植物铵态氮同化及其调控机制的研究进展

doi:10.11983/CBB15077

摘要/Abstract

摘要： 氮是维持植物生长发育最重要的矿质营养元素之一, 在植物整个生命进程中发挥着重要作用。在植物体内, 氮同化既是植物利用氮素的一个中心环节, 也是导致植物氮利用效率不高的因素之一。氮同化主要分为硝态氮(NO₃^-)和铵态氮(NH₄⁺)同化, 其中铵态氮同化是氮同化中最为关键的一步。按照不同来源, 植物体内铵态氮同化又可分为一次同化和二次同化, 但两者都是通过谷氨酰胺/谷氨酸合成酶(GS/GOGAT)途径进行。植物铵态氮同化不仅需要大量的能量, 而且需要大量的碳源, 所以其在转录、转录后以及翻译后等各个水平上都受到严格调控。该文综述了目前关于植物铵态氮同化及其调控机制的最新研究进展。

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关键词: 铵态氮, 一次同化, 二次同化, GS/GOGAT, 调控机制

Abstract: Nitrogen is one of the most important mineral nutrient elements for plant growth and development, playing an essential role in the whole process of plant life. Nitrogen assimilation is a central link for plants to utilize nitrogen and also a factor in low nitrogen use efficiency in plants. It includes two types: assimilation of nitrate (NO₃^-) and ammonium (NH₄⁺); the latter is the critical step in the process of nitrogen assimilation. The source of NH₄⁺ during ammonium assimilation can be divided into 2 types—primary assimilation and secondary assimilation—but both proceed in the glutamine/glutamate (GS/GOGAT) pathway. Ammonium assimilation requires much energy resources but also consumes abundant carbon skeletons, so it is strictly regulated at different levels including transcription, post-transcription, and post-translation. We review the current progress in study of ammonium assimilation and its regulatory mechanism in plants.

Key words: ammonium, primary assimilation, secondary assimilation, GS/GOGAT, regulatory mechanism

徐晓鹏, 傅向东, 廖红. 植物铵态氮同化及其调控机制的研究进展. 植物学报, 2016, 51(2): 152-166.

Xiaopeng Xu, Xiangdong Fu, Hong Liao. Advances in Study of Ammonium Assimilation and its Regulatory Mechanism in Plants. Chinese Bulletin of Botany, 2016, 51(2): 152-166.

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https://www.chinbullbotany.com/CN/Y2016/V51/I2/152

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物种	基因名称	表达部位	基因主要功能
水稻 (Oryza sativa)	OsGS1;1	维管束特异定位(根部、茎和叶片)	参与水稻衰老过程中氮再利用过程, 为籽粒发育提供氮源^b
	OsGS1;2	根系表皮、皮层细胞, 茎, 叶鞘	水稻根部同化NH₄⁺的一个关键基因^a
	OsGS1;3	种子	可能主要参与种子发育时氮的供应或种子萌发时氮的再利用过程^b
	OsNADH-GOGAT1	根系表皮、皮层细胞, 幼嫩叶片维管束	与OsGS1;2一起参与水稻根部NH₄⁺的同化^a
	OsNADH-GOGAT2	成熟叶片维管束	为OsGS1;1反应提供底物Glu, 参与水稻氮的再利用过程^b
拟南芥 (Arabidopsis thaliana)	AtGln1;1	根(表皮和根尖)	同化根部吸收的NH₄⁺; 对根构型具有显著影响, 突变体主根生长受抑制^a
拟南芥 (Arabidopsis thaliana)	AtGln1;2	维管组织中(根、叶片、萼片、花瓣和雄蕊)	参与根系和叶片缓解氨毒^a; 参与地上部同化由NO₃^-还原产生的NH₄⁺(供应高浓度NO₃^-时)^a; 参与为种子萌发和种子发育提供氮源的过程^b
	AtGln1;3	根部维管组织	未知
	AtGln1;4	侧根形成区的中柱鞘细胞	可能在缺氮条件下参与同化根部吸收的NH₄⁺或由NO₃^-还原产生的NH₄^+a
	AtNADH-GOGAT	根、花粉、柱头和顶端分生组织	负责拟南芥根部NH₄⁺的同化^a
玉米	ZmGln1;1	根表皮、皮层细胞	参与同化根系中由NO₃^-还原产生的NH₄^+a
(Zea mays)	ZmGln1;2	根系和叶片韧皮部	参与玉米体内氮的再利用及转运^b
	ZmGln1;3	叶肉细胞	同化叶肉细胞中由NO₃^-还原产生的NH₄⁺, 对玉米籽粒数目有显著影响^a
	ZmGln1;4	叶片维管束鞘细胞	参与重新同化叶片产生的NH₄⁺, 对玉米籽粒大小有显著影响^b
大麦 (Hordeum	HvGS1;1	维管束特异定位(根部、茎和叶片)	可能与OsGS1;1具有类似的功能, 在氮再利用方面起重要作用^b
vulgare)	HvGS1;2	叶肉细胞, 根部的皮层和中柱鞘细胞	可能主要参与同化叶肉细胞中由NO₃^-还原产生的NH₄^+a
	HvGS1;3	谷粒, 根部	可能参与谷粒发育中氮源的供应^b; 参与根系缓解NH₄⁺的毒害^a
苜蓿 (Medicago truncatula)	MtGS1a	维管束定位(根瘤、根、茎和叶)	同化侵染细胞中类菌体释放到细胞质中的NH₄⁺(来源于固氮酶固定的氮)^a; 提供体内氮运转的来源^b
苜蓿 (Medicago truncatula)	MtGS1b	根瘤、根、茎和叶	可能参与同化根系吸收的氮^a
	MtGS2b	籽粒	可能为籽粒发育提供氮源, 表达量随着籽粒的发育而升高^b
大豆 (Glycine max)	GmGlnα	子叶、新根、发育中的根瘤和花	未知
	GmGlnβ(β1和β2)	组成型表达, 根瘤中较高	可能参与同化根瘤固氮产生的NH₄^+a
	GmGlnγ1	根瘤特异	未知
	GmGlnγ2	根瘤增强, 子叶和花	未知

物种	基因名称	表达部位	基因主要功能
水稻 (Oryza sativa)	OsGS1;1	维管束特异定位(根部、茎和叶片)	参与水稻衰老过程中氮再利用过程, 为籽粒发育提供氮源^b
	OsGS1;2	根系表皮、皮层细胞, 茎, 叶鞘	水稻根部同化NH₄⁺的一个关键基因^a
	OsGS1;3	种子	可能主要参与种子发育时氮的供应或种子萌发时氮的再利用过程^b
	OsNADH-GOGAT1	根系表皮、皮层细胞, 幼嫩叶片维管束	与OsGS1;2一起参与水稻根部NH₄⁺的同化^a
	OsNADH-GOGAT2	成熟叶片维管束	为OsGS1;1反应提供底物Glu, 参与水稻氮的再利用过程^b
拟南芥 (Arabidopsis thaliana)	AtGln1;1	根(表皮和根尖)	同化根部吸收的NH₄⁺; 对根构型具有显著影响, 突变体主根生长受抑制^a
拟南芥 (Arabidopsis thaliana)	AtGln1;2	维管组织中(根、叶片、萼片、花瓣和雄蕊)	参与根系和叶片缓解氨毒^a; 参与地上部同化由NO₃^-还原产生的NH₄⁺(供应高浓度NO₃^-时)^a; 参与为种子萌发和种子发育提供氮源的过程^b
	AtGln1;3	根部维管组织	未知
	AtGln1;4	侧根形成区的中柱鞘细胞	可能在缺氮条件下参与同化根部吸收的NH₄⁺或由NO₃^-还原产生的NH₄^+a
	AtNADH-GOGAT	根、花粉、柱头和顶端分生组织	负责拟南芥根部NH₄⁺的同化^a
玉米	ZmGln1;1	根表皮、皮层细胞	参与同化根系中由NO₃^-还原产生的NH₄^+a
(Zea mays)	ZmGln1;2	根系和叶片韧皮部	参与玉米体内氮的再利用及转运^b
	ZmGln1;3	叶肉细胞	同化叶肉细胞中由NO₃^-还原产生的NH₄⁺, 对玉米籽粒数目有显著影响^a
	ZmGln1;4	叶片维管束鞘细胞	参与重新同化叶片产生的NH₄⁺, 对玉米籽粒大小有显著影响^b
大麦 (Hordeum	HvGS1;1	维管束特异定位(根部、茎和叶片)	可能与OsGS1;1具有类似的功能, 在氮再利用方面起重要作用^b
vulgare)	HvGS1;2	叶肉细胞, 根部的皮层和中柱鞘细胞	可能主要参与同化叶肉细胞中由NO₃^-还原产生的NH₄^+a
	HvGS1;3	谷粒, 根部	可能参与谷粒发育中氮源的供应^b; 参与根系缓解NH₄⁺的毒害^a
苜蓿 (Medicago truncatula)	MtGS1a	维管束定位(根瘤、根、茎和叶)	同化侵染细胞中类菌体释放到细胞质中的NH₄⁺(来源于固氮酶固定的氮)^a; 提供体内氮运转的来源^b
苜蓿 (Medicago truncatula)	MtGS1b	根瘤、根、茎和叶	可能参与同化根系吸收的氮^a
	MtGS2b	籽粒	可能为籽粒发育提供氮源, 表达量随着籽粒的发育而升高^b
大豆 (Glycine max)	GmGlnα	子叶、新根、发育中的根瘤和花	未知
	GmGlnβ(β1和β2)	组成型表达, 根瘤中较高	可能参与同化根瘤固氮产生的NH₄^+a
	GmGlnγ1	根瘤特异	未知
	GmGlnγ2	根瘤增强, 子叶和花	未知