Using stable isotopes to investigate interactions between the forest carbon and nitrogen cycles

Abstract

Nitrogen (N) fertilization due to atmospheric deposition (NDEP ) may explain some of the net carbon (C) sink (0.6-0.7 Pg y-1) in temperate forests, but estimates of the additional C uptake due to atmospheric N additions (∆C/∆N) can vary by over an order of magnitude (5 to 200 ∆C/∆N). High estimates from several recent studies [e.g. Magnani (2007), Nature 447 848-850], deriving ∆C/∆N from regional correlations between NDEP and measures of C uptake (such as eddy covariance -derived net ecosystem production, or forest inventory data) contradict estimates from other studies, particularly those involving 15N tracer applications added as fertilizer to the forest floor. A strong ∆C/∆N effect requires nitrogen to be efficiently acquired by trees and allocated to high C:N, long-lived woody tissues, but these isotope experiments typically report relatively little (~ 20 %) of 15N added is found above-ground, with less than 5 % of the total 15N applied found in wood. Consequently the high correlation-derived ∆C/∆N estimates are often attributed to co-variation with other factors across the range of sites investigated. However, 15N-fertilization treatments often impose considerably higher total N loads than ambient NDEP , while almost all exclusively only apply mineral 15N treatments to the soil, often in a limited number of treatment events over relatively short periods of time. Excessive N deposition loads can induce negative physiological effects and limit the resulting ∆C/∆N observed, and applying treatments to the soil ignores canopy nitrogen uptake, which has been demonstrated in numerous studies. As canopies can directly take up nitrogen, the chronic, (relatively) low levels of ambient NDEP inputs from pollution may be acquired without some of the effects of heavy N loads, with trees obtaining this N before it reaches the soil, allowing canopies to substitute for, or supplement, edaphic N nutrition. The strength of this effect depends on how much N uptake can occur across the canopy under field conditions, and if this extra N supplies growth in woody tissues such as the stem, as well as the canopy. Similarly, such mineral fertilizer isotope trace experiments are also unable to trace N in the decomposing litter and humus layers of the soil, which even under heavy NDEP loading contribute most of the N utilised for forest growth. Recent literature suggests that some organic (early decomposition) forms of N may be taken up by roots. If this litter N is not retained or distributed in the same way as mineral fertilizers, its contribution to plant nutrition and ∆C/∆N may need to be reassessed under nitrogen deposition.

We tested some of these assumptions in the nursery and the field. In order to facilitate litter 15N tracing, we conducted an experiment injecting large trees with 15N-NH4NO3 to create 15N-labelled litter, tracing the applied isotope into a full harvest of the canopy. Such labelled litter substitute was used to replace the litter layer in a Sitka Spruce plantation (Picea sitchensis L. (Bong.)), where the fate of this 15N from litter decomposition in the soil system was compared against the fate of 15N in deposition. Similarly, in potted Sitka Spruce saplings, we used combination treatments of 15N-labelled litter, soil-targeted 15N-deposition, and canopy targeted 15N-deposition, investigating 15N return in different age classes of above and below ground biomass.

We found that i) 15N recovery in canopies (needles and branches) in our injected trees was almost all of the injected 15N five months after injection, ii) canopy application of NDEP led to 60 % 15N return in above-ground parts of saplings compared to 21 % in soil applications and iii) a litter-derived 15N source was retained 55 % more in topsoil, and 36 % more in roots, than a similar deposition 15N source applied as mineral fertilizer.

We discuss the implications of such findings in the context of 15N return in different plant organs and ecosystem pools, seasonal variation in N content, and overall inferences of a forest ∆C/∆N effect. Our results suggest that the total ∆C/∆N effect driven by a high N sequestration from canopy uptake in wood is ~ 114:1, more than double that of 15N tracer experiments but not as high as upper estimates from correlative studies, and that litter-derived organic N is better retained in trees and soils in excess of similar amounts of mineral 15N from deposition. Existing forest 15N-fertilization experiments could under-estimate the overall ∆C/∆N effect of atmospheric N deposition.