Nitrogen nutrition and adaptation of glycophytes to saline environment: a review

Relations between nitrogen (N) nutrition and salinity tolerance in plants are multifaceted and varies significantly depending on many soil and plant factors. Saline environment might experience an N dilemma due to the opposing effects of salt ions on N uptake, translocation and metabolism within the plant body. Adequate regulation of N under saline conditions can be a promising approach to alleviate salinity’s effects on plants by ameliorating ion toxicity and nutrient imbalances through its impacts on the uptake and redistribution of salt ions within the plant. Certain N-containing compounds including proline, glycine betaine, proteins and polyamines help the plants to tolerate salinity through their involvement in improving water uptake and water use efficiency, membrane integrity, enzyme activation, hormonal balance, chlorophyll synthesis, stimulation of photosystems and CO₂ assimilation under salinity stress. Nitrogen, particularly NO₃^? represents a stress signal that triggers the activation of antioxidant enzymes to protect the plants against salinity-induced oxidative damage. Furthermore, the source/form of N application can affect not only N-interactions but also the behavior of other nutrients in stress environment. The present review deals with N-salinity relations in plants, particularly glycophytes, emphasizing on N-induced mechanisms which can improve plant adaptation to saline environment.     

Nitrogen assimilation studies using 15N in soybean plants treated with imazethapyr, an inhibitor of branched-chain amino acid biosynthesis

The pattern of nitrogen assimilation in soybean plants treated with a herbicide that inhibits branched-chain amino acid biosynthesis was evaluated by (15)N isotopic analysis. The herbicide imazethapyr caused a strong decrease in nitrate uptake by roots, partly due to a reduced stomatal conductance. The inhibition of (15)N uptake was accompanied by a decrease in the (15)N content in the plant and, concomitantly, an inhibition of translocation to the shoot. Imazethapyr inhibited nitrate reductase activity in leaves and roots. Among all parameters studied, "de novo" synthesis of proteins was the first parameter of the N assimilation metabolism affected by the herbicide. These results show that this class of herbicides totally damages N metabolism and indicates a regulatory effect on N uptake and translocation that would be mediated by the increase in free amino acid pool provoked by the inhibition of branched-chain amino acid biosynthesis.     

植物氮素吸收利用相关NPF基因家族研究进展

氮(N)是植物生长发育需要量最大的矿质营养元素,也是作物产量的限制因子。硝态氮(NO₃^--N)是植物吸收利用氮素的主要形态之一。目前,植物中已报道4个基因家族(NPF、NRT2、CLC和SLAC1/SLAH)参与硝态氮的吸收和利用,其中NPF基因家族成员数量众多且功能多样化,近年来获得较多关注和深入研究。模式植物拟南芥和主要粮食作物水稻、玉米和小麦中,分别含有53、93、79和331个NPF基因。拟南芥NPF家族中已有超过一半成员(31/53)的生物学功能被解析,粮食作物水稻中NPF基因功能亦有较多报道。研究表明,NPF基因广泛参与了植物对氮素的吸收及其调控、转运、分配/再分配等过程,一些成员对于改良和提高作物氮素利用率(nitrogen use efficiency, NUE)具有重要作用。因此,从氮素进入植物体及其在植物体内流动的层面出发,发掘具有重要功能的候选NPF基因,对于解析植物利用氮素的分子机制及其遗传改良具有重要意义。本文综述了模式植物拟南芥以及粮食作物中已报道的NPF基因在氮素吸收和利用中的生物学功能。目前粮食作物玉米中仅有4...     

Toxicity of cadmium and its competition with mineral nutrients for uptake by plants: A review

Cadmium(Cd)is a toxic heavy metal occurring in the environment naturally and is also generated through various anthropogenic sources and acts as a pollutant.Human health is affected by Cd pollution in farmland soils because food is the main source of Cd intake in the non-smoking population.For crops,Cd toxicity may result from a disturbance in uptake and translocation of mineral nutrients and disturbance in plant metabolism,inhibiting plant growth and development.However,plants have Cd tolerance mechanisms,including restricted Cd uptake,decreased Cd root-to-shoot translocation,enhanced antioxidant enzyme activities,and increased production of phytochelatins.Furthermore,optimal supply of mineral nutrients is one of the strategies to alleviate the damaging effects of Cd on plants and to avoid its entry into the food chain.The emerging molecular knowledge contributes to understanding Cd uptake,translocation,and remobilization in plants.In this review,Cd toxicity and tolerance mechanisms,agricultural practices to minimize Cd accumulation,Cd competition with essential elements(calcium,copper,iron,zinc,and manganese),and genes associated with Cd uptake are discussed in detail,especially regarding how these mineral nutrients and genes play a role in decreasing Cd uptake and accumulation in crop plants.     

Negative and positive contributions of arbuscular mycorrhizal fungal taxa to wheat production and nutrient uptake efficiency in organic and conventional systems in the Canadian prairie

Improving technologies and the challenge of producing more bio-products while reducing the environmental footprint of humans are shifting paradigms in agricultural research. Harnessing the microbial resources of arable soils is a new avenue to improve the efficiency of nutrient use in agriculture. The objective of this study was to define how crop management influences the contribution of resident AM fungi to nutrient efficiency and crop productivity. The AM fungal communities of 72 organically and 78 conventionally managed wheat fields of the Canadian prairie were described by 454 pyrosequencing and related to crop productivity and N and P use efficiency. Conventional management reduces soil pH and increases the fluxes of all soil nutrients except S, B, and K. Organic management increased the abundance of Claroideoglomus reads. The efficiency of N and P uptake from soil by organic wheat was 2.3 and 1.8 times higher than that of conventional systems. This high N and P uptake efficiency in organic wheat crops was mainly attributable to the low soil fertility of organic fields, as wheat biomass production was 1.44 times greater in conventional than organic systems. Overall, the amounts of N and P taken up by conventional and organic wheat crops were similar. Plant nutrient balance and the abundance of Paraglomus drove conventional wheat production, whereas organic production depended mainly on soil moisture, plant nutrient balance, and abundance of Glomus, which was associated with reduced and nutrient-inefficient wheat production. The high nutrient concentrations at maturity and the low productivity of organic wheat fit a model of limiting CO₂-assimilation. The trade-off between nutrient use efficiency and productivity in low input wheat production could be relieved by reducing the abundance of Glomus species, increasing soil moisture and early N availability, or by improving the inherent CO₂ assimilation capacity of wheat.     

Leaf nitrogen determination using non-destructive techniques–A review

The optimisation of plant nitrogen-use-efficiency (NUE) has a direct impact on increasing crop production by optimising use of nitrogen fertiliser. Moreover, it protects environment from negative effects of nitrate leaching and nitrous oxide production. Accordingly, nitrogen (N) management in agriculture systems has been major focus of many researchers. Improvement of NUE can be achieved through several methods including more accurate measurement of foliar N contents of crops during different growth phases. There are two types of methods to diagnose foliar N status: destructive and non-destructive. Destructive methods are expensive and time-consuming, as they require tissue sampling and subsequent laboratory analysis. Thus, many farmers find destructive methods to be less attractive. Non-destructive methods are rapid and less expensive but are usually less accurate. Accordingly, improving the accuracy of non-destructive N estimations has become a common goal of many researchers, and various methods varying in complexity and optimality have been proposed for this purpose. This paper reviews various commonly used non-destructive methods for estimating foliar N status of plants.     

Perspectives of Using L-Tryptophan for Improving Productivity of Agricultural Crops: A Review

Plant growth regulators are biologically active signaling molecules that regulate a number of plant physiological processes. Auxin (indole-3-acetic acid) is an important plant growth regulator and is synthesized within plant tissues through L-tryptophan (L-TRP)-dependent and -independent pathways. It has been found that plants respond to exogenously applied L-TRP due to insufficient endogenous auxin biosynthesis. The exogenous application of L-TRP is highly significant for normal plant growth and development. L-tryptophan is applied through foliar spray, seed priming, and soil application. Soil-applied L-TRP is either directly taken up by plants or metabolized to auxin by soil microbiota and then absorbed by plant roots. Similarly, foliar spray and seed priming with L-TRP stimulates auxin synthesis within plants and improves the growth and productivity of agricultural crops. Furthermore, L-TRP contains approximately 14% nitrogen (N) in its composition, which is released upon its metabolism within a plant or in the rhizosphere and plays a role in enhancing crop productivity. This review deals with assessing crop responses under the exogenous application of L-TRP in normal and stressed environments, mode of action of L-TRP, advantages of using L-TRP over other auxin precursors, and role of the simultaneous use of L-TRP and auxin-producing microbes in improving the productivity of agricultural crops. To the best of our knowledge, this is the first review reporting the importance of the use of L-TRP in agriculture.     

不同氮肥水平下水稻产量以及氮素吸收、利用的基因型差异比较

采用田间试验在施氮量为06、0、120、1802、40、3003、60.kg/hm27个水平下研究了不同水稻子粒产量、产量构成因子以及氮素吸收和利用的差异。结果表明,水稻品种4007的子粒产量在各个施氮水平下显著高于品种ELIO对氮肥的响应度高。施氮水平显著影响子粒产量构成因子。有效穗数与子粒产量存在显著正相关:ELIO和4007的相关系数(r)分别为0.839**和0.933**,表明有效穗数对水稻子粒产量起着非常重要的作用。本试验条件下,ELIO和4007获得最高产量所需的有效穗数分别为332、561个/m2;两者的氮素吸收效率在各施氮素水平下差异很小,均随着施氮量的增加而增加,而氮素利用效率均随着施氮量的增加而下降。4007的氮素利用效率在各个施氮水平下显著高于ELIO,较高的氮素收获指数(NHI)是主要原因之一。水稻氮素利用效率与成熟期茎秆、叶片的氮含量显著负相关,说明开花期后植物将吸收的氮素从营养器官有效地转运到子粒中是氮素利用效率高的重要原因之一。     

A review of nitrogen translocation and nitrogen-use efficiency

Nitrogen (N) is a primary nutrient for crop growth. In most agricultural areas, crop production relies heavily on the supply of exogenous N fertilizers. Globally, a large amount of nitrogenous fertilizer is applied to achieve maximum crop productivity. However, N fertilizer application is costly and negatively impacts the environment. One way around these problems is to provide a new N-use efficiency (NUE) framework that promotes the efficient acquisition and use of N and high N translocation rates from vegetative organs to reproductive organs under reduced N inputs. This review integrates the current knowledge about the direct and indirect effects of soil properties, crop varieties, soil water status, soil N fertilizers, and soil microbes on N translocation. In addition, we explore strategies that specifically accomplish N translocation via mycorrhizal fungi. Finally, we discuss recent advances in plant physiology, demonstrating that N translocation is an effective target to improve cereal yields and NUE. Although the relationship between crop variety and N translocation is mostly unknown, understanding this relationship is necessary to select and breed crop varieties that exhibit high yields and N translocation efficiency.     

植物吸收转运硝态氮及其信号调控研究进展

硝态氮是植物吸收利用的主要氮源,其吸收利用是一个高度协调复杂的调控过程。植物为了在各种变化的环境中生存,进化出了适宜不同环境的硝态氮吸收利用机制。植物根系中存在不同类型的硝态氮受体,可以感受外界硝态氮浓度变化,并启用高亲和力或低亲和力硝态氮吸收系统,从而吸收硝态氮;硝态氮进入根系后,大部分被运输到地上部进行同化作用,合成大分子物质,以促进植物生长;如果地上部硝态氮含量过多,植物可把多余的硝态氮运送到液泡内储存,待需要时再从液泡转运至细胞质中利用。植物生长发育过程中,老叶和成熟叶片中的硝态氮可被转运到新生组织中,促进新生组织生长。硝态氮吸收利用过程中大量硝态氮吸收、转运、储存、同化和信号调控基因被有序激活并协调工作,促进植物高效吸收利用硝态氮。本文主要针对NRT1和NRT2硝态氮吸收转运相关基因及其功能,以及参与初级硝态氮反应的相关转录因子和小信号多肽在硝态氮信号传导和组织间的信号交流进行综述,以便深入理解植物吸收利用硝态氮的机理,为高效利用氮素的作物育种和栽培技术的创建提供新的思路。     

种植密度对冬小麦氮素吸收利用和分配的影响

为了探讨实现冬小麦籽粒产量与氮素利用效率协同提高的途径,为制定高产、高效栽培管理措施提供理论依据,在大田条件下,以大穗型小麦品种"泰农18"和中穗型小麦品种"山农15"为试验材料,根据品种特性分别设置4个种植密度("泰农18":135万苗.hm 2、270万苗.hm 2、405万苗.hm 2和540万苗.hm 2;"山农15":172.5万苗.hm 2、345万苗.hm 2、517.5万苗.hm 2和690万苗.hm 2),研究了种植密度对籽粒产量、氮素吸收积累和运转分配、氮素利用效率以及土壤中硝态氮、铵态氮和无机态氮总积累量的影响。研究结果表明,随种植密度增加,两种穗型冬小麦品种成熟期植株氮素积累量、籽粒产量、氮肥吸收利用效率和氮肥偏生产力均表现为先增加后降低,籽粒氮积累量、氮素收获指数和籽粒氮含量下降,花前营养器官氮素转运量和对籽粒氮的贡献率升高。随种植密度的增加,"泰农18"的氮素利用效率随密度的增大先增大后减小,"山农15"随密度的增大而减小。土壤中硝态氮、铵态氮和无机态氮总积累量随密度增加而降低。在本试验条件下,"泰农18"和"山农15"兼顾高产和高效利用氮素的适宜种植密度分别为270万苗.hm 2和345万苗.hm 2。     

Chlorophyll meter readings,N leaf concentration and their relationship with N use efficiency in oregano

Plant-based diagnostic techniques such as chlorophyll meter (CM) readings and nitrogen (N) leaf concentration are used to determine the level of crop N nutrition, but research is limited on perennial crops and especially on aromatic and medicinal plants such as Greek oregano (Origanum vulgare spp. hirtum). The objectives of this study were to determine in a perennial crop species whether there is a relationship between the CM readings, N leaf concentration, and N use efficiency (NUE) and to use CM readings as a diagnostic tool for predicting dry matter yield in response to N fertilization. The CM readings varied across environments, growth stages, and fertilization treatments and were correlated with NUE. NUE was also affected by the environment, N fertilization, and interactions among these variables. This study provides new information about the effect of N application on CM readings, N leaf concentration, and NUE in a perennial crop such as oregano.     

氮素管理的指标

用什么指标(indicators或indexes)来衡量不同尺度或生产体系氮素管理的优劣,是提高氮素管理水平的核心问题。近年来,国际上在氮素管理方面提出了许多新概念、方法和指标,以客观评价和改进氮素管理水平,为生产者和政策制定者提供评判标准。本文系统介绍了国际上氮素管理指标的概念、含义、计算方法及对结果的解析,包括:(1)氮素投入、收益与环境效应的关系;(2)氮素收支(budget)、氮素平衡(balance)和氮素盈余(surplus);(3)氮素利用率(Nitrogen Use Efficiency,NUE);(4)氮素利用率与氮素输入、输出及盈余的关系;(5)旱地作物收获后土体硝态氮允许残留量(Residual nitrate-N)。在此基础上,汇总了我国在不同尺度、不同土壤-作物体系氮素研究结果,初步建立了我国典型农田的氮素管理指标,为实现优良的氮素管理提供科学依据。     

施氮量对氮高效水稻种质4007的氮素吸收、转运和利用的影响

田间条件下,以当地水稻品种武运粳15(WJ15)为对照,对氮高效水稻种质4007在不同施氮水平(0、100、150、200和250 kg hm-2)下的氮素吸收、累积、转运、产量及氮肥利用率进行了研究。结果表明施氮量显著促进水稻各生育期地上部氮素的累积,水稻从分蘖盛期到拔节期植株氮素累积量最大,占总生育期的32.7%～41.6%。当施氮量从0增加至250 kg hm-2,水稻种质4007的氮素转运量的从72.0上升至104kg hm-2,氮素转运率从66.2%下降至51.3%;而WJ15的氮素转运量从57.0上升至96.5 kg hm-2,氮素转运率也从57.1%下降至46.8%。籽粒中的氮素65.3%～87.6%来自营养器官的转运,仅12.4%～34.7%是后期从土壤吸收所得。由于较高的收获指数(HI)和氮收获指数(NHI),水稻种质4007的平均氮肥表观回收率(REN)和氮肥农学利用率(AEN)分别较品种WJ15高出24.5%和95.6%。本试验结果还显示4007和WJ15的适宜施氮量分别是150 kg hm-2和200 kg hm-2,此时,水稻可维持较高的产量和保持较高的氮肥利用率。     

The significance of plant energy status for the uptake and incorporation of NH4-nitrogen by young rice plants

Uptake and assimilation of inorganic N in young rice plants has been studied with labelled N (N-15). Depletion of the plants' carbohydrate content, obtained by a preceding dark period, resulted in a drastic reduction of NH₄ ⁺-N uptake. Plants exposed to low light intensity showed diminishing NH₄ ⁺-N uptake rates as compared with plants exposed to full light intensity, the latter showing constant NH₄ ⁺-N uptake rates during the whole experimental period. The percentage of labelled insoluble N in total labelled N was not significantly affected by a preceding dark period, whereas the low light intensity resulted in a lower proportion of insoluble N in roots and shoots. The incorporation of labelled N into the insoluble fraction (proteins, nucleic acids) was higher in plants fed with NH₄ ⁺-N than in those fed with NO₃ ^-.

The uptake of NH₄ ⁺-N was not significantly affected by NO₃ ^-, whereas the NO₃- uptake rate was considerably reduced in the presence of NH^{4 +}-N. Low energy status of plants affected the nitrate uptake more than the uptake of NH₄ ⁺-N. The results show that uptake and assimilation of inorganic N depend much on the energetic status of plants. Nitrate uptake and assimilation is more sensitive to low energy conditions than NH₄ ⁺-N.     

Interspecific facilitation between intercropped millets and peanuts: insights from root proteomics analysis

Intercropping is practised globally because of its advantages in terms of productivity and resource use efficiency. However, our knowledge on the molecular mechanisms underlying belowground interspecific interactions in intercropping systems is still very limited. Pot experiments involving both intercropped millet and peanut were conducted to quantify the differentially expressed proteins in each component crop under conditions of complete, partial and no interspecific interactions based on tandem mass tag (TMT) labelling. The results showed that the yields of both crops in the intercropping system increased in response to complete root interactions due to increases in nutrient acquisition as well as increases in root length and surface area. There were 73 differentially expressed proteins in the millet roots and 41 in the peanut roots, most of which were involved in C metabolism, N metabolism, transport and signal transduction. Additional bioinformatic analyses revealed that root interactions improved N and P assimilation via relatively high amounts of proteins such as urease and inorganic phosphate transporter in the millet roots and malate dehydrogenase increased P assimilation related proteins in the peanut roots. These results would contribute to a comprehensive understanding at molecular level in cereal/legume intercropping systems in response to interspecific root interactions.     

Cytokinin-dependent improvement in transgenic P(SARK)::IPT tobacco under nitrogen deficiency

Wild-type (WT) and transgenic tobacco plants overexpressing isopentenyltransferase (IPT), a gene coding the rate-limiting step in cytokinin (CKs) synthesis, were grown under limited nitrogen (N) conditions to evaluate the role of CKs in NUE (N-use efficiency) and in different parameters that determine the quality of tobacco leaves. The results indicate that WT tobacco plants submitted to N deficiency show a decline in the leaf/root ratio, associated with a decrease in the NUE and in tobacco-leaf quality, defined by an increase in the quantity of nicotine. On the contrary, the transgenic plants submitted to N deficiency maintained the leaf/root ratio, presenting a higher NUE and greater quality of tobacco leaves than the WT plants, as the latter showed reduced nicotine and an increase in reducing sugars under severe N-deficiency conditions. Therefore, the overexpression of CKs under N deficiency could be a useful tool to improve tobacco cultivation, given that it could reduce N-fertilizer application and thereby provide economic savings and environmental benefits, maintaining yield and improving tobacco leaf quality.     

不同氮效率油菜SPS和PEPC活性差异及其对籽粒产量与油分含量的影响

【目的】蔗糖磷酸合成酶(sucrose phosphate synthase,SPS)与磷酸烯醇式丙酮酸羧化酶(phosphoenolpyruvate carboxylase,PEPC)分别控制着植物体内的碳骨架向碳代谢和氮代谢的流转,影响作物的产量与品质。为探明氮高效油菜品种在高效利用氮素的同时协调籽粒蛋白与油分累积矛盾的机理,研究了不同氮效率油菜品种的SPS与PEPC活性差异及其对籽粒油分含量的影响。【方法】采用土培试验,以氮高效品种27号(H)与氮低效品种6号(L)为试验材料,在常氮(N)与低氮(S)条件下,研究不同氮效率油菜品种苗期到花期叶片与角果发育初期的角果、角果发育中期的角果皮与籽粒中SPS与PEPC活性变化及差异、生长后期碳素转运量与转运率以及收获期籽粒油分含量的差异。【结果】两种供氮水平下,氮高效品种27号的产量与籽粒油分含量均显著高于氮低效品种6号,品种优势明显;且氮高效品种27号苗期到花期叶片与角果发育初期的角果、角果发育中期的角果皮与籽粒中的SPS与PEPC活性均高于氮低效品种6号,两种供氮水平的规律相同,但是SPS与PEPC活性的比值(SPS/PEPC)却因生育期不同而异,营养生长期叶片中氮高效品种27号的SPS/PEPC高于氮低效品种6号,开花期品种间叶片SPS/PEPC相近,角果发育期主要生殖器官中的SPS/PEPC值氮高效品种反而低于氮低效品种。说明氮高效品种向碳代谢和氮代谢输送的碳骨架在全生育期均多于氮低效品种,而碳代谢对氮代谢的响应只在生育前期强于氮低效品种,生育后期则相反。碳素转运量与转运率、籽粒油分含量与产量也是氮高效品种大于氮低效品种,这可能为氮高效品种协调籽粒蛋白与油分累积矛盾的重要生理机制。供氮水平对上述各指标有不同的影响,籽粒产量、PEPC活性、碳素转运量及转运率以常氮处理高于低氮处理,而油分含量、SPS活性及SPS/PEPC以常氮处理低于低氮处理,但不改变以上指标的品种间差异。【结论】与氮低效品种相比,氮高效品种全生育期向碳、氮代谢均输送更多的碳骨架,这是氮高效品种缓解碳、氮代谢矛盾的重要前提;碳代谢对氮代谢的响应生育前期较高、生育后期较低,同时生育后期有更多营养器官的碳素转运到籽粒,也为油菜生育后期满足籽粒碳、氮代谢所需要的碳骨架,并协调籽粒油分与蛋白质含量的矛盾提供了条件。     

施用方式和氮肥种类对砂姜黑土氮素迁移的影响

采用田间微区试验,在砂姜黑土中研究了施肥方式(上层12 cm土混施、土下12 cm点施、土下12 cm条施)和氮肥种类(尿素、磷酸氢二铵)对氮素垂直运移和水平迁移动态的影响。不同施用方式试验结果表明,在处理的90 d内,砂姜黑土中土壤NH_4~+-N和NO_3~–-N含量均呈现土下12 cm点施土下12 cm条施上层12 cm土混施的趋势。尿素在土下12 cm点施条件下,土壤NH_4~+-N主要集中在垂直方向6~18 cm土层和水平距离0~7 cm范围内;而NO_3~–-N的分布核心区土层超过21 cm,水平距离大于15 cm;NH_4~+-N和NO_3~–-N核心区浓度均随处理时间延长而明显下降。土下12 cm点施90 d后,尿素和磷酸铵的氮素养分在砂姜黑土中的横向移动距离为5~7 cm,垂直方向上养分主要集中在6~18 cm的土层范围;点施90 d时,磷酸铵处理在土下18 cm和水平距离12 cm处无机态氮(NH_4~+-N和NO_3~–-N)含量分别为148.9和77.4 mg/kg,其含量远大于尿素处理(96.3和53.2 mg/kg),而在施肥点两种氮肥处理土壤无机态氮含量差异更大,说明磷酸铵较尿素具有更高的保肥性。研究表明:点施延缓了NH_4~+-N向NO_3~–-N转化速率,提高了肥际养分供应浓度。结合作物生长和需肥特性,预示通过优化施肥位置和氮肥种类,采用一次施肥可以实现90 d持续供应高浓度养分以满足旱地作物生长发育的养分需求。