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作物学报

作物学报 ›› 2024, Vol. 50 ›› Issue (10): 2435-2446.doi: 10.3724/SP.J.1006.2024.32060

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

利用基因编辑技术创制低谷蛋白水稻种质

周田田1,2(), 唐兆成1, 李笑1, 朱鹏1, 邓晶晶1, 杨郁文1, 张保龙1,2, 郭冬姝1,*()   

  1. 1江苏省农业科学院种质资源与生物技术研究所，江苏南京 210014
    2南京农业大学生命科学学院，江苏南京 210014
  • 收稿日期:2023-12-22 接受日期:2024-05-21 出版日期:2024-10-12 网络出版日期:2024-06-21
  • 通讯作者: *郭冬姝, E-mail: guods_cau@163.com
  • 作者简介:E-mail: 2021116027@stu.njau.edu.cn
  • 基金资助:
    江苏省重点研发计划现代农业项目(BE2022365);江苏省重点研发计划现代农业项目(cx(21)1002)

Development of low-glutelin rice germplasm by gene editing technology

ZHOU Tian-Tian1,2(), TANG Zhao-Cheng1, LI Xiao1, ZHU Peng1, DENG Jing-Jing1, YANG Yu-Wen1, ZHANG Bao-Long1,2, GUO Dong-Shu1,*()   

  1. 1Institute of Germplasm Resources and Biotechnology, Jiangsu Academy of Agricultural Sciences, Nanjing 210014, Jiangsu, China
    2College of Life Sciences, Nanjing Agricultural University, Nanjing 210014, Jiangsu, China
  • Received:2023-12-22 Accepted:2024-05-21 Published:2024-10-12 Published online:2024-06-21
  • Contact: E-mail: guods_cau@163.com
  • Supported by:
    Jiangsu Key Research and Development Program(BE2022365);Jiangsu Agricultural Science and Technology Innovation Fund (cx(cx(21)1002)

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摘要：

水稻(Oryza sativa L.)是重要的粮食作物，蛋白质是稻米的第二大营养物质。肾脏病人通常需要限制蛋白质的摄入量以减轻肾脏的负担和控制病情进展。稻米蛋白中谷蛋白含量最高，且易被人体消化。本研究以LGC-1作为供体亲本培育出的粳稻品种为转基因受体，利用CRISPR/Cas9介导的基因编辑技术一次性敲除谷蛋白A (Glutelin A，GluA)亚家族编码基因GluA1、GluA2和GluA3，获得了不含转基因元件的谷蛋白含量约为1.8%的低谷蛋白种质，并对所得种质的稻米品质和农艺性状进行了全面评估。所得低谷蛋白种质外观品质较好，相比于受体品种垩白度降低、糙米率和精米率提高。本研究为创制低谷蛋白水稻品种提供了一种高效、简便的方法，为培育适合肾病患者的功能性水稻品种提供了一种新的遗传材料。

关键词: 水稻, 基因编辑, 蛋白, 低谷蛋白水稻

Abstract:

Rice (Oryza sativa L.) is a crucial cereal crop worldwide, with protein being its second-most significant nutritional component. Patients with kidney diseases are required to limit their protein intake to alleviate the metabolic burden on their kidneys and control disease progression. In regular rice cultivars, glutelin is the predominant protein component and is easily digested by the human body. In this study, simultaneous mutations were introduced into Glutelin A1 (GluA1), GluA2, and GluA3 using CRISPR/Cas9-mediated targeted mutagenesis in a japonica rice cultivar derived from Low Glutelin Content-1 (LGC-1). Consequently, a low-glutelin rice germplasm with approximately 1.8% glutelin content, free from transgenic elements, was generated. The quality and agronomic traits of this germplasm were further comprehensively evaluated. The low-glutelin germplasms generated in this study exhibited significantly lower chalkiness degree compared to the recipient cultivar, while the brown rice rate and milled rice rate were significantly higher. This study presents a highly efficient and convenient method for generating rice germplasm with reduced glutelin content and offers new genetic materials for the cultivation of functional rice cultivars suitable for patients with kidney diseases.

Key words: rice, gene editing, protein, low-glutelin rice

表1

载体构建及基因型鉴定相关引物"

引物名称
Primer name		 引物序列
Primer sequence (5'-3')		 功能
Function
GluA1/2-1F GGCAtaggccagagcactagtcaa 谷蛋白A敲除载体构建Glutelin A CRISPR vector construction
GluA1/2-1R AAACttgactagtgctctggccta 谷蛋白A敲除载体构建Glutelin A CRISPR vector construction
GluA1/2-2F TGTGcggagtgtgaggtctcaagc 谷蛋白A敲除载体构建Glutelin A CRISPR vector construction
GluA1/2-2R AAACgcttgagacctcacactccg 谷蛋白A敲除载体构建Glutelin A CRISPR vector construction
GluA3-1F GGCAcaatggcaaagttctcgccg 谷蛋白A敲除载体构建Glutelin A CRISPR vector construction
GluA3-1R AAACcggcgagaactttgccattg 谷蛋白A敲除载体构建Glutelin A CRISPR vector construction
GluA3-2F TGTGctagtgagtcacttcatatg 谷蛋白A敲除载体构建Glutelin A CRISPR vector construction
GluA3-2R AAACcatatgaagtgactcactag 谷蛋白A敲除载体构建Glutelin A CRISPR vector construction
LGC-F1 ACCACAAGACAACACTGTCACACC LGC-1突变体鉴定正向引物
Genotyping forward primer of Lgc1 locus
LGC-R1 TATGGTCGCTCAATCGAGCAACAC LGC-1突变体鉴定反向引物
Genotyping reverse primer of Lgc1 locus
WT-R TAGTGTTTCCTAGGAGTGGG 野生型位点鉴定反向引物(正向引物LGC-F1)
Genotyping reverse primer of wild-type LGC1 locus (the forward primer is LGC-F1)
GluA1-F CATGATCACCAACAACCATG GluA1基因型鉴定正向引物
Genotyping forward primer of GluA1 gene
GluA1/2-R CTAGAGATGCACCATTAGTG GluA1和GluA2基因型鉴定反向引物
Genotyping reverse primer of GluA1 and GluA2 gene
GluA2-F CAAAAAGAGGAGGGCTTTAC GluA2基因型鉴定正向引物
Genotyping forward primer of GluA2 gene
GluA3-F CACTAAAGCAACACACAACG GluA3基因型鉴定正向引物
Genotyping forward primer of GluA3 gene
GluA3-R CAAGTGTGGATTGCCATGTT GluA3基因型鉴定反向引物
Genotyping reverse primer of GluA1 gene
35S-TF1 ACAATCCCACTATCCTTCGCAAG 潮霉素基因表达盒检测 hpt cassette detection
HYG-TR GTACTTCTACACAGCCATCGGTC 潮霉素基因表达盒检测 hpt cassette detection
UBI-F1 TTTCCCCAACCTCGTGTTGTTC Cas9基因表达盒检测 Cas9 cassette detection
Cas9-R2 GAGGTTCTTCTTGATGGAGTGG Cas9基因表达盒检测 Cas9 cassette detection

图1

谷蛋白编码基因靶标位点及载体T-DNA区段示意图 A: GluA1和GluA2基因结构，黄色和桔黄色方框分别代表GluA1和GluA2基因的外显子，外显子间的黑色短线代表内含子，蓝色和绿色线显示基因编辑靶标位点的位置，带下画线的红色字母显示基因编辑靶标位点序列，带下画线的黑色字母显示PAM (protospacer adjacent motif)序列; B: GluA3基因结构，浅蓝色方框代表GluA3基因的外显子，外显子间的黑色短线代表内含子，枚红色短线和粉色线显示基因编辑靶标位点的位置，带下划线的红色字母显示基因编辑靶标位点序列，带下画线的黑色字母显示PAM序列; C: 基因编辑载体T-DNA区段示意图。ZmpUBI: 玉米Ubiquitin1基因启动子; SpCas9: 针对水稻进行密码子优化后的来源于酿脓链球菌的Cas9基因; NOST: NOS终止子; 3: 水稻U3b启动子; 6: 水稻U6b启动子; GluA1/2-1: GluA1和GluA2基因的第1个靶标位点序列; GluA1/2-2: GluA1和GluA2基因的第2个靶标位点序列; GluA3-1: GluA3基因的第一个靶标位点序列; GluA3-2: GluA3基因的第2个靶标位点序列; 35S Promoter: 花椰菜花叶病毒(Cauliflower mosaic virus) 35S启动子; HygR: 潮霉素抗性基因; 35ST: 35S终止子; LB: T-DNA左边界; RB: T-DNA右边界。"

图2

基因型鉴定及Sanger测序峰图 A: 琼脂糖凝胶电泳鉴定基因型; B: Sanger测序峰图; C: 转基因元件检测。Marker: DNA分子量标准; WT: 野生型粳稻苏秀867; LGC-1: 受体品种; +CK: 基因编辑质粒阳性对照; -CK: PCR阴性对照; LGC1: 野生型LGC1位点; Lgc1: 突变型Lgc1位点; GluA1: GluA1基因位点; GluA2: GluA2基因位点; GluA3: GluA3基因位点; Cas9: Cas9基因表达盒; Hyg: 潮霉素抗性基因表达盒。A中GluA1、GluA2和GluA3基因的较长条带代表野生型基因型，较短条带代表突变型基因型; B中“*”和“·”分别标注基因序列缺失区段两侧的碱基; A和C中左侧箭头标注分子量标准对应的条带大小。"

图3

水稻SDS-PAGE和氨基酸组成测定 A: 糙米蛋白聚丙烯酰胺凝胶电泳; B: 精米蛋白聚丙烯酰胺凝胶电泳; C: 稻米氨基酸组成测定结果。M: 蛋白marker; WT: 野生型粳稻苏秀867; LGC-1: 受体品种; GluA-2-11和GluA-3-6: 突变体; A和B右侧竖线表示不同贮藏蛋白组分; C中Asp~Lys代表不同氨基酸，不同字母表示P < 0.01水平差异显著。"

图4

稻米主要营养成分含量检测结果 A: 糙米总蛋白含量; B: 精米总蛋白含量; C: 糙米总淀粉含量; D: 糙米总脂肪酸含量。A~D: 每个样品重复测定3次, 不同小写字母代表显著性差异(P < 0.01); E~G: LGC-1、GluA-2-11和GluA-3-6成熟种子横截面扫描电镜观察结果, 标尺为10 μm; H~J: LGC-1、GluA-2-11和GluA-3-6成熟种子总脂肪酸检测的总离子流图。"

表2

外观品质统计结果"

指标Index LGC-1 GluA-2-11 GluA-3-6
长/宽均值 Average of length/width 1.83 ± 0.02 1.88 ± 0.01 1.84 ± 0.01
糙米率 Brown rice yield 76.19 ± 0.68 a 83.22 ± 0.07 b 82.73 ± 0.39 b
精米率 Milled rice yield 68.33 ± 0.58 a 72.91 ± 0.93 b 72.83 ± 0.82 b
整精米率 Head rice yield 61.09 ± 4.34 62.54 ± 2.17 66.02 ± 2.57
垩白粒率 Chalky kernel percentage 7.52 ± 1.24 7.28 ± 0.78 6.45 ± 0.16
垩白度 Chalkiness 2.75 ± 0.90 A 1.23 ± 0.31 B 1.23 ± 0.17 B
透明度 Transparency 5.00 ± 0.00 5.00 ± 0.00 5.00 ± 0.00

表3

食味品质统计结果"

指标 Index LGC-1 GluA-2-11 GluA-3-6
外观 Appearance 4.10 ± 0.35 4.80 ± 0.17 5.23 ± 0.44
硬度 Hardness 8.07 ± 0.26 7.77 ± 0.09 7.50 ± 0.10
黏度 Viscosity 4.67 ± 0.26 5.40 ± 0.12 5.70 ± 0.59
平衡度 Balance degree 3.93 ± 0.33 4.67 ± 0.15 5.03 ± 0.44
食味值 Comprehensive 53.4 ± 2.10 58.03 ± 0.91 60.50 ± 2.82

图5

水稻株型和稻米外观表型 A: LGC-1、GluA-2-11和GluA-3-6成株外观; B~D: LGC-1、GluA-2-11和GluA-3-6种子外观; E~G: LGC-1、GluA-2-11和GluA-3-6精米外观。A中标尺为20 cm; B~G中标尺为1 cm。"

表4

农艺性状统计结果"

指标 Index LGC-1 GluA-2-11 GluA-3-6
株高 Plant height (cm) 86.16 ± 0.70 ab 87.74 ± 0.71 a 84.72 ± 0.72 b
分蘖数 Tiller number 14.80 ± 0.51 15.90 ± 0.60 16.20 ± 0.70
每穗粒数 Number of spikelets per panicle 105.00 ± 1.73 102.00 ± 1.53 104.33 ± 0.88
千粒重 1000-grain weight (g) 21.34 ± 0.22 a 20.55 ± 0.11 b 20.46 ± 0.03 b
结实率 Filled grain percentage (%) 89.68 ± 0.92 88.08 ± 0.18 88.90 ± 0.43
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