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作物学报 ›› 2023, Vol. 49 ›› Issue (2): 365-376.doi: 10.3724/SP.J.1006.2023.12076

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

OsNAC2d基因编辑水稻突变体的创建及其对干旱胁迫的响应

李兆伟1,2(), 莫祖意1,2, 孙聪颖1,2, 师宇1,2, 尚平2,3, 林伟伟1,2, 范凯2,3, 林文雄1,2,*()   

  1. 1福建农林大学生命科学学院, 福建福州 350002
    2福建农林大学 / 福建省农业生态过程与安全监控重点实验室, 福建福州 350002
    3福建农林大学农学院, 福建福州 350002
  • 收稿日期:2021-11-05 接受日期:2022-06-07 出版日期:2022-07-08 网络出版日期:2022-11-21
  • 通讯作者: 林文雄
  • 作者简介:E-mail: lizw197@163.com
  • 基金资助:
    福建省自然科学基金项目(2021J01098);福建省自然科学基金项目(2022J01142)

Construction of rice mutants by gene editing of OsNAC2d and their response to drought stress

LI Zhao-Wei1,2(), MO Zu-Yi1,2, SUN Cong-Ying1,2, SHI Yu1,2, SHANG Ping2,3, LIN Wei-Wei1,2, FAN Kai2,3, LIN Wen-Xiong1,2,*()   

  1. 1College of Life Sciences, Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
    2Fujian Provincial Key Laboratory of Agroecological Processing and Safety Monitoring, Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
    3College of Agriculture, Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
  • Received:2021-11-05 Accepted:2022-06-07 Published:2022-07-08 Published online:2022-11-21
  • Contact: LIN Wen-Xiong
  • Supported by:
    Natural Science Foundation of Fujian Province(2021J01098);Natural Science Foundation of Fujian Province(2022J01142)

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摘要：

为探究转录因子OsNAC2d的生物学功能及其对水稻耐旱性的影响, 本研究利用CRISPR/Cas9编辑技术对粳稻中花11中的OsNAC2d基因进行突变, 并考察osnac2d突变体在田间种植下的农艺性状, 以及突变体幼苗在干旱胁迫下的生长情况和OsNAC2d基因表达水平。结果表明, OsNAC2d基因主要在水稻成熟籽粒、叶片和花药中表达, 在根系和茎中的表达量较低, 并且受干旱胁迫诱导, 在10株阳性OsNAC2d基因突变株系的T2代植株中, 筛选出6种纯合osnac2d突变体, 田间试验调查表明, osnac2d突变体与野生型中花11相比, 株高、有效穗、穗长、穗粒数、结实率、千粒重和单株产量等农艺性状无显著差异。在干旱胁迫时, osnac2d突变体幼苗的根系与植株生长、根系和地上部生物量的积累均受到抑制, 且OsNAC2d在osnac2d突变体中的表达量维持在较低水平, 而野生型水稻的OsNAC2d表达则受干旱胁迫诱导而增强, 植株生长与生物量积累未受到显著抑制, 表明转录因子OsNAC2d正调控水稻响应干旱胁迫。创建的osnac2d突变体材料为进一步揭示OsNAC2d的生物学功能及其响应干旱胁迫的精细调控机制提供了优质种质资源。

关键词: 水稻, 干旱胁迫, 基因编辑, OsNAC2d基因

Abstract:

The main objective of this study is to explore the biological function of transcription factor OsNAC2d and its effect on drought resistance in rice. The sequence of OsNAC2d was firstly edited in japonica variety Zhonghua 11 through the CRISPR/Cas9 technology. The agronomic traits of osnac2d mutants were investigated in normal field cultivation, and the growth status and relative expression level of OsNAC2d genes were performed in the young mutant seedlings under the drought condition. There was the higher expression level of OsNAC2d in grain, leaf, and anther than those in root and stem, and the relative expression of OsNAC2d was enhanced by drought stress. Six homozygous T2 osnac2d lines were identified in two editing target sites of OsNAC2d genomic sequence through the nucleotide sequencing technology. There was no significant difference between osnac2d mutants and wild type in the agronomic traits including plant height, effective panicle number per plant, panicle length, grain number per panicle, seed setting rate, 1000-grain weight, and grain yield per plant. Under the drought condition, the growth of roots and seedling of osnac2d mutants were depressed, and the biomass of roots and above-ground were also decreased. Meanwhile, the relative expression of OsNAC2d in the osnac2d mutant retained as extremely low level as those in the normal cultivating condition. However, the relative expression of OsNAC2d in wild type was enhanced by drought stress, and the growth and biomass accumulation of wild type were not evidently hindered by drought stress. These results indicated that OsNAC2d positively regulated the drought response in rice. The osnac2d mutants in this present study provided a precious genetic resource for deeply revealing the biological function of OsNAC2d and its fine regulating mechanism in response to drought stress.

Key words: rice, drought stress, gene editing, OsNAC2d gene

图2

OsNAC2d的gRNA靶点和pYLCRISPR/Cas9-OsNAC2d-T12表达载体重组示意图 A: OsNAC2d的2个gRNA靶点位置和碱基序列; B: 2个靶点gRNA表达盒与pYLCRISPR/Cas9-MT重组示意图。"

表1

本研究采用的核苷酸序列及相应引物"

引物名称Primer name 引物序列Primer sequence (5'-3') 用途Usage
OsNAC2d-T1-fwd gccgTCCATGGCGGCACGCTCAG 靶点1序列
Sequence of target site 1
OsNAC2d-T1-rev aaacCTGAGCGTGCCGCCATGGA
OsNAC2d-T2-fwd gttgGTGGCACTAGAGCTGCAGT 靶点2序列
Sequence of target site 2
OsNAC2d-T2-rev aaacACTGCAGCTCTAGTGCCAC
OsNAC2d-T1-F TCGTCTCCCGCTACCTTCT 靶点1检测
Sequencing for target 1
OsNAC2d-T1-R GCGTACTCGTGCATAACCC
OsNAC2d-T2-F AGAACTCCTACGACCTAATGGC 靶点2检测
Sequencing for target 2
OsNAC2d-T2-R ATGTCCGACGCACGCAGAG
Hyp-F ACGGTGTCGTCCATCACAGTTTGCC 阳性转基因植株鉴定
Identification of the positive transgenic plant
Hyp-R TTCCGGAAGTGCTTGACATTGGGGA
OsNAC2d-RT-F GATCAGATCGTTCCCCCAGC OsNAC2d表达量分析
Relative expression analysis of OsNAC2d
OsNAC2d-RT-R TGCGACACAGACAGGTCATC

图1

水稻OsNAC2d的表达量分析 A: OsNAC2d在野生型ZH11水稻的根、茎、叶、花药和籽粒中的表达水平; B: 叶片和根系中OsNAC2d在15%的PEG-6000胁迫下的动态表达分析, 取样时间分别为处理后0、0.5、3、6、12和24 h; 不同字母表示在P < 0.05水平的显著性差异。"

图3

pYLCRISPR/Cas9-OsNAC2d-T12重组表达载体的2个靶点测序检测"

图4

阳性转基因株系鉴定 -: 阴性对照, 为等体积的水代替模板; +: 阳性对照, 为等体积阳性质粒作模板; 1~11: 转基因株系; M: 2 kb DNA ladder。"

图5

osnac2d突变体与野生型植株中的OsNAC2d基因序列比对分析 蓝色字母表示靶点序列, 黄色高亮为PAM序列, 删除线代表缺失碱基, 红色小写字母为插入碱基, -表示缺失, +表示插入, WT代表野生型(中花11)。"

图6

osnac2d突变体与野生型水稻在含15%的PEG-6000营养液胁迫下的表型 A和D为野生型水稻中花11, B和E为osnac2d-6突变体, C和F为osnac2d-4-2突变体, 对照组为完全营养液培养, PEG-6000处理时长为14 d。"

图7

osnac2d突变体与野生型ZH11在含15%的PEG-6000营养液中的长势情况 A为根系长度, B为地上部长度, C为植株总长度, 对照组为完全营养液培养, PEG-6000处理时长为14 d, 不同字母表示在P < 0.05水平的显著性差异。"

图8

osnac2d突变体与野生型ZH11在含15%的PEG-6000营养液中的生物量 A为根系生物量, B为地上部生物量, C为植株总生物量, 对照组为完全营养液培养, PEG-6000处理时长为14 d, 不同字母表示在P < 0.05水平的显著性差异。"

图9

osnac2d-6、osnac2d-4-2突变体和野生型ZH11中OsNAC2d基因在15%的PEG-6000胁迫下的表达水平 对照组为完全营养液培养, 不同字母表示在P < 0.05水平的显著性差异。"

表2

不同类型osnac2d突变体植株及其野生型水稻(WT)的农艺性状"

株系
Line		 株高
Plant height (cm)		 有效穗数
Effective panicle number per plant		 穗长
Length of main panicle (cm)		 穗粒数
Grain number per panicle		 结实率
Seed-setting rate (%)		 千粒重
1000-grain weight (g)		 单株产量
Grain yield per plant (g)
osnac2d-1 68.0±1.0 ab 10.0±2.0 a 23.33±2.75 a 107.25±16.82 ab 77.99±12.71 ab 22.70±1.04 ab 11.14±1.55 a
osnac2d-2-1 64.7±0.6 b 9.0±1.0 a 22.52±1.57 a 115.67±17.59 ab 72.32±13.44 ab 24.30±1.00 a 12.58±2.27 a
osnac2d-2-2 67.3±1.2 ab 10.7±1.5 a 23.43±0.98 a 120.00±19.03 ab 59.83±14.19 b 21.38±0.71 b 11.51±0.90 a
osnac2d-4-1 67.0±2.0 ab 11.3±2.1 a 22.53±0.48 a 117.00±7.81 ab 76.96±2.55 ab 21.84±0.41 ab 14.63±1.04 a
osnac2d-4-2 66.0±2.0 ab 9.3±2.1 a 22.33±0.99 a 88.02±11.79 b 80.67±0.94 ab 22.20±0.26 ab 9.26±2.32 a
osnac2d-6 67.0±1.0 ab 10.7±1.5 a 23.94±1.09 a 112.33±9.18 ab 72.46±8.94 ab 23.20±0.20 ab 12.77±1.84 a
WT(ZH11) 69.2±0.5 a 8.7±2.1 a 21.98±0.70 a 123.11±9.47 a 83.76±6.85 a 22.33±1.55 ab 9.96±3.06 a
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