Biomass allocation and canopy development in spruce model ecosystems under elevated CO2 and increased N deposition.

Ecosystem-level experiments on the effects of atmospheric CO2 enrichment and N deposition on forest trees are urgently needed. Here we present data for nine model ecosystems of spruce (Picea abies) on natural nutrient-poor montane forest soil (0.7 m2 of ground and 350 kg weight). Each system was composed of six 7-year-old (at harvest) trees each representing a different genotype, and a herbaceous understory layer (three species). The model ecosystems were exposed to three different CO2 concentrations (280, 420, 560 μl l-1) and three different rates of wet N deposition (0, 30, 90 kg ha-1 year-1) in a simulated annual course of Swiss montane climate for 3 years. The total ecosystem biomass was not affected by CO2 concentration, but increased with increasing N deposition. However, biomass allocation to roots increased with increasing CO2 leading to significantly lower leaf mass ratios (LMRs) and leaf area ratios (LARs) in trees grown at elevated CO2. In contrast to CO2 enrichment, N deposition increased biomass allocation to the aboveground plant parts, and thus LMR and LAR were higher with increasing N deposition. We observed no CO2 ×  N interactions on growth, biomass production, or allocation, and there were also no genotype × treatment interactions. The final leaf area index (LAI) of the spruce canopies was 19% smaller at 420 and 27% smaller at 560 than that measured at 280 μl CO2 l-1, but was not significantly altered by increasing N deposition. Lower LAIs at elevated CO2 largely resulted from shorter branches (less needles per individual tree) and partially from increased needle litterfall. Independently of N deposition, total aboveground N content in the spruce communities declined with increasing CO2 (-18% at 420 and -31% at 560 compared to 280 μl CO2 l-1). N deposition had the opposite effect on total above ground N content (+18% at 30 and +52% at 90 compared to 0 kg N ha-1 year-1). Our results suggest that under competitive conditions on natural forest soil, atmospheric CO2 enrichment may not lead to higher ecosystem biomass production, but N deposition is likely to do so. The reduction in LAI under elevated CO2 suggests allometric down-regulation of photosynthetic carbon uptake at the canopy level. The strong decline in the tree nitrogen mass per unit ground area in response to elevated CO2 may indicate CO2-induced reductions of soil N availability.