Carbon fluxes in a mature deciduous forest under elevated CO₂

This PhD thesis addressed several major aspects of the carbon (C) cycle in a c. 100-year-old, mixed deciduous forest under elevated CO₂ with an emphasis on below-ground processes. The aim was to assess the responses of tree fine roots and soil respiration to canopy CO₂ enrichment (? 550 ppm) in this tallest forest studied to date. Furthermore, leaf gas-exchange of the five study species was examined to ascertain the long-term response of photosynthetic carbon uptake to elevated atmospheric CO₂. Investigations at the Swiss Canopy Crane (SCC) experimental site were guided by the following key questions: (1) Does below-ground C allocation to fine root production increase in response to CO₂ enrichment in order to acquire more nutrients to match the enhanced C supply in the forest canopy? (2) Is below-ground metabolism enhanced and therefore forest soil respiration stimulated by canopy CO₂ enrichment? (3) Is leaf-level photosynthesis persistently stimulated by elevated CO₂ in this stand or had these mature broad-leaved trees reduced their carbon up- take by photosynthetic down-regulation under long-term CO₂ enrichment?
Findings from earlier studies at the SCC site, including 13C isotope tracing, all point towards an in- creased flux of C through CO₂-enriched trees to the soil but neither fine root biomass nor soil respiration were stimulated by elevated CO₂. Surprisingly, fine root biomass in bulk soil and ingrowth cores showed strong reductions by ? 30% in year five and six but were unaffected in the following seventh year of CO₂ enrichment. Given the absence of a positive biomass response of fine roots, we assumed that the extra C assimilated in the CO₂-enriched forest canopy was largely respired back to the atmosphere via increases in fine root and rhizosphere respiration and the metabolization of increased root derived exudates by soil microbes. Indeed, 52% higher soil air CO₂ concentration during the growing season and 14% greater soil microbial biomass both in- dicated enhanced below-ground metabolism in soil under CO₂-enriched trees. However, this did not translate into a persistent stimulation of soil respiration. At times of high or continuous precipitation soil water savings under CO₂-exposed trees (resulting from reduced sapflow) led to excessive soil moisture (> 45 vol.-%) impeding soil gas-exchange and thus soil respiration. Depending on the interplay between soil temperature and the consistently high soil water content in this stand, instantaneous rates of soil respiration were periodically reduced or increased under elevated CO₂ but on a diel scale and integrated over the growing season soil CO₂ emissions were similar under CO₂-enriched and control trees. Soil respiration could therefore not explain the fate of the extra C. The lacking sink capacity for additional assimilates led us to assume downward adjustment of photosynthetic capacity in CO₂-enriched trees thereby reducing carbon uptake in the forest canopy. Photosynthetic acclimation cannot completely eliminate the CO₂-driven stimulation in carbon uptake, but a reduction could hamper the detection of a CO₂ effect considering the low statistical power inevitably involved with such large-scale experiments. However, after eight years of CO₂ enrichment we found sustained stimulation in leaf photosynthesis (42-49%) indicating a lack of closure in the carbon budget for this stand under elevated atmospheric CO₂.