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作物学报 ›› 2026, Vol. 52 ›› Issue (4): 1046-1056.doi: 10.3724/SP.J.1006.2026.52033

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

利用CRISPR/Cas9编辑Hd6基因创制优质早熟水稻新种质

陈伟1,2(), 卫万娟1, 赵其兵1, 常东伟1, 余凌波2, 翟鹏飞2, 冯志明2,4, 陈宗祥2,4, 任仰涛1, 杨鹏1, 刘海浪1, 李珍富1, 杨永乐1,2, 金彦刚1,*(), 左示敏2,3,4,*()   

  1. 1江苏瑞华农业科技有限公司, 江苏宿迁 223800
    2江苏省作物基因组学和分子育种重点实验室 / 生物育种钟山实验室 / 植物功能基因组学教育部重点实验室 / 扬州大学, 江苏扬州 225009
    3江苏省粮食作物现代产业技术协同创新中心 / 江苏省作物遗传生理重点实验室, 扬州大学, 江苏扬州 225009
    4扬州现代种子创新研究所, 江苏高邮 225600
  • 收稿日期:2025-09-11 接受日期:2026-01-22 出版日期:2026-04-12 网络出版日期:2026-02-05
  • 通讯作者: *左示敏, E-mail: smzuo@yzu.edu.cn; 金彦刚, E-mail: 17949721@qq.com
  • 作者简介:E-mail: 864579662@qq.com
  • 基金资助:
    江苏省种业振兴揭榜挂帅项目(JBGS [2021] 035);扬州市现代农业项目(YZ2024045)

Developing new germplasm of high-quality and early-maturing rice by editing Hd6 via CRISPR/Cas9

Chen Wei1,2(), Wei Wan-Juan1, Zhao Qi-Bing1, Chang Dong-Wei1, Yu Ling-Bo2, Zhai Peng-Fei2, Feng Zhi-Ming2,4, Chen Zong-Xiang2,4, Ren Yang-Tao1, Yang Peng1, Liu Hai-Lang1, Li Zhen-Fu1, Yang Yong-Le1,2, Jin Yan-Gang1,*(), Zuo Shi-Min2,3,4,*()   

  1. 1Jiangsu Ruihua Agricultural Technology Co. Ltd, Suqian 223800, Jiangsu, China
    2Jiangsu Key Laboratory of Crop Genomics and Molecular Breeding / Zhongshan Biological Breeding Laboratory / Key Laboratory of Plant Functional Genomics of the Ministry of Education / Yangzhou University, Yangzhou 225009, Jiangsu, China
    3Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops / Jiangsu Key Laboratory of Crop Genetics and Physiology, Yangzhou University, Yangzhou 225009, Jiangsu, China
    4Yangzhou Modern Seed Innovation Research Institute, Gaoyou 225600, Jiangsu, China
  • Received:2025-09-11 Accepted:2026-01-22 Published:2026-04-12 Published online:2026-02-05
  • Contact: *E-mail: smzuo@yzu.edu.cn; E-mail: 17949721@qq.com
  • Supported by:
    Jiangsu Provincial Seed Industry Revitalization Project(JBGS [2021] 035);Yangzhou Modern Agriculture Project(YZ2024045)

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摘要：

抽穗期是决定水稻品种季节适应性与区域适应性的关键农艺性状。Hd6编码CK2α亚基, 通过磷酸化成花调控因子来调控水稻抽穗期, 其功能缺失突变体在长日照条件下抽穗期缩短。利用CRISPR/Cas9基因编辑技术对优良食味晚粳品种南粳46 (NJ46)的Hd6基因进行敲除, 以缩短NJ46的生育期, 扩大其种植区域。利用CRISPR/Cas9基因编辑的原理, 在Hd6的第2外显子处设计靶点, 通过BLAST分析确定其特异性并构建pCAMBIA1305- Actin:Cas9-sgRNAHd6表达载体, 利用农杆菌介导法转化NJ46愈伤组织, 通过PCR技术筛选不含外源成分的纯合突变系, 分别在苏中和苏北区域测定各纯合突变株系的抽穗期、主要农艺性状、产量性状及品质性状, 并与相应生态区主栽品种南粳9108 (NJ9108)、南粳518 (NJ518)进行比较。在T0代获得了18株转基因阳性植株, 在T2代获得了3个无外源成分、不同突变类型的纯合突变株系: NJ46-hd6ko-1 (缺失TTGG)、NJ46-hd6ko-2 (缺失25个碱基)、NJ46-hd6ko-3 (缺失G), 其抽穗期分别比NJ46提早21.1、24.7和21.4 d。其中, NJ46-hd6ko-3的抽穗期、理论产量、精米率、直链淀粉含量、胶稠度、食味值均与NJ9108相近, 但株高和碱消值则分别低于和高于NJ9108, 综合性状更优, 适宜在苏中地区种植; NJ46-hd6ko-2的抽穗期、理论产量、碱消值等与NJ518均无显著差异, 但精米率和食味值均显著高于NJ518, 而且株高矮于NJ518, 综合性状更优, 适宜在苏北地区种植。定向编辑NJ46中的Hd6基因, 创制了分别适宜苏中和苏北生态区种植的优良食味粳稻新种质, 扩展了优良食味水稻品种的种植区域。

关键词: 水稻, CRISPR/Cas9, 基因编辑, 抽穗期, Hd6

Abstract:

Heading date is a key agronomic trait that determines the seasonal and regional adaptability of rice varieties. Hd6 encodes the CK2α subunit, which regulates heading date in rice by phosphorylating florigen regulators. Loss-of-function mutants of Hd6 exhibit earlier heading under long-day conditions. To shorten the growth duration of the elite, tasty, late-season japonica cultivar Nanjing 46 (NJ46) and expand its cultivation range, the Hd6 gene in NJ46 was knocked out using CRISPR/Cas9- mediated gene editing. A target site was designed within the second exon of Hd6, and its specificity was validated by BLAST analysis. The expression vector pCAMBIA1305-Actin:Cas9-sgRNAHd6 was constructed and introduced into NJ46 callus via Agrobacterium-mediated transformation. Homozygous mutant lines without exogenous components were screened using PCR. The heading date, major agronomic traits, yield-related traits, and grain quality traits of these lines were evaluated in both central and northern Jiangsu and compared with those of local benchmark cultivars Nanjing 9108 (NJ9108) and Nanjing 518 (NJ518), respectively. In the T0 generation, 18 transgenic positive plants were obtained. In the T2 generation, three homozygous mutant lines with different mutation types and free of exogenous sequences were identified: NJ46-hd6ko-1 (TTGG deletion), NJ46-hd6ko-2 (25 bp deletion), and NJ46-hd6ko-3 (G deletion). Their heading dates were 21.1, 24.7, and 21.4 d earlier than that of NJ46, respectively. Among them, NJ46-hd6ko-3 showed similar heading date, theoretical yield, milled rice rate, amylose content, gel consistency, and taste value to NJ9108, but had a shorter plant height and a higher alkali spreading value, indicating superior overall performance and suitability for cultivation in the central Jiangsu region. NJ46-hd6ko-2 showed no significant differences in heading date, theoretical yield, or alkali spreading value compared to NJ518, but demonstrated significantly higher milled rice rate and taste value, along with a shorter plant height, suggesting enhanced overall traits and suitability for the northern Jiangsu region. By precisely editing the Hd6 gene in NJ46, we developed new elite, tasty japonica rice germplasms adapted to the central and northern Jiangsu ecological zones, thereby expanding the cultivation area for high-quality, palatable rice varieties.

Key words: rice, CRISPR/Cas9, genetic editing, heading date, Hd6

表1

本研究所用的引物"

引物名称
Primer name		 引物序列
Primer sequence (5'-3')		 用途
Usage
Hd6-gF GGCAGTTGTCAGGAAAGTTGGAAGAGG Hd6靶序列构建Construct of Hd6 site
Hd6-gR AAACCCTCTTCCAACTTTCCTGACAAC
Hpt-F CGGAAGTGCTTGACATTG 转基因检测Transgenic detection
Hpt-R GACCTCGTATTGGGAATCC
SgRNA-F AACTTATAAACCCGCGCGCTG 单克隆测序鉴定Sequencing for monoclonal
SgRNA-R CAAGTTGATAACGGACTAGC
Cas9-F ACTCGAACGCGACCAACTTA 载体检测Vector assay
Cas9-R ATCTGCAGAATTGCCCTTCG
Hd6-F TGACTTACACAATACCTCGCTAACT Hd6 敲除靶点测序鉴定
Sequencing for Hd6 knockout target
Hd6-R CACAGGCTTGAGTATCTTGATGATG

图1

Hd6基因敲除靶点位置及终载体pCAMBIA1305-Actin:Cas9-sgRNAHd6组成示意图 PAM: 原间隔邻接基序。"

图2

gRNA-F/R引物对T0代转基因阳性植株进行PCR检测 1 : 阴性对照; 2: 阳性对照; 3-24: T0代转基因植株。"

图3

T0代转基因阳性植株突变类型分析 蓝色字体: 靶位点序列; 红色字体: 原间隔邻接基序(PAM)序列; 黑色虚线: 碱基缺失; WT: 野生型; NJ46: 南粳46。括号内的数字表示植株数。"

图4

PCR筛选无外源转基因成分的突变植株 Hpt和Cas9分别表示利用Hpt引物和Cas9引物通过PCR筛选无外源成分的突变植株; “+”: 阳性对照; “-”: 阴性对照; 1-3: NJ46-hd6ko-1-3突变系。"

图5

NJ46-hd6ko-1-3突变株系靶基因敲除位点测序分析 PAM: 原间隔邻接基序; NJ46: 南粳46; WT: 野生型。"

图6

NJ46-hd6ko-1-3突变株系Hd6编码区的氨基酸序列比对分析 红色区域表示NJ46-hd6ko-1-3突变系Hd6编码区氨基酸序列相同的区域。NJ46: 南粳46; WT: 野生型。"

表2

NJ46突变体表型分离比χ2测验"

T1代基因型
Genotype of T1 generation		 T2代分离群体抽穗期的变化统计
Statistics on the changes in heading date of T2 isolated population plants		 1:3分离比的渐进显著性
Progressive significance of the separation ratio of 1:3
早
Earlier		 无变化
No change		 晚
Later
Aa1 47 135 0 0.797
Aa2 54 184 0 0.410
Aa3 56 162 0 0.637

图7

NJ46-hd6ko-1-3突变株系的抽穗期表型 图A、B、C分别为NJ46-hd6ko-1-3纯合突变系在扬州沿江区域、金湖、宿迁的抽穗期表型与相应对照品种的抽穗期表型比较。基于LSD多重比较, 不同字母代表不同株系间差异显著(P < 0.05)。NJ46: 南粳46; NJ9108: 南粳9108; NJ518: 南粳518; WT: 野生型。图中标尺表示30 cm。"

表3

各早抽穗突变系与NJ9108和NJ518的农艺性状比较"

地点
Region		 名称
Name		 株高
Plant height (cm)		 剑叶长
Flag leaf length (cm)		 剑叶宽
Flag leaf width (cm)		 穗长
Panicle length (cm)		 每公顷有效穗数
Effective panicles per hectare
金湖Jinhu NJ9108 102.67±1.53 b 25.43±2.71 a 1.67±0.13 a 18.37±1.64 b 405.00±23.10 b
NJ46-hd6ko-1 101.67±0.58 ab 29.83±1.67 a 1.77±0.08 a 15.63±0.25 ab 336.90±27.30 a
NJ46-hd6ko-3 98.00±1.73 a 31.53±1.60 a 1.83±0.03 a 16.77±0.45 ab 380.85±5.10 ab
宿迁Suqian NJ518 102.33±1.53 c 27.23±1.37 a 1.62±0.13 b 16.13±0.67 ab 349.50±10.05 a
NJ46-hd6ko-2 97.00±1.73 b 27.17±4.14 a 1.83±0.15 b 15.83±1.40 ab 348.30±19.95 a
地点
Region		 名称
Name		 每穗总粒数
Grain number per panicle		 结实率
Seed-setting rate (%)		 千粒重
1000-grain weight (g)		 理论产量
Theoretical yield
(kg hm-2)
金湖Jinhu NJ9108 164.33±7.23 a 84.87±3.77 b 21.08±0.34 b 11,886.45±396.75 c
NJ46-hd6ko-1 141.67±19.14 a 87.87±1.26 b 24.93±0.26 c 10,379.85±531.75 b
NJ46-hd6ko-3 153.00±5.00 a 82.57±0.39 ab 24.44±0.47 c 11,756.25±335.85 c
宿迁Suqian NJ518 148.33±10.26 a 84.29±0.33 b 27.56±0.24 d 12,022.20±394.35 b
NJ46-hd6ko-2 147.00±9.85 a 86.31±2.04 b 26.49±0.49 c 11,679.75±449.40 b

表4

各早抽穗突变株系与对照品种NJ9108或NJ518的主要品质性状比较"

地点
Region		 名称
Name		 精米率
Milled rice rate (%)		 垩白粒率
Chalky rice rate (%)		 垩白度
Chalkiness (%)		 直链淀粉含量
Amylose content (%)
金湖 Jinhu NJ9108 71.43±0.43 b 40.93±2.05 a 12.20±0.53 b 9.80±0.90 a
NJ46-hd6ko-1 71.78±0.35 b 54.59±5.51 b 17.20±0.55 c 10.07±0.49 a
NJ46-hd6ko-3 70.90±0.53 b 34.21±1.86 a 8.80±0.97 a 10.31±0.97 a
宿迁 Suqian NJ518 67.71±0.36 a 47.81±2.21 c 15.66±0.62 b 9.70±0.79 a
NJ46-hd6ko-2 69.34±0.07 b 37.68±2.01 b 13.71±0.56 b 10.58±0.13 a
地点
Region		 名称
Name		 胶稠度
Gel consistency (mm)		 碱消值级
Alkali digestion grade		 食味值
Taste value
金湖 Jinhu NJ9108 89.00±1.10 b 6.53±0.36 a 76.00±5.29 ab
NJ46-hd6ko-1 87.20±1.20 ab 6.67±0.50 ab 80.33±1.15 b
NJ46-hd6ko-3 88.40±2.10 b 7.00±0.15 b 79.67±1.15 ab
宿迁 Suqian NJ518 87.00±1.40 bc 6.87±0.41 ab 77.00±6.08 a
NJ46-hd6ko-2 88.20±1.50 c 6.67±0.22 a 87.00±1.00 b

图8

NJ46-hd6ko-1-3突变株系的品质比较 图A、B分别为纯合突变株系在金湖、宿迁与相应对照品种的品质比较。NJ46: 南粳46; NJ9108: 南粳9108; NJ518: 南粳518。"

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