Progressive N limitation in forests: review and implications for long-term responses to elevated CO2.

Field studies have shown that elevated CO2 can cause increased forest growth over the short term (<6 years) even in the face of N limitation. This is facilitated to some degree by greater biomass production per unit N uptake (lower tissue N concentrations), but more often than not, N uptake is increased with elevated CO2 as well. Some studies also show that N sequestration in the forest floor is increased with elevated CO2. These findings raise the questions of where the "extra" N comes from and how long such growth increases can continue without being truncated by progressive N limitation (PNL). This paper reviews some of the early nutrient cycling literature that describes PNL during forest stand development and attempts to use this information, along with recent developments in soil N research, to put the issue of PNL with elevated CO2 into perspective. Some of the early studies indicated that trees can effectively "mine" N from soils over the long term, and more recent developments in soil N cycling research suggest mechanisms by which this might have occurred. However, both the early nutrient cycling literature and more recent simulation modeling suggest that PNL will at some point truncate the observed increases in growth and nutrient uptake with elevated CO2, unless external inputs of N are increased by either N fixation or atmospheric deposition.