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[植物氮代谢及其环境调控的研究进展]

[Research advance in nitrogen metabolism of plant and its environmental regulation].

作者信息

Xu Zhenzhu, Zhou Guangsheng

机构信息

Laboratory of Quantitative Vegetation Ecology, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China.

出版信息

Ying Yong Sheng Tai Xue Bao. 2004 Mar;15(3):511-6.

Abstract

Nitrogen metabolism is not only one of the basic processes of plant physiology, but also one of the important parts of global chemical cycle. Plant nitrogen assimilation directly takes part in the synthesis and conversion of amino acid through the reduction of nitrate. During this stage, some key enzymes, e.g., nitrate reductase (NR), glutamine synthetase (GS), glutamate dehydrogenase (GDH), glutamine synthase (GOGAT), aspargine synthetase (AS), and asparate aminotransferase (AspAT) participate these processes. The protein is assimilated in plant cell through amino acid, and becomes a part of plant organism through modifying, classifying, transporting and storing processes, etc. The nitrogen metabolism is associated with carbonic metabolism through key enzyme regulations and the conversion of products, which consists of basic life process. Among these amino acids in plant cell, glutamic acid (Glu), glutamine (Gln), aspartic acid (Asp) and asparagines (Asn), etc., play a key role, which regulates their conversion each other and their contents in the plant cell through regulating formation and activity of those key enzymes. Environmental factors also affect the conversion and recycle of the key amino acids through regulating gene expression of the key enzymes and their activities. Nitrate and light intensity positively regulate the gene transcription of NR, but ammonium ions and Glu, Gln do the negative way. Water deficit is a very serious constraint on N2 fixation rate and soybean (Glycine max Merr.) grain yield, in which, ureide accumulation and degradation under water deficit appear to be the key issues of feedback mechanism on nitrogen fixation. Water stress decreases NR activity, but increases proteinase activity, and thus, they regulate plant nitrogen metabolism, although there are some different effects among species and cultivars. Water stress also decreases plant tissue protein content, ratio of protein and amino acid, and reduces the absorption of amino acid by plant. On the contrary, soil flooding decreases the content and accumulation amount of root nitrogen in winter wheat by 11.9% from booting to flowering stages and 39.1% during grain filling stage, and reduces the ratio of carbon and nitrogen by 79.6%. The results misadjust the metabolism between carbon and nitrogen, and result in the end of the root growth. Elevated CO2 level could decrease plant leaf nitrogen content under well-watered condition, but almost maintain stable under water deficit condition. The radiation of UV-B significantly reduces the partitioning coefficient and synthetic rate of Rubisco, which significantly decreases the photosynthetic rate. This paper reviewed the pathway of plant nitrogen assimilation, characteristics of key enzymes and their regulating mechanisms with picturing the regulating mode of NR, and described the signal sensing and conduct of plant nitrogen metabolism and the formation, transportation, storage and degradation of plant cell protein with picturing the schedule of protein transport of membrane system in plant cell. Seven key tasks are emphasized in this paper in terms of the review on the effects and mechanisms of key ecological factors including water stress on plant nitrogen metabolism. They are: 1) the absorption mechanism of plant based on different nitrogen sources and environmental regulations, 2) the localization and compartmentalization of the key enzymes of nitrogen mechanism in plant cell, 3) the gene and environmental regulating model and their relationships in various key enzymes of nitrogen metabolism, 4) the function of main cell organs and their responses to environmental factors in nitrogen metabolism process, 5) physiological and chemical mechanism of nitrogen and the relationship between the mechanism and protein formation during crop grain filling, 6) improving gene structure of special species or cultivars using gene engineering methods to enhance the resistance to environmental factor stress and the efficiency of absorption and transportation of nitrogen, and 7) the mechanism of natural nitrogen cycle and its response to human activity disturbance.

摘要

氮代谢不仅是植物生理学的基本过程之一,也是全球化学循环的重要组成部分。植物氮同化通过硝酸盐的还原直接参与氨基酸的合成与转化。在此阶段,一些关键酶,如硝酸还原酶(NR)、谷氨酰胺合成酶(GS)、谷氨酸脱氢酶(GDH)、谷氨酰胺合成酶(GOGAT)、天冬酰胺合成酶(AS)和天冬氨酸转氨酶(AspAT)参与这些过程。蛋白质在植物细胞中通过氨基酸被同化,并通过修饰、分类、运输和储存等过程成为植物有机体的一部分。氮代谢通过关键酶调控和产物转化与碳代谢相关联,这构成了基本的生命过程。在植物细胞中的这些氨基酸中,谷氨酸(Glu)、谷氨酰胺(Gln)、天冬氨酸(Asp)和天冬酰胺(Asn)等起关键作用,它们通过调节这些关键酶的形成和活性来调控彼此在植物细胞中的转化及其含量。环境因素也通过调节关键酶的基因表达及其活性来影响关键氨基酸的转化和循环利用。硝酸盐和光照强度对NR的基因转录起正调控作用,而铵离子以及Glu、Gln则起负调控作用。水分亏缺对大豆(Glycine max Merr.)的固氮率和籽粒产量是非常严重的限制因素,其中,水分亏缺下脲类物质的积累和降解似乎是固氮反馈机制的关键问题。水分胁迫降低NR活性,但增加蛋白酶活性,因此,它们调控植物氮代谢,尽管不同物种和品种之间存在一些不同的影响。水分胁迫还降低植物组织蛋白质含量、蛋白质与氨基酸的比例,并减少植物对氨基酸的吸收。相反,土壤淹水使冬小麦从孕穗期到开花期根系氮含量及其积累量降低11.9%,灌浆期降低39.1%,碳氮比降低79.6%。这些结果使碳氮代谢失调,导致根系生长停止。在水分充足条件下,CO₂浓度升高会降低植物叶片氮含量,但在水分亏缺条件下几乎保持稳定。UV - B辐射显著降低Rubisco的分配系数和合成速率,从而显著降低光合速率。本文通过绘制NR的调控模式综述了植物氮同化途径、关键酶特性及其调控机制,并通过绘制植物细胞膜系统蛋白质运输示意图描述了植物氮代谢的信号感知与传导以及植物细胞蛋白质的形成、运输、储存和降解。本文在综述包括水分胁迫在内的关键生态因子对植物氮代谢的影响及其机制方面强调了七项关键任务。它们是:1)基于不同氮源和环境调控的植物吸收机制;2)植物细胞中氮代谢关键酶的定位和区室化;3)氮代谢各种关键酶中的基因与环境调控模式及其关系;4)氮代谢过程中主要细胞器官的功能及其对环境因子的响应;5)作物灌浆期氮的生理生化机制及其与蛋白质形成的关系;6)利用基因工程方法改善特定物种或品种的基因结构以增强对环境因子胁迫的抗性以及氮的吸收和运输效率;7)自然氮循环机制及其对人类活动干扰的响应。

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