中国农业科学 ›› 2019, Vol. 52 ›› Issue (14): 2436-2449.doi: 10.3864/j.issn.0578-1752.2019.14.005
收稿日期:
2019-03-22
接受日期:
2019-04-24
出版日期:
2019-07-16
发布日期:
2019-07-26
通讯作者:
王跃进
作者简介:
刘梦琦,E-mail: mengqil@163.com。
基金资助:
LIU MengQi,WU FengYing,WANG YueJin()
Received:
2019-03-22
Accepted:
2019-04-24
Online:
2019-07-16
Published:
2019-07-26
Contact:
YueJin WANG
摘要:
【目的】葡萄是世界性重要果树,欧洲葡萄品种因其优质高产被广泛栽培,但其突出缺点是抗病性弱,尤其易受白粉病危害。葡萄中芪合成酶(stilbene synthase,STS)的代谢产物白藜芦醇是植物体内具有抗病功能的植保素,果实中的白藜芦醇对人具有保健作用。中国野生毛葡萄‘丹凤-2’抗病性强且白藜芦醇含量高。论文旨在研究中国野生毛葡萄‘丹凤-2’芪合成酶基因(STS)及其功能,应用于抗病育种来提高欧洲葡萄抗病性及果实中芪类物质含量。【方法】同源克隆中国野生毛葡萄‘丹凤-2’芪合成酶基因VqSTS26和VqSTS32,构建pCAMBIA35S::VqSTSs::GFP过表达载体;以器官发生途径诱导的无核白分生愈伤组织作为受体材料,采用农杆菌介导法进行遗传转化,获得转基因葡萄植株;分别从转录水平和代谢产物水平比较转基因与野生型无核白在自然生长条件与人工接种葡萄白粉病菌(Uncinula necator)诱导下STS表达及芪类物质产生与积累的差异;通过显微观察白粉病菌在转基因与野生型无核白叶片上的生长发育进程,统计孢子萌发、菌丝生长与分生孢子梗形成数目,对转基因植株进行抗病性分析。【结果】通过PCR检测和Western blot鉴定,获得了稳定转化VqSTS26植株8株和稳定转化VqSTS32植株5株。在自然生长条件下实时荧光定量PCR分析表明VqSTS26、VqSTS32转基因无核白STS的表达量显著提高,芪合成酶上游基因PAL与下游基因RSGT的表达量上调,而与芪合成酶存在底物竞争关系的CHS表达下调;液相色谱分析表明芪类物质主要以糖苷的反式云杉新苷形式存在,VqSTS26、VqSTS32转基因无核白芪类物质的含量极显著高于野生型无核白。STS的表达及其产物合成受白粉病菌诱导,随白粉病菌诱导,STS表达量在1—2 dpi时显著升高,至7 dpi时表达量达最高;诱导表达产生的芪类物质由原来的反式云杉新苷新增加了反式白藜芦醇和葡萄素,且含量增加;转基因植株在STS表达量、芪类物质的积累方面均极显著高于野生型无核白。显微观察白粉病菌在葡萄叶片上的生长发育状态,对比野生型无核白,转基因植株白粉病菌生长受到抑制,菌丝发育更迟,7 dpi时转基因葡萄叶片上的分生孢子梗数量低于野生型无核白。【结论】过表达中国野生毛葡萄‘丹凤-2’VqSTS26和VqSTS32可以提高无核白中STS的表达量,促进芪类物质的形成与积累,抑制转基因无核白叶片上白粉病菌的生长。因此,中国野生毛葡萄‘丹凤-2’与其携带的STS及其产物,是定向改良欧洲葡萄品种白粉病抗性与芪类物质含量的重要种质资源与基因资源。
刘梦琦,吴凤颖,王跃进. 中国野生毛葡萄芪合成酶基因表达与抗白粉病分析[J]. 中国农业科学, 2019, 52(14): 2436-2449.
LIU MengQi,WU FengYing,WANG YueJin. Expression of Stilbene Synthase Gene and Resistance to Powdery Mildew Analysis of Chinese Wild Vitis quinquangularis[J]. Scientia Agricultura Sinica, 2019, 52(14): 2436-2449.
表1
本研究所用引物"
基因 Gene | 引物序列 Primer sequence | 目标大小 Target size (bp) |
---|---|---|
VqSTS26 (Vector construction, PCR detection) | F: AGAACACGGGGGACGAGCTCATGGCTTCAGTTGAGGAATTTAGAAACG R: ACCATGGTGTCGACTCTAGAATTTGTAACTGTAGGAATGCTATGCAGC | 1179 |
VqSTS32 (Vector construction, PCR detection) | F: AGAACACGGGGGACGAGCTCATGGCTTCAATTGAGGAAATTAGAAACGC R: ACCATGGTGTCGACTCTAGAATTTGTAACCATAGGAACGCTATGCAG | 1179 |
图1
‘丹凤-2’VqSTS26、VqSTS32的定位、序列比对及聚类分析 A:VqSTS26、VqSTS32的染色体定位分析Chromosome localization analysis of VqSTS26, VqSTS32;B:VqSTS26、VqSTS32与欧洲葡萄同源蛋白氨基酸序列比对;阴影部分为差异位点,方框部分为保守序列Amino acid sequence alignment of homologous proteins between VqSTS26, VqSTS32 and STS genes from V. vinifera, the shading is the difference site and the box part is the conservative sequence;C:VqSTS26、VqSTS32与欧洲葡萄、高粱、花生、白松、樟子松和松叶兰芪合成酶基因氨基酸序列聚类分析,VqSTS26(AFM56643.1)、VqSTS32(AFM56649.1)、VvSTS15(XP_002268756.1)、VvSTS48(NP_001267934.1)、SbSTS(AAL49965)、AhSTS(BAA78617)、PstrSTS(CAA87012)、PsylSTS(CAA43165)、PnSTS(BAA87924)Cluster analysis of STS amino acid sequence from VqSTS26, VqSTS32 and V. vinifera, Sorghum bicolor, Arachis hypogaea, Pinus strobus, Pinus sylvestris, Psilotum nudum"
图2
转VqSTS26、VqSTS32无核白的遗传转化过程与转基因植株的鉴定 A:基因克隆与过表达载体构建Gene cloning and vector construction。a:pCAMBIA35S:: VqSTSs:: GFP载体 pCAMBIA35S:: VqSTSs:: GFP vector;b:VqSTS26、VqSTS32目的基因扩增Gene amplification of VqSTS26 and VqSTS32;c:VqSTS26、VqSTS32基因连接表达载体pCAMBIA2300的双酶切检测,CK为pCAMBIA2300的空载体对照 Double enzyme digestion detection of recombinant plasmid, CK was the empty vector as control;d:VqSTS26、VqSTS32转化农杆菌GV3101的PCR检测,CK1、CK2分别为每个基因重组质粒的阳性对照 PCR detection of agrobacterium GV3101 transformed with VqSTS26 and VqSTS32. CK1 and CK2 were positive controls of recombinant plasmid for each gene。M:Trans 2k plus DNA maker。B:无核白分生愈伤组织的诱导(a—d)、遗传转化(e—h)与转化后的诱导成苗、扩繁与移栽炼苗(i—n) Induction (a-d) and genetic transformation (e-h) of Thompson Seedless meristem callus and induction into seedlings after transformation (i-n)。C:无核白葡萄转VqSTS26、VqSTS32抗性株系的PCR检测和Western blot检测PCR amplification and Western blot detection of Thompson Seedless transgenic lines with VqSTS26 and VqSTS32"
表2
葡萄叶片每100个孢子萌发数、初级菌丝数、次级菌丝数和分生孢子梗数统计"
株系 Line | 1 dpi | 2 dpi | 3 dpi | 7 dpi | |||||||
---|---|---|---|---|---|---|---|---|---|---|---|
萌发数 Germination | 初级菌 丝数 Primary hyphae | 次级菌 丝数 Secondary hyphae | 萌发数 Germination | 初级菌 丝数 Primary hyphae | 次级菌 丝数 Secondary hyphae | 萌发数 Germination | 初级菌 丝数 Primary hyphae | 次级菌丝数 Secondary hyphae | 分生孢子梗数 Conidiophore | 分生孢子 梗数 Conidiophore | |
野生型 Wide type | 42.67± 3.51a | 21.67± 2.08a | 4.33± 1.53a | 22.67± 2.08a | 8.67± 1.15c | 50.33± 3.51a | 6.33± 1.53c | 12.67± 1.53c | 69.00± 4.00a | 15.33±1.53a | 195.33± 12.66a |
OEVqSTS26-L6 | 32.33± 2.52bc | 7.33± 1.53b | 0.33± 0.58b | 16.33± 0.58b | 11.67± 1.53c | 14.33± 2.08c | 30.67± 1.53a | 13.33± 1.53c | 16.33± 2.52c | 0 | 92.67± 7.37b |
OEVqSTS26-L8 | 26.00± 1.00c | 12.00± 2.00b | 0.33± 0.58b | 17.33± 1.53b | 17.67± 2.52b | 10.33± 1.53c | 31.67± 3.51a | 11.00± 1.73c | 23.00± 2.65c | 0 | 42.00± 3.00c |
OEVqSTS32-L3 | 35.00± 1.53b | 17.67± 2.52a | 3.33± 1.53ab | 17.00± 1.00b | 25.00± 1.00a | 22.67± 1.53b | 13.33± 2.52b | 18.00± 1.00b | 34.33± 3.06b | 0 | 110.33± 14.29b |
OEVqSTS32-L4 | 35.33± 1.53b | 19.67± 1.53a | 3.67± 1.53a | 13.67± 1.53b | 21.00± 2.00ab | 26.33± 1.53b | 18.33± 1.53b | 23.00± 2.00a | 32.33± 2.08b | 0 | 58.33± 2.08c |
图4
人工接种白粉病菌后转基因植株与野生型无核白STS表达及芪类物质含量分析 A、B:qRT-PCR分析VqSTS26(A)和VqSTS32(B)转基因与野生型无核白接菌0—7 dpi叶片STS的表达量 qRT-PCR analysis of STS expression between transgenic grapes with overexpression VqSTS26 (A) and VqSTS32 (B) and wild-type Thompson Seedless under U. necator induction at 0-7 dpi;C、D:HPLC分析VqSTS26(C)和VqSTS32(D)转基因与野生型无核白接菌前后芪类物质含量的变化HPLC detection of stilbenoids content in transgenic grapes with overexpression VqSTS26 (C) and VqSTS32 (D) and wild-type Thompson Seedless under U. necator induction at 0 and 7 dpi. *P<0.05,**P<0.01"
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