Abstract
Human actions are causing declines in plant biodiversity, increases in atmospheric CO2 concentrations and increases in nitrogen deposition; however, the interactive effects of these factors on ecosystem processes are unknown1,2. Reduced biodiversity has raised numerous concerns, including the possibility that ecosystem functioning may be affected negatively1,2,3,4, which might be particularly important in the face of other global changes5,6. Here we present results of a grassland field experiment in Minnesota, USA, that tests the hypothesis that plant diversity and composition influence the enhancement of biomass and carbon acquisition in ecosystems subjected to elevated atmospheric CO2 concentrations and nitrogen deposition. The study experimentally controlled plant diversity (1, 4, 9 or 16 species), soil nitrogen (unamended versus deposition of 4 g of nitrogen per m2 per yr) and atmospheric CO2 concentrations using free-air CO2 enrichment (ambient, 368 µmol mol-1, versus elevated, 560 µmol mol-1). We found that the enhanced biomass accumulation in response to elevated levels of CO2 or nitrogen, or their combination, is less in species-poor than in species-rich assemblages.
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Acknowledgements
We thank the US Department of Energy for financial support of this project, with additional support from the US National Science Foundation, Long-Term Ecological Research Program. (Please note - This acknowledgement was omitted in error from the printed journal. A correction will be noted in a subsequent issue.)
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CO2 enrichment.
The three ambient and three elevated CO2 treatment rings at the BioCON facility have identical free-air delivery systems (i.e., plenums, vent pipes and blowers) operating simultaneously, so the only difference among the treatments involves the additional CO2 released from the vent pipes in the elevated treatment rings. The target concentration during the CO2 enrichment periods at plant canopy height in the center of each ring was 550 µmol mol-1. During all daylight hours during the growing season, the 5 minute average was within 5% of the target concentration for 92-93% of the time (including times when the system was inoperable due to technical difficulties). The mean CO2 concentrations during enrichment over the two years varied by <1 µmol mol-1 among rings.
To assess spatial variation in CO2 concentrations within rings, a multiport sampler sequentially sampled CO2 concentrations at 32 points in a ring. Samples were averaged over 30 seconds intervals, and measurements were recorded from mid-June through mid-October of 1998 and 1999. Values for CO2 concentrations at the centers of the 61 individual plots was calculated using an inverse distance method and no plot center was farther than 2 meters from a sampling port. The CO2 concentrations were slightly higher on average near the edge than the center of the ring. Assuming a similar spatial distribution of CO2 concentrations in the other elevated rings, we estimated the mean concentrations for all treatment combinations. Given the large number of replicates for each treatment and their random location within and among rings, the mean concentrations varied little among species richness levels (560, 561, 561, and 559 µmol mol-1 for 1, 4, 9 and 16 species, respectively; mean standard deviation was 10 µmol mol-1) or among N levels (560 and 561 µmol mol-1 for ambient and enriched N treatment plots, respectively; mean standard deviation was 10 µmol mol-1).
Biomass sampling and biogeochemistry measurements.
At each harvest in every plot, aboveground biomass was harvested by clipping a 10 x 100 cm strip just above the soil surface. All biomass was collected, sorted to live material and senesced litter, dried and weighed. Total belowground biomass (fine roots, coarse roots and crowns) was sampled at 0-20 cm depth using three 5 cm dia. cores in the area used for the aboveground biomass sampling. Roots were thoroughly washed, sorted into fine (<1 mm diameter) and coarse classes and crowns, dried and weighed. Any given area was sampled only once during the four harvests. All biomass was ground and analyzed separately for aboveground and belowground components for C and N concentrations using a CHN analyzer (Carlo-Erba Strumatzione, Milan, Italy). Soil solution N concentrations were also assessed at the 0-20 cm depth at all four harvests following extraction using 0.01 mol KCl. Soil net N mineralization was measured in every plot using one-month in situ incubations at 0-20 cm depth during midsummer of each year.
Data.
Figure 1:
Total aboveground and belowground (0-20 cm depth) biomass (∓ one standard error) for plots planted with either 1, 4, 9 or 16 species, grown at four combinations of ambient (368 µmol mol-1) and elevated (560 µmol mol-1) concentrations of CO2, and ambient N and N addition (4 g N m-2 yr-1) treatments. Data are shown for each of four harvests (June and August in both 1998 and 1999).
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Reich, P., Knops, J., Tilman, D. et al. Plant diversity enhances ecosystem responses to elevated CO2 and nitrogen deposition. Nature 410, 809–810 (2001). https://doi.org/10.1038/35071062
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DOI: https://doi.org/10.1038/35071062
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