中国水稻科学 ›› 2025, Vol. 39 ›› Issue (3): 331-342.DOI: 10.16819/j.1001-7216.2025.2401010
马顺婷1, 胡运高1, 高方远2, 刘利平2, 牟昌铃2, 吕建群2, 苏相文2, 刘松2, 梁毓玉1, 任光俊2,*(
), 郭鸿鸣2,*()
收稿日期:2024-10-25
修回日期:2025-03-27
出版日期:2025-05-10
发布日期:2025-05-21
通讯作者:
*email: guangjun61@sina.com;hongmingguo552@163.com基金资助:
MA Shunting1, HU Yungao1, GAO Fangyuan2, LIU Liping2, MOU Changling2, LÜ Jianqun2, SU Xiangwen2, LIU Song2, LIANG Yuyu1, REN Guangjun2,*(), GUO Hongming2,*()
Received:2024-10-25
Revised:2025-03-27
Online:2025-05-10
Published:2025-05-21
Contact:
*email: guangjun61@sina.com;hongmingguo552@163.com摘要：
【目的】本研究旨在通过反向遗传学策略，深入探究水稻真核翻译起始因子OseIF6.2在水稻生长发育中的具体功能，以期为OseIF6.2功能的进一步解析奠定基础。【方法】首先，利用生物信息学方法进行多重序列比对及进化树的构建，以分析OseIF6.2与其他真核生物中eIF6蛋白的系统发育关系；运用CRISPR/Cas9基因编辑技术，对水稻(日本晴)中的OseIF6.2基因进行精准编辑。其次，通过qRT-PCR技术，检测OseIF6.2在水稻不同组织中的表达水平；借助激光共聚焦显微镜，观察OseIF6.2在细胞内的精确定位。然后对野生型与oseif6.2突变体的粒长、粒宽、粒厚及千粒重等关键农艺性状进行比较分析。最后，通过酵母双杂交(Y2H)及双分子荧光互补(BiFC)实验，验证OseIF6.2与其潜在互作蛋白OseIF6.1的相互作用。【结果】系统发育分析结果表明，OseIF6.2与其他真核生物中的eIF6蛋白具有高度保守性，凸显了其在真核生物翻译调控中的普遍重要性。qRT-PCR结果显示，OseIF6.2在水稻茎中的表达水平较高。通过将OseIF6.2蛋白与GFP融合后注入本氏烟叶片细胞，并在激光共聚焦显微镜下观察，发现OseIF6.2在细胞核及细胞质区域均有荧光信号分布，明确了其亚细胞定位。此外，与野生型相比，oseif6.2突变体在粒长、粒厚及千粒重等性状上均表现为显著降低。Y2H及BiFC实验均成功验证了OseIF6.2与OseIF6.1蛋白之间的直接相互作用。【结论】综上所述，OseIF6.2作为水稻真核翻译起始因子，其功能异常显著影响水稻的粒型特征，进一步揭示了该因子在水稻生长发育及粒型形成中的关键作用。
马顺婷, 胡运高, 高方远, 刘利平, 牟昌铃, 吕建群, 苏相文, 刘松, 梁毓玉, 任光俊, 郭鸿鸣. 水稻真核翻译起始因子OseIF6.2调控粒型的功能研究[J]. 中国水稻科学, 2025, 39(3): 331-342.
MA Shunting, HU Yungao, GAO Fangyuan, LIU Liping, MOU Changling, LÜ Jianqun, SU Xiangwen, LIU Song, LIANG Yuyu, REN Guangjun, GUO Hongming. Functional Study of Rice Eukaryotic Translation Initiation Factor OseIF6.2 in Grain Size Regulation[J]. Chinese Journal OF Rice Science, 2025, 39(3): 331-342.
| 引物名称 Primer name | 引物序列 Sequence (5＇-3＇) | 用途 Purpose |
|---|---|---|
| sgRNA序列 | GCGAGATTGGAGTGTTCGCGAGG | OseIF6.2基因sgRNA序列 |
| eIF6.2-CDS-F | ATGATTTTTTTTTTCAAAAATCGGTTCTGCATTGCA | CDS 扩增 |
| eIF6.2-CDS-R | CTATAGGTAGCTGCTGACGACGAGCGA | CDS amplification |
| K21-F | AGTTTGTAACAGCTGCTGGGATTAC | 亚细胞定位载体 |
| K21-NOSR-R | CCATCTCATAAATAACGTCATGCATTACATGTT | Subcellular localization vector |
| eIF6.2-Cas9-F | AGGAGTTCTTGGTTGGCTGACTGCGA | 基因编辑材料鉴定 |
| eIF6.2-Cas9-R | ACGCACATCCTCCCGATGATCCT | Identification of gene editing materials |
| OsCas9-1000-F | GAGGAGACAATCACCCCCTGGAACT | Cas9蛋白鉴定 |
| OsCas9-1000-R | TTATCGGACTTGCCCCTGTTCTTGT | Identification of Cas9 protein |
| eIF6.1-2YC-767-F | ACGATAGTTAATTAACATGGCGACCCGTATTCAGTTTG | BiFC载体 |
| eIF6.1-2YC-767-R | GATCGACAGTTATGTGCACTAGTGGCGCGCCC | Bimolecular fluorescence complementation vector |
| eIF6.1-CDS-759-F | CCTTAATTAAGGATGGCGACCCGTATTCAGTTTGAGAAC | CDS 扩增 |
| eIF6.1-CDS-759-R | TTGGCGCGCCAACACATAACTGTCGATCAAAG | CDS amplification |
| eIF6.2-BD-798-F | GAGGACCTGCATATGATGATTTTTTTTTTCAAAAATCG | 酵母双杂载体 |
| eIF6.2-BD-798-R | GATCCCCGGGAATTCCTATAGGTAGCTG | Construction of yeast two-hybrid vector |
| eIF6.2-2YN-797-F | AC GATAGTTAATTAACATGATTTTTTTTTTCAAAAATCGG | BiFC载体构建 |
| eIF6.2-2YN-797-R | CACTAGTGGCGCGCCCTAGGTAGCTGC | Constructionof of bimolecular fluorescence complementation vector |
| Actin-qPCR-F | TGTATGCCAGTGGTCGTACCA | 内参基因定量引物 |
| Actin-qPCR-R | CCAGCAAGGTCGAGACGAA | Primers of internal reference gene |
| eIF6.2-qPCR-148-F | AGAGAACTTCTTCAGCGTGTTC | OseIF6.2 定量引物 |
| eIF6.2-qPCR-148-R | CTCTTGGTCTGTGGTGGTATG | qRT-PCR primers of OseIF6.2 |
| OsEXPA5-F | AAGGCTGTGGCTTGATTGACA | 细胞膨大相关基因定量引物 |
| OsEXPA5-R | TTAGGCCCAATTTTGCTATTTTG | qRT-PCR primers of genes related to cell expansion |
| OsEXPA8-F | GCGATGAGCCGCAACTG | 细胞膨大相关基因定量引物 |
| OsEXPA8-R | CTCTTCCATCCTATGCCACG | qRT-PCR primers of genes related to cell expansion |
| OsEXPA30-F | CCAAGTTCAGGGCGATGCAG | 细胞膨大相关基因定量引物 |
| OsEXPA30-R | ACGGTGATCGCCGGCGAGCC | qRT-PCR primers of genes related to cell expansion |
| OsEXPB3-F | CTTTGAGTGGTTGGAGTGGTGG | 细胞膨大相关基因定量引物 |
| OsEXPB3-R | GCAGCCTTCTTGGAGATGGAA | qRT-PCR primers of genes related to cell expansion |
表1 本研究中使用的引物
Table 1. Primers used in this study
| 引物名称 Primer name | 引物序列 Sequence (5＇-3＇) | 用途 Purpose |
|---|---|---|
| sgRNA序列 | GCGAGATTGGAGTGTTCGCGAGG | OseIF6.2基因sgRNA序列 |
| eIF6.2-CDS-F | ATGATTTTTTTTTTCAAAAATCGGTTCTGCATTGCA | CDS 扩增 |
| eIF6.2-CDS-R | CTATAGGTAGCTGCTGACGACGAGCGA | CDS amplification |
| K21-F | AGTTTGTAACAGCTGCTGGGATTAC | 亚细胞定位载体 |
| K21-NOSR-R | CCATCTCATAAATAACGTCATGCATTACATGTT | Subcellular localization vector |
| eIF6.2-Cas9-F | AGGAGTTCTTGGTTGGCTGACTGCGA | 基因编辑材料鉴定 |
| eIF6.2-Cas9-R | ACGCACATCCTCCCGATGATCCT | Identification of gene editing materials |
| OsCas9-1000-F | GAGGAGACAATCACCCCCTGGAACT | Cas9蛋白鉴定 |
| OsCas9-1000-R | TTATCGGACTTGCCCCTGTTCTTGT | Identification of Cas9 protein |
| eIF6.1-2YC-767-F | ACGATAGTTAATTAACATGGCGACCCGTATTCAGTTTG | BiFC载体 |
| eIF6.1-2YC-767-R | GATCGACAGTTATGTGCACTAGTGGCGCGCCC | Bimolecular fluorescence complementation vector |
| eIF6.1-CDS-759-F | CCTTAATTAAGGATGGCGACCCGTATTCAGTTTGAGAAC | CDS 扩增 |
| eIF6.1-CDS-759-R | TTGGCGCGCCAACACATAACTGTCGATCAAAG | CDS amplification |
| eIF6.2-BD-798-F | GAGGACCTGCATATGATGATTTTTTTTTTCAAAAATCG | 酵母双杂载体 |
| eIF6.2-BD-798-R | GATCCCCGGGAATTCCTATAGGTAGCTG | Construction of yeast two-hybrid vector |
| eIF6.2-2YN-797-F | AC GATAGTTAATTAACATGATTTTTTTTTTCAAAAATCGG | BiFC载体构建 |
| eIF6.2-2YN-797-R | CACTAGTGGCGCGCCCTAGGTAGCTGC | Constructionof of bimolecular fluorescence complementation vector |
| Actin-qPCR-F | TGTATGCCAGTGGTCGTACCA | 内参基因定量引物 |
| Actin-qPCR-R | CCAGCAAGGTCGAGACGAA | Primers of internal reference gene |
| eIF6.2-qPCR-148-F | AGAGAACTTCTTCAGCGTGTTC | OseIF6.2 定量引物 |
| eIF6.2-qPCR-148-R | CTCTTGGTCTGTGGTGGTATG | qRT-PCR primers of OseIF6.2 |
| OsEXPA5-F | AAGGCTGTGGCTTGATTGACA | 细胞膨大相关基因定量引物 |
| OsEXPA5-R | TTAGGCCCAATTTTGCTATTTTG | qRT-PCR primers of genes related to cell expansion |
| OsEXPA8-F | GCGATGAGCCGCAACTG | 细胞膨大相关基因定量引物 |
| OsEXPA8-R | CTCTTCCATCCTATGCCACG | qRT-PCR primers of genes related to cell expansion |
| OsEXPA30-F | CCAAGTTCAGGGCGATGCAG | 细胞膨大相关基因定量引物 |
| OsEXPA30-R | ACGGTGATCGCCGGCGAGCC | qRT-PCR primers of genes related to cell expansion |
| OsEXPB3-F | CTTTGAGTGGTTGGAGTGGTGG | 细胞膨大相关基因定量引物 |
| OsEXPB3-R | GCAGCCTTCTTGGAGATGGAA | qRT-PCR primers of genes related to cell expansion |
图1 OseIF6.2的表达模式及亚细胞定位 A：对不同真核生物物种中eIF6蛋白的系统发育比较分析。B：OseIF6.2在幼苗根、茎、叶、鞘和4~5、8~10、10~12 cm幼穗中的表达情况，OsActin作为内参。数据是来自三组独立实验的平均值±标准差。C：OseIF6.2在烟草叶中的亚细胞定位。GFP：绿色荧光蛋白，RFP：红色荧光蛋白。标尺=20 μm。
Fig. 1. Expression pattern and subcellular localization of OseIF6.2 A, Comparative phylogenetic analysis of eIF6 protein cross diverse eukaryotic species. B, Relative expression of OseIF6.2 in root, stem, leaf, and sheath of young seedlings and developing panicles at lengths of 4-5, 8-10, and 10-12 cm. OsActin served as the internal control. Values represent means ± SE from at least three independent experiments. C, Subcellular localization of OseIF6.2 in N. benthamiana leaves. YFP: Yellow fluorescent protein, RFP: Red fluorescent protein. Scale bars = 20 μm.
图2 OseIF6.2突变体类型及表型分析 A：OseIF6.2突变类型。B：野生型和oseif6.2突变体植株成熟期的形态和穗部结构。株高比例尺=10 cm，穗长比例尺=1 cm。C：野生型和oseif6.2突变体植株的穗长。D：野生型和oseif6.2突变体穗的一次枝梗数。E：野生型和oseif6.2突变体穗的二次枝梗数。F：野生型和oseif6.2突变体植株的结实率。数据是来自三组独立实验的平均值±标准差, t检验:*P< 0.05, **P< 0.01, ***P< 0.001。
Fig. 2. Types and phenotypic analysis of OseIF6.2 mutants A, Mutation types of OseIF6.2. B, Plants and panicles of WT and oseif6.2 mutant at the mature stage. Scale bars = 10 cm for plant height and 1 cm for panicle length. C, Panicle length of WT and oseif6.2 mutant. D, Number of primary branches per panicle of WT and oseif6.2 mutant. E, Number of secondary branches of WT and oseif6.2 mutant. F, Seed setting rate of WT and oseif6.2 mutant. Values represent means ± SE from at least three independent experiments, Student's t-test: *P< 0.05, **P< 0.01, ***P< 0.001.
图3 OseIF6.2的突变影响水稻粒型 A: 野生型和oseif6.2突变体株系的成熟水稻籽粒。比例尺=1 cm。B: 野生型和oseif6.2突变体株系的粒长；C: 粒宽；D: 粒厚；E: 千粒重；F: 每穗粒数；G: 单株穗数；H: 单株产量。数据是来自三组独立实验的平均值±标准差，t检验:* P< 0.05, **P< 0.01, ***P< 0.001。
Fig. 3. Mutation of OseIF6.2 alters grain shape A, Mature grains of WT and oseif6.2 mutants. Scale bars =1 cm. B, Grain length of WT and oseif6.2 mutants; C, Grain width; D, Grain thickness; E, 1000-grain weight; F, Filled grains per panicle; G, Panicle number per plant; H, Grain yield per plant. Values represent means ± SE from at least three independent experiments, Student's t-test: *P < 0.05, **P < 0.01, ***P< 0.001.
图4 OseIF6.2通过影响细胞膨大调节水稻粒型 A：野生型和oseif6.2突变体成熟谷粒的颖壳扫描电子显微镜图像。比例尺=100 μm。野生型和oseif6.2突变体颖壳细胞长度（B）和宽度（C）。D：野生型和oseif6.2突变体颖壳细胞面积。数据是来自三组独立实验的平均值±标准差。t检验: *P< 0.05, **P< 0.01, ***P< 0.001。
Fig. 4. OseIF6.2 regulates grain size by affecting cell expansion A, Scanning electron microscopy images of the glume outer surfaces of WT and oseif6.2 mutants. Scale bars = 100 μm. Cell length (B) and width (C) of the outer epidermal of WT and oseif6.2 mutant lemmas. D, Cell area of the outer epidermal of WT and oseif6.2 mutant lemmas. Values represent means ± SE from at least three independent experiments. Student's t-test: *P < 0.05, **P < 0.01, ***P < 0.001.
图5 野生型和oseif6.2突变体株系细胞膨大相关基因的表达水平 数据是来自三组独立实验的平均值±标准差。t检验: *P< 0.05, **P< 0.01, ***P< 0.001。
Fig. 5. Expression levels of genes related to cell expansion in WT and oseif6.2 mutant lines The values represent means ± SE derived from at least three independent experiments. Student's t-test: *P < 0.05, **P < 0.01, ***P < 0.001.
图6 OseIF6.2与OseIF6.1互作 A：OseIF6.2在酵母细胞中与OseIF6.1相互作用。转化在DDO或QDO/X/A培养基上培养。B：双分子荧光互补分析验证烟草中OseIF6.2与OseIF6.1的相互作用。OseIF6.2-nYFP与OseIF6.1-cYFP在烟草细胞中共表达。比例尺=10 μm。
Fig. 6. OseIF6.2 physically interacts with OseIF6.1 A, OseIF6.2 interacts with OseIF6.1 in yeast cells. Transformed cells were cultured on DDO or QDO/X/A media. B, Bimolecular fluorescence complementation assays verifies the interaction between OseIF6.2 and OseIF6.1 in N. benthamiana. OseIF6.2-nYFP was coexpressed with OseIF6.1-cYFP in cells of N. benthamiana. Scale bars = 10 μm.
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