金粳818抗咪唑啉酮类除草剂基因的功能标记开发与应用

doi:10.16819/j.1001-7216.2020.9134

中国水稻科学 ›› 2020, Vol. 34 ›› Issue (4): 316-324.DOI: 10.16819/j.1001-7216.2020.9134

金粳818抗咪唑啉酮类除草剂基因的功能标记开发与应用

王广达¹, 高鹏^1,², 杨文艳¹, 崔傲¹, 赵剑华¹, 冯志明^1,², 曹文磊^1,², 陈宗祥^1,², 左示敏^1,^2,^*()

¹扬州大学农学院江苏省作物遗传生理重点实验室/植物功能基因组学教育部重点实验室/江苏省作物基因组学和分子育种重点实验室,江苏扬州 225009
²扬州大学江苏省粮食作物现代产业技术协同创新中心,江苏扬州 225009

收稿日期:2019-12-13 修回日期:2020-02-19 出版日期:2020-07-10 发布日期:2020-07-10
通讯作者: 左示敏
基金资助:
江苏省农业重大新品种创制项目(PZCZ201703);江苏省大学生创新创业项目(201911117114Y);江苏省重点研发计划资助项目(现代农业, BE2018351-1);江苏高校优势学科建设工程资助项目

Development and Utilization of Functional Markers for Imidazolinone Herbicides Resistance Gene in japonica Rice Variety Jinjing 818

Guangda WANG¹, Peng GAO^1,², Wenyan YANG¹, Ao CUI¹, Jianhua ZHAO¹, Zhiming FENG^1,², Wenlei CAO^1,², Zongxiang CHEN^1,², Shimin ZUO^1,^2,^*()

¹Jiangsu Key Laboratory of Crop Genetics and Physiology / Key Laboratory of Plant Functional Genomics of Ministry of Education / Jiangsu Key Laboratory of Crop Genomics and Molecular Breeding, College of Agriculture, Yangzhou University, Yangzhou 225009, China
²Jiangsu Collaborative Innovation Center for Modern Industrial Technology of Grain Crops, Yangzhou University, Yangzhou 225009, China

Received:2019-12-13 Revised:2020-02-19 Online:2020-07-10 Published:2020-07-10
Contact: Shimin ZUO

摘要/Abstract

摘要：

【目的】培育抗除草剂品种是水稻育种中的一个重要方向,开发高效的功能标记可以加快抗除草剂性状的筛选工作。【方法】利用开发的dCAPS和KASP分子标记对试验材料的基因型分型,另结合除草剂喷施试验进行表型鉴定。【结果】测序结果验证了前人发现的金粳818中乙酰乳酸合成酶（acetolactate synthase,ALS）基因第627位氨基酸由丝氨酸（serine,S）突变为天冬酰胺（asparagine,N）是其抗咪唑啉酮类除草剂的主要原因,本文将其命名为ALS^627N。根据此变异位点的DNA序列差异,开发了两个新的功能标记dC-ALS-627和K-ALS-627,可有效区分ALS^627N纯合型、杂合型及非突变纯合型。【结论】两个标记均可以广泛用于ALS^627N基因的标记辅助选择育种,有助于加快培育抗咪唑啉酮类除草剂新品种,具有重要的实际应用价值。

关键词: 水稻, 咪唑啉酮类除草剂, 乙酰乳酸合成酶, 功能标记, 辅助选择

Abstract:

【Objective】Cultivating herbicide resistant rice varieties is an important direction in rice breeding. Developing efficient functional markers can accelerate the screening of herbicide resistance trait. 【Method】dCAPS and KASP techniques were used to analyze the genotypes of the experimental materials. In addition, the phenotypes were identified by herbicide spraying test. 【Result】Through sequence alignment analysis, we further confirmed that the mutation of 627 amino acid of acetolactate synthase (ALS) gene from serine (S) into asparagine (N) in Jinjing 818 (here after named ALS^627N) produces strong resistance to imidazolinone herbicides. According to the DNA sequence flanking the mutation site, we developed two novel functional markers, dC-ALS-627 and K-ALS-627, for detecting this resistant allele ALS^627N. The two markers can effectively distinguish homozygous ALS^627N type, heterozygous type and homozygous wild type. Combined with herbicide spraying test, we confirmed that ALS^627N is a dominant herbicide resistant gene. 【Conclusion】 We validated that the two markers have great potential in marker-assisted breeding for ALS^627N gene. Totally, this study will accelerate the development of new varieties with resistance to imidazolinone herbicides, and have significance in breeding practice.

Key words: rice, imidazolinone herbicide, acetolactate synthase, functional marker, assisted selection

中图分类号:

S511.032

王广达, 高鹏, 杨文艳, 崔傲, 赵剑华, 冯志明, 曹文磊, 陈宗祥, 左示敏. 金粳818抗咪唑啉酮类除草剂基因的功能标记开发与应用[J]. 中国水稻科学, 2020, 34(4): 316-324.

Guangda WANG, Peng GAO, Wenyan YANG, Ao CUI, Jianhua ZHAO, Zhiming FENG, Wenlei CAO, Zongxiang CHEN, Shimin ZUO. Development and Utilization of Functional Markers for Imidazolinone Herbicides Resistance Gene in japonica Rice Variety Jinjing 818[J]. Chinese Journal OF Rice Science, 2020, 34(4): 316-324.

图/表 7

表1 本研究涉及的引物序列

Table 1 Primer sequences involved in this study.

标记名称 Marker name	引物名称 Primer name	引物序列(5＇-3＇) Primer sequence (5＇-3＇)
dC-ALS-627	ALS-F ALS-R dC-ALS-627-F	CTCGCCCAAACCCAGAAACC CACATACAAACATCATAGGCATACCACTC GAGCATGTGCTGCCTATGATCCTAA
dC-ALS-627	dC-ALS-627-R	TAGAGCACATACAAACATCAT
K-ALS-627	K-ALS^627N-HEX	GAAGGTCGGAGTCAACGGATT ATCATGTCCTTGAATGCGCCCCCAT
	K-ALS^627S-FAM	GAAGGTGACCAAGTTCATGCT ATCATGTCCTTGAATGCGCCCCCAC
	K-ALS^627S-N	TGTTGGATATCATCGTCCCGCAC

图1 ALS基因序列比对结果

Fig. 1. ALS gene sequence alignment results.

图2 两种标记的相关信息灰色背景部分为非编码区序列;方框内连续的3个碱基为ALS第627位氨基酸的密码子,金粳818为AAT,9311和日本晴为AGT,分别编码氨基酸N和S;限制性核酸内切酶DdeⅠ的识别基序为CTNAG（N为任意碱基）;在标记dC-ALS-627的引物dC-ALS-627-F中引入了突变碱基T（模板序列中的碱基为C,T和C均用下划线标注）,当9311或日本晴模板序列中的碱基C被替换成T时,即在扩增产物中存在一个DdeⅠ的识别基序CTAAG,金粳818中则没有;在K-ALS-627标记中,引物K-ALS627N- HEX 的3＇端末碱基为T,5＇端连上HEX荧光标签序列（斜体部分）,引物K-ALS627S- FAM的3＇端末碱基为C,5＇端连上FAM荧光标签序列（斜体部分）;引物前端的箭头表示扩增的方向。

Fig. 2. Information of the two molecular markers. The gray background part is the non-coding region sequence. The three consecutive bases in the box are the codons of amino acids at position 627 of ALS, ‘AAT’ in Jinjing 818 and ‘AGT’ in 9311 and Nipponbare, encoding amino acid ‘N’ and ‘S’ respectively. The recognition sequence of restriction endonuclease DdeⅠ is ‘CTNAG’ (‘N’ for any base). The mutant base ‘T’ was introduced into primer dC-ALS-627-F of the marker dC-ALS-627 (The base in template sequenceis ‘C’, ‘T’ and ‘C’ are highlighted with an underline). When the base ‘C’ in the template sequence of 9311 or Nipponbare is replaced with ‘T’, there is a DdeⅠ recognition sequence ‘CTAAG’ in the amplification product, which doesn’t exist in Jinjing 818. In the marker K-ALS-627, primer K-ALS627N- HEX has a base ‘T’ at the 3＇end, and a HEX fluorescent label sequence (in italics) at the 5＇end, and primer K-ALS627S- FAM has a base ‘C’ at the 3＇end, and a FAM fluorescent label sequence (in italics) at the 5' end. The arrow at the front of primers indicates the direction of amplification.

图3 三种分子标记检测结果 A图为dCAPS标记dC-ALS-627检测结果;B图为KASP标记K-ALS-627检测结果;C图为SSR标记RM7413检测结果。M1为20 bp DNA Ladder;M2为DL500 DNA标记;P1为金粳818,P2为南粳505,P3为华粳0029;1~3为南粳505/金粳818的杂交种,4~6为华粳0029/金粳818的杂交种。菱形代表无DNA模板的空白对照,正方形代表金粳818纯合基因型（变异位点碱基为A）,圆圈代表南粳505和华粳0029纯合基因型（变异位点碱基为G）,三角代表ALS基因的杂合带型（变异位点碱基为G/A）。

Fig. 3. Detection results of three molecular markers. A, Detection results of dC-ALS-627; B, Detection results of K-ALS-627; C, Detection results of RM7413. M1, 20 bp DNA Ladder, M2, DL500 DNA Marker; P1, Jinjing 818, P2, Nanjing 505, P3, Huajing 0029; 1-3, Hybrid of Nanjing 505 / Jinjing 818; 4-6, Hybrid of Huajing 0029 / Jinjing 818. The diamond represents a blank control without the DNA template; the square represents the homozygous genotype of Jinjing 818 (variation site base A); the circle represents the homozygous genotypes of Nanjing 505 and Huajing 0029 (variation site base G); and the triangle represents the heterozygous type of ALS gene (with a G/A mutation site base).

图4 dC-ALS-627和K-ALS-627标记对BC2F2水稻植株的基因型检测结果 A图为dC-ALS-627标记检测结果。M为DL1000 DNA标记;P1为金粳818,P2为南粳505,1~10为南粳505/金粳818的BC2F2部分单株带型。B图为K-ALS-627标记检测结果。黑色方框代表无DNA模板的空白对照,蓝色方框代表金粳818纯合基因型（变异位点碱基为A）,黄色圆圈代表南粳505纯合基因型（变异位点碱基为G）,绿色三角代表ALS基因的杂合带型（变异位点碱基为G/A）。

Fig. 4. Genotyping of BC2F2 generation rice plants using dC-ALS-627 and K-ALS-627 markers. A, Detection results of dC-ALS-627. M, DL1000 DNA marker; P1, Jinjing 818; P2, Nanjing 505; 1-10, Partial of BC2F2 individuals of Nanjing 505 / Jinjing 818.B, Detection results of K-ALS-627. The black box represents a blank control without the DNA template; the blue box represents the homozygous genotype of Jinjing 818 (variation site base A); the yellow circle represents the homozygous genotypes of Nanjing 505 and Huajing 0029 (variation site base G); and the green triangle represents the heterozygous type of ALS gene (with a G/A mutation site base).

图5 咪唑乙烟酸处理后的BC2F2水稻植株表型 P1为金粳818,P2为南粳505,1~10为南粳505/金粳818的部分BC2F2单株。

Fig. 5. Phenotypes of BC2F2 populations after treatment with imazethapyr. P1, Jinjing 818, P2, Nanjing 505. 1-10, Partial BC2F2 plants of Nanjing 505/ Jinjing 818.

图6 利用dC-ALS-627和K-ALS-627标记筛选抗除草剂水稻资源 A图为dC-ALS-627标记检测结果。M为BM2000 DNA标记,J为金粳818,1~40分别为感咪唑啉酮类除草剂的常规粳稻禾粳1819、沭粳1901、华粳1803、南粳46、南粳52、武运粳23、镇稻11、镇稻14、镇稻18、淮稻5号、扬粳103、扬育粳2号、扬育粳3号、华粳8号、淮119、武运粳27号、武运粳32、武运粳80、扬农稻1号、盐粳11、盐粳13、盐粳15、盐粳16、盐粳6号、镇稻99、大粮202、泗稻12、泗稻15、新科稻21、徐稻5号、原旱稻3号、金粳18、金粳787、连粳10、连粳4号、苏垦118、苏秀867、扬粳818、武粳66、宁5718。B图为K-ALS-627标记检测结果。黑色方框代表无DNA模板的空白对照,蓝色方框代表金粳818纯合基因型（变异位点碱基为A）,黄色圆圈代表感咪唑啉酮类除草剂品种的纯合基因型（变异位点碱基为G）。

Fig. 6. Screening herbicide resistant rice resources using dC-ALS-627 and K-ALS-627 markers. A, Detection results of dC-ALS-627. M, BM2000 DNA marker; J refers to Jinjing 818 and 1-40 refer to rice varieties sensitive to imidazolinone herbicides: Hejing 1819, Shujing 1901, Huajing 1803, Nanjing 46, Nanjing 52, Wuyunjing 23, Zhendao 11, Zhendao 14, Zhendao 14, Zhendao 18, Huaidao 5, Yangjing 103, Yangyujing 2, Yangyujing 3, Huajing 8, Huai 119, Wuyunjing 27, Wuyunjing 32, Wuyunjing 80, Yangnongdao 1, Yanjing 11, Yanjing 13, Yanjing 15, Yanjing 16, Yanjing 6, Zhendao 99, Daliang 202, Sidao 12, Sidao 15, Xinkedao 21, Xudao 5, Yuanhandao 3, Jinjing 18, Jinjing 787, Lianjing 10, Lianjing 4, Suken 118, Suxiu 867, Yangjing 818, Wujing 66, Ning 5718. B, Detection results of K-ALS-627. The black box represents a blank control without the DNA template; the blue box represents the homozygous genotype of Jinjing 818 (variation site base A); the yellow circle represents the homozygous genotypes of Nanjing 505 and Huajing 0029 (variation site base G).

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金粳818抗咪唑啉酮类除草剂基因的功能标记开发与应用

Development and Utilization of Functional Markers for Imidazolinone Herbicides Resistance Gene in japonica Rice Variety Jinjing 818

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