HydroShoot: a functional-structural plant model for simulating hydraulic structure, gas and energy exchange dynamics of complex plant canopies under water deficit - application to grapevine (Vitis vinifera L.)

This paper aims at presenting HydroShoot, a functional-structural plant model (FSPM) that is developed to simulate gas-exchange rates of complex plant canopies under water deficit conditions, by scaling up gas-exchange rates from the leaf to the canopy levels. The main hypothesis is that simulating both the hydraulic structure of the shoot together with the energy budget of individual leaves is the asset for successfully achieving this up-scaling task. HydroShoot was hence built as the ensemble of three interacting modules: hydraulic which calculates the distribution of xylem water potential across shoot hydraulic segments, energy which calculates the complete energy budget of individual leaves, and exchange which calculates net assimilation and transpiration rates of individual leaves. HydroShoot was coupled with irradiance interception and soil water balance models, and was evaluated on virtual and real grapevines having strongly contrasted canopies, under well-watered and water-deficit conditions. HydroShoot captured accurately the impact of canopy architecture and the varying soil water deficit conditions on plant-scale gas-exchange rates and leaf-scale temperature and water potential distributions. Both shoot hydraulic structure and leaf energy budget simulations were, as postulated, required to adequately scaling-up leaf to canopy gas-exchange rates. Notwithstanding, simulating the hydraulic structure of the shoot was found far more necessary to adequately performing this scaling task than simulating leaf energy balance. That is, the intra-canopy variability of leaf water potential was a better predictor of the reduction of whole plant gas-exchange rates under water deficit than the intra-canopy variability of leaf temperature. We conclude therefore that simulating the shoot hydraulic structure is a prerequisite if FSPM9s are to be used to assess gas-exchange rates of complex plant canopies as those of grapevines. Finally HydroShoot is available through the OpenAlea platform (https://github.com/openalea/hydroshoot) as a set of reusable modules.