High levels of atmospheric nitrogen (N) deposition in Europe and North America were maintained throughout the 1990s, and global N deposition is expected to increase by a factor of 2.5 over the next century. Available soil N limits primary production in many terrestrial ecosystems, and some computer simulation models have predicted that increasing atmospheric N deposition may result in greater terrestrial carbon (C) storage in woody biomass. However, empirical evidence demonstrating widespread increases in woody biomass C storage due to atmospheric N deposition is uncommon. Increased C storage in soil organic matter due to chronic N inputs has rarely been reported and is often not considered in computer simulation models of N deposition effects. Since 1994, we have experimentally simulated chronic N deposition by adding 3 g N m À2 yr À1 to four different northern hardwood forests, which span a 500 km geographic gradient in Michigan. Each year we measured tree growth. In 2004, we also examined soil C content to a depth of 70 cm. When we compared the control treatment with the NO 3 À deposition treatment after a decade of experimentation, ecosystem C storage had significantly increased in both woody biomass (500 g C m À2 ) and surface soil (0-10 cm) organic matter (690 g C m À2 ). The increase in surface soil C storage was apparently driven by altered rates of organic matter decomposition, rather than an increase in detrital inputs to soil. Our results, for study locations stretching across hundreds of kilometers, support the hypothesis that chronic N deposition may increase C storage in northern forests, potentially contributing to a sink for anthropogenic CO 2 in the northern Hemisphere.

Elevated CO 2 (eCO 2) experiments provide critical information to quantify the effects of rising CO 2 on vegetation 1,2,3,4,5,6. Many eCO 2 experiments suggest that nutrient limitations modulate the local magnitude of the eCO 2 effect on plant biomass 1,3,5 , but the global extent of these limitations has not been empirically quantified, complicating projections of the capacity of plants to take up CO 2 7,8. Here, we present a data-driven global quantification of the eCO 2 effect on biomass based on 138 eCO 2 experiments. The strength of CO 2 fertilization is primarily driven by nitrogen (N) in ~65% of global vegetation and by phosphorus (P) in ~25% of global vegetation, with Nor Plimitation modulated by mycorrhizal association. Our approach suggests that CO 2 levels expected by 2100 can potentially enhance plant biomass by 12 ± 3% above current values, equivalent to 59 ± 13 PgC. The global-scale response to eCO 2 we derive from experiments is similar to past changes in greenness 9 and biomass 10 with rising CO 2 , suggesting that CO 2 will continue to stimulate plant biomass in the future despite the constraining effect of soil nutrients. Our research reconciles conflicting evidence on CO 2 fertilization across scales and provides an empirical estimate of the biomass sensitivity to eCO 2 that may help to constrain climate projections.

Presently, there is uncertainty regarding the degree to which anthropogenic N deposition will foster C storage in the N-limited forests of the Northern Hemisphere, ecosystems which are globally important sinks for anthropogenic CO2. We constructed organic matter and N budgets for replicate northern hardwood stands (n = 4) that have received ambient (0.7-1.2 g N x m(-2) x yr(-1) and experimental NO3- deposition (ambient plus 3 g NO3(-)-N x m(-2) x yr(-1)) for a decade; we also traced the flow of a 15NO3- pulse over a six-year period. Experimental NO3- deposition had no effect on organic matter or N stored in the standing forest overstory, but it did significantly increase the N concentration (+19%) and N content (+24%) of canopy leaves. In contrast, a decade of experimental NO3- deposition significantly increased amounts of organic matter (+12%) and N (+9%) in forest floor and mineral soil, despite no increase in detritus production. A greater forest floor (Oe/a) mass under experimental NO3- deposition resulted from slower decomposition, which is consistent with previously reported declines in lignolytic activity by microbial communities exposed to experimental NO3- deposition. Tracing 15NO3- revealed that N accumulated in soil organic matter by first flowing through soil microorganisms and plants, and that the shedding of 15N-labeled leaf litter enriched soil organic matter over a six-year duration. Our results demonstrate that atmospheric NO3- deposition exerts a direct and negative effect on microbial activity in this forest ecosystem, slowing the decomposition of aboveground litter and leading to the accumulation of forest floor and soil organic matter. To the best of our knowledge, this mechanism is not represented in the majority of simulation models predicting the influence of anthropogenic N deposition on ecosystem C storage in northern forests.

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