Increases in the flux of carbon belowground stimulate nitrogen uptake and sustain the long-term enhancement of forest productivity under elevated CO2.

The earth's future climate state is highly dependent upon changes in terrestrial C storage in response to rising concentrations of atmospheric CO2. Here we show that consistently enhanced rates of net primary production (NPP) are sustained by a C-cascade through the root-microbe-soil system; increases in the flux of C belowground under elevated CO2 stimulated microbial activity, accelerated the rate of soil organic matter decomposition and stimulated tree uptake of N bound to this SOM. This process set into motion a positive feedback maintaining greater C gain under elevated CO2 as a result of increases in canopy N content and higher photosynthetic N-use efficiency. The ecosystem-level consequence of the enhanced requirement for N and the exchange of plant C for N belowground is the dominance of C storage in tree biomass but the preclusion of a large C sink in the soil.

Increased belowground biomass and soil CO2 fluxes after a decade of carbon dioxide enrichment in a warm-temperate forest.

Atmospheric CO2 concentrations have risen 40% since the start of the industrial revolution. Beginning in 1996, the Duke Free-Air CO2 Enrichment experiment has exposed plots in a loblolly pine forest to an additional 200 microL/L CO2 compared to trees growing in ambient CO2. This paper presents new belowground data and a synthesis of results through 2008, including root biomass and nutrient concentrations, soil respiration rates, soil pore-space CO2 concentrations, and soil-solution chemistry to 2 m depth. On average in elevated CO2, fine-root biomass in the top 15 cm of soil increased by 24%, or 59 g/m2 (26 g/m2 C). Coarse-root biomass sampled in 2008 was twice as great in elevated CO2 and suggests a storage of approximately 20 g C x m(-2) x yr(-1). Root C and N concentrations were unchanged, suggesting greater belowground plant demand for N in high CO2. Soil respiration was significantly higher by 23% on average as assessed by instantaneous infrared gas analysis and 24-h integrated estimates. N fertilization decreased soil respiration and fine-root biomass by approximately 10-20% in both ambient and elevated CO2. In recent years, increases in root biomass and soil respiration grew stronger, averaging approximately 30% at high CO2. Peak changes for root biomass, soil respiration, and other variables typically occurred in midsummer and diminished in winter. Soil CO2 concentrations between 15 and 100 cm depths increased 36-60% in elevated CO2. Differences from 30 cm depth and below were still increasing after 10 years' exposure to elevated CO2, with soil CO2 concentrations >10000 microL/L higher at 70- and 100-cm depths, potentially influencing soil acidity and rates of weathering. Soil solution Ca2+ and total base cation concentrations were 140% and 176% greater, respectively, in elevated CO2 at 200 cm depth. Similar increases were observed for soil-solution conductivity and alkalinity at 200 cm in elevated CO2. Overall, the effect of elevated CO2 belowground shows no sign of diminishing after more than a decade of CO2 enrichment.

The effect of carbon dioxide enrichment on apparent stem respiration from Pinus taeda L. is confounded by high levels of soil carbon dioxide.

Respiration supports growth and maintenance processes and returns a substantial portion of the CO(2) fixed by photosynthesis to the atmosphere each year. Investigating stem respiration using CO(2) flux measurements is complicated by uncertainty surrounding the source of CO(2) diffusing from tree stems. Over 2 years we measured the stem efflux from 24 trees exposed to ambient or elevated CO(2). The rate of stem CO(2) efflux increased with annual tree diameter increment and the estimated uptake of dissolved CO(2) from the soil. To determine the source of CO(2) diffusing from tree stems, we used the fumigation gas at the Duke Forest Atmosphere Carbon Transfer and Storage-1 elevated-CO(2) experiment as a (13)C tracer and measured the presence of soil CO(2) in stem efflux on a subset of these trees. The isotopic composition of soil CO(2) explained a considerable portion of the variation in the composition of CO(2) in stem efflux. We also found that direct measurements of the isotopic composition of phloem-respired CO(2), unlike the CO(2) found in stem efflux, was less variable and distinct from the isotopic composition of soil CO(2). Tree growth rates and soil CO(2) concentrations found at the site together explained 56% of the variance in stem CO(2) efflux among trees. These results suggest that the uptake of CO(2) dissolved in soil water and transported through the vascular system can potentially confound efforts to interpret stem efflux measurements in trees exposed to elevated CO(2) and that previous studies may have overestimated the effects of elevated CO(2) on autotrophic respiration in tree stems.

Progressive nitrogen limitation of ecosystem processes under elevated CO2 in a warm-temperate forest.

A hypothesis for progressive nitrogen limitation (PNL) proposes that net primary production (NPP) will decline through time in ecosystems subjected to a step-function increase in atmospheric CO2. The primary mechanism driving this response is a rapid rate of N immobilization by plants and microbes under elevated CO2 that depletes soils of N, causing slower rates of N mineralization. Under this hypothesis, there is little long-term stimulation of NPP by elevated CO2 in the absence of exogenous inputs of N. We tested this hypothesis using data on the pools and fluxes of C and N in tree biomass, microbes, and soils from 1997 through 2002 collected at the Duke Forest free-air CO2 enrichment (FACE) experiment. Elevated CO2 stimulated NPP by 18-24% during the first six years of this experiment. Consistent with the hypothesis for PNL, significantly more N was immobilized in tree biomass and in the O horizon under elevated CO2. In contrast to the PNL hypothesis, microbial-N immobilization did not increase under elevated CO2, and although the rate of net N mineralization declined through time, the decline was not significantly more rapid under elevated CO2. Ecosystem C-to-N ratios widened more rapidly under elevated CO2 than ambient CO2 indicating a more rapid rate of C fixation per unit of N, a processes that could delay PNL in this ecosystem. Mass balance calculations demonstrated a large accrual of ecosystem N capital. Is PNL occurring in this ecosystem and will NPP decline to levels under ambient CO2? The answer depends on the relative strength of tree biomass and O-horizon N immobilization vs. widening C-to-N ratios and ecosystem-N accrual as processes that drive and delay PNL, respectively. Only direct observations through time will definitively answer this question.