CN114127299B - 一种增强植物香味的方法 - Google Patents
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Abstract
提供了一种增强植物香味的方法,还提供了BADH2基因或其编码蛋白的抑制剂的用途,用于增强植物的香味;或制备用于增强植物香味的组合物或制剂,其中所述BADH2基因包括BADH2a和BADH2b基因。抑制BADH2基因或其编码蛋白的表达可显著增强植物,尤其是玉米的香味。
Description
技术领域
本发明涉及农学领域,具体地,涉及一种增强植物香味的方法。
背景技术
香稻以其独特浓郁的香味备受消费者的青睐,因此香味成为水稻一个重要的品质性状指标。香米的价格是普通大米的2-3倍,而产自印度和巴基斯坦的BASMATI香米价格更是要高出许多。香稻的挥发性香味物质包含100多种化合物,其中2-乙酰基-1-吡咯啉(2-acetyl-1-pyrroline,2AP)是最主要的香味物质。
我国每年有6亿多亩的玉米种植面积,其中约有两千万亩的鲜食玉米种植面积,玉米种子的市值每年为200亿元左右。在目前的各类报道中,无论是普通籽粒玉米还是鲜食玉米中都没有发现香味玉米的种质。因此,结合现有报道的生物技术,改进玉米香味,提高玉米的附加值,对于丰富玉米种质资源、扩大玉米种植面积、增加玉米适用范围具有重要意义。
因此,本领域迫切需要开发一种新的增强植物香味的方法。
发明内容
本发明的目的在于提供一种新的增强植物香味的方法。
本发明第一方面提供了一种增强植物香味或者赋予植物产生香味的方法,包括步骤:
降低或抑制所述植物中BADH2基因或其编码蛋白的表达量和/或活性,从而增强植物的香味,其中,所述BADH2基因包括BADH2b基因。
在另一优选例中,所述BADH2基因还包括BADH2a基因。
在另一优选例中,所述BADH2基因包括BADH2b和BADH2a基因。
在另一优选例中,所述方法包括给予植物BADH2基因或其编码蛋白的抑制剂。
在另一优选例中,所述增强植物的香味包括提高所述植物的4-氨基丁醛或2-乙酰基-1-吡咯啉(2AP)的含量。
在另一优选例中,所述增强植物的香味为提高所述植物的4-氨基丁醛或2-乙酰基-1-吡咯啉(2AP)的含量。
在另一优选例中,所述植物包括农作物、林业植物、蔬菜、瓜果、花卉、牧草(包括草坪草)。
在另一优选例中,所述的植物包括单子叶植物和双子叶植物。
在另一优选例中,所述植物选自下组:禾本科、豆科、藜科、十字花科植物、或其组合。
在另一优选例中,所述的植物选自下组:拟南芥、水稻、烟草、玉米、高粱、大麦、小麦、小米、大豆、番茄、马铃薯、藜麦、生菜、油菜、白菜、菠菜、甜菜、草莓、或其组合。
在另一优选例中,所述植物选自玉米。
在另一优选例中,所述植物组织或植物细胞中BADH2基因或其编码蛋白的表达量或活性降低了≥30%,≥50%,≥70%,更佳地,≥80%。
在另一优选例中,所述“降低或抑制”是指BADH2基因或其编码蛋白的表达或活性降低满足以下条件:
使所述植物中BADH2基因或其编码蛋白完全失去活性或者失去部分活性,或者,A1/A0的比值≤80%,较佳地≤60%,更佳地≤40%,最佳地为0-30%;其中,A1为所述植物中BADH2基因或其编码蛋白的表达或活性;A0为野生型同种类型植物中相同BADH2基因或其编码蛋白的表达或活性。
在另一优选例中,所述的降低或抑制指与野生型植株中BADH2基因或其编码蛋白的表达水平E0相比,所述植株中BADH2基因或其编码蛋白的表达水平E1为野生型的0-80%,较佳地0-60%,更佳地0-40%,更佳地,0-30%。
在另一优选例中,所述降低或抑制BADH2基因或其蛋白的表达和/或活性通过选自下组的方法实现:基因突变、基因敲除、基因中断、RNA干扰技术、基因编辑技术、导入基因或蛋白的抑制剂、或其组合。
在另一优选例中,所述的基因突变通过以下一种或多种方法获得:自然变异、物理诱变(如紫外线诱变、X射线或Y射线诱变)、化学诱变(如亚硝酸、羟胺、EMS、亚硝基胍等)、生物诱变(如病毒或细菌介导的诱变)、基因编辑或生物合成。
在另一优选例中,所述的突变区域包括外显子和/或内含子区。
在另一优选例中,所述抑制剂选自下组:反义核酸、抗体、小分子化合物、Crispr试剂、小分子配体、或其组合。
在另一优选例中,所述的基因编辑技术选自下组:CRISPR技术、TALEN技术、ZFN技术、或其组合。
在另一优选例中,所述方法包括步骤:
(i)提供一植物或植物细胞;和
(ii)将BADH2基因或其编码蛋白的抑制剂导入所述植物或植物细胞,从而获得改造的植物或植物细胞。
在另一优选例中,所述方法包括步骤:
(i)提供一植物或植物细胞;和
(ii)利用靶向BADH2基因的gRNA以及相应的Cas蛋白导入所述植物或植物细胞;在优选的实施方式中,向所述植物或植物细胞中导入含有所述gRNA和Cas蛋白的表达载体。
在另一优选例中,所述BADH2基因包括野生型BADH2基因和突变型BADH2基因。
在另一优选例中,所述的突变型包括突变后编码蛋白的功能未发生改变的突变形式(即功能与野生型编码蛋白相同或基本相同)。
在另一优选例中,所述的突变型BADH2基因编码的多肽与野生型BADH2基因所编码的多肽相同或基本相同。
在另一优选例中,所述的突变型BADH2基因包括与野生型BADH2基因相比,同源性≥80%(较佳地≥90%,更佳地≥95%,更佳地,≥98%或99%)的多核苷酸。
在另一优选例中,所述的突变型BADH2基因包括在野生型BADH2基因的5′端和/或3′端截短或添加1-60个(较佳地1-30,更佳地1-10个)核苷酸的多核苷酸。
在另一优选例中,所述的BADH2基因包括cDNA序列、CDS序列、基因组序列、或其组合。
在另一优选例中,所述BADH2基因来源于选自下组的一种或多种植物:禾本科、豆科、藜科、十字花科植物。
在另一优选例中,所述的BADH2基因来源于选自下组的一种或多种植物:拟南芥、水稻、烟草、玉米、高粱、大麦、小麦、小米、大豆、番茄、马铃薯、藜麦、生菜、油菜、白菜、菠菜、甜菜、草莓。
在另一优选例中,所述BADH2基因来源于玉米。
在另一优选例中,所述BADH2a基因的编码蛋白的氨基酸序列选自下组:
(i)具有SEQ ID NO.:1所示氨基酸序列的多肽;
(ii)将如SEQ ID NO.:1所示的氨基酸序列经过一个或几个(如1-10个)氨基酸残基的取代、缺失或添加而形成的,具有所述相同或相近功能(甜菜碱醛脱氢酶活性)由(i)衍生的多肽;
或(iii)氨基酸序列与SEQ ID NO.:1所示氨基酸序列具有50%或以上(优选60%以上,70%以上,80%以上,更优选90%以上,更优选95%以上,最优选98%以上,如99%或100%)的同源性、具有相同或相近功能的多肽。
在另一优选例中,所述BADH2b基因的编码蛋白的氨基酸序列选自下组:
(i)具有SEQ ID NO.:2所示氨基酸序列的多肽;
(ii)将如SEQ ID NO.:2所示的氨基酸序列经过一个或几个(如1-10个)氨基酸残基的取代、缺失或添加而形成的,具有所述相同或相近功能(甜菜碱醛脱氢酶活性)由(i)衍生的多肽;
或(iii)氨基酸序列与SEQ ID NO.:2所示氨基酸序列有50%或以上(优选60%以上,70%以上,80%以上,更优选90%以上,更优选95%以上,最优选98%以上,如99%或100%)的同源性、具有相同或相近功能的多肽。
在另一优选例中,所述BADH2a基因的核苷酸序列选自下组:
(a)编码如SEQ ID NO.:1所示多肽的多核苷酸;
(b)序列如SEQ ID NO.:3所示的多核苷酸;
(c)核苷酸序列与SEQ ID NO.:3所示序列的同源性≥95%(较佳地≥98%,更佳地≥99%)的多核苷酸;
(d)在SEQ ID NO.:3所示多核苷酸的5’端和/或3’端截短或添加1-60个(较佳地1-30,更佳地1-10个)核苷酸的多核苷酸;
(e)与(a)-(d)任一所述的多核苷酸互补的多核苷酸。
在另一优选例中,所述BADH2b基因的核苷酸序列选自下组:
(a)编码如SEQ ID NO.:2所示多肽的多核苷酸;
(b)序列如SEQ ID NO.:4所示的多核苷酸;
(c)核苷酸序列与SEQ ID NO.:4所示序列的同源性≥95%(较佳地≥98%,更佳地≥99%)的多核苷酸;
(d)在SEQ ID NO.:4所示多核苷酸的5’端和/或3’端截短或添加1-60个(较佳地1-30,更佳地1-10个)核苷酸的多核苷酸;
(e)与(a)-(d)任一所述的多核苷酸互补的多核苷酸。
在另一优选例中,所述降低或抑制所述BADH2b基因或其编码蛋白的表达量和/或活性通过对BADH2b基因进行突变来实现。
在另一优选例中,所述降低或抑制所述BADH2b基因以及BADH2a基因或其编码蛋白的表达量和/或活性通过对BADH2b基因和BADH2a基因同时进行突变来实现。
在另一优选例中,所述突变包括插入突变、缺失突变、移码突变、取代突变。
在另一优选例中,所述方法还包括步骤:测试植物中香味物质含量的增加量。
在另一优选例中,所述植物中的香味物质的含量为0.01-20mg/kg,较佳地,0.05-10mg/kg,更佳地,0.1-5mg/kg,更佳地,0.1-2mg/kg,更佳地,0.2-1mg/kg。
在另一优选例中,所述香味物质包括2AP。
本发明第二方面提供了一种BADH2基因或其编码蛋白的抑制剂的用途,用于增强植物的香味或者用于赋予植物产生香味;或制备用于增强植物香味的组合物或制剂或制备用于赋予植物产生香味的组合物或制剂,其中所述BADH2基因包括BADH2b基因。
在另一优选例中,所述BADH2基因还包括BADH2a基因。
在另一优选例中,所述BADH2基因包括BADH2b和BADH2a基因。
在另一优选例中,所述组合物包括农用组合物。
在另一优选例中,所述制剂包括农用制剂。
在另一优选例中,所述增强植物的香味包括提高植物中香味物质的含量。
在另一优选例中,所述香味物质包括2AP。
在另一优选例中,所述组合物包含(a)BADH2基因或其编码蛋白的抑制剂,其中所述BADH2基因包括BADH2b基因;和(b)农学上可接受的载体。
在另一优选例中,所述组合物还包括BADH2a基因。
在另一优选例中,所述组合物或制剂的剂型选自下组:溶液剂、乳剂、混悬剂、粉剂、泡沫剂、糊剂、颗粒剂、气雾剂、或其组合。
在另一优选例中,所述抑制剂选自下组:反义核酸、抗体、小分子化合物、Crispr试剂、小分子配体、或其组合。
在另一优选例中,所述反义核酸选自下组:反义RNA、反义DNA、干扰RNA、核酶、或其组合。
在另一优选例中,所述干扰RNA选自下组:siRNA、shRNA、RNAi、miRNA、dsRNA、hpRNA、ihpRNA、或其组合。
在另一优选例中,所述组合物还包括其他增强农作物的香味的物质。
本发明第三方面提供了一种用于增强植物香味或赋予植物产生香味的组合物,包括:
(a)BADH2基因或其编码蛋白的抑制剂,所述BADH2基因包括BADH2b基因;和
(b)农学上可接受的载体。
在另一优选例中,所述组合物还包括BADH2a基因。
在另一优选例中,所述组合物包括农用组合物。
在另一优选例中,所述组合物的剂型选自下组:溶液剂、乳剂、混悬剂、粉剂、泡沫剂、糊剂、颗粒剂、气雾剂、或其组合。
在另一优选例中,所述组合物中,含有0.0001-99wt%,较佳地0.1-90wt%的组分(a),以所述组合物的总重量计。
在另一优选例中,所述抑制剂选自下组:基因编辑试剂、反义核酸、抗体、小分子化合物、Crispr试剂、小分子配体、或其组合。
在另一优选例中,所述反义核酸选自下组:反义RNA、反义DNA、干扰RNA、核酶、或其组合。
在另一优选例中,所述干扰RNA选自下组:siRNA、shRNA、RNAi、miRNA、dsRNA、hpRNA、ihpRNA、或其组合。
在另一优选例中,所述组合物还包括其他增强植物的香味的物质。
本发明第四方面提供了一种本发明第三方面所述的组合物的用途,用于增强植物的香味或赋予植物产生香味。
本发明第五方面提供了一种制备基因工程的植物组织或植物细胞的方法,包括步骤:
降低或抑制植物组织或植物细胞中的BADH2基因或其编码蛋白的表达和/或活性,从而获得基因工程的植物组织或植物细胞。在另一优选例中,所述降低BADH2基因或其蛋白的表达和/或活性通过选自下组的方法实现:基因突变、基因敲除、基因中断、RNA干扰技术、基因编辑技术、导入基因或蛋白抑制剂、或其组合。
本发明第六方面提供了一种制备基因工程植物的方法,包括步骤:
将本发明第五方面所述方法制备的基因工程的植物组织或植物细胞再生为植物体,从而获得基因工程植物。
本发明第七方面提供了一种改良植物的方法,所述的方法包括步骤:
(a)提供一植物细胞、植物组织、植物部分,向所述植物细胞、植物组织、植物部分中导入BADH2基因或其编码蛋白的抑制剂;或者,降低或抑制植物细胞、植物组织、植物部分中的BADH2基因或其编码蛋白的表达和/或活性;其中所述BADH2基因包括BADH2b基因;
(b)将步骤(a)中的植物细胞、植物组织、植物部分再生成植株。
在另一优选例中,在步骤(a)中,利用基因编辑技术改造所述植物细胞、植物组织、植物部分,从而使所述植物细胞、植物组织、植物部分中的BADH2基因或其编码蛋白的表达或活性降低。
在另一优选例中,所述BADH2基因还包括BADH2a基因。
在另一优选例中,所述的基因编辑技术选自下组:CRISPR基因编辑体系、易错PCR、基因重组、TALEN和ZFN。
在另一优选例中,所述方法用于增强植物的香味。
在另一优选例中,所述方法用于提高香味物质的含量。
在另一优选例中,所述方法还包括步骤:对所述植物细胞、植物组织、植物部分或植物,测试香味物质含量的增加量。
在另一优选例中,所述改良植物为增强/提高植物的香味或赋予植物产生香味。
本发明第八方面提供了一种基因工程植物,所述的植物是用本发明第七方面所述的方法制备的。
本发明第九方面提供了一种筛选或鉴定香味植物的方法,检测植物中BADH2基因和/或其蛋白的表达量,其中所述BADH2基因包括BADH2b基因。
在另一优选例中,所述BADH2基因还包括BADH2a基因。
在另一优选例中,所述检测植物的检测部位包括植物的愈伤组织、果实、种子、花、茎、叶、穗、根。
本发明第十方面提供了一种赋予植物产生香味的方法,包括步骤:
降低或抑制所述植物中BADH2基因或其编码蛋白的表达量和/或活性,从而赋予植物产生香味,其中,所述BADH2基因包括BADH2a和BADH2b基因;所述植物为玉米。
在另一优选例中,所述降低或抑制BADH2基因或其蛋白的表达和/或活性通过选自下组的方法实现:基因突变、基因敲除、基因中断、RNA干扰技术、基因编辑技术、导入基因或蛋白的抑制剂、或其组合。
应理解,在本发明范围内中,本发明的上述各技术特征和在下文(如实施例)中具体描述的各技术特征之间都可以互相组合,从而构成新的或优选的技术方案。限于篇幅,在此不再一一累述。
附图说明
图1显示了郑58背景下的C43植株的香味基因的编辑方式,其中A,ZmBADH2a和ZmBADH2b两个基因在DNA序列的编辑形式;B,ZmBADH2a和ZmBADH2b两个基因在cDNA序列的编辑形式;C,编辑后的ZmBADH2a和ZmBADH2b两个蛋白的推导的蛋白序列。蓝色标记的为与郑58相同的氨基酸序列,红色标记的为移码突变的氨基酸序列。郑58:玉米自交系郑58;C43/郑58:郑58背景下的基因编辑植株C43。
图2显示了郑58背景下的P64植株的香味基因的编辑方式。ZmBADH2a基因在DNA序列的编辑形式;B,ZmBADH2a基因在cDNA序列的编辑形式;C,编辑后的ZmBADH2a推导蛋白序列。蓝色标记的为与郑58相同的氨基酸序列,红色标记的为移码突变的氨基酸序列。郑58:玉米自交系郑58;P64/郑58:郑58背景下的基因编辑植株P64。。
图3显示了郑58背景下的C43和P64种子中2AP的含量;其中,1为C43/郑58,2为P64/郑58,3为未编辑的郑58,图中灰色虚线所指示的为2AP的出峰时间。
图4显示了XCW175背景下的P302植株的香味基因的编辑方式,其中A,ZmBADH2a和ZmBADH2b两个基因在DNA序列的编辑形式;B,ZmBADH2a和ZmBADH2b两个基因在cDNA序列的编辑形式;C,编辑后的ZmBADH2a和ZmBADH2b两个蛋白的推导的蛋白序列。蓝色标记的为与郑58相同的氨基酸序列,红色标记的为移码突变的氨基酸序列。XCW175:玉米自交系XCW175;P302/XCW175:XCW175背景下的基因编辑植株P302。
图5显示了XCW175背景下的P302植株的2AP的含量。图中灰色虚线所指示的为2AP的出峰时间,其中,1为P302,2XCW175。
图6显示了郑58背景下的Z82植株的香味基因的编辑方式;(A)Zm00001d050339的DNA序列编辑形式;(B)Zm00001d050339的cDNA序列编辑形式;(C)编辑后的Zm00001d050339的推导蛋白序列。郑58:玉米自交系郑58;Z82/郑58:郑58背景下的基因编辑植株Z82。
图7显示了郑58背景下的Z10植株的香味基因的编辑方式;(A)Zm00001d032257的DNA序列编辑形式;(B)Zm00001d032257的cDNA序列编辑形式;(C)编辑后的Zm00001d032257的推导蛋白序列。郑58:玉米自交系郑58;Z10/郑58:郑58背景下的基因编辑植株Z10。
图8显示了郑58背景下的Z54植株的香味基因的编辑方式;(A)Zm00001d050339和Zm00001d032257两个基因在DNA序列的编辑形式;(B)Zm00001d050339和Zm00001d032257两个基因在cDNA序列的编辑形式;(C)编辑后的Zm00001d050339和Zm00001d032257两个蛋白的推导的蛋白序列。郑58:玉米自交系郑58;Z54/郑58:郑58背景下的基因编辑植株Z54。
图9显示了郑58背景下的Z10、Z82和Z54种子中2AP的含量。图中灰色虚线所指示的为2AP的出峰时间。
图10显示了XCW175背景下的X651植株的香味基因的编辑方式;(A)Zm00001d050339的DNA序列编辑形式;(B)Zm00001d050339的cDNA序列编辑形式;(C)编辑后的Zm00001d050339的推导蛋白序列。XCW175:玉米自交系XCW175;X651/XCW175:XCW175背景下的基因编辑植株X651。
图11显示了XCW175背景下的X610植株的香味基因的编辑方式;(A)Zm00001d032257的DNA序列编辑形式;(B)Zm00001d032257的cDNA序列编辑形式;(C)编辑后的Zm00001d032257的推导蛋白序列。XCW175:玉米自交系XCW175;X610/XCW175:XCW175背景下的基因编辑植株X610。
图12显示了XCW175背景下的X447植株的香味基因的编辑方式;(A)Zm00001d050339和Zm00001d032257两个基因在DNA序列的编辑形式;(B)Zm00001d050339和Zm00001d032257两个基因在cDNA序列的编辑形式;(C)编辑后的Zm00001d050339和Zm00001d032257两个蛋白的推导的蛋白序列。CW175:玉米自交系XCW175;X447/XCW175:XCW175背景下的基因编辑植株X447。
图13显示了XCW175背景下的X610、X651和X447植株的2AP的含量。图中灰色虚线所指示的为2AP的出峰时间。
图14为本实施方式中利用的基因编辑载体示意图。
具体实施方式
经过广泛而深入的研究,本发明人通过对大量的植物性状位点的研究和筛选,首次意外地发现,当同时抑制BADH2a、BADH2b基因或其编码蛋白的表达时,可显著增强植物的香味。本发明人在此基础上完成了本发明。
除非本申请定义,本发明中所使用的科学术语或专业名词具有本领域技术人员所理解的含义,当本领域技术人员理解的含义与本申请所定义的含义出现矛盾时,以本申请所定义的含义为准。
如本文所用,所述“增强植物香味”含义包括赋予没有香味的植物以香味,也包括提高香味植物的香味。
如本文所用,所述“Cripsr制剂”是指可以实现基因编辑效果的各有效成分的组合,包括gRNA(向导RNA)或其编码序列和Cas蛋白或其编码序列,还可以进一步包括载体,以及有利于同源重组或基因表达的元件。
术语“同源性”或“同一性”用于指两个多肽之间或两个核酸之间序列的匹配情况。当两个进行比较的序列中的某个位置都被相同的碱基或氨基酸单体亚单元占据时(例如,两个DNA分子的每一个中的某个位置都被腺嘌呤占据,或两个多肽的每一个中的某个位置都被赖氨酸占据),那么各分子在该位置上是同一的。两个序列之间。通常,在将两个序列比对以产生最大同一性时进行比较。比对方法为本领域技术人员已知的常规方法,比如BLAST运算法则。
术语“基因工程”是指通过人工干预的方式对控制生物遗传信息的核苷酸进行改造和利用,从而获得新的遗传特性、或新品种、或新产品的技术,包括本领域所公开的所有基因改造技术,如基因诱变、转基因或基因编辑等方法。基因诱变的方法包括但不限于物理诱变(比如紫外线诱变)、化学诱变(比如吖啶类染料)、生物诱变(如病毒、噬菌体诱变)等。在一优选实施方式中,本发明的基因工程改造包括用一个或多个sgRNA介导的Cas核酸酶对BADH2基因家族的成员进行基因编辑。
BADH2基因
Badh2是甜菜碱醛脱氢酶的编码基因,甜菜碱醛脱氢酶具有醛脱氢酶活性,可能催化甜菜碱醛、4-氨基丁醛(AB-ald)和3-氨基丙醛的氧化,而4-氨基丁醛是2-乙酰基-1-吡咯啉(2AP)的合成前体,2AP与植物香味密切相关。在本发明中,BADH2基因家族包括BADH2a、BADH2b。
如本文所用,术语“本发明的BADH2基因”包括单子叶或双子叶植物,如玉米、水稻中的BADH2基因或其变体。在一优选实施方式中,本发明的BADH2基因的核苷酸序列如SEQID NO.:3或4所示。
>Zm00001d050339(ZmBADH2a)基因组DNA序列:
>Zm00001d032257(ZmBADH2b)基因组DNA序列:
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本发明还包括与本发明的优选基因序列(SEQ ID NO.:3或4)具有50%或以上(优选60%以上,70%以上,80%以上,更优选90%以上,更优选95%以上,最优选98%以上,如99%,或100%)同源性的核酸,所述核酸也能有效地增强植物的香味。“同源性”是指按照位置相同的百分比,两条或多条核酸之间的相似水平(即序列相似性或同一性)。在本文中,所述基因的变体可以通过插入或删除调控区域,进行随机或定点突变等来获得。
在本发明中,SEQ ID NO).:3或4中的核苷酸序列可以经过取代、缺失或添加一个或多个,生成SEQ ID NO.:3或4的衍生序列,由于密码子的简并性,即使与SEQ ID NO.:3或4的同源性较低,也能基本编码出如SEQ ID NO.:1或2所示的氨基酸序列。另外,“在SEQ IDNO.:3或4中的核苷酸序列经过取代、缺失或添加至少一个核苷酸衍生序列”的含义还包括能在中度严谨条件下,更佳的在高度严谨条件下与SEQ ID NO.:3或4所示的核苷酸序列杂交的核苷酸序列。这些变异形式包括(但并小限于):若干个(通常为1-90个,较佳地1-60个,更佳地1-20个,最佳地1-10个)核苷酸的缺失、插入和/或取代,以及在5’和/或3’端添加数个(通常为60个以内,较佳地为30个以内,更佳地为10个以内,最佳地为5个以内)核苷酸。
应理解,尽管本发明的实例中提供的基因来源于玉米,但是来源于其它类似的植物的、与本发明的序列(优选地,序列如SEQ ID NO.:3或4所示)具有一定同源性(如具有80%以上,如85%,90%,95%甚至98%,99%,或100%序列相同性)的BADH2的基因序列,也包括在本发明的范围内,只要本领域技术人员在阅读了本申请后根据本申请提供的信息可以方便地从其它植物中分离得到该序列。比对序列相同性的方法和工具也是本领域周知的,例如BLAST。
本发明的多核苷酸可以是DNA形式或RNA形式。DNA形式包括:DNA、基因组DNA或人工合成的DNA,DNA可以是单链的或是双链的。DNA可以是编码链或非编码链。编码成熟多肽的编码区序列可以与SEQ ID NO.:3或4所示的编码区序列相同或者是简并的变异体。
编码成熟多肽的多核苷酸包括:只编码成熟多肽的编码序列;成熟多肽的编码序列和各种附加编码序列;成熟多肽的编码序列(和任选的附加编码序列)以及非编码序列。
术语“编码多肽的多核苷酸”可以是包括编码此多肽的多核苷酸,也可以是还包括附加编码和/或非编码序列的多核苷酸。本发明还涉及上述多核苷酸的变异体,其编码与本发明有相同的氨基酸序列的多苷或多肽的片段、类似物和衍生物。此多核苷酸的变异体可以是天然发生的等位变异体或非天然发生的变异体。这些核苷酸变异体包括取代变异体、缺失变异体和插入变异体。如本领域所知的,等位变异体是一个多核苷酸的替换形式,它可能是一个或多个核苷酸的取代、缺失或插入,但不会从实质上改变其编码的多肽的功能。
本发明还涉及与上述的序列杂交且两个序列之间具有至少50%,较佳地至少70%,更佳地至少80%相同性的多核苷酸。本发明特别涉及在严格条件下与本发明所述多核苷酸可杂交的多核苷酸。在本发明中,“严格条件”是指:(1)在较低离子强度和较高温度下的杂交和洗脱,如0.2×SSC,0.1%SDS,60℃;或(2)杂交时加有变性剂,如50%(v/v)甲酞胺,0.1%小牛血清/0.1%Ficoll,42℃等;或(3)仅在两条序列之间的相同性至少在90%以上,更好是95%以上时才发生杂交。
本发明的BADH2核苷酸全长序列或其片段通常可以用PCR扩增法、重组法或人工合成的方法获得。对于PCR扩增法,可根据本发明所公开的有关核苷酸序列,尤其是开放阅读框序列来设计引物,并用市售的DNA库或按本领域技术人员已知的常规方法所制备的cDNA库作为模板,扩增而得有关序列。当序列较长时,常常需要进行两次或多次PCR扩增,然后再将各次扩增出的片段按正确次序拼接在一起。一旦获得了有关的序列,就可以用重组法来大批量地获得有关序列。通常是将其克隆入载体,再转入细胞,然后通过常规方法从增殖后的宿主细胞中分离得到有关序列。
此外,还可用人工合成的方法来合成有关序列,尤其是片段长度较短时。通常,通过先合成多个小片段,然后再进行连接可获得序列很长的片段。目前,已经可以完全通过化学合成来得到编码本发明蛋白(或其片段,或其衍生物)的DNA序列。然后可将该DNA序列引入本领域中已知的各种现有的DNA分子(或如载体)和细胞中。此外,还可通过化学合成将突变引入本发明蛋白序列中。
BADH2基因编码的多肽
如本文所用,术语“本发明多肽”、“BADH2基因的编码蛋白”、可以互换使用,都是指来源于植物(如玉米)的BADH2的多肽及其变体。在一优选实施方式中,本发明多肽的一种典型的氨基酸序列如SEQ ID NO.:1或2所示。
>Zm00001d050339(ZmBADH2a)蛋白序列:
>Zm00001d032257((ZmBADH2b)蛋白序列:
本发明还包括与本发明的SEQ ID NO.:1或2所示序列具有50%或以上(优选60%以上,70%以上,80%以上,更优选90%以上,更优选95%以上,最优选98%以上,如99%或100%)同源性的具有相同或相似功能的多肽或蛋白。
所述“相同或相似功能”主要是指具有甜菜碱醛脱氢酶活性。
本发明的多肽可以是重组多肽、天然多肽、合成多肽。本发明的多肽可以是天然纯化的产物,或是化学合成的产物,或使用重组技术从原核或真核宿主(例如,细菌、酵母、高等植物、昆虫和哺乳动物细胞)中产生。根据重组生产方案所用的宿主,本发明的多肽可以是糖基化的,或可以是非糖基化的。本发明的多肽还可包括或不包括起始的甲硫氨酸残基。
本发明还包括具有BADH2蛋白活性的BADH2蛋白片段和类似物。如本文所用,术语“片段”和“类似物”是指基本上保持本发明的天然BADH2蛋白相同的生物学功能或活性的多肽。
本发明的多肽片段、衍生物或类似物可以是:(i)有一个或多个保守或非保守性氨基酸残基(优选保守性氨基酸残基)被取代的多肽,而这样的取代的氨基酸残基可以是也可以不是由遗传密码编码的;或(ii)在一个或多个氨基酸残基中具有取代基团的多肽;或(iii)成熟多肽与另一个化合物(比如延长多肽半衰期的化合物,例如聚乙二醇)融合所形成的多肽;或(iv)附加的氨基酸序列融合到此多肽序列而形成的多肽(如前导序列或分泌序列或用来纯化此多肽的序列或蛋白原序列,或融合蛋白)。根据本文的定义这些片段、衍生物和类似物属于本领域熟练技术人员公知的范围。
本发明中,所述的多肽变体是如SEQ ID NO.:1或2所示的氨基酸序列,经过若干个(通常为1-60个,较佳地1-30个,更佳地1-20个,最佳地1-10个)取代、缺失或添加至少一个氨基酸所得的衍生序列,以及在C末端和/或N末端添加一个或数个(通常为20个以内,较佳地为10个以内,更佳地为5个以内)氨基酸。例如,在所述蛋白中,用性能相近或相似的氨基酸进行取代时,通常不会改变蛋白质的功能,在C末端和/或末端添加一个或数个氨基酸通常也不会改变蛋白质的功能。这些保守性变异最好根据表1进行替换而产生。
表1
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本发明还包括所要求保护的蛋白的类似物。这些类似物与天然SEQ ID NO.:1或2差别可以是氨基酸序列上的差异,也可以是不影响序列的修饰形式上的差异,或者兼而有之。这些蛋白的类似物包括天然或诱导的遗传变异体。诱导变异体可以通过各种技术得到,如通过辐射或暴露于诱变剂而产生随机诱变,还可通过定点诱变法或其他已知分了生物学的技术。类似物还包括具有不同于天然L-氨基酸的残基(如D-氨基酸)的类似物,以及具有非天然存在的或合成的氨基酸(如β、γ-氨基酸)的类似物。应理解,本发明的蛋白并不限于上述例举的代表性的蛋白。
修饰(通常不改变一级结构)形式包括:体内或体外蛋白的化学衍生形式如乙酸化或羧基化。修饰还包括糖基化,如那些在蛋白质合成和加工中进行糖基化修饰。这种修饰可以通过将蛋白暴露于进行糖基化的酶(如哺乳动物的糖基化酶或去糖基化酶)而完成。修饰形式还包括具有磷酸化氨基酸残基(如磷酸酪氨酸,磷酸丝氨酸,磷酸苏氨酸)的序列。
农用制剂
可将本发明的活性物质(如BADH2基因或其编码蛋白的抑制剂)以常规的方法制备成农用制剂,例如溶液剂、乳剂、混悬剂、粉剂、泡沫剂、糊剂、颗粒剂、气雾剂、用活性物质浸渍的天然的和合成的材料、在多聚物中的微胶囊、用于种子的包衣剂。
这些制剂可用已知的方法生产,例如,将活性物质与扩充剂混合,这些扩充剂就是液体的或液化气的或固体的稀释剂或载体,并可任意选用表面活性剂即乳化剂和/或分散剂和/或泡沫形成剂。例如在用水作扩充剂时,有机溶剂也可用作助剂。
用液体溶剂作稀释剂或载体时,基本上是合适的,如:芳香烃类,例如二甲苯,甲苯或烷基萘;氯化的芳香或氯化的脂肪烃类,例如氯苯,氯乙烯或二氯甲烷;脂肪烃类,例如环己烷或石蜡,例如矿物油馏分;醇类,例如乙醇或乙二醇以及它们的醚和脂类;酮类,例如丙酮,甲乙酮,甲基异丁基酮或环己酮;或不常用的极性溶剂,例如二甲基甲酰胺和二甲基亚砜,以及水。
就液化气的稀释剂或载体说,指的是在常温常压下将成为气体的液体,例如气溶胶推进剂,如卤化的烃类以及丁烷,丙烷,氮气和二氧化碳。
固体载体可用研磨的天然矿物质,例如高岭土,粘土,滑石,石英,活性白土,蒙脱土,或硅藻土,和研磨合成的矿物质,例如高度分散的硅酸,氧化铝和硅酸盐。供颗粒用的固体载体是碾碎的和分级的天然锆石,例如方解石,大理石,浮石,海泡石和白云石,以及无机和有机粗粉合成的颗粒,和有机材料例如锯木屑,椰子壳,玉米棒子和烟草梗的颗粒等。
非离子的和阴离子的乳化列可用作乳化剂和/或泡沫形成剂。例如聚氧乙烯-脂肪酸酯类,聚氧乙烯-脂肪醇醚类,例如烷芳基聚乙二醇醚类,烷基磺酸酯类,烷基硫酸酯类,芳基磺酸酯类以及白蛋白水解产物。分散剂包括,例如木质素亚硫酸盐废液和甲基纤维素。
在制剂中可以用粘合剂,例如羧甲基纤维素和以粉末,颗粒或乳液形式的天然和合成的多聚物,例如阿拉伯胶,聚乙烯基醇和聚乙烯醋酸酯。
可以用着色剂例如无机染料,如氧化铁,氧化钻和普鲁士蓝;有机染料,如有机染料,如偶氮染料或金属钛菁染料;和用痕量营养剂,如铁,猛,硼,铜,钴,铝和锌的盐等。
在本发明中,所述“农用制剂”通常是农用植物生长调节剂,其含有BADH2基因或其编码蛋白的抑制剂作为改良植物性状(如,增强植物香味)的活性成分;以及农业上可接受的载体。
如本文所用,所述“农业上可接受的载体”是用于将本发明的活性物质传送给植物的农药学上可接受的溶剂、悬浮剂或赋形剂。载体可以是液体或固体。适用于本发明的农业上可接受的载体选自下组:水、缓冲液、DMSO、表面活性剂如Tween-20、或其组合。任何本领域技术人员已知的农业上可接受的载体均可用于本发明中。
本发明的农用制剂可包括农用组合物。
本发明的农用制剂可与其他增强植物香味的物质联用。所述其他增强香味的物质可以是本领域技术人员已知的植物生长调节剂。
本发明所述的农用制剂的剂型可以是多种多样的,只要能够使活性成分有效地到达植物体内的剂型都是可以的,从易于制备和施用的立场看,优选的农用制剂是一种喷雾剂或溶液制剂。
本发明所述的农用制剂通常含有占所述农用制剂总重量的0.0001-99wt%,较佳地0.1-90wt%的本发明的活性成分。商品制剂或使用剂型中的本发明的活性成分的浓度可在广阔的范围内变动。商品制剂或使用剂型中的本发明的活性成分的浓度可从0.0000001-100%(g/v),最好在0.0001与50%(g/v)之间。
植物性状的改良
本发明还提供了一种改良植物性状的方法,所述的改良包括:增强植物的香味,包括步骤:降低所述植物中BADH2基因或其编码蛋白的表达量和/或活性,或添加BADH2基因或其编码蛋白的抑制剂。
在本发明中,还可进一步用常规方法将其他可以增强植物香味的物质处理植物或植物种子,从而改良对应植物的性状。
本发明的主要优点包括:
(1)本发明首次发现,通过抑制BADH2a和BADH2b基因或其编码蛋白的表达或活性,可显著增强植物香味。
(2)本发明首次发现,通过抑制BADH2a和BADH2b基因或其编码蛋白的表达或活性,可提高香味物质(如2AP)的含量。
(3)本发明提供了一种快速改良玉米香味的方法,改进的玉米香味(2AP含量)与野生型相比,有大幅度的增加。
(4)本发明创造了一个新的玉米种质资源,具有重要的理论和实际应用意义。
下面结合具体实施例,进一步阐述本发明。应理解,这些实施例仅用于说明本发明而不用于限制本发明的范围。下列实施例中未注明具体条件的实验方法,通常按照常规条件如Sambrook等人,分子克隆:实验室手册(New York:Cold Spring Harbor LaboratoryPress,1989)中所述的条件,或按照制造厂商所建议的条件。除非有特别说明,否则实施例中所用的材料和试剂均为市售产品。
实施例
1、靶点设计及载体构建
通过NCBI(https://www.ncbi.nlm.nih.gov)网站获得OsBADH2基因的蛋白序列,然后与玉米蛋白数据库进行Blast后发现了两个同源基因,分别是位于1号染色体的Zm00001d032257(ZmBADH2b)和位于4号染色体的Zm00001d050339(ZmBADH2a)。
本实施例中利用Cas9和靶向ZmBADH2b和ZmBADH2a的sgRNA在玉米中针对上述两个基因进行编辑,具体操作方法可按照本领域常规的方式进行;本实施方式中,所构建的基因编辑载体的示意图如图14所示;其中,ZmU6pro为U6启动子,Gl y-tRNA为甘氨酸tRNA,ZmU6Ter为终止子,UBIpro为UBI启动子,NLS为核定位信号;载体构建亦可借鉴参考文献(“High-efficiency CRISPR/Cas9 multiplex gene editing using the glycine tRNA-processing system-based strategy in maize”,Weiwei Qi等,《BMC Biotechnology》,2016);本实施例中,Cas9采用植物密码子优化的Cas9,在其他的实施方式中,也可以采用其他方式优化的Cas9。
具体而言,本实施方式中,利用target Design(http://skl.scau.edu.cn/targetdesign/)设计gRNA,其中位于第一个外显子区的ZmBADH2-T1(CGCCAGCGATGGTCCCGCTG(SEQ ID NO.:5)),位于第二个外显子区的ZmBADH2-T2(AGTCGCGGCCACGGTTCCTC(SEQ ID NO.:6))和位于与第二个内含子区的ZmBADH2-T3(AATTAGGCTAGAGCAAAGAG(SEQ ID NO.:7)都是ZmBADH2a基因特异性的序列,其中ZmBADH2-T1与ZmBADH2b的对应序列存在2个碱基的错配,ZmBADH2-T2与ZmBADH2b的对应序列存在2个碱基的错配,ZmBADH2-T3与ZmBADH2b的对应序列存在3个碱基的错配。将ZmBADH2-T1分别与ZmBADH2-T2和ZmBADH2-T3组合构建入U6启动子启动的利用甘氨酸tRNA间隔的基因编辑载体中,分别构建成载体P0195和P0196。另外将ZmBADH2-T1和ZmBADH2-T2构建入载体P0077中,两个gRNA用甘氨酸tRNA间隔开。
位于第一个外显子区的ZmBADH2-T4(GGTTGACGACGGGGAGGCGG(SEQ ID NO.:8))和位于第二个外显子区的ZmBADH2-T5(GCGGCGCTCAAGAGGAACCG(SEQ ID NO.:9))为两个基因的共有序列。将这两个gRNA构建入U6启动子启动的利用甘氨酸tRNA间隔的基因编辑载体中,构建成载体P0593。
位于第一个外显子区的ZmBADH2-T6(CAGCGGTACCATCGCTTGCG)和位于第二个外显子区的ZmBADH2-T7(GAGGAACCGCGGCCGCGATT)都是Zm00001d032257基因特异性的序列,其中ZmBADH2-T6F与Zm00001d050339的对应序列存在2个碱基的错配,ZmBADH2-T7R与Zm00001d050339的对应序列存在2个碱基的错配。将这两个gRNA构建入U6启动子启动的利用甘氨酸tRNA间隔的基因编辑载体中,构建成载体P1801。
具体构建方法如下:
1)、分别利用引物对
ZmBADH2-T1F(TAGGTCTCTTGCACGCCAGCGATGGTCCCGCTGGTTTTAGAGCTAGAAATAGCAAGT(SEQ ID NO.:10))和
ZmBADH2--T2R(TAGGTCTCTAAACAGTCGCGGCCACGGTTCCTCTGCACCAGCCGGGAATCG(SEQID NO.:11)),ZmBADH2-T 1F(TAGGTCTCTTGCACGCCAGCGATGGTCCCGCTGGTTTTAGAGCTAGAAATAGCAAGT(SEQ ID NO.:12))和ZmBADH2-T3R(TAGGTCTCTAAACAATTAGGCTAGAGCAAAGAGTGCACCAGCCGGGAATCG(SEQ ID NO.:13)),ZmBADH2-T4F(TAGGTCTCTTGCAGGTTGACGACGGGGAGGCGGGTTTTAGAGCTAGAAATAGCAAGT(SEQ ID NO.:14))和ZmBADH2-T5R(TAGGTCTCTAAACGCGGCGCTCAAGAGGAACCG TGGTGCACCAGCCGGGAATCG(SEQ ID NO.:15)),ZmBADH2-T6F(TAGGTCTCTTGCACAGCGGTACCATCGCTTGCGGTTTTAGAGCTAGAAATAGCAAGT)和ZmBADH2-T7R(TAGGTCTCTAAACAATCGCGGCCGCGGTTCCTCTGCACCAGCCGGGAATCG);
以质粒P0055为模板扩增,分别割胶回收约200bp的片段为ZmBADH2-T1&2,ZmBADH2-T1&3和ZmBADH2-T4&5,ZmBADH2-T6&7对回收片段用BsaI酶切,并利用试剂盒回收;
2)、分别对骨架载体P0174和P0522进行BsaI酶切,分别回收23Kb片段;
3)、分别将ZmBADH2-T1&2,ZmBADH2-T1&3与P0174用T4连接酶进行连接,构建成最终载体P0195和P0196;将ZmBADH2-T4&5,ZmBADH2-T6&7与P0522进行连接,构建成最终载体P0593和P1801;
4)、将质粒P0055用BsaI酶切,并通过试剂盒进行液体回收,并与BsaI酶切好的ZmBADH2-T1&3进行连接,经质粒扩增后,对质粒用EcoRV和Ps tI进行双酶切,割胶回收1020bp的片段;将P0185质粒利用SmaI和PstI进行双酶切,回收载体片段;将1020bp的DNA片段与P0185的载体用T4DNA连接酶进行连接,构建成最终载体P0077。
5)、将上述3)和4)中的连接产物分别转化大肠杆菌感受态Trans T1,涂布于Kan平板培养,挑8个菌落液体培养2h,PCR菌液检测,选择正确的单克隆2个,进行菌液送测。
6)、选择测序正确的单克隆进行扩繁、保菌、提质粒,转化农杆菌EHA105,挑取5个单菌落培养,PCR检测正确后保菌备用。
2、遗传转化
2.1转化农杆菌
将1中的载体利用热击法转入农杆菌菌株EHA105中,挑取单克隆经液体培养以及PCR鉴定后保存于-80℃冰箱中备用。
2.2菌种活化
从-80冰箱取出菌,在YEP固体培养基上划线,28℃暗培养,通常1~2天可以长出菌落,取菌落重新涂布在新的YEP平板上于28℃暗培养,12~24小时平板菌可长好。
2.3准备农杆菌侵染液
从新活化的菌板上刮取新鲜菌体,重悬到预先加入乙酰丁香酮(AS)的侵染液中,用1ml移液器将菌体打散至看不到颗粒状菌体,调整0D550至0.3~0.4,在摇床上室温低速振荡培养2~4h。
2.4取玉米幼胚
取授粉后10天左右的玉米果穗,拨去苞叶和花丝,浸没在75%酒精中消毒5~10min,期间晃动2~3次,挑取1.5~2mm的幼胚,放入加有AS的侵染培养基(不含农杆菌)中,每管放50~70个幼胚。
2.5侵染
将待转化幼胚用侵染液洗3遍,至侵染液澄清,将侵染液倒干净加入1ml菌液,温和颠倒10下,静置5~10min。取干净培养皿,放3张灭菌滤纸,侵染完毕后颠倒几下后迅速将菌液倒在滤纸上,手持培养皿,变换方向使携带幼胚的菌液在滤纸上均匀分布。
2.6共培养
待最上一层滤纸看不到菌液时用镊子将上层滤纸夹起,沾有幼胚的一面贴在共培养培养基上,用镊子驱赶滤纸和培养基之间的气泡后,用镊子夹住滤纸一角快速揭下,留在滤纸上的幼胚用剥胚刀转移到培养基上,幼胚盾面朝上,22℃暗培养3天。
2.7恢复培养
共培养3天后,将幼胚转移到恢复培养基,每皿30~40个幼胚,25℃暗培养10~14天,单转P0077载体的幼胚恢复培养7天。
2.8筛选
恢复结束后,将转P0077载体的幼胚转移到带双丙铵磷的相应筛选压的筛选一培养基上,25~28℃暗培养14天后,换较高筛选压的筛选二培养基上,25~28℃暗培养14天。
转P0195,P0196,P0593,和P1801这4个载体的幼胚无需愈伤筛选阶段。
2.9分化
将转P0077载体得到的抗性愈伤放到分化培养基上进行分化,夹成小块后平铺在培养基上,25~28℃暗培养,14天换一次培养基。
转P0195,P0196,P0593,和P1801载体额幼胚直接于分化培养基上分化。
2.10生根
将分化产生的小苗转接到生根培养基,每瓶3~4个小苗,25~28℃光照培养直至长成完整植株,生根培养7天后将有白色小根长出,可取样检测。
3、阳性苗的筛选
对于再生苗,取少量叶片以CTAB法提取基因组DNA。
对于转P0077载体的植株以引物对ZmCas9-jc-F2(TCACCGACGAGTACAAGGTC(SEQ IDNO.:16))和ZmCas9-jc-R1(GTCTGGACGAGCTGGATGAA(SEQ ID NO.:17)),ZmCas9-jc-F5(TTCAAGGAGGACATCCAGAAG(SEQ ID NO.:18))和ZmCas9-jc-R5(TCTCGTCGTACTTGGTGTTCAT(SEQ ID NO.:19)),35S-F2(TCATTTGGAGAGGACACGCT(SEQ ID NO.:20))和sp110(GGAGAAACTCGAGTCAAATCTCG(SEQ ID NO.:21))等3对引物分别检测,任何一对引物可以扩增到目的产物,则该再生苗即为转基因阳性苗。
对于转P0195和P0196的植株分别用引物对Cas9-jc-F1(AAGAAGCGGAAGGTCGGTAT(SEQ ID NO.:22))和Cas9-jc-R1(CTCAGGTGGTAGATGGTGGG(SEQ ID NO.:23)),Cas9-jc-F3(CAGAAAGAGCGAGGAAACCA(SEQ ID NO.:24))和Cas9-jc-R3(CCTCAAACAGTGTCAGGGTCA(SEQID NO.:25)),SIE1-jc-F1(CCCGAGGATAATGAGCAGAA(SEQ ID NO.:26))和NOS-jc-R1(CCGATCTAGTAACATAGATGACA(SEQ ID NO.:27))等3对引物分别检测,任何一对引物可以扩增到目的产物,则该再生苗即为转基因阳性苗。
对于转P0593和P1801的植株分别用引物对Cas9-jc-F1(AAGAAGCGGAAGGTCGGTAT(SEQ ID NO.:28))和Cas9-jc-R1(CTCAGGTGGTAGATGGTGGG(SEQ ID NO.:29)),Cas9-jc-F3(CAGAAAGAGCGAGGAAACCA(SEQ ID NO.:30))和Cas9-jc-R3(CCTCAAACAGTGTCAGGGTCA(SEQID NO.:31)),35S-F2(TCATTTGGAGAGGACACGCT(SEQ ID NO.:32))和sp110(GGAGAAACTCGAGTCAAATCTCG(SEQ ID NO.:33))等3对引物分别检测,任何一对引物可以扩增到目的产物,则该再生苗即为转基因阳性苗。
4、基因检测
对于转基因阳性苗以引物对ZmBADH2a-73F(GAGACGTCCTCGCTTTCCAC(SEQ ID NO.:34))和ZmBADH2a-904R(ATGTGCACGCTGCGTTTTAC(SEQ ID NO.:35))扩增ZmBADH2a基因的相应片段,以引物对BADH2b-jc-F1(GAAGTCCACTGCCGAGTTGC(SEQ ID NO.:36))和BADH2b-jc-R1(CGACTGAGTTGTCTCACACTGA(SEQ ID NO.:37))扩增ZmBADH2b基因的相应片段。扩增产物经sanger法进行测序,确认编辑形式,若为测序结果显示双峰,则将PCR产物连接入T载体,选取5个克隆进行测序,确认编辑形式。
5、实验结果
利用P0077转化郑58的幼胚获得了编辑植株C43,其编辑结果是在ZmBADH2a CDS的+24nt后插入一个A,导致其后的编码区发生移码突变,并在新CDS的265-267nt形成一个终止密码子TGA,将会导致蛋白翻译的提前终止;在ZmBADH2b的CDS+27nt处删除一个碱基G,导致其后发生移码突变,并且在新的CDS的262-264nt形成一个终止密码子TAA,导致将会导致蛋白翻译的提前终止。具体编辑方式参见图1(A-C)。
利用P0077转化郑58的幼胚获得了编辑植株P64,编辑植株P64由于其编辑导致在ZmBADH2a CDS的+24nt后插入一个C,导致其后的编码区发生移码突变(图2,A-C),并在新CDS的265-267nt形成一个终止密码子TGA,将会导致蛋白翻译的提前终止;而在P64植株中的ZmBADH2b基因没有发生编辑。具体编辑方式参见图2。
综上,C43植株在ZmBADH2a和ZmBADH2b两个基因同时发生编辑,而P64植株仅在ZmBADH2a基因发生编辑。对C43和P64两个植株的种子提取物通过质谱分析其香味物质2AP的含量;如图3所示,C43植株的种子中2AP的含量为0.648mg/kg、其可以产生香味,而郑58野生型以及P64植株的种子中检测不到2AP、也不会产生香味(图3)。
利用P0593转化糯玉米自交系XCW175的幼胚获得了编辑植株P302,该编辑植物分别在ZmBADH2a、ZmBADH2b的各两个位置发生了编辑,其编辑结果是在ZmBADH2a CDS的+77nt后插入一个C以及+188nt后插入一个A,由编辑后CDS的推导出的蛋白质在第27位氨基酸开始发生移码突变,并在新CDS的262-264nt形成一个终止密码子TGA,将会导致蛋白翻译的提前终止;在ZmBADH2b的CDS+80nt处插入AA以及+191nt处缺失一个A,由编辑后CDS的推导出的蛋白质在第27位氨基酸开始发生移码突变,并在新CDS的265-267nt形成一个终止密码子TAA,将会导致蛋白翻译的提前终止。具体编辑方式参见图4(A-C)。
P302种子提取物通过质谱分析其香味物质2AP含量为0.266mg/kg,并且其种子可以产生香味。而XCW175野生型不产生香味,也检测不到2AP(图5)。
另外,利用P0196转化糯玉米自交系N355的幼胚获得了编辑植株282,将其自交并在其后代植株中鉴定到3株编辑植株为288-4、288-5和288-6,这三株在ZmBADH2a和ZmBADH2b都发生纯合编辑,并且编辑方式相同。其编辑方式为:在ZmBADH2a CDS的+24nt后插入一个A,导致其后的编码区发生移码突变,并在新CDS的265-267nt形成一个终止密码子TGA,将会导致蛋白翻译的提前终止;在ZmBADH2b的CDS+27nt处删除一个碱基G,导致其后发生移码突变,并且在新的CDS的262-264nt形成一个终止密码子TAA,导致将会导致蛋白翻译的提前终止。将三株植株自交,并取授粉后27天处于糯玉米鲜食期未成熟籽粒,通过质谱分析其香味物质2AP含量,三株2AP的含量分别为0.774mg/kg、0.826mg/kg、0.503mg/kg。而对照的N355自交系相同时期的籽粒中检测不到2AP。
利用P0195、P1801和P0593分别转化郑58的幼胚获得了编辑植株Z82(图6A)、Z10(图7A)和Z54(图8A),其中Z82只在Zm00001d050339(ZmBADH2a)上发生编辑,其编辑导致Zm00001d050339CDS的+24nt处缺失一个G,导致其后的编码区发生移码突变(图6B),并在新CDS的259-261nt形成一个终止密码子TGA,将会导致蛋白翻译的提前终止(图6C);Z10只在Zm00001d032257(ZmBADH2b)发生编辑,其编辑导致Zm00001d032257CDS的+15nt到+203nt处缺失188bp,导致其后的编码区发生移码突变(图7B),并在新CDS的79-81nt形成一个终止密码子TAG,将会导致蛋白翻译的提前终止(图7C);Z54在Zm00001d050339和Zm00001d032257都发生编辑,其编辑导致Zm00001d050339 CDS 188-189nt缺失2bp(图8B),并在新CDS的262-264nt形成一个终止密码子TGA,导致蛋白翻译的提前终止(图8C),Zm00001d032257CDS+191nt后插入一个A(图8B),并在新CDS的268-270nt形成一个终止密码子TGA,导致蛋白翻译的提前终止(图8C)。
Z10、Z82和Z54的种子提取物通过质谱分析香味物质2AP含量,结果发现在野生型、Z10和Z82的种子中检测不到2AP,而Z54种子中的2AP含量为0.288mg/kg、并且可以产生香味(图9)。
利用P0593转化糯玉米自交系XCW175的幼胚获得了编辑植株X651(图10A)、X610(图11A)以及X447(图12A)。其中X651只在Zm00001d050339发生编辑,其编辑导致Zm00001d050339 CDS 187-188nt缺失2bp(图10B),并在新CDS的262-264nt形成一个终止密码子TGA,导致蛋白翻译的提前终止(图10C);X610只在Zm00001d032257发生编辑,其编辑导致Zm00001d032257CDS的+191nt处缺失一个A,导致其后的编码区发生移码突变(图11B),并在新CDS的262-264nt形成一个终止密码子TAA,将会导致蛋白翻译的提前终止(图11C);X447植株的Zm00001d050339和Zm00001d032257基因同时发生编辑,编辑分别导致在Zm00001d050339 CDS的+188nt处缺失了一个A(图12B),并在新CDS的199-201nt形成一个终止密码子TGA,将会导致蛋白翻译的提前终止(图12C);在Zm00001d032257的CDS+83nt到+193nt缺失110bp(图12B),并在新CDS的157-159nt形成一个终止密码子TGA,将会导致蛋白翻译的提前终止(图12C)。
对X610、X651和X447的种子提取物通过质谱分析香味物质2AP含量,结果发现在野生型、X610和X651的种子中检测不到2AP,而X447种子中的2AP含量为为0.126mg/kg、并且可以产生香味(图13)。
在本发明提及的所有文献都在本申请中引用作为参考,就如同每一篇文献被单独引用作为参考那样。此外应理解,在阅读了本发明的上述讲授内容之后,本领域技术人员可以对本发明作各种改动或修改,这些等价形式同样落于本申请所附权利要求书所限定的范围。
序列表
<110> 山东舜丰生物科技有限公司
<120> 一种增强植物香味的方法
<130> P2021-1394
<150> CN202010519874.3
<151> 2020-06-09
<160> 37
<170> PatentIn version 3.5
<210> 1
<211> 505
<212> PRT
<213> 玉米(Zea mays)
<400> 1
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aaccgccacg tgctgaaccg cctgccgcac cgcactagcg tccgcaacag caggagacgt 60
cctcgctttc cacctgcaca gagcgccagg gccgtagcgt gccgatcgct cgatcccacc 120
aactgaggcc actgtctagt tgccgactct cctcgcaggc cgcaatctcg cagacacgca 180
ccgtcgcggt atccgtatcg ccagagtcct cacggatggc ctcgccagcg atggtcccgc 240
tgcggcagct cttcgtcgac ggcgagtggc gcccgcccgc gcagggccgc cgcctccccg 300
tcgtcaaccc caccaccgag gctcacatcg gtgagtaagc tcccccgatc actgtcgcct 360
tctattatta tactctacta gctttttact tggcctggcg ctgatgcgct gggtttcgcg 420
tgcgcgcgca ggcgagatcc ctgcgggcac agcggaggac gtggacgccg cggtggcggc 480
ggcgcgggcg gcgctcaaga ggaaccgtgg ccgcgactgg gcgcgcgccc cgggggccgt 540
ccgggccaag tacctccgcg ctatcgccgc caaggtacgc tggtcctctt tgctctagcc 600
taattgggcc agaaaccact cctgtgctgt gcctctgctt ccaatggttg gtacctacta 660
gcccctaagc tagtaagtta ttcagagtgg attgaatgtg tgacaactaa gtagggggtg 720
tttgggagtg aagttttttc aaagttttag aagaatactg cagtattctc aaaaactata 780
gtattatata ctaaaaaggt gtttggcaag ccagctaaaa tctctgtttt caaaactaaa 840
gtattacaaa tactgcagct cttttgaagt attctaaact tatgtttata cctcagtttt 900
ctctgtaaaa cgcagcgtgc acatctctcc cacgtgcaaa aaaatacttt gcttccaaac 960
acctctatgt attcaatact gtggtattct aaaactgtag tattttaaac tgtggtattg 1020
tcatgactaa aatataccgt agtatttcaa aaaccgtggt ttccaaaaac tttgttccca 1080
aacaggcccg tacttgtgaa atatacgttg gtttccgtac actccattac tgtttctgcg 1140
ttacatgagc agaggcacag aaagtacaaa ttaaactcct ccaattgtaa accaatctat 1200
tcaggggaaa tccttatgtt ctgcttttgg tgtgaacaaa ctattgcact tcccatcaat 1260
acagggtgtg ttagctttat tatcaactcc aatttcaaaa cacctcaaaa gatctttctc 1320
tttaaacttc aattgagaaa gtcttgtaac atttggctgg gccatcaaag ttttgttcaa 1380
atatattcac atatgtcaca ttttgtgtat aagaatgtct tcttattcag gtgattgaga 1440
ggaaacctga gctggctaag ctagaggcac ttgattgtgg gaagccttat gatgaagcag 1500
catgggacat ggtatgtgac accttatgga agtgcaattt attttcacta gcttgcaatt 1560
acaatctgtt tggtattcca tttatcagga tgatgttgct gggtgctttg agtactttgc 1620
ggatcaggca gaagccttgg ataaaaggca aaattcccca gtttctcttc caatggaaac 1680
ttttaaatgt catctccgaa gagagcctat tggggtagtt gggctgataa ctccttggta 1740
ttttcatttc cccctcactt agctgttact tgtatttatg accaaaactc acaggcgagt 1800
gccttattca gtaatacaac ttcatttctc atagatctca gggttgtata aatgtatagc 1860
gaataaactg catattctct aaccacacac agtcttttaa acctaatttc gataactttc 1920
ctcatcacca ttctgaaata tttccagtat tcagatttgt acttcctatc cagtgttgaa 1980
tctcacctat atatccaagt taatgtcaat tttttttgct tgtgtcaatg tttggccaat 2040
atcaagtatt ctttctatgt tgatatgaca gtgagctttg tcctagtata agtgtacgct 2100
tcttttctag aaactcacta ttgtgttact atctttgttc acataattgt tgtgcgggcc 2160
atttcagact attatgtcta tatataagat gctgttgact aagtaattat ttacttatat 2220
ttatgttttc ttcatatgtt cttgttttac ttattctgat tctgactcta aagcattata 2280
ctttgtactt gtatttgtgt tgcaggaact atcctctact gatggctaca tggaagatag 2340
cccctgcatt ggctgctggt tgtacagctg tgctgaagcc atctgaactg gcttctgtgt 2400
aagtattagc atgttatatc atgttcatgc tacaagcaac agcattttat gtgcctacgt 2460
tttgaattac caggacttgc ttagagcttg ctgatatctg taaagaagtc ggtctccctt 2520
ctggtgtctt gaacattgtg acaggattag gtcctgatgc tggcgctcct ttgtcagcac 2580
acccagatgt tgacaaggta aactattctg catataatga cataatcatg catgagcact 2640
cctcatagca tgggttttct gtttatttag gtcgctttta ctgggagttt tgaaactggc 2700
aagaaaatta tggcatcggc agctcctatg gtcaaggttc gtctgtgaat tatgttttat 2760
ttatagtaat gtgctgtgcc gtgttgtttt gctctcttct agttgataaa ttatcagaaa 2820
gtatgatttg agtgccctat tttaagagga aattctgttg cattccttaa cattgcattt 2880
ttcttgacag cctgttacac tggaacttgg tggaaaaagt cctatagtag tatttgatga 2940
tgttgacatt gacaaaggta cgtttatatt ctaaggatta caaaattcct gaagttcatt 3000
ggtatttgtg gattgttgca atttgcagag aatttaactg atgagaatgc taaatgccta 3060
aacccttcaa agcccttcat cggtttctgg gattttcata gaaatactca gctgagcctg 3120
atcaggctac tacaataaga attattttgg tttgaaaatt gattgttagg atttgtaaaa 3180
tacagtccat aaagttgtgg ggaaaaccca catgcttgct atagtttgga cttttgacca 3240
cagcaccaga ctttttatta gctgcatttt gcacaaaatt tagagatgca gcatgccagt 3300
tgaataaaaa gatgaaacag aaagtatttg atgcattatg tatatgttac atgatttgat 3360
ttcatcatag accttgagaa atatagggct gaacttcttt gtagaacata agttttagtt 3420
ccaagaaagg gaacttttac aaattcactg accaacttac tttagttatg ctgatacttt 3480
gctcattgtg gaattatgga ttatgctata cttccatcat tgatcattcg ttctaaatat 3540
aaaacagctg tcgagtggac tctgtttggg tgcttttgga ccaatggtca gatttgcagc 3600
gcaacatctc gtcttcttat ccatgtaagt ttcatatgtc attgccagta gcagttggga 3660
tacttttcac acataatagc tgtcaaccat aatatatttc cttgagtggt gttgctttag 3720
ctttaccccg ccgtgccgtt aacttctgat tcgagaaggc taccatttca aaatcgctgt 3780
actgaatata tagcagtata ctatgaggtt tacacctata ttgaacaata tatgtgcccc 3840
attgcctaag gttcacactc ttcaatttgt tgtgaactgt gcactttggt ttggtgtata 3900
tttagaagta tatattctca atgacttgat tgtttttact ccctctgata ccaaatgcaa 3960
gtagtttagg acatagtcct gatctccaat gtgacactgt ttgaccgact tgaacaaaaa 4020
acaacctaac aattgggatt ggaaggaata ggttaacaat ggctatttac tgctatatat 4080
tactctttca aatcagatct caattatgaa tcagttatgt tatgtactta tgttattgtt 4140
gtcagtaaca cagtaaccaa gttctgcctc caatgaaacc gttgcagaca aaaattgcta 4200
aaaaatttaa tgagaggatg gttgcatggg ccaaaaatat taaggtttcg gatccacttg 4260
aagagggttg caggcttggg ccagttgtta gtgaaggaca ggtactacat gtcaactttg 4320
tctaaatgta aattcaccat gccaatgcta tgatttgtag cttctttctt atttgacaat 4380
ttggtacttc tgatatagta gctgtaaact ttgtctaaat gtaaattcac catgcgaatg 4440
ctataatttg tagcttcttt ctgatttgac aatttggtac ttctgattta gtagttgtac 4500
actcttgttt ttgctagttt ctacaggatg tagcttcctc tgattttgag aggatctgtg 4560
ttcagatgca ctgaccaaca ccacctttag atttactttg ttatatttta gtatgttttt 4620
acagcaagag ctatggtatg gattaatgtt tatggtcata ctggattatg agaaaatgac 4680
accctaatca ccaccatatt ggtaaccata cacatagttt cagattacct tgtttcatct 4740
atagttgttc ataggagcct tatatatatt tttgtgttct cagtatgaga agattaagaa 4800
gttcatatcg aatgccaaaa gccaaggtgc tactattctg actggaggtg ttagacctgc 4860
ggtaaggcct gcatttggat ttacaaagat ccacatgcta aaagcatgct atagagagat 4920
agaactaatg tttcatgttt tacagcatct tgagaagggg ttctttattg aaccaactat 4980
tattactgat atcaccacat caatggaaat ctggagggag gaagtctttg gtccagtcct 5040
gtgcgttaaa gaatttagca ctgaagatga agccatcgaa cttgccaacg atacacagtg 5100
agctgattct ttttagcgca gtttcgatgt cttttttgca ggcttcacag gtcatgtgta 5160
gtccatagct tatgaattta tgccagtatc cttcttttga atttttgata ggtatggttt 5220
ggctggtgct gtaatttctg gtgatcggga gcgctgccag aggttatctg aggtacgtat 5280
aagtgaagag gtccacagta tttggttggc aaattgactg catcagactg atactcagag 5340
tgcacttttg tgtgtgtgtg tgtgtgtatg tggggtgggg ggggataagg ggtcaccatt 5400
atagctctgt gcttctgtcc atgctatcac tcgtttgaag ggttcaattt ggtacttttg 5460
ataccatacc actcaattcg attttcgtgt tgcaacatgg aattcgtggt gcacttgctc 5520
tcagaactgg ttttgggtat ttctactaca catctctaat gaaacttgcg caacaggaga 5580
ttgacgctgg atgcatctgg gtaaactgct cgcaaccctg cttctgccaa gctccctggg 5640
gcgggaacaa gcgcagtgga tttggacgtg agcttggaga agggtgggta acatggaacg 5700
acaacatttc taacagatat tgctatggga tctgaatgac actggctgtt tatctgtagg 5760
ggcattgata actacctgag cgtcaagcaa gtcacggagt acatctctga tgagccgtgg 5820
ggatggtacc aatccccctc caagctgtaa actggcaaga cgaaaatttg tctgtttcgg 5880
tagaataaat atatctcgat gctatacaga acgaacccgt gtatcacatt gaa 5933
<210> 4
<211> 5807
<212> DNA
<213> 玉米(Zea mays)
<400> 4
atctgtgtag tatttccgtt aagcctgaat ggagtcgcgg acagcgtgcc gataaagcgt 60
tccagttata tctctatctg tatagcctac actacacagg gccgcagctt gccgatccct 120
agcgacgaag tccactgccg agttgccgac tcaccgactc ccctcgcaag tcgcaatctc 180
cactctccag agtccaaact ccactcctca ccgatgatgg cctcgcaagc gatggtaccg 240
ctgcggcagc tctttgtcga cggcgagtgg cgcccgcccg cgcagggccg ccgcctcccc 300
gtcgtcaacc ctaccactga ggctcacatc ggtgagtgac cgccccccga tcgcaatcat 360
ccctgttttc ttctattata cttgttgctt ggcctggcgc tgatgcgcta ggtctcgcgt 420
gcacgcgcag gcgagatccc ggcgggcacg gcggaggacg tggacgccgc ggtggcggcg 480
gcgcgcgcgg cgctcaagag gaaccgcggc cgcgattggg cgcgcgcgcc gggggccgtt 540
cgggccaagt acctccgcgc catcgccgcc aaggtacgct ggtcctcttt gctctagcta 600
atttaattgg gtcaaaaacg actccagtgc tgtgcttctg cttccaagca ttggtatcta 660
ctccctccgc cccaatataa atgtacatct cgcttaggaa tactaccata attcggatgt 720
gaattcagtg tgagacaact cagtcgcggc atctattttt tgtgctttga tgggtatgat 780
tggtgactgg atctgtactt ggggatgcta ggagtgaact tctcagcacc tttgctatat 840
gcgttttcct tacacttcaa cactgttact gttttacatg cacagaggca cagaaagtac 900
aaattaagtt ccttctattc aggagaagtc cttatgttcc gcttttgttg tgaacaagct 960
attgctttct gccaatacag ggtgttttga cctctagtat caactccaaa ttcaaaacac 1020
cttaaaagat ctttctcttt aaacttccat ttgtaaagtc ttgtaacatt tggttgggtc 1080
atcaaagttt tgttgaaata tatttacaca tgtcacattt tgcatgtaaa aatgtcttct 1140
tattcaggta attgagagga aacaagagct agctaagcta gaggcacttg attgtgggaa 1200
gccttatgat gaagctgcat gggacatggt atgtgacaca ccttatggaa gtgtgcaatt 1260
tttttttcca gtttgcaatg acaatctgtc tggtattcca tttatcagga tgatgttgct 1320
gggtgctttg agtacttcgc agatcaggca gaagctttgg acaaaagaca aaattcccca 1380
gtttctcttc cgatggaaac ttttaaatgc catctccgaa gagagcctat tggggtagtt 1440
gggctgataa ctccttggta ttttcatacc tacctcacct tgctgttatt cgtatttatg 1500
atcaaagctc acaggcaagc gacctgtatt cagtagtaca gcttcgtttc ctgtaagacc 1560
tcagagttct atagcaaaaa aatctgcata ttctctaacc acacaatttt cttaaactat 1620
tttttgacaa ctttcctcat catcgtcctg aaatatttcc agtgttcata tttgtacttc 1680
ctatccagtt tttaatctca cctatgatct atatatccaa gttaatgtca aattatttgc 1740
ttgtgtgaat gtttggccaa tgtcaatatg ctatcttggc acaacgttgt gctttgtccc 1800
agtacaactg ttgcttctgc tccataagct cactattctg ctattatcct ttttcacgta 1860
tttgttgggc cggggcattt cactttattg tgtctatata taaaatgttg ttgactaagt 1920
aactgattta tctatttttt catatgtttt tgttttactt attctgattc taaagtatta 1980
tgctttgtac ttgtatttgt attgcaggaa ctatcctctc ctgatggcta catggaaggt 2040
agctcctgct ctggctgctg gttgtgcagc tgtgctaaag ccatctgaat tggcttctgt 2100
gtaagtatta gcattctata ttctcatgct agaagcaatg gcatttgaag tgcctacctt 2160
ttgaattccc aggacttgct tagagcttgc tgatatctgt aaagaagtcg gtcttcctcc 2220
cggtgtctta aacattgtga caggattagg tcctgatgct ggtgctcctt tgtcagcaca 2280
cccagatgtt gacaaggtaa actgttctgc atataatgac ataatcatgc atgtgcatga 2340
cacatcaatg aaaacattat ttctcatatc atgtgttttc tgtgtttgta ggtcgctttt 2400
actgggagtt ttgaaactgg taagaagatt atggcggctg cagctcctat ggttaaggtt 2460
tgtctgtgaa tttatgtttt atttcaagta atgaactgtg cattattcat tttgctctct 2520
tttagttgat aaattatcag aagtatgatt tggataccct acgattaaga ggaaattcta 2580
ttgcattctt taacattgca tttttcttga cagcctgtta cactggaact tggtggaaaa 2640
agtcctatag tagtatttga tgatgttgac attgacaaag gtacatgtat attctaaggc 2700
ttactaaatt cctaaatttc attgatattt gtggattgtt gcaaattgca aagaattgaa 2760
ctgatcacaa tgctaaacgc ctaaaactcc tcaaagccct tcttggtatc tgagcttttc 2820
gtagaaatac tcaactgagc aggctactac aacaagaatt gctttggtat tgaaaattga 2880
ttgttaggat ttgtaaaata cagtacataa aattgcgggg gaaactacat gcttgctatg 2940
gttttgacca gagtaccaga ctttgtgttg gttgtctgca tctgacacaa aatttggaga 3000
tgcattatgc cagttgaata aaaagatgaa atcgaaagta ttttatgcat tatgtacatg 3060
ttatatgatt tcatatttca tcattgtttt tgagaaatat agggctgaac ttctgtggag 3120
aacataagct ttagttccaa gaaaggggat tttataaatt tactgattct gacttgcttc 3180
agttatgctg atactttgct caacatggaa ttgtggattc tgatatgctt ccatcgttga 3240
tccttctaaa taacagctgt tgagtggact ctgtttgggt gcttttggac caatggtcag 3300
atttgcagcg caacatctcg tcttcttgtc catgtaagtg ccacatgtta ttgcttgtat 3360
tgtcacttgg aatacttctc acacttaata gttgtcaacc atgatctatt tcttggagtg 3420
gtgtcactta gcttgtcctt gccctgatgt taacttctga taatcttctg cgggagtgat 3480
tttaaaattg cctcacagaa taaatagcag catatatgtg ctttacgcct gttttgaata 3540
aatagctggt gtgtgtgttg tgtttcctag tgttgttttt tccttacctt tgctacctta 3600
atgaaatgac atgcaactct tgcattgttt gagaaaaaaa aacaatatat gcgttccatt 3660
gcttaaggtc tgggtttgaa ctttgaactc ttcaatttgc gcacttaaaa aaaaagtctt 3720
ctatttgctg tgaagtgtgt gcttccatct ggtgcatatt tagaaatata atctcaatga 3780
cttgattgtt ttttactcat ctgatacgaa acctaagatg tttaggacat tgccacggtc 3840
tccaatgtaa cactttgact ttagttttgt aagttatatt atttaaaagg gacacaccta 3900
tatattttat gaaattattt ttcatgatga actagtaaca ttggtcatgc tttactaatc 3960
tatgcaactt gacatatatt gatgagtcaa gaaagaagtt tgaccgactt gaatcgaaaa 4020
atgacctaca attgggacca aaaggagtag gttaaccatg gctatttatc agtctatatt 4080
actatttgtc tcttccaaat cacatttcag ttatgaatca gttatgcact atgttattgt 4140
tgtcagtaac atagtaatca agttttgcct tcaatgaaac tgttgcagac aaaaattgct 4200
aaagaattta atgagaagat ggttgcatgg gccaagaata ttaaggtttc tgatccgctt 4260
gaagagggtt gcagacttgg gccagttgtt agtgaagggc aggtattaca tgttaacttc 4320
atctaaatct aacgtcacca ttgacaatgc tatgattttt gtctcctttc tgatttgaca 4380
gttcggtact tcctatttag ttgctataca ctcttgttct ttctagtttc tacgggctac 4440
aggaccagga tgtagctccc gccgattttg agaggatcta ttttgatatg cactgactga 4500
caccaccttc tttatattac tttgttatat tacagtattt ttgttacaga aagagctatg 4560
gtatagatta atgtttatgg tcatatatat aacacaagac tgagaaagtg gcaccctaat 4620
cgccactatc ttggcaacca tatacatagt ttcagattac cttgttgctt tatagttgtt 4680
cacgagtctt attcatgttt ttgcgttctc agtatgagaa gattaaaaag ttcatattga 4740
atgccaaaag cgaaggtgct actattctga ctggaggtgt tagacctgcg gtaaggcctg 4800
catttggatt tacatggacc acatgttata gtgagacata actaatgttt tatattttac 4860
agcatcttga gaaggggttc tttattgaac caacaatcat tactgatatc accacatcaa 4920
tggaaatttg gagggaggaa gtcttcggtc cagtcctgtg tgttaaagaa tttagcactg 4980
aagatgaagc cattgaactg gccaatgata cacagtgagc tacttatttt tagtacaact 5040
tcaatgctct tttgcaggct ttgcatctca tgcgtagtca gtagcttaag aattcatgtc 5100
actatccttt cttttatagg tatggtttag ctggtgctgt aatttctggt gatcgtgagc 5160
gctgccagag attatctgag gtatgtttag tgaaggggtt cacagtattt ggctcgtggt 5220
ttgatgactg aatcagactg atactcggag tctctctctc tctctctctc tcttgctctg 5280
tccatgttgc cacattgact tgctctcaga actgtttttg ggtactccta ctagacatct 5340
ctaatgaaaa cttgcgcaac aggagattga tgctggaatt atctgggtaa actgctcaca 5400
gccctgcttc tgccaggctc cctggggcgg taacaagcgt agtggatttg gacgtgagct 5460
tggagaaggg tgggtaacat ggaacgacaa cattttaaga cgtctaacag ctatatttgg 5520
gatctgactg acactgcctg tttacctgta ggggcattga taactacctg agcgtcaagc 5580
aagtcacgga gtacatctct gatgagccgt ggggatggta tcgatccccc tccaagctgt 5640
aaactggcat gaaattttgt ttgttccatt agaataaata tatctcgatg ctatacagaa 5700
cgaacccgtg tatcaaatgg agtatgacat tgtgaaaaag gcacatcagc ggaacaaatg 5760
tcagagttga caattccctc taatgcaaat agttaactcg cccttga 5807
<210> 5
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 5
cgccagcgat ggtcccgctg 20
<210> 6
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 6
agtcgcggcc acggttcctc 20
<210> 7
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 7
aattaggcta gagcaaagag 20
<210> 8
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 8
ggttgacgac ggggaggcgg 20
<210> 9
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 9
gcggcgctca agaggaaccg 20
<210> 10
<211> 57
<212> DNA
<213> 人工序列(artificial sequence)
<400> 10
taggtctctt gcacgccagc gatggtcccg ctggttttag agctagaaat agcaagt 57
<210> 11
<211> 51
<212> DNA
<213> 人工序列(artificial sequence)
<400> 11
taggtctcta aacagtcgcg gccacggttc ctctgcacca gccgggaatc g 51
<210> 12
<211> 57
<212> DNA
<213> 人工序列(artificial sequence)
<400> 12
taggtctctt gcacgccagc gatggtcccg ctggttttag agctagaaat agcaagt 57
<210> 13
<211> 51
<212> DNA
<213> 人工序列(artificial sequence)
<400> 13
taggtctcta aacaattagg ctagagcaaa gagtgcacca gccgggaatc g 51
<210> 14
<211> 57
<212> DNA
<213> 人工序列(artificial sequence)
<400> 14
taggtctctt gcaggttgac gacggggagg cgggttttag agctagaaat agcaagt 57
<210> 15
<211> 54
<212> DNA
<213> 人工序列(artificial sequence)
<400> 15
taggtctcta aacgcggcgc tcaagaggaa ccgtggtgca ccagccggga atcg 54
<210> 16
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 16
tcaccgacga gtacaaggtc 20
<210> 17
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 17
gtctggacga gctggatgaa 20
<210> 18
<211> 21
<212> DNA
<213> 人工序列(artificial sequence)
<400> 18
ttcaaggagg acatccagaa g 21
<210> 19
<211> 22
<212> DNA
<213> 人工序列(artificial sequence)
<400> 19
tctcgtcgta cttggtgttc at 22
<210> 20
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 20
tcatttggag aggacacgct 20
<210> 21
<211> 23
<212> DNA
<213> 人工序列(artificial sequence)
<400> 21
ggagaaactc gagtcaaatc tcg 23
<210> 22
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 22
aagaagcgga aggtcggtat 20
<210> 23
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 23
ctcaggtggt agatggtggg 20
<210> 24
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 24
cagaaagagc gaggaaacca 20
<210> 25
<211> 21
<212> DNA
<213> 人工序列(artificial sequence)
<400> 25
cctcaaacag tgtcagggtc a 21
<210> 26
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 26
cccgaggata atgagcagaa 20
<210> 27
<211> 23
<212> DNA
<213> 人工序列(artificial sequence)
<400> 27
ccgatctagt aacatagatg aca 23
<210> 28
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 28
aagaagcgga aggtcggtat 20
<210> 29
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 29
ctcaggtggt agatggtggg 20
<210> 30
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 30
cagaaagagc gaggaaacca 20
<210> 31
<211> 21
<212> DNA
<213> 人工序列(artificial sequence)
<400> 31
cctcaaacag tgtcagggtc a 21
<210> 32
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 32
tcatttggag aggacacgct 20
<210> 33
<211> 23
<212> DNA
<213> 人工序列(artificial sequence)
<400> 33
ggagaaactc gagtcaaatc tcg 23
<210> 34
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 34
gagacgtcct cgctttccac 20
<210> 35
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 35
atgtgcacgc tgcgttttac 20
<210> 36
<211> 20
<212> DNA
<213> 人工序列(artificial sequence)
<400> 36
gaagtccact gccgagttgc 20
<210> 37
<211> 22
<212> DNA
<213> 人工序列(artificial sequence)
<400> 37
cgactgagtt gtctcacact ga 22
Claims (11)
1.一种增强植物香味的方法,其特征在于,包括步骤:
降低或抑制所述植物中BADH2基因或其编码蛋白的表达量和/或活性,从而增强植物的香味,其中,所述BADH2基因包括BADH2a和BADH2b基因;所述BADH2a基因编码蛋白的氨基酸序列如SEQ ID NO.1所示;所述BADH2b基因编码蛋白的氨基酸序列如SEQ ID NO.2所示;所述植物为玉米;
所述降低或抑制BADH2基因或其蛋白的表达和/或活性通过选自下组的方法实现:基因突变、基因敲除、基因中断、RNA干扰技术、或其组合。
2.一种赋予植物产生香味的方法,其特征在于,包括步骤:
降低或抑制所述植物中BADH2基因或其编码蛋白的表达量和/或活性,从而赋予植物产生香味,其中,所述BADH2基因包括BADH2a和BADH2b基因;所述BADH2a基因编码蛋白的氨基酸序列如SEQ ID NO.1所示;所述BADH2b基因编码蛋白的氨基酸序列如SEQ ID NO.2所示;所述植物为玉米;
所述降低或抑制BADH2基因或其蛋白的表达和/或活性通过选自下组的方法实现:基因突变、基因敲除、基因中断、RNA干扰技术、或其组合。
3.根据权利要求1或2所述的方法,其特征在于,所述降低或抑制所述BADH2基因或其编码蛋白的表达量和/或活性通过对BADH2b基因和BADH2a基因同时进行突变来实现;
所述BADH2a基因的核苷酸序列的CDS的+24nt后插入一个A,所述BADH2b基因的核苷酸序列的CDS+27nt处删除一个碱基G,所述玉米为郑58或糯玉米自交系N355。
4.一种BADH2基因或其编码蛋白的抑制剂的用途,其特征在于,用于增强植物的香味或赋予植物产生香味;或者,制备用于增强植物香味或赋予植物产生香味的组合物或制剂,其中所述BADH2基因包括BADH2a和BADH2b基因;所述BADH2a基因编码蛋白的氨基酸序列如SEQID NO.1所示;所述BADH2b基因编码蛋白的氨基酸序列如SEQ ID NO.2所示;所述植物为玉米;
所述抑制剂选自下组:基因编辑试剂、反义核酸、或其组合。
5.根据权利要求4所述的用途,其特征在于,所述基因编辑试剂为Crispr试剂。
6.一种用于增强植物香味或赋予植物产生香味的组合物,其特征在于,包括:
(a)BADH2基因或其编码蛋白的抑制剂,所述BADH2基因包括BADH2a和BADH2b基因;所述BADH2a基因编码蛋白的氨基酸序列如SEQ ID NO.1所示;所述BADH2b基因编码蛋白的氨基酸序列如SEQ ID NO.2所示;和
(b)农学上可接受的载体;
所述植物为玉米;
所述抑制剂选自下组:基因编辑试剂、反义核酸、或其组合。
7.根据权利要求6所述的组合物,其特征在于,所述基因编辑试剂为Crispr试剂。
8.一种权利要求6-7任一所述的组合物的用途,其特征在于,用于增强植物的香味或赋予植物产生香味,所述植物为玉米。
9.一种制备基因工程的植物组织或植物细胞的方法,其特征在于,包括步骤:
降低或抑制植物组织或植物细胞中的BADH2基因或其编码蛋白的表达和/或活性,从而获得基因工程的植物组织或植物细胞,所述BADH2基因包括BADH2a和BADH2b基因,所述BADH2a基因编码蛋白的氨基酸序列如SEQ ID NO.1所示;所述BADH2b基因编码蛋白的氨基酸序列如SEQ ID NO.2所示;所述植物为玉米;
所述降低或抑制BADH2基因或其蛋白的表达和/或活性通过选自下组的方法实现:基因突变、基因敲除、基因中断、RNA干扰技术、或其组合。
10.根据权利要求9所述的方法,其特征在于,所述降低或抑制所述BADH2基因或其编码蛋白的表达量和/或活性通过对BADH2b基因和BADH2a基因同时进行突变来实现;
所述BADH2a基因的核苷酸序列的CDS的+24nt后插入一个A,所述BADH2b基因的核苷酸序列的CDS+27nt处删除一个碱基G,所述玉米为郑58或糯玉米自交系N355。
11.一种制备基因工程植物的方法,其特征在于,包括步骤:
将权利要求9-10任一所述方法制备的基因工程的植物组织或植物细胞再生为植物体,从而获得基因工程植物;所述植物为玉米。
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