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作物学报 ›› 2026, Vol. 52 ›› Issue (2): 349-362.doi: 10.3724/SP.J.1006.2026.55035

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

Bna-miR1040-EIF3A模块调控油菜开花时间的功能研究

马毅娜(), 吴晓明玉, 李藕琪, 王圆, 陈丽, 张盈川, 赵伦, 文静, 傅廷栋, 沈金雄()   

  1. 华中农业大学作物遗传改良全国重点实验室 / 国家油菜工程技术研究中心, 湖北武汉 430070
  • 收稿日期:2025-06-03 接受日期:2025-10-30 出版日期:2026-02-12 网络出版日期:2025-11-05
  • 通讯作者: *沈金雄 E-mail:jxshen@mail.hzau.edu.cn
  • 作者简介:E-mail: mayina202202@163.com
  • 基金资助:
    国家自然科学基金项目(31930032);财政部和农业农村部国家现代农业产业技术体系建设专项(CARS-12)

Functional analysis of the Bna-miR1040-EIF3A module in regulating flowering time in rapeseed (Brassica napus)

Ma Yi-Na(), Wu Xiao-Ming-Yu, Li Ou-Qi, Wang Yuan, Chen Li, Zhang Ying-Chuan, Zhao Lun, Wen Jing, Fu Ting-Dong, Shen Jin-Xiong()   

  1. National Key Laboratory of Crop Genetic Improvement, Huazhong Agricultural University / National Engineering Research Center of Rapeseed, Wuhan 430070, Hubei, China
  • Received:2025-06-03 Accepted:2025-10-30 Published:2026-02-12 Published online:2025-11-05
  • Contact: *Shen Jin-Xiong E-mail:jxshen@mail.hzau.edu.cn
  • Supported by:
    National Natural Science Foundation of China project(31930032);China Agriculture Research System of MOF and MARA(CARS-12)

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摘要：

开花时间是作物重要发育性状, 对作物产量有显著影响。miRNA是一类由生物体内源MIR基因转录后加工而形成的小分子RNA, 可通过切割靶基因影响靶基因的表达。前期研究基础上在甘蓝型油菜中发现一个未知生物学功能的新的小分子RNA, novel-Bna-miR1040, 本研究过表达Bna-miR1040的甘蓝型油菜较野生型花期延迟。降解组测序表明BnaEIF3A是Bna-miR1040的潜在靶基因, 5'RACE和烟草瞬时转化试验表明, Bna-miR1040可以切割BnaEIF3A, 使BnaEIF3A的表达量降低; 进化分析结果表明, BnaEIF3A在不同物种中具有保守性; 亚细胞定位结果显示BnaEIF3A定位在细胞核上; 通过CRISPR/Cas9创建BnaEIF3A的功能缺失突变体, 突变体较野生型也表现为花期延迟。本研究结果为甘蓝型油菜miRNA调控开花时间提供了数据, 为甘蓝型油菜不同花期品种选育提供了理论参考。

关键词: 甘蓝型油菜, miRNA, Bna-miR1040, EIF3A, 开花时间

Abstract:

Flowering time is a critical developmental trait in crops that significantly influences yield. MicroRNAs (miRNAs), a class of small endogenous RNAs processed from MIR gene transcripts, regulate gene expression by mediating transcript cleavage. In this study, we identified a novel miRNA—novel-Bna-miR1040—in rapeseed (Brassica napus), whose biological function had not been previously characterized. Transgenic rapeseed plants overexpressing Bna-miR1040 exhibited delayed flowering compared to wild-type plants. Degradome sequencing identified BnaEIF3A as a potential target of Bna-miR1040. This interaction was validated by 5′RACE and transient expression assays in tobacco leaves, which confirmed that Bna-miR1040 cleaves BnaEIF3A, significantly reducing its transcript levels. Evolutionary analysis showed that BnaEIF3A is highly conserved across diverse plant species. Subcellular localization analysis revealed that the BnaEIF3A protein is exclusively localized in the nucleus. To further investigate its function, we generated BnaEIF3A loss-of-function mutants using CRISPR/Cas9 genome editing. These mutants also exhibited delayed flowering phenotypes compared to wild-type controls. Together, our findings demonstrate that Bna-miR1040 regulates flowering time in rapeseed by suppressing BnaEIF3A expression, providing novel insights for molecular breeding strategies aimed at optimizing flowering time in rapeseed.

Key words: Brassica napus, miRNA, Bna-miR1040, EIF3A, flowering time

附表1

载体构建扩增引物"

引物名称
Primer name		 引物序列
Primer sequence (5′-3′)
Bna-miR1040-F CAGGTCGACTCTAGAGGATCCACGCTAACGATTCCAGCTAA
Bna-miR1040-R CGAGAATTCGAGCTCGGTACCGACGCTAACGCTTCCAACGAA
BnaC09.EIF3A-GFP-F CTTGCATGCCTGCAGGTCGACATGGCGAATTTTGCCAAACCTG
BnaC09.EIF3A-GFP-R TCTACCGGTACCCGGGGATCCCCACGCTGTGGCCTGGGC
m7BnaC09.EIF3A-GFP-F CTTGCATGCCTGCAGGTCGACATGGCGAATTTTGCCAAACCTG
m7BnaC09.EIF3A-R TCGGCTCCAAGTTAAAGCCAATGAGGTTGGCCATCCTCAA
m7BnaC09.EIF3A-F GCTTTAACTTGGAGCCGAAATTCGAGGGCAGAGATATGGT
C83-R CTTGCATGCCTGCAGGTCGACCCACGCTGTGGCCTGGGC
bnaeif3a-DT1-BsF ATATATGGTCTCGATTGGCGAACAACAGAGTTCCGTGTT
bnaeif3a-DT1-F0 TGGCGAACAACAGAGTTCCGTGTTTTAGAGCTAGAAATAGC
bnaeif3a-DT2-R0 AACTTGGTCTCTGTACTTGTTCCAATCTCTTAGTCGACTCTAC
bnaeif3a-DT2-BsR ATTATTGGTCTCGAAACTTGGTCTCTGTACTTGTTCCAA
BnaC09.EIF3A-S1-F ggagtgagtacggtgtgcCCTCTCATCTTTAGTTGATTAAGCTCTA
BnaC09.EIF3A-S1-R gagttggatgctggatggTCAGGTTTGCCAAATGGTTTCTGATA
BnaA09.EIF3A-S1-F ggagtgagtacggtgtgcCCGTAAATCTTTAGTTGATTAAGGTC
BnaA09.EIF3A-S1-R gagttggatgctggatggCAGGTTCGCCAAATGGTTCCTG
BnaA02.EIF3A-S1-F ggagtgagtacggtgtgcGATTTGTTCATTGTGTAGATGACAGC
BnaA02.EIF3A-S1-R gagttggatgctggatggGAGCTGCAAGCTCTCTGGG
BnaC02.EIF3A-S1-F ggagtgagtacggtgtgcCGTGTAGATGACCGCACATAAAGCCTTT
BnaC02.EIF3A-S1-R gagttggatgctggatggCAGCTGCAAGCTCTCCGGG
BnaA02.EIF3A-S1-F ggagtgagtacggtgtgcGATTTGTTCATTGTGTAGATGACAGC
BnaA02.EIF3A-S1-R gagttggatgctggatggGAGCTGCAAGCTCTCTGGG
BnaC02.EIF3A-S1-F ggagtgagtacggtgtgcCGTGTAGATGACCGCACATAAAGCCTTT
BnaC02.EIF3A-S1-R gagttggatgctggatggCAGCTGCAAGCTCTCCGGG
BnaA03a.EIF3A-S2-F ggagtgagtacggtgtgcCGAGTTCCGTAGGCTTTGTGAAATC
BnaA03a.EIF3A-S2-R gagttggatgctggatggAGGTAAAGCTGCAAGCTCTCAGGA
BnaC03.EIF3A-S2-F ggagtgagtacggtgtgcGAGTTCCGCAGGCTTTGTGAAATC
BnaC03.EIF3A-S2-R gagttggatgctggatggAAGCTGCAAGCTCTCTGGG
RT Bna-miR1040-primer GTCGTATCCAGTGCAGGGTCCGAGGTATTCGCACTGGATACGACAACCCA
qRT Bna-miR1040-F CGCGCGTAGGGTCTAAGACT
qRT Bna-miR1040-R AGTGCAGGGTCCGAGGTATT
RT U6-primer TTGGACCATTTCTCGATTTTTG
qRT U6-F TTGGAACGATACAGAGAAGATTAGCA
qRT U6-R TTGGACCATTTCTCGATTTTTG
qRT BnaC09.EIF3A -F GACCGAAAGCCTGAGGATCTG
qRT BnaC09.EIF3A -R CGCTTTATGTGCAGTCATCGCG
qRT BnaA09.EIF3A -F CCAGGCACTTCATGATCTCATCACG
qRT BnaA09.EIF3A -R GGAAAGGTGCAGGAAGTGCTTGATA
qRT BnaA02.EIF3A -F GTCGTTTCGCCAAGGACGGT
qRT BnaA02.EIF3A -R CGGCCTTGTCGGTAGAGAGATGA
qRT BnaC02.EIF3A -F GGGTCGTTTCGCTAAGGACGGA
qRT BnaC02.EIF3A -R TGCATCAGCCTGAGAACGAGCCA
qRT BnaA03a.EIF3A -F GTGGCAGAAGCCGCTCGAGAA
qRT BnaA03a.EIF3A -R GCTCACATTCACTTGCTGGCATACA
qRT BnaC03.EIF3A -F CATGGCAGAAGCCGCTCGAAA
qRT BnaC03.EIF3A -R GCTGGCAAACGATACGGTACTGG
qRT BnaA03b.EIF3A -F TAGATGATCTTGAGGTTACACCATGG
qRT BnaA03b.EIF3A -R GTACTTCTTGCAGAACTGGAAGGCT
qRT BnaC07.EIF3A -F CCGTAATTCTTGTCAGCAAGTGAACC
qRT BnaC07.EIF3A -R CTCCCACAAGAACTTGAACCATGG
qRT BnaACT7-F CTATCCTCCGTCTCGATCTCGC
qRT BnaACT7-R CTTAGCCGTCTCCAGCTCTTGC
Bna-miR1040-4-JCF TCTAGACCCTAAACCCAAACCATAAACT
BnaC09.EIF3A-JCF TTGAGTTGAAGTGTATGAGGTCCA
C83-GFP-R CAAGAATTGGGACAACTCCAGTG

图1

Bna-miR1040前体二级结构预测图 箭头所指处为成熟体序列。"

附图1

不同甘蓝型油菜品种中Bna-pre-miR1040序列比对图 框内为成熟体序列。Bna-pre-miR1040在ZS11、Darmor、Quinta、Tapidor、Westar、No2127六种油菜品种中对应的核苷酸序列比对。"

图2

Bna-miR1040过表达材料构建及初花期考察 A: 生长46 d的WT及Bna-miR1040过表达材料。标尺为10 cm。B: Bna-miR1040过表达材料表达量鉴定。C: WT及Bna-miR1040过表达材料初花期统计。每株系考察5~7株, 3次生物学重复, 3次技术重复。采用t测验分析; **表示P < 0.01。"

表1

Bna-miR1040降解组测序预测的8个候选靶基因"

miRNA名称
miRNA name		 靶基因
Target gene		 拟南芥同源基因
Arabidopsis thaliana homologous gene		 靶基因功能
Function of target gene
Bna-miR1040 BnaA09g21660D AT4G11420 Eukaryotic translation initiation factor 3 subunit A
BnaC09g23910D
BnaA02g21720D
BnaCnng08330D
BnaA07g32230D AT1G75950 SKP1-like protein 1A
BnaA02g04280D AT5G19530 Thermospermine synthase
BnaC09g54340D AT5G60690 Homeobox-leucine zipper protein
BnaCnng09000D AT1G01960 Brefeldin A-inhibited guanine nucleotide-exchange protein 3

图3

Bna-miR1040靶基因验证 A: p35S::Bna-miR1040、p35S::BnaC09.EIF3A-GFP、p35S::m7BnaC09.EIF3A-GFP载体构建图。“∶”表示碱基配对, “.”表示G/C: U配对, 箭头表示BnaC09.EIF3A具体被切割的位置, 3/3右边数字表示总挑斑克隆数、左边数字表示BnaC09.EIF3A在该位点被切割后所获得的克隆数。B: Bna-miR1040过表达材料中靶基因BnaEIF3A四个拷贝表达量检测图, 3次生物学重复, 3次技术学重复。采用t测验分析, *表示P < 0.05。C: BnaEIF3A亚细胞定位。标尺为20 μm。D: 共聚焦显微镜下不同浓度p35S::Bna-miR1040农杆菌液分别与p35S::BnaC09.EIF3A-GFP、p35S::m7BnaC09.EIF3A-GFP共侵染烟草叶片后表达GFP荧光信号图像, 标尺为20 μm。"

图4

BnaEIF3家族系统发育树"

图5

BnaEIF3A生物信息学分析 A: EIF3A在不同物种间的进化关系。B: BnaEIF3A及拟南芥同源基因Motif分析。C: BnaEIF3A及拟南芥同源基因功能结构域。"

图6

不同拷贝BnaEIF3A在Westar各组织中的相对表达量 不同字母表示BnaEIF3A同一拷贝中不同组织间差异显著(P < 0.05)。"

图7

BnaEIF3A敲除材料T1代初花期统计 A: BnaEIF3A敲除载体靶点分布图。B: 生长46 d的WT、CR#23、CR#100、CR#32。标尺为10 cm。C: WT、CR#23、CR#67、CR#15、CR#100、CR#32、CR#102初花期统计图, 每株系统计5~7株。采用t测验分析, **表示P < 0.01。"

附表2

bnaeif3a敲除材料T0代编辑情况统计"

载体Vectors 拷贝数
Copy
number		 野生型
WT
(%)		 单拷贝突变
One copy
mutated (%)		 两拷贝突变Two copies mutated (%) 三拷贝突变Three copies mutated (%) 四拷贝突变Four copies mutated (%) 五拷贝突变Five copies mutated (%) 六拷贝突变
Six copies
mutated (%)
bnaeif3a 6 35.88 5.34 1.53 26.72 9.92 10.69 9.92

附图2

T1代bnaeif3编辑情况"

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