CN114450404A - 来自荞麦的c-糖苷化酶基因及其应用 - Google Patents
来自荞麦的c-糖苷化酶基因及其应用 Download PDFInfo
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Abstract
本发明提供一种具有蓝色系花色的转化植物或其自繁或者杂交后代,或者它们的营养繁殖体、植物体的一部分、组织或细胞。本发明通过将来自荞麦的C‑糖苷化酶(CGT)基因或其同源基因导入宿主植物,使花翠素型花色素苷与黄酮单‑C‑糖苷在植物的细胞内共存。
Description
技术领域
本发明涉及来自荞麦的C-糖苷化酶(CGT)基因或其同源基因,及包含使用这些而使花翠素型花色素苷(Anthocyanin)与黄酮单-C-糖苷在植物的细胞内共存的工序的用于制作具有蓝色系花色的转化植物的方法。
背景技术
蔷薇、菊花、康乃馨等为世界性产业上重要的花卉。尤其蔷薇为最受欢迎的花卉植物,有从公元前开始就已经进行栽培的记录,跨越数百年经历了人为的品种改良。然而,由于在可杂交的近缘种中没有蓝色系的花色的野生种等问题,在以往的杂交育种、突变育种中制作具有蓝色系花色的蔷薇品种存在困难。全新的蓝色系花色的创造,伴随花卉的利用场合的扩大而唤起的新需要,且关系到生产或消费的扩大。于是,尝试了通过基因工程的方法创造具有蓝色系花色的蔷薇。
例如,已知在由紫色至蓝色的花中,富含以花翠素、矮牵牛素及二甲花翠素为骨架的花翠素型花色素苷,然而由于蔷薇等花卉无法生产这样的花翠素型花色素苷,因此正在进行通过使他们的合成所必需的类黄酮3’,5’-羟化酶基因表达,而人为地生产花翠素的研究(非专利文献1)。然而,即使为了在转基因植物中使生产目标物质的酶基因进行表达而人为地改变植物的代谢,但多数情况下目标物质的累积完全或者几乎不发生。
进一步,花色除了根据花色素苷自身的结构,还根据共存的类黄酮(称作辅色素)或金属离子、液泡的pH等而变化。黄酮或黄酮醇为代表性的辅色素,通过与花色素苷堆积成三明治状,具有使花色素苷变蓝,看起来浓的效果(非专利文献2)。已知这是辅色素效果。尤其已知黄酮表现出强辅色素效果,例如报告称在转基因康乃馨的分析中黄酮表现出显著的辅色素效果(非专利文献3)。此外,报告称在荷兰鸢尾中,相对于总花翠素量的总黄酮量的比越高越表现出辅色素效果,颜色越蓝(非专利文献4)。
然而,并不是所有的植物都能够生产黄酮,蔷薇或矮牵牛等并不累积黄酮。从而,正在进行使编码具有由黄烷酮合成黄酮的活性的蛋白质的基因在这样的植物中表达进而改变花色的尝试(专利文献1)。
此外,已知在植物中,黄酮除游离状态以外还以糖苷的形式分布,主要生成黄酮O-糖苷和黄酮C-糖苷,特别是黄酮C-糖苷表现出强辅色素效果。例如,报告称作为黄酮C-糖苷的一种的异牡荆素,在花菖蒲(Iris ensata Thunb.)中对于花色素苷表现出辅色素效果,通过对花色素苷进行稳定化,可使花色蓝色化(非专利文献5)。作为黄酮C-糖苷的生物合成路径之一,已知通过黄烷酮2-羟化酶(F2H)及C-糖苷化酶(CGT)、脱水酶(FDH)催化的反应由黄烷酮合成(非专利文献6)。
至今为止报告了将来自风铃草的F3’5’H基因和来自蓝猪耳的MT基因、来自甘草的F2H基因、来自稻子的CGT基因、来自百脉根的FDH基因导入,通过使花翠素型花色素苷与黄酮C-糖苷在植物细胞内共存,从而制作具有蓝色系花色的蔷薇(专利文献2)。然而,这样制作的蔷薇系统的花色有强烈的发红的倾向,从而依然期望开发用于可制作均匀且稳定地具有更蓝的花色的蔷薇的蓝色表达控制技术。
专利文献
专利文献1:日本特开2000-279182号公报
专利文献2:国际公开第2019/069946号
专利文献3:国际公开第2008/156206号
非专利文献
非专利文献1:Phytochemistry Reviews 5,283-291
非专利文献2:Prog.Chem.Org.Natl.Prod.52
非专利文献3:Phytochemistry,63,15-23(2003)
非专利文献4:Plant Physiol.Bioch.72,116-124(2013)
非专利文献5:Euphytica 115,1-5(2000)
非专利文献6:FEBS Lett.589,182-187(2015)
发明内容
本发明所要解决的课题在于,通过查明花色中发红的原因,而制作均匀且稳定地具有蓝色系花色(RHS色谱标准第5版:Violet-Blue组/Blue组且/或色相角度:339.7°~270.0°)的转化植物。
本申请发明者等,为了解决上述课题进行了深入研究,重复进行试验,结果发现,发红的原因在于黄酮二-C-糖苷,就与花翠素型花色素苷的辅色素效果而言,黄酮单-C-糖苷显著高于黄酮二-C-糖苷。进一步,本申请发明者等,在各种CGT基因之中,通过导入来自荞麦的CGT基因,成功地在植物的花瓣中,仅使黄酮单-C-糖苷显著累积。基于这样的见识,从而完成了本发明。
本发明如下所示。
[1]一种来自荞麦的CGT基因或其同源基因,其为来自荞麦的C-糖苷化酶,即CGT基因或其同源基因,其特征在于,所述来自荞麦的CGT基因或其同源基因选自:
(1-a)由序列号11的碱基序列构成的多聚核苷酸;
(1-b)在严谨条件下与由与序列号11的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(1-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(1-c)编码由序列号12的氨基酸序列构成的蛋白质的多聚核苷酸。
[2]根据1所述的来自荞麦的CGT基因或其同源基因,其特征在于,附加有来自拟南芥ADH基因的非翻译区域(5’-UTR),即序列号15,或来自拟南芥HSPRO基因的非翻译区域(5’-UTR),即序列号13。
[3]一种载体,其为含有来自荞麦的CGT基因或其同源基因的载体,其特征在于,所述来自荞麦的CGT基因或其同源基因选自:
(1-a)由序列号11的碱基序列构成的多聚核苷酸;
(1-b)在严谨条件下与由与序列号11的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(1-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(1-c)编码由序列号12的氨基酸序列构成的蛋白质的多聚核苷酸;
(1-d)编码由在序列号12的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(1-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(1-e)编码具有相对于序列号12的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(1-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸。
[4]根据3所述的载体,其特征在于,在所述来自荞麦的CGT基因或其同源基因上附加有来自拟南芥ADH基因的非翻译区域(5’-UTR),即序列号15,或来自拟南芥HSPRO基因的非翻译区域(5’-UTR),即序列号13。
[5]根据3或4所述的载体,其特征在于,进一步含有黄烷酮2-羟化酶,即F2H基因或其同源基因,以及脱水酶,即FDH基因或其同源基因。
[6]根据5所述的载体,其特征在于,所述F2H基因或其同源基因选自:
(2-a)由序列号5的碱基序列构成的多聚核苷酸;
(2-b)在严谨条件下与由与序列号5的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(2-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(2-c)编码由序列号6的氨基酸序列构成的蛋白质的多聚核苷酸;
(2-d)编码由在序列号6的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(2-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(2-e)编码具有相对于序列号6的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(2-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸,
所述FDH基因或其同源基因选自:
(3-a)由序列号9的碱基序列构成的多聚核苷酸;
(3-b)在严谨条件下与由与序列号9的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(3-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(3-c)编码由序列号10的氨基酸序列构成的蛋白质的多聚核苷酸;
(3-d)编码由在序列号10的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(3-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(3-e)编码具有相对于序列号10的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(3-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸。
[7]根据3~6中任一项所述的载体,其特征在于,进一步含有类黄酮F3’5’羟化酶,即F3’5’H基因或其同源基因,以及甲基转移酶,即MT基因或其同源基因。
[8]根据7所述的载体,其特征在于,所述F3’5’H基因或其同源基因选自:
(4-a)由序列号1的碱基序列构成的多聚核苷酸;
(4-b)在严谨条件下与由与序列号1的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(4-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(4-c)编码由序列号2的氨基酸序列构成的蛋白质的多聚核苷酸;
(4-d)编码由在序列号2的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(4-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(4-e)编码具有相对于序列号2的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(4-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸,且,
所述MT基因或其同源基因选自:
(5-a)由序列号3的碱基序列构成的多聚核苷酸;
(5-b)在严谨条件下与由与序列号3的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(5-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(5-c)编码由序列号4的氨基酸序列构成的蛋白质的多聚核苷酸;
(5-d)编码由在序列号4的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(5-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(5-e)编码具有相对于序列号4的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(5-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸。
[9]一种转化植物或其自繁或者杂交后代,其特征在于,含有1或2所述的来自荞麦的CGT基因或其同源基因,或者3~8中任一项所述的载体。
[10]根据9所述的转化植物或其自繁或者杂交后代,其特征在于,所述植物选自蔷薇、菊花、康乃馨或百合。
[11]根据10所述的转化植物或其自繁或者杂交后代,其特征在于,所述植物为蔷薇。
[12]一种营养繁殖体、植物体的一部分、组织或细胞,其特征在于,为9~11中任一项所述的转化植物或其自繁或者杂交后代的营养繁殖体、植物体的一部分、组织或细胞。
[13]一种切花,或由该切花制成的加工品,其特征在于,为9~11中任一项所述的转化植物或其自繁或者杂交后代的切花,或由该切花制成的加工品。
[14]一种方法,其为用于制作具有蓝色系花色的转化植物的方法,其特征在于,包含通过将来自荞麦的C-糖苷化酶,即CGT基因或其同源基因导入宿主植物,使花翠素型花色素苷与黄酮单-C-糖苷在植物的细胞内共存的工序。
[15]根据14所述的方法,其特征在于,所述黄酮单-C-糖苷为芹菜素6-C-糖苷、木犀草素6-C-糖苷、三粒小麦黄酮6-C-糖苷、芹菜素8-C-糖苷、木犀草素8-C-糖苷、三粒小麦黄酮8-C-糖苷或它们的衍生物。
[16]根据14或15所述的方法,其特征在于,所述花翠素型花色素苷选自二甲花翠素、花翠素、矮牵牛素及它们的组合。
[17]根据14~16中任一项所述的方法,其特征在于,包含将1或2所述的来自荞麦的CGT基因或其同源基因,或者3~8中任一项所述的载体导入宿主植物的细胞内的工序。
[18]根据14~17中任一项所述的方法,其特征在于,所述植物选自蔷薇、菊花、康乃馨或百合。
[19]根据18所述的方法,其特征在于,所述植物为蔷薇。
根据本发明,可制作均匀且稳定地具有蓝色系花色(RHS色谱标准第5版:Violet-Blue组/Blue组且/或色相角度:339.7°~270.0°)的植物品种。
附图说明
图1表示植物中的黄酮单-C-糖苷的生物合成路径。
图2表示pSPB6486的结构。
图3表示pSPB7473的结构。
图4表示pSPB7473的详细结构。
图5表示pSPB7472的详细结构。
图6表示pSPB7808的详细结构。
图7表示pSPB7809的详细结构。
具体实施方式
已知花色素苷是植物中广泛存在的色素群,呈现红色、蓝色、紫色的花色。根据作为糖苷配基的花青素(Anthocyanidins)部位的B环的羟基数,分类为天竺葵素、矢车菊素及花翠素的3个系统。发色团为糖苷配基部分,天竺葵素型花色素苷呈现橙色、矢车菊素型花色素苷呈现红色、花翠素型花色素苷呈现紫~蓝色。本申请说明书中,例如“花翠素型花色素苷”可列举以花翠素、二甲花翠素、矮牵牛素为骨架的它们的衍生物,优选为二甲花翠素。
通过使花翠素型花色素苷与黄酮、黄酮醇、有机酸酯类、单宁类等的物质共存,与它们进行分子间相互作用从而有时表达带蓝色的颜色。该现象被称作辅色素作用,引起该现象的物质被称作辅色素(辅助色素)。在辅色素作用中不仅有引起蓝色表达的深色效果,还有浓色效果、提高颜色稳定性的效果。本发明者们确认通过花翠素型花色素苷和黄酮C-糖苷的辅色素作用,蔷薇的花瓣表达蓝色(专利文献2)。
黄酮是有机化合物的一种,为黄烷衍生物的环状酮,在植物内主要作为糖苷存在。黄酮狭义上是指化学式C15H10O2、分子量222.24的化合物,2,3-二脱氢黄烷-4-酮,广义的黄酮(黄酮类)是类黄酮的分类之一,在类黄酮中以黄酮结构为基本骨架,进一步在3位上不具有羟基的类黄酮被分类为“黄酮”。本申请说明书中,“黄酮C-糖苷”是指广义的黄酮,即,属于黄酮类的衍生物的糖苷中,醛糖的端基碳上直接键合糖苷配基的糖苷。作为黄酮C-糖苷,可列举木犀草素C-糖苷、三粒小麦黄酮C-糖苷、芹菜素C-糖苷、金合欢素C-糖苷,但并不限定于此。黄酮C-糖苷另外还包含芹菜素、木犀草素、三粒小麦黄酮、金合欢素衍生物的糖苷。在植物中,作为黄酮C-糖苷的生物合成路径之一,已知示于图1的路径。该路径中,通过F2H、CGT及FDH的作用,来生产黄酮C-糖苷。
在这样的合成路径中,除黄酮单-C-糖苷外,维采宁(Vicenin)-2(芹菜素6,8-二-C-糖苷)等黄酮二-C-糖苷也被合成,而发明者等,此次惊人地发现发红的原因在于黄酮二-C-糖苷,就与花翠素型花色素苷的辅色素效果而言,黄酮单-C-糖苷高于黄酮二-C-糖苷。为了制作均匀且稳定地具有蓝色系花色的转化植物,需要极力减少黄酮二-C-糖苷的累积,而仅使黄酮单-C-糖苷在花瓣中显著累积。黄酮单-C-糖苷,典型而言为黄酮6-C-糖苷或黄酮8-C-糖苷,优选为黄酮6-C-糖苷。作为这样的黄酮单-C-糖苷,例如可列举芹菜素6-C-糖苷(异牡荆素)、木犀草素6-C-糖苷(异荭草素)、三粒小麦黄酮6-C-糖苷、芹菜素8-C-糖苷(牡荆素)、木犀草素8-C-糖苷(荭草素)、三粒小麦黄酮8-C-糖苷或它们的衍生物。
在植物的细胞内,黄酮C-糖苷的累积可通过以含有上述合成路径中所必需的基因(即,F2H基因、CGT基因及FDH基因)或它们的同源基因的载体对宿主植物进行转化来达成。且,通过作为CGT基因,使用来自荞麦的CGT基因或其同源基因,尤其是作为翻译增强序列附加有来自拟南芥ADH基因的非翻译区域(5’-UTR)(序列号15)或来自拟南芥HSPRO基因的非翻译区域(5’-UTR)(序列号13)的来自荞麦的CGT基因或其同源基因,与黄酮二-C-糖苷相比,可使黄酮单-C-糖苷在花瓣中显著累积。
所述来自荞麦的CGT基因或其同源基因选自以下多聚核苷酸:
(1-a)由序列号11的碱基序列构成的多聚核苷酸;
(1-b)在严谨条件下与由与序列号11的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(1-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(1-c)编码由序列号12的氨基酸序列构成的蛋白质的多聚核苷酸;
(1-d)编码由在序列号12的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(1-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(1-e)编码具有相对于序列号12的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(1-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸。
所述F2H基因或其同源基因,只要具有所期望的功能即可,关于其起源并无特别限制,优选为来自甘草的F2H基因或其同源基因,选自以下多聚核苷酸:
(2-a)由序列号5的碱基序列构成的多聚核苷酸;
(2-b)在严谨条件下与由与序列号5的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(2-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(2-c)编码由序列号6的氨基酸序列构成的蛋白质的多聚核苷酸;
(2-d)编码由在序列号6的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(2-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(2-e)编码具有相对于序列号6的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(2-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸。
所述FDH基因或其同源基因,只要具有所期望的功能即可,关于其起源并无特别限制,优选为来自百脉根的FDH基因或其同源基因,选自以下多聚核苷酸:
(3-a)由序列号9的碱基序列构成的多聚核苷酸;
(3-b)在严谨条件下与由与序列号9的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(3-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(3-c)编码由序列号10的氨基酸序列构成的蛋白质的多聚核苷酸;
(3-d)编码由在序列号10的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(3-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(3-e)编码具有相对于序列号10的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(3-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸。
植物的细胞内的花翠素型花色素苷的累积,可通过将类黄酮F3’5’羟化酶(F3’5’H)基因或其同源基因及甲基转移酶(MT)基因或其同源基因编入宿主植物来达成(专利文献3)。从而,除上述黄酮单-C-糖苷的合成路径中的必需基因或它们的同源基因外,通过以进一步含有F3’5’H基因或其同源基因及MT基因或其同源基因的载体对宿主植物进行转化,可使花翠素型花色素苷与黄酮单-C-糖苷在宿主植物的细胞内共存。
F3’5’H基因或其同源基因,只要具有所期望的功能即可,关于其起源并无特别限制,优选为来自风铃草的F3’5’H基因或其同源基因,选自以下多聚核苷酸:
(4-a)由序列号1的碱基序列构成的多聚核苷酸;
(4-b)在严谨条件下与由与序列号1的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(4-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(4-c)编码由序列号2的氨基酸序列构成的蛋白质的多聚核苷酸;
(4-d)编码由在序列号2的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(4-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(4-e)编码具有相对于序列号2的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(4-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸。
MT基因或其同源基因,只要具有所期望的功能即可,关于其起源并无特别限制,优选为来自蓝猪耳的MT基因或其同源基因,选自以下多聚核苷酸:
(5-a)由序列号3的碱基序列构成的多聚核苷酸;
(5-b)在严谨条件下与由与序列号3的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(5-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(5-c)编码由序列号4的氨基酸序列构成的蛋白质的多聚核苷酸;
(5-d)编码由在序列号4的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(5-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(5-e)编码具有相对于序列号4的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(5-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸。
所述MT基因或其同源基因中,作为翻译增强序列可附加来自拟南芥ADH基因的非翻译区域(5’-UTR)(序列号15)或来自拟南芥HSPRO基因的非翻译区域(5’-UTR)(序列号13)。
本说明书中,用语“多聚核苷酸”是指DNA或RNA。
本说明书中,所谓用语“严谨条件”为使多聚核苷酸或寡核苷酸能与基因组DNA进行选择性且可检出的特异性结合的条件。严谨条件通过盐浓度、有机溶剂(例如,甲酰胺)、温度及其他公知的条件的适当组合进行定义。即,通过降低盐浓度,或增加有机溶剂浓度,或者提高杂交温度增加严谨度(stringency)。进一步杂交后的洗净条件也影响严谨度。该洗净条件也可根据盐浓度和温度进行定义,通过盐浓度的减少和温度的上升增加洗净的严谨度。从而,所谓用语“严谨条件”,是指仅在各碱基序列间的“一致性”程度,例如,全体平均为约80%以上,优选为约90%以上,更优选为约95%以上,进一步优选为97%以上,最优选为98%以上的具有高度一致性的碱基序列之间,进行特异性杂交的条件。作为“严谨条件”,例如可列举在温度60℃~68℃下,钠浓度150~900mM,优选600~900mM,pH6~8的条件,作为具体例可列举,在5×SSC(750mM NaCl、75mM柠檬酸三钠)、1%SDS、5×Denhard溶液50%甲醛及42℃的条件下进行杂交,在0.1×SSC(15mM NaCl、1.5mM柠檬酸三钠)、0.1%SDS及55℃的条件下进行洗净。
杂交可根据例如,分子生物学现代实验室指南(Current protocols inmolecular biology(edited by Frederick M.Ausubel et al.,1987))中记载的方法等业界公知的方法或以其为基准的方法进行实施。此外,使用市售的库时,可根据附加的使用说明书中记载的方法进行实施。作为由这样的杂交而被选择的基因,可为来自天然的基因,例如来自植物的基因,也可为来自植物以外的基因。此外,由杂交而被选择的基因可为cDNA,也可为基因组DNA,还可为化学合成的DNA。
本说明书中,所谓“1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列”,是指例如1~20个,优选1~5个,更优选1~3个任意数量的氨基酸缺失、取代、插入及/或附加而得的氨基酸序列。基因工程的方法之一的部位特定诱导突变法为在特定的位置导入特定的突变的方法,因而有用,可依据Molecular Cloning:A Laboratory Manual,2ndEd.,Cold Spring Harbor Laboratory Press,Cold Spring Harbor,N.Y.,1989等中记载的方法来实施。通过使用适当的表达系使该突变DNA表达,可得到由1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成的蛋白质。
此外,多聚核苷酸可通过本领域技术人员所公知的方法,例如通过亚磷酰胺法等进行化学合成的方法,以植物的核酸样本为模板,使用基于目标基因的核苷序列而设计的引物的核酸扩增法等来获得。
本说明书中,所谓用语“一致性”,是指多肽序列(或氨基酸序列)或多聚核苷酸序列(或碱基序列)中的2条链之间,构成该链的各氨基酸残基彼此或者各碱基彼此的适合关系中可确定为一致的物质的量(数目),为表示两个多肽序列或两个多聚核苷酸序列间的序列相关性的程度的用语,“一致性”可简单地计算。已知许多测定两个多聚核苷酸序列或多肽序列间的一致性的方法,用语“一致性”对于本领域技术人员是周知的(例如,参考Lesk,A.M.(Ed.),Computational Molecular Biology,Oxford University Press,New York,(1988);Smith,D.W.(Ed.),Biocomputing:Informatics and Genome Projects,AcademicPress,New York,(1993);Grifin,A.M.&Grifin,H.G.(Ed.),Computer Analysis ofSequence Data:Part I,Human Press,New Jersey,(1994);von Heinje,G.,SequenceAnalysis in Molecular Biology,Academic Press,New York,(1987);Gribskov,M.&Devereux,J.(Ed.),Sequence Analysis Primer,M-Stockton Press,New York,(1991)等)。
此外,本说明书中记载的“一致性”的数值,除特别明示时以外,可为使用本领域技术人员公知的一致性检索程序进行计算的数值,优选为使用MacVector应用(版本9.5Oxford Molecular Ltd.,Oxford,England)的ClustalW程序计算的数值。在本发明中,各多聚核苷酸序列间或各氨基酸序列间的“一致性”的程度例如为约90%以上,优选为约95%以上,进一步优选为约97%以上,最优选为约98%以上。
本发明的多聚核苷酸(核酸、基因)为“编码”着眼的蛋白质的物质。在此,所谓“编码”是指使着眼的蛋白质以具备其活性的状态进行表达。此外,所谓“编码”含有将着眼的蛋白质作为连续的结构序列(外显子)进行编码或通过间插序列(内含子)进行编码两种意义。
具有天然的碱基序列的基因例如可通过由DNA测序仪进行解析来得到。此外,编码具有经修饰的氨基酸序列的酶的DNA能够以具有天然的碱基序列的DNA作为基础,使用常用的部位特定诱导突变法、PCR法来合成。例如,通过天然的cDNA或基因组DNA的限制酶处理来得到欲修饰的DNA片段,并以其为模板使用导入了所期望的突变的引物来实施部位特定诱导突变法、PCR法,从而得到所期望的修饰后的DNA片段。其后,只要将导入了该突变的DNA片段与编码作为目标的酶的其他部分的DNA片段进行连接即可。
或者为了得到编码由缩短的氨基酸序列组成的酶的DNA,例如,将编码比作为目标的氨基酸序列长的氨基酸序列,例如,通过所期望的限制酶对编码全长氨基酸序列的DNA进行切断,其结果所得到的DNA片段不能编码作为目标的氨基酸序列整体时,只要合成由不足部分的序列组成的DNA片段并进行连接即可。
此外,通过使用大肠杆菌及酵母中的基因表达系使得到的多聚核苷酸表达并测定酶活性,可确认得到的多聚核苷酸对具有期望的活性的蛋白质进行编码。
本发明还涉及含有上述多聚核苷酸的(转基因)载体,特别是表达载体,进一步涉及通过该载体而转化的植物。
此外,本发明的载体,依赖于导入他们的宿主植物的种类而含有表达控制区域,例如,启动子、终止子、复制起点等。作为使在植物细胞内多聚核苷酸组成型表达的启动子的例子,可列举花椰菜花叶病毒的35S启动子、连接两个35S启动子的增强区域的El235S启动子、rd29A基因启动子、rbcS启动子、mac-1启动子等。此外,为了组织特异性的基因表达,可使用在该组织中进行特异性表达的基因的启动子。
载体的制作,可依照使用限制酶、连接酶等的常用方法进行实施。此外,基于表达载体的宿主植物的转化也可依照常用方法进行实施。
在目前的技术水平下,可利用将多聚核苷酸导入植物并使该多聚核苷酸组成型或组织特异性表达的技术。向植物中的DNA的导入,可通过本领域技术人员公知的方法例如农杆菌法、双元载体法、电穿孔法、PEG法、粒子枪法等来进行。
本发明中,可用作宿主的植物并无特别限定,可使用蔷薇科蔷薇属、菊花科菊花属、石竹科石竹属(康乃馨等)、百合科百合属的植物,特别优选为蔷薇科蔷薇属的栽培蔷薇(学名Rosa hybrida)。另外,所谓本说明书中使用的用语“蔷薇植物”,为在分类学上的定位中蔷薇科蔷薇属的栽培蔷薇(学名Rosa hybrida)。蔷薇根据树形和花的大小,主要分为杂交茶香月季(Hybrid tea)类、丰花月季(Floribunda)类、西洋樱草(Polyantha)类等,在任一系统中花瓣中所含的主要的色素(花色素苷)皆仅为矢车菊素型和天竺葵素型这两种。在本发明中,对于可用作宿主的蔷薇植物的种类无特别限定,可适当使用这些品种或系统。例如,作为可用作宿主的蔷薇品种可列举海洋之歌(Ocean song)、贵族(Noblesse)、Rita·Perfumera、冷水(Cool water)、Fame、Topless、蜜桃雪山(Peach avalanche)等。
根据本发明可制作均匀且稳定地具有蓝色系花色的转化植物,优选为蔷薇科蔷薇属、菊花科菊花属或石竹科石竹属(康乃馨等),特别优选为蔷薇植物。获得的转化植物表现出RHS色谱标准下的Blue组或Violet-Blue组,且/或CIEL*a*b*表色系的色相角度339.7°~270.0°,优选315°以下的花色。
进一步,本发明还涉及通过上述方法获得的转化植物或其自繁或者杂交后代的切花、它们的营养繁殖体、植物的一部分、组织或细胞、或者由该切花制成的加工品(尤其是切花加工品)。在此作为切花加工品,包括使用该切花的压花、保鲜花、干花、树脂密封品等,但并不限定于此。
以下通过实施例对本发明进行具体地说明。
实施例
[实施例1:黄酮C-糖苷相对于花色素苷(二甲花翠苷)的辅色素效果的模拟]
为了进行黄酮C-糖苷相对于花色素苷(二甲花翠苷)的辅色素效果的模拟,对二甲花翠苷及黄酮C-糖苷进行调整。本实验中使用的二甲花翠苷(二甲花翠素3,5-二糖苷)及黄酮C-糖苷(牡荆素(芹菜素8-C-糖苷)、异牡荆素(芹菜素6-C-糖苷)、荭草素(木犀草素8-C-糖苷)、异荭草素(木犀草素6-C-糖苷)、维采宁-2(芹菜素6,8-二-C-糖苷))由NacalaiTesque株式会社购入。
相对于如此获得的二甲花翠苷,在pH5.0的缓冲液中,以5当量的摩尔浓度比添加各黄酮C-糖苷(牡荆素、异牡荆素、荭草素、异荭草素、维采宁-2),测定吸收光谱。二甲花翠苷的浓度设为0.5mM。
[表1]
添加黄酮C-糖苷时的二甲花翠苷水溶液的吸收极值(λmax)及色相角度(°)
通过添加黄酮C-糖苷,二甲花翠苷水溶液的吸光度增加,添加任一种黄酮C-糖苷时,与二甲花翠苷单独的情况相比,吸收极值(λmax)均向长波长侧移动。此外,其效果为确认以异牡荆素>异荭草素>荭草素>牡荆素>维采宁-2的顺序,吸收极值向长波长侧移动。进一步确认,关于色相角度同样以异牡荆素>异荭草素>荭草素>牡荆素>维采宁-2的顺序变小。由上明确黄酮单-C-糖苷的辅色素效果比黄酮二-C-糖苷高。进一步明确黄酮单-C-糖苷中,黄酮6-C-糖苷的辅色素效果尤其高。
[实施例2:向蔷薇品种“海洋之歌”中导入来自风铃草的F3’5’H基因和来自蓝猪耳的MT基因、来自甘草的F2H基因、来自稻子的改变密码子使用的CGT基因、来自百脉根的FDH基因]
pSPB6486以pBINPLUS为基本骨架,并含有以下5种表达盒。
(1)El235S启动子和来自风铃草的F3’5’H全长cDNA(序列号1)和D8终止子
(2)El235S启动子和来自蓝猪耳的MT全长cDNA(序列号3)和NOS终止子
(3)35S启动子和来自甘草的F2H全长cDNA(序列号5)和来自紫苏的AT终止子
(4)35S启动子和来自稻子的改变密码子使用的CGT全长cDNA(序列号7)和来自拟南芥的HSP终止子
(5)35S启动子和来自百脉根的FDH全长cDNA(序列号9)和来自拟南芥的HSP终止子
本质粒在植物中,对风铃草的F3’5’H基因和蓝猪耳的MT基因、甘草的F2H基因、稻子的改变密码子使用的CGT基因、百脉根的FDH基因进行组成型表达。
将如此制作的pSPB6486导入蓝色系蔷薇品种“海洋之歌”中,获得总计27个体的转化体。对它们进行色素分析的结果,在26个体中确认二甲花翠素的累积,二甲花翠素含有率最高为74.5%(平均57.0%)。进一步,作为黄酮C-糖苷,实施异牡荆素(芹菜素6-C-糖苷)、牡荆素(芹菜素8-C-糖苷)、异荭草素(木犀草素6-C-糖苷)、荭草素(木犀草素8-C-糖苷)、维采宁-2(芹菜素6,8-C-二糖苷)的鉴定及定量。在检测出二甲花翠素的所有个体中可检出黄酮C-糖苷,作为主成分检测出作为黄酮二-C-糖苷的维采宁-2。生成量的顺序为维采宁-2>异牡荆素>牡荆素>异荭草素>荭草素,总量最高为每花瓣新鲜重量1g中1.563mg。此外,黄酮C-糖苷的总量相对于二甲花翠素为约10倍以上的生成量。
代表性转化体的分析值示于下表2。
[表2]
宿主:海洋之歌
Del:花翠素、Cya:矢车菊素、Pet:矮牵牛素、Pel:天竺葵素、Mal:二甲花翠素
M:杨梅素、Q:槲皮素、K:山奈酚
Tri:五羟黄酮、Lut:木犀草素、Api:芹菜素、Vic2:维采宁-2、VX:牡荆素、IVX:异牡荆素、Ori:荭草素、Iori:异荭草素
Mal(%):总花青素中的二甲花翠素的比例
[实施例3:向蔷薇品种“海洋之歌”中导入来自风铃草的F3’5’H基因和来自蓝猪耳的MT基因、来自甘草的F2H基因、来自荞麦的改变密码子使用的CGT基因、来自百脉根的FDH基因]
pSPB7473以pBINPLUS为基本骨架,并含有以下5种表达盒。
(1)El235S启动子和来自风铃草的F3’5’H全长cDNA(序列号1)和D8终止子
(2)El235S启动子和来自蓝猪耳的MT全长cDNA(序列号3)(在5’位侧附加来自拟南芥HSPRO基因的5’-UTR(序列号13))和来自拟南芥的HSP终止子
(3)El235S启动子和来自甘草的F2H全长cDNA(序列号5)和来自紫苏的AT终止子
(4)El235S启动子和来自荞麦的改变密码子使用的CGT全长cDNA(序列号11)(在5’位侧附加来自拟南芥HSPRO基因的5’-UTR(序列号13))和来自拟南芥的HSP终止子
(5)El235S启动子和来自百脉根的FDH全长cDNA(序列号9)和来自拟南芥的HSP终止子
本质粒在植物中,对风铃草的F3’5’H基因和蓝猪耳的MT基因、甘草的F2H基因、荞麦的改变密码子使用的CGT基因、百脉根的FDH基因进行组成型表达。
将如此制作的pSPB7473导入蓝色系蔷薇品种“海洋之歌”中,获得总计35个体的转化体。对它们进行色素分析的结果,在21个体中确认二甲花翠素的累积,二甲花翠素含有率最高为20.8%(平均9.1%)。进一步,作为黄酮C-糖苷,实施异牡荆素(芹菜素6-C-糖苷)、牡荆素(芹菜素8-C-糖苷)、异荭草素(木犀草素6-C-糖苷)、荭草素(木犀草素8-C-糖苷)、维采宁-2(芹菜素6,8-C-二糖苷)的鉴定及定量。在检测出二甲花翠素的所有个体中可检出黄酮C-糖苷,作为主成分检测出作为黄酮6-C-糖苷的异牡荆素。生成量的顺序为异牡荆素>牡荆素>维采宁-2>异荭草素>荭草素,总量最高为每花瓣新鲜重量1g中7.418mg,为非常高的含量。与实施例2中所示的OS/6486系统相比,几乎所有个体表现出黄酮C-糖苷的总量为每花瓣新鲜重量1g中3mg以上的高含量,相对于花翠素为约100倍以上的生成量。
代表性转化体的分析值示于下表3。
[表3]
宿主:海洋之歌
Del:花翠素、Cya:矢车菊素、Pet:矮牵牛素、Pel:天竺葵素、Mal:二甲花翠素
M:杨梅素、Q:槲皮素、K:山奈酚
Tri:五羟黄酮、Lut:木犀草素、Api:芹菜素、Vic2:维采宁-2、VX:牡荆素、IVX:异牡荆素、Ori:荭草素、Iori:异荭草素
Mal(%):总花青素中的二甲花翠素的比例
[实施例4:向蔷薇品种“海洋之歌”中导入来自风铃草的F3’5’H基因和来自蓝猪耳的MT基因、来自甘草的F2H基因、来自荞麦的CGT基因、来自百脉根的FDH基因]
pSPB7472以pBINPLUS为基本骨架,并含有以下5种表达盒。
(1)El235S启动子和来自风铃草的F3’5’H全长cDNA(序列号1)和D8终止子
(2)El235S启动子和来自蓝猪耳的MT全长cDNA(序列号3)(在5’位侧附加来自拟南芥HSPRO基因的5’-UTR(序列号13))和来自拟南芥的HSP终止子
(3)El235S启动子和来自甘草的F2H全长cDNA(序列号5)和来自紫苏的AT终止子
(4)El235S启动子和来自荞麦的CGT全长cDNA(序列号14)(在5’位侧附加来自拟南芥HSPRO基因的5’-UTR(序列号13))和来自拟南芥的HSP终止子
(5)El235S启动子和来自百脉根的FDH全长cDNA(序列号9)和来自拟南芥的HSP终止子
本质粒在植物中,对风铃草的F3’5’H基因和蓝猪耳的MT基因、甘草的F2H基因、荞麦的CGT基因、百脉根的FDH基因进行组成型表达。
将如此制作的pSPB7472导入蓝色系蔷薇品种“海洋之歌”中,获得总计33个体的转化体。接着,作为黄酮C-糖苷,实施异牡荆素(芹菜素6-C-糖苷)、牡荆素(芹菜素8-C-糖苷)、异荭草素(木犀草素6-C-糖苷)、荭草素(木犀草素8-C-糖苷)、维采宁-2(芹菜素6,8-C-二糖苷)的鉴定及定量,作为花青素,实施花翠素、矢车菊素、矮牵牛素、天竺葵素、二甲花翠素的鉴定及定量。其结果在11个体中确认黄酮C-糖苷及二甲花翠素的累积。黄酮C-糖苷的平均含量为每花瓣新鲜重量1g中3.75mg,作为主成分检测出作为黄酮6-C-糖苷的异牡荆素。此外,二甲花翠素的含有率最高为15.6%(平均8.8%)。
如以上所示,每花瓣新鲜重量1g中的黄酮C-糖苷的平均含量在实施例3所示的OS/7473系统一方高达4.19mg。即,表明与原始的来自荞麦的CGT基因相比改变密码子的CGT基因一方可更有效地生成黄酮C-糖苷。
代表性转化体的分析值示于下表4。
[表4]
宿主:海洋之歌
Del:花翠素、Cya:矢车菊素、Pet:矮牵牛素、Pel:天竺葵素、Mal:二甲花翠素
M:杨梅素、Q:槲皮素、K:山奈酚
Tri:五羟黄酮、Lut:木犀草素、Api:芹菜素、Vic2:维采宁-2、VX:牡荆素、IVX:异牡荆素、Ori:荭草素、Iori:异荭草素
Mal(%):总花青素中的二甲花翠素的比例
[实施例5:向蔷薇品种“海洋之歌”中导入来自风铃草的F3’5’H基因和来自蓝猪耳的MT基因、来自甘草的F2H基因、来自荞麦的CGT基因、来自百脉根的FDH基因]
pSPB7808以pBINPLUS为基本骨架,并含有以下5种表达盒。
(1)El235S启动子和来自风铃草的F3’5’H全长cDNA(序列号1)和D8终止子
(2)El235S启动子和来自蓝猪耳的MT全长cDNA(序列号3)和来自拟南芥的HSP终止子
(3)El235S启动子和来自甘草的F2H全长cDNA(序列号5)和来自紫苏的AT终止子
(4)El235S启动子和来自荞麦的CGT全长cDNA(序列号14)(在5’位侧附加来自拟南芥醇脱氢酶(ADH)基因的5’-UTR(序列号15))和来自拟南芥的HSP终止子
(5)El235S启动子和来自百脉根的FDH全长cDNA(序列号9)和来自拟南芥的HSP终止子
本质粒在植物中,对风铃草的F3’5’H基因和蓝猪耳的MT基因、甘草的F2H基因、荞麦的CGT基因、百脉根的FDH基因进行组成型表达。
将如此制作的pSPB7808导入蓝色系蔷薇品种“海洋之歌”中,获得总计65个体的转化体。接着,作为黄酮C-糖苷,实施异牡荆素(芹菜素6-C-糖苷)、牡荆素(芹菜素8-C-糖苷)、异荭草素(木犀草素6-C-糖苷)、荭草素(木犀草素8-C-糖苷)、维采宁-2(芹菜素6,8-C-二糖苷)的鉴定及定量,作为花青素,实施花翠素、矢车菊素、矮牵牛素、天竺葵素、二甲花翠素的鉴定及定量。其结果在32个体中确认黄酮C-糖苷及二甲花翠素的累积。黄酮C-糖苷的平均含量为每花瓣新鲜重量1g中3.00mg,作为主成分检测出作为黄酮6-C-糖苷的异牡荆素。此外,二甲花翠素的含有率最高为69.2%(平均43.9%)。
代表性转化体的分析值示于下表5。
[表5]
宿主:海洋之歌
Del:花翠素、Cya:矢车菊素、Pet:矮牵牛素、Pel:天竺葵素、Mal:二甲花翠素
M:杨梅素、Q:槲皮素、K:山奈酚
Tri:五羟黄酮、Lut:木犀草素、Api:芹菜素、Vic2:维采宁-2、VX:牡荆素、IVX:异牡荆素、Ori:荭草素、Iori:异荭草素
Mal(%):总花青素中的二甲花翠素的比例
[实施例6:向蔷薇品种“海洋之歌”中导入来自风铃草的F3’5’H基因和来自蓝猪耳的MT基因、来自甘草的F2H基因、来自荞麦的改变密码子使用的CGT基因、来自百脉根的FDH基因]
pSPB7809以pBINPLUS为基本骨架,并含有以下5种表达盒。
(1)El235S启动子和来自风铃草的F3’5’H全长cDNA(序列号1)和D8终止子
(2)El235S启动子和来自蓝猪耳的MT全长cDNA(序列号3)和来自拟南芥的HSP终止子
(3)El235S启动子和来自甘草的F2H全长cDNA(序列号5)和来自紫苏的AT终止子
(4)El235S启动子和来自荞麦的改变密码子使用的CGT全长cDNA(序列号11)(在5’位侧附加来自拟南芥醇脱氢酶(ADH)基因的5’-UTR(序列号15))和来自拟南芥的HSP终止子
(5)El235S启动子和来自百脉根的FDH全长cDNA(序列号9)和来自拟南芥的HSP终止子
本质粒在植物中,对风铃草的F3’5’H基因和蓝猪耳的MT基因、甘草的F2H基因、荞麦的改变密码子使用的CGT基因、百脉根的FDH基因进行组成型表达。
将如此制作的pSPB7809导入蓝色系蔷薇品种“海洋之歌”中,获得总计143个体的转化体。接着,作为黄酮C-糖苷,实施异牡荆素(芹菜素6-C-糖苷)、牡荆素(芹菜素8-C-糖苷)、异荭草素(木犀草素6-C-糖苷)、荭草素(木犀草素8-C-糖苷)、维采宁-2(芹菜素6,8-C-二糖苷)的鉴定及定量,作为花青素,实施花翠素、矢车菊素、矮牵牛素、天竺葵素、二甲花翠素的鉴定及定量。其结果在58个体中确认黄酮C-糖苷的累积。黄酮C-糖苷的平均含量为每花瓣新鲜重量1g中3.24mg,作为主成分检测出作为黄酮6-C-糖苷的异牡荆素。此外,二甲花翠素的含有率最高为80.3%(平均46.6%)。
如以上所示,每花瓣新鲜重量1g中的黄酮C-糖苷的平均含量在本实施例所示的OS/78093系统一方较高。从而,与实施例3及4中所得的结果相同,表明与原始的来自荞麦的CGT基因相比改变密码子的CGT基因一方可更有效地生成黄酮C-糖苷。
代表性转化体的分析值示于下表6。
[表6-1]
[表6-2]
宿主:海洋之歌
Del:花翠素、Cya:矢车菊素、Pet:矮牵牛素、Pel:天竺葵素、Mal:二甲花翠素
M:杨梅素、Q:槲皮素、K:山奈酚
Tri:五羟黄酮、Lut:木犀草素、Api:芹菜素、Vic2:维采宁-2、VX:牡荆素、IVX:异牡荆素、Ori:荭草素、Iori:异荭草素
Mal(%):总花青素中的二甲花翠素的比例
[实施例7:含有黄酮C-糖苷的蔷薇的花色的评价]
对于在实施例2及3中制作的转化体(以蔷薇品种“海洋之歌”为宿主),使用分光光度计CM-2022(美能达株式会社)在10度视野、D65光源下分别对花瓣的色彩进行测定,通过色彩管理软件SpectraMagicTM(美能达株式会社)进行解析。
在色相角度的平均值的对比中,CGT基因为来自荞麦的(实施例3)的蔷薇与来自稻子的(实施例2)的蔷薇之前的花瓣的色相角度未见差异。然而,若另外从个体方面来看,导入来自荞麦的CGT基因的蔷薇中色相角度表现为315°以下的个体多,其中还获得了表现出294.5°的至今为止变得最蓝的个体。由以上的结果,确认通过利用来自荞麦的CGT基因,可显著增加黄酮C-糖苷的生成量,尤其是单-C-糖苷的生成量,而这些与花色素苷的共存可使花瓣的色彩变蓝。
结果示于表7。
[表7]
序列表
<110> 三得利控股株式会社
<120> 来自荞麦的CGT基因及其应用
<130> P200445WO
<160> 15
<170> PatentIn version 3.5
<210> 1
<211> 1927
<212> DNA
<213> 风铃草
<400> 1
accaaatgag ctttgtaatt tgagattaat cataattgca tgctcaacta acattctgta 60
ttcatatatc catatgtatt ttgacctata gatattacat tacaccttga ggcctttata 120
tatagagagt gtatctactt cccttaatat caccttttca ttcaacaagt gaagccacca 180
tgtctataga catatccacc ctcttctatg aacttgttgc agcaatttca ctctacttag 240
ctacctactc tttcattcgt ttcctcttca aaccctctca ccaccaccac ctccctcccg 300
gcccaaccgg atggccgatc atcggagccc ttccactctt aggcaccatg ccacatgttt 360
ccttagccga catggccgtt aaatacggtc ctataatgta cctaaaactt ggttcaaagg 420
gcaccgtcgt ggcctcaaat ccaaaagccg cccgagcctt cttgaaaacc catgatgcca 480
atttttctaa ccgtccgatt gatgggggcc ctacctacct cgcgtataat gcacaagaca 540
tggtttttgc agaatatggc ccaaaatgga agcttttgcg aaagctatgt agcttgcaca 600
tgttaggccc gaaggcactc gaggattggg ctcatgtcaa agtttcagag gtcggtcata 660
tgctcaaaga aatgtacgag caatcgagta agtcagtgcc agtgccagtg gtggtgccag 720
agatgttaac ttatgccatg gctaatatga ttggacgaat catactcagc cgacgccctt 780
ttgttatcac gagcaaatta gactcgtctg cttctgcttc tgcttctgtt agtgaattcc 840
aatatatggt tatggagctc atgaggatgg cagggttgtt caatattggt gatttcatac 900
catatattgc atggatggat ttgcaaggca ttcaacgtga tatgaaggtt atacagaaaa 960
agtttgatgt cttgttgaac aaaatgatca aggaacatac agaatccgct catgatcgca 1020
aagataatcc tgattttctt gatattctta tggcggctac ccaagaaaac acggagggaa 1080
ttcagcttaa tcttgtaaat gttaaggcac ttcttttgga tttattcacg gcgggcacgg 1140
atacatcatc aagtgtgatc gaatgggcac tagccgaaat gttgaaccat cgacagatcc 1200
taaaccgggc ccacgaagaa atggaccaag tcattggcag aaacagaaga ctagaacaat 1260
ctgacatacc aaacttgcca tatttccaag ccatatgcaa agaaacattc cgaaaacacc 1320
cttccacgcc cttaaacctc ccaagaatct caacagaagc atgtgaagtg gacggatttc 1380
acataccaaa aaacactaga ctaatagtga acatatgggc aatagggagg gaccctaaag 1440
tgtgggaaaa tccattagat tttactccgg aacgtttctt gagtgaaaaa cacgcgaaaa 1500
ttgatccgcg aggtaatcat tttgagttaa tcccatttgg ggctggacga aggatatgtg 1560
caggggctag aatgggagcg gcctcggtcg agtacatatt aggtacattg gtgcactcat 1620
ttgattggaa attgcctgat ggagttgtgg aagttaatat ggaagagagc tttgggatcg 1680
cattgcaaaa aaaagtgcct ctttctgcta ttgttactcc aagattgcct ccaagttctt 1740
acactgtcta ggcaaatgct tatatatatg aataattgat tgagttgttt agttgtatga 1800
aagatttgag aaaataaatt attaggtttt gcaccattat gttgagatgg ttgttgttag 1860
tgttaaggaa gtcgattgta gtaataataa ttttattttt ttcgaaaaaa aaaaaaaaaa 1920
aaaaaaa 1927
<210> 2
<211> 523
<212> PRT
<213> 风铃草
<400> 2
Met Ser Ile Asp Ile Ser Thr Leu Phe Tyr Glu Leu Val Ala Ala Ile
1 5 10 15
Ser Leu Tyr Leu Ala Thr Tyr Ser Phe Ile Arg Phe Leu Phe Lys Pro
20 25 30
Ser His His His His Leu Pro Pro Gly Pro Thr Gly Trp Pro Ile Ile
35 40 45
Gly Ala Leu Pro Leu Leu Gly Thr Met Pro His Val Ser Leu Ala Asp
50 55 60
Met Ala Val Lys Tyr Gly Pro Ile Met Tyr Leu Lys Leu Gly Ser Lys
65 70 75 80
Gly Thr Val Val Ala Ser Asn Pro Lys Ala Ala Arg Ala Phe Leu Lys
85 90 95
Thr His Asp Ala Asn Phe Ser Asn Arg Pro Ile Asp Gly Gly Pro Thr
100 105 110
Tyr Leu Ala Tyr Asn Ala Gln Asp Met Val Phe Ala Glu Tyr Gly Pro
115 120 125
Lys Trp Lys Leu Leu Arg Lys Leu Cys Ser Leu His Met Leu Gly Pro
130 135 140
Lys Ala Leu Glu Asp Trp Ala His Val Lys Val Ser Glu Val Gly His
145 150 155 160
Met Leu Lys Glu Met Tyr Glu Gln Ser Ser Lys Ser Val Pro Val Pro
165 170 175
Val Val Val Pro Glu Met Leu Thr Tyr Ala Met Ala Asn Met Ile Gly
180 185 190
Arg Ile Ile Leu Ser Arg Arg Pro Phe Val Ile Thr Ser Lys Leu Asp
195 200 205
Ser Ser Ala Ser Ala Ser Ala Ser Val Ser Glu Phe Gln Tyr Met Val
210 215 220
Met Glu Leu Met Arg Met Ala Gly Leu Phe Asn Ile Gly Asp Phe Ile
225 230 235 240
Pro Tyr Ile Ala Trp Met Asp Leu Gln Gly Ile Gln Arg Asp Met Lys
245 250 255
Val Ile Gln Lys Lys Phe Asp Val Leu Leu Asn Lys Met Ile Lys Glu
260 265 270
His Thr Glu Ser Ala His Asp Arg Lys Asp Asn Pro Asp Phe Leu Asp
275 280 285
Ile Leu Met Ala Ala Thr Gln Glu Asn Thr Glu Gly Ile Gln Leu Asn
290 295 300
Leu Val Asn Val Lys Ala Leu Leu Leu Asp Leu Phe Thr Ala Gly Thr
305 310 315 320
Asp Thr Ser Ser Ser Val Ile Glu Trp Ala Leu Ala Glu Met Leu Asn
325 330 335
His Arg Gln Ile Leu Asn Arg Ala His Glu Glu Met Asp Gln Val Ile
340 345 350
Gly Arg Asn Arg Arg Leu Glu Gln Ser Asp Ile Pro Asn Leu Pro Tyr
355 360 365
Phe Gln Ala Ile Cys Lys Glu Thr Phe Arg Lys His Pro Ser Thr Pro
370 375 380
Leu Asn Leu Pro Arg Ile Ser Thr Glu Ala Cys Glu Val Asp Gly Phe
385 390 395 400
His Ile Pro Lys Asn Thr Arg Leu Ile Val Asn Ile Trp Ala Ile Gly
405 410 415
Arg Asp Pro Lys Val Trp Glu Asn Pro Leu Asp Phe Thr Pro Glu Arg
420 425 430
Phe Leu Ser Glu Lys His Ala Lys Ile Asp Pro Arg Gly Asn His Phe
435 440 445
Glu Leu Ile Pro Phe Gly Ala Gly Arg Arg Ile Cys Ala Gly Ala Arg
450 455 460
Met Gly Ala Ala Ser Val Glu Tyr Ile Leu Gly Thr Leu Val His Ser
465 470 475 480
Phe Asp Trp Lys Leu Pro Asp Gly Val Val Glu Val Asn Met Glu Glu
485 490 495
Ser Phe Gly Ile Ala Leu Gln Lys Lys Val Pro Leu Ser Ala Ile Val
500 505 510
Thr Pro Arg Leu Pro Pro Ser Ser Tyr Thr Val
515 520
<210> 3
<211> 720
<212> DNA
<213> 蓝猪耳杂交品种
<400> 3
atgaaagata agttctatgg caccattttg cagagcgaag ccctcgcaaa gtatctgtta 60
gagacaagtg cctatccacg agaacatccg cagctcaaag aactaaggag cgcaactgtg 120
gacaagtatc aatattggag cttgatgaat gttccagctg atgaggggca gttcatttca 180
atgttactga aaattatgaa cgcaaaaaag acaattgaag ttggagtttt cacaggctac 240
tcactcctat caactgctct ggctctacct gatgatggca aaatcgttgc cattgatcct 300
gatagagaag cttatgagac tggtttgcca tttatcaaga aagcaaacgt ggctcataaa 360
atccaataca tacaatctga tgccatgaaa gtcatgaatg acctcattgc tgccaaggga 420
gaagaagaag aggggagctt tgactttggg ttcgtggatg cagacaaaga aaactacata 480
aactaccacg agaaactgtt gaagctggtt aaggttggag ggatcatagg atacgacaac 540
actctgtggt ctggaacagt tgctgcatct gaagacgatg agaataatat gcaagactac 600
ttaagaggtt gcagagggca tatcctcaaa ctaaactcct ttctcgcaaa cgatgatcgg 660
attgaattgg ctcacctctc tattggagat ggactcacct tgtgcaaacg tctcaaataa 720
<210> 4
<211> 239
<212> PRT
<213> 蓝猪耳杂交品种
<400> 4
Met Lys Asp Lys Phe Tyr Gly Thr Ile Leu Gln Ser Glu Ala Leu Ala
1 5 10 15
Lys Tyr Leu Leu Glu Thr Ser Ala Tyr Pro Arg Glu His Pro Gln Leu
20 25 30
Lys Glu Leu Arg Ser Ala Thr Val Asp Lys Tyr Gln Tyr Trp Ser Leu
35 40 45
Met Asn Val Pro Ala Asp Glu Gly Gln Phe Ile Ser Met Leu Leu Lys
50 55 60
Ile Met Asn Ala Lys Lys Thr Ile Glu Val Gly Val Phe Thr Gly Tyr
65 70 75 80
Ser Leu Leu Ser Thr Ala Leu Ala Leu Pro Asp Asp Gly Lys Ile Val
85 90 95
Ala Ile Asp Pro Asp Arg Glu Ala Tyr Glu Thr Gly Leu Pro Phe Ile
100 105 110
Lys Lys Ala Asn Val Ala His Lys Ile Gln Tyr Ile Gln Ser Asp Ala
115 120 125
Met Lys Val Met Asn Asp Leu Ile Ala Ala Lys Gly Glu Glu Glu Glu
130 135 140
Gly Ser Phe Asp Phe Gly Phe Val Asp Ala Asp Lys Glu Asn Tyr Ile
145 150 155 160
Asn Tyr His Glu Lys Leu Leu Lys Leu Val Lys Val Gly Gly Ile Ile
165 170 175
Gly Tyr Asp Asn Thr Leu Trp Ser Gly Thr Val Ala Ala Ser Glu Asp
180 185 190
Asp Glu Asn Asn Met Gln Asp Tyr Leu Arg Gly Cys Arg Gly His Ile
195 200 205
Leu Lys Leu Asn Ser Phe Leu Ala Asn Asp Asp Arg Ile Glu Leu Ala
210 215 220
His Leu Ser Ile Gly Asp Gly Leu Thr Leu Cys Lys Arg Leu Lys
225 230 235
<210> 5
<211> 1920
<212> DNA
<213> 甘草
<400> 5
ccccaatttc ctctatcata agccattccg ttgattgagc ttcctttccg tgaaaaaaat 60
aactaagcga tatggaacct caactcgtag cagtgtctgt gttggtttca gcacttatct 120
gctacttctt tttccggcca tatttccacc gttacggaaa aaaccttcca ccatctcctt 180
ttttccggct tccaataatt ggccacatgc acatgttagg tccccttctt caccaatcct 240
tccacaacct ctcccaccgt tacggtcctc tgttttcact taactttggc tctgttctct 300
gtgtcgttgc ttcaacccct cactttgcca aacaactcct tcaaaccaac gaactcgcct 360
ttaactgtcg cattgaatca accgccgtta aaaaactcac ttacgagtct tccttggcct 420
tcgcacctta cggtgattac tggaggttca ttaagaagct gagcatgaac gagcttttgg 480
gctctcgtag cataaacaac ttccaacacc tacgagcaca agagacccat caattgttaa 540
ggcttttgtc caacagggca agagcgtttg aggccgtgaa tatcaccgag gagcttctta 600
agttgaccaa caacgttatt tctataatga tggttgggga ggcagaggag gcaagggatg 660
tggtgcgtga tgtgacggag atatttggag agtttaatgt ttcggatttt atttggttgt 720
ttaagaagat ggacttgcag gggtttggga agaggattga ggatttgttt cagaggtttg 780
atacgttggt ggaaaggatt attagcaagc gggagcagac gaggaaagac agaaggagga 840
atgggaagaa gggtgagcag gggagtggtg atgggatcag agactttctt gatatcttgc 900
ttgactgtac tgaggatgag aattccgaga ttaaaatcca aagggttcac attaaggcct 960
tgattatgga tttcttcact gcagggacgg ataccacagc gatttcaaca gagtgggcat 1020
tagtggagct cgtcaagaaa ccctccgtgc tacaaaaagt tcgtgaagag atagacaatg 1080
tcgtaggaaa agacagactt gttgaggaat ctgattgtcc taatctccca tatctccaag 1140
ccattcttaa agaaacattc cgtttgcacc caccggttcc tatggttaca agaagatgcg 1200
tggcagagtg cacggtagag aattacgtca tcccagaaga ctcacttctc tttgtgaatg 1260
tttggtccat cggaagaaac ccaaagtttt gggacaaccc attggagttt cgccccgaac 1320
gattcttaaa actagaagga gattccagtg gagttgttga tgtgagggga agccattttc 1380
agcttctgcc atttgggtct ggaaggagga tgtgccctgg tgtgtccttg gctatgcaag 1440
aggtgccagc actacttggt gctataatcc agtgctttga tttccacgtt gtgggtccca 1500
aaggtgagat tttgaagggt gatgacatag tcattaatgt ggatgaaagg ccaggattga 1560
cggctccaag ggcacataac cttgtgtgtg ttcccgttga tagaacaagt ggcggtggac 1620
ccctcaaaat cattgaatgt tgattattcg tctcttgaat ttggatctgt gtgtggcttt 1680
gaataacatg tatggtgtat gtatgtatgt gttcttttct ttctttctaa ttctgatcag 1740
tagcgtacac taggcactag ccttcgttag tggcaataac ttcggcaaat taacgaacat 1800
atgctgttca gagatatttt ttgccatgta tcgtcattct attctaggtt gtttttccgt 1860
tttccttatt acattctatg ataaatataa taaattgagt aatattatag tctcttaatt 1920
<210> 6
<211> 523
<212> PRT
<213> 甘草
<400> 6
Met Glu Pro Gln Leu Val Ala Val Ser Val Leu Val Ser Ala Leu Ile
1 5 10 15
Cys Tyr Phe Phe Phe Arg Pro Tyr Phe His Arg Tyr Gly Lys Asn Leu
20 25 30
Pro Pro Ser Pro Phe Phe Arg Leu Pro Ile Ile Gly His Met His Met
35 40 45
Leu Gly Pro Leu Leu His Gln Ser Phe His Asn Leu Ser His Arg Tyr
50 55 60
Gly Pro Leu Phe Ser Leu Asn Phe Gly Ser Val Leu Cys Val Val Ala
65 70 75 80
Ser Thr Pro His Phe Ala Lys Gln Leu Leu Gln Thr Asn Glu Leu Ala
85 90 95
Phe Asn Cys Arg Ile Glu Ser Thr Ala Val Lys Lys Leu Thr Tyr Glu
100 105 110
Ser Ser Leu Ala Phe Ala Pro Tyr Gly Asp Tyr Trp Arg Phe Ile Lys
115 120 125
Lys Leu Ser Met Asn Glu Leu Leu Gly Ser Arg Ser Ile Asn Asn Phe
130 135 140
Gln His Leu Arg Ala Gln Glu Thr His Gln Leu Leu Arg Leu Leu Ser
145 150 155 160
Asn Arg Ala Arg Ala Phe Glu Ala Val Asn Ile Thr Glu Glu Leu Leu
165 170 175
Lys Leu Thr Asn Asn Val Ile Ser Ile Met Met Val Gly Glu Ala Glu
180 185 190
Glu Ala Arg Asp Val Val Arg Asp Val Thr Glu Ile Phe Gly Glu Phe
195 200 205
Asn Val Ser Asp Phe Ile Trp Leu Phe Lys Lys Met Asp Leu Gln Gly
210 215 220
Phe Gly Lys Arg Ile Glu Asp Leu Phe Gln Arg Phe Asp Thr Leu Val
225 230 235 240
Glu Arg Ile Ile Ser Lys Arg Glu Gln Thr Arg Lys Asp Arg Arg Arg
245 250 255
Asn Gly Lys Lys Gly Glu Gln Gly Ser Gly Asp Gly Ile Arg Asp Phe
260 265 270
Leu Asp Ile Leu Leu Asp Cys Thr Glu Asp Glu Asn Ser Glu Ile Lys
275 280 285
Ile Gln Arg Val His Ile Lys Ala Leu Ile Met Asp Phe Phe Thr Ala
290 295 300
Gly Thr Asp Thr Thr Ala Ile Ser Thr Glu Trp Ala Leu Val Glu Leu
305 310 315 320
Val Lys Lys Pro Ser Val Leu Gln Lys Val Arg Glu Glu Ile Asp Asn
325 330 335
Val Val Gly Lys Asp Arg Leu Val Glu Glu Ser Asp Cys Pro Asn Leu
340 345 350
Pro Tyr Leu Gln Ala Ile Leu Lys Glu Thr Phe Arg Leu His Pro Pro
355 360 365
Val Pro Met Val Thr Arg Arg Cys Val Ala Glu Cys Thr Val Glu Asn
370 375 380
Tyr Val Ile Pro Glu Asp Ser Leu Leu Phe Val Asn Val Trp Ser Ile
385 390 395 400
Gly Arg Asn Pro Lys Phe Trp Asp Asn Pro Leu Glu Phe Arg Pro Glu
405 410 415
Arg Phe Leu Lys Leu Glu Gly Asp Ser Ser Gly Val Val Asp Val Arg
420 425 430
Gly Ser His Phe Gln Leu Leu Pro Phe Gly Ser Gly Arg Arg Met Cys
435 440 445
Pro Gly Val Ser Leu Ala Met Gln Glu Val Pro Ala Leu Leu Gly Ala
450 455 460
Ile Ile Gln Cys Phe Asp Phe His Val Val Gly Pro Lys Gly Glu Ile
465 470 475 480
Leu Lys Gly Asp Asp Ile Val Ile Asn Val Asp Glu Arg Pro Gly Leu
485 490 495
Thr Ala Pro Arg Ala His Asn Leu Val Cys Val Pro Val Asp Arg Thr
500 505 510
Ser Gly Gly Gly Pro Leu Lys Ile Ile Glu Cys
515 520
<210> 7
<211> 1416
<212> DNA
<213> 粳稻组
<400> 7
atgccgagct ctggcgacgc ggcgggcagg cggccgcatg tggtgctcat cccgagcgcc 60
ggcatgggcc acctcgtccc cttcggccgc ctcgccgtgg cgctctcctc cggccacggc 120
tgcgacgtct ccctcgtcac ggtgctcccc acggtgtcca ccgcggagtc gaagcacctc 180
gacgcgctgt tcgacgcgtt cccggcggtg cggcggctcg acttcgagct cgcgccgttc 240
gacgcgtcgg agttccccag cgccgacccg ttcttcctcc ggttcgaggc catgcggcgg 300
tcggcgccgc tgctcggccc gctcctcacc ggcgccggcg catcggcgct cgccacggac 360
atcgcgctga catccgtcgt catacccgtg gcgaaggagc agggcctccc gtgccacatc 420
ctcttcaccg cctccgccgc gatgctctcc ctctgcgcct acttccccac atacctcgac 480
gccaacgctg gcgacggcgg cggcgtcggc gacgtcgaca tccccggcgt gtaccgcatc 540
cccaaggcct ccatcccgca ggcgctgcac gatcccaacc acctcttcac ccgccagttc 600
gtcgccaacg gccggagcct cacgagcgcc gccggcatcc tcgtcaacac gttcgacgcc 660
ttggagccgg aggccgtcgc ggccctgcag cagggcaagg tcgcctccgg cttcccgccg 720
gtgttcgccg tggggccact tctcccggcg agcaaccagg caaaagatcc gcaggcaaac 780
tacatggagt ggctcgacgc gcagccggcg cggtcggtgg tgtacgtgag cttcggcagc 840
cgcaaggcca tctcagggga gcagctcagg gagctcgccg ccgggctgga gaccagcggc 900
cacaggttcc tgtgggtggt gaagagcacc gtcgtggaca gggacgacgc cgccgagctc 960
ggcgagctgc tcggcgaggg gttcttgaag cgggtggaga agcgaggcct cgtcaccaag 1020
gcatgggtgg atcaggaaga ggtcctgaag cacgagtccg tggcgctgtt cgtgagccac 1080
tgcggctgga actcggtgac ggaggcggcg gcgagcggcg tcccggtgct ggcgctgccg 1140
aggttcggcg accagcgggt gaactccggc gtggtggcgc gcgccgggct cggcgtgtgg 1200
gcggacacct ggagctggga gggggaagcc ggggtgatcg gcgcagagga gatatcggag 1260
aaggtgaagg cggcgatggc ggacgaggcg ttgcgtagga aggcggcgag cctcgccaag 1320
gccgccgcga aggccgtcgc cggcggtgga tcgagccacc gttgtctggt cgagttcgcg 1380
cggctgtgcc aagggggaac atgtcgcact aattga 1416
<210> 8
<211> 471
<212> PRT
<213> 粳稻组
<400> 8
Met Pro Ser Ser Gly Asp Ala Ala Gly Arg Arg Pro His Val Val Leu
1 5 10 15
Ile Pro Ser Ala Gly Met Gly His Leu Val Pro Phe Gly Arg Leu Ala
20 25 30
Val Ala Leu Ser Ser Gly His Gly Cys Asp Val Ser Leu Val Thr Val
35 40 45
Leu Pro Thr Val Ser Thr Ala Glu Ser Lys His Leu Asp Ala Leu Phe
50 55 60
Asp Ala Phe Pro Ala Val Arg Arg Leu Asp Phe Glu Leu Ala Pro Phe
65 70 75 80
Asp Ala Ser Glu Phe Pro Ser Ala Asp Pro Phe Phe Leu Arg Phe Glu
85 90 95
Ala Met Arg Arg Ser Ala Pro Leu Leu Gly Pro Leu Leu Thr Gly Ala
100 105 110
Gly Ala Ser Ala Leu Ala Thr Asp Ile Ala Leu Thr Ser Val Val Ile
115 120 125
Pro Val Ala Lys Glu Gln Gly Leu Pro Cys His Ile Leu Phe Thr Ala
130 135 140
Ser Ala Ala Met Leu Ser Leu Cys Ala Tyr Phe Pro Thr Tyr Leu Asp
145 150 155 160
Ala Asn Ala Gly Asp Gly Gly Gly Val Gly Asp Val Asp Ile Pro Gly
165 170 175
Val Tyr Arg Ile Pro Lys Ala Ser Ile Pro Gln Ala Leu His Asp Pro
180 185 190
Asn His Leu Phe Thr Arg Gln Phe Val Ala Asn Gly Arg Ser Leu Thr
195 200 205
Ser Ala Ala Gly Ile Leu Val Asn Thr Phe Asp Ala Leu Glu Pro Glu
210 215 220
Ala Val Ala Ala Leu Gln Gln Gly Lys Val Ala Ser Gly Phe Pro Pro
225 230 235 240
Val Phe Ala Val Gly Pro Leu Leu Pro Ala Ser Asn Gln Ala Lys Asp
245 250 255
Pro Gln Ala Asn Tyr Met Glu Trp Leu Asp Ala Gln Pro Ala Arg Ser
260 265 270
Val Val Tyr Val Ser Phe Gly Ser Arg Lys Ala Ile Ser Gly Glu Gln
275 280 285
Leu Arg Glu Leu Ala Ala Gly Leu Glu Thr Ser Gly His Arg Phe Leu
290 295 300
Trp Val Val Lys Ser Thr Val Val Asp Arg Asp Asp Ala Ala Glu Leu
305 310 315 320
Gly Glu Leu Leu Gly Glu Gly Phe Leu Lys Arg Val Glu Lys Arg Gly
325 330 335
Leu Val Thr Lys Ala Trp Val Asp Gln Glu Glu Val Leu Lys His Glu
340 345 350
Ser Val Ala Leu Phe Val Ser His Cys Gly Trp Asn Ser Val Thr Glu
355 360 365
Ala Ala Ala Ser Gly Val Pro Val Leu Ala Leu Pro Arg Phe Gly Asp
370 375 380
Gln Arg Val Asn Ser Gly Val Val Ala Arg Ala Gly Leu Gly Val Trp
385 390 395 400
Ala Asp Thr Trp Ser Trp Glu Gly Glu Ala Gly Val Ile Gly Ala Glu
405 410 415
Glu Ile Ser Glu Lys Val Lys Ala Ala Met Ala Asp Glu Ala Leu Arg
420 425 430
Arg Lys Ala Ala Ser Leu Ala Lys Ala Ala Ala Lys Ala Val Ala Gly
435 440 445
Gly Gly Ser Ser His Arg Cys Leu Val Glu Phe Ala Arg Leu Cys Gln
450 455 460
Gly Gly Thr Cys Arg Thr Asn
465 470
<210> 9
<211> 990
<212> DNA
<213> 百脉根
<400> 9
atggcttctg caacaaccac cccaaccaaa gagatagaca gagacctccc tcctcttctc 60
cgagtctaca aagatggaac cgtggaacgt ctcctaggct ctcctgtcgt tccagcaatc 120
cctcatgacc cagaaacaga ggtctcatca aaagacatag tcatctcaca aaccccctta 180
atctctgctc gtatccacct cccaaaacag agcaaccccc aaaacccaaa ggttccaata 240
ttgatctact accatggtgg tgcgttttgc cttgaatcag ctttctcctt cctccaccaa 300
cgctacctca acatcgtggc ttcacgatca aacgttgtgg tggtttccgt cgagtacagg 360
ctcgcgccag agcatcctct gcctgcagca tatgaagatg gttgggaagc tctgaaatgg 420
gttacctctc attccaccga caacaaaccc atcaactctg agccatggtt gatcgaacat 480
ggtgatttca gcagattcta catcggaggt gacacttcag gtgccaacat cgcataccat 540
gtgggtctcc gtgtcggtgg tggagttgag aaattgccag gggatgtgaa aattgcaggg 600
gcgttacttg cttttcccct gttttggagt tcataccctg ttttggaaga acctgttgag 660
gggtttgaac agagtttgag caggaaggtt tggaactttg tgtacccaga tgcacctggt 720
gggatcgaca accctctgat caatcctttg gctgatgggg ctccaagctt gacaacgttt 780
ggaagcaaca agatgttgat ctttgttgca gggaatgatg aactgagaga cagaggaatc 840
tggttctatg aggctgtgaa gaagagtgag tgggaaggtg atgtggaact cattcgagtg 900
gatggagagg agcattgctt ccagatttac catcctgaat ctgagaattc taaagacatg 960
atgaagcgca tagcttcttt ccttgtttga 990
<210> 10
<211> 329
<212> PRT
<213> 百脉根
<400> 10
Met Ala Ser Ala Thr Thr Thr Pro Thr Lys Glu Ile Asp Arg Asp Leu
1 5 10 15
Pro Pro Leu Leu Arg Val Tyr Lys Asp Gly Thr Val Glu Arg Leu Leu
20 25 30
Gly Ser Pro Val Val Pro Ala Ile Pro His Asp Pro Glu Thr Glu Val
35 40 45
Ser Ser Lys Asp Ile Val Ile Ser Gln Thr Pro Leu Ile Ser Ala Arg
50 55 60
Ile His Leu Pro Lys Gln Ser Asn Pro Gln Asn Pro Lys Val Pro Ile
65 70 75 80
Leu Ile Tyr Tyr His Gly Gly Ala Phe Cys Leu Glu Ser Ala Phe Ser
85 90 95
Phe Leu His Gln Arg Tyr Leu Asn Ile Val Ala Ser Arg Ser Asn Val
100 105 110
Val Val Val Ser Val Glu Tyr Arg Leu Ala Pro Glu His Pro Leu Pro
115 120 125
Ala Ala Tyr Glu Asp Gly Trp Glu Ala Leu Lys Trp Val Thr Ser His
130 135 140
Ser Thr Asp Asn Lys Pro Ile Asn Ser Glu Pro Trp Leu Ile Glu His
145 150 155 160
Gly Asp Phe Ser Arg Phe Tyr Ile Gly Gly Asp Thr Ser Gly Ala Asn
165 170 175
Ile Ala Tyr His Val Gly Leu Arg Val Gly Gly Gly Val Glu Lys Leu
180 185 190
Pro Gly Asp Val Lys Ile Ala Gly Ala Leu Leu Ala Phe Pro Leu Phe
195 200 205
Trp Ser Ser Tyr Pro Val Leu Glu Glu Pro Val Glu Gly Phe Glu Gln
210 215 220
Ser Leu Ser Arg Lys Val Trp Asn Phe Val Tyr Pro Asp Ala Pro Gly
225 230 235 240
Gly Ile Asp Asn Pro Leu Ile Asn Pro Leu Ala Asp Gly Ala Pro Ser
245 250 255
Leu Thr Thr Phe Gly Ser Asn Lys Met Leu Ile Phe Val Ala Gly Asn
260 265 270
Asp Glu Leu Arg Asp Arg Gly Ile Trp Phe Tyr Glu Ala Val Lys Lys
275 280 285
Ser Glu Trp Glu Gly Asp Val Glu Leu Ile Arg Val Asp Gly Glu Glu
290 295 300
His Cys Phe Gln Ile Tyr His Pro Glu Ser Glu Asn Ser Lys Asp Met
305 310 315 320
Met Lys Arg Ile Ala Ser Phe Leu Val
325
<210> 11
<211> 1374
<212> DNA
<213> 荞麦
<400> 11
atgatgggcg acttaaccac atcttttcca gcaaccacac tcaccacgaa tgatcaacct 60
cacgtggtgg tttgctcagg agctggtatg gggcatttga caccgttcct taacttggcc 120
tctgctcttt catctgcgcc gtataactgc aaggttactc tgttaatcgt gattccgctt 180
atcacagatg ccgaaagtca ccacattagc agtttcttct cttcacatcc cactatccac 240
aggctagact tccacgtgaa tttaccggct ccaaagccca atgtggatcc gttctttctc 300
cgttacaaga gcatctcaga ctctgcacat agacttcctg tccacttatc ggccttgagt 360
cccccgatat ctgccgtttt ctctgacttc ctcttcactc agggtctgaa cactactttg 420
cctcatcttc ccaactacac gtttactacg acttctgctc ggtttttcac cttgatgtcc 480
tatgttcctc acttggccaa atcgagctca tcaagtcctg tagagattcc tggccttgag 540
ccgtttccaa ctgacaacat ccctccccct tttttcaacc cagagcacat ctttaccagt 600
ttcaccatat ccaatgccaa gtacttctcg ttgagcaaag ggattttggt gaataccttc 660
gactcctttg agcccgagac actctctgca ttgaactcgg gagacacatt gtcagatctc 720
cctccggtca tccctattgg cccactgaac gaactcgagc ataacaaaca ggaggaactc 780
ctgccatggc tcgaccaaca acctgagaaa tccgtccttt acgtctcttt tgggaatcga 840
acagccatga gtagcgatca gattctcgag ctaggaatgg gattagagcg tagtgattgt 900
cgctttattt gggtcgttaa gacgtccaag atagacaagg acgataagag tgagctacgt 960
aaactctttg gtgaagagct ctaccttaaa ctgtccgaga agggaaagct tgtgaaatgg 1020
gtgaaccaaa ccgagattct cgggcatact gctgtaggtg gtttcctgtc tcattgtggg 1080
tggaattcgg tcatggaagc agccagaaga ggtgttccga tattggcttg gccacaacac 1140
ggtgatcagc gggaaaatgc atgggttgtt gaaaaggctg gcttgggagt gtgggaaagg 1200
gagtgggcaa gcgggattca ggcagccatt gttgagaagg tcaagatgat catgggcaat 1260
aacgaccttc gcaaaagcgc tatgaaggtg ggtgaagagg caaagcgagc atgtgatgta 1320
ggtgggagtt ccgctaccgc cttgatgaat attatcgggt cattgaagag gtaa 1374
<210> 12
<211> 457
<212> PRT
<213> 荞麦
<400> 12
Met Met Gly Asp Leu Thr Thr Ser Phe Pro Ala Thr Thr Leu Thr Thr
1 5 10 15
Asn Asp Gln Pro His Val Val Val Cys Ser Gly Ala Gly Met Gly His
20 25 30
Leu Thr Pro Phe Leu Asn Leu Ala Ser Ala Leu Ser Ser Ala Pro Tyr
35 40 45
Asn Cys Lys Val Thr Leu Leu Ile Val Ile Pro Leu Ile Thr Asp Ala
50 55 60
Glu Ser His His Ile Ser Ser Phe Phe Ser Ser His Pro Thr Ile His
65 70 75 80
Arg Leu Asp Phe His Val Asn Leu Pro Ala Pro Lys Pro Asn Val Asp
85 90 95
Pro Phe Phe Leu Arg Tyr Lys Ser Ile Ser Asp Ser Ala His Arg Leu
100 105 110
Pro Val His Leu Ser Ala Leu Ser Pro Pro Ile Ser Ala Val Phe Ser
115 120 125
Asp Phe Leu Phe Thr Gln Gly Leu Asn Thr Thr Leu Pro His Leu Pro
130 135 140
Asn Tyr Thr Phe Thr Thr Thr Ser Ala Arg Phe Phe Thr Leu Met Ser
145 150 155 160
Tyr Val Pro His Leu Ala Lys Ser Ser Ser Ser Ser Pro Val Glu Ile
165 170 175
Pro Gly Leu Glu Pro Phe Pro Thr Asp Asn Ile Pro Pro Pro Phe Phe
180 185 190
Asn Pro Glu His Ile Phe Thr Ser Phe Thr Ile Ser Asn Ala Lys Tyr
195 200 205
Phe Ser Leu Ser Lys Gly Ile Leu Val Asn Thr Phe Asp Ser Phe Glu
210 215 220
Pro Glu Thr Leu Ser Ala Leu Asn Ser Gly Asp Thr Leu Ser Asp Leu
225 230 235 240
Pro Pro Val Ile Pro Ile Gly Pro Leu Asn Glu Leu Glu His Asn Lys
245 250 255
Gln Glu Glu Leu Leu Pro Trp Leu Asp Gln Gln Pro Glu Lys Ser Val
260 265 270
Leu Tyr Val Ser Phe Gly Asn Arg Thr Ala Met Ser Ser Asp Gln Ile
275 280 285
Leu Glu Leu Gly Met Gly Leu Glu Arg Ser Asp Cys Arg Phe Ile Trp
290 295 300
Val Val Lys Thr Ser Lys Ile Asp Lys Asp Asp Lys Ser Glu Leu Arg
305 310 315 320
Lys Leu Phe Gly Glu Glu Leu Tyr Leu Lys Leu Ser Glu Lys Gly Lys
325 330 335
Leu Val Lys Trp Val Asn Gln Thr Glu Ile Leu Gly His Thr Ala Val
340 345 350
Gly Gly Phe Leu Ser His Cys Gly Trp Asn Ser Val Met Glu Ala Ala
355 360 365
Arg Arg Gly Val Pro Ile Leu Ala Trp Pro Gln His Gly Asp Gln Arg
370 375 380
Glu Asn Ala Trp Val Val Glu Lys Ala Gly Leu Gly Val Trp Glu Arg
385 390 395 400
Glu Trp Ala Ser Gly Ile Gln Ala Ala Ile Val Glu Lys Val Lys Met
405 410 415
Ile Met Gly Asn Asn Asp Leu Arg Lys Ser Ala Met Lys Val Gly Glu
420 425 430
Glu Ala Lys Arg Ala Cys Asp Val Gly Gly Ser Ser Ala Thr Ala Leu
435 440 445
Met Asn Ile Ile Gly Ser Leu Lys Arg
450 455
<210> 13
<211> 71
<212> DNA
<213> 拟南芥
<400> 13
acaaattcaa aatttaacac acaaacacaa acacacacac caaaaaaaac acagacctta 60
aaaaaataaa a 71
<210> 14
<211> 1374
<212> DNA
<213> 荞麦
<400> 14
atgatgggag atttaacaac ttcttttccg gcaaccacat taaccaccaa tgaccaaccc 60
catgttgtcg tttgttcggg tgcggggatg ggccacttaa ccccattcct caacctagct 120
tccgcccttt cctccgcacc ctacaactgt aaagtcaccc tactcattgt catccctctt 180
atcaccgatg ctgaatccca ccatatctcg tccttctttt cctctcaccc caccatccac 240
cgcctcgact tccacgtcaa cctccccgcc cccaaaccta acgtcgaccc tttcttctta 300
cgctacaaaa gcatctccga ctcggcccac cgcctccccg tccatctctc agccctctcg 360
ccacccatct ccgccgtgtt ttccgacttt ctattcacac aaggactcaa cactactctc 420
cctcacctcc ctaactacac cttcaccacc acctccgcaa ggttcttcac tctcatgtct 480
tacgttcctc acttggctaa atcgtcatcg tcctcgcccg tcgagattcc cggccttgaa 540
ccttttccga ccgacaacat ccctcctcct ttcttcaatc ccgaacacat cttcacttcc 600
ttcacaatct ccaacgctaa gtatttttct ctttccaaag ggattctcgt caacacattc 660
gactccttcg aaccggaaac attatcggcg ctcaattccg gcgatactct ttccgatctc 720
cctccggtaa tccctatagg gcctcttaat gaacttgaac ataataaaca agaggagtta 780
ctcccttggt tggatcaaca accggagaaa tccgtactgt acgtatcatt cgggaatagg 840
acggcgatgt cgagcgatca gatacttgag ctggggatgg gactggagag gagtgattgc 900
aggttcattt gggtggtgaa aaccagcaag attgacaagg atgataaatc ggagctacgg 960
aagctattcg gcgaggagtt gtacctgaag ctaagtgaga aagggaagtt agtgaaatgg 1020
gtgaatcaaa cggagatttt agggcatacg gcggtcggag ggtttttgag ccattgtggg 1080
tggaactccg tgatggaagc tgctcggcgc ggagttccga ttctagcatg gccgcagcac 1140
ggggatcaga gagagaatgc ttgggtggtg gagaaggcgg ggcttggagt ttgggagaga 1200
gagtgggcgt cggggattca ggcggcgatt gtggagaagg tgaagatgat tatgggaaat 1260
aatgatctga gaaagagtgc aatgaaggtt ggggaggaag cgaagagggc atgtgatgtt 1320
ggtggaagct ctgcaactgc attgatgaac atcatcggaa gtcttaaacg ttga 1374
<210> 15
<211> 58
<212> DNA
<213> 拟南芥
<400> 15
tacatcacaa tcacacaaaa ctaacaaaag atcaaaagca agttcttcac tgttgata 58
Claims (19)
1.一种来自荞麦的CGT基因或其同源基因,其为来自荞麦的C-糖苷化酶,即CGT基因或其同源基因,其特征在于,所述来自荞麦的CGT基因或其同源基因选自:
(1-a)由序列号11的碱基序列构成的多聚核苷酸;
(1-b)在严谨条件下与由与序列号11的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(1-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸。
2.根据权利要求1所述的来自荞麦的CGT基因或其同源基因,其特征在于,附加有来自拟南芥ADH基因的非翻译区域(5’-UTR),即序列号15,或来自拟南芥HSPRO基因的非翻译区域(5’-UTR),即序列号13。
3.一种载体,其为含有来自荞麦的CGT基因或其同源基因的载体,其特征在于,所述来自荞麦的CGT基因或其同源基因选自:
(1-a)由序列号11的碱基序列构成的多聚核苷酸;
(1-b)在严谨条件下与由与序列号11的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(1-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(1-c)编码由序列号12的氨基酸序列构成的蛋白质的多聚核苷酸;
(1-d)编码由在序列号12的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(1-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(1-e)编码具有相对于序列号12的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(1-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸。
4.根据权利要求3所述的载体,其特征在于,在所述来自荞麦的CGT基因或其同源基因上附加有来自拟南芥ADH基因的非翻译区域(5’-UTR),即序列号15,或来自拟南芥HSPRO基因的非翻译区域(5’-UTR),即序列号13。
5.根据权利要求3或4所述的载体,其特征在于,进一步含有黄烷酮2-羟化酶,即F2H基因或其同源基因,以及脱水酶,即FDH基因或其同源基因。
6.根据权利要求5所述的载体,其特征在于,所述F2H基因或其同源基因选自:
(2-a)由序列号5的碱基序列构成的多聚核苷酸;
(2-b)在严谨条件下与由与序列号5的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(2-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(2-c)编码由序列号6的氨基酸序列构成的蛋白质的多聚核苷酸;
(2-d)编码由在序列号6的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(2-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(2-e)编码具有相对于序列号6的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(2-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸,
所述FDH基因或其同源基因选自:
(3-a)由序列号9的碱基序列构成的多聚核苷酸;
(3-b)在严谨条件下与由与序列号9的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(3-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(3-c)编码由序列号10的氨基酸序列构成的蛋白质的多聚核苷酸;
(3-d)编码由在序列号10的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(3-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(3-e)编码具有相对于序列号10的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(3-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸。
7.根据权利要求3~6中任一项所述的载体,其特征在于,进一步含有类黄酮F3’5’羟化酶,即F3’5’H基因或其同源基因,以及甲基转移酶,即MT基因或其同源基因。
8.根据权利要求7所述的载体,其特征在于,所述F3’5’H基因或其同源基因选自:
(4-a)由序列号1的碱基序列构成的多聚核苷酸;
(4-b)在严谨条件下与由与序列号1的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(4-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(4-c)编码由序列号2的氨基酸序列构成的蛋白质的多聚核苷酸;
(4-d)编码由在序列号2的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(4-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(4-e)编码具有相对于序列号2的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(4-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸,且,
所述MT基因或其同源基因选自:
(5-a)由序列号3的碱基序列构成的多聚核苷酸;
(5-b)在严谨条件下与由与序列号3的碱基序列互补的碱基序列构成的多聚核苷酸进行杂交,且与(5-a)中记载的多聚核苷酸具有同样活性的多聚核苷酸;
(5-c)编码由序列号4的氨基酸序列构成的蛋白质的多聚核苷酸;
(5-d)编码由在序列号4的氨基酸序列中,1个或多个氨基酸缺失、取代、插入及/或附加而得的氨基酸序列构成,且与(5-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸;
(5-e)编码具有相对于序列号4的氨基酸序列具有90%以上的一致性的氨基酸序列,且与(5-c)中记载的多聚核苷酸所编码的蛋白质具有同样活性的蛋白质的多聚核苷酸。
9.一种转化植物或其自繁或者杂交后代,其特征在于,含有权利要求1或2所述的来自荞麦的CGT基因或其同源基因,或者权利要求3~8中任一项所述的载体。
10.根据权利要求9所述的转化植物或其自繁或者杂交后代,其特征在于,所述植物选自蔷薇、菊花、康乃馨或百合。
11.根据权利要求10所述的转化植物或其自繁或者杂交后代,其特征在于,所述植物为蔷薇。
12.一种营养繁殖体、植物体的一部分、组织或细胞,其特征在于,为权利要求9~11中任一项所述的转化植物或其自繁或者杂交后代的营养繁殖体、植物体的一部分、组织或细胞。
13.一种切花,或由该切花制成的加工品,其特征在于,为权利要求9~11中任一项所述的转化植物或其自繁或者杂交后代的切花,或由该切花制成的加工品。
14.一种方法,其为用于制作具有蓝色系花色的转化植物的方法,其特征在于,包含通过将来自荞麦的C-糖苷化酶,即CGT基因或其同源基因导入宿主植物,使花翠素型花色素苷与黄酮单-C-糖苷在植物的细胞内共存的工序。
15.根据权利要求14所述的方法,其特征在于,所述黄酮单-C-糖苷为芹菜素6-C-糖苷、木犀草素6-C-糖苷、三粒小麦黄酮6-C-糖苷、芹菜素8-C-糖苷、木犀草素8-C-糖苷、三粒小麦黄酮8-C-糖苷或它们的衍生物。
16.根据权利要求14或15所述的方法,其特征在于,所述花翠素型花色素苷选自二甲花翠素、花翠素、矮牵牛素及它们的组合。
17.根据权利要求14~16中任一项所述的方法,其特征在于,包含将权利要求1或2所述的来自荞麦的CGT基因或其同源基因,或者权利要求3~8中任一项所述的载体导入宿主植物的细胞内的工序。
18.根据权利要求14~17中任一项所述的方法,其特征在于,所述植物选自蔷薇、菊花、康乃馨或百合。
19.根据权利要求18所述的方法,其特征在于,所述植物为蔷薇。
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YOSHIHISA NAGATOMO: "Purification, molecular cloning and functional characterization of flavonoid C-glucosyltransferases from Fagopyrum esculentum M. (buckwheat) cotyledon", 《PLANT J》, 26 September 2014 (2014-09-26), pages 1 - 48 * |
佚名: "GenBank: AB909375.1", 《NCBI》, 18 November 2014 (2014-11-18) * |
佚名: "NCBI Reference Sequence: NM_129558.3", 《NCBI》, 20 March 2017 (2017-03-20) * |
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