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The Jasper Ridge CO2experiment in northern California3 exposed two natural annual grassland ecosystems (sandstone and serpentine) to increased atmospheric CO2for six growing seasons by using cylindrical, open-top chambers (1 m tall, 0.33 m2, n = 10). In both grasslands, a higher proportion of soil was found in aggregates 1-2 mm across in elevated CO2, and the proportion of aggregates of 0.25-1 mm was significantly increased in the sandstone grassland (Table 1). The water stability of both size classes followed a pattern similar to the mass of aggregates. This suggested that the higher mass of aggregates could be explained by an increase in the water stability of aggregates (Table 1).

Table 1 Effects of increased CO2on aggregates and glomalin in two annual grasslands

Although soil aggregation is a complex hierarchical process4, the soil concentration of the glycoprotein glomalin5 is tightly correlated with aggregate stability across many soils6. Glomalin is produced mainly by hyphae of arbuscular mycorrhizal fungi5, which form symbiotic associations with plant roots. The length of the hyphae in these fungi increases with elevated CO2in the sandstone grassland, but not in the serpentine grassland, with root biomass and length showing the opposite pattern7. Total glomalin and immunoreactive glomalin concentrations in soil increased in both grasslands with elevated CO2 (Table 1). Glomalin concentration in aggregates (from a separate extraction) increased under elevated CO2for aggregates of 0.25-1 mm in both communities, but this was not the case for those of 1-2 mm (Table 1). The water stability of that fraction may be under different control.

The Sky Oaks CO2study in southern California used 12 greenhouses (2×2×2 m) with controlled CO2, ambient lighting and controlled temperature at six CO2concentrations from a pre-industrial level of 250 μl l−1 to 750 μl l−1 at intervals of 100 μl l−1 (n = 2). The chambers were built around Adenostoma fasciculatum (chamise) shrubs in chaparral vegetation recovering from an experimental burn. Soil samples were taken after three years of treatment and analysed for soil aggregation and glomalin concentration to see whether the patterns in the grasslands also existed in a different vegetation type. The proportion of soil mass in aggregates of 0.25-1 mm showed a linear increase (linear regression, P = 0.03, r 2 = 0.74) along the CO2gradient, but the 1-2 mm aggregate mass did not (P = 0.68, r 2 = 0.04). Glomalin concentrations followed a pattern similar to that of the small aggregate size class (P = 0.03, r 2 = 0.71).

The carbon sink represented by glomalin over the experimental period for Jasper Ridge was 8.29 g C m−2 in the serpentine and 4.25 g C m−2 in the sandstone grassland. These are very small amounts compared with the large organic carbon stocks in these soils, and are on the order of 5% of the total calculated litter and soil accumulation under elevated CO2on an annual basis8. Glomalin therefore seems to be more important in carbon sequestration by virtue of its function in soil aggregation (which has been linked with carbon stabilization) than by acting as a carbon sink itself.

Our results indicate that changes in soil structure in response to CO2enrichment should be incorporated into global research because soil structure has a strong effect on soil processes and organisms. On a global scale, the extent of soil degradation and erosion is severe9 and is accelerated by changes in many global factors, including climate and land use10. Our finding that an increase in soil aggregation could be brought about by atmospheric change may have implications for studies of soil stabilization in ecosystems.