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Plant growth and resilience to abiotic stresses, such as soil salinity and drought, depend intricately on nitrogen metabolism. This review explores nitrogen's regulatory role in plant responses to these challenges, unveiling a dynamic interplay between nitrogen availability and abiotic stress. In the context of soil salinity, a nuanced relationship emerges, featuring both antagonistic and synergistic interactions between salinity and nitrogen levels. Salinity-induced chlorophyll depletion in plants can be alleviated by optimal nitrogen supplementation; however, excessive nitrogen can exacerbate salinity stress. We delve into the complexities of this interaction and its agricultural implications. Nitrogen, a vital element within essential plant structures like chloroplasts, elicits diverse responses based on its availability. This review comprehensively examines manifestations of nitrogen deficiency and toxicity across various crop types, including cereals, vegetables, legumes, and fruits. Furthermore, we explore the broader consequences of nitrogen products, such as N2O, NO2, and ammonia, on human health. Understanding the intricate relationship between nitrogen and salinity, especially chloride accumulation in nitrate-fed plants and sodium buildup in ammonium-fed plants, is pivotal for optimizing crop nitrogen management. However, prudent nitrogen use is essential, as overapplication can exacerbate nitrogen-related issues. Nitrogen Use Effi- ciency (NUE) is of paramount importance in addressing salinity challenges and enhancing sustainable crop productivity. Achieving this goal requires advancements in crop varieties with efficient nitrogen utilization, precise timing and placement of nitrogen fertilizer application, and thoughtful nitrogen source selection to mitigate losses, particularly urea-based fertilizer volatilization. This review article delves into the multifaceted world of plant nitrogen metabolism and its pivotal role in enabling plant resilience to nutritional stress and abiotic challenges. It offers insights into future directions for sustainable agriculture. [ABSTRACT FROM AUTHOR]

Nitrogen (N) is a crucial nutrient that plays a significant role in enhancing crop yield. Its availability, including both supply and deficiency, serves as a crucial signal for plant development. However, excessive N use in agriculture leads to environmental and economic issues. Enhancing nitrogen use efficiency (NUE) is, therefore, essential to minimize negative impacts. Prior studies have investigated the genetic factors involved in N responses and the process of low-nitrogen (LN) adaptation. In this review, we discuss recent advances in understanding how epigenetic modifications, including DNA methylation, histone modification, and small RNA, participate in the regulation of N response and LN adaptation. We highlight the importance of decoding the epigenome at various levels to accelerate the functional study of how plants respond to N availability. Understanding the epigenetic control of N signaling and adaptation can lead to new strategies to improve NUE and enhance crop productivity sustainably. [ABSTRACT FROM AUTHOR]

Nitrogen (N) is a crucial nutrient that plays a significant role in enhancing crop yield. Its availability, including both supply and deficiency, serves as a crucial signal for plant development. However, excessive N use in agriculture leads to environmental and economic issues. Enhancing nitrogen use efficiency (NUE) is, therefore, essential to minimize negative impacts. Prior studies have investigated the genetic factors involved in N responses and the process of low-nitrogen (LN) adaptation. In this review, we discuss recent advances in understanding how epigenetic modifications, including DNA methylation, histone modification, and small RNA, participate in the regulation of N response and LN adaptation. We highlight the importance of decoding the epigenome at various levels to accelerate the functional study of how plants respond to N availability. Understanding the epigenetic control of N signaling and adaptation can lead to new strategies to improve NUE and enhance crop productivity sustainably.

C-terminally encoded peptides (CEP), post-translationally modified peptide hormones, play important roles in plant development. CEP peptides are known as regulators of the root system in response to abiotic stress factors, in particular, to a nitrogen deficiency. However, the members of the CEP family in potato were not yet characterized. In our work, we identified 17 CEP genes in the potato (Solanum tuberosum) genome. Using available transcriptomic data, we estimated the expression levels of the CEP genes in different organs and found that the expression levels of some of the CEP genes depend on the amount of nitrogen in the medium. Further study of the CEP genes in potato will expand our understanding of the role of these signal peptides in potato development. [ABSTRACT FROM AUTHOR]

Plant developmental processes are very flexible and highly depend on environmental factors. This is largely due to the existence of regulatory mechanisms that systemically control development on the whole plant level. In plants, regulatory peptides produced in the roots have been identified that are activated in response to different factors influencing root system, such as variation in the level of macronutrients (first of all, nitrogen and phosphorus) in the soil, influence of symbiotic microorganisms (soil rhizobial bacteria and arbuscular mycorrhiza fungi), and water deficiency. Among the systemically acting peptides, the most thoroughly investigated are CLE (CLAVATA3/EMBRYO SURROUNDING REGION-related) and CEP (C-TERMINALLY ENCODED PEPTIDES) peptides that are capable of travelling through the xylem from the roots to the shoot and triggering responses via binding to specific receptors operating in the phloem of the leaf. This review focuses on the role of these two groups of peptides in molecular dialog between the root and shoot. [ABSTRACT FROM AUTHOR]

Among all essential mineral elements, nitrogen (N) is required in the largest amounts and thus is often a limiting factor for plant growth. N is taken up by plant roots in the form of water-soluble nitrate, ammonium, and, depending on abundance, low-molecular weight organic N. In soils, the availability and composition of these N forms can vary over space and time, which exposes roots to various local N signals that regulate root system architecture in combination with systemic signals reflecting the N nutritional status of the shoot. Uncovering the molecular mechanisms underlying N-dependent signaling provides great potential to optimize root system architecture for the sake of higher N uptake efficiency in crop breeding. In this review, we summarize prominent signaling mechanisms and their underlying molecular players that derive from external N forms or the internal N nutritional status and modulate root development including root hair formation and gravitropism. We also compare the current state of knowledge of these pathways between Arabidopsis and graminaceous plant species. [ABSTRACT FROM AUTHOR]

While decades of research have considered redox metabolism as purely defensive, recent results show that reactive oxygen species (ROS) are necessary for growth and development. Close relationships have been found between the regulation of nitrogen metabolism and ROS in response to both carbon and nitrogen availability. Root nitrate uptake and nitrogen metabolism have been shown to be regulated by a signal from the oxidative pentose phosphate pathway (OPPP) in response to carbon signaling. As a major source of NADP(H), the OPPP is critical to maintaining redox balance under stress situations. Furthermore, recent results suggest that at least part of the regulation of the root nitrate transporter by nitrogen signaling is also linked to the redox status of the plant. This leads to the question of whether there is a more general role of redox metabolism in the regulation of nitrogen metabolism by carbon and nitrogen. This review highlights the role of the OPPP in carbon signaling and redox metabolism, and the interaction between redox and nitrogen metabolism. We discuss how redox metabolism could be an important player in the regulation of nitrogen metabolism in response to carbon/nitrogen interaction and the implications for plant adaptation to extreme environments and future crop development. [ABSTRACT FROM AUTHOR]

The control of cell fate, growth and proliferation in response to nitrogen availability is a tightly controlled process, with the two TOR complexes (TORC1 and TORC2) and their effectors playing a central role. PP2A-B55 has recently been shown to be a key element in this response in fission yeast, where it regulates cell cycle progression and sexual differentiation. Importantly, a recent study from our group has shown that PP2A-B55 acts as a mediator between the activities of the two TOR signaling modules, enabling a crosstalk that is required to engage in the differentiation program. In this review, we recapitulate the studies that have led to our current understanding of the interplay between TOR complexes. Moreover, we discuss several aspects of the response to nitrogen availability that still require further attention, and which will be important in the future to fully realize the implications of phosphatase activity in the context of TOR signaling. [ABSTRACT FROM AUTHOR]

Ammonium is a major inorganic nitrogen source for plants. At low external supplies, ammonium promotes plant growth, while at high external supplies it causes toxicity. Ammonium triggers rapid changes in cytosolic pH, in gene expression, and in post-translational modifications of proteins, leading to apoplastic acidification, co-ordinated ammonium uptake, enhanced ammonium assimilation, altered oxidative and phytohormonal status, and reshaped root system architecture. Some of these responses are dependent on AMT-type ammonium transporters and are not linked to a nutritional effect, indicating that ammonium is perceived as a signaling molecule by plant cells. This review summarizes current knowledge of ammonium-triggered physiological and morphological responses and highlights existing and putative mechanisms mediating ammonium signaling and sensing events in plants. We put forward the hypothesis that sensing of ammonium takes place at multiple steps along its transport, storage, and assimilation pathways. [ABSTRACT FROM AUTHOR]

In non-climacteric fruit tree species, color change is primarily controlled by the interplay between environmental conditions, nutritional factors (nitrogen and sugars) and hormones, mainly abscisic acid and gibberellins (GA), through a complex mechanism which is not completely understood. Nitrogen has a strong impact on color change, influenced by environmental changes, either locally or at the whole tree level. We use Citrus trees, as a non-climacteric model species, to understand the long-distance nitrogen signaling mechanism delaying the chloroplast-to-chromoplast transition at the molecular level. It is unknown whether nitrogen regulates the branch point of geranylgeranyl diphosphate (GGPP), the shared precursor for gibberellin, chlorophyll and carotenoid synthesis. We used 15N to trace the root-to-fruit nitrogen flux in trees grown under soils differing in nitrogen content and temperature, and treated with NH 4 NO 3 and GA. Key genes involved in nitrogen signaling, transport and metabolism, and those from the GGPP branch point were analyzed in the fruit. Results explain how soil temperature modifies 15N transport to play a key role in signaling citrus color change, and show that fruit-nitrogen remobilization is required for triggering degreening. Nitrogen content in the fruit modulates the onset of glutamate deamination, asparagine synthesis, nitrite assimilation and GA 1 depletion. Expression of the nitrate transporter NRT1.2, glutamate dehydrogenase and asparagine synthetase genes was high right before the start of degreening, together with NH 4 + concentration. Nitrogen delayed carotenoid synthesis (phytoene synthase gene expression) without modifying gibberellin synthesis (ent -kaurene synthase and oxidase) at the chloroplast level. [Display omitted] • N and GA 1 depletion from the fruit is a prerequisite for citrus color change. • Cold-soil temperatures reduce 15N translocation and trigger fruit color change. • Low N flux correlates fruit protein hydrolysis, upregulating GDH, AS and NRT1.2. • GA treatment delays protein hydrolysis, NRT1.2 expression and N depletion. • The first steps of synthesis of GA in the chloroplast are not modified by N. [ABSTRACT FROM AUTHOR]

Genes containing domains related to glutamine synthetase of the prokaryotic type (GSI-like) are widespread in higher plants, but their function is currently unknown. To gain insights into the possible role of GSI-like proteins, we characterized the GSI-like gene family of Medicago truncatula and investigated the functionality of the encoded proteins. M. truncatula contains two-expressed GSI-like genes, MtGSIa and MtGSIb , encoding polypeptides of 454 and 453 amino acids, respectively. Heterologous complementation assays of a bacterial glnA mutant indicate that the proteins are not catalytically functional for glutamine synthesis. Gene expression was investigated by qRT-PCR and western blot analysis in different organs of the plant and under different nitrogen (N) regimes, revealing that both genes are preferentially expressed in roots and root nodules, and that their expression is influenced by the N-status of the plant. Analysis of transgenic plants expressing MtGSI-like -promoter- gusA fusion, indicate that the two genes are strongly expressed in the root pericycle, and interestingly, the expression is enhanced at the sites of nodule emergence being particularly strong in specific cells located in front of the protoxylem poles. Taken together, the results presented here support a role of GSI-like proteins in N sensing and/or signaling, probably operating at the interface between perception of the N-status and the developmental processes underlying both root nodule and lateral root formation. This study indicates that GSI-like genes may represent a novel class of molecular players of the N-mediated signaling events. [ABSTRACT FROM AUTHOR]

The dynamic nature of gene regulatory networks allows cells to rapidly respond to environmental change. However, the underlying temporal connections are missed, even in kinetic studies as transcription factor (TF) binding within at least one time point is required to identify primary targets. The TF-regulated but unbound genes are dismissed as secondary targets. Instead we report that these genes comprise transient TF-target interactions most relevant to rapid signal transduction. We temporally perturbed a master TF (Basic Leucine Zipper 1, bZIP1) and the nitrogen (N) signal it transduces and integrated TF regulation and binding data from the same cell samples. Our enabling approach could identify primary TF targets based solely on gene regulation in the absence of TF binding. We uncovered three classes of primary TF targets: (i) poised (TF-bound but not TF-regulated), (ii) stable (TF-bound and TF-regulated), and (iii) transient (TF-regulated but not TF-bound), the largest class. Unexpectedly, the transient bZIPI targets are uniquely relevant to rapid N signaling in planta enriched in dynamic N-responsive genes, and regulated by TF and N signal interactions. These transient targets include early N responders nitrate transporter 2.1 and NIN-like protein 3, bound by bZIPI at 1-5 min, but not at later time points following TF perturbation Moreover, promoters of these transient targets are uniquely enriched with os-regulatory motifs coinherited with bZIPI binding sites, suggesting a recruitment role for bZIPL This transient mode of TF action supports a classic, but forgotten, "hit-and-run" transcription model, which enables a "catalyst TF" to activate a large set of targets within minutes of signal perturbation. [ABSTRACT FROM AUTHOR]

Morphological plasticity of root systems is critically important for plant survival because it allows plants to optimize their capacity to take up water and nutrients from the soil environment. Here we show that a signaling module composed of nitrogen (N)-responsive CLE (CLAVATA3/ESR-related) peptides and the CLAVATA1 (CLV1) leucine-rich repeat receptor-like kinase is expressed in the root vasculature in Arabidopsis thaliana and plays a crucial role in regulating the expansion of the root system under N-deficient conditions. CLE1, -3, -4, and -7 were induced by N deficiency in roots, predominantly expressed in root pericycle cells, and their overexpression repressed the growth of lateral root primordia and their emergence from the primary root. In contrast, clv1 mutants showed progressive outgrowth of lateral root primordia into lateral roots under N-deficient conditions. The clv1 phenotype was reverted by introducing a CLV1 promoter-driven CLV1:GFP construct producing CLV1:GFP fusion proteins in phloem companion cells of roots. The overaccumulation of CLE2, -3, -4, and -7 in clv1 mutants suggested the amplitude of the CLE peptide signals being feedback-regulated by CLV1. When CLE3 was overexpressed under its own promoter in wild-type plants, the length of lateral roots was negatively correlated with increasing CLE3 mRNA levels; however, this inhibitory action of CLE3 was abrogated in the clv1 mutant background. Our findings identify the N-responsive CLECLV1 signaling module as an essential mechanism restrictively controlling the expansion of the lateral root system in N-deficient environments. [ABSTRACT FROM AUTHOR]

In plants, root nitrate uptake systems are under systemic feedback repression by the N satiety of the whole organism, thus adjusting the N acquisition capacity to the N demand for growth; however, the underlying molecular mechanisms are largely unknown. We previously isolated the Arabidopsis high nitrogen-insensitive 9-1 (hni9-1) mutant, impaired in the systemic feedback repression of the root nitrate transporter NRT2.1 by high N supply. Here, we show that HNI9 encodes Arabidopsis INTERACT WITH SPT6 (AtIWS1), an evolutionary conserved component of the RNA polymerase II complex. HNI9/AtIWS1 acts in roots to repress NRT2.1 transcription in response to high N supply. At a genomic level, HNI9/AtIWS1 is shown to play a broader role in N signaling by regulating several hundred N-responsive genes in roots. Repression of NRT2.1 transcription by high N supply is associated with an HNI9/AtIWS1-dependent increase in histone H3 lysine 27 trimethylation at the NRT2.1 locus. Our findings highlight the hypothesis that posttranslational chromatin modifications control nutrient acquisition in plants. [ABSTRACT FROM AUTHOR]

Activation of phospholipase D (PLD) produces phosphatidic acid (PA), a lipid messenger implicated in cell growth and proliferation, but direct evidence for PLD and PA promotion of growth at the organism level is lacking. Here we characterize a new PLD gene, PLDε, and show that it plays a role in promoting Arabidopsis growth. PLDε is mainly associated with the plasma membrane, and is the most permissive of all PLDs tested with respect to its activity requirements. Knockout (KO) of PLDε decreases root growth and biomass accumulation, whereas over-expression (OE) of PLDε enhances root growth and biomass accumulation. The level of PA was higher in OE plants, but lower in KO plants than in wild-type plants, and suppression of PLD-mediated PA formation by alcohol alleviated the growth-promoting effect of PLDε. OE and KO of PLDε had opposite effects on lateral root elongation in response to nitrogen. Increased expression of PLDε also promoted root hair elongation and primary root growth under severe nitrogen deprivation. The results suggest that PLDε and PA promote organism growth and play a role in nitrogen signaling. The lipid-signaling process may play a role in connecting membrane sensing of nutrient status to increased plant growth and biomass production. [ABSTRACT FROM AUTHOR]

Summary: This work aimed at identifying a possible role of phytohormones in long-distance (root–shoot) signaling under nitrogen deficiency. Three-months old pineapple plants were transferred from Murashige and Skoog (MS) medium to nitrogen-free MS (−N). During the first 24h on −N, 20 plants were harvested every 4h. After 30 days in −N, the remaining plants were transferred back to regular MS (+N) and 20 plants harvested every 4h for the first 24h. Following the harvests, endogenous levels of nitrate (NO3 −), indole-3-acetic acid (IAA), isopentenyladenine (iP), isopentenyladenine riboside (iPR), zeatin (Z) and zeatin riboside (ZR) were analyzed in roots and leaves. In N-starved plants, the NO3 − level dropped by 20% in roots between the first (4h) and the second harvest (8h). In leaves a reduction of 20% was found 4h later. Accumulation of IAA peaked in leaves at 16h. In roots, the accumulation of IAA only started at 16h while the leaf content was already in decline, which suggests that the hormone might have traveled from the leaves to the roots, communicating N-shortage. The contents of the four cytokinins were generally low in both, shoot and roots, and remained almost unchanged during the 24h of analysis. After N re-supply, roots showed a NO3 − peak at 8h whereas the foliar concentration increased 4h later. Hormone levels in roots climaxed at 8h, this coinciding with the highest NO3 − concentration. In leaf tissue, a dramatic accumulation was only observed for Z and ZR, and the peak was seen 4h later than in roots, suggesting that Z-type cytokinins might have traveled from the roots to the leaves. These findings provide evidence that there is a signaling pathway for N availability in pineapple plants, communicated upwards through cytokinins (N-supplemented plants) and downwards through auxin (N-starved plants). [Copyright &y& Elsevier]

Under nitrogen (N) starvation, certain microalgae increase carbon storage in the form of lipid droplets while also downregulating photosynthesis and eventually terminating growth. To improve lipid yield, we asked whether lipid droplets and N starvation responses can be induced without limiting growth or photosynthesis. In the chlorophyte Chlamydomonas reinhardtii , N starvation induces gametogenesis alongside lipid droplet accumulation, and gametogenesis has been observed in arginine-fed cultures wherein the arginine was provided as the sole N source. We therefore assessed whether arginine-fed cultures displayed N starvation responses other than gametogenesis in mixo- and phototrophic conditions, representing two primary modes of nutrition that may affect N starvation responses. We showed that arginine-fed cultures using two lab strains of C. reinhardtii supported normal mixotrophic growth, constitutively turned on N starvation-induced genes at the equivalent level to N-starved cells, and increased the triacylglycerol content of total fatty acids by 125–400% relative to ammonium-fed cultures. The lipid profile of triacylglycerol in these arginine-fed mixotrophic cultures exhibited 3 to 5.5-fold enrichment of saturated and monounsaturated fatty acids, a preferred characteristic of biodiesel precursors. Arginine-fed phototrophic cultures likewise turned on N starvation-induced genes, accumulated lipid droplets, and led to a 50% reduced growth rate per day while reaching a 3 to 6.5-fold more cell density at the stationary phase relative to ammonium-fed cultures. To test the applicability of our result to algae outside the green lineage, we conducted similar experiments with the diatom Phaeodactylum tricornutum , which also accumulated 2 to 7-fold more neutral lipids in arginine-fed phototrophic cultures without growth impairment relative to nitrate-fed cultures. We document a system wherein N starvation responses are induced without compromising photosynthesis or growth, thereby suited to the production of valuable chemicals and biofuel precursors without requiring stressors in microalgae. • Arginine catabolism leads to the activation of nitrogen starvation responses. • Arginine-fed cultures grow up to six times higher density than ammonium-fed cultures. • Triacylglycerol from arginine-fed culture is enriched in saturated and monounsaturated fatty acids. [ABSTRACT FROM AUTHOR]

Ammonium is a major inorganic nitrogen source for plants. At low external supplies, ammonium promotes plant growth, while at high external supplies it causes toxicity. Ammonium triggers rapid changes in cytosolic pH, in gene expression, and in post-translational modifications of proteins, leading to apoplastic acidification, co-ordinated ammonium uptake, enhanced ammonium assimilation, altered oxidative and phytohormonal status, and reshaped root system architecture. Some of these responses are dependent on AMT-type ammonium transporters and are not linked to a nutritional effect, indicating that ammonium is perceived as a signaling molecule by plant cells. This review summarizes current knowledge of ammonium-triggered physiological and morphological responses and highlights existing and putative mechanisms mediating ammonium signaling and sensing events in plants. We put forward the hypothesis that sensing of ammonium takes place at multiple steps along its transport, storage, and assimilation pathways., (© The Author 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)