac aaagatttca cattacagat cgacatggct 660
ttcaacgtgt tcttccttct ctacttcggc ttgcggttta ttgcagccaa cgataaattg 720
tggttctggc tggaagtgaa ctctgtagtg gatttcttca cggtgccccc cgtgtttgtg 780
tctgtgtact taaacagaag ttggcttggt ttgagatttt taagagctct gagactgata 840
cagttttcag aaattttgca gtttctgaat attcttaaaa caagtaattc catcaagctg 900
gtgaatctgc tctccatatt tatcagcacg tggctgactg cagccgggtt catccatttg 960
gtggagaatt caggggaccc atgggaaaat ttccaaaaca accaggctct cacctactgg 1020
gaatgtgtct atttactcat ggtcacaatg tccaccgttg gttatgggga tgtttatgca 1080
aaaaccacac ttgggcgcct cttcatggtc ttcttcatcc tcgggggact ggccatgttt 1140
gccagctacg tccctgaaat catagagtta ataggaaacc gcaagaaata cgggggctcc 1200
tatagtgcgg ttagtggaag aaagcacatt gtggtctgcg gacacatcac tctggagagt 1260
gtttccaact tcctgaagga ctttctgcac aaggaccggg gtgacgtcaa tgtggagatc 1320
gtttttcttc acaacatctc ccccaacctg gagcttgaag ctctgttcaa acgacatttt 1380
actcaggtgg aattttatca gggttccgtc ctcaatccac atgatcttgc aagagtcaag 1440
atagagtcag cagatgcatg cctgatcctt gccaacaagt actgcgctga cccggatgcg 1500
gaggatgcct cgaatatcat gagagtaatc tccataaaga actaccatcc gaagataaga 1560
atcatcactc aaatgctgca gtatcacaac aaggcccatc tgctaaacat cccgagctgg 1620
aattggaaag aaggtgatga cgcaatctgc ctcgcagagt tgaagttggg cttcatagcc 1680
cagagctgcc tggctcaagg cctctccacc atgcttgcca acctcttctc catgaggtca 1740
ttcaaaacgt cacaaaccac accggactgg ctgaacctgt acctttgcgg tgccggaatg 1800
gaaatgtaca ccgacacgct atcccactcg ttcgtcggaa tgacttttcc tgaagctgtc 1860
gacctattat tcaaccgcct ggggcttctt cttctggcga ttgagcttaa agatgaggaa 1920
aataaagaat gcaatattgc aatcaatcca ggcccacata ttgtcatcca gcctcaaact 1980
caaggatttt tcattgctca gagcgctgac gaggtgaagc gggcgttttt ctggtgtaag 2040
cagtgtcatg acgatattaa ggacgtttcc ttgatcaaaa aatgtaaatg caaaaatttg 2100
gccctcttcc gtcgcaacac caaacactcg acagcggccc gtgcaagagc caccgatgtg 2160
ctccagcagt tccagccaca agcgccagca ggtccgatgg gccatttggg ccaacaagtt 2220
cagatgatta atcaacagag ttcgacgagt gatactcatc tcaacacgaa atcccttcga 2280
tttgcctatg aaatcaaaaa actcatgcca tcttccggag gacgccgaaa ttcgatgagt 2340
attccgccag acggcagggg tgtcgatttt agcaaggatt ttgagcagca attccaagac 2400
atgaaatatg acagtactgg aatgttccac tggtgtccat cccgtaacct agaggactgt 2460
gttctggagc gccaccaggc cgctatgacc gtactcaacg gccacgtggt ggtttgtttg 2520
ttcgctgatc aggattctcc gctaattggg ctccggaact tcataatgcc actaagaagc 2580
tcaaacttcc attatcatga gctcaaacac gtggttattg tgggagacct ggaatatttg 2640
aggaaagagt ggaagacgct gtacaacttg ccgaaaatct cgattttgaa tggctccccg 2700
ctatctcgag cagatttacg cgccgtcaac attaacctct gcgacatgtg tgtcataatc 2760
tctgcccggg tcccaaatac cgaagatacc actttggcag ataaggaagc gattctggca 2820
tcattgaaca ttaaagctat gcaatttgat gatacccttg gctttttccc gatgaggcat 2880
caaactggcg atcggagccc gctgggctcc ccgatttcca tgcagaagaa gggcgccaag 2940
tttgggacca atgtgccgat gattacggag cttgttaacg actcaaatgt gcaattcctc 3000
gaccaggacg acgacgacga tccggacacc gagctctacc tcacgcagcc cttcgcctgc 3060
ggaactgcct tcgccatttc agtgttggat tctttgatga gtactaccta cttcaacgac 3120
tcggccctga ccctcatccg aactctggtc actggtggag ccacaccgga gcttgaactt 3180
attctggcag aaggcgccgg gctccgaggt ggctacagta ccccggagac cctctcaaac 3240
agagatcggt gccgaattgc tcagatttcg ctgcaggata acccgtacga tggagttgtg 3300
cacaacacta cctatggtgc aatgttcacg atagctctga ggaggtacgg ccaactttgt 3360
atcgggctgt ataggcttca cgaccaggat aatccggact ctatgaagcg atatgttatc 3420
actaatcctc cagcagaatt gaggattaaa aatactgatt acgtctacgt acttgaacaa 3480
ttcgatccgg gactggagta cgagccgggc aaacgacact tt 3522
<210> 19
<211> 4491
<212> DNA
<213> 人工序列(Artificial Sequence)
<220>
<221> misc_feature
<222> (1)..(4491)
<223> pPD95.75表达载体
<400> 19
aagcttgcat gcctgcaggt cgactctaga ggatccccgg gattggccaa aggacccaaa 60
ggtatgtttc gaatgatact aacataacat agaacatttt caggaggacc cttgcttgga 120
gggtaccggt agaaaaaatg agtaaaggag aagaactttt cactggagtt gtcccaattc 180
ttgttgaatt agatggtgat gttaatgggc acaaattttc tgtcagtgga gagggtgaag 240
gtgatgcaac atacggaaaa cttaccctta aatttatttg cactactgga aaactacctg 300
ttccatgggt aagtttaaac atatatatac taactaaccc tgattattta aattttcagc 360
caacacttgt cactactttc tgttatggtg ttcaatgctt ctcgagatac ccagatcata 420
tgaaacggca tgactttttc aagagtgcca tgcccgaagg ttatgtacag gaaagaacta 480
tatttttcaa agatgacggg aactacaaga cacgtaagtt taaacagttc ggtactaact 540
aaccatacat atttaaattt tcaggtgctg aagtcaagtt tgaaggtgat acccttgtta 600
atagaatcga gttaaaaggt attgatttta aagaagatgg aaacattctt ggacacaaat 660
tggaatacaa ctataactca cacaatgtat acatcatggc agacaaacaa aagaatggaa 720
tcaaagttgt aagtttaaac atgattttac taactaacta atctgattta aattttcaga 780
acttcaaaat tagacacaac attgaagatg gaagcgttca actagcagac cattatcaac 840
aaaatactcc aattggcgat ggccctgtcc ttttaccaga caaccattac ctgtccacac 900
aatctgccct ttcgaaagat cccaacgaaa agagagacca catggtcctt cttgagtttg 960
taacagctgc tgggattaca catggcatgg atgaactata caaatagcat tcgtagaatt 1020
ccaactgagc gccggtcgct accattacca acttgtctgg tgtcaaaaat aataggggcc 1080
gctgtcatca gagtaagttt aaactgagtt ctactaacta acgagtaata tttaaatttt 1140
cagcatctcg cgcccgtgcc tctgacttct aagtccaatt actcttcaac atccctacat 1200
gctctttctc cctgtgctcc caccccctat ttttgttatt atcaaaaaac ttctcttaat 1260
ttctttgttt tttagcttct tttaagtcac ctctaacaat gaaattgtgt agattcaaaa 1320
atagaattaa ttcgtaataa aaagtcgaaa aaaattgtgc tccctccccc cattaataat 1380
aattctatcc caaaatctac acaatgttct gtgtacactt cttatgtttt ttacttctga 1440
taaatttttt tgaaacatca tagaaaaaac cgcacacaaa ataccttatc atatgttacg 1500
tttcagttta tgaccgcaat ttttatttct tcgcacgtct gggcctctca tgacgtcaaa 1560
tcatgctcat cgtgaaaaag ttttggagta tttttggaat ttttcaatca agtgaaagtt 1620
tatgaaatta attttcctgc ttttgctttt tggggtttcc cctattgttt gtcaagattt 1680
cgaggacggc gtttttcttg ctaaaatcac aagtattgat gagcacgatg caagaaagat 1740
cggaagaagg tttgggtttg aggctcagtg gaaggtgagt agaagttgat aatttgaaag 1800
tggagtagtg tctatggggt ttttgcctta aatgacagaa tacattccca atataccaaa 1860
cataactgtt tcctactagt cggccgtacg ggccctttcg tctcgcgcgt ttcggtgatg 1920
acggtgaaaa cctctgacac atgcagctcc cggagacggt cacagcttgt ctgtaagcgg 1980
atgccgggag cagacaagcc cgtcagggcg cgtcagcggg tgttggcggg tgtcggggct 2040
ggcttaacta tgcggcatca gagcagattg tactgagagt gcaccatatg cggtgtgaaa 2100
taccgcacag atgcgtaagg agaaaatacc gcatcaggcg gccttaaggg cctcgtgata 2160
cgcctatttt tataggttaa tgtcatgata ataatggttt cttagacgtc aggtggcact 2220
tttcggggaa atgtgcgcgg aacccctatt tgtttatttt tctaaataca ttcaaatatg 2280
tatccgctca tgagacaata accctgataa atgcttcaat aatattgaaa aaggaagagt 2340
atgagtattc aacatttccg tgtcgccctt attccctttt ttgcggcatt ttgccttcct 2400
gtttttgctc acccagaaac gctggtgaaa gtaaaagatg ctgaagatca gttgggtgca 2460
cgagtgggtt acatcgaact ggatctcaac agcggtaaga tccttgagag ttttcgcccc 2520
gaagaacgtt ttccaatgat gagcactttt aaagttctgc tatgtggcgc ggtattatcc 2580
cgtattgacg ccgggcaaga gcaactcggt cgccgcatac actattctca gaatgacttg 2640
gttgagtact caccagtcac agaaaagcat cttacggatg gcatgacagt aagagaatta 2700
tgcagtgctg ccataaccat gagtgataac actgcggcca acttacttct gacaacgatc 2760
ggaggaccga aggagctaac cgcttttttg cacaacatgg gggatcatgt aactcgcctt 2820
gatcgttggg aaccggagct gaatgaagcc ataccaaacg acgagcgtga caccacgatg 2880
cctgtagcaa tggcaacaac gttgcgcaaa ctattaactg gcgaactact tactctagct 2940
tcccggcaac aattaataga ctggatggag gcggataaag ttgcaggacc acttctgcgc 3000
tcggcccttc cggctggctg gtttattgct gataaatctg gagccggtga gcgtgggtct 3060
cgcggtatca ttgcagcact ggggccagat ggtaagccct cccgtatcgt agttatctac 3120
acgacgggga gtcaggcaac tatggatgaa cgaaatagac agatcgctga gataggtgcc 3180
tcactgatta agcattggta actgtcagac caagtttact catatatact ttagattgat 3240
ttaaaacttc atttttaatt taaaaggatc taggtgaaga tcctttttga taatctcatg 3300
accaaaatcc cttaacgtga gttttcgttc cactgagcgt cagaccccgt agaaaagatc 3360
aaaggatctt cttgagatcc tttttttctg cgcgtaatct gctgcttgca aacaaaaaaa 3420
ccaccgctac cagcggtggt ttgtttgccg gatcaagagc taccaactct ttttccgaag 3480
gtaactggct tcagcagagc gcagatacca aatactgtcc ttctagtgta gccgtagtta 3540
ggccaccact tcaagaactc tgtagcaccg cctacatacc tcgctctgct aatcctgtta 3600
ccagtggctg ctgccagtgg cgataagtcg tgtcttaccg ggttggactc aagacgatag 3660
ttaccggata aggcgcagcg gtcgggctga acggggggtt cgtgcacaca gcccagcttg 3720
gagcgaacga cctacaccga actgagatac ctacagcgtg agcattgaga aagcgccacg 3780
cttcccgaag ggagaaaggc ggacaggtat ccggtaagcg gcagggtcgg aacaggagag 3840
cgcacgaggg agcttccagg gggaaacgcc tggtatcttt atagtcctgt cgggtttcgc 3900
cacctctgac ttgagcgtcg atttttgtga tgctcgtcag gggggcggag cctatggaaa 3960
aacgccagca acgcggcctt tttacggttc ctggcctttt gctggccttt tgctcacatg 4020
ttctttcctg cgttatcccc tgattctgtg gataaccgta ttaccgcctt tgagtgagct 4080
gataccgctc gccgcagccg aacgaccgag cgcagcgagt cagtgagcga ggaagcggaa 4140
gagcgcccaa tacgcaaacc gcctctcccc gcgcgttggc cgattcatta atgcagctgg 4200
cacgacaggt ttcccgactg gaaagcgggc agtgagcgca acgcaattaa tgtgagttag 4260
ctcactcatt aggcacccca ggctttacac tttatgcttc cggctcgtat gttgtgtgga 4320
attgtgagcg gataacaatt tcacacagga aacagctatg accatgatta cgccaagctg 4380
taagtttaaa catgatctta ctaactaact attctcattt aaattttcag agcttaaaaa 4440
tggctgaaat cactcacaac gatggatacg ctaacaactt ggaaatgaaa t 4491
<210> 20
<211> 33
<212> DNA
<213> 引物(Primer)
<400> 20
gcatgcctgc aggccgaagg aattatcaca gca 33
<210> 21
<211> 34
<212> DNA
<213> 引物(Primer)
<400> 21
ccccccggga tcttcaaaat tgaaactttg actg 34
<210> 22
<211> 40
<212> DNA
<213> 引物(Primer)
<400> 22
ttcaattttg aagatcccgg gatgtctcaa gcttcacaag 40
<210> 23
<211> 27
<212> DNA
<213> 引物(Primer)
<400> 23
cagcttgcga aatcgttgat cagtacg 27
<210> 24
<211> 27
<212> DNA
<213> 引物(Primer)
<400> 24
aacgatttcg caagctgcag aatttcc 27
<210> 25
<211> 29
<212> DNA
<213> 引物(Primer)
<400> 25
gcagcagagt agaatctcca ggccgactg 29
<210> 26
<211> 30
<212> DNA
<213> 引物(Primer)
<400> 26
gagattctac tctgctgcac cggaatccac 30
<210> 27
<211> 44
<212> DNA
<213> 引物(Primer)
<400> 27
ctcatttttt ctaccggtac cctacaactt gaagtaccgg gacc 44
<210> 28
<211> 29
<212> DNA
<213> 引物(Primer)
<400> 28
gctccggagc ctgtgcttcc tgcagattc 29
<210> 29
<211> 29
<212> DNA
<213> 引物(Primer)
<400> 29
gaatctgcag gaagcacagg ctccggagc 29
<210> 30
<211> 32
<212> DNA
<213> 引物(Primer)
<400> 30
catgcatgca cagcgttctc acctaaaagg cc 32
<210> 31
<211> 28
<212> DNA
<213> 引物(Primer)
<400> 31
cgggatcccg ggtcgtaatg accgcctg 28
<210> 32
<211> 40
<212> DNA
<213> 引物(Primer)
<400> 32
caggtcgact ctagaggatc catgcgattc ggtggtcaac 40
<210> 33
<211> 34
<212> DNA
<213> 引物(Primer)
<400> 33
cttgtctctg ggttttatta tttggcatca tcac 34
<210> 34
<211> 28
<212> DNA
<213> 引物(Primer)
<400> 34
aataaaaccc agagacaagt gtggcttc 28
<210> 35
<211> 43
<212> DNA
<213> 引物(Primer)
<400> 35
ctcatttttt ctaccggtac cctgacatca gttaacattt tgg 43
<210> 36
<211> 28
<212> DNA
<213> 引物(Primer)
<400> 36
cctccgtgtc atccagttgg taagagtc 28
<210> 37
<211> 28
<212> DNA
<213> 引物(Primer)
<400> 37
gactcttacc aactggatga cacggagg 28
<210> 38
<211> 31
<212> DNA
<213> 引物(Primer)
<400> 38
cgggatccag cgcagacagc tagagtgtct g 31
<210> 39
<211> 28
<212> DNA
<213> 引物(Primer)
<400> 39
cgggatcccg tgtcgacgaa gggtccac 28
<210> 40
<211> 38
<212> DNA
<213> 引物(Primer)
<400> 40
caggtcgact ctagaggatc catggcaaat ggtggcgg 38
<210> 41
<211> 30
<212> DNA
<213> 引物(Primer)
<400> 41
gacgttttga atgacctcat ggagaagagg 30
<210> 42
<211> 29
<212> DNA
<213> 引物(Primer)
<400> 42
ggtcattcaa aacgtcacaa accacaccg 29
<210> 43
<211> 41
<212> DNA
<213> 引物(Primer)
<400> 43
attctacgaa tgaccggtac cctaaaagtg tcgtttgccc g 41
<210> 44
<211> 29
<212> DNA
<213> 引物(Primer)
<400> 44
gcacaaggac cggggtgacg tcaatgtgg 29
<210> 45
<211> 29
<212> DNA
<213> 引物(Primer)
<400> 45
ccacattgac gtcaccccgg tccttgtgc 29
<210> 46
<211> 20
<212> DNA
<213> 引物(Primer)
<400> 46
aatggttgat ttcgtttcgc 20
<210> 47
<211> 20
<212> DNA
<213> 引物(Primer)
<400> 47
tgcagtcgct tttcactttg 20
<210> 48
<211> 23
<212> DNA
<213> 引物(Primer)
<400> 48
gatcccttcc aagaacaaga gtt 23
<210> 49
<211> 23
<212> DNA
<213> 引物(Primer)
<400> 49
atcgtctttt tctccatttc ctc 23
Claims (23)
1.钾离子通道蛋白的融合蛋白，其特征在于，其具有N端区、跨膜区以及C端区，包括：
(a)线虫KQT-1的N端区、人KCNQ2的S1-S6跨膜区和线虫KQT-1的C端区；或
(b)线虫SHK-1的N端区、人KV1.2的S1-S6跨膜区和C端区；或
(c)人BK的N端区和跨膜区S0-S6、线虫SLO-1的C端区。
2.如权利要求1所述的融合蛋白，其特征在于，(a)中：
所述线虫KQT-1的N端区包含其N端第1～145位氨基酸残基中长120～145个氨基酸残基的序列；较佳地为包含N端第1～130位氨基酸残基或由第1～130位氨基酸残基组成的序列；
所述人KCNQ2的S1-S6跨膜区包含自其第66～75氨基酸残基中的任一位起至第347～360位氨基酸中的任一位氨基酸残基的序列；较佳地为包含第75～347位氨基酸残基或由第75～347位氨基酸残基组成的序列；
所述线虫KQT-1的C端区包含其C端第379～739位氨基酸残基中长330～361个氨基酸残基的序列；较佳地为其C端第379～406氨基酸残基中的任一位起至第730～739位氨基酸中的任一位氨基酸残基的序列；更佳地为至少包含C端第405～739位氨基酸残基或由第405～739位氨基酸残基组成的序列。
3.如权利要求2所述的融合蛋白，其特征在于，所述融合蛋白为线虫KQT-1第1～130位氨基酸残基、人KCNQ2第75～347位氨基酸残基与KQT-1蛋白C端第405～739氨基酸残基组成的融合蛋白。
4.如权利要求1～3任一所述的融合蛋白，其特征在于，(a)中所述的跨膜区序列中存在通道功能获得性突变，其中所述突变包括：在人KCNQ2蛋白第201位上的R201C突变。
5.如权利要求4所述的融合蛋白，其特征在于，所述的融合蛋白的氨基酸序列为SEQ IDNO:3所示；其cDNA序列为SEQ ID NO:5所示；或，含有功能获得性突变的融合蛋白的氨基酸序列为SEQ ID NO:4所示，其cDNA序列为SEQ ID NO:6所示。
6.如权利要求1所述的融合蛋白，其特征在于，(b)中：
所述线虫SHK-1的N端区包含其N端第1～260位氨基酸残基之间，长240～260个氨基酸残基的序列，较佳地为包含第1～245位氨基酸残基，或由第1～245位氨基酸残基组成的序列；
所述人KV1.2的S1-S6跨膜区和C端区包含其第135～449位氨基酸残基之间，长为295～315个氨基酸残基的片段；较佳地为其第135～150氨基酸残基中的任一位起到第435～449位氨基酸中的任一位氨基酸残基的序列；更佳地为至少包含其第146～435位氨基酸残基或由其第146～435位氨基酸残基组成。
7.如权利要求6所述的融合蛋白，其特征在于，所述融合蛋白为线虫SHK-1的N端区第1～245位氨基酸残基，人KV1.2第146～449位氨基酸残基组成的融合蛋白。
8.如权利要求1、6～7任一所述的融合蛋白，其特征在于，(b)中所述的跨膜区序列中存在通道功能获得性突变，其中所述突变包括：在人KV1.2蛋白第297位上的R297Q突变。
9.如权利要求8所述的融合蛋白，其特征在于，所述融合蛋白的氨基酸序列为SEQ IDNO:9；其cDNA序列为SEQ ID NO:11所示；或，含有功能获得性突变的融合蛋白的氨基酸序列为SEQ ID NO:10所示；其cDNA序列为SEQ ID NO:12所示。
10.如权利要求1所述的融合蛋白，其特征在于，(c)中：
所述人BK的N端区和跨膜区S0-S6包含其第1～604位氨基酸残基之间，长570～604个氨基酸残基的片段；较佳地为其第1～20位氨基酸残基中的任一位起到第581～604位氨基酸中的任一位氨基酸残基的序列；更佳地为至少包含其第1～581位氨基酸残基或由其第1～581位氨基酸残基组成；
所述线虫SLO-1的C端区包含其第488～1140位氨基酸残基之间，长595～653个氨基酸残基的片段；较佳地为其第488～546氨基酸残基中的任一位起到第1120～1140位氨基酸中的任一位氨基酸残基的序列；更佳地为至少包含其第546～1140位氨基酸残基或由其第546～1140位氨基酸残基组成。
11.如权利要求10所述的融合蛋白，其特征在于，所述融合蛋白为人BK的N端区和跨膜区S0-S6第1～581位氨基酸残基，线虫SLO-1的C端区第546～1140位氨基酸残基组成的融合蛋白。
12.如权利要求1、10～11任一所述的融合蛋白，其特征在于，所述的跨膜区序列中存在通道功能获得性突变，其中所述突变包括：在人BK蛋白第434位上的D434G突变。
13.如权利要求12所述的融合蛋白，其特征在于，(c)中所述融合蛋白的氨基酸序列为SEQ ID NO:15所示，其cDNA序列为SEQ ID NO:17所示；或，含有功能获得性突变的融合蛋白的氨基酸序列为SEQ ID NO:16所示，其cDNA序列为SEQ ID NO:18所示。
14.如权利要求1所述的融合蛋白，其特征在于，其还含有可被检测的蛋白质或酶。
15.分离的多核苷酸序列，其特征在于，所述多核苷酸序列选自：
(a)编码权利要求1～12中任一项所述的融合蛋白的多核苷酸序列；和
(b)与(a)的多核苷酸序列互补的序列。
16.一种表达构建物或表达载体，其包含权利要求15所述的多核苷酸序列。
17.基因工程细胞或转基因线虫，其包含权利要求15所述的多核苷酸序列或转入了权利要求16所述的表达载体，优选地，所述转基因线虫具有以下一项或多项特征：
(i)所述多核苷酸序列整合入所述转基因线虫的基因组中；
(ii)所述转基因线虫表达权利要求4、8或12所述的融合蛋白；和
(iii)所述转基因线虫为acs-20基因缺陷型线虫。
18.如权利要求17所述的基因工程细胞或转基因线虫，其特征在于，所述的基因工程细胞或转基因线虫中，ifd-2和/或c15c7.5基因或其编码的蛋白被下调；较佳地，所述ifd-2和/或c15c7.5基因或其编码的蛋白藉由RNAi干扰、CRISPR/Cas9基因编辑、基因敲除、抗体靶向结合的方式被下调。
19.权利要求1～12中任一项所述的融合蛋白、权利要求15所述的多核苷酸、权利要求16所述的表达构建物或表达载体或权利要求17或18所述的基因工程细胞或转基因线虫的用途，用于筛选人KCNQ2、KV1.2或BK通道的抑制剂或激活剂，或用于制备筛选人KCNQ2、KV1.2或BK通道的抑制剂或激活剂的筛选模型。
20.一种筛选人钾离子通道抑制剂和激活剂的方法，其中所述钾离子通道为KCNQ2、KV1.2或BK通道，该方法包括：
所述筛选人KCNQ2通道抑制剂和激活剂的方法包括：使待筛选的化合物与表达权利要求1中(a)所述的融合蛋白的转基因线虫共同孵育，并观察、比较共同孵育前和共同孵育后线虫的行为表现；与共同孵育前相比，共同孵育后使共同孵育前的线虫的行为缺陷得以改善或恢复的化合物鉴别为KCNQ2通道的抑制剂；共同孵育后使共同孵育前的线虫的行为缺陷加重的化合物鉴别为KCNQ2通道的激活剂；或
所述筛选人KV1.2通道抑制剂和激活剂的方法包括：使待筛选的化合物与表达权利要求1中(2)所述的融合蛋白的转基因线虫共同孵育，并观察、比较共同孵育前和共同孵育后线虫的行为表现；与共同孵育前相比，共同孵育后使共同孵育前的线虫的行为缺陷得以改善或恢复的化合物鉴别为KV1.2通道的抑制剂；共同孵育后使共同孵育前的线虫的行为缺陷加重的化合物鉴别为KV1.2通道的激活剂。
所述筛选人BK通道抑制剂和激活剂的方法包括：使待筛选的化合物与表达权利要求1中(3)所述的融合蛋白的转基因线虫共同孵育，并观察、比较共同孵育前和共同孵育后线虫的行为表现；与共同孵育前相比，共同孵育后使共同孵育前的线虫的行为缺陷得以改善或恢复的化合物鉴别为BK通道的抑制剂；共同孵育后使共同孵育前的线虫的行为缺陷加重的化合物鉴别为BK通道的激活剂；
优选的，上述方法使用权利要求17或18所述的转基因线虫进行筛选。
21.如权利要求20所述的方法，其特征在于，所述的转基因线虫中，ifd-2和/或c15c7.5基因被下调；较佳地，藉由RNAi干扰、CRISPR/Cas9基因编辑、基因敲除、抗体靶向结合的方式下调ifd-2和/或c15c7.5基因。
22.如权利要求20或21所述的方法，其特征在于，在含诱导剂和抗生素的线虫培养基中接种所述靶向ifd-2和c15c7.5基因的RNAi菌，培养后将待筛选的化合物加入，之后接种所述的转基因线虫，优选为L3-L4时期的线虫，观察转基因线虫行为的变化。
23.用于筛选人KCNQ2、KV1.2或BK通道的抑制剂或激活剂的试剂盒，其中含有：权利要求1～12中任一项所述的融合蛋白、权利要求15所述的多核苷酸、权利要求16所述的表达构建物或表达载体或权利要求17或18所述的基因工程细胞或转基因线虫。
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