CN107190015B - 玉米糖基转移酶基因ufgt2在提高植物体内黄酮含量中的应用 - Google Patents
玉米糖基转移酶基因ufgt2在提高植物体内黄酮含量中的应用 Download PDFInfo
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- CN107190015B CN107190015B CN201710562086.0A CN201710562086A CN107190015B CN 107190015 B CN107190015 B CN 107190015B CN 201710562086 A CN201710562086 A CN 201710562086A CN 107190015 B CN107190015 B CN 107190015B
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Abstract
本发明公开了一种玉米糖基转移酶基因UFGT2在提高植物体内黄酮含量中的应用。其中所述玉米糖基转移酶基因UFGT2的核苷酸序列如SEQ ID No.1所示,其是通过RT‑PCR技术从玉米中克隆获得。本发明利用基因UFGT2构建植物过表达载体,进行植物转基因操作,获得转基因植物。通过检测表明:获得的转基因植物中黄酮含量得到显著提高。预示本发明实施后有望能显著提升作物品质,可用于后续的作物品种改良,对我国农业生产和人类健康具有重大意义。
Description
技术领域
本发明涉及一种糖基转移酶基因的应用,尤其涉及一种玉米糖基转移酶基因UFGT2在提高植物体内黄酮含量中的应用,属于基因工程领域。
背景技术
黄酮类化合物(flavonoids,又称类黄酮),是植物为适应环境而合成的一大类次生代谢产物,其数量列为天然酚类化合物之首。研究表明,类黄酮在植物体内可以有效清除活性氧自由基,保护植物抵御紫外线伤害。另外,对动物和人而言,类黄酮具有很好的保健价值,适量摄入黄酮类化合物除了可抗衰老之外,还可以减少癌症、肿瘤、心血管疾病、抑郁症的发病率。黄酮类化合物已被我国卫生部批准为新型抗氧化剂。在自然界中,黄酮类化合物大多以糖苷的形式存在,所以黄酮的糖基化修饰对其储存与运输有重要意义。
糖基转移酶是专门负责催化糖基化修饰反应的酶类,它将活性糖基从供体(通常是UDP-glucose)转移到受体分子上。糖基化修饰往往会改变植物分子的生物活性、水溶性、在细胞内和整体植株中的转运特性、亚细胞定位以及与受体的相互识别与结合特性,另外还能降低或消除内源和外源物质的毒性(Lim and Bowles,2004;Bowles et al.,2006;Wang and Hou,2009)。因此,糖基转移酶基因在调节植物细胞代谢平衡、维持植物正常生长发育等方面有重要意义。例如,已有报道糖基转移酶基因参与植物激素平衡调节、植物防御反应、植物次生代谢物合成以及植物信号转导等(Wang and Hou,2009)。但是有关糖基转移酶在提高植物黄酮含量方面的应用却未见报道。
玉米是世界上种植最广的农作物之一,在作为主要的粮食作物的同时,其茎杆也可被加工成饲料饲喂牲畜。如果通过基因工程技术提高玉米中黄酮的含量,将对于提高玉米的营养品质有十分重要的意义。玉米UFGT2是糖基转移酶家族1中的一个成员,目前它的基因序列已公开于核酸序列数据库中,但是经过检索,有关玉米糖基转移酶基因UFGT2在调控黄酮的代谢途径、提高玉米等植物中的黄酮含量方面的应用尚未见报道。
发明内容
针对现有技术的不足,本发明的目的是提供一种玉米糖基转移酶基因UFGT2在提高植物体内黄酮含量中的应用。
本发明所述玉米糖基转移酶基因UFGT2在提高植物体内黄酮含量中的应用。
其中:所述玉米糖基转移酶基因UFGT2的核苷酸序列如SEQ ID No.1所示,所述玉米糖基转移酶UFGT2的氨基酸序列如SEQ ID No.2所示。所述植物优选是单子叶植物或双子叶植物;所述单子叶植物为粮食作物,优选是玉米、水稻或小麦,所述双子叶植物是蔬菜类植物或中药植物。
本发明所述玉米糖基转移酶基因UFGT2是利用SEQ ID No.3和SEQ ID No.4所示的引物序列,通过RT-PCR技术从玉米中克隆得到。应用该基因构建植物过表达载体,进行转基因操作,可获得转基因植物。通过检测表明:获得的转基因植物中黄酮含量得到显著提高。
本发明的有益效果是:首次证明玉米糖基转移酶基因UFGT2能够提高植物体内的黄酮含量。实验证实,应用本发明所述玉米糖基转移酶基因UFGT2进行植物转基因操作,可以显著提高转基因植物体内的黄酮含量(见附图1、附图2、附图3和附图4)。预示本发明实施后有望能显著提升作物品质,可用于后续的作物品种改良,对我国农业生产具有重大意义。
附图说明
图1.UFGT2体外糖基化黄酮化合物的酶促反应。将UFGT2cDNA克隆至PGEX原核表达载体中,转化XL1-blue大肠杆菌,纯化蛋白,进行体外酶促反应。实验结果显示,UFGT2对黄酮类化合物槲皮素、山奈酚有很高的催化活性,能够催化形成黄酮的糖苷。图中UGT89B1作为酶促反应的阳性对照,因为UGT89B1已被证明能够糖基化修饰槲皮素和山奈酚(文献:Limet al.,2004,Biotechnology and Bioengineering)。
图2.玉米ufgt2突变体中黄酮积累量降低。申请人从美国UniformMu突变体库购买了两个UFGT2敲除突变体Mu689和Mu943,分别提取WT和突变体玉米的幼苗、种子中的总黄酮,发现ufgt2突变体中两种主要的黄酮醇(槲皮素、山奈酚)含量是降低的。说明UFGT2基因与黄酮含量有关。
图3.玉米UFGT2基因在转基因拟南芥不同株系中的表达水平。WT为野生型拟南芥对照,OE1,OE2,OE3为转入玉米UFGT2的转基因拟南芥。
图4.UFGT2转基因拟南芥中黄酮积累量升高。其中WT为对照,OE1、OE2和OE3为三个过表达株系。将拟南芥培养至5日龄,收集幼苗,提取总黄酮,用HPLC检测,发现过表达UFGT2的拟南芥中黄酮含量比WT显著增高。并且发现UFGT2在OE2、OE3中表达水平比OE1的高,黄酮含量也是在OE2、OE3中比OE1更高。实验结果表明UFGT2可以提高植物中的黄酮含量。
具体实施方式
实施例1玉米糖基转移酶基因UFGT2的克隆
通过公开网站http://plants.ensembl.org/Zea_mays获得玉米UFGT2基因的cDNA序列。
根据cDNA序列设计引物,正向引物为UFGT2-F:5’-GGATCCATGGCCGAAGGGGACGGTCCA-3’,反向引物为UFGT2-R:5GTCGACTCAAAGAGAAAGTATGTGATCCACC-3’。
利用TRIzol试剂盒提取拟南芥RNA,RT-PCR方法扩增UFGT2基因的全长cDNA序列。将cDNA克隆的过程是先经过BamHI和SacⅠ酶切,之后连入相应酶切的pBluescript II SK(+)克隆载体中,记为pK-UFGT2,通过PCR扩增和BamHI和SacⅠ酶切验证,最后进行序列测定,验证克隆序列的正确性。
玉米糖基转移酶基因UFGT2的序列信息与特性分析:UFGT2基因的编码区cDNA为1410bp(如SEQ ID No.1所示),编码469个氨基酸的蛋白(如SEQ ID No.2所示),C端具有44个氨基酸的PSPG盒,为植物次生代谢物糖基转移酶所共同具有的保守序列。
实施例2玉米糖基转移酶基因UFGT2转入拟南芥获得转基因植物
1.含有UFGT2编码区cDNA表达载体的构建
含有UFGT2编码框的克隆载体pK-UFGT2经过BamHI和SacⅠ双酶切后,获得带有粘性酶切位点末端的全长cDNA序列,将此基因片段与用BamHI和SacI双酶切后的pBI121载体部分相连,得到以CaMV 35S启动子驱动糖基转移酶基因过表达的植物表达载体,称为pBI-UFGT2。
2.农杆菌介导植物转化
农杆菌GV3101具有侵染植物和转移基因的能力,故将构建的pBI-UFGT2植物表达载体转入农杆菌,然后进行PCR验证验证。利用浸花法(一种公开的通用方法),使含有植物表达载体的农杆菌GV3101浸染拟南芥花蕾。待其长出的角果成熟之后,收集T1代种子并在筛选培养基上进行筛选,将能够正常生长的绿色转化苗移栽至营养土中培养,分别收获其T2代种子再进行下一轮筛选,挑选出绿苗:白苗为3:1的培养皿。将此培养皿上的绿苗移栽,单株收获种子(T3代)。对每一单株的种子部分用于筛选,直到选出在筛选培养基上为全绿的株系,即为纯合转基因株系。
3.过表达转基因植株分子鉴定
对上述拟南芥转基因植株进行基因表达水平的检测。分别提取转基因植株和野生型植株的RNA,进行qRT-PCR扩增,分析过表达植株和野生型植株的基因表达差异。玉米UFGT2在拟南芥过表达植株OE1、OE2、OE3中的表达量都明显高于野生型植株(理论上野生型没有UFGT2的表达)。
利用三个转基因拟南芥株系,来进行后续的分析工作。
实施例3玉米突变体和转基因拟南芥中黄酮含量的分析测定
玉米UFGT2的突变体从美国UniformMu突变体库购买,两个突变株系的编号为Mu689和Mu943。转基因拟南芥为实施例2制备的过表达株系OE1、OE2、OE3。利用这些材料分析玉米UFGT2基因对植物体内黄酮含量的影响。
黄酮的提取方法:主要针对槲皮素和山奈酚两类黄酮醇的含量进行了测定。基本按照参考文献(Ruohe et al.,2012)进行。
(1)称取0.1g样品放入2ml的Eppendorf管中,加入钢珠后经液氮速冻,在研磨仪上研磨1min至粉末;
(2)加入1ml的甲醇充分溶解,在4℃温和旋转提取1h;
(3)4℃,12,000rpm离心10min;
(4)将上清转至新的离心管中,加入1/3体积的蒸馏水,4℃,12,000rpm离心10min,去除叶绿素和油脂;
(5)取100ul上清,加入等体积2N HCl,70℃酸解40min,反应结束后加入100ul100%甲醇以防止苷元的沉降;
(6)4℃,12,000rpm离心15min;
(7)上清直接用于高效液相色谱分析。分析条件为:色谱仪是岛津LC-20AT(Shimadzu,日本)。色谱柱为Ultimate XB-C18(150mm×4.6mm,5μm)反相柱。每个物质峰的检测波长在190-430nm之间。流动相为乙腈和水(均含有0.1%甲酸),二元高压浓度梯度设置为10%-100%乙腈,流速为1ml/min,洗脱时间为35min,检测波长设置为270nm。
结果和结论:检测结果发现在玉米UFGT2的两个突变体中黄酮含量显著降低(附图2)。而在UFGT2的转基因拟南芥中,黄酮含量都明显提高。并且发现UFGT2的表达水平越高(OE2和OE3比OE1表达水平高),黄酮含量也越高(OE2、OE3比OE1黄酮含量更高)。
以上这些实验结果表明玉米的UFGT2可以影响植物体内的黄酮含量,通过转移和表达UFGT2基因可以实现在它种植物中提高黄酮的含量。
序列表
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<222>(1)…(31)
<400> 4
gtcgactcaa agagaaagta tgtgatccac c 31
Claims (3)
1.玉米糖基转移酶基因UFGT2在提高植物体内黄酮含量中的应用;
所述玉米糖基转移酶基因UFGT2的核苷酸序列如SEQ ID No.1所示,所述植物是单子叶植物或双子叶植物。
2.如权利要求1所述的应用,其特征在于:所述单子叶植物是粮食作物,所述双子叶植物是蔬菜类植物或中药植物。
3.如权利要求2所述的应用,其特征在于:所述粮食作物是玉米、水稻或小麦。
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NM_001153075.2;Schnable PS,等;《NCBI》;20170423;CDS序列部分 * |
The maize secondary metabolism glycosyltransferase UFGT2 modifies flavonols and contributes to plant acclimation to abiotic stresses;Li Yan-jie,等;《Annals of Botany》;20180704;第122卷;摘要 * |
糖基转移酶oled对于黄酮类化合物的底物特异性研究;xx1276243;《豆丁网》;20160617;第6页倒数第一到第二段 * |
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