A dynamic whole-plant model of integrated metabolism of nitrogen and carbon. 2. Balanced growth driven by C fluxes and regulated by signals from C and N substrate

A whole-plant model of C and N metabolism is presented for the juvenile stage. It is aimed at comparing the growth performance of (wild) plant species in a range of environments with respect to irradiance and availability of nitrate (NO(3)(-)) and ammonium (NH(4)(+)). State variables are the structural masses of leaves, stem and root, NO(3)(-) concentrations in root and shoot, non-structural carbohydrate (C) densities in leaves, stem and root and non-structural organic N concentration in the whole plant. Explicit expressions for NO(3)(-) influx, efflux, translocation and assimilation, and for NH(4)(+) uptake and assimilation have been formulated in an accompanying paper. Photosynthetic rate is derived from electron-transport rate which depends on irradiance and chlorophyll concentration on a leaf-area basis. The latter is proportional to non-structural organic N concentration. Photosynthetic N is considered non-structural. Unique features of the model are the use of metabolite signals and the treatment of C allocation and balanced growth. Metabolite signals are dimensionless functions of non-structural compounds (NO(3)(-), C, organic N) and modify rate variables involved in N uptake and assimilation, C allocation and growth. Carbon allocation is driven by concentration differences of the cytosolic C pools in stem and root and is modified by the N status of the plant such that a high N status increases the apparent size of the shoot. Photosynthate is unloaded into C buffers which degrade at a constant specific rate. The sugar fluxes which arise from these buffers drive the growth rate of stem and root. No parameters are included for maximum specific growth or for activity or strength of sinks. Primary stem growth is proportional to growth of the leaf compartment: leaves arise from stems in a modular fashion. Leaves are autonomous with respect to their C balance. The model is presented as a system of differential equations which is integrated numerically. Parameter values, e.g., for uptake and assimilation capacities and costs of uptake, assimilation, maintenance and growth, are estimated for a grass species, Dactylis glomerata. Juvenile growth is simulated under optimal conditions with respect to irradiance and NO(3)(-) availability and compared with literature data. Diurnal and daily patterns of C utilisation and respiration, expressed as percentages of gross photosynthetic rate, are discussed. The model satisfactorily simulates typical responses to nutrient and light limitation and pruning, such as redirected C allocation, adjusted root and leaf weight ratios and compensatory growth. A sensitivity analysis is included for selected parameters.