Physiological measurements of nitrate (NO₃⁻) uptake by roots have defined two systems of high and low affinity uptake. In Arabidopsis, genes encoding both of these two uptake systems have been identified. Most is known about the high affinity transport system (HATS) and its regulation and yet measurements of soil NO₃⁻ show that it is more often available in the low affinity range above 1 mM concentration. Several different regulatory mechanisms have been identified for AtNRT2.1, one of the membrane transporters encoding HATS; these include feedback regulation of expression, a second component protein requirement for membrane targeting and phosphorylation, possibly leading to degradation of the protein. These various changes in the protein may be important for a second function in sensing NO₃⁻ availability at the surface of the root. Another transporter protein, AtNRT1.1 also has a role in NO₃⁻ sensing that, like AtNRT2.1, is independent of their transport function. From the range of concentrations present in the soil it is proposed that the NO₃⁻-inducible part of HATS functions chiefly as a sensor for root NO₃⁻ availability. Two other key NO₃⁻ transport steps for efficient nitrogen use by crops, efflux across membranes and vacuolar storage and remobilization, are discussed. Genes encoding vacuolar transporters have been isolated and these are important for manipulating storage pools in crops, but the efflux system is yet to be identified. Consideration is given to how well our molecular and physiological knowledge can be integrated as well to some key questions and opportunities for the future.

Cellular pH homeostasis is fundamental for life, and all cells adapt to maintain this balance. In plants, the chemical form of nitrogen supply, nitrate and ammonium, is one of the cellular pH dominators. We report that the rice nitrate transporter OsNRT2.3 is transcribed into two spliced isoforms with a natural variation in expression ratio. One splice form, OsNRT2.3b is located on the plasma membrane, is expressed mainly in the phloem, and has a regulatory motif on the cytosolic side that acts to switch nitrate transport activity on or off by a pH-sensing mechanism. High OsNRT2.3b expression in rice enhances the pH-buffering capacity of the plant, increasing N, Fe, and P uptake. In field trials, increased expression of OsNRT2.3b improved grain yield and nitrogen use efficiency (NUE) by 40%. These results indicate that pH sensing by the rice nitrate transporter OsNRT2.3b is important for plant adaption to varied N supply forms and can provide a target for improving NUE. Significance Significant progress has been made in our understanding of plant adaptive responses to maintain cellular pH under varied N supply forms. Rice is a plant adapted to grow in waterlogged or dryland environments, in contrast to other crops, such as wheat, soybean, and maize. The nitrate transporter OsNRT2.3b provides a molecular mechanism explaining plant adaptation to the ammonium-nitrate supply shift between the waterlogged and drained soil environments. The sensing of cytosolic pH by OsNRT2.3b can function to improve rice nitrogen use efficiency and pH balance, providing an explanation for plant adaptation to changes in the form of N supply.

Organic acids (OAs) are central to cellular metabolism. Many plant stress responses involve the exudation of OAs at the root–soil interface, which can improve soil mineral acquisition and toxic metal tolerance. Because of their simple structure, the low-molecular-weight OAs are widely studied. We discuss the conventional roles of OAs, and some newly emerging roles in plant stress tolerance. OAs are more versatile in their role in plant stress tolerance and are more efficient chelating agents than other acids, such as amino acids. Root OA exudation is important in soil carbon sequestration. These functions are key processes in combating climate change and helping with more sustainable food production. We briefly review the mechanisms behind enhanced biosynthesis, secretion, and regulation of these activities under different stresses, and provide an outline of the transgenic approaches targeted towards the enhanced production and secretion of OAs. A recurring theme of OAs in plant biology is their role as ‘acids’ modifying pH, as ‘chelators’ binding metals, or as ‘carbon sources’ for microbes. We argue that these multiple functions are key factors for understanding these molecules’ important roles in plant stress biology. Finally, we discuss how the functions of OAs in plant stress responses could be used, and identify the important unanswered questions.

The identification of a family of NAR2-type genes in higher plants showed that there was a homolog in Arabidopsis (Arabidopsis thaliana), AtNAR2.1. These genes encode part of a two-component nitrate high-affinity transport system (HATS). As the Arabidopsis NRT2 gene family of nitrate transporters has been characterized, we tested the idea that AtNAR2.1 and AtNRT2.1 are partners in a two-component HATS. Results using the yeast split-ubiquitin system and Xenopus oocyte expression showed that the two proteins interacted to give a functional HATS. The growth and nitrogen (N) physiology of two Arabidopsis gene knockout mutants, atnrt2.1-1 and atnar2.1-1, one for each partner protein, were compared. Both types of plants had lost HATS activity at 0.2 mm nitrate, but the effect was more severe in atnar2.1-1 plants. The relationship between plant N status and nitrate transporter expression revealed a pattern that was characteristic of N deficiency that was again stronger in atnar2.1-1. Plants resulting from a cross between both mutants (atnrt2.1-1 × atnar2.1-1) showed a phenotype like that of the atnar2.1-1 mutant when grown in 0.5 mm nitrate. Lateral root assays also revealed growth differences between the two mutants, confirming that atnar2.1-1 had a stronger phenotype. To show that the impaired HATS did not result from the decreased expression of AtNRT2.1, we tested if constitutive root expression of a tobacco (Nicotiana plumbaginifolia) gene, NpNRT2.1, previously been shown to complement atnrt2.1-1, can restore HATS to the atnar2.1-1 mutant. These plants did not recover wild-type nitrate HATS. Taken together, these results show that AtNAR2.1 is essential for HATS of nitrate in Arabidopsis.

Despite being some of the most important crops globally, there has been limited research on forages when compared with cereals, fruits and vegetables. This review summarises the literature highlighting the significance of forage crops, the current improvements and some of future directions for improving yield and nutritional quality. We make the point that the knowledge obtained from model plant and grain crops can be applied to forage crops. The timely development of genomics and bioinformatics together with genome editing techniques offer great scope to improve forage crops. Given the social, environmental and economic importance of forage across the globe and especially in poorer countries, this opportunity has enormous potential to improve food security and political stability.

Ion concentrations in the roots of two barley (Hordeum vulgare) varieties that differed in NaCl tolerance were compared after exposure to NaCl. Triple-barreled H+-, K+-, and Na+-selective microelectrodes were used to measure cytosolic activities of the three ions after 5 and 8 d of NaCl stress. In both varieties of barley, it was only possible to record successfully from root cortical cells because the epidermal cells appeared to be damaged. The data show that from the 1st d of full NaCl stress, there were differences in the way in which the two varieties responded. At 5 d, the tolerant variety maintained a 10-fold lower cytosolic Na+ than the more sensitive variety, although by 8 d the two varieties were not significantly different. At this time, the more tolerant variety was better at maintaining root cytosolic K+ in the high-NaCl background than was the more sensitive variety. In contrast to earlier work on K+-starved barley (Walker et al., 1996), there was no acidification of the cytosol associated with the decreased cytosolic K+ activity during NaCl stress. These single-cell measurements of cytosolic and vacuolar ion activities allow calculation of thermodynamic gradients that can be used to reveal (or predict) the type of active transporters at both the plasma membrane and tonoplast.