水稻氮高效相关基因的研究进展

doi:10.3864/j.issn.0578-1752.2022.08.001

摘要/Abstract

摘要：

提高氮肥利用率,降低农业生产中的氮肥投入,对实现中国提出的“化肥零增长”目标至关重要。氮素可通过植物根部以硝态氮或者铵态氮的形式从土壤中被吸收,并在植物体内转运合成氨基酸、核苷酸等生命必需物质,是农作物生长和产量形成的必要性元素。然而,氮肥过量则会破坏土壤理化性质,导致土壤盐碱化、生态环境污染;减少氮肥用量难以发挥当前水稻、小麦等大田农作物品种的高产潜力,极大地制约了农作物的产量,同时也会过度消耗土壤中的氮素,威胁中国的粮食安全。为提高氮肥利用效率,确保粮食安全,挖掘氮高效基因（如NRT1.1B、OsGRF4等）,通过分子设计育种对当前现有品种进行遗传改良,有助于培育高效利用氮素的水稻新品系,挖掘当前水稻品种的生产潜力,提升中国农业可持续绿色发展水平。本文基于目前水稻研究中挖掘的氮高效基因,根据其功能和特点将其分为四类：NRT/PTR类、AMT类（铵转运蛋白家族）、NLP家族类及其他种类;并对具有潜在育种价值的氮高效基因的利用及当前存在的问题进行分析与展望。

关键词: 氮, 水稻, 氮肥, 氮高效基因

Abstract:

Over the last few years, China government has put forward a strategy to achieve the goal of “zero growth of chemical fertilizer”. It is particularly important to reduce the input of nitrogen fertilizer in agricultural production and enhance the nitrogen use efficiency in crops. Nitrogen is mostly absorbed from the soil by plant roots in the form of nitrate nitrogen (NO₃^-) and ammonium nitrogen (NH₄^-). It is transported from roots in plants to synthesize essential life substances, such as amino acids and nucleotides. Nitrogen is used as a basic element for crop growth and yield formation. However, excessive application of nitrogen fertilizer destroys the physical and chemical properties of the soil, causes undesirable changes to soil salinization, and pollutes the ecological environment, and pollutes the ecological environment. By reducing the quantity of nitrogen fertilizer, will destabilize the yield potential of field crops including rice and wheat which is being used as a staple food in China. It can threaten food security of the country. To improve the nitrogen use efficiency (NUE) and stabilize the food security, mining nitrogen-efficient genes, such as NRT1.1B, OsGRF4 etc., genetic improvement of current existing varieties through molecular design breeding will help to cultivate new nitrogen efficiently rice lines. Tapping the productive potential of current rice varieties will improve the level of sustainable agricultural development in our country. In this article, based on the nitrogen-efficient genes excavated in the current rice research, this article reviews the PTR (polypeptide transporter) family, NRT (nitrate transporter) family, AMT family (ammonium transporter family), NLP family and other types of rice nitrogen-efficient genes. The future prospects of gene utilization have been prospected. Based on the nitrogen-efficient genes excavated in the current rice research, they are divided into four categories: NRT/PTR, AMT (ammonium transporter), NLP and other types, and summarize their functions and characteristics, and analysis the utilization prospect and existing problems of nitrogen-efficient genes with potential breeding value.

Key words: nitrogen, rice, nitrogen fertilizer, nitrogen efficient gene

桑世飞, 曹梦雨, 王亚男, 王君怡, 孙晓涵, 张文玲, 姬生栋. 水稻氮高效相关基因的研究进展[J]. 中国农业科学, 2022, 55(8): 1479-1491.

SANG ShiFei, CAO MengYu, WANG YaNan, WANG JunYi, SUN XiaoHan, ZHANG WenLing, JI ShengDong. Research Progress of Nitrogen Efficiency Related Genes in Rice[J]. Scientia Agricultura Sinica, 2022, 55(8): 1479-1491.

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图/表 7

表1

NPF家族的基因种类及特征"

基因名称 Gene name	基因登录号 Gene accession No.	功能 Features	验证方法 Authentication method	参考文献 References
OsNPF2.2/OsPTR2	LOC_Os12g44100	低亲和力硝酸盐转运蛋白,参与从根到茎的硝酸盐转运和维管发育 Low-affinity nitrate transporter, involved in nitrate transport from root to stem and vascular development	T-DNA插入突变体 T-DNA insertion mutant	[12,16]
OsNPF2.4	LOC_Os03g48180	pH依赖性低亲和力硝酸盐转运蛋白,参与获取和长距离硝酸盐运输 pH-dependent low-affinity nitrate transporter, involved in acquisition and long-distance nitrate transport	T-DNA插入突变体,过表达 T-DNA insertion mutant, overexpression	[17]
OsNPF4.5	LOC_Os01g54515	低亲和硝酸盐转运蛋白,增强表达显著提高氮素吸收效率并促进水稻生长 Low-affinity nitrate transporter, enhanced expression, significantly improves nitrogen absorption efficiency and promotes rice growth	敲除突变体 Knockout mutant	[18]
OsNPF6.1	LOC_Os01g01360	编码硝酸盐转运蛋白,可能直接参与硝酸盐的吸收和再分配 Encodes a nitrate transporter, which may be directly involved in the absorption and redistribution of nitrate	T-DNA插入突变体,过表达 T-DNA insertion mutant, overexpression	[23]
OsNRT1	LOC_Os03g13274	低亲和力硝酸盐转运蛋白,增加氮积累 Low-affinity nitrate transporter, increasing nitrogen accumulation	突变体 Mutant	[28]
OsNPF6.3/OsNRT1.1A	LOC_Os08g05910	定位于液泡膜的硝酸盐转运蛋白,参与调节氮素利用 Nitrate transporter located in the vacuole membrane, involved in regulating nitrogen utilization	过表达 Overexpression	[29]
OsNPF6.5/OsNRT1.1B	LOC_Os10g40600	通过改变根际微环境影响水稻籼粳亚种间的氮肥利用效率 Influence of NUE between indica and japonica rice subspecies by changing the rhizosphere microenvironment	过表达 Overexpression	[30]
OsNPF7.2	LOC_Os02g47090	低浓度硝酸盐转运蛋白,参与根细胞中硝酸盐的分配 Low-concentration nitrate transporter, involved in the distribution of nitrate in root cells	敲除突变体 Knockout mutant	[19]
OsNPF7.3/OsPTR6	LOC_Os04g50950	高表达可提高水稻的氮素利用效率 High expression can improve the NUE of rice	过表达 Overexpression	[20,31 -32]
OsNPF7.7/OsPTR10	LOC_Os10g42870	调节枝条的分支和氮素的利用效率 Regulates the branching of branches and NUE	过表达 Overexpression	[33]
OsNPF8.1/OsPTR7	LOC_Os01g04950	参与了稻谷中砷酸二甲酯的积累 Participated in the accumulation of dimethyl arsenate in rice	MiRNA	[34]
OsNRT8.2/OsPTR1	LOC_Os07g01070	干旱胁迫诱导 Drought and Salt Stress induced	表达分析 Expression patterns	[26]
OsNPF8.20/OsPTR9	LOC_Os06g49250	正调节铵吸收、侧根形成和籽粒产量 Positive regulation of ammonium absorption, lateral root formation and grain yield	T-DNA插入突变体,RNAi T-DNA insertion mutant, RNAi	[27]

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基因名称 Gene name	基因登录号 Gene accession No.	功能 Features	验证方法 Authentication method	参考文献 References
OsNRT2.4	LOC_Os01g36720	双重亲和力硝酸盐转运蛋白 Dual affinity nitrate transporter	敲除突变体 Knockout mutant	[38]
OsNRT2.1	LOC_Os02g02170	高亲和力硝酸盐转运蛋白 High affinity nitrate transporter	过表达 Overexpression	[39,41⇓ -43]
OsNRT2.2	LOC_Os02g02190	与OsNAR2.1相互作用以进行高亲和力的硝酸盐转运 Interacts with OsNAR2.1 for high-affinity nitrate transport	过表达 Overexpression	[40,44]
OsNRT2.3a	LOC_Os01g50820	参与根到茎的远距离硝态氮运输 Participate in long-distance nitrate nitrogen transport from root to stem	过表达 Overexpression	[37,40]
OsNRT2.3b	LOC_Os01g50820	正向调节pH缓冲和氮素利用效率 Positive adjustment of pH buffer and nitrogen utilization efficiency	过表达 Overexpression	[31]

基因名称 Gene name	基因登录号 Gene accession No.	功能 Features	验证方法 Authentication method	引用文献 References
OsAMT1.1	LOC_Os04g43070	铵吸收和铵钾稳态 Ammonium absorption and ammonium potassium homeostasis	过表达 Overexpression	[47⇓-49,57]
OsAMT1.2	LOC_Os02g40730	功能性铵转运体 Functional ammonium transporter	突变体 Mutant	[49,58]
OsAMT1.3	LOC_Os02g40710	高亲和力铵转运蛋白,参与根系形态和碳氮代谢 High affinity ammonium transporter, involved in root morphology and carbon and nitrogen metabolism	过表达 Overexpression	[50-51]
OsAMT2.1	LOC_Os05g39240	功能性铵转运蛋白,组成型表达 Functional ammonium transporter, constitutively expressed	突变体 Mutant	[54]
OsAMT2.2	LOC_Os01g61510	铵转运蛋白 Ammonium transporter	突变体 Mutant	[59]
OsAMT2.3	LOC_Os01g61550	铵转运蛋白 Ammonium transporter	突变体 Mutant	[59]
OsAMT3.1	LOC_Os01g65000	介导菌根铵转移 Mediated Mycorrhizal Ammonium Transfer	突变体 Mutant	[56,60]
OsAMT3.2	LOC_Os03g62200	铵转运蛋白 Ammonium transporter	突变体 Mutant	[59]
OsAMT3.3	LOC_Os02g34580	铵转运蛋白 Ammonium transporter	突变体 Mutant	[59]
OsAMT4	LOC_Os03g53780	铵转运蛋白 Ammonium transporter	突变体 Mutant	[59]
OsAMT5.1	LOC_Os12g01420	铵转运蛋白 Ammonium transporter	突变体 Mutant	[59]
OsAMT5.2	LOC_Os11g01410	铵转运蛋白 Ammonium transporter	突变体 Mutant	[59]

基因名称 Gene name	基因登录号 Gene accession No.	功能 Features	验证方法 Authentication method	备注 Remark	引用文献 References
OsNLP1	LOC_Os03g03900	初级硝酸盐反应中具有潜在作用 Potential role in primary nitrate reactions	过表达 Overexpression	无 None	[62]
OsNLP2	LOC_Os04g41850	无 None	无 None	无 None	[60]
OsNLP3	LOC_Os01g13540	硝酸盐信号传导的核心转录因子 Core transcription factor for nitrate signaling	转基因系 Transgenic lines	无 None	[61]
OsNLP4	LOC_Os09g37710	通过结合启动子上的顺式元件调控N的吸收同化 Regulate N uptake and assimilation by binding the cis-elements on the promoter	过表达 Overexpression	与野生型相比,过表达系产量提高30%,氮素利用率提高47% Compared with the wild type, the yield of the overexpression line is increased by 30%, and the nitrogen utilization rate is increased by 47%	[63]
OsNLP5	LOC_Os11g16290	无 None	无 None	无 None	[60]
OsNLP6	LOC_Os02g04340	无 None	无 None	无 None	[60]

基因名称 Gene name	基因登录号 Gene accession No.	功能 Features	验证方法 Authentication method	引用文献 References
OsMADS25	LOC_Os04g23910	正调控硝酸盐转运蛋白基因表达 Positive regulation of nitrate transporter gene expression	过表达 Overexpression	[64]
OsNAR2.1	LOC_Os02g38230	能与OsNRT2.1/2.2和OsNRT2.3a互作 Interacts with OsNRT2.1/2.2 and OsNRT2.3a	RNAi	[40,44]
OsGRF4	LOC_Os02g47280	正调控植物碳-氮代谢,促进氮素吸收、同化和转运 Positively regulate plant carbon-nitrogen metabolism, promote nitrogen absorption, assimilation and transport	突变体 Mutant	[67]
TOND1	LOC_Os12g43440	过表达TOND1能增加水稻对低氮的耐性 Overexpression of TOND1 can increase the tolerance of rice to low nitrogen	突变体 Mutant	[65]
OsGOGAT1	LOC_Os01g48960	与OsAMT1.2同时激活,提高氮素利用效率 Activate simultaneously with OsAMT1.2 to improve NUE	突变体 Mutant	[70]
Ghd7	LOC_Os07g15770	抑制ARE1的表达,调节氮素利用效率 Inhibit the expression of ARE1 and regulate NUE	突变体 Mutant	[69]
OsTCP19	LOC_Os06g12230	OsTCP19-H有提高氮肥利用率的潜力 OsTCP19-H has the potential to improve nitrogen fertilizer utilization	突变体 Mutant	[71]
qNGR9/DEP1	LOC_Os09g26999	株高对氮的响应 Response of plant height to nitrogen	定位克隆与遗传互补 Positional cloning and genetic complementation	[72]