Eco-Evolutionary Optimal Carbon Allocation in a MechanisticCrop Growth Model: Theory and Application to Wheat

Carbon allocation governs how plants partition assimilates among leaves, roots, stems, and reproductive organs, directly shaping growth and yield. Most crop models represent this partitioning using fixed or empirically derived coefficients, which limits their ability to predict how allocation responds to genotype, environment, or their interaction. Eco-evolutionary optimality theory offers an alternative where the allocation strategy is allowed to emerge from the marginal costs and benefits of investing carbon in each organ. While this approach has been applied successfully to trees, it has not previously been developed for herbaceous plants. Here, we present DAESIM2-Plant, a mechanistic plant growth model that couples physiological processes including photosynthesis, stomatal conductance, plant hydraulics, and canopy radiative transfer, with an eco-evolutionary optimal allocation scheme for leaves and roots. This is applied to wheat by coupling it to a source-sink grain production module. Using a series of idealized sensitivity experiments, we show that the model reproduces well-documented patterns of plasticity in carbon allocation: diminishing returns on leaf investment as canopy closure and water limitation are approached, a shift in allocation toward roots under drier conditions, and a root:shoot response that depends jointly on soil moisture and the plant's existing root:leaf balance. Simulating a full growing season across a gradient of soil moisture levels reveals a threshold-like response in canopy development, biomass accumulation, and grain yield, with yield constrained by the same assimilate supply that governs vegetative growth during the critical period and grain filling. This provides a promising basis for generalizing model behaviour across a wide range of environmental conditions. Evaluating these results against experimental and field trial data remains an important next step.