Increased understanding of nutrient immobilization in soil organic matter is critical for predicting the carbon sink strength of forest ecosystems over the next 100 years.

The terrestrial biosphere is currently thought to be a significant sink for atmospheric carbon (C). However, the future course of this sink under rising [CO2] and temperature is uncertain. Some contrasting possibilities that have been suggested are: that the sink is currently increasing through CO2 fertilization of plant growth but will decline over the next few decades because of CO2 saturation and soil nutrient constraints; that the sink will continue to increase over the next century because rising temperature will stimulate the release of plant-available soil nitrogen (N) through increased soil decomposition; that, alternatively, the sink will not be sustained because the additional soil N released will be immobilized in the soil rather than taken up by plants; or that the sink will soon become negative because loss of soil C through temperature stimulation of soil respiration will override any CO2 or temperature stimulation of plant growth. Soil N immobilization is thus a key process; however, it remains poorly understood. In this paper we use a forest ecosystem model of plant-soil C and N dynamics to gauge the importance of this uncertainty for predictions of the future C sink of forests under rising [CO2] and temperature. We characterize soil N immobilization by the degree of variability of soil N:C ratios assumed in the model. We show that the modeled C sink of a stand of Norway spruce (Picea abies (L.) Karst.) in northern Sweden is highly sensitive to this assumption. Under increasing temperature, the model predicts a strong C sink when soil N:C is inflexible, but a greatly reduced C sink when soil N:C is allowed to vary. In complete contrast, increasing atmospheric [CO2] leads to a much stronger C sink when soil N:C is variable. When both temperature and [CO2] increase, the C sink strength is relatively insensitive to variability in soil N:C; significantly, however, with inflexible soil N:C the C sink is primarily a temperature response whereas with variable soil N:C, it is a combined temperature-CO2 response. Simulations with gradual increases of temperature and [CO2] indicate a sustained C sink over the next 100 years, in contrast to recent claims that the C sink will decline over the next few decades. Nevertheless, in using a relatively simple model, our primary aim is not to make precise predictions of the C sink over the next 100 years, but rather to highlight key areas of model uncertainty requiring further experimental clarification. Here we show that improved understanding of the processes underlying soil N immobilization is essential if we are to predict the future course of the forest carbon sink.

Foliage, fine-root, woody-tissue and stand respiration in Pinus radiata in relation to nitrogen status.

We measured respiration of 20-year-old Pinus radiata D. Don trees growing in control (C), irrigated (I), and irrigated + fertilized (IL) stands in the Biology of Forest Growth experimental plantation near Canberra, Australia. Respiration was measured on fully expanded foliage, live branches, boles, and fine and coarse roots to determine the relationship between CO(2) efflux, tissue temperature, and biomass or nitrogen (N) content of individual tissues. Efflux of CO(2) from foliage (dark respiration at night) and fine roots was linearly related to biomass and N content, but N was a better predictor of CO(2) efflux than biomass. Respiration (assumed to be maintenance) per unit N at 15 degrees C and a CO(2) concentration of 400 micro mol mol(-1) was 1.71 micro mol s(-1) mol(-1) N for foliage and 11.2 micro mol s(-1) mol(-1) N for fine roots. Efflux of CO(2) from stems, coarse roots and branches was linearly related to sapwood volume (stems) or total volume (branches + coarse roots) and growth, with rates for maintenance respiration at 15 degrees C ranging from 18 to 104 micro mol m(-3) s(-1). Among woody components, branches in the upper canopy and small diameter coarse roots had the highest respiration rates. Stem maintenance respiration per unit sapwood volume did not differ among treatments. Annual C flux was estimated by summing (1) dry matter production and respiration of aboveground components, (2) annual soil CO(2) efflux minus aboveground litterfall, and (3) the annual increment in coarse root biomass. Annual C flux was 24.4, 25.3 and 34.4 Mg ha(-1) year(-1) for the C, I and IL treatments, respectively. Total belowground C allocation, estimated as the sum of (2) and (3) above, was equal to the sum of root respiration and estimated root production in the IL treatment, whereas in the nutrient-limited C and I treatments, total belowground C allocation was greater than the sum of root respiration and estimated root production, suggesting higher fine root turnover or increased allocation to mycorrhizae and root exudation. Carbon use efficiency, the ratio of net primary production to assimilation, was similar among treatments for aboveground tissues (0.43-0.50). Therefore, the proportion of assimilation used for construction and maintenance respiration on an annual basis was also similar among treatments.

Modeling productivity and transpiration of Pinus radiata: climatic effects.

Climatic effects on annual net carbon gain, stem biomass and annual transpiration were simulated for Pinus radiata D. Don at Canberra and Mt. Gambier. Simulations were conducted with an existing process-based forest growth model (BIOMASS, Model 1) and with a modified version of the BIOMASS model (Model 2) in which response functions for carbon assimilation and leaf conductance were replaced with those derived from field gas exchange data collected at Mt. Gambier. Simulated carbon gain was compared with a published report stating that mean annual stem volume increment (MAI) at Mt. Gambier was 1.8 times greater than at Canberra and that the difference could be the result solely of differences in climate. Regional differences in climate resulted in a 20% greater simulated annual transpiration at Canberra than at Mt. Gambier but only small differences in simulated productivity, indicating that climatic differences did not account for the reported differences in productivity. With Model 1, simulated annual net carbon gain and annual increase in stem biomass were greater at Canberra than at Mt. Gambier, whereas Model 2 indicated a similar annual net carbon gain and annual stem biomass increase in both regions.

Aboveground net primary production decline with stand age: potential causes.

Aboveground net primary production (ANPP) commonly reaches a maximum in young forest stands and decreases by 0-76% as stands mature. However, the mechanism(s) responsible for the decline are not well understood. Current hypotheses for declining ANPP with stand age include: (1) an altered balance between photosynthetic and respiring tissues, (2) decreasing soil nutrient availability, and (3) increasing stomatal limitation leading to reduced photosynthetic rates. Recent empirical and modeling studies reveal that mechanisms (2) and (3) are largely responsible for age-related decline in ANPP for forests in cold environments. Increasing respiratory costs appear to be relatively unimportant in explaining declining productivity in ageing stands.

Long-Term Response of Nutrient-Limited Forests to CO"2 Enrichment; Equilibrium Behavior of Plant-Soil Models.

Established process-based models of forest biomass production in relation to atmospheric CO"2 concentration (McMurtrie 1991) and soil carbon/nutrient dynamics (Parton et al. 1987) are integrated to derive the @'Generic Decomposition and Yield@' model (G'DAY). The model is used to describe how photosynthesis and nutritional factors interact to determine the productivity of forests growing under nitrogen-limited conditions. A simulated instantaneous doubling of atmospheric CO"2 concentration leads to a growth response that is initially large (27% above productivity at current CO"2) but declines to <10% elevation within 5 yr. The decline occurs because increases in photosynthetic carbon gain at elevated CO"2 are not matched by increases in nutrient supply. Lower foliar N concentrations at elevated CO"2 have two countervailing effects on forest production: decreased rates of N cycling between vegetation and soils (with negative consequences for productivity), and reduced rates of N loss through gaseous emission, fire, and leaching. Theoretical analysis reveals that there is an enduring response to CO"2 enrichment, but that the magnitude of the long-term equilibrium response is extremely sensitive to the assumed rate of gaseous emission resulting from mineralization of nitrogen. Theory developed to analyze G'DAY is applicable to other published production-decomposition models describing the partitioning of soil carbon among compartments with widely differing decay-time constants.