The role of root nitrate reduction in the systemic control of biomass partitioning between leaves and roots in accordance to the C/N-status of tobacco plants

The root/shoot-ratio is a simple parameter to describe the systemic response of plants to alterations of their nutritional status, as indicated by the C/N-balance of leaves. The ‘functional equilibrium hypothesis’ holds that leaf growth is limited by the supply of nitrogen from the roots, whereas root growth depends on the carbon supply from leaves. The nature of the systemic control that balances root and shoot growth is not fully understood. Previous experiments have shown that root growth of transformed tobacco plants, which lack functional root nitrate reductase, was severely impeded, when plants were grown on NO3− as the sole N-source. In these experiments, the root/shoot-ratio was correlated with the Glutamate/Glutamine-ratio of roots. In the present study we tested the hypothesis that high internal Glu contents (in relation to Gln) inhibit root growth. Wild type and transformed tobacco plants were given access to both NH4 and NO3, and were cultivated at ambient and elevated pCO2 in order to vary carbon availability. The uptake and assimilation of NH4+ by the root was significantly higher in transformed than in wild type tobacco, in particular at elevated pCO2. Consequently, the Glu/Gln-ratio in the root of transformants was significantly lower than in NO3− -grown plants, and was, in the present study, not different from the wild type. However, we failed to observe a correlation between plant architecture and the Glu/Gln-ratio of roots, suggesting that signals arising from the immediate products of nitrate reduction (nitrite) are involved in the systemic control of root growth. Furthermore the synthesis of root-derived signals, which affect N-turnover, starch re-mobilization and the growth of leaves, appears to be associated with root nitrate reduction. This enzymatic step seems to be indispensable for the systemic control of biomass partitioning, and plays a crucial role for the integration of carbon and nitrogen metabolism at the whole plant level.