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The objective was to examine the effects of optimal leaf nitrogen levels > 2.0% and suboptimal levels ˂ 2.0%, nitrogen nutrition on net photosynthetic rate, stem diameter increment, height growth increment and acorn mass of pedunculate oak during 2010 in the absence of drought stress and during 2011 under the impact of moderate drought stress. According to the results, moderate drought stress significantly reduced net photosynthetic rate, stem diameter increment and height growth increment, while acorn mass was not affected. Suboptimal nitrogen nutrition significantly reduced the net photosynthetic rate and stem diameter increment only in the wet year, acorn mass in both wet and dry years, while height growth increment was not significantly reduced by suboptimal nitrogen nutrition in either year. The results indicate that optimal nitrogen levels can stimulate photosynthetic rate and stem diameter increment of pedunculate oak only in the absence of moderate drought stress. Moreover, the results show that moderate drought stress is a more dominant stressor for photosynthesis and growth of pedunculate oak than suboptimal nitrogen nutrition, while for acorn development, it is the more dominant stressor.

Climate projections predict higher precipitation variability with more frequent dry extremes(1). CO2 assimilation of forests decreases during drought, either by stomatal closure(2) or by direct environmental control of sink tissue activities(3). Ultimately, drought effects on forests depend on the ability of forests to recover, but the mechanisms controlling ecosystem resilience are uncertain(4). Here, we have investigated the effects of drought and drought release on the carbon balances in beech trees by combining CO2 flux measurements, metabolomics and (13)CO2 pulse labelling. During drought, net photosynthesis (AN), soil respiration (RS) and the allocation of recent assimilates below ground were reduced. Carbohydrates accumulated in metabolically resting roots but not in leaves, indicating sink control of the tree carbon balance. After drought release, RS recovered faster than AN and CO2 fluxes exceeded those in continuously watered trees for months. This stimulation was related to greater assimilate allocation to and metabolization in the rhizosphere. These findings show that trees prioritize the investment of assimilates below ground, probably to regain root functions after drought. We propose that root restoration plays a key role in ecosystem resilience to drought, in that the increased sink activity controls the recovery of carbon balances.