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作物学报 ›› 2025, Vol. 51 ›› Issue (12): 3184-3197.doi: 10.3724/SP.J.1006.2025.55032

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

过表达BnNRT2.3-like对油菜氮素吸收利用及产量的影响

黄绒, 周渠晨, 陈楚铭, 罗倩, 易东, 杜常欢, 黄祥宇, 盛锋(), 杜雪竹()   

  1. 湖北大学生命科学学院, 湖北武汉 430062
  • 收稿日期:2025-05-15 接受日期:2025-09-10 出版日期:2025-12-12 网络出版日期:2025-09-17
  • 通讯作者: *盛锋, E-mail: shengfsk@163.com;杜雪竹, E-mail: duxuezhusk@163.com
  • 基金资助:
    本研究由湖北省科技创新计划项目(2025BBB012);科技创新2030—重大项目(2023ZD04070)

Effects of BnNRT2.3-like overexpression on nitrogen uptake, utilization efficiency, and yield in Brassica napus L.

HUANG Rong, ZHOU Qu-Chen, CHEN Chu-Ming, LUO Qian, YI Dong, DU Chang-Huan, HUANG Xiang-Yu, SHENG Feng(), DU Xue-Zhu()   

  1. School of Life Sciences, Hubei University, Wuhan 430062, Hubei, China
  • Received:2025-05-15 Accepted:2025-09-10 Published:2025-12-12 Published online:2025-09-17
  • Contact: *E-mail: shengfsk@163.com;E-mail: duxuezhusk@163.com
  • Supported by:
    Hubei Provincial Science and Technology Innovation Plan Project(2025BBB012);Science and Technology Innovation 2030-Major Project(2023ZD04070)

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摘要： 硝酸盐转运蛋白2 (NRT2)是一种高亲和硝酸盐转运蛋白, 在植物硝酸盐的吸收和转运过程中发挥重要作用。本研究利用实时荧光定量PCR (qRT-PCR)鉴定到响应低氮胁迫的关键基因BnNRT2.3-like, 对其进行生物信息学分析, 构建BnNRT2.3-like过表达载体, 获得转基因植株, 通过盆栽试验对油菜成熟期的重要农艺性状进行观察和分析, 并在油菜苗期测定其氮素积累量、氮素利用效率、叶绿素含量、氮素利用相关基因表达量以及硝酸还原酶(NR)和谷氨酰胺合成酶(GS)活性。结果表明, BnNRT2.3-like蛋白分子量为61.87 kD, 等电点为9.08, 具有MFS超家族和NNP家族基因的典型特征。与野生型相比, BnNRT2.3-like过表达植株的株高、千粒重和单株籽粒产量分别提高9.1%、20.0%和62.1%, 有效分枝高度、主轴有效长度、主轴有效角果数、全株有效角果数、每角果长度、平均每角果粒数均显著提高。在低氮处理下, 过表达油菜与野生型相比主根长平均增加1.22 cm, 植株鲜重增加0.633 g, 地上部和根部干重分别增加55.0%和13.6%, 单株总干重增加42.1%; 地上部氮积累量较野生型增加17.0%。此外, 在低氮胁迫下, 过表达植株苗期的氮素积累量、氮素利用效率、硝酸还原酶和谷氨酰胺合成酶活性以及叶绿素含量与野生型相比均显著提高, 氮素利用相关基因BnNPF4.6、BnNPF6.3-like和BnAMT1.1a在叶片中表达水平上升而在根中显著降低。因此, 过表达BnNRT2.3-like可以增强油菜对低氮胁迫的耐受性, 提高油菜产量性状, 为提高油菜氮素利用效率提供了种质资源及理论依据。

关键词: 甘蓝型油菜, BnNRT2.3-like, 硝酸盐, 氮素利用率, 产量

Abstract:

Nitrate transporter 2 (NRT2) is a high-affinity nitrate transporter that plays a crucial role in nitrate uptake and translocation in plants. In this study, we identified a key low-nitrogen-responsive gene, BnNRT2.3-like, using real-time quantitative PCR (RT-qPCR). Bioinformatics analysis was conducted, and an overexpression vector of BnNRT2.3-like was constructed to generate transgenic plants. Key agronomic traits of rapeseed at the mature stage were evaluated through a natural pot experiment. At the seedling stage, we assessed nitrogen accumulation, nitrogen use efficiency (NUE), chlorophyll content, expression levels of nitrogen utilization-related genes, and the activities of nitrate reductase (NR) and glutamine synthetase (GS). The results indicated that BnNRT2.3-like protein was found to have a molecular weight of 61.87 kD and an isoelectric point of 9.08, exhibiting typical features of the MFS superfamily and NNP gene family. Compared to the wild type (WT), BnNRT2.3-like overexpression lines showed increases of 9.1% in plant height, 20.0% in thousand-seed weight, and 62.1% in grain yield per plant. Significant improvements were also observed in effective branch height, main inflorescence length, number of siliques per main inflorescence, total siliques per plant, silique length, and average seeds per silique. Under low-nitrogen (LN) conditions, the overexpression lines exhibited an average increase of 1.22 cm in primary root length, a 0.633 g increase in fresh weight, and 55.0% and 13.6% increases in shoot and root dry weights, respectively, resulting in a 42.1% increase in total dry weight per plant. Shoot nitrogen accumulation was also 17.0% higher than in WT. Furthermore, under LN stress at the seedling stage, the overexpression lines demonstrated significant enhancements in nitrogen accumulation, NUE, NR and GS activities, and chlorophyll content compared to WT plants. Notably, expression levels of nitrogen utilization-related genes (BnNPF4.6, BnNPF6.3-like, and BnAMT1.1a) were upregulated in the leaves but markedly downregulated in the roots. Together, these findings demonstrate that BnNRT2.3-like overexpression enhances low-nitrogen tolerance and yield-related traits in rapeseed, providing valuable germplasm resources and a theoretical foundation for improving nitrogen use efficiency in Brassica napus.

Key words: Brassica napus L., BnNRT2.3-like, nitrate, nitrogen use efficiency, yield

表1

本研究所用的引物序列"

引物名称
Primer name		 引物序列
Primer sequence (5'-3')
BnNRT2.3-like-F TCTCGAGCTTTCGCGAGCTCATGGCTTCTAATGAAGAA
BnNRT2.3-like-R AGGTCGACTCTAGAGGATCCTTACGGAAACGTGAAACA
Bn-Kan-F ACTGGGCACAACAGACAATCG
Bn-Kan-R GCATCAGCCATGATGGATACTTT
Q-BnActin7F TCTTCCTCACGCTATCCTCCG
Q-BnActin7R AGCCGTCTCCAGCTCTTGC
Q-BnNRT2.3L-F ATGTTTTTGAGACCGTCTAGCG
Q-BnNRT2.3L-R GGTTCG CATCAGAGTTCCAG
Q-BnAMT1.12-F GAAACTTCTTGGGGCTCAGC
Q-BnAMT1.12-R TGCCGGGTCATATCCATACC
Q-BnNPF4.6-F GTCATTACCGTCGCATGGAG
Q-BnNPF4.6-R TCCCCATTCCCATCCTTTGT
Q-BnNPF6.4-F CGCATACCAATCAGTTCCGG
Q-BnNPF6.4-R CAATGTGACGGTCCATGGTC

图1

BnNRT2基因家族表达模式分析 A: 低氮处理下(0.3 mmol L-1 NO3-) Westar油菜叶片BnNRT2s表达量; B: 低氮处理下(0.3 mmol L-1 NO3-) Westar油菜根部BnNRT2s表达量。"

图2

BnNRT2.3-like基因结构及蛋白序列比对 A: BnNRT2.3-like基因结构示意图; B: BnNRT2.3-like和拟南芥AtNRT2.3蛋白序列比对结果, 其中120至140位氨基酸处的黑色框部分为MFS家族的保守序列(G-x-x-x-D-x-x-G-x-R), 第200位氨基酸处黑色框部分为NPP家族的保守序列(G-W/LG-N-M/A-G)。"

图3

不同物种的NRT2.3基因进化树"

图4

BnNRT2.3-like蛋白跨膜结构模式图"

图5

BnNRT2.3-like转基因株系阳性鉴定及表达量分析 A: T0代阳性转化苗PCR鉴定; B: 对照Westar与转基因株系中BnNRT2.3-like表达量的定量检测。WT: Westar; OE-18、OE-25、OE-36为过表达BnNRT2.3-like的T2代转基因株系。***表示在0.001水平差异显著。"

图6

BnNRT2.3-like过表达油菜表型分析 A: 野生型油菜和过表达BnNRT2.3-like油菜表型; B: 过表达BnNRT2.3-like油菜叶面积, ***表示在0.001水平差异显著。缩写同图5。"

表2

转基因植株与野生型(WT)农艺性状分析"

性状
Character		 WT OE-18 OE-25 OE-36
株高Plant height (cm) 150.43±9.85 b 164.52±6.78 a 166.73±4.71 a 161.30±7.73 ab
一次有效分枝数First effective branch number 6.14±1.36 a 7.80±3.60 a 6.60±2.06 a 7.20±2.13 a
有效分枝高度Effective branch height (cm) 65.86±9.72 a 26.84±2.71 b 26.68±2.26 b 28.70±2.30 b
一次分枝角度First branch angle 33.69±3.79 a 32.26±6.53 a 29.62±6.86 a 36.00±5.83 a
主轴有效长度Effective length of spindle (cm) 49.43±5.48 c 63.00±5.87 b 54.20±4.41 bc 75.60±7.36 a
主轴有效角果数Effective pod number of main shaft 16.43±5.85 b 26.20±2.03 a 23.83±1.83 a 26.80±1.32 a
全株有效角果数Effective pod number of whole plant 191.29±43.33 b 256.20±24.32 a 244.25±27.84 ab 247.62±43.26 ab
每角果长度Length per pod (cm) 4.30±0.93 b 5.40±0.19 a 5.48±0.63 a 5.56±0.35 a
平均每角粒数Average number of grains per pod 21.59±4.68 b 30.69±1.56 a 31.16±2.23 a 30.42±4.15 a
千粒重1000-grain weight (g) 4.40±0.64 b 4.86±0.42 ab 5.64±0.82 a 4.95±0.29 ab
单株籽粒产量Seed yield per plant (g) 19.70±6.45 b 32.88±6.45 a 33.13±5.17 a 29.77±3.71 a

表3

不同氮处理下的油菜主根长及鲜重"

株系
Variety		 主根长
Main shoot length (cm)		 鲜重
Fresh weight (g)
-N +N -N +N
WT 32.27±0.59 c 22.35±0.39 b 1.11±0.14 c 1.88±0.03 a
OE-18 34.62±0.10 a 24.40±0.13 a 1.83±0.08 a 1.91±0.08 a
OE-25 33.63±0.15 b 22.35±0.29 b 1.60±0.07 b 1.85±0.03 a
OE-36 35.23±0.21 a 22.39±0.21 b 1.79±0.07 ab 2.04±0.16 a

表4

不同氮处理下的油菜干重"

株系Variety 地上部
Overground part (g)		 地下部
Underground portion (g)		 单株干重
Total dry weight (g)
-N +N -N +N -N +N
WT 0.227±0.006 c 0.410±0.020 a 0.105±0.005 a 0.089±0.0015 c 0.332±0.011 b 0.499±0.005 b
OE-18 0.355±0.005 a 0.430±0.020 a 0.126±0.090 a 0.090±0.001 bc 0.481±0.009 a 0.522±0.008 a
OE-25 0.345±0.001 b 0.425±0.005 a 0.112±0.002 a 0.098±0.005 a 0.470±0.015 a 0.520±0.005 a
OE-36 0.358±0.002 a 0.410±0.010 a 0.120±0.005 a 0.095±0.002 ab 0.465±0.014 a 0.528±0.010 a

图7

不同氮处理下油菜硝态氮、地上/下部氮含量及氮素利用率 A: 不同氮处理7 d后油菜下部叶片硝态氮含量; B: 不同氮处理7 d后油菜地上部氮素积累量; C: 不同氮处理7 d后油菜根中氮素积累量; D: 不同氮处理7 d后BnNRT2.3-like过表达油菜植株的氮素利用率。***表示在0.001水平差异显著。缩写同图5。-N: 0.3 mmol L-1 N; +N: 6 mmol L-1 N。"

图8

油菜氮代谢相关基因在BnNRT2.3-like过表达油菜中的表达 A: 0.3 mmol L-1 NO3-处理7 d后油菜上部(叶片)氮素利用相关基因的表达量; B: 0.3 mmol L-1 NO3-处理7 d后油菜根部相关基因的表达量。****表示在0.0001水平差异显著。缩写同图5。"

图9

不同氮处理下油菜叶片NR和GS活性 A: 不同氮处理7 d后油菜下部叶片硝酸还原酶活性; B: 不同氮处理7 d后油菜下部叶片谷氨酰胺合成酶活性。****表示过表达材料与WT间在0.0001水平差异显著。缩写同图5。-N: 0.3 mmol L-1 N; +N: 6 mmol L-1 N。"

图10

不同氮处理下油菜叶绿素含量 A: 6 mmol L-1 NO3-处理7 d后油菜下部叶片叶绿素含量; B: 0.3 mmol L-1 NO3-处理7 d后油菜下部叶片叶绿素含量。**、***、****分别表示过表达材料与WT在0.01、0.001和0.0001概率水平差异显著。缩写同图5。"

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