Increasing nitrogen supply to phosphorus-deficient Medicago sativa decreases shoot growth and enhances root exudation of tartrate to discharge surplus carbon dependent on nitrogen form

Aims: Carboxylate release by roots has been considered a strategy for mobilisation and acquisition of phosphorus (P). However, recently, it was argued that carboxylate release may be a way to discharge surplus carbon produced under conditions that limit plant growth. Plant P status may not be the main factor driving carboxylate release by roots. Instead, plant nitrogen (N) status and/or N:P ratio of the soil or plant may play a more important role in enhancing carboxylate release. Methods: A greenhouse pot experiment was performed to grow alfalfa in a P-deficient soil, supplied with two rates of P (0 and 20 mg kg⁻¹) in combination with four forms of nitrogen (N) at five rates (0, 25, 50, 75, and 100 mg kg⁻¹), to explore the effects of P rate, N form, N rate, and their interactions on plant growth, P and N status, and carboxylate release, and to determine the factors driving carboxylate release. Results: Nitrogen addition weakened the positive effect of P addition on plant growth, and increased plant N and P concentrations; P addition increased plant P concentration, but weakened the effect of N addition on plant N concentration. The amount of tartrate increased dramatically with increasing N rate, which decreased shoot growth, depending on N form. At high P supply, tartrate exudation correlated negatively with shoot biomass. Conclusions: Nitrogen addition to P-deficient alfalfa decreased shoot growth and enhanced the release of tartrate, likely to discharge surplus carbon; and the effects varied with N form.