NRT1.1 Dual-Affinity Nitrate Transport/Signalling and its Roles in Plant Abiotic Stress Resistance

NRT1.1 is the first nitrate transport protein cloned in plants and has both high- and low-affinity functions. It imports and senses nitrate, which is modulated by the phosphorylation on Thr101 (T101). Structural studies have revealed that the phosphorylation of T101 either induces dimer decoupling or increases structural flexibility within the membrane, thereby switching the NRT1.1 protein from a low- to high-affinity state. Further studies on the adaptive regulation of NRT1.1 in fluctuating nitrate conditions have shown that, at low nitrate concentrations, nitrate binding only at the high-affinity monomer initiates NRT1.1 dimer decoupling and priming of the T101 site for phosphorylation activated by CIPK23, which functions as a high-affinity nitrate transceptor. However, nitrate binding in both monomers retains the unmodified NRT1.1, maintaining the low-affinity mode. This NRT1.1-mediated nitrate signalling and transport may provide a key to improving the efficiency of plant nitrogen use. However, recent studies have revealed that NRT1.1 is extensively involved in plant tolerance of several adverse environmental conditions. In this context, we summarise the recent progress in the molecular mechanisms of NRT1.1 dual-affinity nitrate transport/signalling and focus on its expected and unexpected roles in plant abiotic stress resistance and their regulation processes.

Regulation of seed dormancy and germination by nitrate

Nitrate promotes seed germination at low concentrations in many plant species, and functions as both a nutrient and a signal. As a nutrient, it is assimilated via nitrite to ammonium, which is then incorporated into amino acids. Nitrate reductase (NR) catalyses the reduction of nitrate to nitrite, the committed step in the assimilation. Seed sensitivity to nitrate is affected by other environmental factors, such as light and after-ripening, and by genotypes. Mode of nitrate action in seed germination has been well documented in Arabidopsis thaliana and the hedge mustard Sisymbrium officinale. In these species nitrate promotes seed germination independent of its assimilation by NR, suggesting that it acts as a signal to stimulate germination. In Arabidopsis, maternally applied nitrate affects the degree of primary dormancy in both wild-type and mutants defective in NR. This indicates that nitrate acts not only during germination, but also during seed development to negatively regulate primary dormancy. Functional genomics studies in Arabidopsis have revealed that nitrate elicits downstream events similar to other germination stimulators, such as after-ripening, light and stratification, suggesting that these distinct environmental signals share the same target(s). In Arabidopsis, the NIN-like protein 8 (NLP8) transcription factor, which acts downstream of nitrate signalling, induces nitrate-dependent gene expression. In particular, a gene encoding the abscisic acid (ABA) catabolic enzyme CYP707A2 is directly regulated by NLP8. This regulation triggers a nitrate-induced ABA decrease that permits seed germination. This review article summarizes an update of our current understanding of the regulation of seed dormancy and germination by nitrate.