植物寡肽运输与硝酸根运输基因家族的研究进展

氮素是植物生长发育的重要营养元素,也是限制植物生物量尤其是经济产量的关键营养元素之一.植物不仅能从外界获取无机氮素(硝酸根、铵根和尿素等),还能以氨基酸、寡肽等形式获取有机氮素.植物已进化出复杂的运输系统来吸收与运输这些含氮化合物.硝酸根运输基因家族分为低亲和力硝酸根运输基因(low-affmity nitrate t...     

植物K+通道AKT1的研究进展

钾(K)是植物生长发育必需的大量营养元素之一, 主要通过根细胞的K⁺通道及转运蛋白介导吸收。AKT1是Shaker型K⁺通道家族的重要成员, 在植物根吸收K⁺和体内跨膜转运中发挥重要作用。该文综述了植物AKT1的分子结构、组织特异性表达、调控机制及生物学功能等方面的研究进展, 并对该通道今后的研究方向进行了展望。     

云锦杜鹃NRT基因序列的生物信息学分析

通过转录组测序,获得在接种 ERM 真菌的云锦杜鹃苗根系中显著差异表达的基因,其中硝酸根转运蛋白(NRT )基因是硝态氮吸收转运的关键基因。利用生物信息学方法,分析云锦杜鹃根转录组的硝酸根转运蛋白(NRT )基因序列,对其推导的氨基酸的理化性质、亲水性/疏水性、跨膜结构、导肽、二级结构、高级结构进行预测,并对硝酸根转运蛋白的氨基酸做进化发育分析。为进一步了解 NRT 基因在云锦杜鹃接种苗根系氮素吸收的作用奠定了基础。     

C3和C4植物的氮素利用机制

提高植物的氮素利用效率(NUE)不仅有利于保障全球粮食安全, 也是实现农业可持续发展的重要途径。近半个世纪以来, 植物氮素利用机理研究已取得重要进展, 但NUE的调控机制仍不明确, NUE的提高仍然十分有限。高等植物集光合碳素同化和氮素同化于一体, 只有碳氮代谢相互协调, 才能维持植物体内的碳氮平衡, 保证植物正常生长发育。由于C₃和C₄植物的光合氮素利用率(PNUE)存在差异, 对氮素的利用效率也会存在差异。为了更有效地提高作物的NUE, 须更全面地了解C₃和C₄植物对氮素吸收、转运、同化和信号转导等关键因子的功能和调控机制。此外, 面对大气CO₂浓度增高和全球气候变暖条件下的植物碳氮同化及其机理的研究也不容忽视。该文综述了C₃和C₄植物氮素利用关键因素的差异及其调控机制, 并对提高C₃禾本科作物氮素利用效率的遗传改良途径进行了展望。     

耕作方式与施氮量对小麦-玉米复种系统玉米季土壤氮素转化及产量的影响

在黄淮砂姜黑土区冬小麦-夏玉米复种两熟种植体系中,研究了小麦季3种耕作方式(常规翻耕、旋耕和深松)结合夏玉米播前3个施氮量(120、225和330 kg·hm^-2)对玉米季主要生育时期根际土壤氮素转化微生物作用强度及酶活性、无机氮含量和产量的影响.结果表明: 旋耕方式下氨化作用强度最高,且随着施氮量的增加,土壤氮素转化微生物作用强度及酶活性增强.深松方式下根际土壤硝化、反硝化作用强度与脲酶活性明显高于常规与旋耕方式.增施氮肥可加强深松方式对土壤氮素转化的促进作用,而过量施氮虽然提高了土壤无机氮含量及玉米产量,但会对土壤氮素转化微生物作用强度及酶活性产生抑制.深松方式结合225 kg·hm^-2施氮量更有利于砂姜黑土区夏玉米土壤氮素转化,而深松方式结合330 kg·hm^-2施氮处理下产量最高.     

氮添加对盐渍化草地根际土壤理化性质的影响

为探究氮输入和根际效应对盐渍化草地土壤理化性质的影响,对8个水平氮添加处理下(0、1、2、4、8、16、24和32 g N·m^-2·a^-1)晋北盐渍化草地根际和非根际土壤理化性质进行研究。结果表明: 氮添加显著降低根际土壤pH,显著增加根际和非根际土壤Ca²⁺、NO₃^--N和无机氮含量;随氮添加量的增加,根际和非根际土壤Ca²⁺、NO₃^--N、无机氮含量以及根际土壤全氮含量呈逐渐升高的趋势,而根际土壤Na⁺、K⁺、Mg²⁺、NH₄⁺-N和氨基酸含量以及非根际土壤全氮含量呈先升高后降低的趋势。主成分分析表明,根际土壤理化性质对低氮(≤8 g·m^-2·a^-1)和高氮添加(>8 g·m^-2·a^-1)的响应具有明显差异。根际土壤pH、有机酸和氨基酸含量分别比非根际土壤低0.71、44.3%和9.8%,而K⁺、Ca²⁺、Mg²⁺、NH₄⁺-N、无机氮、全碳和全氮含量分别比非根际土壤高51.0%、47.6%、20.8%、215.5%、139.3%、31.7%和65.3%,表明根际效应对盐渍化草地土壤理化性质的影响大于氮输入的影响。     

油蒿叶片氮分配对其最大净光合速率季节变异的影响

植物最大净光合速率的季节变异性及其氮调控机制是近年来植物生理生态领域研究热点,对荒漠植物光合最大净光合速率季节动态及其叶氮影响,特别是叶氮分配对最大净光合速率调控机制的了解仍非常有限。2018年5—10月在宁夏盐池毛乌素沙地,对当地主要建群种油蒿(Artemisia ordosica)进行生长季原位观测,测定其叶光合光响应曲线(A-PAR)、CO₂响应曲线(A-Ci)和叶氮含量,结合环境观测数据,分析A-PAR关键参数最大净光合速率(A_max)的季节变异和叶片氮分配相关参数对A_max的调控。结果表明,油蒿叶片A_max在生长期季节变异系数(Cv)为14%,在完全展叶中期,光合氮利用效率(PNUE)有最大值11.82μmolCO₂ gN^-1 s^-1,此时叶片氮素在光合系统中的分配比例最大,A_max有最大值29.48μmol CO₂ m^-2s^-1,油蒿光...     

两种豆科植物及各器官对不同形态氮的吸收、分配研究

豆科植物在氮素缺乏的荒漠生态系统中大量存在，是该生态系统提供有效氮的中心，也是这一区域重要的先锋物种。该文选择古尔班通古特沙漠广泛分布的弯花黄芪(Astragalus flexus)和镰荚黄芪(Astragalus arpilobus)作为研究对象，分别在0～5、5～15 cm土层添加3种不同形态氮(¹⁵N-NH₄⁺、¹⁵N-NO₃^-、¹⁵N-glycine),研究两种植物及各器官对不同形态氮素的吸收、分配策略。结果表明：(1)在不同土层中，两种植物均偏好吸收硝态氮，并且弯花黄芪、镰荚黄芪对硝态氮的最高吸收速率均为3.26、2.59μg·g^-1·h^-1。(2)在不同土层中，植物各器官间均对不同氮源吸收及分配有显著性差异(P<0.05),弯花黄芪根的¹⁵N吸收量均大于镰荚黄芪的，3种不同形态氮主要分配于叶。(3)在不同土层中，不同氮源对两种植物的贡献率均为^...     

水氮耦合对旱地胡麻产量形成与花后氮素积累转运的影响

为明确旱地胡麻在有限灌水条件下的最佳水氮耦合管理模式,采用完全随机裂区试验设计,以灌水(I₀: 0 m³·hm^-2; I₁₂₀₀: 1200 m³·hm^-2; I₁₈₀₀: 1800 m³·hm^-2)为主区,施氮量(N₀: 0 kg·hm^-2; N₆₀₀: 60 kg·hm^-2; N₁₂₀: 120 kg·hm^-2)为副区,测定胡麻不同生育阶段氮素积累量、花后氮素转运特征、产量和氮肥利用率。结果表明: 不同水氮处理对旱地胡麻不同生育时期各器官氮素吸收、积累及产量的耦合效应不同。不灌水条件下,施氮有利于胡麻花期和成熟期茎秆对氮素的吸收,不同灌水水平下N₁₂₀均抑制了茎秆对氮的吸收;I₁₂₀₀水平下,花期叶片氮含量随施氮量的增加先升高后下降,N₆₀较N₀和N₁₂₀高11.0%和28.9%;I₁₈₀₀水平下,施氮提高了成熟期胡麻叶片中氮含量,N₆₀和N₁₂₀较N₀高39.7%和26.9%。水氮对胡麻阶段氮素积累量影响的耦合效应主要表现在现蕾期以后,同一灌水水平下,N₆₀促进了胡麻现蕾期以后各阶段氮素积累量,而N₁₂₀具有抑制作用。施氮分别提高了I₁₂₀₀和I₁₈₀₀水平下叶片和茎秆氮素转运率和贡献率。灌水1800 m³·hm^-2、施氮60 kg·hm^-2显著增加了胡麻单株有效蒴果数和籽粒产量(6.6%~22.8%),是试验区比较适宜的水氮耦合管理模式。     

不同供氮量对二月兰产量、土壤无机氮残留及氮平衡的影响

研究华北冬绿肥二月兰对不同供氮水平的响应特征,确定实现绿肥高产高效的土壤适宜供氮量,可为华北集约化农田最大化发挥绿肥生态效应和优化春玉米/冬绿肥轮作体系氮素管理提供理论依据和技术参考.选取多年不施肥试验地设置供氮梯度试验,研究了不同供氮水平对冬绿肥二月兰翻压前地上部生物量累积、氮素吸收、土壤无机氮残留和冬绿肥季土壤氮素平衡的影响.结果表明: 在土壤无机氮含量较低(0～90 cm土层15 kg·hm^-2)条件下,施氮显著提高二月兰生物量和吸氮量.其中,施氮90 kg·hm^-2处理表现最高,绿肥生物量(干质量)和吸氮量分别为2031.0和42.0 kg·hm^-2;土壤无机氮残留量随施氮量增加而增加,且在施氮量高于60 kg·hm^-2后呈现快速增加趋势;随施氮量增加二月兰生长季的表观氮平衡表现出由亏缺到盈余的变化特征,在施氮量为60～90 kg·hm^-2条件下氮收支基本平衡.土壤供氮量(绿肥播前0～90 cm土壤无机氮含量与施氮量之和)与二月兰生物量、吸氮量和绿肥翻压前土壤无机氮含量的关系可以分别用二次、线性加平台和指数方程进行模拟,依据模型计算二月兰生物量最高值(2010 kg·hm^-2)时的播前土壤供氮量和绿肥翻压前土壤无机氮残留量分别是136和78 kg·hm^-2;而在二月兰吸氮量最高值40 kg·hm^-2时,二月兰生物量为1919 kg·hm^-2,相当于最高生物量的95%,绿肥翻压前土壤残留无机氮降低至57 kg·hm^-2,与之对应的播前土壤供氮量为105 kg·hm^-2,该值与目前华北地区优化施氮下玉米收获后土壤残留无机氮推荐含量(100 kg·hm^-2)基本相当.综合考虑绿肥的农学和环境效应,春玉米/冬绿肥轮作体系中二月兰播前土壤供氮量应控制在100～105 kg·hm^-2.     

Plant acquisition of organic nitrogen in boreal forests

Research on plant nitrogen (N) uptake and metabolism has more or less exclusively concerned inorganic N, particularly nitrate. Nevertheless, recent as well as older studies indicate that plants may have access to organic N sources. Laboratory studies have shown that ectomycorrhizal and ericoid mycorrhizal plants can degrade polymeric N and absorb the resulting products. Recent studies have also shown that some non‐mycorrhizal plants are able to absorb amino acids. Moreover, amino acid transporters have been shown to be present in both plant roots and in mycorrhizal hyphae. Although both mycorrhizal and non‐mycorrhizal plants appear to have a capacity for absorbing a range of organic N compounds, is this capacity realized in the field? Several lines of evidence show that plants are outcompeted by microorganisms for organic N sources. Such studies, however, have not addressed the issue of spatial and temporal separation between plants and microorganisms. Moreover, competition studies have not been able to separate uptake by symbiotic and non‐symbiotic microorganisms. Qualitative assessment of organic N uptake by plants has been performed with dual‐labelled glycine in several studies. These studies arrive at different conclusions: some indicate that plants do not absorb this organic N source when competing with other organisms in soil, while others conclude that significant fractions of amino acid N are absorbed as intact amino acid. These variable results may reflect species differences in the ability to absorb glycine as well as differences in experimental conditions and analytical techniques. Although theoretical calculations indicate that organic N might add significant amounts of N to plant N uptake, direct quantitative assessment of the fraction of plant N derived from uptake by organic N sources is a challenge for future research.     

Regulation of genes involved in nitrogen utilization on different C/N ratios and nitrogen sources in the model ectomycorrhizal fungus Hebeloma cylindrosporum

Nitrogen (N) utilization by ectomycorrhizal fungi is an essential aspect of their ecosystem function. N deposition changes both the N pools and the carbon/nitrogen (C/N) ratio of the substrates where ectomycorrhizal fungi are found, and it is important to understand how these changes affect the N forms used by ectomycorrhizal fungi. To overcome the difficulties of studying ectomycorrhizal fungi in situ, we investigated all known N genes in the model fungus, Hebeloma cylindrosporum in a culture study. In addition to studying the regulation of all known N utilization genes, we aimed to understand whether there are gene clusters that undergo similar regulation. Lastly we studied how C/N ratio, N transporter type, and N source affected relative gene expression levels. We grew the D2 strain of H. cylindrosporum on a range of inorganic and organic N sources under low, medium, and high C/N ratios. We found three gene clusters that were regulated in a similar pattern. Lastly, we found C/N ratio, N source and N transporter type all affected gene expression levels. Relative expression levels were highest on the high C/N ratio, BSA and diLeucine N sources, and inorganic N transporters were always expressed at higher levels than organic N transporters. These results suggest that inorganic N sources may always the default preference for H. cylindrosporum, regardless of both the N sources and the C/N ratio of the substrate.     

Peptide uptake in the ectomycorrhizal fungus Hebeloma cylindrosporum: characterization of two di- and tripeptide transporters (HcPTR2A and B)

Constraints on plant growth imposed by low availability of nitrogen are a characteristic feature of ecosystems dominated by ectomycorrhizal plants. Ectomycorrhizal fungi play a key role in the N nutrition of plants, allowing their host plants to access decomposition products of dead plant and animal materials. Ectomycorrhizal plants are thus able to compensate for the low availability of inorganic N in forest ecosystems. The capacity to take up peptides, as well as the transport mechanisms involved, were analysed in the ectomycorrhizal fungus Hebeloma cylindrosporum. The present study demonstrated that H. cylindrosporum mycelium was able to take up di- and tripeptides and use them as sole N source. Two peptide transporters (HcPTR2A and B) were isolated by yeast functional complementation using an H. cylindrosporum cDNA library, and were shown to mediate dipeptide uptake. Uptake capacities and expression regulation of both genes were analysed, indicating that HcPTR2A was involved in the high-efficiency peptide uptake under conditions of limited N availability, whereas HcPTR2B was expressed constitutively.     

Structure, function and regulation of ammonium transporters in plants

Ammonium is an important source of nitrogen for plants. It is taken up by plant cells via ammonium transporters in the plasma membrane and distributed to intracellular compartments such as chloroplasts, mitochondria and vacuoles probably via different transporters in each case. Ammonium is generally not used for long-distance transport of nitrogen within the plant. Instead, most of the ammonium transported into plant cells is assimilated locally via glutamine synthetases in the cytoplasm and plastids. Ammonium is also produced by plant cells during normal metabolism, and ammonium transporters enable it to be moved from intracellular sites of production to sites of consumption. Ammonium can be generated de novo from molecular nitrogen (N(2)) by nitrogen-fixing bacteria in some plant cells, such as rhizobia in legume root nodule cells, and at least one ammonium transporter is implicated in the transfer of ammonium from the bacteria to the plant cytoplasm. Plant physiologists have described many of these ammonium transport processes over the last few decades. However, the genes and proteins that underlie these processes have been isolated and studied only recently. In this review, we consider in detail the molecular structure, function and regulation of plant ammonium transporters. We also attempt to reconcile recent discoveries at the molecular level with our knowledge of ammonium transport at the whole plant level.     

Molecular characterization, expression and functional analysis of the amino acid transporter gene family (OsAATs) in rice

Nitrogen (N) is one of the most important limiting factors for plant growth and development. Amino acids are the major source of organic N, which is converted from inorganic N absorbed by plant roots from the soil. Amino acid transporters are the principal mediators of organic N distribution and important regulators of resource allocation in plants. Although the complete genomic sequence of rice has already been released, there is still little known about amino acid transporter genes in rice. In this study, 79 OsAAT genes were identified by a database search of the rice genome based upon HMM profiles. A bioinformatics analysis of the complete set of OsAAT genes is presented, including chromosomal location, phylogenetic analysis, gene structure, protein analysis, conserved motifs, protein structures and cis-element analysis of the promoters. In addition, the comprehensive expression profile of OsAAT genes in rice tissues/organs under N starvation conditions was investigated by real-time PCR analysis. Diverse expression patterns of OsAAT genes indicated diverse biological functions of the amino acid transporters and the important roles of OsAAT genes in N uptake, metabolism and distribution during N starvation. The evaluation of yield and carbon (C) and N content of osaat knockout mutants also suggested the important roles of the OsAAT5, OsAAT7, OsAAT24, OsAAT49 and OsAAT60 genes in yield and biomass production and C and N metabolism and distribution in rice plants.     

Challenging the paradigm of nitrogen cycling: no evidence of in situ resource partitioning by coexisting plant species in grasslands of contrasting fertility

In monoculture, certain plant species are able to preferentially utilize different nitrogen (N) forms, both inorganic and organic, including amino acids and peptides, thus forming fundamental niches based on the chemical form of N. Results from field studies, however, are inconsistent: Some showing that coexisting plant species predominantly utilize inorganic N, while others reveal distinct interspecies preferences for different N forms. As a result, the extent to which hypothetical niches are realized in nature remains unclear. Here, we used in situ stable isotope tracer techniques to test the idea, in temperate grassland, that niche partitioning of N based on chemical form is related to plant productivity and the relative availability of organic and inorganic N. We also tested in situ whether grassland plants vary in their ability to compete for, and utilize peptides, which have recently been shown to act as an N source for plants in strongly N-limited ecosystems. We hypothesized that plants would preferentially use NO₃⁻-N and NH₄⁺-N over dissolved organic N in high-productivity grassland where inorganic N availability is high. On the other hand, in low-productivity grasslands, where the availability of dissolved inorganic N is low, and soil availability of dissolved organic N is greater, we predicted that plants would preferentially use N from amino acids and peptides, prior to microbial mineralization. Turves from two well-characterized grasslands of contrasting productivity and soil N availability were injected, in situ, with mixtures of ¹⁵N-labeled inorganic N (NO₃⁻ and NH₄⁺) and ¹³C¹⁵N labeled amino acid (l-alanine) and peptide (l-tri-alanine). In order to measure rapid assimilation of these N forms by soil microbes and plants, the uptake of these substrates was traced within 2.5 hours into the shoots of the most abundant plant species, as well as roots and the soil microbial biomass. We found that, contrary to our hypothesis, the majority of plant species across both grasslands took up most N in the form of NH₄⁺, suggesting that inorganic N is their predominant N source. However, we did find that organic N was a source of N which could be utilized by plant species at both sites, and in the low-productivity grassland, plants were able to capture some tri-alanine-N directly. Although our findings did not support the hypothesis that differences in the availability of inorganic and organic N facilitate resource partitioning in grassland, they do support the emerging view that peptides represent a significant, but until now neglected, component of the terrestrial N cycle.     

植物蔗糖转运蛋白表达的调控因素与分子机制

植物光合作用的产物主要以蔗糖的形式在植物体内进行从源到库的运输。蔗糖转运蛋白是此过程的重要参与者,其表达和调控与植物中光合作用产物的分配紧密关联,从而调控着植物的生长发育、结果结实、抗逆抗病等性状。蔗糖转运蛋白的表达受到植物发育时期、外界环境条件及激素的影响。蔗糖转运蛋白的调控机制有转录因子的调节、基因内部序列调控、蛋白质的磷酸化、蛋白之间的相互作用及质子转运体的活性调节等。综述了国内外对蔗糖转运蛋白表达与活性的调控因素及机制等最新的研究内容,以期为从多角度上探索植物蔗糖转运蛋白的功能和调控机制提供相关研究信息和思路。     

The carbon bonus of organic nitrogen enhances nitrogen use efficiency of plants

The importance of organic nitrogen (N) for plant nutrition and productivity is increasingly being recognized. Here we show that it is not only the availability in the soil that matters, but also the effects on plant growth. The chemical form of N taken up, whether inorganic (such as nitrate) or organic (such as amino acids), may significantly influence plant shoot and root growth, and nitrogen use efficiency (NUE). We analysed these effects by synthesizing results from multiple laboratory experiments on small seedlings (Arabidopsis, poplar, pine and spruce) based on a tractable plant growth model. A key point is that the carbon cost of assimilating organic N into proteins is lower than that of inorganic N, mainly because of its carbon content. This carbon bonus makes it more beneficial for plants to take up organic than inorganic N, even when its availability to the roots is much lower – up to 70% lower for Arabidopsis seedlings. At equal growth rate, root:shoot ratio was up to three times higher and nitrogen productivity up to 20% higher for organic than inorganic N, which both are factors that may contribute to higher NUE in crop production.     

The roles of biotic resistance and nitrogen deposition in regulating non-native understory plant diversity

Understanding the mechanisms that allow for plant invasions is important for both ecologists and land managers, due to both the environmental and economic impacts of native biodiversity losses. We conducted an observational field study in 2008 to examine the relationship between native and non-native forest understory plant species and to investigate the influence of soil nitrogen (N) on plant community richness and diversity. In 2009, we conducted a companion fertilization experiment to investigate how various forms of N deposition (inorganic and organic) influenced native and non-native species richness and diversity. We found that native species richness and diversity were negatively correlated with 1) non-native species richness and diversity and 2) higher total soil inorganic N. In the deposition experiment, adding organic N fertilizers decreased native richness and diversity compared to inorganic N fertilizers. Together, these results indicate that increasing soil N can be detrimental to native species; however, native species richness and diversity may counteract the N-stimulation of non-native species. Furthermore, the negative effects of organic N deposition on native plants may be just as strong, if not stronger, than the effects of inorganic N deposition.