Productivity and carbon allocation in European forests: a process-based modelling approach
The processes that underlie forest productivity and C allocation dynamics in trees are still poorly understood. Forest growth has for long been thought to be C limited, through a hypothesized causal link between C supply and growth (source control). This C-centric paradigm underlies most of the C allocation rules formalized in process-based models (PBMs). However, the source limitation of growth has been questioned by several authors, arguing that meristem activities are more sensitive than C assimilation to environmental stresses (e.g., water deficit and low temperatures). Moreover, the effect of management, which strongly affects forest functioning and wood growth, is not accounted for in most of the PBMs used to project the future terrestrial C sink. Our main objective in this thesis was to move forward into our understanding of the constraints that affect - or will affect - the wood productivity in European forests, from present to the end of the 21st century. We addressed this objective through the improvement of the representation of the forest productivity and C allocation in the CASTANEA PBM, building on a detailed analysis of the key drivers of annual wood productivity in French forests over the last 30 years (the species studied are Fagus sylvatica, Quercus ilex, Quercus petraea, Quercus robur and Picea abies). Our results supported the premise that the annual wood growth of the studied species is under a complex control including both source and sink limitations. The inter-site variability in the fraction of C allocatedto stand wood growth was predominantly driven by an age-related decline. At the tree level, we showed that annual wood growth was well predicted by the individual size. The size-asymmetry of growth, i.e., the advantage of big trees in the competition for resources, increased consistently with the whole stand productivity at both inter-site and inter-annual scales. On the basis of our findings, we developed a new C allocation scheme in the CASTANEA PBM, which integrate a combined source-sink limitation of wood growth. The new calibrated model captured both the inter-annual and inter-site changes in stand wood growth that was observed across national environmental gradients. The model was also successfully evaluated against a meta-analysis of carbohydrate reserve pools in trees and satellite-derived leaf area index estimates. Our results indicated that the representation of the environmental control of sink activity does not affect the qualitative predictions of the future of the European forest productivity previously obtained from source-driven PBMs. However, the current, source-driven generation of PBMs probably underestimates the spatial heterogeneity of the effects of climate change on forest growth that arise from sink limitations. Further, we successfully used our findings regarding the dependences of annual wood growth at tree level (i.e., empirical rules of tree growth competition) to calibrate a module for the simulation of the individual growth of trees in the CASTANEA model. The coupled model was used to assess the potential effects of management on forest functioning and wood growth across France. We identified the areas where management efforts may be concentrated in order to mitigate near-future drought impact on national forest productivity. Around a quarter of the French temperate oak and beech forests are currently in zones of high vulnerability, where management could thus mitigate the influence of climate change on forest yield.