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作物学报 ›› 2024, Vol. 50 ›› Issue (2): 414-424.doi: 10.3724/SP.J.1006.2024.32015

• 耕作栽培·生理生化 • 上一篇    下一篇

低氮条件下超级杂交稻苗期根系特征的变化及与其高氮素积累的关系

吴宇**(), 刘磊**(), 崔克辉*(), 齐晓丽, 黄见良, 彭少兵   

  1. 作物遗传改良全国重点实验室 / 农业农村部长江中游作物生理生态与耕作重点实验室 / 华中农业大学植物科学技术学院, 湖北武汉 430070
  • 收稿日期:2023-04-24 接受日期:2023-09-13 出版日期:2024-02-12 网络出版日期:2023-10-07
  • 通讯作者: *崔克辉, E-mail: cuikehui@mail.hzau.edu.cn, Tel: 027-87288380
  • 作者简介:吴宇, E-mail: 1172608346@qq.com;刘磊, E-mail: 18790091496@163.com
    **同等贡献
  • 基金资助:
    国家自然科学基金项目(31671598)

Changes of root characteristics of super hybrid rice variety contributing to high nitrogen accumulation under low nitrogen application at seedling stage

WU Yu**(), LIU Lei**(), CUI Ke-Hui*(), QI Xiao-Li, HUANG Jian-Liang, PENG Shao-Bing   

  1. National Key Laboratory of Crop Genetic Improvement / Key Laboratory of Crop Ecophysiology and Farming System in the Middle Reaches of the Yangtze River, Ministry of Agricultural and Rural Affairs / College of Plant Science and Technology, Huazhong Agricultural University, Wuhan 430070, Hubei, China
  • Received:2023-04-24 Accepted:2023-09-13 Published:2024-02-12 Published online:2023-10-07
  • Contact: *E-mail: cuikehui@mail.hzau.edu.cn, Tel: 027-87288380
  • About author:**Contributed equally to this work
  • Supported by:
    National Natural Science Foundation of China(31671598)

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摘要：

作物根系在氮素吸收与分配中起着重要作用。本研究选用2个根系特征具有差异的水稻品种黄华占(Huanghuazhan, HHZ, 常规稻)和扬两优6号(Yangliangyou 6, YLY6, 超级杂交稻), 在水培条件和2个氮素浓度(低氮, low nitrogen, LN; 高氮, high nitrogen, HN)处理下, 研究根系形态和解剖结构特征的变化及其与LN下氮素积累的关系。与HN处理相比, LN处理下HHZ总氮积累量显著下降了19.7%, YLY6没有显著下降; YLY6根干重、总根长、根表面积、根尖数分别显著增加了41.3%、57.1%、74.9%和20.6%, 而HHZ没有显著变化。与HN处理相比, LN处理下YLY6的根直径和根皮层面积分别显著增加了12.4%和24.2%, 而HHZ根直径和根皮层面积、中柱直径分别显著降低了12.0%、21.9%和11.4%。LN处理下, YLY6根系铵转运蛋白基因AMT2;1、AMT2;3、AMT3;1、AMT3;2的表达量分别显著增加了195.6%、29.3%、314.9%和388.9%, 谷氨酰胺合成酶基因GS1;1表达量增加了158.2%, HHZ中这5个基因的表达量没有显著变化。LN下总氮积累量与根系性状显著正相关(根厚壁组织厚度除外), 与部分铵转运和同化相关基因的表达量显著正相关。这些结果表明超级杂交稻YLY6幼苗在LN下根系特征变化有利于增加氮积累。选育根系对减氮表现出有利变化的水稻品种应是绿色水稻生产的一个重要途径。

关键词: 水稻, 低氮, 根系特征变化, 氮积累, 铵转运蛋白, 谷氨酰胺合成酶

Abstract:

Crop root system plays an important role in nitrogen uptake. In this study, two rice varieties, conventional rice variety Huanghuazhan (HHZ) and super hybrid rice variety Yangliangyou 6 (YLY6), were grew hydroponically under high nitrogen (HN) and low nitrogen (LN) treatments to investigate the changes of root characteristics and their relationships with nitrogen accumulation at seedling stage. Compared to HN, LN significantly decreased the total N accumulation in HHZ by 19.7% and had no substantial effect in YLY6. Under LN, root dry weight, the total root length, root surface area, and root tips in YLY6 significantly increased by 41.3%, 57.1%, 74.9%, and 20.6%, respectively. However, these four morphological parameters had no significant change in HHZ. Under LN, root diameter and root cortical area in YLY6 significantly increased by 12.4% and 24.2%, respectively. However, the two parameters and root stele diameter in HHZ significantly reduced by 12.0%, 21.9%, and 11.4%, respectively. In YLY6, LN significantly up-regulated the relative expression level of root ammonium transporter genes (AMT2;1, AMT2;3, AMT3;1, and AMT3;2) by 195.6%, 29.3%, 314.9%, and 388.9%, respectively, and increased the relative expression level of glutamine synthetase gene GS1;1 by 158.2%. However, LN had no effect on the relative expression level of the five genes in HHZ. Total nitrogen accumulation was significantly and positively correlated with the root characteristics (but thickness of root cortical sclerenchyma) and expression of above-mentioned genes under LN. These results indicated that the enhanced positive responses of above-mentioned root characteristics contributed to high nitrogen accumulation in YLY6 seedlings under LN. Developing varieties with root positive responses to nitrogen reduction should be a considerable target for green rice production.

Key words: rice (Oryza sativa L.), low nitrogen application, changes of root characteristics, nitrogen accumulation, ammonium transporter, glutamine synthetase

表1

AMT、GS、GOGAT和Actin基因扩增引物序列"

基因
Gene name		 上游引物
Forward sequences (5'-3')		 下游引物
Reverse sequences (5'-3')
OsAMT1.1 TTTTGCTGGGCTTCTCTTGT ACCATTCCACCACACCCTTA
OsAMT1.2 CTTCATCGGGAAGCAGTTCT TGAGGAAGGCGGAGTAGATG
OsAMT1.3 CGGCTTCGACTACAGCTTCT GACCAGATCCAGTGGGACAC
OsAMT2.1 CTGGCTCCTCCTCTCCTACA CAGGATGTTGTTCGGTGAGA
OsAMT2.2 GCCTCGACGTCATCTTCTTC TTGTGGAGGATCATCATGGA
OsAMT2.3 GCCTCGACGTCATCTTCTTC GGAAGGTGGATTTCTTGTGC
OsAMT3.1 ACCAAGGACAGGGAGAGGTT AAGATGACGTCGAGGCAA
OsAMT3.2 GCACAGAAGGACAGGGAGAG GCAGATGTTGGTGTTGAGGA
OsAMT3.3 CGAGCATCACCATCATCATC ATGACACCCCACTGGAAGAG
OsAMT4 CTGGCCTCAAGAAGATGGACA AGCTGCTTCACGTACTTGATCG
NADH-GOGAT1 GTGCAGCCTGTTGCAGCATAAA CGGCATTTCACCATGCAAATC
NADH-GOGAT2 CCTGTCGAAGGATGATGAAGGTGAAACC TGCATGGCCCTACTATCTTCGCATCA
Fd-GOGAT AAACAGGCAGCGAGAAAGGTG AAACTCGGCACAAGCTTCAGG
GS1.1 GAGTCGTCGTCTCATTTGACCC GTAGCCACCATCGTTCCTCATC
GS1.2 TTTTCAAGGACCCGTTCAGGA CGGCACTGTGCCTCTTGTTAGT
GS1.3 TCAAGCCATCTTCAGAGACCCA TACCGGTTGTTCGTCGGAATC
GS2 TCACTTCGCCATGACTTGCA CCCCATGAGAAATTGTCAATGC
Actin ATGAAGATCAAGGTGGTCGC GATCTCAGCCTTGGCAATCC

表2

低氮处理对水稻生物量和氮积累的影响"

品种
Variety		 氮处理
Nitrogen
treatment		 根干重
Root dry weight
(g plant-1)		 地上部分干重
Shoot dry weight
(g plant-1)		 根氮含量
Root N content
(mg g-1)		 地上部分氮含量
Shoot N content
(mg g-1)		 总氮积累量
Total N accumulation
(mg plant-1)
HHZ HN 0.192±0.019 a 0.683±0.070 a 29.47±1.42 a 43.63±1.93 a 35.36±2.71 a
LN 0.230±0.026 a 0.672±0.052 a 21.63±2.25 b 34.92±1.06 b 28.40±1.59 b
YLY6 HN 0.208±0.017 b 1.017±0.030 b* 30.98±0.83 a 47.42±1.89 a 54.70±3.73 a*
LN 0.415±0.027 a* 1.388±0.158 a* 19.38±0.48 b 32.24±1.33 b 52.64±3.82 a*
ANOVA 氮Nitrogen (N) +++ ++ ++ +++ +
品种Variety (V) +++ +++ ns ns +++
氮×品种N×V ++ ++ + + ns

表3

低氮处理对根形态特征的影响"

品种
Variety		 氮处理
Nitrogen
treatment		 总根长
Total root length
(cm plant-1)		 根表面积
Root surface area
(cm2 plant-1)		 根体积
Root volume
(cm3 plant-1)		 根尖数
Root tips
(No. plant-1)
HHZ HN 1822±269 a 181±20 a 1.42±0.11 b 26,678±5793 a
LN 1809±159 a 216±16 a 2.09±0.21 a 22,397±2879 a
YLY6 HN 2321±221 b 228±19 b 1.79±0.16 b 34,354±4079 b
LN 3280±256 a* 358±29 a* 3.12±0.28 a* 41,429±3964 a*
ANOVA 氮Nitrogen (N) + ++ ++ ns
品种Variety (V) +++ +++ +++ ++
氮×品种N×V ++ ++ ++ +

表4

低氮处理对根解剖特征的影响"

品种
Variety		 氮处理
Nitrogen
treatment		 根直径
Root diameter
(mm)		 通气组织面积比例
Proportion of root cortical aerenchyma area (%)		 根皮层面积
Root cortical area
(mm2)		 根皮层厚壁组织厚度
Thickness of root
cortical sclerenchyma (mm)		 根中柱直径
Root stele diameter (mm)
HHZ HN 989±33 a 0.62±0.53 b 0.64±0.04 a 6.50±0.26 a 228±8 a
LN 870±41 b 5.86±3.13 a 0.50±0.05 b 6.01±0.42 b 202±4 b
YLY6 HN 883±78 b 6.08±5.11 b* 0.52±0.09 b 8.26±0.38 a* 215±6 a
LN 993±25 a* 27.69±2.49 a* 0.64±0.04 a* 6.18±0.28 b 221±4 a*
ANOVA 氮Nitrogen (N) ns ++ ns ++ +
品种Variety (V) ns +++ ns ++ ns
氮×品种N×V ++ ++ + ++ ++

图1

低氮处理对根铵转运基因表达量的影响 HHZ为黄华占, YLY6为扬两优6号。HN为高氮处理, LN为低氮处理。数据为平均值±标准差(STDEV); 不同小写字母表示同一品种不同氮处理在0.05概率水平差异显著(LSD test); *表示同一氮处理下HHZ和YLY6在0.05概率水平差异显著(LSD检验)。"

图2

低氮处理对根谷氨酸合酶(GOGAT)和谷氨酰胺合成酶(GS)基因表达量的影响 HHZ为黄华占, YLY6为扬两优6号。HN为高氮处理, LN为低氮处理。数据为平均值±标准差(STDEV); 不同小写字母表示同一品种不同氮处理在0.05概率水平差异显著(LSD test); *表示同一氮处理下HHZ和YLY6在0.05概率水平差异显著(LSD检验)。"

表5

总氮积累与根特征的相关性分析"

氮处理
Nitrogen
treatment		 根干重
RDW		 总根长
TRL		 根表面积
RS		 根体积
RV		 根尖数
RT		 根直径
RD		 通气组织
比例
RCA		 根皮层面积
CCA		 根皮层厚壁
组织厚度
SCT		 根中柱
直径
RSD
高氮处理HN 0.61 0.68 0.70 0.71* 0.57 -0.70 0.65 -0.70 0.94*** -0.70
低氮处理LN 0.99*** 0.96*** 0.95*** 0.92** 0.94*** 0.92** 0.96*** 0.91** 0.37 0.97***

表6

总氮积累与铵转运基因表达量的相关性分析"

氮处理Nitrogen treatment AMT1;1 AMT1;2 AMT1;3 AMT2;1 AMT2;2 AMT2;3 AMT3;1 AMT3;2 AMT3;3 AMT4
高氮处理HN 0.75* 0.11 -0.48 -0.31 -0.82* 0.31 0.26 0.05 0.63 0.44
低氮处理LN 0.89** 0.27 0.71 0.68 -0.79* 0.79* 0.86** 0.85** 0.39 0.48

表7

总氮积累与铵同化相关基因表达量的相关性分析"

氮处理
Nitrogen treatment		 GS1;1 GS1;2 GS1;3 GS2 NADH-GOGAT1 NADH-GOGAT2 Fd-GOGAT
高氮处理HN -0.17 0.08 -0.42 -0.93*** -0.05 -0.84** 0.54
低氮处理LN 0.77* 0.50 0.73* -0.76* 0.61 -0.55 0.92**
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