一种提高植物对双生病毒抗性的方法与流程

本发明涉及分子生物学领域，特别涉及一种改造的抗双生病毒的vtale分子，尤其涉及一种对双生病毒科病毒具有广谱抗性且有效稳定的vtale分子。本发明还涉及所述vtale分子的应用以及利用所述vtale分子培育具有稳定的广谱双生病毒抗性的植物的方法。
背景技术：
：双生病毒(geminivirus)为孪生颗粒形态的单链dna病毒，病毒颗粒大小约为20×30nm。依据病毒的基因组结构、寄主范围、传播介体以及基因组序列的相关性将双生病毒分为七个属，即菜豆金色花叶病毒属(begomovirus)、玉米线条病毒属(mastrevirus)、甜菜曲顶病毒属(curtovirus)、番茄伪曲顶病毒属(topocuvirus)、伊朗甜菜曲顶病毒属(becurtovirus)、芜菁曲顶病毒属(turncurtovirus)和弯叶画眉草条纹病毒属(eragrovirus)。双生病毒高度依赖于寄主植物细胞的dna复制和rna转录系统来完成自身的复制和转录过程。在最近20年间，双生病毒引起的病害由局部发生的小病害衍变成目前全球性的最重要的植物病毒病害之一，危害玉米、小麦、棉花、木薯、番茄等重要农作物和经济作物，仅对于木薯，每年在全球造成的经济损失就达数十亿美元之巨。在我国，随着入侵烟粉虱的不断蔓延，导致由其传播的双生病毒病害——番茄黄化曲叶病在18个省市多个番茄产区发生，成为我国番茄生产上最重要的病害之一。2009年经济损失超过50亿元，2010--2012年危害进一步扩大，病害发生面积超过100万亩。因此，对双生病毒病害的有效防治已经成为亟待解决的实际生产问题。目前，在实际生产中仍普遍采用化学防治的方法来直接或者间接地防控双生病毒病害，但效果并不理想，同时化学防治对环境及食品安全也构成了严重的威胁。随着分子生物学的不断发展，科学家们对植物抗双生病毒的分子机制有了进一步的解析，研究较清楚的机制主要有两类：抗性较专一的抗性基因(rgene)介导的抗性和抗性谱较宽的rna沉默(rnasilencing)介导的抗性。利用抗性基因选育抗性品种来防控双生病毒的方法具有育种周期长、抗病谱较窄的弱点，而双生病毒的突变率高、病毒同入侵型介体昆虫形成互惠共生的关系等原因导致该种方法未能快速、有效地发挥防控作用。而利用rna沉默机制来防控双生病毒的方法由于存在抗性不稳定且易丢失，此外双生病毒同其他病毒一样也进化出了有效的抑制植物rna沉默的基因沉默抑制子(suppressorofrnasilencing)，使得该方法也未能在实际生产中得到广泛应用。最近发展出来一种利用crispr/cas技术使植物获得抗双生病毒的新方法。此系统的工作原理是crrna(crispr-derivedrna)通过碱基配对与tracrrna(trans-activatingrna)结合形成tracrrna/crrna复合物，此复合物引导核酸酶cas9蛋白在与crrna配对的序列靶位点剪切双链dna。而通过人工设计这两种rna，可以改造形成具有引导作用的sgrna(shortguiderna)，足以引导cas9对dna的定点切割。但是研究表明通过该方法植物获得的对双生病毒的抗性并不稳定且常有脱靶的现象。综上所述，利用分子生物学技术发展一种有效并且抗性稳定的抗双生病毒的新方法对于农作物及经济作物的高产和稳产以及环境和食品的安全都是十分必要的。技术实现要素：本发明的目的是提供一种对双生病毒科病毒具有广谱抗性且有效稳定的vtale分子。本发明提供的对双生病毒科病毒具有广谱抗性且有效稳定的vtale分子是一种改造的抗双生病毒的vtale分子，其以植物病原细菌水稻黄单胞杆菌(xanthomonasoryzaepv.oryzae，缩写为xoo)三型分泌系统效应因子avrxa27基因序列为骨架序列，保留avrxa27基因的n端和c端序列，然后对avrxa27基因的dna识别/结合结构域序列和功能输出结构域序列进行改造而获得。其中改造的部分涉及dna识别/结合结构域序列和功能输出结构域序列两部分(下文将这两部分分别称为“改造后的dna识别/结合结构域序列”和“改造后的功能输出结构域序列”)。vtale分子的dna识别/结合结构域经人工设计能够特异性地识别并结合任意指定的dna序列，在功能输出结构域的作用下从而可以实现对靶基因的调控与修饰。通过对双生病毒基因组特定区域进行人工设计的表观遗传学修饰从而干扰双生病毒的正常复制及转录过程，达到提高植物对双生病毒的免疫能力的目的。本发明分别针对双生病毒科多株双生病毒高度保守的ir(intergeneticregion)区域设计了vtale靶点ir-vtale及针对菜豆金色花叶病毒属病毒甘蓝曲叶病毒(cabbageleafcurlvirus，calcuv)复制相关蛋白结合位点(replicationassociatedproteinbinding，repbinding)设计了特异性的rb-vtale靶点为vtale的dna识别/结合结构域的靶点序列，功能输出结构域融合表达表观遗传学修饰相关因子。在第一方面，本发明提供一种改造的抗双生病毒的vtale分子，其以植物病原细菌水稻黄单胞杆菌(xanthomonasoryzaepv.oryzae)三型分泌系统效应因子avrxa27基因序列为骨架序列，保留avrxa27基因的n端和c端序列，然后对avrxa27基因的dna识别/结合结构域序列和功能输出结构域序列进行改造而获得，其中所述改造后的dna识别/结合结构域序列靶向seqidno：1所示的ir-vtale靶点序列或seqidno：2所示的rb-vtale靶点序列，并且所述改造后的功能输出结构域序列选自由编码seqidnos：8-12所示的氨基酸序列的核苷酸序列组成的组中的任一种。这两部分序列可以通过特异的限制性内切酶处理后连接。本领域技术人员能够选择这两个序列中合适的特异性限制性内切酶酶切位点经限制性内切酶处理后将二者连接在一起。优选地，所述改造后的dna识别/结合结构域序列靶向ir-vtale靶点序列，其序列为5’-taatattaccggatggcc-3’(seqidno：1)。优选地，所述改造后的dna识别/结合结构域序列靶向rb-vtale靶点序列，其序列为5’-tatatattggacaccaag-3’(seqidno：2)。优选地，所述改造后的功能输出结构域序列为编码atkyp(其氨基酸序列为seqidno：8)的核苷酸序列。优选地，所述改造后的功能输出结构域序列为编码athdac19(其氨基酸序列为seqidno：9)的核苷酸序列。优选地，所述改造后的功能输出结构域序列为编码atmet1(其氨基酸序列为seqidno：10)的核苷酸序列。优选地，所述改造后的功能输出结构域序列为编码atsrdx(其氨基酸序列为seqidno：11)的核苷酸序列。优选地，所述改造后的功能输出结构域序列为编码atsuvh2(其氨基酸序列为seqidno：12)的核苷酸序列。优选地，所述改造后的功能输出结构域序列是选自由seqidnos：3-7所示的核苷酸序列组成的组的任意一种核苷酸序列。根据本发明第一方面构建的抗双生病毒的vtale分子包括，但不限于，seqidnos：17-19和26-30。其中，优选的vtale分子的序列为ir-vtale-atkyp(seqidno：17)和rb-vtale-atkyp(seqidno：26)。其中ir-vtale-atkyp(seqidno：17)包括来源于xoo的avrxa27基因n端序列、靶向ir-vtale靶点序列的序列、来源于xoo的avrxa27基因c端序列和atkyp氨基酸序列(见图13)；rb-vtale-atkyp(seqidno：26)包括来源于xoo的avrxa27基因n端序列、靶向rb-vtale靶点序列的序列、来源于xoo的avrxa27基因c端序列和atkyp氨基酸序列(见图22)。本领域技术人员知晓，现有技术中已有的一般tale分子主要由两部分组成：dna识别/结合结构域(recognition/bindingdomain)和激活结构域(activationdomain，ad)(图1a)。其中dna识别/结合结构域是由一段串联排列的重复序列组成，能够特异性识别并结合靶基因序列，每个重复序列由长度为33-35个氨基酸组成，除了第12和13位的氨基酸可变外，其他氨基酸序列高度保守，而重复序列可变的双氨基酸残基(repeatvariabledi-residue，rvd)是tale实现靶向识别特异dna碱基的关键位点，不同的rvd能够相对特异地识别a、t、c、g四种碱基中的一种或多种，其中分别与这4种碱基相对应的最常见的4种rvd分别是ni＝a，ng＝t，hd＝c和nn＝g或a。激活结构域能够在tale特异性识别并结合靶基因后激活靶基因的表达。本发明设计的vtale分子结构示意图显示在图1b中。本发明设计的vtale分子的dna识别/结合结构域所结合的靶点序列选自下述中的任一种：针对双生病毒科多株病毒高度保守的ir(intergeneticregion)区域序列设计的广谱抗双生病毒科病毒的intergeneticregion-vtale(ir-vtale)靶点和针对菜豆金色花叶病毒属病毒甘蓝曲叶病毒(cabbageleafcurlvirus，calcuv)复制相关蛋白(replicationassociatedprotein，rep)结合位点设计的vtale靶点repbinding-vtale(rb-vtale)。此外，由于tale蛋白所识别的靶点dna序列必须以t开始，所以，虽然两个靶点序列都只含有16.5个串联排列的重复序列，但是都可以特异识别并结合18bp的靶点dna序列。根据本发明人设计的靶点dna序列(即，ir-vtale靶点序列或rb-vtale靶点序列)，本领域技术人员能够设计相应的氨基酸重复序列进行串联，构成改造的dna识别/结合结构域，这是在本领域技术人员的技能范围之内的。本发明人的创造性劳动体现在对avrxa27基因的dna识别/结合结构域序列所靶向的靶点序列的选择。在第二方面，本发明提供本发明改造的抗双生病毒的vtale分子在培育具有稳定的广谱双生病毒抗性的植物的方法中的应用。在第三方面，本发明提供一种用于培育具有稳定的广谱双生病毒抗性的植物的方法，所述方法包括用第一方面的改造的抗双生病毒的vtale分子转化待培育的植物，使vtale在所述植物中稳定表达。也就是说，使用第一方面的改造的抗双生病毒的vtale分子对植物进行转基因处理，获得具有稳定的广谱双生病毒抗性的植物。在一个优选的实施方案中，本发明提供一种用于培育具有稳定的广谱双生病毒抗性的植物的方法，所述方法包括下述步骤：(1)vtale分子的构建：所述vtale分子以植物病原细菌水稻黄单胞杆菌(xanthomonasoryzaepv.oryzae)三型分泌系统效应因子avrxa27基因序列为骨架序列，保留avrxa27基因的n端和c端序列，然后对avrxa27基因的dna识别/结合结构域序列和功能输出结构域序列进行改造而获得，其中所述改造后的dna识别/结合结构域序列靶向seqidno：1所示的ir-vtale靶点序列或seqidno：2所示的rb-vtale靶点序列；并且所述改造后的功能输出结构域序列选自由编码seqidnos：8-12所示的氨基酸序列的核苷酸序列组成的组中的任一种；(2)植物转化：将步骤(1)构建的vtale分子，转化到所述植物中进行稳定表达。优选地，所述改造后的dna识别/结合结构域序列靶向seqidno：1所示的ir-vtale靶点序列或seqidno：2所示的rb-vtale靶点序列。具体地，在初步验证实验中，本发明人发现，在本氏烟草中瞬时表达步骤(1)构建的vtale分子的优选组合ir-vtale-atkyp分子能够使本氏烟草获得对双生病毒科病毒的广谱抗性。将含有ir-vtale-atkyp分子的农杆菌注射本氏烟草24h后，再接种双生病毒科不同属病毒的侵染性克隆，包括玉米线条病毒属(mastrevirus)小麦矮缩病毒(wheatdwarfvirus，wdv，中国农业科学院植物保护研究所王锡峰课题组惠赠)；甜菜曲顶病毒属(curtovirus)甜菜曲顶病毒(beetcurlytopvirus，bctv，本实验室保藏)；菜豆金色花叶病毒属(begomovirus)单组分病毒番茄黄化曲叶病毒(tomatoyellowleafcurlvirus，tylcv-sh2，中国农业科学院植物保护研究所周雪平课题组惠赠)和双组分病毒印度木薯花叶病毒(indiancassavamosaicvirus，icmv-sg，本实验室分离鉴定并保存，gangwangetal.，2014，virusgenes)。本氏烟草的发病症状和病毒滴度两方面数据均表明瞬时表达ir-vtale-atkyp分子对双生病毒科病毒具有广谱抗性。在小麦中瞬时表达步骤(1)构建的vtale分子的优选组合ir-vtale-atkyp分子能够使小麦获得对小麦矮缩病毒的抗性。将分别含有ir-vtale-mcs，ir-vtale-atkyp分子和wdv的农杆菌活化后，使其浓度达到od600＝2.0，按vtale分子和wdv体积比3∶1混合，穿刺接种小麦jn17(中国科学院遗传与发育生物学研究所李振生课题组惠赠)的胚及其周围位置，然后将接种的小麦种子种在土中，观察小麦的发病情况，发病症状和病毒滴度两方面结果均表明在小麦中瞬时表达ir-vtale-atkyp分子对wdv具有抗性。在本氏烟中分别瞬时表达步骤(1)构建的vtale分子和crispr/cas9分子，接种甘蓝曲叶病毒后发现vtale分子的优选组合rb-vtale分子与crispr/cas9分子相比能够使本氏烟草获得更好的抗性效果。将分别含有vtale和crispr/cas9分子的农杆菌注射本氏烟24h后，再接种甘蓝曲叶病毒的侵染性克隆，本氏烟的发病症状和病毒滴度两方面数据表明rb-vtale分子具有比crispr/cas9分子更好的抗calcuv的能力。本发明人发现，在所选的植物物种中瞬时表达步骤(1)构建的vtale分子能够赋予植物对双生病毒科病毒的广谱抗性。进一步地，本发明人验证了在植物中稳定表达步骤(1)构建的rb-vtale分子的效果。具体地，rb-vtale-atkyp分子转基因拟南芥植物具有良好的抗甘蓝曲叶病毒的效果。依照文献(xiurenzhangetal.，2006，natureprotocol)报道的拟南芥转化方法，将含有步骤(1)构建的vtale分子的优选组合rb-vtale分子的农杆菌菌株分别转化columbia生态型拟南芥野生型植物col-0并收获t1代种子，用抗生素筛选t1代转基因种子从而成功获得转化rb-vtale分子的t1代转基因植物株系。然后对转基因植物接种calcuv侵染性克隆，发病症状、病情指数和病毒滴度三方面数据表明rb-vtale-atkyp的转基因植物对甘蓝曲叶病毒具有很好的抗性。rb-vtale-atkyp转基因拟南芥植物对甘蓝曲叶病毒(calcuv)的抗性效果可以稳定遗传给t3代，且这种抗性效果与atkyp基因的表达水平呈正相关。对t3代rb-vtale-atkyp转基因拟南芥植物注射接种calcuv侵染性克隆，从发病症状、病情指数和病毒滴度三方面数据说明rb-vtale-atkyp的转基因植物对甘蓝曲叶病毒的抗性可以稳定遗传给下一代；对atkyp基因的表达谱分析的结果表明rb-vtale-atkyp的转基因植物对甘蓝曲叶病毒的抗性与atkyp基因的表达呈正相关。t3代rb-vtale-atkyp转基因拟南芥植物中接种的甘蓝曲叶病毒基因组rb-vtale靶点所在的核小体组蛋白和dna均具有高水平的甲基化修饰，此外病毒相关基因的表达受到明显抑制。利用染色质免疫沉淀(chromatinimmuno-precipitation，chip)技术检测t3代rb-vtale-atkyp转基因拟南芥植物中接种的甘蓝曲叶病毒基因组rb-vtale靶点所在的核小体组蛋白的甲基化水平，发现组蛋白h3第9位赖氨酸甲基化水平明显提高；利用基于对甲基化敏感的限制性内切酶酶切消化的方法检测rb-vtale靶点所在dna的甲基化水平，发现dna甲基化水平明显提高。t4代rb-vtale-atkyp转基因拟南芥植物对甘蓝曲叶病毒(calcuv)具有很好的抗性效果，而且植物内源转座子mu1和休眠相关基因的表达没有受rb-vtale-atkyp分子的影响。对t4代rb-vtale-atkyp转基因拟南芥植物注射接种calcuv侵染性克隆，从发病症状、病情指数和病毒滴度三方面数据说明rb-vtale-atkyp的转基因植物对甘蓝曲叶病毒的抗性可以稳定遗传到第四代；对植物内源转座子mu1和休眠相关基因的表达谱分析的结果表明rb-vtale-atkyp的转基因植物中植物内源转座子mu1和休眠相关基因的表达并没有明显变化，说明rb-vtale-atkyp分子并未脱靶作用于植物内源转座子mu1和休眠相关基因上。所述植物可以选自拟南芥、本氏烟草或小麦，但不限于此。利用本发明的方法培育的广谱双生病毒抗性的植物可以抗多个双生病毒属的双生病毒，包括菜豆金色花叶病毒属中多种重要的病原甘蓝曲叶病毒(cabbageleafcurlvirus，calcuv)，印度木薯花叶病毒(indiancassavamosaicvirus，icmv)，番茄黄化曲叶病毒(tomatoyellowleafcurlvirus，tylcv)；甜菜曲顶病毒属的甜菜曲顶病毒(beetcurlytopvirus，bctv)，或玉米线条病毒属的小麦矮缩病毒(wheatdwarfvirus，wdv)，但不限于这些病毒。通过本发明的方法能使植物获得对双生病毒的广谱抗性，也可以使植物获得对某种双生病毒特异的、稳定的高抗效果，这两种抗性效果都是基于对双生病毒染色质进行表观遗传学修饰从而抑制病毒相关基因的复制或转录过程的策略。综上所述，本发明提供下述技术方案：1.一种改造的抗双生病毒的vtale分子，其以植物病原细菌水稻黄单胞杆菌(xanthomonasoryzaepv.oryzae)三型分泌系统效应因子avrxa27基因序列为骨架序列，保留avrxa27基因的n端和c端序列，然后对avrxa27基因的dna识别/结合结构域序列和功能输出结构域序列进行改造而获得，其中所述改造后的dna识别/结合结构域序列靶向seqidno：1所示的ir-vtale靶点序列或seqidno：2所示的rb-vtale靶点序列，并且所述改造后的功能输出结构域序列选自由编码seqidnos：8-12所示的氨基酸序列的核苷酸序列组成的组中的任一种。2.第1项所述的改造的抗双生病毒的vtale分子，其中所述改造后的dna识别/结合结构域序列靶向seqidno：1所示的ir-vtale靶点序列。3.第1项所述的改造的抗双生病毒的vtale分子，其中所述改造后的dna识别/结合结构域序列靶向seqidno：2所示的rb-vtale靶点序列。4.第1项所述的改造的抗双生病毒的vtale分子，其中所述改造后的功能输出结构域序列编码seqidno：8所示的atkyp蛋白。5.第1项所述的改造的抗双生病毒的vtale分子，其中所述改造后的功能输出结构域序列编码seqidno：9所示的athdac19蛋白。6.第1项所述的改造的抗双生病毒的vtale分子，其中所述改造后的功能输出结构域序列编码seqidno：10所示的atmet1蛋白。7.第1项所述的改造的抗双生病毒的vtale分子，其中所述改造后的功能输出结构域序列编码seqidno：11所示的atsrdx蛋白。8.第1项所述的改造的抗双生病毒的vtale分子，其中所述改造后的功能输出结构域序列编码seqidno：12所示的atsuvh2蛋白。9.第1项所述的改造的抗双生病毒的vtale分子，其中所述改造后的功能输出结构域序列是选自由seqidnos：3-7所示的核苷酸序列组成的组的任一种核苷酸序列。10.第1项所述的改造的抗双生病毒的vtale分子，其中所述vtale分子的核苷酸序列如seqidnos：14-16和21-25所示。11.第10项所述的改造的抗双生病毒的vtale分子，其核苷酸序列如seqidno：14所示，命名为ir-vtale-atkyp。12.第10项所述的改造的抗双生病毒的vtale分子，其核苷酸序列如seqidno：21所示，命名为rb-vtale-atkyp。13.一种培育具有稳定的广谱双生病毒抗性的植物的方法，所述方法包括下述步骤：(1)vtale分子的构建：所述vtale分子以植物病原细菌水稻黄单胞杆菌(xanthomonasoryzaepv.oryzae)三型分泌系统效应因子avrxa27基因序列为骨架序列，保留avrxa27基因的n端和c端序列，然后对avrxa27基因的dna识别/结合结构域序列和功能输出结构域序列进行改造而获得，其中所述改造后的dna识别/结合结构域序列靶向seqidno：1所示的ir-vtale靶点序列或seqidno：2所示的rb-vtale靶点序列；并且所述改造后的功能输出结构域序列选自由编码seqidnos：8-12所示的氨基酸序列的核苷酸序列组成的组中的任一种；(2)植物转化：将步骤(1)构建的vtale分子，转化到所述植物中进行稳定表达。14.第13项所述的方法，其中所培育的植物抗多个双生病毒属的双生病毒，包括菜豆金色花叶病毒属中多种重要的病原甘蓝曲叶病毒(cabbageleafcurlvirus，calcuv)，印度木薯花叶病毒(indiancassavamosaicvirus，icmv)，番茄黄化曲叶病毒(tomatoyellowleafcurlvirus，tylcv)；甜菜曲顶病毒属的甜菜曲顶病毒(beetcurlytopvirus，bctv)，或玉米线条病毒属的小麦矮缩病毒(wheatdwarfvirus，wdv)。附图说明从下面结合附图的详细描述中，本发明的上述特征和优点将更明显，其中：图1显示vtale分子构建示意图。图1a.tale分子结构示意图。现有技术中已有的一般tale分子主要由两部分组成：dna识别/结合结构域(recognition/bindingdomain)和激活结构域(activationdomain，ad)。其中dna识别/结合结构域是由一段串联排列的重复序列组成，能够特异性识别并结合靶基因序列，每个重复序列由长度为33-35个氨基酸组成，除了第12和13位的氨基酸可变外，其他氨基酸序列高度保守，而重复序列可变的双氨基酸残基(repeatvariabledi-residue，rvd)是tale实现靶向识别特异dna碱基的关键位点，不同的rvd能够相对特异地识别a、t、c、g四种碱基中的一种或多种，其中分别与这4种碱基相对应的最常见的4种rvd分别是ni＝a，ng＝t，hd＝c和nn＝g或a。激活结构域能够在tale特异性识别并结合靶基因后激活靶基因的表达。图1b.本发明设计的vtale分子结构示意图。本发明设计的vtale分子的dna识别/结合结构域所结合的靶点序列选自下述中的任一种：针对双生病毒甘蓝曲叶病毒(cabbageleafcurlvirus，calcuv)基因组共同区(commonregion，cr)序列设计的双生病毒科病毒高度保守的vtale靶点intergeneticregion-vtale(ir-vtale)和calcuv特异的含有复制相关蛋白(replicationassociatedprotein，rep)结合位点的vtale靶点repbinding-vtale(rb-vtale)。此外，由于tale蛋白所识别的靶点dna序列必须以t开始，所以，虽然两个靶点序列都只含有16.5个串联排列的重复序列，但是都可以特异识别并结合18bp的靶点dna序列。激活结构域被替换为分别融合表达拟南芥(arabidopsisthaliana)不同的表观遗传学修饰相关效应因子(epi-effector)，包括组蛋白甲基转移酶atkyp，组蛋白去乙酰化酶athdac19，dna甲基转移酶atmet1，转录抑制基序atsrdx和虽同atkyp同属一个家族，但是功能上不同的组蛋白甲基转移酶atsuvh2。图1c.双生病毒科不同属病毒的ir-vtale靶点序列比对结果。begomovirus，菜豆金色花叶病毒属；turncurtovirus，芜菁曲顶病毒属；curtovirus，甜菜曲顶病毒属；topocuvirus，番茄伪曲顶病毒属；mastrevirus，玉米线条病毒属。calcuv：甘蓝曲叶病毒，cabbageleafcurlvirus；icmv：印度木薯花叶病毒，indiancassavamosaicvirus；tylcv：番茄黄化曲叶病毒，tomatoyellowleafcurlvirus；tctv：芜菁曲顶病毒，turnipcurlytopvirus；bctv：甜菜曲顶病毒，beetcurlytopvirus；tpctv：番茄伪曲顶病毒，tomatopseudo-curlytopvirus；wdv：小麦矮缩病毒，wheatdwarfvirus；acmv：非洲木薯花叶病毒，africancassavamosaicvirus；eacmv：东非木薯花叶病毒，eastafricancassavamosaicvirus；eacmzv：东非木薯花叶病毒桑给巴尔分离株，eastafricancassavamosaiczanzibarvirus；eacmkv：东非木薯花叶病毒肯尼亚分离株，eastafricancassavamosaickenyavirus。图1d.本发明设计的vtale和crispr/cas9的靶点序列在calcuv基因组dna-a链共同区中的位置示意图。两个vtale靶点(ir-vtale和rb-vtale)及两个crispr/cas9靶点(rb-sgrna和tata-sgrna)均位于calcuv基因组dna-a链的共同区内。蓝色框内序列为推测的calcuv由ac1编码的复制相关蛋白(replication-associatedprotein，rep)结合位点，其中下划线部分为保守序列；黄色框内序列为calcuv基因组中转录因子结合位点tatabox。蓝色曲线位置序列为双生病毒科病毒高度保守的ir-vtale靶点，其转录方向同毒粒链方向，位于发夹环内(haripin)；红色箭头位置序列为calcuv特异的rb-vtale靶点，其转录方向同互补链方向，含有tatabox和推测的calcuvrep蛋白结合位点序列。绿色箭头位置序列为crispr/cas9的rb-sgrna靶点，其转录方向同互补链方向，含有推测的calcuvrep蛋白结合位点序列。紫色箭头位置序列为crispr/cas9的tata-sgrna靶点，其转录方向同毒粒链方向，含有calcuv基因组中转录因子结合位点tatabox。图2显示在本氏烟草中瞬时表达ir-vtale-atkyp分子对双生病毒科不同病毒的抗性效果。图2a.显示在本氏烟草中瞬时表达空白载体对照ir-vtale-mcs和ir-vtale-atkyp分子24h后接种双生病毒科不同病毒的发病症状。包括玉米线条病毒属(mastrevirus)小麦矮缩病毒(wheatdwarfvirus，wdv)21dpi的症状；甜菜曲顶病毒属(curtovirus)甜菜曲顶病毒(beetcurlytopvirus，bctv)5dpi的症状；菜豆金色花叶病毒属(begomovirus)单组分病毒番茄黄化曲叶病毒(tomatoyellowleafcurlvirus，tylcv-sh2)10dpi的症状和双组分病毒印度木薯花叶病毒(indiancassavamosaicvirus，icmv-sg)7dpi的症状。bar：1厘米。图2b.显示在本氏烟草中瞬时表达空白载体对照ir-vtale-mcs和ir-vtale-atkyp分子24h后接种双生病毒科不同病毒的病毒滴度。包括21dpiwdv，5dpibctv，10dpitylcv和7dpiicmv-sg的病毒滴度。图3显示在小麦中瞬时表达ir-vtale-atkyp分子对小麦矮缩病毒(wdv)的抗性效果。图3a.显示分别将ir-vtale-mcs，ir-vtale-atkyp，rb-vtale-mcs和rb-vtale-atkyp分子与wdv混合接种于小麦品种jn17种子胚及其周围位置10d的发病症状。bar：1厘米。图3b.显示分别将ir-vtale-mcs和ir-vtale-atkyp分子与wdv混合接种于小麦品种jn17种子胚及其周围位置6个月后的发病症状。bar：1厘米。图3c.显示分别将ir-vtale-mcs，ir-vtale-atkyp和rb-vtale-atkyp分子与wdv混合接种于小麦品种jn17种子胚及其周围位置10d后的病毒滴度。图4显示在本氏烟中瞬时表达vtale和crispr/cas9分子对甘蓝曲叶病毒(calcuv)的抗性效果的比较。图4a.显示注射mma缓冲液的mock对照。bar：1厘米。图4b-d.显示在本氏烟中瞬时表达载体对照sgrna-vector(b)，tata-sgrna(c)和rb-sgrna(d)分子24h后接种calcuv14dpi的发病症状。bar：1厘米。图4e-i.显示在本氏烟中瞬时表达载体对照rb-vtale-mcs(e)，rb-vtale-atkyp(f)，rb-vtale-athdac19(g)，rb-vtale-atmet1(h)和rb-vtale-atsrdx(i)分子24h后接种calcuv14dpi的发病症状。bar：1厘米。图4j-l.显示在本氏烟中瞬时表达ir-vtale-atmet1(j)，ir-vtale-athdac19(k)和ir-vtale-atkyp(l)分子24h后接种calcuv14dpi的发病症状。bar：1厘米。图4m.显示在本氏烟中瞬时表达vtale和crispr/cas9分子24h后接种calcuv14dpi的病毒滴度。图5显示rb-vtale-atkyp转化拟南芥的植物对甘蓝曲叶病毒(calcuv)的抗性效果。图5a.显示rb-vtale-mcs，rb-vtale-atkyp，rb-vtale-athdac19，rb-vtale-atmet1和rb-vtale-atsuvh2转化拟南芥的t1代植物和col-0对照接种calcuv两个月后的病情指数。图5b-d.显示rb-vtale-atkyp(b)转化拟南芥的t2代植物和载体对照rb-vtale-mcs(c和d)接种calcuv1个月后的发病症状。图5e.显示rb-vtale-mcs和rb-vtale-atkyp(#1，#2，#4，#5，#6和#8)转化拟南芥的t2代植物和col-0对照接种calcuv1个月后的病毒滴度。图6显示t3代rb-vtale-atkyp转基因拟南芥植物对甘蓝曲叶病毒(calcuv)的抗性效果及atkyp基因的表达谱分析。图6a.显示t3代rb-vtale-mcs(#1)和rb-vtale-atkyp(#1和#4)转基因拟南芥植物接种calcuv32天后的发病症状。图6b.显示t3代rb-vtale-mcs(#1)和rb-vtale-atkyp(#1，#2，#4，#8，#11，#13，#15和#18)转基因拟南芥和col-0对照接种calcuv18天、25天及32天后的病情指数。图6c.显示t3代rb-vtale-mcs(#1)和rb-vtale-atkyp(#1和#4)转基因拟南芥接种calcuv18天、25天及32天时的病毒滴度。图6d.显示在t3代rb-vtale-atkyp(#1-3，#13-1，#15-1和#18-1)转基因拟南芥植物中atkyp基因的表达谱分析。图7显示t3代rb-vtale-atkyp转基因拟南芥植物中接种的甘蓝曲叶病毒基因组特定位置的dna与组蛋白甲基化水平及病毒相关基因的表达谱的分析。图7a.显示本发明中使用的calcuv基因组dna-a链中核小体的分布情况，rb：rb-vtale靶点，cr：共同区。图7b.显示t3代rb-vtale-atkyp转基因植物对接种的calcuv基因组dna-a链中#1核小体的组蛋白h3第9位和27位赖氨酸甲基化以及第14位赖氨酸乙酰化的影响。图7c.显示t3代rb-vtale-atkyp转基因植物对接种的calcuv基因组dna-a链中#5，#6，#8和#9核小体的组蛋白h3第9位和27位赖氨酸甲基化的影响。图7d.显示t3代rb-vtale-atkyp转基因植物对接种的calcuv基因组dna-a链中#1，#5，#6，#8和#9核小体的dna甲基化的影响。图7e.显示t3代rb-vtale-atkyp转基因植物对接种的calcuv的共同区(commonregion，cr)，互补链编码复制酶蛋白的基因acl和互补链编码参与病毒复制或转录调控蛋白的基因ac4的表达谱的影响。图8显示t4代rb-vtale-atkyp转基因拟南芥植物对甘蓝曲叶病毒(calcuv)的抗性效果、植物内源转座子mu1和休眠相关基因的表达谱的影响。图8a和8b.显示t4代rb-vtale-atkyp(b)转基因拟南芥植物和载体对照rb-vtale-mcs(a)接种calcuv21天时的发病症状。bar：1厘米。图8c.显示t4代rb-vtale-atkyp(#1-4和#1-12)转基因拟南芥植物和载体对照rb-vtale-mcs(#1)接种calcuv13天、21天及35天时的病情指数。图8d.显示t4代rb-vtale-atkyp(#1-4和#1-12)转基因拟南芥植物和载体对照rb-vtale-mcs(#1)接种calcuv21天时的病毒滴度。图8e.显示t4代rb-vtale-atkyp(#1-4和#1-12)和rb-vtale-mcs(#1)转基因拟南芥叶片中植物内源转座子mu1的表达谱分析。图8f.显示t4代rb-vtale-atkyp(#1-12)和rb-vtale-mcs(#1)转基因拟南芥种子中休眠相关基因(atabi3、atdog1和atnced6)的表达谱分析。图9显示ir-vtale-mcs核苷酸序列(seqidno：13)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向ir-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列。图10显示ir-vtale-atkyp核苷酸序列(seqidno：14)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向ir-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表atkyp核苷酸序列。图11显示ir-vtale-athdac19核苷酸序列(seqidno：15)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向ir-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表athdac19核苷酸序列。图12显示ir-vtale-atmet1核苷酸序列(seqidno：16)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向ir-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表atmet1核苷酸序列。图13显示ir-vtale-atkyp氨基酸序列(seqidno：17)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向ir-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表atkyp氨基酸序列。图14显示ir-vtale-athdac19氨基酸序列(seqidno：18)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向ir-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表athdac19氨基酸序列。图15显示ir-vtale-atmet1氨基酸序列(seqidno：19)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向ir-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表atmet1氨基酸序列。图16显示rb-vtale-mcs核苷酸序列(seqidno：20)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向rb-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列。图17显示rb-vtale-atkyp核苷酸序列(seqidno：21)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向rb-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表atkyp核苷酸序列。图18显示rb-vtale-athdac19核苷酸序列(seqidno：22)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向rb-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表athdac19核苷酸序列。图19显示rb-vtale-atmet1核苷酸序列(seqidno：23)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向rb-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表atmet1核苷酸序列。图20显示rb-vtale-atsrdx核苷酸序列(seqidno：24)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向rb-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表atsrdx核苷酸序列。图21显示rb-vtale-atsuvh2核苷酸序列(seqidno：25)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向rb-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表atsuvh2核苷酸序列。图22显示rb-vtale-atkyp氨基酸序列(seqidno：26)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向rb-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表atkyp氨基酸序列。图23显示rb-vtale-athdac19氨基酸序列(seqidno：27)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向rb-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表athdac19氨基酸序列。图24显示rb-vtale-atmet1氨基酸序列(seqidno：28)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向rb-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表atmet1氨基酸序列。图25显示rb-vtale-atsrdx氨基酸序列(seqidno：29)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向rb-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表atsrdx氨基酸序列。图26显示rb-vtale-atsuvh2氨基酸序列(seqidno：30)，其中阴影代表来源于xoo的avrxa27基因n端序列，下划线代表靶向rb-vtale靶点的16.5个重复序列，斜体代表来源于xoo的avrxa27基因c端序列，粗体代表atsuvh2氨基酸序列。具体实施方式下面参照具体的实施例进一步描述本发明，但是本领域技术人员应该理解，本发明并不限于这些具体的实施例。实施例1.构建vtale分子转录激活效应子(transcriptionactivator-likeeffector，tale)是一类来源于植物病原细菌黄单胞杆菌属(xanthomonasspp.)三型分泌系统的效应因子，能够特异性识别并结合水稻的r基因从而激发水稻的抗病性(keyuguetal.，2005，nature)。但直到2009年，两个独立的实验室才解析了tale特异性识别并结合靶基因的分子机制(moscouetal.，2009，science和bochetal.，2009，science)。如图1a所示，一般tale蛋白主要由两部分组成：由串联排列的重复序列组成的dna识别/结合结构域(recognition/bindingdomain)和激活结构域(activationdomain，ad)。其中组成dna识别/结合结构域的串联重复序列由长度为33-35个氨基酸组成，除了第12和13位的氨基酸可变外，每个重复序列的氨基酸序列高度保守，而可变的第12和13位的氨基酸被称为重复序列可变的双氨基酸残基(repeatvariabledi-residue，rvd)，是tale特异性识别并结合dna序列的关键位点，不同的rvd能够相对特异地识别a、t、c、g四种碱基中的一种或多种，其中分别与这4种碱基相对应的最常见的4种rvd分别是ni＝a，ng＝t，hd＝c和nn＝g或a。激活结构域能够在tale特异性识别并结合靶基因后激活靶基因的表达。本发明以双生病毒科病毒为研究对象，期望利用上述的tale技术发展出一种提高植物对双生病毒抗性的新方法。我们的策略是人工设计全新的vtale分子，该分子的dna识别/结合结构域能够特异性识别并结合双生病毒基因组的一段序列，而激活结构域则被置换为具有抑制作用的表观遗传学修饰相关效应因子(epi-effector)。因此当在植物中表达这种vtale分子时，入侵的双生病毒就会被vtale分子识别并特异性结合在病毒基因组靶点位置，然后融合表达的表观遗传学修饰相关效应因子就会对靶点位置的病毒染色质进行表观遗传学修饰，从而抑制双生病毒基因组dna的复制和转录，达到提高植物对双生病毒抗性的目的(图1b)。所有双生病毒基因组中均含有一段共同保守的非编码的基因间区序列——共同区(commonregion，cr)，该区域富含病毒复制和转录的各种控制元件。如双生病毒科病毒番茄金色花叶病毒(tomatogoldenmosaicvirus，tgmv)共同区第54-153bp的序列中含有复制相关蛋白(replicationassociatedprotein，rep)的结合位点5’-ggtagtaaggtag-3’。rep蛋白与病毒的复制相关，可以起始毒粒链dna的合成，rep蛋白结合位点是病毒特异性的，但具有一些重复的保守序列如tgmv的ggtag序列，此外重复序列间的间隔序列也很重要如tgmv的间隔序列taa。在rep蛋白结合位点下游是转录因子结合位点tata框(tatabox)，它与病毒的转录有关。在tata框下游，有段序列能形成发夹环的结构，该结构与病毒的复制有关，该环中存在保守的序列5’-taatattac-3’(马修斯植物病毒学第八章第八节d3)。对双生病毒科不同属病毒含有发夹环序列的18bpdna序列进行比对分析，发现该段序列在双生病毒科内高度保守(图1c)，因此我们将该段18bpdna序列作为vtale分子的intergenicregion-vtale(ir-vtale)靶点，希望通过序列高度保守的ir-vtale靶点获得对双生病毒科病毒的广谱抗性。另外我们分析了calcuv基因组共同区第91-192bp的序列，结果如图1d所示，在calcuv共同区中存在可能的rep蛋白结合位点5’-ggtgtcttggtgt-3’，tata框和发夹环结构，然后我们将calcuv含有tata框和部分rep蛋白结合位点的18bpdna序列作为vtale分子的repbinding-vtale(rb-vtale)靶点，希望通过病毒特异性的rb-vtale靶点获得对calcuv更特异性的抗性(图1b和1d)。与ir-vtale和rb-vtale靶点融合表达的表观遗传学修饰相关效应因子均来自于模式植物拟南芥(arabidopsisthaliana)，包括组蛋白甲基转移酶atkyp，组蛋白去乙酰化酶athdac19，dna甲基转移酶atmet1，转录抑制基序atsrdx和虽同atkyp同属一个家族，但是功能上不同的组蛋白甲基转移酶atsuvh2。参照文献(fengzhangetal.，2011，naturebiotechnology)报道的基于层级连接的策略构建tale蛋白dna识别/结合结构域的串联排列重复序列的方法，构建3个ir-vtale-epi-effector分子、5个rb-vtale-epi-effector分子及其对应的载体对照分子ir-vtale-mcs和rb-vtale-mcs，详细信息见表1。将构建好的10个vtale分子分别转化农杆菌菌株c58c1的感受态细胞(购自北京博迈德生物技术有限公司)，利用含有相应抗生素的lb固体平板筛选阳性克隆。实施例2.在本氏烟草中瞬时表达ir-vtale-atkyp分子对双生病毒科病毒具有广谱抗性。将双子叶植物本氏烟草(nicotianabenthamiana)种子均匀播种于灭菌的营养土表面，浇水并覆盖塑料薄膜以充分保湿，置于25℃植物培养箱中14小时光照/10小时黑暗培养10天左右，然后将萌发的小苗单株移栽到含有灭菌营养土的营养钵中，再生长3周左右即可用于实验。将含有ir-vtale-mcs(ir-vtale分子空白载体对照)和ir-vtale-atkyp的农杆菌接种于含有相应抗生素的lb液体培养基中活化培养24-36小时(抗生素见表1)，然后取约1ml新鲜菌液接种于含有相应抗生素、mes(10mm)和as(40μm)的20mllb液体培养基中培养16-24小时，待od600＝1.0-1.5之间时，离心收集菌体，再用mma缓冲液重悬(10mmmes，10mmmgcl2，200μmas)，调od600＝1.0，静止2h后用1ml注射器接种本氏烟草叶片背面，使整片叶子充分浸润菌液。注射后的本氏烟在25℃，14h光照/10h黑暗条件下的培养箱中生长24小时后，使用相同方法按照表1的抗生素种类及浓度分别重新接种双生病毒科不同属病毒的侵染性克隆，包括玉米线条病毒属(mastrevirus)小麦矮缩病毒(wheatdwarfvirus，wdv，中国农业科学院植物保护研究所王锡峰课题组惠赠)；甜菜曲顶病毒属(curtovirus)甜菜曲顶病毒(beetcurlytopvirus，bctv，本实验室保藏)；菜豆金色花叶病毒属(begomovirus)单组分病毒番茄黄化曲叶病毒(tomatoyellowleafcurlvirus，tylcv-sh2，中国农业科学院植物保护研究所周雪平课题组惠赠)和双组分病毒印度木薯花叶病毒(indiancassavamosaicvirus，icmv-sg，本实验室分离鉴定并保存，gangwangetal.，2014，virusgenes)。接种后的本氏烟草置于25℃，14h光照/10h黑暗条件下的培养箱中生长，观察发病情况。bctv，icmv-sg，tylcv和wdv分别于接种后的第5天，第7天，第10天和第21天在表达空白载体对照ir-vtale-mcs分子的本氏烟中出现明显的发病症状，其中接种wdv的叶片明显增厚和皱缩，接种bctv的叶片出现半透明状沿叶脉扩展的症状，接种tylcv的叶片出现明显的增厚、皱缩和向下扭曲的症状，接种icmv-sg的叶片出现严重的增厚、皱缩和向下扭曲的症状，而表达ir-vtale-atkyp分子的本氏烟中未见明显的发病症状(图2a)。为了进一步验证观察到的结果，我们取发病的样品进行病毒滴度的测定。每个处理至少取四个生物学重复，每一个样品100mg，使用植物基因组dna提取试剂盒(购自qiagen公司)提取基因组dna。取2μldna为模板，0.4μl10μm的引物，10μlsybrgreen(toyobo)，然后用ddh2o补足至20μl。实时pcr程序为预变性95℃10min，变性95℃15s，退火58℃30s，延伸72℃30s，40个循环，补平延伸5min。所有样品都做三次技术重复。nbef1a为内参基因(引物见表2)，进行实时pcr分析。如图2b所示，bctv，icmv-sg，tylcv和wdv在表达空白载体对照ir-vtale-mcs分子的本氏烟中的病毒滴度明显高于表达ir-vtale-atkyp分子的本氏烟，其中wdv在瞬时表达ir-vtale-mcs分子的本氏烟中的病毒滴度是瞬时表达ir-vtale-atkyp分子的本氏烟的3倍多，bctv在瞬时表达ir-vtale-mcs分子的本氏烟中的病毒滴度是瞬时表达ir-vtale-atkyp分子的本氏烟的1倍多，tylcv在瞬时表达ir-vtale-mcs分子的本氏烟中的病毒滴度是瞬时表达ir-vtale-atkyp分子的本氏烟的1倍多，icmv-sg在瞬时表达ir-vtale-mcs分子的本氏烟中的病毒滴度是瞬时表达ir-vtale-atkyp分子的本氏烟的约500000倍，这与我们观察到的发病结果相吻合，说明瞬时表达ir-vtale-atkyp分子的本氏烟表现出抗多种双生病毒侵染的能力。结论：针对双生病毒calcuv基因组共同区序列设计的双生病毒科病毒序列高度保守的vtale靶点ir-vtale融合表达拟南芥组蛋白甲基转移酶kyp的vtale分子ir-vtale-atkyp在双子叶植物本氏烟草中瞬时表达24小时后即能够增强本氏烟对多种双生病毒的抗性能力，也就是说ir-vtale-atkyp分子对双生病毒科病毒具有广谱抗性的效果。表2.本发明中病毒滴度实验所用到的引物实施例3.在小麦中瞬时表达ir-vtale-atkyp分子对小麦矮缩病毒具有抗性。将分别含有ir-vtale-mcs(ir-vtale空白载体对照)，ir-vtale-atkyp，rb-vtale-mcs(rb-vtale空白载体对照)，rb-vtale-atkyp和wdv侵染性克隆的农杆菌接种于含有相应抗生素的3mllb液体培养基中(抗生素见表1)，活化培养至od600＝2.0，离心收集菌体，再用mma缓冲液浓缩至0.75ml(10mmmes，10mmmgcl2，200μmas)，静止2h后，分别将vtale分子与wdv侵染性克隆按3∶1的比例混匀，然后用昆虫针蘸取混好的菌液，穿刺接种于灭菌水浸泡过夜的小麦(triticumaestivumlinn)品种jn17(中国科学院遗传与发育生物学研究所李振生课题组惠赠)的胚及其周围位置，每个种子从不同角度接种五次，然后将接种的种子置于不含抗生素的lb固体平板上28℃过夜培养，第二天将其均匀播种于灭菌的营养土中，25℃植物培养箱中18小时光照/6小时黑暗培养2-4周，观察发病情况。在vtale和wdv混合接种10天后，wdv分别与ir-vtale-mcs(ir-vtale载体对照)，rb-vtale-mcs(rb-vtale载体对照)和rb-vtale-atkyp混合接种的小麦与空白对照mock植物相比植株明显矮小，生长状态不佳；而wdv与ir-vtale-atkyp接种的小麦表现出比其他vtale分子接种的小麦更健康的表型，植株明显高大、健壮，生长状态更接近于mock对照(图3a)。为了进一步验证观察到的结果，我们按照实施例2中所述的病毒滴度测定方法以小麦actin为内参基因(引物见表2)对混合接种vtale和wdv的小麦进行病毒滴度的测定。如图3c所示，wdv在与ir-vtale-mcs(ir-vtale载体对照)混合接种的小麦中的病毒滴度是与ir-vtale-atkyp混合接种的小麦的约4倍，而wdv在与rb-vtale-atkyp混合接种的小麦中的病毒滴度是与ir-vtale-mcs(ir-vtale载体对照)混合接种的小麦的约1.5倍，这与我们观察到的发病结果相吻合，说明瞬时表达ir-vtale-atkyp分子的小麦表现出高抗wdv侵染的能力，而rb-vtale-atkyp分子没有抵抗wdv侵染的能力。为了更好地检验瞬时表达ir-vtale-atkyp分子的小麦抗小麦矮缩病毒(wheatdwarfvirus，wdv)侵染的效果，我们将vtale分子和wdv混合接种的小麦移栽到大钵中，在适宜小麦生长的季节将其置于室外自然条件下生长，在注射接种后的6个月，我们发现wdv与ir-vtale-atkyp混合接种的小麦植株明显高大、健壮、能够正常抽穗，生长状态更接近于mock对照；而wdv与ir-vtale-mcs(ir-vtale载体对照)混合接种的小麦植株矮小、几乎绝产(图3b)。说明瞬时表达ir-vtale-atkyp分子的小麦对wdv的侵染具有高抗且持久的效果。结论：针对双生病毒calcuv基因组共同区序列设计的双生病毒科病毒高度保守的tale靶点ir-vtale融合表达拟南芥组蛋白甲基转移酶kyp的vtale分子ir-vtale-atkyp在单子叶作物小麦中瞬时表达后能够增强小麦对wdv的抗性能力，这一研究结果在培育高抗植物双生病毒的小麦品种方面具有很好的应用前景。而calcuv特异的tale靶点rb-vtale融合表达拟南芥组蛋白甲基转移酶kyp的vtale分子rb-vtale-atkyp没有抵抗wdv侵染的能力。实施例4.在本氏烟中瞬时表达vtale和crispr/cas9分子对甘蓝曲叶病毒的抗性效果比较。首先，我们针对calcuv基因组dna-a链共同区序列设计包括tata-sgrna(a2)和rb-sgrna(a1)在内的8个crispr/cas9靶点(引物序列见表3)，然后使用annealingbuffer(购自碧云天生物技术有限公司)将每个靶点的正、反向引物退火形成互补的双链dna分子，最后将annealing产物与被bsai消化过的载体phsn401(中国科学院遗传与发育生物学研究所高彩霞课题组惠赠)进行连接反应，4小时后将连接产物转化大肠杆菌dh5α感受态(购自北京博迈德生物技术有限公司)，复苏1小时后涂布于含有卡那霉素(50mg/l)的lb固体平板上筛选阳性克隆，挑取阳性克隆提取质粒送测，测序结果比对正确的克隆转化农杆菌c58c1感受态(购自北京博迈德生物技术有限公司)，利用含有卡那霉素(50mg/l)，利福平(20mg/l)和四环素(10mg/l)的固体lb平板筛选阳性克隆。其次，按照实施例2中所述的方法对双子叶植物本氏烟进行vtale和crispr/cas9分子的瞬时表达，24h后再重新注射接种calcuv侵染性克隆(本实验室保存)，观察发病情况。接种calcuv侵染性克隆14天时我们发现与mock植物相比(图4a)，瞬时表达crispr/cas9载体对照sgrna-vector的本氏烟叶片上有明显的calcuv感染的症状，包括叶片上呈现褪绿斑点，叶片明显增厚并向下卷曲(图4b)；而除了瞬时表达rb-sgrna(a1)靶点的本氏烟叶片上未见明显的calcuv感染的症状外(图4d)，瞬时表达其他crispr/cas9靶点的本氏烟均具有与载体对照类似的明显的calcuv感染的症状(图4c，更具体的数据未提供)，说明我们设计的8个crispr/cas9靶点中只有rb-sgrna(a1)靶点具有相对比较好的抗calcuv侵染的能力。另一方面，瞬时表达vtale载体对照rb-vtale-mcs的本氏烟叶片上有明显的与sgrna-vector类似的calcuv感染的症状(图4e)，而瞬时表达rb-vtale-epi-effector【rb-vtale-atkyp(图4f)，rb-vtale-athdac19(图4g)，rb-vtale-atmet1(图4h)和rb-vtale-atsrdx(图4i)】的本氏烟叶片上未见明显的calcuv感染的症状，与瞬时表达crispr/cas9靶点rb-sgrna(a1)的本氏烟类似；而ir-vtale-epi-effector中除了瞬时表达ir-vtale-atkyp的本氏烟叶片上有比较微弱的calcuv感染的症状外(图4l)，瞬时表达ir-vtale-atmet1(图4j)和ir-vtale-athdac19(图4k)的本氏烟叶片上有明显的calcuv感染的症状，其发病程度与载体对照rb-vtale-mcs类似(图4e)，说明从总体来讲rb-vtale分子具有比crispr/cas9分子更强的抗calcuv侵染的效果。为了进一步验证观察到的结果，我们按照实施例2中所述的病毒滴度测定方法以nbef1a为内参基因(引物见表2)检测分别瞬时表达vtale和crispr-cas9分子的本氏烟中calcuv的病毒滴度。如图4m所示，calcuv在本氏烟叶片中的病毒滴度由低到高依次为瞬时表达rb-vtale-epi-effector(rb-vtale-atmet1，rb-vtale-atkyp，rb-vtale-atsrdx和rb-vtale-athdac19)的本氏烟叶片，平均病毒滴度不到载体对照的万分之一；其次是瞬时表达crispr/cas9靶点rb-sgrna(a1)的本氏烟叶片，病毒滴度比载体对照的万分之一略高；最后是瞬时表达ir-vtale-atkyp的本氏烟叶片，病毒滴度约为载体对照的千分之一；而瞬时表达其他vtale和crispr/cas9分子的本氏烟中calcuv的病毒滴度与瞬时表达相应的载体对照相类似。说明rb-vtale-epi-effectors分子对calcuv的侵染具有比crispr/cas9分子更好的抗性效果，此外ir-vtale-atkyp对calcuv的侵染也具有较好的抗性效果，这一结果与我们观察到的发病情况一致。结论：calcuv特异性的靶点rb-vtale融合表达表观遗传学修饰相关效应因子的vtale分子对calcuv的侵染具有比crispr/cas9分子更好的抗性效果，此外对双生病毒科病毒具有广谱抗性的ir-vtale靶点融合表达拟南芥组蛋白甲基转移酶kyp的vtale分子对calcuv的侵染也具有较好的抗性效果。表3.本发明中针对calcuv设计的crispr/cas9靶点序列引物引物名称引物序列(5’-3’)目的rb-sgrna(a1)fattgaccaagacaccaaaaacaatcrispr/cas9rb-sgrna(a1)raaacattgtttttggtgtcttggtcrispr/cas9tata-sgrna(a2)fattgaatatatactaaaagcctctcrispr/cas9tata-sgrna(a2)raaacagaggcttttagtatatattcrispr/cas9a3fattgaaagcctctggggaacaccacrispr/cas9a3raaactggtgttccccagaggctttcrispr/cas9a4fattggggaacaccaggggcaaaagcrispr/cas9a4raaaccttttgcccctggtgttccccrispr/cas9a5fattggccatccgcaataatattaccrispr/cas9a5raaacgtaatattattgcggatggccrispr/cas9a6fattgcggatggccgcttttgcccccrispr/cas9a6raaacggggcaaaagcggccatccgcrispr/cas9a7fattgtccggtaatattattgcggacrispr/cas9a7raaactccgcaataatattaccggacrispr/cas9a8fattgccggatggccgcgattttttcrispr/cas9a8raaacaaaaaatcgcggccatccggcrispr/cas9实施例5.rb-vtale-atkyp转化拟南芥的植物对甘蓝曲叶病毒的抗性效果分析。由于在本氏烟中瞬时表达不同的rb-vtale-epi-effector对calcuv的侵染均具有很好的抗性效果，因此我们想检测稳定表达不同的rb-vtale-epi-effector对calcuv的抗性效果。依照文献(xiurenzhangetal.，2006，natureprotocol)报道的拟南芥转化方法分别将含有不同rb-vtale-epi-effector分子的农杆菌菌株c58c1转化野生型拟南芥col-0，再对t1代种子进行转化子筛选，从而成功获得rb-vtale-mcs，rb-vtale-atkyp，rb-vtale-athdac19，rb-vtale-atmet1和rb-vtale-atsuvh2转基因拟南芥植物。然后，按照实施例2中所述的注射接种本氏烟草的方法将calcuv侵染性克隆接种到4周左右的rb-vtale-mcs，rb-vtale-atkyp，rb-vtale-athdac19，rb-vtale-atmet1和rb-vtale-atsuvh2转基因拟南芥的轮作叶片上，同时接种col-0做对照，观察发病情况。接种calcuv2个月后，我们将拟南芥发病情况按由轻到重的顺序划分为6个等级(0-5级)，利用病情指数这一指标初步统计每个vtale分子转基因植物对calcuv的抗性效果，结果如图5a所示，t1代rb-vtale-athdac19，rb-vtale-atmet1和rb-vtale-atkyp转基因拟南芥都表现出抗calcuv侵染的效果，但是rb-vtale-atsuvh2转基因植物表现出与载体对照和col-0相一致的表型，说明atsuvh2虽为拟南芥的另一个组蛋白甲基转移酶，同atkyp同属一个家族，但是功能上不同，对calcuv的侵染没有抗性效果。待种子成熟后，挑选载体对照rb-vtale-mcs和抗性比较好的rb-vtale-atkyp转基因株系的种子萌发t2代植株，按照实施例2中所述的注射接种本氏烟草的方法将calcuv侵染性克隆接种到4周左右的转基因拟南芥轮作叶上，观察发病情况。接种calcuv1个月后，t2代rb-vtale-atkyp转基因株系表现出高抗表型，植株生长健康，正常开花，无明显的叶片斑驳或褪绿的症状(图5b)，而t2代载体对照rb-vtale-mcs转基因株系表现出易感表型，包括植株矮小，花絮扭曲，花叶等症状(图5c和d)。为了进一步验证观察到的结果，我们按照实施例2中所述的病毒滴度测定方法以拟南芥tubulin2为内参基因(引物见表2)对接种calcuv的转基因拟南芥进行病毒滴度的测定。如图5e所示，病毒滴度分析的结果表明与载体对照rb-vtale-mcs转基因拟南芥植物相比rb-vtale-atkyp转基因拟南芥植物株系(#1，#4，#8)对calcuv具有高抗的表型，其中rb-vtale-atkyp转基因拟南芥植物株系#1的病毒滴度约为载体对照植物的1/14，#4和#8的病毒滴度约为载体对照植物的1/3，这一结果与观察到的发病症状的结果相一致。结论：calcuv特异的靶点rb-vtale融合表达拟南芥组蛋白甲基转移酶kyp的rb-vtale-atkyp分子在双子叶植物拟南芥中稳定表达后对calcuv的侵染具有很好的抗性效果。实施例6.t3代rb-vtale-atkyp转基因拟南芥植物对甘蓝曲叶病毒(calcuv)的抗性效果及atkyp基因的表达谱分析。待t3代转基因植物种子成熟后，挑选载体对照rb-vtale-mcs和抗性比较好的rb-vtale-atkyp转基因株系的种子萌发t3代植株，按照实施例2中所述的注射接种本氏烟草的方法将calcuv侵染性克隆接种到4周左右的转基因拟南芥轮作叶上，观察发病情况。接种calcuv32天后，t3代rb-vtale-atkyp转基因株系(#1和#4)表现出高抗表型，植株生长健康，正常开花，无明显的叶片斑驳或褪绿的症状，而t3代载体对照rb-vtale-mcs#1转基因株系表现出易感表型，包括植株矮小，花絮扭曲，花叶等症状，如图6a所示，其中左上角为rb-vtale-mcs#1转基因株系发病症状的俯视放大图。利用病情指数这一指标按发病早、中和晚期三个时间点统计的rb-vtale-atkyp转基因株系对calcuv的抗性效果如图6b所示，rb-vtale-atkyp转基因株系(#1，#4，#8和#18)在发病的早、中和晚期均对calcuv具有很好的抗性效果，说明rb-vtale-atkyp转基因拟南芥对calcuv的抗性效果可以稳定遗传给下一代。为了进一步验证观察到的结果，我们按照实施例2中所述的病毒滴度测定方法以拟南芥tubulin2为内参基因(引物见表2)对接种calcuv的转基因拟南芥进行病毒滴度的测定。如图6c所示，病毒滴度分析的结果表明rb-vtale-atkyp转基因植物株系(#1和#4)在发病的早、中和晚期对calcuv都具有高抗的效果，发病中期的抗性效果尤为显著，这一结果与观察到的发病症状及病情指数调查的结果相一致。为了检测不同rb-vtale-atkyp转基因植物株系中atkyp基因的表达水平，我们选取高抗株系#1-3和#18-1及易感株系#13-1和#15-1为待测样品。首先按atkyp基因核苷酸序列为模板设计扩增引物(见表4，atkyp-f和atkyp-r)，然后每种植物至少三个生物学重复，每一个样品取100mg，使用rna提取试剂盒(购自qiagen)提取总rna并对基因组dna进行消化处理，nanodrop定量后取2μg总rna为模板，mmlv反转录酶合成cdna。取2μlcdna为模板，0.4μl10μm的引物，10μlsybrgreen(toyobo)，然后用ddh2o补足至20μl。实时pcr程序为预变性95℃10min，变性95℃15s，退火58℃30s，延伸72℃30s，40个循环，补平延伸5min。所有样品都做三次技术重复。atactin2为内参基因(引物见表4，atactin2-f和atactin2-r)，进行实时pcr分析。如图6d所示，rb-vtale-atkyp转基因植物高抗株系#1-3和#18-1中atkyp基因的表达水平是易感株系#13-1和#15-1的约7倍。结论：calcuv特异的靶点rb-vtale融合表达拟南芥组蛋白甲基转移酶kyp的rb-vtale-atkyp分子在双子叶植物拟南芥中稳定表达后对calcuv的高抗表型能够稳定遗传给下一代，且与atkyp基因表达水平呈正相关。表4.本发明中用于基因表达分析的引物实施例7.t3代rb-vtale-atkyp转基因拟南芥植物中接种的甘蓝曲叶病毒基因组特定位置的组蛋白与dna甲基化水平及病毒相关基因的表达谱的分析由于atkyp是来源于拟南芥的组蛋白h3第9位赖氨酸甲基化转移酶基因，而前期的研究结果已经证明rb-vtale-atkyp转基因拟南芥植物对calcuv的高抗表型与atkyp基因表达水平呈正相关，且能够稳定遗传给下一代。这就暗示我们rb-vtale-atkyp转基因拟南芥植物对calcuv的高抗表型可能是通过atkyp甲基化修饰calcuv基因组rb-vtale靶点所在的核小体组蛋白，进而引起dna的甲基化修饰，使得calcuv基因组不能正常复制和转录，最终使得rb-vtale-atkyp转基因拟南芥植物获得对calcuv的抗性。为了验证这一推论，首先我们利用染色质免疫沉淀(chromatinimmuno-precipitation，chip)技术来检测calcuv基因组rb-vtale靶点所在核小体即#1核小体组蛋白的甲基化水平(图7a)。根据文献(in-cheoljangetal.，2011，theplantcell)报道的方法，首先取相同叶位的发病叶片3g，每个处理至少三个生物学重复，将样品浸润在终浓度为1％甲醛中，抽真空10min使染色质结合蛋白与dna交联，然后加入终浓度为0.125m的甘氨酸抽真空5min以终止交联反应，将样品用灭菌水冲洗三遍后，用液氮速冻并磨成粉末，再用一系列抽提缓冲液处理后，将收集的染色质经过超声波打断，再用识别特异组蛋白修饰的抗体(anti-h3k9me2，anti-h3k27me3和anti-h3k14ac均购自millipore公司)将核小体沉淀下来，缠绕在核小体上的dna也随之沉淀，解交联用蛋白酶k除去蛋白质后用pcr纯化试剂盒(购自qiagen公司)纯化出dna，再设计基因组特异位点的引物用定量pcr的方法检测dna的量(引物序列见表5)，dna的量的多少就可以反映出该位点对应的组蛋白修饰的相对富集程度。结果如图7b所示，rb-vtale-atkyp转基因拟南芥植物株系#1-12中calcuv基因组的#1核小体的组蛋白h3第9位赖氨酸的甲基化水平与载体对照植物中的相比提高10倍，说明rb-vtale-atkyp转基因拟南芥植物株系#1-12中rb-vtale-atkyp分子结合于calcuv基因组的rb-vtale靶点，然后atkyp对靶点所在的#1核小体的组蛋白h3第9位赖氨酸进行甲基化修饰。但是与此同时我们发现calcuv基因组的#1核小体所在的组蛋白h3第27位赖氨酸的甲基化水平与载体对照植物中的相比也提高5倍多，这可能也是由atkyp引起的，但是具体的机制我们还不清楚。此外我们发现calcuv基因组的#1核小体所在的组蛋白h3第14位赖氨酸的乙酰化水平与载体对照植物中的相比没有明显变化，说明rb-vtale-atkyp转基因拟南芥植物株系#1-12中rb-vtale-atkyp分子对calcuv基因组的#1核小体的组蛋白乙酰化没有作用，而只特异性地作用于甲基化修饰上。既然rb-vtale-atkyp转基因拟南芥植物中rb-vtale-atkyp分子能够甲基化修饰calcuv基因组的rb-vtale靶点所在的#1核小体的组蛋白，那么rb-vtale-atkyp分子是否对calcuv基因组中其他核小体的组蛋白也具有甲基化修饰的作用呢？带着这个问题我们选择了与calcuv基因组rb-vtale靶点平面距离较近的#8和#9核小体及平面距离较远的#5和#6核小体(图7a)来检测它们的组蛋白的甲基化修饰水平。结果如图7c所示，rb-vtale-atkyp转基因拟南芥植物中calcuv基因组的#9核小体的组蛋白h3第9位赖氨酸的甲基化水平与载体对照植物中的相比提高16倍，而#5、#6和#8核小体的组蛋白h3第9位赖氨酸的甲基化水平与载体对照植物中的相比并没有明显的变化。说明rb-vtale-atkyp转基因拟南芥植物中rb-vtale-atkyp分子能够甲基化修饰与其识别位点rb-vtale靶点平面距离最近的#9核小体的组蛋白，但不能甲基化修饰与rb-vtale靶点平面距离较远的#5、#6和#8核小体的组蛋白。此外我们发现rb-vtale-atkyp转基因拟南芥植物中calcuv基因组的#5核小体的组蛋白h3第27位赖氨酸的甲基化水平与载体对照植物中的相比提高12倍，而#6、#8和#9核小体的组蛋白h3第27位赖氨酸的甲基化水平并没有明显的变化，这可能是由atkyp引起的，虽然#5核小体与rb-vtale靶点所在的#1核小体平面距离最远，但是calcuv基因组dna在复制过程中会形成双链结构的复制中间体，进而招募寄主植物的组蛋白形成类似植物染色体结构的minichromosome，这种结构可能会拉近二者的空间距离，从而使得atkyp影响#5核小体的组蛋白h3第27位赖氨酸甲基化修饰的水平，但是具体原因我们还不能很好地解释。据文献(jiamuduetal.，2014，molecularcell)报道，拟南芥核小体组蛋白h3第9位赖氨酸在被kyp修饰带有甲基化marker后，就会招募cmt3(dna甲基转移酶chromomethylase3)对其dna进行甲基化修饰，带有甲基化marker的dna又会招募kyp对其所在核小体的组蛋白或与其相毗邻的核小体组蛋白h3第9位赖氨酸进行甲基化修饰，二者作为分子伴侣通过这样的方式共同控制核小体组蛋白h3第9位赖氨酸及其dna的甲基化修饰。前面结果显示rb-vtale-atkyp转基因拟南芥植物中calcuv基因组的#1和#9核小体的组蛋白h3第9位赖氨酸的甲基化水平与载体对照植物中的相比有明显提高，因此我们采用以下方法来检测rb-vtale-atkyp转基因拟南芥植物中calcuv基因组rb-vtale靶点所在#1核小体及其他核小体(#5，#6，#8和#9)内dna甲基化水平。取相同叶位的发病叶片，每一个样品100mg，每个处理至少三个生物学重复，使用基因组dna提取试剂盒(购自qiagen)进行高纯度基因组的提取；每个样品取400ng的基因组dna，分装两管，向处理管中加入3u甲基化敏感性限制性内切酶hpaii，相应的对照管中加入等体积的ddh2o，每管中加入3ul酶切缓冲液后用ddh2o定容至30ul，37℃单酶切消化5h后，加热至80℃失活20min；向酶切体系中加入50ulddh2o稀释，取2ul作为模板，以attubulin2为内参基因按照实施例2中所述的测定病毒滴度的反应体系及程序进行real-timepcr分析，所用的引物序列见表5。结果如图7d所示，rb-vtale-atkyp转基因拟南芥植物中calcuv基因组rb-vtale靶点所在#1核小体及与其平面距离最近的#9核小体内的dna甲基化水平与载体对照植物中的相比有所提升，而与rb-tale靶点平面距离较远的#5，#6和#8核小体内的dna甲基化水平与载体对照植物中的相比并没有明显的差异。说明rb-vtale-atkyp转基因拟南芥植物中rb-vtale-atkyp分子能够对其靶点所在及其毗邻的核小体组蛋白进行甲基化修饰，进而招募cmt3使其对dna进行甲基化修饰，但是对于其他核小体(#5，#6和#8核小体)的dna没有作用。由于rb-vtale靶点所在的#1核小体dna序列位于calcuv的共同区，calcuv的复制及转录起始相关酶均结合在共同区内特异位点上，而我们的数据表明rb-vtale-atkyp转基因拟南芥植物中calcuv基因组#1核小体dna甲基化水平有所提高，因此想要检测一下病毒相关基因的表达水平是否受到影响。我们按照实施例6中所述的检测目的基因表达水平的方法对calcuv的共同区(commonregion，cr)，互补链编码复制酶蛋白的基因ac1和互补链编码参与病毒复制或转录调控蛋白的基因ac4的表达水平进行检测。结果如图7e所示，rb-vtale-atkyp转基因拟南芥植物中calcuv的共同区(commonregion，cr)，互补链编码复制酶蛋白的基因ac1和互补链编码参与病毒复制或转录调控蛋白的基因ac4的表达水平至少降到载体对照植物中的1/10，说明rb-vtale-atkyp转基因拟南芥植物中rb-vtale-atkyp分子通过对组蛋白h3第9位赖氨酸进行甲基化修饰从而提高了dna甲基化修饰的水平，最终抑制了病毒相关基因的表达，使得植物获得抗病毒的能力。结论：calcuv特异的靶点rb-vtale融合表达拟南芥组蛋白甲基转移酶kyp的rb-vtale-atkyp分子在双子叶植物拟南芥中稳定表达后对入侵的calcuv基因组中的rb-vtale靶点所在核小体及其毗邻的核小体组蛋白进行甲基化修饰，使得核小体dna受到进一步的甲基化修饰，从而抑制calcuv基因组的正常复制和转录过程。表5.本发明中用于测定组蛋白和dna甲基化程度的引物序列实施例8.t4代rb-vtale-atkyp转基因拟南芥植物对甘蓝曲叶病毒(calcuv)的抗性效果、植物内源转座子mu1和休眠相关基因的表达谱的影响。待t4代转基因植物种子成熟后，挑选载体对照rb-vtale-mcs和抗性比较好的rb-vtale-atkyp转基因株系的种子萌发t4代植株，按照实施例2中所述的注射接种本氏烟草的方法将calcuv侵染性克隆接种到4周左右的转基因拟南芥轮作叶上，观察发病情况。接种calcuv21天后，t4代rb-vtale-atkyp转基因株系#1-12表现出高抗表型，植株生长健康，叶片无明显的斑驳或褪绿的症状(图8b)，而t4代载体对照rb-tale-mcs#1转基因株系表现出易感表型，花叶及明显的褪绿等症状(图8a)。利用病情指数这一指标按发病的早、中和晚期三个时间点统计的rb-vtale-atkyp转基因株系对calcuv的抗性效果如图8c所示，rb-vtale-atkyp转基因株系(#1-4和#1-12)在发病的早、中和晚期三个时间点均表现出高抗calcuv侵染的表型。为了进一步验证观察到的结果，我们按照实施例2中所述的病毒滴度测定方法以拟南芥tubulin2为内参基因(引物见表2)对接种calcuv的转基因拟南芥进行病毒滴度的测定。如图8d所示，病毒滴度分析的结果表明rb-vtale-atkyp转基因植物株系(#1-4和#1-12)中calcuv的病毒滴度明显低于载体对照植物，是其1/16，说明rb-vtale-atkyp转基因植物株系(#1-4和#1-12)对calcuv的侵染具有抗性效果，这一结果与观察到的发病症状及病情指数调查的结果相一致。在拟南芥基因组内部含有丰富的转座子，为了检测rb-vtale-atkyp转基因植物株系中atkyp的组蛋白甲基化修饰及由其引发的dna甲基化修饰是否会脱靶作用在转座子上，我们按照实施例6中检测atkyp基因表达水平的方法，检测拟南芥基因组中的转座子mu1在t4代rb-vtale-atkyp转基因植物株系(#1-4和#1-12)中的表达水平(引物序列见表4)。如果rb-vtale-atkyp转基因植物株系中rb-vtale-atkyp分子脱靶即没有结合在rb-tale靶点位置而是随机结合在mu1的启动子区，那么atkyp就会对mu1启动子区进行组蛋白甲基化修饰，进而引起dna甲基化修饰，最终抑制mu1的转录，但是实际上拟南芥基因组中的转座子mu1在rb-vtale-atkyp转基因植物株系(#1-4和#1-12)中和载体对照植物中的表达水平并无明显差异(图8e)，说明rb-vtale-atkyp转基因植物中rb-vtale-atkyp分子并未脱靶作用于拟南芥基因组的转座子mu1上。据文献(jianzhengetal.，2012，newphytologist)报道，atkyp与种子休眠呈负相关，为了进一步验证rb-vtale-atkyp转基因植物株系中rb-vtale-atkyp分子是否存在脱靶的可能，我们检测t4代rb-vtale-atkyp转基因植物株系#1-12中与种子休眠相关基因的表达水平(引物序列见表4)。如图8f所示，种子休眠相关基因(atabi3，atdog1和atnced6)在rb-vtale-atkyp转基因植物株系#1-12中和载体对照植物中的表达水平并无明显差异，说明rb-vtale-atkyp转基因植物中rb-vtale-atkyp分子并未脱靶作用于拟南芥基因组种子休眠相关基因。结论：calcuv特异的靶点rb-vtale融合表达拟南芥组蛋白甲基转移酶基因atkyp的rb-vtale-atkyp分子在双子叶植物拟南芥中稳定表达后对calcuv的高抗表型能够稳定遗传给第四代，且rb-vtale-atkyp分子并未脱靶作用于拟南芥的转座子mu1和种子休眠相关的基因上。应该理解，尽管参考其示例性的实施方案，已经对本发明进行具体地显示和描述，但是本领域的普通技术人员应该理解，在不背离由后附的权利要求所定义的本发明的精神和范围的条件下，可以在其中进行各种形式和细节的变化，可以进行各种实施方案的任意组合。序列表<110>中国科学院微生物研究所<120>一种提高植物对双生病毒抗性的方法<130>ib168044<160>30<170>patentinversion3.1<210>1<211>18<212>dna<213>ir-vtale靶点序列<400>1taatattaccggatggcc18<210>2<211>18<212>dna<213>rb-vtale靶点序列<400>2tatatattggacaccaag18<210>3<211>1875<212>dna<213>来自拟南芥的atkyp核苷酸序列<400>3atggctggaaaaaggaaacgagctaatgctcctgaccaaacagagcgaagatcgagtgtt60cgggttcagaaagtgagacagaaagcgttagatgagaaggcgcgtttagtacaggagagg120gttaagctcctcagtgacagaaagagtgaaatttgtgtcgatgacactgagttacatgag180aaagaagaggaaaatgtcgatgggagccctaaacgaagaagccctccaaagctaaccgca240atgcagaaaggaaagcagaaattgagtgtttctctgaatggtaaggacgtgaacttggaa300cctcatctcaaagtgacaaagtgtctgaggttatttaacaagcaatatctcctctgtgtc360caggctaagttgagcaggcctgatttgaagggtgtaactgagatgataaaagctaaggcg420atattgtacccaagaaaaataatcggtgaccttccaggtatagacgttggacaccgtttt480ttttcaagagctgaaatgtgtgctgtaggattccataaccattggctaaatggcattgat540tatatgtcaatggaatacgaaaaagagtatagtaactacaaattaccgcttgctgtttct600attgttatgtcgggccagtacgaggatgatctagacaatgcagatacagtgacttacact660ggacagggagggcataacttaactggtaataaacgtcagataaaggatcaacttttagaa720cgagggaatttggcgctaaagcactgctgcgaatataatgtgcctgtcagagtaactcgt780ggtcacaattgcaaaagtagctataccaaacgagtatacacttatgatggactgtacaag840gttgaaaagttctgggcacaaaagggcgtttcaggatttacagtgtataagtaccgactg900aaacgattggaggggcaaccagaactaactactgatcaggtcaactttgttgctggacgc960ataccaacgagtacttcagaaattgagggtttggtatgtgaggacatctccggagggcta1020gaatttaagggtatccccgccactaatcgtgttgatgattcaccagtttcaccaacatct1080ggtttcacatacatcaaatctttgattattgagcctaatgtcataattccaaagagttca1140actgggtgtaactgccgaggcagctgcactgactcaaagaaatgtgcatgtgctaagctt1200aatgggggtaactttccatatgttgaccttaatgatggcagattaattgagtctcgagat1260gttgtatttgaatgtggtcctcactgtgggtgtgggccaaaatgtgtcaaccgaacttct1320cagaagcgtctaagattcaatcttgaggttttccgctctgcaaagaagggttgggcagtt1380agatcatgggagtacataccagctggttcaccagtatgtgagtacataggagttgtcagg1440agaactgctgatgtggatactatctctgacaatgaatacatatttgagattgactgccaa1500cagacaatgcaaggtcttggtggaagacagagaagactaagagatgttgctgtaccaatg1560aataatggagtcagtcagagcagtgaagatgagaatgcgccagagttctgcattgatgct1620ggttcaacaggaaactttgctaggtttataaatcacagttgtgaaccaaacctatttgtt1680cagtgcgtcctgagttctcaccaggatataaggcttgcccgtgtggttcttttcgcagct1740gacaacatttccccaatgcaggagctcacttacgactatggatatgcgcttgatagcgtt1800catggaccggatgggaaggtgaagcagctcgcttgctactgtggagcgctaaattgtagg1860aaacgcctttactaa1875<210>4<211>1410<212>dna<213>来源于拟南芥的athdac19核苷酸序列<400>4atggatactggcggcaattcgctggcgtccggacctgatggtgtgaagaggaaagtttgt60tatttctatgaccctgaggtcggcaattactactatggccaaggtcatcccatgaagccc120catcgcatccgcatgacccatgccctcctcgctcactacggtctccttcagcatatgcag180gttctcaagcccttccctgcccgcgaccgtgatctctgccgcttccacgccgacgactat240gtctcttttctccgcagcattacccctgaaacccagcaagatcagattcgccaacttaag300cgcttcaatgttggtgaagactgtcccgtctttgacggcctttattccttttgccagacc360tatgctggaggatctgttggtggctctgtcaagcttaaccacggcctctgcgatattgcc420atcaactgggctggtggtctccatcacgctaagaagtgcgaggcctctggcttctgttac480gtcaatgatatcgtcttagctatcctagagctccttaagcagcatgagcgtgttctttat540gtcgatattgatatccaccacggggatggagtggaggaggcattttatgctactgacagg600gttatgactgtctcgtttcataaatttggtgattactttcccggtacaggtcacattcag660gatataggttatggtagcggaaagtactattctctcaatgtaccactggatgatggaatc720gatgatgagagctatcatctgttattcaagcccatcatggggaaagttatggaaattttc780cgaccaggggctgtggtattgcaatgtggtgctgattcattgtctggtgataggttgggg840tgctttaatctttcaatcaaaggtcatgctgagtgcgtcaaatttatgagatcgttcaat900gttcccctactgctcttgggtggtggtggttacactatccgcaatgttgcccgttgctgg960tgctacgagactggagttgcacttggagttgaagttgaagacaagatgccggagcatgaa1020tattatgaatactttggtccagactatacacttcacgttgctccaagtaacatggaaaat1080aagaattctcgtcagatgcttgaagagattcgcaatgaccttctccacaatctctctaag1140cttcagcatgctccaagtgtaccatttcaggaaagaccacctgatacagagactcccgag1200gttgatgaagaccaagaagatggggataaaagatgggatccggattcagacatggatgtt1260gatgatgaccgtaaacctataccaagcagagtaaaaagagaagctgttgaaccagataca1320aaggacaaggatggactgaaaggaattatggagcgtggaaaaggttgtgaggtggaggtg1380gatgagagtggaagcactaaggtaaaatga1410<210>5<211>4605<212>dna<213>来源于拟南芥的atmet1核苷酸序列<400>5atggtggaaaatggggctaaagctgcgaagcgaaagaagagaccacttccagagattcaa60gaggtagaagatgtacctaggacgaggagaccaaggcgtgctgcagcgtgtaccagtttc120aaggagaaatctattcgagtctgtgagaaatctgctactattgaagtaaagaaacagcag180attgtggaggaagagtttctcgcgttacggttaacggctctggaaactgatgttgaagat240cgtccaaccaggagactgaatgattttgttttgtttgattcagatggagttccacaacct300ctggagatgttggagattcatgacatattcgtttcaggtgctatcttaccttcagatgtg360tgtactgataaggagaaagagaagggtgtgaggtgtacatcgtttggacgggttgagcat420tggagtatctctggttatgaagatggttcccctgttatttggatctcaacggaattggcg480gattatgattgtcgtaaacctgctgctagctacaggaaggtttatgattacttctatgag540aaagctcgtgcttcagtggctgtgtataagaaattgtccaagtcatctggtggggatcct600gatataggtcttgaggagttacttgcggcggttgtcagatcaatgagcagtggaagcaag660tacttttctagtggtgcggcaatcatcgattttgttatatcccagggagattttatatat720aaccaactcgctggtttggatgagacagccaagaaacatgaatcaagctatgttgagatt780cctgttcttgtagctctcagagagaagagtagtaagattgacaagcctctgcagagggaa840agaaacccatctaatggtgtgaggattaaagaagtttctcaagttgcggagagcgaggcc900ttgacatctgatcaactggttgatggtactgatgatgacagaagatatgctatactctta960caagacgaagagaataggaaatctatgcaacagcccagaaaaaacagcagctcaggttct1020gcttcaaatatgttctacattaagataaatgaagatgagattgccaatgattatcctctc1080ccatcgtactataagacctccgaagaagaaacagatgaacttatactttatgatgcttcc1140tatgaggttcaatctgaacacctgcctcacaggatgcttcacaactgggctctttataac1200tctgatttacgattcatatcactggaacttctaccgatgaaacaatgtgatgatattgat1260gtcaacatttttgggtcaggtgtggtgactgatgataatggaagttggatttctttaaac1320gatcctgacagcggttctcagtcacacgatcctgatgggatgtgcatattcctcagtcaa1380attaaagaatggatgattgagtttgggagcgatgatattatctccatttctatacgaaca1440gatgtggcctggtaccgtcttgggaaaccatcaaaactttatgccccttggtggaaacct1500gttctgaaaacagcaagggttgggataagcattcttacttttcttagggtggaaagtagg1560gttgctaggctttcatttgcagatgtcacaaaaagactgtctgggttacaggcgaatgat1620aaagcttacatttcttctgaccccttggctgttgagagatatttggtcgtccatgggcaa1680attattttacagctttttgcagtttatccggacgacaatgtcaaaaggtgtccatttgtt1740gttggtcttgcaagcaaattggaggataggcaccacacaaaatggatcatcaagaagaag1800aaaatttcgctgaaggaactgaatctgaatccaagggcaggcatggcaccagtagcatcg1860aagaggaaagctatgcaagcaacaacaactcgcctggtcaacagaatttggggagagttt1920tactccaattactctccagaggatccattgcaggcgactgctgcagaaaatggggaggat1980gaggtggaagaggaaggcggaaatggggaggaagaggttgaagaggaaggtgaaaatggt2040ctcacagaggacactgtaccagaacctgttgaggttcagaagcctcatactcctaagaaa2100atccgaggcagttctggaaaaagggaaataaaatgggatggtgagagtctaggaaaaact2160tctgctggcgagcctctctatcaacaagcccttgttggaggggaaatggtggctgtaggt2220ggcgctgtcaccttggaagttgatgatccagatgaaatgccggccatctattttgtggag2280tacatgttcgaaagtacagatcactgcaaaatgttacatggtagattcttacaaagagga2340tctatgactgttctggggaatgctgctaacgagagggaactattcctgactaatgaatgc2400atgactacacagctcaaggacattaaaggagtagccagttttgagattcgatcaaggcca2460tgggggcatcagtataggaaaaagaacatcactgcggataagcttgactgggctagagca2520ttagaaagaaaagtaaaagatttgccaacagagtattactgcaaaagcttgtactcacct2580gagagagggggattctttagtcttccactaagtgatattggtcgcagttctgggttctgc2640acttcatgtaagataagggaggatgaagagaagaggtctacaattaaactaaatgtttca2700aagacaggctttttcatcaatgggattgagtattctgttgaggattttgtctatgtcaac2760cctgactctattggtgggttgaaggagggtagtaaaacttcttttaagtctgggcgaaac2820attgggttaagagcgtatgttgtttgccaattgctggaaattgttccaaaggaatctaga2880aaggctgatttgggttcctttgatgttaaagtgagaaggttttataggcctgaggatgtt2940tctgcagagaaggcctatgcttcagacatccaagaattgtatttcagccaggacacagtt3000gttctccctccaggtgctctagagggaaaatgtgaagtaagaaagaaaagtgatatgccc3060ttatcccgtgaatatccaatatcagaccatattttcttctgtgatcttttctttgacacc3120tccaaaggttctctcaagcagctgcccgccaatatgaagccaaagttctctactattaag3180gacgacacacttttaagaaagaaaaagggaaagggagtagagagtgaaattgagtctgag3240attgtcaagcctgttgagccacctaaagagattcgtctggctactctagatatttttgct3300ggttgtggtggcctgtctcatggactgaaaaaggcgggtgtatctgatgcaaagtgggcg3360attgagtatgaagagccagctgggcaggcttttaaacaaaaccatcctgagtcaacagtt3420tttgttgacaactgcaatgtgattcttagggctataatggagaaaggtggagatcaagat3480gattgtgtctctactacagaggcaaatgaattagcagctaaactaactgaggagcagaag3540agtactctgccactgcctggtcaagtggacttcatcaatggtggacctccatgtcaggga3600ttttctggtatgaacaggttcaaccaaagctcttggagtaaagttcagtgtgaaatgata3660ttagcattcttgtcctttgctgactatttccggccaaggtattttcttctggagaacgtg3720aggacctttgtgtcattcaataaagggcagacatttcagcttactttggcttcccttctc3780gaaatgggttaccaggtgagatttggaatcctggaggccggtgcatatggagtatcccaa3840tctcgtaaacgagctttcatttgggctgctgcaccagaagaagttctccctgaatggcct3900gagccgatgcatgtctttggtgttccaaagttgaaaatctcactatctcaaggtttacat3960tatgctgctgttcgtagtactgcacttggtgcccctttccgtccaatcaccgtgagagac4020acaattggtgatcttccatcagtagaaaacggagactctaggacaaacaaagagtataaa4080gaggttgcagtctcgtggttccaaaaggagataagaggaaacacgattgctctcactgat4140catatctgcaaggctatgaatgagcttaacctcattcgatgcaaattaatcccaactagg4200cctggggctgattggcatgacttgccaaagagaaaggttacgttatctgatgggcgcgta4260gaagaaatgattcctttttgtctcccaaacacagctgagcgccacaacggttggaaggga4320ctatatgggagattagattggcaaggaaactttccgacttccgtcacggatcctcagccc4380atgggtaaggttggaatgtgctttcatcctgaacagcacagaatccttacagtccgtgaa4440tgcgcccgatctcaggggtttccggatagctacgagtttgcagggaacataaatcacaag4500cacaggcagattgggaatgcagtccctccaccattggcatttgctctaggtcgtaagctc4560aaagaagccctacatctcaagaagtctcctcaacaccaaccctag4605<210>6<211>36<212>dna<213>来源于拟南芥的atsrdx核苷酸序列<400>6cttgatctggatctggaactccgtttgggtttcgct36<210>7<211>1956<212>dna<213>来源于拟南芥的atsuvh2核苷酸序列<400>7atgagtacattgttaccatttcctgacctcaacctcatgccggattctcaatcctccacc60gccggaaccacagccggcgacactgtagtcaccggaaagttagaagtgaaatcggaacca120attgaggaatggcaaactccaccatcgtctacctccgatcaatcagccaataccgatctc180atcgccgagtttattcgtatctcagagctcttccgctcggcattcaagccactgcaggtc240aaaggattagacggagtctccgtatacggattggattctggtgccatcgttgctgtaccg300gagaaagaaaaccgggaattgattgaaccgcctccaggatttaaggataatcgagtctcc360accgtcgttgtctcgccgaaattcgagagaccgagagagctagcgagaattgcaattcta420ggtcatgaacaacggaaggaactccgacaagtcatgaaacgaactaggatgacttatgag480tctcttcggattcatctaatggcggagagtatgaagaatcatgtacttggtcaagggcgt540agacggaggagcgatatggcggctgcgtatataatgagggatcggggactgtggttgaac600tatgataagcacatagttggtccagtcacaggggttgaagtaggggatatattcttctac660cgtatggaattgtgtgtgctgggcttacacggccagacacaagccgggattgattgttta720acggctgaacggagcgccactggagagcctatagctaccagcattgttgtctcaggtggt780tatgaggatgatgaagatacaggagatgttttggtttatactggtcacggtggtcaggat840catcaacacaagcagtgtgataaccagaggctagtaggtgggaatctgggaatggagaga900agtatgcactacggtatcgaggtacgtgtgattagaggtattaagtatgagaacagtata960agctcgaaggtgtatgtatatgatggtttgtataagatagtagattggtggtttgctgtg1020gggaagtctggttttggggtttttaaatttaggttagtaagaattgaagggcaaccgatg1080atgggcagtgcggtaatgaggtttgcacaaactcttagaaataagccatcgatggttagg1140cctactggttatgttagctttgacctttctaacaagaaggagaatgtacccgtgtttcta1200tataatgatgtagatggtgatcaagaaccgagacattatgagtacattgcaaaagctgtc1260tttcctcctggaatttttggtcagggggggatcagcaggactggctgcgagtgtaagctc1320tcttgtactgatgattgtctctgtgcaaggaagaacggtggtgagtttgcatatgatgat1380aacgggcatctattgaaaggaaagcatgtggtatttgaatgtggggaattctgcacttgt1440ggtccgagctgtaagagccgtgtgacacagaagggattgaggaacaggctagaggttttc1500agatcgaaggaaaccggttggggtgtcaggacacttgatctaattgaagctggtgctttc1560atatgcgaatatgcaggtgtagttgtcacgagacttcaagccgagattctgtcaatgaat1620ggggatgttatggtctatcctggtcggttcacagaccaatggcgtaactggggtgattta1680tctcaagtatacccagattttgttaggccgaattatccatctctccctccactggatttc1740tcaatggatgtgtcaaggatgaggaacgtggcttgctacattagtcacagcaaagaacca1800aatgtgatggtgcaatttgttttgcatgaccacaatcacctgatgttcccccgtgtgatg1860ctttttgcgctggagaatatctcaccgttggccgagctaagcctggattacggcttggca1920gatgaagtgaacggcaagctcgccatctgcaactag1956<210>8<211>624<212>prt<213>来源于拟南芥的atkyp氨基酸序列<400>8metalaglylysarglysargalaasnalaproaspglnthrgluarg151015argserservalargvalglnlysvalargglnlysalaleuaspglu202530lysalaargleuvalglngluargvallysleuleuserasparglys354045sergluilecysvalaspaspthrgluleuhisglulysglugluglu505560asnvalaspglyserprolysargargserproprolysleuthrala65707580metglnlysglylysglnlysleuservalserleuasnglylysasp859095valasnleugluprohisleulysvalthrlyscysleuargleuphe100105110asnlysglntyrleuleucysvalglnalalysleuserargproasp115120125leulysglyvalthrglumetilelysalalysalaileleutyrpro130135140arglysileileglyaspleuproglyileaspvalglyhisargphe145150155160pheserargalaglumetcysalavalglyphehisasnhistrpleu165170175asnglyileasptyrmetsermetglutyrglulysglutyrserasn180185190tyrlysleuproleualavalserilevalmetserglyglntyrglu195200205aspaspleuaspasnalaaspthrvalthrtyrthrglyglnglygly210215220hisasnleuthrglyasnlysargglnilelysaspglnleuleuglu225230235240argglyasnleualaleulyshiscyscysglutyrasnvalproval245250255argvalthrargglyhisasncyslyssersertyrthrlysargval260265270tyrthrtyraspglyleutyrlysvalglulysphetrpalaglnlys275280285glyvalserglyphethrvaltyrlystyrargleulysargleuglu290295300glyglnprogluleuthrthraspglnvalasnphevalalaglyarg305310315320ileprothrserthrsergluilegluglyleuvalcysgluaspile325330335serglyglyleugluphelysglyileproalathrasnargvalasp340345350aspserprovalserprothrserglyphethrtyrilelysserleu355360365ileilegluproasnvalileileprolysserserthrglycysasn370375380cysargglysercysthraspserlyslyscysalacysalalysleu385390395400asnglyglyasnpheprotyrvalaspleuasnaspglyargleuile405410415gluserargaspvalvalpheglucysglyprohiscysglycysgly420425430prolyscysvalasnargthrserglnlysargleuargpheasnleu435440445gluvalpheargseralalyslysglytrpalavalargsertrpglu450455460tyrileproalaglyserprovalcysglutyrileglyvalvalarg465470475480argthralaaspvalaspthrileseraspasnglutyrilepheglu485490495ileaspcysglnglnthrmetglnglyleuglyglyargglnargarg500505510leuargaspvalalavalprometasnasnglyvalserglnserser515520525gluaspgluasnalaprogluphecysileaspalaglyserthrgly530535540asnphealaargpheileasnhissercysgluproasnleupheval545550555560glncysvalleuserserhisglnaspileargleualaargvalval565570575leuphealaalaaspasnileserprometglngluleuthrtyrasp580585590tyrglytyralaleuaspservalhisglyproaspglylysvallys595600605glnleualacystyrcysglyalaleuasncysarglysargleutyr610615620<210>9<211>469<212>prt<213>来源于拟南芥的athdac19氨基酸序列<400>9metaspthrglyglyasnserleualaserglyproaspglyvallys151015arglysvalcystyrphetyraspprogluvalglyasntyrtyrtyr202530glyglnglyhisprometlysprohisargileargmetthrhisala354045leuleualahistyrglyleuleuglnhismetglnvalleulyspro505560pheproalaargaspargaspleucysargphehisalaaspasptyr65707580valserpheleuargserilethrprogluthrglnglnaspglnile859095argglnleulysargpheasnvalglygluaspcysprovalpheasp100105110glyleutyrserphecysglnthrtyralaglyglyservalglygly115120125servallysleuasnhisglyleucysaspilealaileasntrpala130135140glyglyleuhishisalalyslyscysglualaserglyphecystyr145150155160valasnaspilevalleualaileleugluleuleulysglnhisglu165170175argvalleutyrvalaspileaspilehishisglyaspglyvalglu180185190glualaphetyralathraspargvalmetthrvalserphehislys195200205pheglyasptyrpheproglythrglyhisileglnaspileglytyr210215220glyserglylystyrtyrserleuasnvalproleuaspaspglyile225230235240aspaspglusertyrhisleuleuphelysproilemetglylysval245250255metgluilepheargproglyalavalvalleuglncysglyalaasp260265270serleuserglyaspargleuglycyspheasnleuserilelysgly275280285hisalaglucysvallysphemetargserpheasnvalproleuleu290295300leuleuglyglyglyglytyrthrileargasnvalalaargcystrp305310315320cystyrgluthrglyvalalaleuglyvalgluvalgluasplysmet325330335progluhisglutyrtyrglutyrpheglyproasptyrthrleuhis340345350valalaproserasnmetgluasnlysasnserargglnmetleuglu355360365gluileargasnaspleuleuhisasnleuserlysleuglnhisala370375380proservalpropheglngluargproproaspthrgluthrproglu385390395400valaspgluaspglngluaspglyasplysargtrpaspproaspser405410415aspmetaspvalaspaspasparglysproileproserargvallys420425430argglualavalgluproaspthrlysasplysaspglyleulysgly435440445ilemetgluargglylysglycysgluvalgluvalaspglusergly450455460serthrlysvallys465<210>10<211>1534<212>prt<213>来源于拟南芥的atmet1氨基酸序列<400>10metvalgluasnglyalalysalaalalysarglyslysargproleu151015progluileglngluvalgluaspvalproargthrargargproarg202530argalaalaalacysthrserphelysglulysserileargvalcys354045glulysseralathrilegluvallyslysglnglnilevalgluglu505560glupheleualaleuargleuthralaleugluthraspvalgluasp65707580argprothrargargleuasnaspphevalleupheaspseraspgly859095valproglnproleuglumetleugluilehisaspilephevalser100105110glyalaileleuproseraspvalcysthrasplysglulysglulys115120125glyvalargcysthrserpheglyargvalgluhistrpserileser130135140glytyrgluaspglyserprovaliletrpileserthrgluleuala145150155160asptyraspcysarglysproalaalasertyrarglysvaltyrasp165170175tyrphetyrglulysalaargalaservalalavaltyrlyslysleu180185190serlysserserglyglyaspproaspileglyleuglugluleuleu195200205alaalavalvalargsermetserserglyserlystyrpheserser210215220glyalaalaileileaspphevalileserglnglyasppheiletyr225230235240asnglnleualaglyleuaspgluthralalyslyshisgluserser245250255tyrvalgluileprovalleuvalalaleuargglulysserserlys260265270ileasplysproleuglnarggluargasnproserasnglyvalarg275280285ilelysgluvalserglnvalalagluserglualaleuthrserasp290295300glnleuvalaspglythraspaspaspargargtyralaileleuleu305310315320glnaspglugluasnarglyssermetglnglnproarglysasnser325330335serserglyseralaserasnmetphetyrilelysileasngluasp340345350gluilealaasnasptyrproleuprosertyrtyrlysthrserglu355360365glugluthraspgluleuileleutyraspalasertyrgluvalgln370375380sergluhisleuprohisargmetleuhisasntrpalaleutyrasn385390395400seraspleuargpheileserleugluleuleuprometlysglncys405410415aspaspileaspvalasnilepheglyserglyvalvalthraspasp420425430asnglysertrpileserleuasnaspproaspserglyserglnser435440445hisaspproaspglymetcysilepheleuserglnilelysglutrp450455460metileglupheglyseraspaspileileserileserileargthr465470475480aspvalalatrptyrargleuglylysproserlysleutyralapro485490495trptrplysprovalleulysthralaargvalglyileserileleu500505510thrpheleuargvalgluserargvalalaargleuserphealaasp515520525valthrlysargleuserglyleuglnalaasnasplysalatyrile530535540serseraspproleualavalgluargtyrleuvalvalhisglygln545550555560ileileleuglnleuphealavaltyrproaspaspasnvallysarg565570575cysprophevalvalglyleualaserlysleugluasparghishis580585590thrlystrpileilelyslyslyslysileserleulysgluleuasn595600605leuasnproargalaglymetalaprovalalaserlysarglysala610615620metglnalathrthrthrargleuvalasnargiletrpglygluphe625630635640tyrserasntyrserprogluaspproleuglnalathralaalaglu645650655asnglygluaspgluvalgluglugluglyglyasnglyglugluglu660665670valgluglugluglygluasnglyleuthrgluaspthrvalproglu675680685provalgluvalglnlysprohisthrprolyslysileargglyser690695700serglylysarggluilelystrpaspglygluserleuglylysthr705710715720seralaglygluproleutyrglnglnalaleuvalglyglyglumet725730735valalavalglyglyalavalthrleugluvalaspaspproaspglu740745750metproalailetyrphevalglutyrmetphegluserthrasphis755760765cyslysmetleuhisglyargpheleuglnargglysermetthrval770775780leuglyasnalaalaasngluarggluleupheleuthrasnglucys785790795800metthrthrglnleulysaspilelysglyvalalaserphegluile805810815argserargprotrpglyhisglntyrarglyslysasnilethrala820825830asplysleuasptrpalaargalaleugluarglysvallysaspleu835840845prothrglutyrtyrcyslysserleutyrserprogluargglygly850855860phepheserleuproleuseraspileglyargserserglyphecys865870875880thrsercyslysilearggluaspgluglulysargserthrilelys885890895leuasnvalserlysthrglyphepheileasnglyileglutyrser900905910valgluaspphevaltyrvalasnproaspserileglyglyleulys915920925gluglyserlysthrserphelysserglyargasnileglyleuarg930935940alatyrvalvalcysglnleuleugluilevalprolysgluserarg945950955960lysalaaspleuglyserpheaspvallysvalargargphetyrarg965970975progluaspvalseralaglulysalatyralaseraspileglnglu980985990leutyrpheserglnaspthrvalvalleuproproglyalaleuglu99510001005glylyscysgluvalarglyslysseraspmetproleuserarg101010151020glutyrproileserasphisilephephecysaspleuphephe102510301035aspthrserlysglyserleulysglnleuproalaasnmetlys104010451050prolyspheserthrilelysaspaspthrleuleuarglyslys105510601065lysglylysglyvalglusergluileglusergluilevallys107010751080provalgluproprolysgluileargleualathrleuaspile108510901095phealaglycysglyglyleuserhisglyleulyslysalagly110011051110valseraspalalystrpalaileglutyrglugluproalagly111511201125glnalaphelysglnasnhisprogluserthrvalphevalasp113011351140asncysasnvalileleuargalailemetglulysglyglyasp114511501155glnaspaspcysvalserthrthrglualaasngluleualaala116011651170lysleuthrglugluglnlysserthrleuproleuproglygln117511801185valasppheileasnglyglyproprocysglnglyphesergly119011951200metasnargpheasnglnsersertrpserlysvalglncysglu120512101215metileleualapheleuserphealaasptyrpheargproarg122012251230tyrpheleuleugluasnvalargthrphevalserpheasnlys123512401245glyglnthrpheglnleuthrleualaserleuleuglumetgly125012551260tyrglnvalargpheglyileleuglualaglyalatyrglyval126512701275serglnserarglysargalapheiletrpalaalaalaproglu128012851290gluvalleuproglutrpproglupromethisvalpheglyval129513001305prolysleulysileserleuserglnglyleuhistyralaala131013151320valargserthralaleuglyalapropheargproilethrval132513301335argaspthrileglyaspleuproservalgluasnglyaspser134013451350argthrasnlysglutyrlysgluvalalavalsertrpphegln135513601365lysgluileargglyasnthrilealaleuthrasphisilecys137013751380lysalametasngluleuasnleuileargcyslysleuilepro138513901395thrargproglyalaasptrphisaspleuprolysarglysval140014051410thrleuseraspglyargvalgluglumetileprophecysleu141514201425proasnthralagluarghisasnglytrplysglyleutyrgly143014351440argleuasptrpglnglyasnpheprothrservalthrasppro144514501455glnprometglylysvalglymetcysphehisprogluglnhis146014651470argileleuthrvalargglucysalaargserglnglyphepro147514801485aspsertyrgluphealaglyasnileasnhislyshisarggln149014951500ileglyasnalavalproproproleualaphealaleuglyarg150515101515lysleulysglualaleuhisleulyslysserproglnhisgln152015251530pro<210>11<211>12<212>prt<213>来源于拟南芥的atsrdx氨基酸序列<400>11leuaspleuaspleugluleuargleuglypheala1510<210>12<211>651<212>prt<213>来源于拟南芥的atsuvh2氨基酸序列<400>12metserthrleuleupropheproaspleuasnleumetproaspser151015glnserserthralaglythrthralaglyaspthrvalvalthrgly202530lysleugluvallyssergluproilegluglutrpglnthrpropro354045serserthrseraspglnseralaasnthraspleuilealagluphe505560ileargilesergluleupheargseralaphelysproleuglnval65707580lysglyleuaspglyvalservaltyrglyleuaspserglyalaile859095valalavalproglulysgluasnarggluleuileglupropropro100105110glyphelysaspasnargvalserthrvalvalvalserprolysphe115120125gluargproarggluleualaargilealaileleuglyhisglugln130135140arglysgluleuargglnvalmetlysargthrargmetthrtyrglu145150155160serleuargilehisleumetalaglusermetlysasnhisvalleu165170175glyglnglyargargargargseraspmetalaalaalatyrilemet180185190argaspargglyleutrpleuasntyrasplyshisilevalglypro195200205valthrglyvalgluvalglyaspilephephetyrargmetgluleu210215220cysvalleuglyleuhisglyglnthrglnalaglyileaspcysleu225230235240thralagluargseralathrglygluproilealathrserileval245250255valserglyglytyrgluaspaspgluaspthrglyaspvalleuval260265270tyrthrglyhisglyglyglnasphisglnhislysglncysaspasn275280285glnargleuvalglyglyasnleuglymetgluargsermethistyr290295300glyilegluvalargvalileargglyilelystyrgluasnserile305310315320serserlysvaltyrvaltyraspglyleutyrlysilevalasptrp325330335trpphealavalglylysserglypheglyvalphelyspheargleu340345350valargilegluglyglnprometmetglyseralavalmetargphe355360365alaglnthrleuargasnlysprosermetvalargprothrglytyr370375380valserpheaspleuserasnlyslysgluasnvalprovalpheleu385390395400tyrasnaspvalaspglyaspglngluproarghistyrglutyrile405410415alalysalavalpheproproglyilepheglyglnglyglyileser420425430argthrglycysglucyslysleusercysthraspaspcysleucys435440445alaarglysasnglyglygluphealatyraspaspasnglyhisleu450455460leulysglylyshisvalvalpheglucysglygluphecysthrcys465470475480glyprosercyslysserargvalthrglnlysglyleuargasnarg485490495leugluvalpheargserlysgluthrglytrpglyvalargthrleu500505510aspleuileglualaglyalapheilecysglutyralaglyvalval515520525valthrargleuglnalagluileleusermetasnglyaspvalmet530535540valtyrproglyargphethraspglntrpargasntrpglyaspleu545550555560serglnvaltyrproaspphevalargproasntyrproserleupro565570575proleuaspphesermetaspvalserargmetargasnvalalacys580585590tyrileserhisserlysgluproasnvalmetvalglnphevalleu595600605hisasphisasnhisleumetpheproargvalmetleuphealaleu610615620gluasnileserproleualagluleuserleuasptyrglyleuala625630635640aspgluvalasnglylysleualailecysasn645650<210>13<211>3330<212>dna<213>ir-vtale-mcs核苷酸序列<400>13atggaccccattcgttcgcgcacgccaagtcctgcccgcgagcttctgcccggaccccaa60ccggatagggttcagccgactgcagatcgggggggggctccgcctgctggcggccccctg120gatggcttgcccgctcggcggacgatgtcccggacccggctgccatctccccctgcgccc180tcgcctgcgttctcggcgggcagcttcaacgatctgctccgtcagttcgatccgtcgctt240cttgatacatcgcttcttgattcgatgcctgccgtcggcacgccgcatacagcggctgcc300ccagcagagtgggatgaggtgcaatcgggtctgcgtgcagccgatgacccgccacccacc360gtgcgtgtcgctgtcactgccgcgcggccgccgcgcgccaagccggccccgcgacggcgt420gcggcgcaaccctccgacgcttcgccggccgcgcaggtggatctacgcacgctcggctac480agtcagcagcagcaagagaagatcaaatcgaaggtgcgttcgacagtggcgcagcaccac540gaggcactggtgggccatgggtttacacacgcgcacatcgttgcgctcagccaacacccg600gcagcgttagggaccgtcgctgtcaagtatcagcacataatcacggcgttgccagaggcg660acacacgaagacatcgttggcgtcggcaaacagtggtccggcgcacgcgccctggaggcc720ttgctcacgaaggcgggggagttgagaggtccgccgttacagttggacacaggccaactt780ctcaagattgcaaaacgtggcggcgtgaccgcagtggaggcagtgcatgcatcgcgcaat840gcactgacgggtgcccccctgaaccttactccagatcaagttgtggcaatcgcttcaaat900atcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagct960catggactcactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc1020ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggcctcactccagat1080caagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaagg1140ctccttcccgtgttgtgtcaagctcatggattaactccagatcaagttgtggcaatcgct1200tcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgt1260caagctcatggcttaactccagatcaagttgtggctatcgctagtaacggtggtggaaaa1320caggcgttagaaactgtccaaaggctccttcccgtgttgtgtcaagctcatggactcact1380ccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtc1440caaaggctccttcccgtgttgtgtcaagctcatggcctcactccagatcaagttgtggca1500atcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtg1560ttgtgtcaagctcatggattaactccagatcaagttgtggccattgcatctcatgatggt1620ggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgtcaagctcatggg1680ctcactccagatcaagttgtggccattgcatctcatgatggtggtaagcaagctctcgaa1740acagtccaaagacttctcccagtgttgtgtcaagctcatggactcactccagatcaagtt1800gtggcaatagcctctaacaatggaggtaaacaagcacttgagactgttcagagattgtta1860ccggtgttgtgtcaagctcatggcctcactccagatcaagttgtggcaatagcctctaac1920aatggaggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgtcaagct1980catggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc2040ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtcttactccagat2100caagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaagg2160ctccttcccgtgttgtgtcaagctcatggactcactccagatcaagttgtggcaatagcc2220tctaacaatggaggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgt2280caagctcatggcctcactccagatcaagttgtggcaatagcctctaacaatggaggtaaa2340caagcacttgagactgttcagagattgttaccggtgttgtgtcaagctcatggattaact2400ccagatcaagttgtggccattgcatctcatgatggtggtaagcaagctctcgaaacagtc2460caaagacttctcccagtgttgtgtcaagctcatggactgaccccggaccaggtcgtggcc2520atcgccagccatgatggcggcaagcaggcgctggagagcattgttgcccagttatctcgc2580cctgatccggcgttggccgcgttgaccaacgaccacctcgtcgccttggcctgcctcggc2640ggacgtcctgccctggatgcagtgaaaaagggattgccgcacgcgccggaattgatcaga2700agaatcaatcgccgtattcccgaacgcacgtcccatcgcgttgccgacctcgcgcacgtg2760gtgcgcgtgcttggttttttccagagccactcccacccagcgcaagcattcgatgacgcc2820atgacgcagttcgggatgagcaggcacgggttggtacagctctttcgcagagtgggcgtc2880accgaattcgaagcccgctgcggaacgctccccccagcctcgcagcgttgggaccgtatc2940ctccaggcatcagggatgaaaagggccaaaccgtcccctacttcagctcaaacgccggat3000caggcgtctttgcatgcattcgccgattcgctggagcgtgaccttgatgcgcccagccca3060atgcacgagggagatcagacgcgggcaagcagccgtaaacggtcccgatcggatcgtgct3120gtcaccggcccctccacacagcaatctttcgaggtgcgcgttcccgaacagcgcgatgcg3180ctgcatttgcccctcagctggagggtaaaacgcccgcgtaccaggatcgggggcggcctc3240ccggatcctggtacgcccatcgctgccgacctggcagcgtccagcaccgtgatgtggtct3300agattaattaaccccggggtcgactagtga3330<210>14<211>5175<212>dna<213>ir-vtale-atkyp核苷酸序列<400>14atggaccccattcgttcgcgcacgccaagtcctgcccgcgagcttctgcccggaccccaa60ccggatagggttcagccgactgcagatcgggggggggctccgcctgctggcggccccctg120gatggcttgcccgctcggcggacgatgtcccggacccggctgccatctccccctgcgccc180tcgcctgcgttctcggcgggcagcttcaacgatctgctccgtcagttcgatccgtcgctt240cttgatacatcgcttcttgattcgatgcctgccgtcggcacgccgcatacagcggctgcc300ccagcagagtgggatgaggtgcaatcgggtctgcgtgcagccgatgacccgccacccacc360gtgcgtgtcgctgtcactgccgcgcggccgccgcgcgccaagccggccccgcgacggcgt420gcggcgcaaccctccgacgcttcgccggccgcgcaggtggatctacgcacgctcggctac480agtcagcagcagcaagagaagatcaaatcgaaggtgcgttcgacagtggcgcagcaccac540gaggcactggtgggccatgggtttacacacgcgcacatcgttgcgctcagccaacacccg600gcagcgttagggaccgtcgctgtcaagtatcagcacataatcacggcgttgccagaggcg660acacacgaagacatcgttggcgtcggcaaacagtggtccggcgcacgcgccctggaggcc720ttgctcacgaaggcgggggagttgagaggtccgccgttacagttggacacaggccaactt780ctcaagattgcaaaacgtggcggcgtgaccgcagtggaggcagtgcatgcatcgcgcaat840gcactgacgggtgcccccctgaaccttactccagatcaagttgtggcaatcgcttcaaat900atcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagct960catggactcactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc1020ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggcctcactccagat1080caagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaagg1140ctccttcccgtgttgtgtcaagctcatggattaactccagatcaagttgtggcaatcgct1200tcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgt1260caagctcatggcttaactccagatcaagttgtggctatcgctagtaacggtggtggaaaa1320caggcgttagaaactgtccaaaggctccttcccgtgttgtgtcaagctcatggactcact1380ccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtc1440caaaggctccttcccgtgttgtgtcaagctcatggcctcactccagatcaagttgtggca1500atcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtg1560ttgtgtcaagctcatggattaactccagatcaagttgtggccattgcatctcatgatggt1620ggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgtcaagctcatggg1680ctcactccagatcaagttgtggccattgcatctcatgatggtggtaagcaagctctcgaa1740acagtccaaagacttctcccagtgttgtgtcaagctcatggactcactccagatcaagtt1800gtggcaatagcctctaacaatggaggtaaacaagcacttgagactgttcagagattgtta1860ccggtgttgtgtcaagctcatggcctcactccagatcaagttgtggcaatagcctctaac1920aatggaggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgtcaagct1980catggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc2040ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtcttactccagat2100caagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaagg2160ctccttcccgtgttgtgtcaagctcatggactcactccagatcaagttgtggcaatagcc2220tctaacaatggaggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgt2280caagctcatggcctcactccagatcaagttgtggcaatagcctctaacaatggaggtaaa2340caagcacttgagactgttcagagattgttaccggtgttgtgtcaagctcatggattaact2400ccagatcaagttgtggccattgcatctcatgatggtggtaagcaagctctcgaaacagtc2460caaagacttctcccagtgttgtgtcaagctcatggactgaccccggaccaggtcgtggcc2520atcgccagccatgatggcggcaagcaggcgctggagagcattgttgcccagttatctcgc2580cctgatccggcgttggccgcgttgaccaacgaccacctcgtcgccttggcctgcctcggc2640ggacgtcctgccctggatgcagtgaaaaagggattgccgcacgcgccggaattgatcaga2700agaatcaatcgccgtattcccgaacgcacgtcccatcgcgttgccgacctcgcgcacgtg2760gtgcgcgtgcttggttttttccagagccactcccacccagcgcaagcattcgatgacgcc2820atgacgcagttcgggatgagcaggcacgggttggtacagctctttcgcagagtgggcgtc2880accgaattcgaagcccgctgcggaacgctccccccagcctcgcagcgttgggaccgtatc2940ctccaggcatcagggatgaaaagggccaaaccgtcccctacttcagctcaaacgccggat3000caggcgtctttgcatgcattcgccgattcgctggagcgtgaccttgatgcgcccagccca3060atgcacgagggagatcagacgcgggcaagcagccgtaaacggtcccgatcggatcgtgct3120gtcaccggcccctccacacagcaatctttcgaggtgcgcgttcccgaacagcgcgatgcg3180ctgcatttgcccctcagctggagggtaaaacgcccgcgtaccaggatcgggggcggcctc3240ccggatcctggtacgcccatcgctgccgacctggcagcgtccagcaccgtgatgtggtct3300agtgctggaaaaaggaaacgagctaatgctcctgaccaaacagagcgaagatcgagtgtt3360cgggttcagaaagtgagacagaaagcgttagatgagaaggcgcgtttagtacaggagagg3420gttaagctcctcagtgacagaaagagtgaaatttgtgtcgatgacactgagttacatgag3480aaagaagaggaaaatgtcgatgggagccctaaacgaagaagccctccaaagctaaccgca3540atgcagaaaggaaagcagaaattgagtgtttctctgaatggtaaggacgtgaacttggaa3600cctcatctcaaagtgacaaagtgtctgaggttatttaacaagcaatatctcctctgtgtc3660caggctaagttgagcaggcctgatttgaagggtgtaactgagatgataaaagctaaggcg3720atattgtacccaagaaaaataatcggtgaccttccaggtatagacgttggacaccgtttt3780ttttcaagagctgaaatgtgtgctgtaggattccataaccattggctaaatggcattgat3840tatatgtcaatggaatacgaaaaagagtatagtaactacaaattaccgcttgctgtttct3900attgttatgtcgggccagtacgaggatgatctagacaatgcagatacagtgacttacact3960ggacagggagggcataacttaactggtaataaacgtcagataaaggatcaacttttagaa4020cgagggaatttggcgctaaagcactgctgcgaatataatgtgcctgtcagagtaactcgt4080ggtcacaattgcaaaagtagctataccaaacgagtatacacttatgatggactgtacaag4140gttgaaaagttctgggcacaaaagggcgtttcaggatttacagtgtataagtaccgactg4200aaacgattggaggggcaaccagaactaactactgatcaggtcaactttgttgctggacgc4260ataccaacgagtacttcagaaattgagggtttggtatgtgaggacatctccggagggcta4320gaatttaagggtatccccgccactaatcgtgttgatgattcaccagtttcaccaacatct4380ggtttcacatacatcaaatctttgattattgagcctaatgtcataattccaaagagttca4440actgggtgtaactgccgaggcagctgcactgactcaaagaaatgtgcatgtgctaagctt4500aatgggggtaactttccatatgttgaccttaatgatggcagattaattgagtctcgagat4560gttgtatttgaatgtggtcctcactgtgggtgtgggccaaaatgtgtcaaccgaacttct4620cagaagcgtctaagattcaatcttgaggttttccgctctgcaaagaagggttgggcagtt4680agatcatgggagtacataccagctggttcaccagtatgtgagtacataggagttgtcagg4740agaactgctgatgtggatactatctctgacaatgaatacatatttgagattgactgccaa4800cagacaatgcaaggtcttggtggaagacagagaagactaagagatgttgctgtaccaatg4860aataatggagtcagtcagagcagtgaagatgagaatgcgccagagttctgcattgatgct4920ggttcaacaggaaactttgctaggtttataaatcacagttgtgaaccaaacctatttgtt4980cagtgcgtcctgagttctcaccaggatataaggcttgcccgtgtggttcttttcgcagct5040gacaacatttccccaatgcaggagctcacttacgactatggatatgcgcttgatagcgtt5100catggaccggatgggaaggtgaagcagctcgcttgctactgtggagcgctaaattgtagg5160aaacgcctttactaa5175<210>15<211>4710<212>dna<213>ir-vtale-athdac19核苷酸序列<400>15atggaccccattcgttcgcgcacgccaagtcctgcccgcgagcttctgcccggaccccaa60ccggatagggttcagccgactgcagatcgggggggggctccgcctgctggcggccccctg120gatggcttgcccgctcggcggacgatgtcccggacccggctgccatctccccctgcgccc180tcgcctgcgttctcggcgggcagcttcaacgatctgctccgtcagttcgatccgtcgctt240cttgatacatcgcttcttgattcgatgcctgccgtcggcacgccgcatacagcggctgcc300ccagcagagtgggatgaggtgcaatcgggtctgcgtgcagccgatgacccgccacccacc360gtgcgtgtcgctgtcactgccgcgcggccgccgcgcgccaagccggccccgcgacggcgt420gcggcgcaaccctccgacgcttcgccggccgcgcaggtggatctacgcacgctcggctac480agtcagcagcagcaagagaagatcaaatcgaaggtgcgttcgacagtggcgcagcaccac540gaggcactggtgggccatgggtttacacacgcgcacatcgttgcgctcagccaacacccg600gcagcgttagggaccgtcgctgtcaagtatcagcacataatcacggcgttgccagaggcg660acacacgaagacatcgttggcgtcggcaaacagtggtccggcgcacgcgccctggaggcc720ttgctcacgaaggcgggggagttgagaggtccgccgttacagttggacacaggccaactt780ctcaagattgcaaaacgtggcggcgtgaccgcagtggaggcagtgcatgcatcgcgcaat840gcactgacgggtgcccccctgaaccttactccagatcaagttgtggcaatcgcttcaaat900atcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagct960catggactcactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc1020ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggcctcactccagat1080caagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaagg1140ctccttcccgtgttgtgtcaagctcatggattaactccagatcaagttgtggcaatcgct1200tcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgt1260caagctcatggcttaactccagatcaagttgtggctatcgctagtaacggtggtggaaaa1320caggcgttagaaactgtccaaaggctccttcccgtgttgtgtcaagctcatggactcact1380ccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtc1440caaaggctccttcccgtgttgtgtcaagctcatggcctcactccagatcaagttgtggca1500atcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtg1560ttgtgtcaagctcatggattaactccagatcaagttgtggccattgcatctcatgatggt1620ggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgtcaagctcatggg1680ctcactccagatcaagttgtggccattgcatctcatgatggtggtaagcaagctctcgaa1740acagtccaaagacttctcccagtgttgtgtcaagctcatggactcactccagatcaagtt1800gtggcaatagcctctaacaatggaggtaaacaagcacttgagactgttcagagattgtta1860ccggtgttgtgtcaagctcatggcctcactccagatcaagttgtggcaatagcctctaac1920aatggaggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgtcaagct1980catggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc2040ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtcttactccagat2100caagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaagg2160ctccttcccgtgttgtgtcaagctcatggactcactccagatcaagttgtggcaatagcc2220tctaacaatggaggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgt2280caagctcatggcctcactccagatcaagttgtggcaatagcctctaacaatggaggtaaa2340caagcacttgagactgttcagagattgttaccggtgttgtgtcaagctcatggattaact2400ccagatcaagttgtggccattgcatctcatgatggtggtaagcaagctctcgaaacagtc2460caaagacttctcccagtgttgtgtcaagctcatggactgaccccggaccaggtcgtggcc2520atcgccagccatgatggcggcaagcaggcgctggagagcattgttgcccagttatctcgc2580cctgatccggcgttggccgcgttgaccaacgaccacctcgtcgccttggcctgcctcggc2640ggacgtcctgccctggatgcagtgaaaaagggattgccgcacgcgccggaattgatcaga2700agaatcaatcgccgtattcccgaacgcacgtcccatcgcgttgccgacctcgcgcacgtg2760gtgcgcgtgcttggttttttccagagccactcccacccagcgcaagcattcgatgacgcc2820atgacgcagttcgggatgagcaggcacgggttggtacagctctttcgcagagtgggcgtc2880accgaattcgaagcccgctgcggaacgctccccccagcctcgcagcgttgggaccgtatc2940ctccaggcatcagggatgaaaagggccaaaccgtcccctacttcagctcaaacgccggat3000caggcgtctttgcatgcattcgccgattcgctggagcgtgaccttgatgcgcccagccca3060atgcacgagggagatcagacgcgggcaagcagccgtaaacggtcccgatcggatcgtgct3120gtcaccggcccctccacacagcaatctttcgaggtgcgcgttcccgaacagcgcgatgcg3180ctgcatttgcccctcagctggagggtaaaacgcccgcgtaccaggatcgggggcggcctc3240ccggatcctggtacgcccatcgctgccgacctggcagcgtccagcaccgtgatgtggtct3300agtgatactggcggcaattcgctggcgtccggacctgatggtgtgaagaggaaagtttgt3360tatttctatgaccctgaggtcggcaattactactatggccaaggtcatcccatgaagccc3420catcgcatccgcatgacccatgccctcctcgctcactacggtctccttcagcatatgcag3480gttctcaagcccttccctgcccgcgaccgtgatctctgccgcttccacgccgacgactat3540gtctcttttctccgcagcattacccctgaaacccagcaagatcagattcgccaacttaag3600cgcttcaatgttggtgaagactgtcccgtctttgacggcctttattccttttgccagacc3660tatgctggaggatctgttggtggctctgtcaagcttaaccacggcctctgcgatattgcc3720atcaactgggctggtggtctccatcacgctaagaagtgcgaggcctctggcttctgttac3780gtcaatgatatcgtcttagctatcctagagctccttaagcagcatgagcgtgttctttat3840gtcgatattgatatccaccacggggatggagtggaggaggcattttatgctactgacagg3900gttatgactgtctcgtttcataaatttggtgattactttcccggtacaggtcacattcag3960gatataggttatggtagcggaaagtactattctctcaatgtaccactggatgatggaatc4020gatgatgagagctatcatctgttattcaagcccatcatggggaaagttatggaaattttc4080cgaccaggggctgtggtattgcaatgtggtgctgattcattgtctggtgataggttgggg4140tgctttaatctttcaatcaaaggtcatgctgagtgcgtcaaatttatgagatcgttcaat4200gttcccctactgctcttgggtggtggtggttacactatccgcaatgttgcccgttgctgg4260tgctacgagactggagttgcacttggagttgaagttgaagacaagatgccggagcatgaa4320tattatgaatactttggtccagactatacacttcacgttgctccaagtaacatggaaaat4380aagaattctcgtcagatgcttgaagagattcgcaatgaccttctccacaatctctctaag4440cttcagcatgctccaagtgtaccatttcaggaaagaccacctgatacagagactcccgag4500gttgatgaagaccaagaagatggggataaaagatgggatccggattcagacatggatgtt4560gatgatgaccgtaaacctataccaagcagagtaaaaagagaagctgttgaaccagataca4620aaggacaaggatggactgaaaggaattatggagcgtggaaaaggttgtgaggtggaggtg4680gatgagagtggaagcactaaggtaaaatga4710<210>16<211>7905<212>dna<213>ir-vtale-atmet1核苷酸序列<400>16atggaccccattcgttcgcgcacgccaagtcctgcccgcgagcttctgcccggaccccaa60ccggatagggttcagccgactgcagatcgggggggggctccgcctgctggcggccccctg120gatggcttgcccgctcggcggacgatgtcccggacccggctgccatctccccctgcgccc180tcgcctgcgttctcggcgggcagcttcaacgatctgctccgtcagttcgatccgtcgctt240cttgatacatcgcttcttgattcgatgcctgccgtcggcacgccgcatacagcggctgcc300ccagcagagtgggatgaggtgcaatcgggtctgcgtgcagccgatgacccgccacccacc360gtgcgtgtcgctgtcactgccgcgcggccgccgcgcgccaagccggccccgcgacggcgt420gcggcgcaaccctccgacgcttcgccggccgcgcaggtggatctacgcacgctcggctac480agtcagcagcagcaagagaagatcaaatcgaaggtgcgttcgacagtggcgcagcaccac540gaggcactggtgggccatgggtttacacacgcgcacatcgttgcgctcagccaacacccg600gcagcgttagggaccgtcgctgtcaagtatcagcacataatcacggcgttgccagaggcg660acacacgaagacatcgttggcgtcggcaaacagtggtccggcgcacgcgccctggaggcc720ttgctcacgaaggcgggggagttgagaggtccgccgttacagttggacacaggccaactt780ctcaagattgcaaaacgtggcggcgtgaccgcagtggaggcagtgcatgcatcgcgcaat840gcactgacgggtgcccccctgaaccttactccagatcaagttgtggcaatcgcttcaaat900atcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagct960catggactcactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc1020ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggcctcactccagat1080caagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaagg1140ctccttcccgtgttgtgtcaagctcatggattaactccagatcaagttgtggcaatcgct1200tcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgt1260caagctcatggcttaactccagatcaagttgtggctatcgctagtaacggtggtggaaaa1320caggcgttagaaactgtccaaaggctccttcccgtgttgtgtcaagctcatggactcact1380ccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtc1440caaaggctccttcccgtgttgtgtcaagctcatggcctcactccagatcaagttgtggca1500atcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtg1560ttgtgtcaagctcatggattaactccagatcaagttgtggccattgcatctcatgatggt1620ggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgtcaagctcatggg1680ctcactccagatcaagttgtggccattgcatctcatgatggtggtaagcaagctctcgaa1740acagtccaaagacttctcccagtgttgtgtcaagctcatggactcactccagatcaagtt1800gtggcaatagcctctaacaatggaggtaaacaagcacttgagactgttcagagattgtta1860ccggtgttgtgtcaagctcatggcctcactccagatcaagttgtggcaatagcctctaac1920aatggaggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgtcaagct1980catggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc2040ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtcttactccagat2100caagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaagg2160ctccttcccgtgttgtgtcaagctcatggactcactccagatcaagttgtggcaatagcc2220tctaacaatggaggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgt2280caagctcatggcctcactccagatcaagttgtggcaatagcctctaacaatggaggtaaa2340caagcacttgagactgttcagagattgttaccggtgttgtgtcaagctcatggattaact2400ccagatcaagttgtggccattgcatctcatgatggtggtaagcaagctctcgaaacagtc2460caaagacttctcccagtgttgtgtcaagctcatggactgaccccggaccaggtcgtggcc2520atcgccagccatgatggcggcaagcaggcgctggagagcattgttgcccagttatctcgc2580cctgatccggcgttggccgcgttgaccaacgaccacctcgtcgccttggcctgcctcggc2640ggacgtcctgccctggatgcagtgaaaaagggattgccgcacgcgccggaattgatcaga2700agaatcaatcgccgtattcccgaacgcacgtcccatcgcgttgccgacctcgcgcacgtg2760gtgcgcgtgcttggttttttccagagccactcccacccagcgcaagcattcgatgacgcc2820atgacgcagttcgggatgagcaggcacgggttggtacagctctttcgcagagtgggcgtc2880accgaattcgaagcccgctgcggaacgctccccccagcctcgcagcgttgggaccgtatc2940ctccaggcatcagggatgaaaagggccaaaccgtcccctacttcagctcaaacgccggat3000caggcgtctttgcatgcattcgccgattcgctggagcgtgaccttgatgcgcccagccca3060atgcacgagggagatcagacgcgggcaagcagccgtaaacggtcccgatcggatcgtgct3120gtcaccggcccctccacacagcaatctttcgaggtgcgcgttcccgaacagcgcgatgcg3180ctgcatttgcccctcagctggagggtaaaacgcccgcgtaccaggatcgggggcggcctc3240ccggatcctggtacgcccatcgctgccgacctggcagcgtccagcaccgtgatgtggtct3300agtgtggaaaatggggctaaagctgcgaagcgaaagaagagaccacttccagagattcaa3360gaggtagaagatgtacctaggacgaggagaccaaggcgtgctgcagcgtgtaccagtttc3420aaggagaaatctattcgagtctgtgagaaatctgctactattgaagtaaagaaacagcag3480attgtggaggaagagtttctcgcgttacggttaacggctctggaaactgatgttgaagat3540cgtccaaccaggagactgaatgattttgttttgtttgattcagatggagttccacaacct3600ctggagatgttggagattcatgacatattcgtttcaggtgctatcttaccttcagatgtg3660tgtactgataaggagaaagagaagggtgtgaggtgtacatcgtttggacgggttgagcat3720tggagtatctctggttatgaagatggttcccctgttatttggatctcaacggaattggcg3780gattatgattgtcgtaaacctgctgctagctacaggaaggtttatgattacttctatgag3840aaagctcgtgcttcagtggctgtgtataagaaattgtccaagtcatctggtggggatcct3900gatataggtcttgaggagttacttgcggcggttgtcagatcaatgagcagtggaagcaag3960tacttttctagtggtgcggcaatcatcgattttgttatatcccagggagattttatatat4020aaccaactcgctggtttggatgagacagccaagaaacatgaatcaagctatgttgagatt4080cctgttcttgtagctctcagagagaagagtagtaagattgacaagcctctgcagagggaa4140agaaacccatctaatggtgtgaggattaaagaagtttctcaagttgcggagagcgaggcc4200ttgacatctgatcaactggttgatggtactgatgatgacagaagatatgctatactctta4260caagacgaagagaataggaaatctatgcaacagcccagaaaaaacagcagctcaggttct4320gcttcaaatatgttctacattaagataaatgaagatgagattgccaatgattatcctctc4380ccatcgtactataagacctccgaagaagaaacagatgaacttatactttatgatgcttcc4440tatgaggttcaatctgaacacctgcctcacaggatgcttcacaactgggctctttataac4500tctgatttacgattcatatcactggaacttctaccgatgaaacaatgtgatgatattgat4560gtcaacatttttgggtcaggtgtggtgactgatgataatggaagttggatttctttaaac4620gatcctgacagcggttctcagtcacacgatcctgatgggatgtgcatattcctcagtcaa4680attaaagaatggatgattgagtttgggagcgatgatattatctccatttctatacgaaca4740gatgtggcctggtaccgtcttgggaaaccatcaaaactttatgccccttggtggaaacct4800gttctgaaaacagcaagggttgggataagcattcttacttttcttagggtggaaagtagg4860gttgctaggctttcatttgcagatgtcacaaaaagactgtctgggttacaggcgaatgat4920aaagcttacatttcttctgaccccttggctgttgagagatatttggtcgtccatgggcaa4980attattttacagctttttgcagtttatccggacgacaatgtcaaaaggtgtccatttgtt5040gttggtcttgcaagcaaattggaggataggcaccacacaaaatggatcatcaagaagaag5100aaaatttcgctgaaggaactgaatctgaatccaagggcaggcatggcaccagtagcatcg5160aagaggaaagctatgcaagcaacaacaactcgcctggtcaacagaatttggggagagttt5220tactccaattactctccagaggatccattgcaggcgactgctgcagaaaatggggaggat5280gaggtggaagaggaaggcggaaatggggaggaagaggttgaagaggaaggtgaaaatggt5340ctcacagaggacactgtaccagaacctgttgaggttcagaagcctcatactcctaagaaa5400atccgaggcagttctggaaaaagggaaataaaatgggatggtgagagtctaggaaaaact5460tctgctggcgagcctctctatcaacaagcccttgttggaggggaaatggtggctgtaggt5520ggcgctgtcaccttggaagttgatgatccagatgaaatgccggccatctattttgtggag5580tacatgttcgaaagtacagatcactgcaaaatgttacatggtagattcttacaaagagga5640tctatgactgttctggggaatgctgctaacgagagggaactattcctgactaatgaatgc5700atgactacacagctcaaggacattaaaggagtagccagttttgagattcgatcaaggcca5760tgggggcatcagtataggaaaaagaacatcactgcggataagcttgactgggctagagca5820ttagaaagaaaagtaaaagatttgccaacagagtattactgcaaaagcttgtactcacct5880gagagagggggattctttagtcttccactaagtgatattggtcgcagttctgggttctgc5940acttcatgtaagataagggaggatgaagagaagaggtctacaattaaactaaatgtttca6000aagacaggctttttcatcaatgggattgagtattctgttgaggattttgtctatgtcaac6060cctgactctattggtgggttgaaggagggtagtaaaacttcttttaagtctgggcgaaac6120attgggttaagagcgtatgttgtttgccaattgctggaaattgttccaaaggaatctaga6180aaggctgatttgggttcctttgatgttaaagtgagaaggttttataggcctgaggatgtt6240tctgcagagaaggcctatgcttcagacatccaagaattgtatttcagccaggacacagtt6300gttctccctccaggtgctctagagggaaaatgtgaagtaagaaagaaaagtgatatgccc6360ttatcccgtgaatatccaatatcagaccatattttcttctgtgatcttttctttgacacc6420tccaaaggttctctcaagcagctgcccgccaatatgaagccaaagttctctactattaag6480gacgacacacttttaagaaagaaaaagggaaagggagtagagagtgaaattgagtctgag6540attgtcaagcctgttgagccacctaaagagattcgtctggctactctagatatttttgct6600ggttgtggtggcctgtctcatggactgaaaaaggcgggtgtatctgatgcaaagtgggcg6660attgagtatgaagagccagctgggcaggcttttaaacaaaaccatcctgagtcaacagtt6720tttgttgacaactgcaatgtgattcttagggctataatggagaaaggtggagatcaagat6780gattgtgtctctactacagaggcaaatgaattagcagctaaactaactgaggagcagaag6840agtactctgccactgcctggtcaagtggacttcatcaatggtggacctccatgtcaggga6900ttttctggtatgaacaggttcaaccaaagctcttggagtaaagttcagtgtgaaatgata6960ttagcattcttgtcctttgctgactatttccggccaaggtattttcttctggagaacgtg7020aggacctttgtgtcattcaataaagggcagacatttcagcttactttggcttcccttctc7080gaaatgggttaccaggtgagatttggaatcctggaggccggtgcatatggagtatcccaa7140tctcgtaaacgagctttcatttgggctgctgcaccagaagaagttctccctgaatggcct7200gagccgatgcatgtctttggtgttccaaagttgaaaatctcactatctcaaggtttacat7260tatgctgctgttcgtagtactgcacttggtgcccctttccgtccaatcaccgtgagagac7320acaattggtgatcttccatcagtagaaaacggagactctaggacaaacaaagagtataaa7380gaggttgcagtctcgtggttccaaaaggagataagaggaaacacgattgctctcactgat7440catatctgcaaggctatgaatgagcttaacctcattcgatgcaaattaatcccaactagg7500cctggggctgattggcatgacttgccaaagagaaaggttacgttatctgatgggcgcgta7560gaagaaatgattcctttttgtctcccaaacacagctgagcgccacaacggttggaaggga7620ctatatgggagattagattggcaaggaaactttccgacttccgtcacggatcctcagccc7680atgggtaaggttggaatgtgctttcatcctgaacagcacagaatccttacagtccgtgaa7740tgcgcccgatctcaggggtttccggatagctacgagtttgcagggaacataaatcacaag7800cacaggcagattgggaatgcagtccctccaccattggcatttgctctaggtcgtaagctc7860aaagaagccctacatctcaagaagtctcctcaacaccaaccctag7905<210>17<211>1724<212>prt<213>ir-vtale-atkyp氨基酸序列<400>17metaspproileargserargthrproserproalaarggluleuleu151015proglyproglnproaspargvalglnprothralaaspargglygly202530alaproproalaglyglyproleuaspglyleuproalaargargthr354045metserargthrargleuproserproproalaproserproalaphe505560seralaglyserpheasnaspleuleuargglnpheaspproserleu65707580leuaspthrserleuleuaspsermetproalavalglythrprohis859095thralaalaalaproalaglutrpaspgluvalglnserglyleuarg100105110alaalaaspaspproproprothrvalargvalalavalthralaala115120125argproproargalalysproalaproargargargalaalaglnpro130135140seraspalaserproalaalaglnvalaspleuargthrleuglytyr145150155160serglnglnglnglnglulysilelysserlysvalargserthrval165170175alaglnhishisglualaleuvalglyhisglyphethrhisalahis180185190ilevalalaleuserglnhisproalaalaleuglythrvalalaval195200205lystyrglnhisileilethralaleuproglualathrhisgluasp210215220ilevalglyvalglylysglntrpserglyalaargalaleugluala225230235240leuleuthrlysalaglygluleuargglyproproleuglnleuasp245250255thrglyglnleuleulysilealalysargglyglyvalthralaval260265270glualavalhisalaserargasnalaleuthrglyalaproleuasn275280285leuthrproaspglnvalvalalailealaserasnileglyglylys290295300glnalaleugluthrvalglnargleuleuprovalleucysglnala305310315320hisglyleuthrproaspglnvalvalalailealaserasnilegly325330335glylysglnalaleugluthrvalglnargleuleuprovalleucys340345350glnalahisglyleuthrproaspglnvalvalalailealaserasn355360365glyglyglylysglnalaleugluthrvalglnargleuleuproval370375380leucysglnalahisglyleuthrproaspglnvalvalalaileala385390395400serasnileglyglylysglnalaleugluthrvalglnargleuleu405410415provalleucysglnalahisglyleuthrproaspglnvalvalala420425430ilealaserasnglyglyglylysglnalaleugluthrvalglnarg435440445leuleuprovalleucysglnalahisglyleuthrproaspglnval450455460valalailealaserasnglyglyglylysglnalaleugluthrval465470475480glnargleuleuprovalleucysglnalahisglyleuthrproasp485490495glnvalvalalailealaserasnileglyglylysglnalaleuglu500505510thrvalglnargleuleuprovalleucysglnalahisglyleuthr515520525proaspglnvalvalalailealaserhisaspglyglylysglnala530535540leugluthrvalglnargleuleuprovalleucysglnalahisgly545550555560leuthrproaspglnvalvalalailealaserhisaspglyglylys565570575glnalaleugluthrvalglnargleuleuprovalleucysglnala580585590hisglyleuthrproaspglnvalvalalailealaserasnasngly595600605glylysglnalaleugluthrvalglnargleuleuprovalleucys610615620glnalahisglyleuthrproaspglnvalvalalailealaserasn625630635640asnglyglylysglnalaleugluthrvalglnargleuleuproval645650655leucysglnalahisglyleuthrproaspglnvalvalalaileala660665670serasnileglyglylysglnalaleugluthrvalglnargleuleu675680685provalleucysglnalahisglyleuthrproaspglnvalvalala690695700ilealaserasnglyglyglylysglnalaleugluthrvalglnarg705710715720leuleuprovalleucysglnalahisglyleuthrproaspglnval725730735valalailealaserasnasnglyglylysglnalaleugluthrval740745750glnargleuleuprovalleucysglnalahisglyleuthrproasp755760765glnvalvalalailealaserasnasnglyglylysglnalaleuglu770775780thrvalglnargleuleuprovalleucysglnalahisglyleuthr785790795800proaspglnvalvalalailealaserhisaspglyglylysglnala805810815leugluthrvalglnargleuleuprovalleucysglnalahisgly820825830leuthrproaspglnvalvalalailealaserhisaspglyglylys835840845glnalaleugluserilevalalaglnleuserargproaspproala850855860leualaalaleuthrasnasphisleuvalalaleualacysleugly865870875880glyargproalaleuaspalavallyslysglyleuprohisalapro885890895gluleuileargargileasnargargileprogluargthrserhis900905910argvalalaaspleualahisvalvalargvalleuglyphephegln915920925serhisserhisproalaglnalapheaspaspalametthrglnphe930935940glymetserarghisglyleuvalglnleupheargargvalglyval945950955960thrglupheglualaargcysglythrleuproproalaserglnarg965970975trpaspargileleuglnalaserglymetlysargalalysproser980985990prothrseralaglnthrproaspglnalaserleuhisalapheala99510001005aspserleugluargaspleuaspalaproserpromethisglu101010151020glyaspglnthrargalaserserarglysargserargserasp102510301035argalavalthrglyproserthrglnglnserphegluvalarg104010451050valprogluglnargaspalaleuhisleuproleusertrparg105510601065vallysargproargthrargileglyglyglyleuproasppro107010751080glythrproilealaalaaspleualaalaserserthrvalmet108510901095trpserseralaglylysarglysargalaasnalaproaspgln110011051110thrgluargargserservalargvalglnlysvalargglnlys111511201125alaleuaspglulysalaargleuvalglngluargvallysleu113011351140leuserasparglyssergluilecysvalaspaspthrgluleu114511501155hisglulysgluglugluasnvalaspglyserprolysargarg116011651170serproprolysleuthralametglnlysglylysglnlysleu117511801185servalserleuasnglylysaspvalasnleugluprohisleu119011951200lysvalthrlyscysleuargleupheasnlysglntyrleuleu120512101215cysvalglnalalysleuserargproaspleulysglyvalthr122012251230glumetilelysalalysalaileleutyrproarglysileile123512401245glyaspleuproglyileaspvalglyhisargphepheserarg125012551260alaglumetcysalavalglyphehisasnhistrpleuasngly126512701275ileasptyrmetsermetglutyrglulysglutyrserasntyr128012851290lysleuproleualavalserilevalmetserglyglntyrglu129513001305aspaspleuaspasnalaaspthrvalthrtyrthrglyglngly131013151320glyhisasnleuthrglyasnlysargglnilelysaspglnleu132513301335leugluargglyasnleualaleulyshiscyscysglutyrasn134013451350valprovalargvalthrargglyhisasncyslyssersertyr135513601365thrlysargvaltyrthrtyraspglyleutyrlysvalglulys137013751380phetrpalaglnlysglyvalserglyphethrvaltyrlystyr138513901395argleulysargleugluglyglnprogluleuthrthraspgln140014051410valasnphevalalaglyargileprothrserthrsergluile141514201425gluglyleuvalcysgluaspileserglyglyleugluphelys143014351440glyileproalathrasnargvalaspaspserprovalserpro144514501455thrserglyphethrtyrilelysserleuileilegluproasn146014651470valileileprolysserserthrglycysasncysargglyser147514801485cysthraspserlyslyscysalacysalalysleuasnglygly149014951500asnpheprotyrvalaspleuasnaspglyargleuilegluser150515101515argaspvalvalpheglucysglyprohiscysglycysglypro152015251530lyscysvalasnargthrserglnlysargleuargpheasnleu153515401545gluvalpheargseralalyslysglytrpalavalargsertrp155015551560glutyrileproalaglyserprovalcysglutyrileglyval156515701575valargargthralaaspvalaspthrileseraspasnglutyr158015851590ilephegluileaspcysglnglnthrmetglnglyleuglygly159516001605argglnargargleuargaspvalalavalprometasnasngly161016151620valserglnsersergluaspgluasnalaprogluphecysile162516301635aspalaglyserthrglyasnphealaargpheileasnhisser164016451650cysgluproasnleuphevalglncysvalleuserserhisgln165516601665aspileargleualaargvalvalleuphealaalaaspasnile167016751680serprometglngluleuthrtyrasptyrglytyralaleuasp168516901695servalhisglyproaspglylysvallysglnleualacystyr170017051710cysglyalaleuasncysarglysargleutyr17151720<210>18<211>1569<212>prt<213>ir-vtale-athdac19氨基酸序列<400>18metaspproileargserargthrproserproalaarggluleuleu151015proglyproglnproaspargvalglnprothralaaspargglygly202530alaproproalaglyglyproleuaspglyleuproalaargargthr354045metserargthrargleuproserproproalaproserproalaphe505560seralaglyserpheasnaspleuleuargglnpheaspproserleu65707580leuaspthrserleuleuaspsermetproalavalglythrprohis859095thralaalaalaproalaglutrpaspgluvalglnserglyleuarg100105110alaalaaspaspproproprothrvalargvalalavalthralaala115120125argproproargalalysproalaproargargargalaalaglnpro130135140seraspalaserproalaalaglnvalaspleuargthrleuglytyr145150155160serglnglnglnglnglulysilelysserlysvalargserthrval165170175alaglnhishisglualaleuvalglyhisglyphethrhisalahis180185190ilevalalaleuserglnhisproalaalaleuglythrvalalaval195200205lystyrglnhisileilethralaleuproglualathrhisgluasp210215220ilevalglyvalglylysglntrpserglyalaargalaleugluala225230235240leuleuthrlysalaglygluleuargglyproproleuglnleuasp245250255thrglyglnleuleulysilealalysargglyglyvalthralaval260265270glualavalhisalaserargasnalaleuthrglyalaproleuasn275280285leuthrproaspglnvalvalalailealaserasnileglyglylys290295300glnalaleugluthrvalglnargleuleuprovalleucysglnala305310315320hisglyleuthrproaspglnvalvalalailealaserasnilegly325330335glylysglnalaleugluthrvalglnargleuleuprovalleucys340345350glnalahisglyleuthrproaspglnvalvalalailealaserasn355360365glyglyglylysglnalaleugluthrvalglnargleuleuproval370375380leucysglnalahisglyleuthrproaspglnvalvalalaileala385390395400serasnileglyglylysglnalaleugluthrvalglnargleuleu405410415provalleucysglnalahisglyleuthrproaspglnvalvalala420425430ilealaserasnglyglyglylysglnalaleugluthrvalglnarg435440445leuleuprovalleucysglnalahisglyleuthrproaspglnval450455460valalailealaserasnglyglyglylysglnalaleugluthrval465470475480glnargleuleuprovalleucysglnalahisglyleuthrproasp485490495glnvalvalalailealaserasnileglyglylysglnalaleuglu500505510thrvalglnargleuleuprovalleucysglnalahisglyleuthr515520525proaspglnvalvalalailealaserhisaspglyglylysglnala530535540leugluthrvalglnargleuleuprovalleucysglnalahisgly545550555560leuthrproaspglnvalvalalailealaserhisaspglyglylys565570575glnalaleugluthrvalglnargleuleuprovalleucysglnala580585590hisglyleuthrproaspglnvalvalalailealaserasnasngly595600605glylysglnalaleugluthrvalglnargleuleuprovalleucys610615620glnalahisglyleuthrproaspglnvalvalalailealaserasn625630635640asnglyglylysglnalaleugluthrvalglnargleuleuproval645650655leucysglnalahisglyleuthrproaspglnvalvalalaileala660665670serasnileglyglylysglnalaleugluthrvalglnargleuleu675680685provalleucysglnalahisglyleuthrproaspglnvalvalala690695700ilealaserasnglyglyglylysglnalaleugluthrvalglnarg705710715720leuleuprovalleucysglnalahisglyleuthrproaspglnval725730735valalailealaserasnasnglyglylysglnalaleugluthrval740745750glnargleuleuprovalleucysglnalahisglyleuthrproasp755760765glnvalvalalailealaserasnasnglyglylysglnalaleuglu770775780thrvalglnargleuleuprovalleucysglnalahisglyleuthr785790795800proaspglnvalvalalailealaserhisaspglyglylysglnala805810815leugluthrvalglnargleuleuprovalleucysglnalahisgly820825830leuthrproaspglnvalvalalailealaserhisaspglyglylys835840845glnalaleugluserilevalalaglnleuserargproaspproala850855860leualaalaleuthrasnasphisleuvalalaleualacysleugly865870875880glyargproalaleuaspalavallyslysglyleuprohisalapro885890895gluleuileargargileasnargargileprogluargthrserhis900905910argvalalaaspleualahisvalvalargvalleuglyphephegln915920925serhisserhisproalaglnalapheaspaspalametthrglnphe930935940glymetserarghisglyleuvalglnleupheargargvalglyval945950955960thrglupheglualaargcysglythrleuproproalaserglnarg965970975trpaspargileleuglnalaserglymetlysargalalysproser980985990prothrseralaglnthrproaspglnalaserleuhisalapheala99510001005aspserleugluargaspleuaspalaproserpromethisglu101010151020glyaspglnthrargalaserserarglysargserargserasp102510301035argalavalthrglyproserthrglnglnserphegluvalarg104010451050valprogluglnargaspalaleuhisleuproleusertrparg105510601065vallysargproargthrargileglyglyglyleuproasppro107010751080glythrproilealaalaaspleualaalaserserthrvalmet108510901095trpserseraspthrglyglyasnserleualaserglyproasp110011051110glyvallysarglysvalcystyrphetyraspprogluvalgly111511201125asntyrtyrtyrglyglnglyhisprometlysprohisargile113011351140argmetthrhisalaleuleualahistyrglyleuleuglnhis114511501155metglnvalleulyspropheproalaargaspargaspleucys116011651170argphehisalaaspasptyrvalserpheleuargserilethr117511801185progluthrglnglnaspglnileargglnleulysargpheasn119011951200valglygluaspcysprovalpheaspglyleutyrserphecys120512101215glnthrtyralaglyglyservalglyglyservallysleuasn122012251230hisglyleucysaspilealaileasntrpalaglyglyleuhis123512401245hisalalyslyscysglualaserglyphecystyrvalasnasp125012551260ilevalleualaileleugluleuleulysglnhisgluargval126512701275leutyrvalaspileaspilehishisglyaspglyvalgluglu128012851290alaphetyralathraspargvalmetthrvalserphehislys129513001305pheglyasptyrpheproglythrglyhisileglnaspilegly131013151320tyrglyserglylystyrtyrserleuasnvalproleuaspasp132513301335glyileaspaspglusertyrhisleuleuphelysproilemet134013451350glylysvalmetgluilepheargproglyalavalvalleugln135513601365cysglyalaaspserleuserglyaspargleuglycyspheasn137013751380leuserilelysglyhisalaglucysvallysphemetargser138513901395pheasnvalproleuleuleuleuglyglyglyglytyrthrile140014051410argasnvalalaargcystrpcystyrgluthrglyvalalaleu141514201425glyvalgluvalgluasplysmetprogluhisglutyrtyrglu143014351440tyrpheglyproasptyrthrleuhisvalalaproserasnmet144514501455gluasnlysasnserargglnmetleuglugluileargasnasp146014651470leuleuhisasnleuserlysleuglnhisalaproservalpro147514801485pheglngluargproproaspthrgluthrprogluvalaspglu149014951500aspglngluaspglyasplysargtrpaspproaspseraspmet150515101515aspvalaspaspasparglysproileproserargvallysarg152015251530glualavalgluproaspthrlysasplysaspglyleulysgly153515401545ilemetgluargglylysglycysgluvalgluvalaspgluser155015551560glyserthrlysvallys1565<210>19<211>2634<212>prt<213>ir-vtale-atmet1氨基酸序列<400>19metaspproileargserargthrproserproalaarggluleuleu151015proglyproglnproaspargvalglnprothralaaspargglygly202530alaproproalaglyglyproleuaspglyleuproalaargargthr354045metserargthrargleuproserproproalaproserproalaphe505560seralaglyserpheasnaspleuleuargglnpheaspproserleu65707580leuaspthrserleuleuaspsermetproalavalglythrprohis859095thralaalaalaproalaglutrpaspgluvalglnserglyleuarg100105110alaalaaspaspproproprothrvalargvalalavalthralaala115120125argproproargalalysproalaproargargargalaalaglnpro130135140seraspalaserproalaalaglnvalaspleuargthrleuglytyr145150155160serglnglnglnglnglulysilelysserlysvalargserthrval165170175alaglnhishisglualaleuvalglyhisglyphethrhisalahis180185190ilevalalaleuserglnhisproalaalaleuglythrvalalaval195200205lystyrglnhisileilethralaleuproglualathrhisgluasp210215220ilevalglyvalglylysglntrpserglyalaargalaleugluala225230235240leuleuthrlysalaglygluleuargglyproproleuglnleuasp245250255thrglyglnleuleulysilealalysargglyglyvalthralaval260265270glualavalhisalaserargasnalaleuthrglyalaproleuasn275280285leuthrproaspglnvalvalalailealaserasnileglyglylys290295300glnalaleugluthrvalglnargleuleuprovalleucysglnala305310315320hisglyleuthrproaspglnvalvalalailealaserasnilegly325330335glylysglnalaleugluthrvalglnargleuleuprovalleucys340345350glnalahisglyleuthrproaspglnvalvalalailealaserasn355360365glyglyglylysglnalaleugluthrvalglnargleuleuproval370375380leucysglnalahisglyleuthrproaspglnvalvalalaileala385390395400serasnileglyglylysglnalaleugluthrvalglnargleuleu405410415provalleucysglnalahisglyleuthrproaspglnvalvalala420425430ilealaserasnglyglyglylysglnalaleugluthrvalglnarg435440445leuleuprovalleucysglnalahisglyleuthrproaspglnval450455460valalailealaserasnglyglyglylysglnalaleugluthrval465470475480glnargleuleuprovalleucysglnalahisglyleuthrproasp485490495glnvalvalalailealaserasnileglyglylysglnalaleuglu500505510thrvalglnargleuleuprovalleucysglnalahisglyleuthr515520525proaspglnvalvalalailealaserhisaspglyglylysglnala530535540leugluthrvalglnargleuleuprovalleucysglnalahisgly545550555560leuthrproaspglnvalvalalailealaserhisaspglyglylys565570575glnalaleugluthrvalglnargleuleuprovalleucysglnala580585590hisglyleuthrproaspglnvalvalalailealaserasnasngly595600605glylysglnalaleugluthrvalglnargleuleuprovalleucys610615620glnalahisglyleuthrproaspglnvalvalalailealaserasn625630635640asnglyglylysglnalaleugluthrvalglnargleuleuproval645650655leucysglnalahisglyleuthrproaspglnvalvalalaileala660665670serasnileglyglylysglnalaleugluthrvalglnargleuleu675680685provalleucysglnalahisglyleuthrproaspglnvalvalala690695700ilealaserasnglyglyglylysglnalaleugluthrvalglnarg705710715720leuleuprovalleucysglnalahisglyleuthrproaspglnval725730735valalailealaserasnasnglyglylysglnalaleugluthrval740745750glnargleuleuprovalleucysglnalahisglyleuthrproasp755760765glnvalvalalailealaserasnasnglyglylysglnalaleuglu770775780thrvalglnargleuleuprovalleucysglnalahisglyleuthr785790795800proaspglnvalvalalailealaserhisaspglyglylysglnala805810815leugluthrvalglnargleuleuprovalleucysglnalahisgly820825830leuthrproaspglnvalvalalailealaserhisaspglyglylys835840845glnalaleugluserilevalalaglnleuserargproaspproala850855860leualaalaleuthrasnasphisleuvalalaleualacysleugly865870875880glyargproalaleuaspalavallyslysglyleuprohisalapro885890895gluleuileargargileasnargargileprogluargthrserhis900905910argvalalaaspleualahisvalvalargvalleuglyphephegln915920925serhisserhisproalaglnalapheaspaspalametthrglnphe930935940glymetserarghisglyleuvalglnleupheargargvalglyval945950955960thrglupheglualaargcysglythrleuproproalaserglnarg965970975trpaspargileleuglnalaserglymetlysargalalysproser980985990prothrseralaglnthrproaspglnalaserleuhisalapheala99510001005aspserleugluargaspleuaspalaproserpromethisglu101010151020glyaspglnthrargalaserserarglysargserargserasp102510301035argalavalthrglyproserthrglnglnserphegluvalarg104010451050valprogluglnargaspalaleuhisleuproleusertrparg105510601065vallysargproargthrargileglyglyglyleuproasppro107010751080glythrproilealaalaaspleualaalaserserthrvalmet108510901095trpserservalgluasnglyalalysalaalalysarglyslys110011051110argproleuprogluileglngluvalgluaspvalproargthr111511201125argargproargargalaalaalacysthrserphelysglulys113011351140serileargvalcysglulysseralathrilegluvallyslys114511501155glnglnilevalglugluglupheleualaleuargleuthrala116011651170leugluthraspvalgluaspargprothrargargleuasnasp117511801185phevalleupheaspseraspglyvalproglnproleuglumet119011951200leugluilehisaspilephevalserglyalaileleuproser120512101215aspvalcysthrasplysglulysglulysglyvalargcysthr122012251230serpheglyargvalgluhistrpserileserglytyrgluasp123512401245glyserprovaliletrpileserthrgluleualaasptyrasp125012551260cysarglysproalaalasertyrarglysvaltyrasptyrphe126512701275tyrglulysalaargalaservalalavaltyrlyslysleuser128012851290lysserserglyglyaspproaspileglyleuglugluleuleu129513001305alaalavalvalargsermetserserglyserlystyrpheser131013151320serglyalaalaileileaspphevalileserglnglyaspphe132513301335iletyrasnglnleualaglyleuaspgluthralalyslyshis134013451350glusersertyrvalgluileprovalleuvalalaleuargglu135513601365lysserserlysileasplysproleuglnarggluargasnpro137013751380serasnglyvalargilelysgluvalserglnvalalagluser138513901395glualaleuthrseraspglnleuvalaspglythraspaspasp140014051410argargtyralaileleuleuglnaspglugluasnarglysser141514201425metglnglnproarglysasnserserserglyseralaserasn143014351440metphetyrilelysileasngluaspgluilealaasnasptyr144514501455proleuprosertyrtyrlysthrsergluglugluthraspglu146014651470leuileleutyraspalasertyrgluvalglnsergluhisleu147514801485prohisargmetleuhisasntrpalaleutyrasnseraspleu149014951500argpheileserleugluleuleuprometlysglncysaspasp150515101515ileaspvalasnilepheglyserglyvalvalthraspaspasn152015251530glysertrpileserleuasnaspproaspserglyserglnser153515401545hisaspproaspglymetcysilepheleuserglnilelysglu155015551560trpmetileglupheglyseraspaspileileserileserile156515701575argthraspvalalatrptyrargleuglylysproserlysleu158015851590tyralaprotrptrplysprovalleulysthralaargvalgly159516001605ileserileleuthrpheleuargvalgluserargvalalaarg161016151620leuserphealaaspvalthrlysargleuserglyleuglnala162516301635asnasplysalatyrileserseraspproleualavalgluarg164016451650tyrleuvalvalhisglyglnileileleuglnleuphealaval165516601665tyrproaspaspasnvallysargcysprophevalvalglyleu167016751680alaserlysleugluasparghishisthrlystrpileilelys168516901695lyslyslysileserleulysgluleuasnleuasnproargala170017051710glymetalaprovalalaserlysarglysalametglnalathr171517201725thrthrargleuvalasnargiletrpglygluphetyrserasn173017351740tyrserprogluaspproleuglnalathralaalagluasngly174517501755gluaspgluvalgluglugluglyglyasnglygluglugluval176017651770gluglugluglygluasnglyleuthrgluaspthrvalproglu177517801785provalgluvalglnlysprohisthrprolyslysilearggly179017951800serserglylysarggluilelystrpaspglygluserleugly180518101815lysthrseralaglygluproleutyrglnglnalaleuvalgly182018251830glyglumetvalalavalglyglyalavalthrleugluvalasp183518401845aspproaspglumetproalailetyrphevalglutyrmetphe185018551860gluserthrasphiscyslysmetleuhisglyargpheleugln186518701875argglysermetthrvalleuglyasnalaalaasngluargglu188018851890leupheleuthrasnglucysmetthrthrglnleulysaspile189519001905lysglyvalalaserphegluileargserargprotrpglyhis191019151920glntyrarglyslysasnilethralaasplysleuasptrpala192519301935argalaleugluarglysvallysaspleuprothrglutyrtyr194019451950cyslysserleutyrserprogluargglyglyphepheserleu195519601965proleuseraspileglyargserserglyphecysthrsercys197019751980lysilearggluaspgluglulysargserthrilelysleuasn198519901995valserlysthrglyphepheileasnglyileglutyrserval200020052010gluaspphevaltyrvalasnproaspserileglyglyleulys201520202025gluglyserlysthrserphelysserglyargasnileglyleu203020352040argalatyrvalvalcysglnleuleugluilevalprolysglu204520502055serarglysalaaspleuglyserpheaspvallysvalargarg206020652070phetyrargprogluaspvalseralaglulysalatyralaser207520802085aspileglngluleutyrpheserglnaspthrvalvalleupro209020952100proglyalaleugluglylyscysgluvalarglyslysserasp210521102115metproleuserargglutyrproileserasphisilephephe212021252130cysaspleuphepheaspthrserlysglyserleulysglnleu213521402145proalaasnmetlysprolyspheserthrilelysaspaspthr215021552160leuleuarglyslyslysglylysglyvalglusergluileglu216521702175sergluilevallysprovalgluproprolysgluileargleu218021852190alathrleuaspilephealaglycysglyglyleuserhisgly219522002205leulyslysalaglyvalseraspalalystrpalaileglutyr221022152220glugluproalaglyglnalaphelysglnasnhisprogluser222522302235thrvalphevalaspasncysasnvalileleuargalailemet224022452250glulysglyglyaspglnaspaspcysvalserthrthrgluala225522602265asngluleualaalalysleuthrglugluglnlysserthrleu227022752280proleuproglyglnvalasppheileasnglyglyproprocys228522902295glnglypheserglymetasnargpheasnglnsersertrpser230023052310lysvalglncysglumetileleualapheleuserphealaasp231523202325tyrpheargproargtyrpheleuleugluasnvalargthrphe233023352340valserpheasnlysglyglnthrpheglnleuthrleualaser234523502355leuleuglumetglytyrglnvalargpheglyileleugluala236023652370glyalatyrglyvalserglnserarglysargalapheiletrp237523802385alaalaalaproglugluvalleuproglutrpproglupromet239023952400hisvalpheglyvalprolysleulysileserleuserglngly240524102415leuhistyralaalavalargserthralaleuglyalaprophe242024252430argproilethrvalargaspthrileglyaspleuproserval243524402445gluasnglyaspserargthrasnlysglutyrlysgluvalala245024552460valsertrppheglnlysgluileargglyasnthrilealaleu246524702475thrasphisilecyslysalametasngluleuasnleuilearg248024852490cyslysleuileprothrargproglyalaasptrphisaspleu249525002505prolysarglysvalthrleuseraspglyargvalgluglumet251025152520ileprophecysleuproasnthralagluarghisasnglytrp252525302535lysglyleutyrglyargleuasptrpglnglyasnpheprothr254025452550servalthraspproglnprometglylysvalglymetcysphe255525602565hisprogluglnhisargileleuthrvalargglucysalaarg257025752580serglnglypheproaspsertyrgluphealaglyasnileasn258525902595hislyshisargglnileglyasnalavalproproproleuala260026052610phealaleuglyarglysleulysglualaleuhisleulyslys261526202625serproglnhisglnpro2630<210>20<211>3330<212>dna<213>rb-vtale-mcs核苷酸序列<400>20atggaccccattcgttcgcgcacgccaagtcctgcccgcgagcttctgcccggaccccaa60ccggatagggttcagccgactgcagatcgggggggggctccgcctgctggcggccccctg120gatggcttgcccgctcggcggacgatgtcccggacccggctgccatctccccctgcgccc180tcgcctgcgttctcggcgggcagcttcaacgatctgctccgtcagttcgatccgtcgctt240cttgatacatcgcttcttgattcgatgcctgccgtcggcacgccgcatacagcggctgcc300ccagcagagtgggatgaggtgcaatcgggtctgcgtgcagccgatgacccgccacccacc360gtgcgtgtcgctgtcactgccgcgcggccgccgcgcgccaagccggccccgcgacggcgt420gcggcgcaaccctccgacgcttcgccggccgcgcaggtggatctacgcacgctcggctac480agtcagcagcagcaagagaagatcaaatcgaaggtgcgttcgacagtggcgcagcaccac540gaggcactggtgggccatgggtttacacacgcgcacatcgttgcgctcagccaacacccg600gcagcgttagggaccgtcgctgtcaagtatcagcacataatcacggcgttgccagaggcg660acacacgaagacatcgttggcgtcggcaaacagtggtccggcgcacgcgccctggaggcc720ttgctcacgaaggcgggggagttgagaggtccgccgttacagttggacacaggccaactt780ctcaagattgcaaaacgtggcggcgtgaccgcagtggaggcagtgcatgcatcgcgcaat840gcactgacgggtgcccccctgaaccttactccagatcaagttgtggcaatcgcttcaaat900atcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagct960catggactcactccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcg1020ttagaaactgtccaaaggctccttcccgtgttgtgtcaagctcatggcctcactccagat1080caagttgtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgt1140cttcttcctgtgttgtgtcaagctcatggattaactccagatcaagttgtggctatcgct1200agtaacggtggtggaaaacaggcgttagaaactgtccaaaggctccttcccgtgttgtgt1260caagctcatggcttaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaa1320caggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggactcact1380ccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtc1440caaaggctccttcccgtgttgtgtcaagctcatggcctcactccagatcaagttgtggct1500atcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaaggctccttcccgtg1560ttgtgtcaagctcatggactcactccagatcaagttgtggcaatagcctctaacaatgga1620ggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgtcaagctcatggc1680ctcactccagatcaagttgtggcaatagcctctaacaatggaggtaaacaagcacttgag1740actgttcagagattgttaccggtgttgtgtcaagctcatggattaactccagatcaagtt1800gtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttctt1860cctgtgttgtgtcaagctcatggtctcactccagatcaagttgtggccattgcatctcat1920gatggtggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgtcaagct1980catggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc2040ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtctcactccagat2100caagttgtggccattgcatctcatgatggtggtaagcaagctctcgaaacagtccaaaga2160cttctcccagtgttgtgtcaagctcatggactcactccagatcaagttgtggccattgca2220tctcatgatggtggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgt2280caagctcatggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaa2340caggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtttaact2400ccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtg2460caacgtcttcttcctgtgttgtgtcaagctcatggtctcactccagatcaagttgtggca2520atagcctctaacaatggaggtaaacaagcacttgagagcattgttgcccagttatctcgc2580cctgatccggcgttggccgcgttgaccaacgaccacctcgtcgccttggcctgcctcggc2640ggacgtcctgccctggatgcagtgaaaaagggattgccgcacgcgccggaattgatcaga2700agaatcaatcgccgtattcccgaacgcacgtcccatcgcgttgccgacctcgcgcacgtg2760gtgcgcgtgcttggttttttccagagccactcccacccagcgcaagcattcgatgacgcc2820atgacgcagttcgggatgagcaggcacgggttggtacagctctttcgcagagtgggcgtc2880accgaattcgaagcccgctgcggaacgctccccccagcctcgcagcgttgggaccgtatc2940ctccaggcatcagggatgaaaagggccaaaccgtcccctacttcagctcaaacgccggat3000caggcgtctttgcatgcattcgccgattcgctggagcgtgaccttgatgcgcccagccca3060atgcacgagggagatcagacgcgggcaagcagccgtaaacggtcccgatcggatcgtgct3120gtcaccggcccctccacacagcaatctttcgaggtgcgcgttcccgaacagcgcgatgcg3180ctgcatttgcccctcagctggagggtaaaacgcccgcgtaccaggatcgggggcggcctc3240ccggatcctggtacgcccatcgctgccgacctggcagcgtccagcaccgtgatgtggtct3300agattaattaaccccggggtcgactagtga3330<210>21<211>5175<212>dna<213>rb-vtale-atkyp核苷酸序列<400>21atggaccccattcgttcgcgcacgccaagtcctgcccgcgagcttctgcccggaccccaa60ccggatagggttcagccgactgcagatcgggggggggctccgcctgctggcggccccctg120gatggcttgcccgctcggcggacgatgtcccggacccggctgccatctccccctgcgccc180tcgcctgcgttctcggcgggcagcttcaacgatctgctccgtcagttcgatccgtcgctt240cttgatacatcgcttcttgattcgatgcctgccgtcggcacgccgcatacagcggctgcc300ccagcagagtgggatgaggtgcaatcgggtctgcgtgcagccgatgacccgccacccacc360gtgcgtgtcgctgtcactgccgcgcggccgccgcgcgccaagccggccccgcgacggcgt420gcggcgcaaccctccgacgcttcgccggccgcgcaggtggatctacgcacgctcggctac480agtcagcagcagcaagagaagatcaaatcgaaggtgcgttcgacagtggcgcagcaccac540gaggcactggtgggccatgggtttacacacgcgcacatcgttgcgctcagccaacacccg600gcagcgttagggaccgtcgctgtcaagtatcagcacataatcacggcgttgccagaggcg660acacacgaagacatcgttggcgtcggcaaacagtggtccggcgcacgcgccctggaggcc720ttgctcacgaaggcgggggagttgagaggtccgccgttacagttggacacaggccaactt780ctcaagattgcaaaacgtggcggcgtgaccgcagtggaggcagtgcatgcatcgcgcaat840gcactgacgggtgcccccctgaaccttactccagatcaagttgtggcaatcgcttcaaat900atcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagct960catggactcactccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcg1020ttagaaactgtccaaaggctccttcccgtgttgtgtcaagctcatggcctcactccagat1080caagttgtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgt1140cttcttcctgtgttgtgtcaagctcatggattaactccagatcaagttgtggctatcgct1200agtaacggtggtggaaaacaggcgttagaaactgtccaaaggctccttcccgtgttgtgt1260caagctcatggcttaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaa1320caggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggactcact1380ccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtc1440caaaggctccttcccgtgttgtgtcaagctcatggcctcactccagatcaagttgtggct1500atcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaaggctccttcccgtg1560ttgtgtcaagctcatggactcactccagatcaagttgtggcaatagcctctaacaatgga1620ggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgtcaagctcatggc1680ctcactccagatcaagttgtggcaatagcctctaacaatggaggtaaacaagcacttgag1740actgttcagagattgttaccggtgttgtgtcaagctcatggattaactccagatcaagtt1800gtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttctt1860cctgtgttgtgtcaagctcatggtctcactccagatcaagttgtggccattgcatctcat1920gatggtggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgtcaagct1980catggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc2040ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtctcactccagat2100caagttgtggccattgcatctcatgatggtggtaagcaagctctcgaaacagtccaaaga2160cttctcccagtgttgtgtcaagctcatggactcactccagatcaagttgtggccattgca2220tctcatgatggtggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgt2280caagctcatggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaa2340caggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtttaact2400ccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtg2460caacgtcttcttcctgtgttgtgtcaagctcatggtctcactccagatcaagttgtggca2520atagcctctaacaatggaggtaaacaagcacttgagagcattgttgcccagttatctcgc2580cctgatccggcgttggccgcgttgaccaacgaccacctcgtcgccttggcctgcctcggc2640ggacgtcctgccctggatgcagtgaaaaagggattgccgcacgcgccggaattgatcaga2700agaatcaatcgccgtattcccgaacgcacgtcccatcgcgttgccgacctcgcgcacgtg2760gtgcgcgtgcttggttttttccagagccactcccacccagcgcaagcattcgatgacgcc2820atgacgcagttcgggatgagcaggcacgggttggtacagctctttcgcagagtgggcgtc2880accgaattcgaagcccgctgcggaacgctccccccagcctcgcagcgttgggaccgtatc2940ctccaggcatcagggatgaaaagggccaaaccgtcccctacttcagctcaaacgccggat3000caggcgtctttgcatgcattcgccgattcgctggagcgtgaccttgatgcgcccagccca3060atgcacgagggagatcagacgcgggcaagcagccgtaaacggtcccgatcggatcgtgct3120gtcaccggcccctccacacagcaatctttcgaggtgcgcgttcccgaacagcgcgatgcg3180ctgcatttgcccctcagctggagggtaaaacgcccgcgtaccaggatcgggggcggcctc3240ccggatcctggtacgcccatcgctgccgacctggcagcgtccagcaccgtgatgtggtct3300agtgctggaaaaaggaaacgagctaatgctcctgaccaaacagagcgaagatcgagtgtt3360cgggttcagaaagtgagacagaaagcgttagatgagaaggcgcgtttagtacaggagagg3420gttaagctcctcagtgacagaaagagtgaaatttgtgtcgatgacactgagttacatgag3480aaagaagaggaaaatgtcgatgggagccctaaacgaagaagccctccaaagctaaccgca3540atgcagaaaggaaagcagaaattgagtgtttctctgaatggtaaggacgtgaacttggaa3600cctcatctcaaagtgacaaagtgtctgaggttatttaacaagcaatatctcctctgtgtc3660caggctaagttgagcaggcctgatttgaagggtgtaactgagatgataaaagctaaggcg3720atattgtacccaagaaaaataatcggtgaccttccaggtatagacgttggacaccgtttt3780ttttcaagagctgaaatgtgtgctgtaggattccataaccattggctaaatggcattgat3840tatatgtcaatggaatacgaaaaagagtatagtaactacaaattaccgcttgctgtttct3900attgttatgtcgggccagtacgaggatgatctagacaatgcagatacagtgacttacact3960ggacagggagggcataacttaactggtaataaacgtcagataaaggatcaacttttagaa4020cgagggaatttggcgctaaagcactgctgcgaatataatgtgcctgtcagagtaactcgt4080ggtcacaattgcaaaagtagctataccaaacgagtatacacttatgatggactgtacaag4140gttgaaaagttctgggcacaaaagggcgtttcaggatttacagtgtataagtaccgactg4200aaacgattggaggggcaaccagaactaactactgatcaggtcaactttgttgctggacgc4260ataccaacgagtacttcagaaattgagggtttggtatgtgaggacatctccggagggcta4320gaatttaagggtatccccgccactaatcgtgttgatgattcaccagtttcaccaacatct4380ggtttcacatacatcaaatctttgattattgagcctaatgtcataattccaaagagttca4440actgggtgtaactgccgaggcagctgcactgactcaaagaaatgtgcatgtgctaagctt4500aatgggggtaactttccatatgttgaccttaatgatggcagattaattgagtctcgagat4560gttgtatttgaatgtggtcctcactgtgggtgtgggccaaaatgtgtcaaccgaacttct4620cagaagcgtctaagattcaatcttgaggttttccgctctgcaaagaagggttgggcagtt4680agatcatgggagtacataccagctggttcaccagtatgtgagtacataggagttgtcagg4740agaactgctgatgtggatactatctctgacaatgaatacatatttgagattgactgccaa4800cagacaatgcaaggtcttggtggaagacagagaagactaagagatgttgctgtaccaatg4860aataatggagtcagtcagagcagtgaagatgagaatgcgccagagttctgcattgatgct4920ggttcaacaggaaactttgctaggtttataaatcacagttgtgaaccaaacctatttgtt4980cagtgcgtcctgagttctcaccaggatataaggcttgcccgtgtggttcttttcgcagct5040gacaacatttccccaatgcaggagctcacttacgactatggatatgcgcttgatagcgtt5100catggaccggatgggaaggtgaagcagctcgcttgctactgtggagcgctaaattgtagg5160aaacgcctttactaa5175<210>22<211>4710<212>dna<213>rb-vtale-athdac19核苷酸序列<400>22atggaccccattcgttcgcgcacgccaagtcctgcccgcgagcttctgcccggaccccaa60ccggatagggttcagccgactgcagatcgggggggggctccgcctgctggcggccccctg120gatggcttgcccgctcggcggacgatgtcccggacccggctgccatctccccctgcgccc180tcgcctgcgttctcggcgggcagcttcaacgatctgctccgtcagttcgatccgtcgctt240cttgatacatcgcttcttgattcgatgcctgccgtcggcacgccgcatacagcggctgcc300ccagcagagtgggatgaggtgcaatcgggtctgcgtgcagccgatgacccgccacccacc360gtgcgtgtcgctgtcactgccgcgcggccgccgcgcgccaagccggccccgcgacggcgt420gcggcgcaaccctccgacgcttcgccggccgcgcaggtggatctacgcacgctcggctac480agtcagcagcagcaagagaagatcaaatcgaaggtgcgttcgacagtggcgcagcaccac540gaggcactggtgggccatgggtttacacacgcgcacatcgttgcgctcagccaacacccg600gcagcgttagggaccgtcgctgtcaagtatcagcacataatcacggcgttgccagaggcg660acacacgaagacatcgttggcgtcggcaaacagtggtccggcgcacgcgccctggaggcc720ttgctcacgaaggcgggggagttgagaggtccgccgttacagttggacacaggccaactt780ctcaagattgcaaaacgtggcggcgtgaccgcagtggaggcagtgcatgcatcgcgcaat840gcactgacgggtgcccccctgaaccttactccagatcaagttgtggcaatcgcttcaaat900atcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagct960catggactcactccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcg1020ttagaaactgtccaaaggctccttcccgtgttgtgtcaagctcatggcctcactccagat1080caagttgtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgt1140cttcttcctgtgttgtgtcaagctcatggattaactccagatcaagttgtggctatcgct1200agtaacggtggtggaaaacaggcgttagaaactgtccaaaggctccttcccgtgttgtgt1260caagctcatggcttaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaa1320caggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggactcact1380ccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtc1440caaaggctccttcccgtgttgtgtcaagctcatggcctcactccagatcaagttgtggct1500atcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaaggctccttcccgtg1560ttgtgtcaagctcatggactcactccagatcaagttgtggcaatagcctctaacaatgga1620ggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgtcaagctcatggc1680ctcactccagatcaagttgtggcaatagcctctaacaatggaggtaaacaagcacttgag1740actgttcagagattgttaccggtgttgtgtcaagctcatggattaactccagatcaagtt1800gtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttctt1860cctgtgttgtgtcaagctcatggtctcactccagatcaagttgtggccattgcatctcat1920gatggtggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgtcaagct1980catggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc2040ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtctcactccagat2100caagttgtggccattgcatctcatgatggtggtaagcaagctctcgaaacagtccaaaga2160cttctcccagtgttgtgtcaagctcatggactcactccagatcaagttgtggccattgca2220tctcatgatggtggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgt2280caagctcatggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaa2340caggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtttaact2400ccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtg2460caacgtcttcttcctgtgttgtgtcaagctcatggtctcactccagatcaagttgtggca2520atagcctctaacaatggaggtaaacaagcacttgagagcattgttgcccagttatctcgc2580cctgatccggcgttggccgcgttgaccaacgaccacctcgtcgccttggcctgcctcggc2640ggacgtcctgccctggatgcagtgaaaaagggattgccgcacgcgccggaattgatcaga2700agaatcaatcgccgtattcccgaacgcacgtcccatcgcgttgccgacctcgcgcacgtg2760gtgcgcgtgcttggttttttccagagccactcccacccagcgcaagcattcgatgacgcc2820atgacgcagttcgggatgagcaggcacgggttggtacagctctttcgcagagtgggcgtc2880accgaattcgaagcccgctgcggaacgctccccccagcctcgcagcgttgggaccgtatc2940ctccaggcatcagggatgaaaagggccaaaccgtcccctacttcagctcaaacgccggat3000caggcgtctttgcatgcattcgccgattcgctggagcgtgaccttgatgcgcccagccca3060atgcacgagggagatcagacgcgggcaagcagccgtaaacggtcccgatcggatcgtgct3120gtcaccggcccctccacacagcaatctttcgaggtgcgcgttcccgaacagcgcgatgcg3180ctgcatttgcccctcagctggagggtaaaacgcccgcgtaccaggatcgggggcggcctc3240ccggatcctggtacgcccatcgctgccgacctggcagcgtccagcaccgtgatgtggtct3300agtgatactggcggcaattcgctggcgtccggacctgatggtgtgaagaggaaagtttgt3360tatttctatgaccctgaggtcggcaattactactatggccaaggtcatcccatgaagccc3420catcgcatccgcatgacccatgccctcctcgctcactacggtctccttcagcatatgcag3480gttctcaagcccttccctgcccgcgaccgtgatctctgccgcttccacgccgacgactat3540gtctcttttctccgcagcattacccctgaaacccagcaagatcagattcgccaacttaag3600cgcttcaatgttggtgaagactgtcccgtctttgacggcctttattccttttgccagacc3660tatgctggaggatctgttggtggctctgtcaagcttaaccacggcctctgcgatattgcc3720atcaactgggctggtggtctccatcacgctaagaagtgcgaggcctctggcttctgttac3780gtcaatgatatcgtcttagctatcctagagctccttaagcagcatgagcgtgttctttat3840gtcgatattgatatccaccacggggatggagtggaggaggcattttatgctactgacagg3900gttatgactgtctcgtttcataaatttggtgattactttcccggtacaggtcacattcag3960gatataggttatggtagcggaaagtactattctctcaatgtaccactggatgatggaatc4020gatgatgagagctatcatctgttattcaagcccatcatggggaaagttatggaaattttc4080cgaccaggggctgtggtattgcaatgtggtgctgattcattgtctggtgataggttgggg4140tgctttaatctttcaatcaaaggtcatgctgagtgcgtcaaatttatgagatcgttcaat4200gttcccctactgctcttgggtggtggtggttacactatccgcaatgttgcccgttgctgg4260tgctacgagactggagttgcacttggagttgaagttgaagacaagatgccggagcatgaa4320tattatgaatactttggtccagactatacacttcacgttgctccaagtaacatggaaaat4380aagaattctcgtcagatgcttgaagagattcgcaatgaccttctccacaatctctctaag4440cttcagcatgctccaagtgtaccatttcaggaaagaccacctgatacagagactcccgag4500gttgatgaagaccaagaagatggggataaaagatgggatccggattcagacatggatgtt4560gatgatgaccgtaaacctataccaagcagagtaaaaagagaagctgttgaaccagataca4620aaggacaaggatggactgaaaggaattatggagcgtggaaaaggttgtgaggtggaggtg4680gatgagagtggaagcactaaggtaaaatga4710<210>23<211>7905<212>dna<213>rb-vtale-atmet1核苷酸序列<400>23atggaccccattcgttcgcgcacgccaagtcctgcccgcgagcttctgcccggaccccaa60ccggatagggttcagccgactgcagatcgggggggggctccgcctgctggcggccccctg120gatggcttgcccgctcggcggacgatgtcccggacccggctgccatctccccctgcgccc180tcgcctgcgttctcggcgggcagcttcaacgatctgctccgtcagttcgatccgtcgctt240cttgatacatcgcttcttgattcgatgcctgccgtcggcacgccgcatacagcggctgcc300ccagcagagtgggatgaggtgcaatcgggtctgcgtgcagccgatgacccgccacccacc360gtgcgtgtcgctgtcactgccgcgcggccgccgcgcgccaagccggccccgcgacggcgt420gcggcgcaaccctccgacgcttcgccggccgcgcaggtggatctacgcacgctcggctac480agtcagcagcagcaagagaagatcaaatcgaaggtgcgttcgacagtggcgcagcaccac540gaggcactggtgggccatgggtttacacacgcgcacatcgttgcgctcagccaacacccg600gcagcgttagggaccgtcgctgtcaagtatcagcacataatcacggcgttgccagaggcg660acacacgaagacatcgttggcgtcggcaaacagtggtccggcgcacgcgccctggaggcc720ttgctcacgaaggcgggggagttgagaggtccgccgttacagttggacacaggccaactt780ctcaagattgcaaaacgtggcggcgtgaccgcagtggaggcagtgcatgcatcgcgcaat840gcactgacgggtgcccccctgaaccttactccagatcaagttgtggcaatcgcttcaaat900atcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagct960catggactcactccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcg1020ttagaaactgtccaaaggctccttcccgtgttgtgtcaagctcatggcctcactccagat1080caagttgtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgt1140cttcttcctgtgttgtgtcaagctcatggattaactccagatcaagttgtggctatcgct1200agtaacggtggtggaaaacaggcgttagaaactgtccaaaggctccttcccgtgttgtgt1260caagctcatggcttaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaa1320caggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggactcact1380ccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtc1440caaaggctccttcccgtgttgtgtcaagctcatggcctcactccagatcaagttgtggct1500atcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaaggctccttcccgtg1560ttgtgtcaagctcatggactcactccagatcaagttgtggcaatagcctctaacaatgga1620ggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgtcaagctcatggc1680ctcactccagatcaagttgtggcaatagcctctaacaatggaggtaaacaagcacttgag1740actgttcagagattgttaccggtgttgtgtcaagctcatggattaactccagatcaagtt1800gtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttctt1860cctgtgttgtgtcaagctcatggtctcactccagatcaagttgtggccattgcatctcat1920gatggtggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgtcaagct1980catggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc2040ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtctcactccagat2100caagttgtggccattgcatctcatgatggtggtaagcaagctctcgaaacagtccaaaga2160cttctcccagtgttgtgtcaagctcatggactcactccagatcaagttgtggccattgca2220tctcatgatggtggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgt2280caagctcatggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaa2340caggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtttaact2400ccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtg2460caacgtcttcttcctgtgttgtgtcaagctcatggtctcactccagatcaagttgtggca2520atagcctctaacaatggaggtaaacaagcacttgagagcattgttgcccagttatctcgc2580cctgatccggcgttggccgcgttgaccaacgaccacctcgtcgccttggcctgcctcggc2640ggacgtcctgccctggatgcagtgaaaaagggattgccgcacgcgccggaattgatcaga2700agaatcaatcgccgtattcccgaacgcacgtcccatcgcgttgccgacctcgcgcacgtg2760gtgcgcgtgcttggttttttccagagccactcccacccagcgcaagcattcgatgacgcc2820atgacgcagttcgggatgagcaggcacgggttggtacagctctttcgcagagtgggcgtc2880accgaattcgaagcccgctgcggaacgctccccccagcctcgcagcgttgggaccgtatc2940ctccaggcatcagggatgaaaagggccaaaccgtcccctacttcagctcaaacgccggat3000caggcgtctttgcatgcattcgccgattcgctggagcgtgaccttgatgcgcccagccca3060atgcacgagggagatcagacgcgggcaagcagccgtaaacggtcccgatcggatcgtgct3120gtcaccggcccctccacacagcaatctttcgaggtgcgcgttcccgaacagcgcgatgcg3180ctgcatttgcccctcagctggagggtaaaacgcccgcgtaccaggatcgggggcggcctc3240ccggatcctggtacgcccatcgctgccgacctggcagcgtccagcaccgtgatgtggtct3300agtgtggaaaatggggctaaagctgcgaagcgaaagaagagaccacttccagagattcaa3360gaggtagaagatgtacctaggacgaggagaccaaggcgtgctgcagcgtgtaccagtttc3420aaggagaaatctattcgagtctgtgagaaatctgctactattgaagtaaagaaacagcag3480attgtggaggaagagtttctcgcgttacggttaacggctctggaaactgatgttgaagat3540cgtccaaccaggagactgaatgattttgttttgtttgattcagatggagttccacaacct3600ctggagatgttggagattcatgacatattcgtttcaggtgctatcttaccttcagatgtg3660tgtactgataaggagaaagagaagggtgtgaggtgtacatcgtttggacgggttgagcat3720tggagtatctctggttatgaagatggttcccctgttatttggatctcaacggaattggcg3780gattatgattgtcgtaaacctgctgctagctacaggaaggtttatgattacttctatgag3840aaagctcgtgcttcagtggctgtgtataagaaattgtccaagtcatctggtggggatcct3900gatataggtcttgaggagttacttgcggcggttgtcagatcaatgagcagtggaagcaag3960tacttttctagtggtgcggcaatcatcgattttgttatatcccagggagattttatatat4020aaccaactcgctggtttggatgagacagccaagaaacatgaatcaagctatgttgagatt4080cctgttcttgtagctctcagagagaagagtagtaagattgacaagcctctgcagagggaa4140agaaacccatctaatggtgtgaggattaaagaagtttctcaagttgcggagagcgaggcc4200ttgacatctgatcaactggttgatggtactgatgatgacagaagatatgctatactctta4260caagacgaagagaataggaaatctatgcaacagcccagaaaaaacagcagctcaggttct4320gcttcaaatatgttctacattaagataaatgaagatgagattgccaatgattatcctctc4380ccatcgtactataagacctccgaagaagaaacagatgaacttatactttatgatgcttcc4440tatgaggttcaatctgaacacctgcctcacaggatgcttcacaactgggctctttataac4500tctgatttacgattcatatcactggaacttctaccgatgaaacaatgtgatgatattgat4560gtcaacatttttgggtcaggtgtggtgactgatgataatggaagttggatttctttaaac4620gatcctgacagcggttctcagtcacacgatcctgatgggatgtgcatattcctcagtcaa4680attaaagaatggatgattgagtttgggagcgatgatattatctccatttctatacgaaca4740gatgtggcctggtaccgtcttgggaaaccatcaaaactttatgccccttggtggaaacct4800gttctgaaaacagcaagggttgggataagcattcttacttttcttagggtggaaagtagg4860gttgctaggctttcatttgcagatgtcacaaaaagactgtctgggttacaggcgaatgat4920aaagcttacatttcttctgaccccttggctgttgagagatatttggtcgtccatgggcaa4980attattttacagctttttgcagtttatccggacgacaatgtcaaaaggtgtccatttgtt5040gttggtcttgcaagcaaattggaggataggcaccacacaaaatggatcatcaagaagaag5100aaaatttcgctgaaggaactgaatctgaatccaagggcaggcatggcaccagtagcatcg5160aagaggaaagctatgcaagcaacaacaactcgcctggtcaacagaatttggggagagttt5220tactccaattactctccagaggatccattgcaggcgactgctgcagaaaatggggaggat5280gaggtggaagaggaaggcggaaatggggaggaagaggttgaagaggaaggtgaaaatggt5340ctcacagaggacactgtaccagaacctgttgaggttcagaagcctcatactcctaagaaa5400atccgaggcagttctggaaaaagggaaataaaatgggatggtgagagtctaggaaaaact5460tctgctggcgagcctctctatcaacaagcccttgttggaggggaaatggtggctgtaggt5520ggcgctgtcaccttggaagttgatgatccagatgaaatgccggccatctattttgtggag5580tacatgttcgaaagtacagatcactgcaaaatgttacatggtagattcttacaaagagga5640tctatgactgttctggggaatgctgctaacgagagggaactattcctgactaatgaatgc5700atgactacacagctcaaggacattaaaggagtagccagttttgagattcgatcaaggcca5760tgggggcatcagtataggaaaaagaacatcactgcggataagcttgactgggctagagca5820ttagaaagaaaagtaaaagatttgccaacagagtattactgcaaaagcttgtactcacct5880gagagagggggattctttagtcttccactaagtgatattggtcgcagttctgggttctgc5940acttcatgtaagataagggaggatgaagagaagaggtctacaattaaactaaatgtttca6000aagacaggctttttcatcaatgggattgagtattctgttgaggattttgtctatgtcaac6060cctgactctattggtgggttgaaggagggtagtaaaacttcttttaagtctgggcgaaac6120attgggttaagagcgtatgttgtttgccaattgctggaaattgttccaaaggaatctaga6180aaggctgatttgggttcctttgatgttaaagtgagaaggttttataggcctgaggatgtt6240tctgcagagaaggcctatgcttcagacatccaagaattgtatttcagccaggacacagtt6300gttctccctccaggtgctctagagggaaaatgtgaagtaagaaagaaaagtgatatgccc6360ttatcccgtgaatatccaatatcagaccatattttcttctgtgatcttttctttgacacc6420tccaaaggttctctcaagcagctgcccgccaatatgaagccaaagttctctactattaag6480gacgacacacttttaagaaagaaaaagggaaagggagtagagagtgaaattgagtctgag6540attgtcaagcctgttgagccacctaaagagattcgtctggctactctagatatttttgct6600ggttgtggtggcctgtctcatggactgaaaaaggcgggtgtatctgatgcaaagtgggcg6660attgagtatgaagagccagctgggcaggcttttaaacaaaaccatcctgagtcaacagtt6720tttgttgacaactgcaatgtgattcttagggctataatggagaaaggtggagatcaagat6780gattgtgtctctactacagaggcaaatgaattagcagctaaactaactgaggagcagaag6840agtactctgccactgcctggtcaagtggacttcatcaatggtggacctccatgtcaggga6900ttttctggtatgaacaggttcaaccaaagctcttggagtaaagttcagtgtgaaatgata6960ttagcattcttgtcctttgctgactatttccggccaaggtattttcttctggagaacgtg7020aggacctttgtgtcattcaataaagggcagacatttcagcttactttggcttcccttctc7080gaaatgggttaccaggtgagatttggaatcctggaggccggtgcatatggagtatcccaa7140tctcgtaaacgagctttcatttgggctgctgcaccagaagaagttctccctgaatggcct7200gagccgatgcatgtctttggtgttccaaagttgaaaatctcactatctcaaggtttacat7260tatgctgctgttcgtagtactgcacttggtgcccctttccgtccaatcaccgtgagagac7320acaattggtgatcttccatcagtagaaaacggagactctaggacaaacaaagagtataaa7380gaggttgcagtctcgtggttccaaaaggagataagaggaaacacgattgctctcactgat7440catatctgcaaggctatgaatgagcttaacctcattcgatgcaaattaatcccaactagg7500cctggggctgattggcatgacttgccaaagagaaaggttacgttatctgatgggcgcgta7560gaagaaatgattcctttttgtctcccaaacacagctgagcgccacaacggttggaaggga7620ctatatgggagattagattggcaaggaaactttccgacttccgtcacggatcctcagccc7680atgggtaaggttggaatgtgctttcatcctgaacagcacagaatccttacagtccgtgaa7740tgcgcccgatctcaggggtttccggatagctacgagtttgcagggaacataaatcacaag7800cacaggcagattgggaatgcagtccctccaccattggcatttgctctaggtcgtaagctc7860aaagaagccctacatctcaagaagtctcctcaacaccaaccctag7905<210>24<211>3339<212>dna<213>rb-vtale-atsrdx核苷酸序列<400>24atggaccccattcgttcgcgcacgccaagtcctgcccgcgagcttctgcccggaccccaa60ccggatagggttcagccgactgcagatcgggggggggctccgcctgctggcggccccctg120gatggcttgcccgctcggcggacgatgtcccggacccggctgccatctccccctgcgccc180tcgcctgcgttctcggcgggcagcttcaacgatctgctccgtcagttcgatccgtcgctt240cttgatacatcgcttcttgattcgatgcctgccgtcggcacgccgcatacagcggctgcc300ccagcagagtgggatgaggtgcaatcgggtctgcgtgcagccgatgacccgccacccacc360gtgcgtgtcgctgtcactgccgcgcggccgccgcgcgccaagccggccccgcgacggcgt420gcggcgcaaccctccgacgcttcgccggccgcgcaggtggatctacgcacgctcggctac480agtcagcagcagcaagagaagatcaaatcgaaggtgcgttcgacagtggcgcagcaccac540gaggcactggtgggccatgggtttacacacgcgcacatcgttgcgctcagccaacacccg600gcagcgttagggaccgtcgctgtcaagtatcagcacataatcacggcgttgccagaggcg660acacacgaagacatcgttggcgtcggcaaacagtggtccggcgcacgcgccctggaggcc720ttgctcacgaaggcgggggagttgagaggtccgccgttacagttggacacaggccaactt780ctcaagattgcaaaacgtggcggcgtgaccgcagtggaggcagtgcatgcatcgcgcaat840gcactgacgggtgcccccctgaaccttactccagatcaagttgtggcaatcgcttcaaat900atcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagct960catggactcactccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcg1020ttagaaactgtccaaaggctccttcccgtgttgtgtcaagctcatggcctcactccagat1080caagttgtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgt1140cttcttcctgtgttgtgtcaagctcatggattaactccagatcaagttgtggctatcgct1200agtaacggtggtggaaaacaggcgttagaaactgtccaaaggctccttcccgtgttgtgt1260caagctcatggcttaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaa1320caggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggactcact1380ccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtc1440caaaggctccttcccgtgttgtgtcaagctcatggcctcactccagatcaagttgtggct1500atcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaaggctccttcccgtg1560ttgtgtcaagctcatggactcactccagatcaagttgtggcaatagcctctaacaatgga1620ggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgtcaagctcatggc1680ctcactccagatcaagttgtggcaatagcctctaacaatggaggtaaacaagcacttgag1740actgttcagagattgttaccggtgttgtgtcaagctcatggattaactccagatcaagtt1800gtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttctt1860cctgtgttgtgtcaagctcatggtctcactccagatcaagttgtggccattgcatctcat1920gatggtggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgtcaagct1980catggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc2040ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtctcactccagat2100caagttgtggccattgcatctcatgatggtggtaagcaagctctcgaaacagtccaaaga2160cttctcccagtgttgtgtcaagctcatggactcactccagatcaagttgtggccattgca2220tctcatgatggtggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgt2280caagctcatggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaa2340caggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtttaact2400ccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtg2460caacgtcttcttcctgtgttgtgtcaagctcatggtctcactccagatcaagttgtggca2520atagcctctaacaatggaggtaaacaagcacttgagagcattgttgcccagttatctcgc2580cctgatccggcgttggccgcgttgaccaacgaccacctcgtcgccttggcctgcctcggc2640ggacgtcctgccctggatgcagtgaaaaagggattgccgcacgcgccggaattgatcaga2700agaatcaatcgccgtattcccgaacgcacgtcccatcgcgttgccgacctcgcgcacgtg2760gtgcgcgtgcttggttttttccagagccactcccacccagcgcaagcattcgatgacgcc2820atgacgcagttcgggatgagcaggcacgggttggtacagctctttcgcagagtgggcgtc2880accgaattcgaagcccgctgcggaacgctccccccagcctcgcagcgttgggaccgtatc2940ctccaggcatcagggatgaaaagggccaaaccgtcccctacttcagctcaaacgccggat3000caggcgtctttgcatgcattcgccgattcgctggagcgtgaccttgatgcgcccagccca3060atgcacgagggagatcagacgcgggcaagcagccgtaaacggtcccgatcggatcgtgct3120gtcaccggcccctccacacagcaatctttcgaggtgcgcgttcccgaacagcgcgatgcg3180ctgcatttgcccctcagctggagggtaaaacgcccgcgtaccaggatcgggggcggcctc3240ccggatcctggtacgcccatcgctgccgacctggcagcgtccagcaccgtgatgtggtct3300agtcttgatctggatctggaactccgtttgggtttcgct3339<210>25<211>5256<212>dna<213>rb-vtale-atsuvh2核苷酸序列<400>25atggaccccattcgttcgcgcacgccaagtcctgcccgcgagcttctgcccggaccccaa60ccggatagggttcagccgactgcagatcgggggggggctccgcctgctggcggccccctg120gatggcttgcccgctcggcggacgatgtcccggacccggctgccatctccccctgcgccc180tcgcctgcgttctcggcgggcagcttcaacgatctgctccgtcagttcgatccgtcgctt240cttgatacatcgcttcttgattcgatgcctgccgtcggcacgccgcatacagcggctgcc300ccagcagagtgggatgaggtgcaatcgggtctgcgtgcagccgatgacccgccacccacc360gtgcgtgtcgctgtcactgccgcgcggccgccgcgcgccaagccggccccgcgacggcgt420gcggcgcaaccctccgacgcttcgccggccgcgcaggtggatctacgcacgctcggctac480agtcagcagcagcaagagaagatcaaatcgaaggtgcgttcgacagtggcgcagcaccac540gaggcactggtgggccatgggtttacacacgcgcacatcgttgcgctcagccaacacccg600gcagcgttagggaccgtcgctgtcaagtatcagcacataatcacggcgttgccagaggcg660acacacgaagacatcgttggcgtcggcaaacagtggtccggcgcacgcgccctggaggcc720ttgctcacgaaggcgggggagttgagaggtccgccgttacagttggacacaggccaactt780ctcaagattgcaaaacgtggcggcgtgaccgcagtggaggcagtgcatgcatcgcgcaat840gcactgacgggtgcccccctgaaccttactccagatcaagttgtggcaatcgcttcaaat900atcggagggaaacaggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagct960catggactcactccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcg1020ttagaaactgtccaaaggctccttcccgtgttgtgtcaagctcatggcctcactccagat1080caagttgtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgt1140cttcttcctgtgttgtgtcaagctcatggattaactccagatcaagttgtggctatcgct1200agtaacggtggtggaaaacaggcgttagaaactgtccaaaggctccttcccgtgttgtgt1260caagctcatggcttaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaa1320caggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggactcact1380ccagatcaagttgtggctatcgctagtaacggtggtggaaaacaggcgttagaaactgtc1440caaaggctccttcccgtgttgtgtcaagctcatggcctcactccagatcaagttgtggct1500atcgctagtaacggtggtggaaaacaggcgttagaaactgtccaaaggctccttcccgtg1560ttgtgtcaagctcatggactcactccagatcaagttgtggcaatagcctctaacaatgga1620ggtaaacaagcacttgagactgttcagagattgttaccggtgttgtgtcaagctcatggc1680ctcactccagatcaagttgtggcaatagcctctaacaatggaggtaaacaagcacttgag1740actgttcagagattgttaccggtgttgtgtcaagctcatggattaactccagatcaagtt1800gtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtgcaacgtcttctt1860cctgtgttgtgtcaagctcatggtctcactccagatcaagttgtggccattgcatctcat1920gatggtggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgtcaagct1980catggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggcc2040ttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtctcactccagat2100caagttgtggccattgcatctcatgatggtggtaagcaagctctcgaaacagtccaaaga2160cttctcccagtgttgtgtcaagctcatggactcactccagatcaagttgtggccattgca2220tctcatgatggtggtaagcaagctctcgaaacagtccaaagacttctcccagtgttgtgt2280caagctcatggattaactccagatcaagttgtggcaatcgcttcaaatatcggagggaaa2340caggccttggaaaccgtgcaacgtcttcttcctgtgttgtgtcaagctcatggtttaact2400ccagatcaagttgtggcaatcgcttcaaatatcggagggaaacaggccttggaaaccgtg2460caacgtcttcttcctgtgttgtgtcaagctcatggtctcactccagatcaagttgtggca2520atagcctctaacaatggaggtaaacaagcacttgagagcattgttgcccagttatctcgc2580cctgatccggcgttggccgcgttgaccaacgaccacctcgtcgccttggcctgcctcggc2640ggacgtcctgccctggatgcagtgaaaaagggattgccgcacgcgccggaattgatcaga2700agaatcaatcgccgtattcccgaacgcacgtcccatcgcgttgccgacctcgcgcacgtg2760gtgcgcgtgcttggttttttccagagccactcccacccagcgcaagcattcgatgacgcc2820atgacgcagttcgggatgagcaggcacgggttggtacagctctttcgcagagtgggcgtc2880accgaattcgaagcccgctgcggaacgctccccccagcctcgcagcgttgggaccgtatc2940ctccaggcatcagggatgaaaagggccaaaccgtcccctacttcagctcaaacgccggat3000caggcgtctttgcatgcattcgccgattcgctggagcgtgaccttgatgcgcccagccca3060atgcacgagggagatcagacgcgggcaagcagccgtaaacggtcccgatcggatcgtgct3120gtcaccggcccctccacacagcaatctttcgaggtgcgcgttcccgaacagcgcgatgcg3180ctgcatttgcccctcagctggagggtaaaacgcccgcgtaccaggatcgggggcggcctc3240ccggatcctggtacgcccatcgctgccgacctggcagcgtccagcaccgtgatgtggtct3300agtagtacattgttaccatttcctgacctcaacctcatgccggattctcaatcctccacc3360gccggaaccacagccggcgacactgtagtcaccggaaagttagaagtgaaatcggaacca3420attgaggaatggcaaactccaccatcgtctacctccgatcaatcagccaataccgatctc3480atcgccgagtttattcgtatctcagagctcttccgctcggcattcaagccactgcaggtc3540aaaggattagacggagtctccgtatacggattggattctggtgccatcgttgctgtaccg3600gagaaagaaaaccgggaattgattgaaccgcctccaggatttaaggataatcgagtctcc3660accgtcgttgtctcgccgaaattcgagagaccgagagagctagcgagaattgcaattcta3720ggtcatgaacaacggaaggaactccgacaagtcatgaaacgaactaggatgacttatgag3780tctcttcggattcatctaatggcggagagtatgaagaatcatgtacttggtcaagggcgt3840agacggaggagcgatatggcggctgcgtatataatgagggatcggggactgtggttgaac3900tatgataagcacatagttggtccagtcacaggggttgaagtaggggatatattcttctac3960cgtatggaattgtgtgtgctgggcttacacggccagacacaagccgggattgattgttta4020acggctgaacggagcgccactggagagcctatagctaccagcattgttgtctcaggtggt4080tatgaggatgatgaagatacaggagatgttttggtttatactggtcacggtggtcaggat4140catcaacacaagcagtgtgataaccagaggctagtaggtgggaatctgggaatggagaga4200agtatgcactacggtatcgaggtacgtgtgattagaggtattaagtatgagaacagtata4260agctcgaaggtgtatgtatatgatggtttgtataagatagtagattggtggtttgctgtg4320gggaagtctggttttggggtttttaaatttaggttagtaagaattgaagggcaaccgatg4380atgggcagtgcggtaatgaggtttgcacaaactcttagaaataagccatcgatggttagg4440cctactggttatgttagctttgacctttctaacaagaaggagaatgtacccgtgtttcta4500tataatgatgtagatggtgatcaagaaccgagacattatgagtacattgcaaaagctgtc4560tttcctcctggaatttttggtcagggggggatcagcaggactggctgcgagtgtaagctc4620tcttgtactgatgattgtctctgtgcaaggaagaacggtggtgagtttgcatatgatgat4680aacgggcatctattgaaaggaaagcatgtggtatttgaatgtggggaattctgcacttgt4740ggtccgagctgtaagagccgtgtgacacagaagggattgaggaacaggctagaggttttc4800agatcgaaggaaaccggttggggtgtcaggacacttgatctaattgaagctggtgctttc4860atatgcgaatatgcaggtgtagttgtcacgagacttcaagccgagattctgtcaatgaat4920ggggatgttatggtctatcctggtcggttcacagaccaatggcgtaactggggtgattta4980tctcaagtatacccagattttgttaggccgaattatccatctctccctccactggatttc5040tcaatggatgtgtcaaggatgaggaacgtggcttgctacattagtcacagcaaagaacca5100aatgtgatggtgcaatttgttttgcatgaccacaatcacctgatgttcccccgtgtgatg5160ctttttgcgctggagaatatctcaccgttggccgagctaagcctggattacggcttggca5220gatgaagtgaacggcaagctcgccatctgcaactag5256<210>26<211>1724<212>prt<213>rb-vtale-atkyp氨基酸序列<400>26metaspproileargserargthrproserproalaarggluleuleu151015proglyproglnproaspargvalglnprothralaaspargglygly202530alaproproalaglyglyproleuaspglyleuproalaargargthr354045metserargthrargleuproserproproalaproserproalaphe505560seralaglyserpheasnaspleuleuargglnpheaspproserleu65707580leuaspthrserleuleuaspsermetproalavalglythrprohis859095thralaalaalaproalaglutrpaspgluvalglnserglyleuarg100105110alaalaaspaspproproprothrvalargvalalavalthralaala115120125argproproargalalysproalaproargargargalaalaglnpro130135140seraspalaserproalaalaglnvalaspleuargthrleuglytyr145150155160serglnglnglnglnglulysilelysserlysvalargserthrval165170175alaglnhishisglualaleuvalglyhisglyphethrhisalahis180185190ilevalalaleuserglnhisproalaalaleuglythrvalalaval195200205lystyrglnhisileilethralaleuproglualathrhisgluasp210215220ilevalglyvalglylysglntrpserglyalaargalaleugluala225230235240leuleuthrlysalaglygluleuargglyproproleuglnleuasp245250255thrglyglnleuleulysilealalysargglyglyvalthralaval260265270glualavalhisalaserargasnalaleuthrglyalaproleuasn275280285leuthrproaspglnvalvalalailealaserasnileglyglylys290295300glnalaleugluthrvalglnargleuleuprovalleucysglnala305310315320hisglyleuthrproaspglnvalvalalailealaserasnglygly325330335glylysglnalaleugluthrvalglnargleuleuprovalleucys340345350glnalahisglyleuthrproaspglnvalvalalailealaserasn355360365ileglyglylysglnalaleugluthrvalglnargleuleuproval370375380leucysglnalahisglyleuthrproaspglnvalvalalaileala385390395400serasnglyglyglylysglnalaleugluthrvalglnargleuleu405410415provalleucysglnalahisglyleuthrproaspglnvalvalala420425430ilealaserasnileglyglylysglnalaleugluthrvalglnarg435440445leuleuprovalleucysglnalahisglyleuthrproaspglnval450455460valalailealaserasnglyglyglylysglnalaleugluthrval465470475480glnargleuleuprovalleucysglnalahisglyleuthrproasp485490495glnvalvalalailealaserasnglyglyglylysglnalaleuglu500505510thrvalglnargleuleuprovalleucysglnalahisglyleuthr515520525proaspglnvalvalalailealaserasnasnglyglylysglnala530535540leugluthrvalglnargleuleuprovalleucysglnalahisgly545550555560leuthrproaspglnvalvalalailealaserasnasnglyglylys565570575glnalaleugluthrvalglnargleuleuprovalleucysglnala580585590hisglyleuthrproaspglnvalvalalailealaserasnilegly595600605glylysglnalaleugluthrvalglnargleuleuprovalleucys610615620glnalahisglyleuthrproaspglnvalvalalailealaserhis625630635640aspglyglylysglnalaleugluthrvalglnargleuleuproval645650655leucysglnalahisglyleuthrproaspglnvalvalalaileala660665670serasnileglyglylysglnalaleugluthrvalglnargleuleu675680685provalleucysglnalahisglyleuthrproaspglnvalvalala690695700ilealaserhisaspglyglylysglnalaleugluthrvalglnarg705710715720leuleuprovalleucysglnalahisglyleuthrproaspglnval725730735valalailealaserhisaspglyglylysglnalaleugluthrval740745750glnargleuleuprovalleucysglnalahisglyleuthrproasp755760765glnvalvalalailealaserasnileglyglylysglnalaleuglu770775780thrvalglnargleuleuprovalleucysglnalahisglyleuthr785790795800proaspglnvalvalalailealaserasnileglyglylysglnala805810815leugluthrvalglnargleuleuprovalleucysglnalahisgly820825830leuthrproaspglnvalvalalailealaserasnasnglyglylys835840845glnalaleugluserilevalalaglnleuserargproaspproala850855860leualaalaleuthrasnasphisleuvalalaleualacysleugly865870875880glyargproalaleuaspalavallyslysglyleuprohisalapro885890895gluleuileargargileasnargargileprogluargthrserhis900905910argvalalaaspleualahisvalvalargvalleuglyphephegln915920925serhisserhisproalaglnalapheaspaspalametthrglnphe930935940glymetserarghisglyleuvalglnleupheargargvalglyval945950955960thrglupheglualaargcysglythrleuproproalaserglnarg965970975trpaspargileleuglnalaserglymetlysargalalysproser980985990prothrseralaglnthrproaspglnalaserleuhisalapheala99510001005aspserleugluargaspleuaspalaproserpromethisglu101010151020glyaspglnthrargalaserserarglysargserargserasp102510301035argalavalthrglyproserthrglnglnserphegluvalarg104010451050valprogluglnargaspalaleuhisleuproleusertrparg105510601065vallysargproargthrargileglyglyglyleuproasppro107010751080glythrproilealaalaaspleualaalaserserthrvalmet108510901095trpserseralaglylysarglysargalaasnalaproaspgln110011051110thrgluargargserservalargvalglnlysvalargglnlys111511201125alaleuaspglulysalaargleuvalglngluargvallysleu113011351140leuserasparglyssergluilecysvalaspaspthrgluleu114511501155hisglulysgluglugluasnvalaspglyserprolysargarg116011651170serproprolysleuthralametglnlysglylysglnlysleu117511801185servalserleuasnglylysaspvalasnleugluprohisleu119011951200lysvalthrlyscysleuargleupheasnlysglntyrleuleu120512101215cysvalglnalalysleuserargproaspleulysglyvalthr122012251230glumetilelysalalysalaileleutyrproarglysileile123512401245glyaspleuproglyileaspvalglyhisargphepheserarg125012551260alaglumetcysalavalglyphehisasnhistrpleuasngly126512701275ileasptyrmetsermetglutyrglulysglutyrserasntyr128012851290lysleuproleualavalserilevalmetserglyglntyrglu129513001305aspaspleuaspasnalaaspthrvalthrtyrthrglyglngly131013151320glyhisasnleuthrglyasnlysargglnilelysaspglnleu132513301335leugluargglyasnleualaleulyshiscyscysglutyrasn134013451350valprovalargvalthrargglyhisasncyslyssersertyr135513601365thrlysargvaltyrthrtyraspglyleutyrlysvalglulys137013751380phetrpalaglnlysglyvalserglyphethrvaltyrlystyr138513901395argleulysargleugluglyglnprogluleuthrthraspgln140014051410valasnphevalalaglyargileprothrserthrsergluile141514201425gluglyleuvalcysgluaspileserglyglyleugluphelys143014351440glyileproalathrasnargvalaspaspserprovalserpro144514501455thrserglyphethrtyrilelysserleuileilegluproasn146014651470valileileprolysserserthrglycysasncysargglyser147514801485cysthraspserlyslyscysalacysalalysleuasnglygly149014951500asnpheprotyrvalaspleuasnaspglyargleuilegluser150515101515argaspvalvalpheglucysglyprohiscysglycysglypro152015251530lyscysvalasnargthrserglnlysargleuargpheasnleu153515401545gluvalpheargseralalyslysglytrpalavalargsertrp155015551560glutyrileproalaglyserprovalcysglutyrileglyval156515701575valargargthralaaspvalaspthrileseraspasnglutyr158015851590ilephegluileaspcysglnglnthrmetglnglyleuglygly159516001605argglnargargleuargaspvalalavalprometasnasngly161016151620valserglnsersergluaspgluasnalaprogluphecysile162516301635aspalaglyserthrglyasnphealaargpheileasnhisser164016451650cysgluproasnleuphevalglncysvalleuserserhisgln165516601665aspileargleualaargvalvalleuphealaalaaspasnile167016751680serprometglngluleuthrtyrasptyrglytyralaleuasp168516901695servalhisglyproaspglylysvallysglnleualacystyr170017051710cysglyalaleuasncysarglysargleutyr17151720<210>27<211>1569<212>prt<213>rb-vtale-athdac19氨基酸序列<400>27metaspproileargserargthrproserproalaarggluleuleu151015proglyproglnproaspargvalglnprothralaaspargglygly202530alaproproalaglyglyproleuaspglyleuproalaargargthr354045metserargthrargleuproserproproalaproserproalaphe505560seralaglyserpheasnaspleuleuargglnpheaspproserleu65707580leuaspthrserleuleuaspsermetproalavalglythrprohis859095thralaalaalaproalaglutrpaspgluvalglnserglyleuarg100105110alaalaaspaspproproprothrvalargvalalavalthralaala115120125argproproargalalysproalaproargargargalaalaglnpro130135140seraspalaserproalaalaglnvalaspleuargthrleuglytyr145150155160serglnglnglnglnglulysilelysserlysvalargserthrval165170175alaglnhishisglualaleuvalglyhisglyphethrhisalahis180185190ilevalalaleuserglnhisproalaalaleuglythrvalalaval195200205lystyrglnhisileilethralaleuproglualathrhisgluasp210215220ilevalglyvalglylysglntrpserglyalaargalaleugluala225230235240leuleuthrlysalaglygluleuargglyproproleuglnleuasp245250255thrglyglnleuleulysilealalysargglyglyvalthralaval260265270glualavalhisalaserargasnalaleuthrglyalaproleuasn275280285leuthrproaspglnvalvalalailealaserasnileglyglylys290295300glnalaleugluthrvalglnargleuleuprovalleucysglnala305310315320hisglyleuthrproaspglnvalvalalailealaserasnglygly325330335glylysglnalaleugluthrvalglnargleuleuprovalleucys340345350glnalahisglyleuthrproaspglnvalvalalailealaserasn355360365ileglyglylysglnalaleugluthrvalglnargleuleuproval370375380leucysglnalahisglyleuthrproaspglnvalvalalaileala385390395400serasnglyglyglylysglnalaleugluthrvalglnargleuleu405410415provalleucysglnalahisglyleuthrproaspglnvalvalala420425430ilealaserasnileglyglylysglnalaleugluthrvalglnarg435440445leuleuprovalleucysglnalahisglyleuthrproaspglnval450455460valalailealaserasnglyglyglylysglnalaleugluthrval465470475480glnargleuleuprovalleucysglnalahisglyleuthrproasp485490495glnvalvalalailealaserasnglyglyglylysglnalaleuglu500505510thrvalglnargleuleuprovalleucysglnalahisglyleuthr515520525proaspglnvalvalalailealaserasnasnglyglylysglnala530535540leugluthrvalglnargleuleuprovalleucysglnalahisgly545550555560leuthrproaspglnvalvalalailealaserasnasnglyglylys565570575glnalaleugluthrvalglnargleuleuprovalleucysglnala580585590hisglyleuthrproaspglnvalvalalailealaserasnilegly595600605glylysglnalaleugluthrvalglnargleuleuprovalleucys610615620glnalahisglyleuthrproaspglnvalvalalailealaserhis625630635640aspglyglylysglnalaleugluthrvalglnargleuleuproval645650655leucysglnalahisglyleuthrproaspglnvalvalalaileala660665670serasnileglyglylysglnalaleugluthrvalglnargleuleu675680685provalleucysglnalahisglyleuthrproaspglnvalvalala690695700ilealaserhisaspglyglylysglnalaleugluthrvalglnarg705710715720leuleuprovalleucysglnalahisglyleuthrproaspglnval725730735valalailealaserhisaspglyglylysglnalaleugluthrval740745750glnargleuleuprovalleucysglnalahisglyleuthrproasp755760765glnvalvalalailealaserasnileglyglylysglnalaleuglu770775780thrvalglnargleuleuprovalleucysglnalahisglyleuthr785790795800proaspglnvalvalalailealaserasnileglyglylysglnala805810815leugluthrvalglnargleuleuprovalleucysglnalahisgly820825830leuthrproaspglnvalvalalailealaserasnasnglyglylys835840845glnalaleugluserilevalalaglnleuserargproaspproala850855860leualaalaleuthrasnasphisleuvalalaleualacysleugly865870875880glyargproalaleuaspalavallyslysglyleuprohisalapro885890895gluleuileargargileasnargargileprogluargthrserhis900905910argvalalaaspleualahisvalvalargvalleuglyphephegln915920925serhisserhisproalaglnalapheaspaspalametthrglnphe930935940glymetserarghisglyleuvalglnleupheargargvalglyval945950955960thrglupheglualaargcysglythrleuproproalaserglnarg965970975trpaspargileleuglnalaserglymetlysargalalysproser980985990prothrseralaglnthrproaspglnalaserleuhisalapheala99510001005aspserleugluargaspleuaspalaproserpromethisglu101010151020glyaspglnthrargalaserserarglysargserargserasp102510301035argalavalthrglyproserthrglnglnserphegluvalarg104010451050valprogluglnargaspalaleuhisleuproleusertrparg105510601065vallysargproargthrargileglyglyglyleuproasppro107010751080glythrproilealaalaaspleualaalaserserthrvalmet108510901095trpserseraspthrglyglyasnserleualaserglyproasp110011051110glyvallysarglysvalcystyrphetyraspprogluvalgly111511201125asntyrtyrtyrglyglnglyhisprometlysprohisargile113011351140argmetthrhisalaleuleualahistyrglyleuleuglnhis114511501155metglnvalleulyspropheproalaargaspargaspleucys116011651170argphehisalaaspasptyrvalserpheleuargserilethr117511801185progluthrglnglnaspglnileargglnleulysargpheasn119011951200valglygluaspcysprovalpheaspglyleutyrserphecys120512101215glnthrtyralaglyglyservalglyglyservallysleuasn122012251230hisglyleucysaspilealaileasntrpalaglyglyleuhis123512401245hisalalyslyscysglualaserglyphecystyrvalasnasp125012551260ilevalleualaileleugluleuleulysglnhisgluargval126512701275leutyrvalaspileaspilehishisglyaspglyvalgluglu128012851290alaphetyralathraspargvalmetthrvalserphehislys129513001305pheglyasptyrpheproglythrglyhisileglnaspilegly131013151320tyrglyserglylystyrtyrserleuasnvalproleuaspasp132513301335glyileaspaspglusertyrhisleuleuphelysproilemet134013451350glylysvalmetgluilepheargproglyalavalvalleugln135513601365cysglyalaaspserleuserglyaspargleuglycyspheasn137013751380leuserilelysglyhisalaglucysvallysphemetargser138513901395pheasnvalproleuleuleuleuglyglyglyglytyrthrile140014051410argasnvalalaargcystrpcystyrgluthrglyvalalaleu141514201425glyvalgluvalgluasplysmetprogluhisglutyrtyrglu143014351440tyrpheglyproasptyrthrleuhisvalalaproserasnmet144514501455gluasnlysasnserargglnmetleuglugluileargasnasp146014651470leuleuhisasnleuserlysleuglnhisalaproservalpro147514801485pheglngluargproproaspthrgluthrprogluvalaspglu149014951500aspglngluaspglyasplysargtrpaspproaspseraspmet150515101515aspvalaspaspasparglysproileproserargvallysarg152015251530glualavalgluproaspthrlysasplysaspglyleulysgly153515401545ilemetgluargglylysglycysgluvalgluvalaspgluser155015551560glyserthrlysvallys1565<210>28<211>2634<212>prt<213>rb-vtale-atmet1氨基酸序列<400>28metaspproileargserargthrproserproalaarggluleuleu151015proglyproglnproaspargvalglnprothralaaspargglygly202530alaproproalaglyglyproleuaspglyleuproalaargargthr354045metserargthrargleuproserproproalaproserproalaphe505560seralaglyserpheasnaspleuleuargglnpheaspproserleu65707580leuaspthrserleuleuaspsermetproalavalglythrprohis859095thralaalaalaproalaglutrpaspgluvalglnserglyleuarg100105110alaalaaspaspproproprothrvalargvalalavalthralaala115120125argproproargalalysproalaproargargargalaalaglnpro130135140seraspalaserproalaalaglnvalaspleuargthrleuglytyr145150155160serglnglnglnglnglulysilelysserlysvalargserthrval165170175alaglnhishisglualaleuvalglyhisglyphethrhisalahis180185190ilevalalaleuserglnhisproalaalaleuglythrvalalaval195200205lystyrglnhisileilethralaleuproglualathrhisgluasp210215220ilevalglyvalglylysglntrpserglyalaargalaleugluala225230235240leuleuthrlysalaglygluleuargglyproproleuglnleuasp245250255thrglyglnleuleulysilealalysargglyglyvalthralaval260265270glualavalhisalaserargasnalaleuthrglyalaproleuasn275280285leuthrproaspglnvalvalalailealaserasnileglyglylys290295300glnalaleugluthrvalglnargleuleuprovalleucysglnala305310315320hisglyleuthrproaspglnvalvalalailealaserasnglygly325330335glylysglnalaleugluthrvalglnargleuleuprovalleucys340345350glnalahisglyleuthrproaspglnvalvalalailealaserasn355360365ileglyglylysglnalaleugluthrvalglnargleuleuproval370375380leucysglnalahisglyleuthrproaspglnvalvalalaileala385390395400serasnglyglyglylysglnalaleugluthrvalglnargleuleu405410415provalleucysglnalahisglyleuthrproaspglnvalvalala420425430ilealaserasnileglyglylysglnalaleugluthrvalglnarg435440445leuleuprovalleucysglnalahisglyleuthrproaspglnval450455460valalailealaserasnglyglyglylysglnalaleugluthrval465470475480glnargleuleuprovalleucysglnalahisglyleuthrproasp485490495glnvalvalalailealaserasnglyglyglylysglnalaleuglu500505510thrvalglnargleuleuprovalleucysglnalahisglyleuthr515520525proaspglnvalvalalailealaserasnasnglyglylysglnala530535540leugluthrvalglnargleuleuprovalleucysglnalahisgly545550555560leuthrproaspglnvalvalalailealaserasnasnglyglylys565570575glnalaleugluthrvalglnargleuleuprovalleucysglnala580585590hisglyleuthrproaspglnvalvalalailealaserasnilegly595600605glylysglnalaleugluthrvalglnargleuleuprovalleucys610615620glnalahisglyleuthrproaspglnvalvalalailealaserhis625630635640aspglyglylysglnalaleugluthrvalglnargleuleuproval645650655leucysglnalahisglyleuthrproaspglnvalvalalaileala660665670serasnileglyglylysglnalaleugluthrvalglnargleuleu675680685provalleucysglnalahisglyleuthrproaspglnvalvalala690695700ilealaserhisaspglyglylysglnalaleugluthrvalglnarg705710715720leuleuprovalleucysglnalahisglyleuthrproaspglnval725730735valalailealaserhisaspglyglylysglnalaleugluthrval740745750glnargleuleuprovalleucysglnalahisglyleuthrproasp755760765glnvalvalalailealaserasnileglyglylysglnalaleuglu770775780thrvalglnargleuleuprovalleucysglnalahisglyleuthr785790795800proaspglnvalvalalailealaserasnileglyglylysglnala805810815leugluthrvalglnargleuleuprovalleucysglnalahisgly820825830leuthrproaspglnvalvalalailealaserasnasnglyglylys835840845glnalaleugluserilevalalaglnleuserargproaspproala850855860leualaalaleuthrasnasphisleuvalalaleualacysleugly865870875880glyargproalaleuaspalavallyslysglyleuprohisalapro885890895gluleuileargargileasnargargileprogluargthrserhis900905910argvalalaaspleualahisvalvalargvalleuglyphephegln915920925serhisserhisproalaglnalapheaspaspalametthrglnphe930935940glymetserarghisglyleuvalglnleupheargargvalglyval945950955960thrglupheglualaargcysglythrleuproproalaserglnarg965970975trpaspargileleuglnalaserglymetlysargalalysproser980985990prothrseralaglnthrproaspglnalaserleuhisalapheala99510001005aspserleugluargaspleuaspalaproserpromethisglu101010151020glyaspglnthrargalaserserarglysargserargserasp102510301035argalavalthrglyproserthrglnglnserphegluvalarg104010451050valprogluglnargaspalaleuhisleuproleusertrparg105510601065vallysargproargthrargileglyglyglyleuproasppro107010751080glythrproilealaalaaspleualaalaserserthrvalmet108510901095trpserservalgluasnglyalalysalaalalysarglyslys110011051110argproleuprogluileglngluvalgluaspvalproargthr111511201125argargproargargalaalaalacysthrserphelysglulys113011351140serileargvalcysglulysseralathrilegluvallyslys114511501155glnglnilevalglugluglupheleualaleuargleuthrala116011651170leugluthraspvalgluaspargprothrargargleuasnasp117511801185phevalleupheaspseraspglyvalproglnproleuglumet119011951200leugluilehisaspilephevalserglyalaileleuproser120512101215aspvalcysthrasplysglulysglulysglyvalargcysthr122012251230serpheglyargvalgluhistrpserileserglytyrgluasp123512401245glyserprovaliletrpileserthrgluleualaasptyrasp125012551260cysarglysproalaalasertyrarglysvaltyrasptyrphe126512701275tyrglulysalaargalaservalalavaltyrlyslysleuser128012851290lysserserglyglyaspproaspileglyleuglugluleuleu129513001305alaalavalvalargsermetserserglyserlystyrpheser131013151320serglyalaalaileileaspphevalileserglnglyaspphe132513301335iletyrasnglnleualaglyleuaspgluthralalyslyshis134013451350glusersertyrvalgluileprovalleuvalalaleuargglu135513601365lysserserlysileasplysproleuglnarggluargasnpro137013751380serasnglyvalargilelysgluvalserglnvalalagluser138513901395glualaleuthrseraspglnleuvalaspglythraspaspasp140014051410argargtyralaileleuleuglnaspglugluasnarglysser141514201425metglnglnproarglysasnserserserglyseralaserasn143014351440metphetyrilelysileasngluaspgluilealaasnasptyr144514501455proleuprosertyrtyrlysthrsergluglugluthraspglu146014651470leuileleutyraspalasertyrgluvalglnsergluhisleu147514801485prohisargmetleuhisasntrpalaleutyrasnseraspleu149014951500argpheileserleugluleuleuprometlysglncysaspasp150515101515ileaspvalasnilepheglyserglyvalvalthraspaspasn152015251530glysertrpileserleuasnaspproaspserglyserglnser153515401545hisaspproaspglymetcysilepheleuserglnilelysglu155015551560trpmetileglupheglyseraspaspileileserileserile156515701575argthraspvalalatrptyrargleuglylysproserlysleu158015851590tyralaprotrptrplysprovalleulysthralaargvalgly159516001605ileserileleuthrpheleuargvalgluserargvalalaarg161016151620leuserphealaaspvalthrlysargleuserglyleuglnala162516301635asnasplysalatyrileserseraspproleualavalgluarg164016451650tyrleuvalvalhisglyglnileileleuglnleuphealaval165516601665tyrproaspaspasnvallysargcysprophevalvalglyleu167016751680alaserlysleugluasparghishisthrlystrpileilelys168516901695lyslyslysileserleulysgluleuasnleuasnproargala170017051710glymetalaprovalalaserlysarglysalametglnalathr171517201725thrthrargleuvalasnargiletrpglygluphetyrserasn173017351740tyrserprogluaspproleuglnalathralaalagluasngly174517501755gluaspgluvalgluglugluglyglyasnglygluglugluval176017651770gluglugluglygluasnglyleuthrgluaspthrvalproglu177517801785provalgluvalglnlysprohisthrprolyslysilearggly179017951800serserglylysarggluilelystrpaspglygluserleugly180518101815lysthrseralaglygluproleutyrglnglnalaleuvalgly182018251830glyglumetvalalavalglyglyalavalthrleugluvalasp183518401845aspproaspglumetproalailetyrphevalglutyrmetphe185018551860gluserthrasphiscyslysmetleuhisglyargpheleugln186518701875argglysermetthrvalleuglyasnalaalaasngluargglu188018851890leupheleuthrasnglucysmetthrthrglnleulysaspile189519001905lysglyvalalaserphegluileargserargprotrpglyhis191019151920glntyrarglyslysasnilethralaasplysleuasptrpala192519301935argalaleugluarglysvallysaspleuprothrglutyrtyr194019451950cyslysserleutyrserprogluargglyglyphepheserleu195519601965proleuseraspileglyargserserglyphecysthrsercys197019751980lysilearggluaspgluglulysargserthrilelysleuasn198519901995valserlysthrglyphepheileasnglyileglutyrserval200020052010gluaspphevaltyrvalasnproaspserileglyglyleulys201520202025gluglyserlysthrserphelysserglyargasnileglyleu203020352040argalatyrvalvalcysglnleuleugluilevalprolysglu204520502055serarglysalaaspleuglyserpheaspvallysvalargarg206020652070phetyrargprogluaspvalseralaglulysalatyralaser207520802085aspileglngluleutyrpheserglnaspthrvalvalleupro209020952100proglyalaleugluglylyscysgluvalarglyslysserasp210521102115metproleuserargglutyrproileserasphisilephephe212021252130cysaspleuphepheaspthrserlysglyserleulysglnleu213521402145proalaasnmetlysprolyspheserthrilelysaspaspthr215021552160leuleuarglyslyslysglylysglyvalglusergluileglu216521702175sergluilevallysprovalgluproprolysgluileargleu218021852190alathrleuaspilephealaglycysglyglyleuserhisgly219522002205leulyslysalaglyvalseraspalalystrpalaileglutyr221022152220glugluproalaglyglnalaphelysglnasnhisprogluser222522302235thrvalphevalaspasncysasnvalileleuargalailemet224022452250glulysglyglyaspglnaspaspcysvalserthrthrgluala225522602265asngluleualaalalysleuthrglugluglnlysserthrleu227022752280proleuproglyglnvalasppheileasnglyglyproprocys228522902295glnglypheserglymetasnargpheasnglnsersertrpser230023052310lysvalglncysglumetileleualapheleuserphealaasp231523202325tyrpheargproargtyrpheleuleugluasnvalargthrphe233023352340valserpheasnlysglyglnthrpheglnleuthrleualaser234523502355leuleuglumetglytyrglnvalargpheglyileleugluala236023652370glyalatyrglyvalserglnserarglysargalapheiletrp237523802385alaalaalaproglugluvalleuproglutrpproglupromet239023952400hisvalpheglyvalprolysleulysileserleuserglngly240524102415leuhistyralaalavalargserthralaleuglyalaprophe242024252430argproilethrvalargaspthrileglyaspleuproserval243524402445gluasnglyaspserargthrasnlysglutyrlysgluvalala245024552460valsertrppheglnlysgluileargglyasnthrilealaleu246524702475thrasphisilecyslysalametasngluleuasnleuilearg248024852490cyslysleuileprothrargproglyalaasptrphisaspleu249525002505prolysarglysvalthrleuseraspglyargvalgluglumet251025152520ileprophecysleuproasnthralagluarghisasnglytrp252525302535lysglyleutyrglyargleuasptrpglnglyasnpheprothr254025452550servalthraspproglnprometglylysvalglymetcysphe255525602565hisprogluglnhisargileleuthrvalargglucysalaarg257025752580serglnglypheproaspsertyrgluphealaglyasnileasn258525902595hislyshisargglnileglyasnalavalproproproleuala260026052610phealaleuglyarglysleulysglualaleuhisleulyslys261526202625serproglnhisglnpro2630<210>29<211>1113<212>prt<213>rb-vtale-atsrdx氨基酸序列<400>29metaspproileargserargthrproserproalaarggluleuleu151015proglyproglnproaspargvalglnprothralaaspargglygly202530alaproproalaglyglyproleuaspglyleuproalaargargthr354045metserargthrargleuproserproproalaproserproalaphe505560seralaglyserpheasnaspleuleuargglnpheaspproserleu65707580leuaspthrserleuleuaspsermetproalavalglythrprohis859095thralaalaalaproalaglutrpaspgluvalglnserglyleuarg100105110alaalaaspaspproproprothrvalargvalalavalthralaala115120125argproproargalalysproalaproargargargalaalaglnpro130135140seraspalaserproalaalaglnvalaspleuargthrleuglytyr145150155160serglnglnglnglnglulysilelysserlysvalargserthrval165170175alaglnhishisglualaleuvalglyhisglyphethrhisalahis180185190ilevalalaleuserglnhisproalaalaleuglythrvalalaval195200205lystyrglnhisileilethralaleuproglualathrhisgluasp210215220ilevalglyvalglylysglntrpserglyalaargalaleugluala225230235240leuleuthrlysalaglygluleuargglyproproleuglnleuasp245250255thrglyglnleuleulysilealalysargglyglyvalthralaval260265270glualavalhisalaserargasnalaleuthrglyalaproleuasn275280285leuthrproaspglnvalvalalailealaserasnileglyglylys290295300glnalaleugluthrvalglnargleuleuprovalleucysglnala305310315320hisglyleuthrproaspglnvalvalalailealaserasnglygly325330335glylysglnalaleugluthrvalglnargleuleuprovalleucys340345350glnalahisglyleuthrproaspglnvalvalalailealaserasn355360365ileglyglylysglnalaleugluthrvalglnargleuleuproval370375380leucysglnalahisglyleuthrproaspglnvalvalalaileala385390395400serasnglyglyglylysglnalaleugluthrvalglnargleuleu405410415provalleucysglnalahisglyleuthrproaspglnvalvalala420425430ilealaserasnileglyglylysglnalaleugluthrvalglnarg435440445leuleuprovalleucysglnalahisglyleuthrproaspglnval450455460valalailealaserasnglyglyglylysglnalaleugluthrval465470475480glnargleuleuprovalleucysglnalahisglyleuthrproasp485490495glnvalvalalailealaserasnglyglyglylysglnalaleuglu500505510thrvalglnargleuleuprovalleucysglnalahisglyleuthr515520525proaspglnvalvalalailealaserasnasnglyglylysglnala530535540leugluthrvalglnargleuleuprovalleucysglnalahisgly545550555560leuthrproaspglnvalvalalailealaserasnasnglyglylys565570575glnalaleugluthrvalglnargleuleuprovalleucysglnala580585590hisglyleuthrproaspglnvalvalalailealaserasnilegly595600605glylysglnalaleugluthrvalglnargleuleuprovalleucys610615620glnalahisglyleuthrproaspglnvalvalalailealaserhis625630635640aspglyglylysglnalaleugluthrvalglnargleuleuproval645650655leucysglnalahisglyleuthrproaspglnvalvalalaileala660665670serasnileglyglylysglnalaleugluthrvalglnargleuleu675680685provalleucysglnalahisglyleuthrproaspglnvalvalala690695700ilealaserhisaspglyglylysglnalaleugluthrvalglnarg705710715720leuleuprovalleucysglnalahisglyleuthrproaspglnval725730735valalailealaserhisaspglyglylysglnalaleugluthrval740745750glnargleuleuprovalleucysglnalahisglyleuthrproasp755760765glnvalvalalailealaserasnileglyglylysglnalaleuglu770775780thrvalglnargleuleuprovalleucysglnalahisglyleuthr785790795800proaspglnvalvalalailealaserasnileglyglylysglnala805810815leugluthrvalglnargleuleuprovalleucysglnalahisgly820825830leuthrproaspglnvalvalalailealaserasnasnglyglylys835840845glnalaleugluserilevalalaglnleuserargproaspproala850855860leualaalaleuthrasnasphisleuvalalaleualacysleugly865870875880glyargproalaleuaspalavallyslysglyleuprohisalapro885890895gluleuileargargileasnargargileprogluargthrserhis900905910argvalalaaspleualahisvalvalargvalleuglyphephegln915920925serhisserhisproalaglnalapheaspaspalametthrglnphe930935940glymetserarghisglyleuvalglnleupheargargvalglyval945950955960thrglupheglualaargcysglythrleuproproalaserglnarg965970975trpaspargileleuglnalaserglymetlysargalalysproser980985990prothrseralaglnthrproaspglnalaserleuhisalapheala99510001005aspserleugluargaspleuaspalaproserpromethisglu101010151020glyaspglnthrargalaserserarglysargserargserasp102510301035argalavalthrglyproserthrglnglnserphegluvalarg104010451050valprogluglnargaspalaleuhisleuproleusertrparg105510601065vallysargproargthrargileglyglyglyleuproasppro107010751080glythrproilealaalaaspleualaalaserserthrvalmet108510901095trpserserleuaspleuaspleugluleuargleuglypheala110011051110<210>30<211>1751<212>prt<213>rb-vtale-atsuvh2氨基酸序列<400>30metaspproileargserargthrproserproalaarggluleuleu151015proglyproglnproaspargvalglnprothralaaspargglygly202530alaproproalaglyglyproleuaspglyleuproalaargargthr354045metserargthrargleuproserproproalaproserproalaphe505560seralaglyserpheasnaspleuleuargglnpheaspproserleu65707580leuaspthrserleuleuaspsermetproalavalglythrprohis859095thralaalaalaproalaglutrpaspgluvalglnserglyleuarg100105110alaalaaspaspproproprothrvalargvalalavalthralaala115120125argproproargalalysproalaproargargargalaalaglnpro130135140seraspalaserproalaalaglnvalaspleuargthrleuglytyr145150155160serglnglnglnglnglulysilelysserlysvalargserthrval165170175alaglnhishisglualaleuvalglyhisglyphethrhisalahis180185190ilevalalaleuserglnhisproalaalaleuglythrvalalaval195200205lystyrglnhisileilethralaleuproglualathrhisgluasp210215220ilevalglyvalglylysglntrpserglyalaargalaleugluala225230235240leuleuthrlysalaglygluleuargglyproproleuglnleuasp245250255thrglyglnleuleulysilealalysargglyglyvalthralaval260265270glualavalhisalaserargasnalaleuthrglyalaproleuasn275280285leuthrproaspglnvalvalalailealaserasnileglyglylys290295300glnalaleugluthrvalglnargleuleuprovalleucysglnala305310315320hisglyleuthrproaspglnvalvalalailealaserasnglygly325330335glylysglnalaleugluthrvalglnargleuleuprovalleucys340345350glnalahisglyleuthrproaspglnvalvalalailealaserasn355360365ileglyglylysglnalaleugluthrvalglnargleuleuproval370375380leucysglnalahisglyleuthrproaspglnvalvalalaileala385390395400serasnglyglyglylysglnalaleugluthrvalglnargleuleu405410415provalleucysglnalahisglyleuthrproaspglnvalvalala420425430ilealaserasnileglyglylysglnalaleugluthrvalglnarg435440445leuleuprovalleucysglnalahisglyleuthrproaspglnval450455460valalailealaserasnglyglyglylysglnalaleugluthrval465470475480glnargleuleuprovalleucysglnalahisglyleuthrproasp485490495glnvalvalalailealaserasnglyglyglylysglnalaleuglu500505510thrvalglnargleuleuprovalleucysglnalahisglyleuthr515520525proaspglnvalvalalailealaserasnasnglyglylysglnala530535540leugluthrvalglnargleuleuprovalleucysglnalahisgly545550555560leuthrproaspglnvalvalalailealaserasnasnglyglylys565570575glnalaleugluthrvalglnargleuleuprovalleucysglnala580585590hisglyleuthrproaspglnvalvalalailealaserasnilegly595600605glylysglnalaleugluthrvalglnargleuleuprovalleucys610615620glnalahisglyleuthrproaspglnvalvalalailealaserhis625630635640aspglyglylysglnalaleugluthrvalglnargleuleuproval645650655leucysglnalahisglyleuthrproaspglnvalvalalaileala660665670serasnileglyglylysglnalaleugluthrvalglnargleuleu675680685provalleucysglnalahisglyleuthrproaspglnvalvalala690695700ilealaserhisaspglyglylysglnalaleugluthrvalglnarg705710715720leuleuprovalleucysglnalahisglyleuthrproaspglnval725730735valalailealaserhisaspglyglylysglnalaleugluthrval740745750glnargleuleuprovalleucysglnalahisglyleuthrproasp755760765glnvalvalalailealaserasnileglyglylysglnalaleuglu770775780thrvalglnargleuleuprovalleucysglnalahisglyleuthr785790795800proaspglnvalvalalailealaserasnileglyglylysglnala805810815leugluthrvalglnargleuleuprovalleucysglnalahisgly820825830leuthrproaspglnvalvalalailealaserasnasnglyglylys835840845glnalaleugluserilevalalaglnleuserargproaspproala850855860leualaalaleuthrasnasphisleuvalalaleualacysleugly865870875880glyargproalaleuaspalavallyslysglyleuprohisalapro885890895gluleuileargargileasnargargileprogluargthrserhis900905910argvalalaaspleualahisvalvalargvalleuglyphephegln915920925serhisserhisproalaglnalapheaspaspalametthrglnphe930935940glymetserarghisglyleuvalglnleupheargargvalglyval945950955960thrglupheglualaargcysglythrleuproproalaserglnarg965970975trpaspargileleuglnalaserglymetlysargalalysproser980985990prothrseralaglnthrproaspglnalaserleuhisalapheala99510001005aspserleugluargaspleuaspalaproserpromethisglu101010151020glyaspglnthrargalaserserarglysargserargserasp102510301035argalavalthrglyproserthrglnglnserphegluvalarg104010451050valprogluglnargaspalaleuhisleuproleusertrparg105510601065vallysargproargthrargileglyglyglyleuproasppro107010751080glythrproilealaalaaspleualaalaserserthrvalmet108510901095trpserserserthrleuleupropheproaspleuasnleumet110011051110proaspserglnserserthralaglythrthralaglyaspthr111511201125valvalthrglylysleugluvallyssergluproilegluglu113011351140trpglnthrproproserserthrseraspglnseralaasnthr114511501155aspleuilealaglupheileargilesergluleupheargser116011651170alaphelysproleuglnvallysglyleuaspglyvalserval117511801185tyrglyleuaspserglyalailevalalavalproglulysglu119011951200asnarggluleuilegluproproproglyphelysaspasnarg120512101215valserthrvalvalvalserprolysphegluargproargglu122012251230leualaargilealaileleuglyhisgluglnarglysgluleu123512401245argglnvalmetlysargthrargmetthrtyrgluserleuarg125012551260ilehisleumetalaglusermetlysasnhisvalleuglygln126512701275glyargargargargseraspmetalaalaalatyrilemetarg128012851290aspargglyleutrpleuasntyrasplyshisilevalglypro129513001305valthrglyvalgluvalglyaspilephephetyrargmetglu131013151320leucysvalleuglyleuhisglyglnthrglnalaglyileasp132513301335cysleuthralagluargseralathrglygluproilealathr134013451350serilevalvalserglyglytyrgluaspaspgluaspthrgly135513601365aspvalleuvaltyrthrglyhisglyglyglnasphisglnhis137013751380lysglncysaspasnglnargleuvalglyglyasnleuglymet138513901395gluargsermethistyrglyilegluvalargvalilearggly140014051410ilelystyrgluasnserileserserlysvaltyrvaltyrasp141514201425glyleutyrlysilevalasptrptrpphealavalglylysser143014351440glypheglyvalphelyspheargleuvalargilegluglygln144514501455prometmetglyseralavalmetargphealaglnthrleuarg146014651470asnlysprosermetvalargprothrglytyrvalserpheasp147514801485leuserasnlyslysgluasnvalprovalpheleutyrasnasp149014951500valaspglyaspglngluproarghistyrglutyrilealalys150515101515alavalpheproproglyilepheglyglnglyglyileserarg152015251530thrglycysglucyslysleusercysthraspaspcysleucys153515401545alaarglysasnglyglygluphealatyraspaspasnglyhis155015551560leuleulysglylyshisvalvalpheglucysglygluphecys156515701575thrcysglyprosercyslysserargvalthrglnlysglyleu158015851590argasnargleugluvalpheargserlysgluthrglytrpgly159516001605valargthrleuaspleuileglualaglyalapheilecysglu161016151620tyralaglyvalvalvalthrargleuglnalagluileleuser162516301635metasnglyaspvalmetvaltyrproglyargphethraspgln164016451650trpargasntrpglyaspleuserglnvaltyrproasppheval165516601665argproasntyrproserleuproproleuaspphesermetasp167016751680valserargmetargasnvalalacystyrileserhisserlys168516901695gluproasnvalmetvalglnphevalleuhisasphisasnhis170017051710leumetpheproargvalmetleuphealaleugluasnileser171517201725proleualagluleuserleuasptyrglyleualaaspgluval173017351740asnglylysleualailecysasn17451750当前第1页12