The dynamic of the annual carbon allocation to wood in European tree species is consistent with a combined source-sink limitation of growth: implications for modelling

The extent to which wood growth is limited by carbon (C) supply (i.e. source control) or by cambial activity (i.e. sink control) will strongly determine the responses of trees to global changes. Nevertheless, the physiological processes that are responsible for limiting forest growth are still a matter of debate. The aim of this study was to evaluate the key determinants of the annual C allocation to wood along large soil and climate regional gradients over France. The study was conducted for five tree species representative of the main European forest biomes (Fagus sylvatica, Quercus petraea, Quercus ilex, Quercus robur and Picea abies). The drivers of stand biomass growth were assessed on both inter-site and inter-annual scales. Our data set comprised field measurements performed at 49 sites (931 site-years) that included biometric measurements and a variety of stand characteristics (e.g. soil water holding capacity, leaf area index). It was complemented with process-based simulations when possible explanatory variables could not be directly measured (e.g. annual and seasonal tree C balance, bioclimatic water stress indices). Specifically, the relative influences of tree C balance (source control), direct environmental control (water and temperature controls of sink activity) and allocation adjustments related to age, past climate conditions, competition intensity and soil nutrient availability on growth were quantified. The inter-site variability in the stand C allocation to wood was predominantly driven by age-related decline. The direct effects of temperature and water stress on sink activity (i.e. effects independent from their effects on the C supply) exerted a strong influence on the annual stand wood growth in all of the species considered, including deciduous temperate species. The lagged effect of the past environmental conditions (e.g. the previous year's water stress and low C uptake) significantly affected the annual C allocation to wood. The C supply appeared to strongly limit growth only in temperate deciduous species. We provide an evaluation of the spatio-temporal dynamics of the annual C allocation to wood in French forests. Our study supports the premise that the growth of European tree species is subject to complex control processes that include both source and sink limitations. The relative influences of the growth drivers strongly vary with time and across spatial ecological gradients. We suggest a straightforward modelling framework with which to implement these combined forest growth limitations into terrestrial biosphere models