Winter climate change implications for decomposition in northeastern forests: comparisons of sugar maple litter with herbivore fecal inputs

Forests in northeastern North America are influenced by varying climatic and biotic factors; however, there is concern that rapid changes in these factors may lead to important changes in ecosystem processes such as decomposition. Climate change (especially warming) is predicted to increase rates of decomposition in northern latitudes. Warming in winter may result in complex effects including decreased levels of snow cover and an increased incidence of soil freezing that will effect decomposition. Along with these changes in climate, moose densities have also been increasing in this region, likely affecting nutrient dynamics. We measured decomposition and N release from ¹⁵N‐labeled sugar maple leaf litter and moose feces over 20 months in reference and snow removal treatment (to induce soil freezing) plots in two separate experiments at the Hubbard Brook Experimental Forest in New Hampshire, USA. Snow removal/soil freezing decreased decomposition of maple litter, but stimulated N transfer to soil and microbial biomass. Feces decomposed more rapidly than maple litter, and feces N moved into the mineral soil more than N derived from litter, likely due to the lower C : N ratio of feces. Feces decomposition was not affected by the snow removal treatment. Total microbial biomass (measured as microbial N and C) was not significantly affected by the treatments in either the litter or feces plots. These results suggest that increases in soil freezing and/or large herbivore populations, increase the transfer rate of N from plant detritus or digested plants into the mineral soil. Such changes suggest that altering the spatial and temporal patterns of soil freezing and moose density have important implications for ecosystem N cycling.     

Influence of experimental snow removal on root and canopy physiology of sugar maple trees in a northern hardwood forest

Due to projected increases in winter air temperatures in the northeastern USA over the next 100 years, the snowpack is expected to decrease in depth and duration, thereby increasing soil exposure to freezing air temperatures. To evaluate the potential physiological responses of sugar maple (Acer saccharum Marsh.) to a reduced snowpack, we measured root injury, foliar cation and carbohydrate concentrations, woody shoot carbohydrate levels, and terminal woody shoot lengths of trees in a snow manipulation experiment in New Hampshire, USA. Snow was removed from treatment plots for the first 6 weeks of winter for two consecutive years, resulting in lower soil temperatures to a depth of 50 cm for both winters compared to reference plots with an undisturbed snowpack. Visibly uninjured roots from trees in the snow removal plots had significantly higher (but sub-lethal) levels of relative electrolyte leakage than trees in the reference plots. Foliar calcium: aluminum (Al) molar ratios were significantly lower, and Al concentrations were significantly higher, in trees from snow removal plots than trees from reference plots. Snow removal also reduced terminal shoot growth and increased foliar starch concentrations. Our results are consistent with previous research implicating soil freezing as a cause of soil acidification that leads to soil cation imbalances, but are the first to show that this translates into altered foliar cation pools, and changes in soluble and structural carbon pools in trees. Increased soil freezing due to a reduced snowpack could exacerbate soil cation imbalances already caused by acidic deposition, and have widespread implications for forest health in the northeastern USA.     

Seasonal variation in nitrogen uptake and turnover in two high-elevation soils: mineralization responses are site-dependent

In arctic and alpine ecosystems, soil nitrogen (N) dynamics can differ markedly between winter and summer months, and nitrogen losses can be measurable during the spring and fall transitions. To explore the effect of seasonality on biogeochemical processes in a temperate alpine environment, we used a combination of field incubations (year-round) and ¹⁵N tracer additions (late fall, early spring, summer) to characterize soil N dynamics in a wet and dry meadow in the Sierra Nevada, California. The snowmelt to early summer season marked a period of high ¹⁵N uptake and turnover in the two soils, coincident with the increase in microbial N pools at the start of snowmelt (wet and dry meadow); an increase in net N mineralization and net nitrification as snowmelt progressed (wet meadow only); and measureable net production of ¹⁵N-NH₄ ⁺ in mid-summer (wet and dry meadow). Whereas fluctuations in microbial biomass were generally synchronous between the wet and dry meadow soils, only wet meadow soils appeared to mineralize N in response to declines in the microbial N pool. Net N mineralization and net nitrification rates in the dry meadow soil were negligible on all but one sampling date, in spite of periodic decreases in biomass of up to 60%. Across both sites, high ¹⁵N recoveries in microbial biomass N, rapid ¹⁵N-NH₄ ⁺ turnover, and low or negative net ¹⁵N-NH₄ ⁺ fluxes suggested tight cycling of N, particularly in the late fall and early spring.     

Inorganic nitrogen and microbial biomass dynamics before and during spring snowmelt

Recent work in seasonally snow covered ecosystems has identifiedthawed soil and high levels of heterotrophic activity throughout the winterunder consistent snow cover. We performed measurements during the winter of1994 to determine how the depth and timing of seasonal snow cover affectsoil microbial populations, surface water NO loss during snowmelt, and plant Navailability early in the growing season. Soil under early accumulating,consistent snow cover remained thawed during most of the winter and bothmicrobial biomass and soil inorganic N pools gradually increased under thesnowpack. At the initiation of snowmelt, microbial biomass N pools increasedfrom 3.0 to 5.9 g n m^-2,concurrent with a decrease in soil inorganic N pools. During the latterstages of snowmelt, microbial biomass N pools decreased sharply without aconcurrent increase in inorganic N pools or significant leaching losses. Incontrast, soil under inconsistent snow cover remained frozen during most ofthe winter. During snowmelt, microbial biomass initially increased from 1.7to 3.1 g N m^-2 and thendecreased as sites became snow-free. In contrast to smaller pool sizes,NO export during snowmeltfrom the inconsistent snow cover sites of 1.14 (±0.511) g N m^-2 was significantly greater (p< 0.001) than the 0.27 (±0.16) g N m^-2 exported from sites with consistent snowcover. These data suggest that microbial biomass in consistentlysnow-covered soil provides a significant buffer limiting the export ofinorganic N to surface water during snowmelt. However, this buffer is verysensitive to changes in snowpack regime. Therefore, interannual variabilityin the timing and depth of snowpack accumulation may explain the year toyear variability in inorganic N concentrations in surface water theseecosystems.     

Interactive effects of warming and increased nitrogen deposition on 15N tracer retention in a temperate old field: seasonal trends

The extent to which increased atmospheric nitrogen (N) deposition will drive changes in plant productivity and species composition over the next century will depend on how other influential global change factors, such as climate warming, affect the N retention of ecosystems. We examined the interactive effects of simulated climate warming and N deposition on the recoveries of ¹⁵N‐labeled ammonium and ¹⁵N‐labeled nitrate tracers added as a pulse to grass‐dominated, temperate old‐field plots at spring thaw. In addition to the year‐round warming treatment, a winter‐only warming treatment was applied to a set of plots to explore the contribution of this component of climate warming to the overall warming effect. By the end of the plant growing season, there was approximately twice as much ¹⁵N enrichment in the plant roots and bulk soil from ¹⁵NH₄⁺‐addition plots than from ¹⁵NO₃^?‐addition plots, but there were no effects of warming or N fertilization on ¹⁵N recovery. Over winter, approximately half of the excess ¹⁵N present in plant shoots was lost, which corresponded with large ¹⁵N losses from bulk soil in N fertilized plots and large ¹⁵N increases in bulk soil in nonfertilized plots. By the next spring, there was decreased ¹⁵N recovery in plants in response to N fertilization, which was largely offset by increases in plant ¹⁵N recovery in response to year‐round warming. However, ¹⁵N retention in bulk soil, where the major part of the ¹⁵N label was recovered, was approximately 40% higher in nonfertilized plots than in N fertilized plots. Overall, our results indicate that climate warming increases plant N sequestration in this system but this effect is overwhelmed by the overall effect of nitrogen deposition on ecosystem N losses.     

Plant and Soil N Response of Southern Californian Semi-arid Shrublands After 1 Year of Experimental N Deposition

Large inputs of atmospheric N from dry deposition accumulate on vegetation and soil surfaces of southern Californian chaparral and coastal sage scrub (CSS) ecosystems during the late-summer and early-fall and become available as a pulse following winter rainfall; however, the fate of this dry season atmospheric N addition is unknown. To assess the potential for dry season atmospheric N inputs to be incorporated into soil and/or vegetation N pools, an in situ N addition experiment was initiated in a post-fire chaparral and a mature CSS stand where 10 × 10 m plots were exposed to either ambient N deposition (control) or ambient +50 kg N ha⁻¹ (added N) added as NH₄NO₃ during a single application in October 2003. After 1 year of N addition, plots exposed to added N had significantly higher accumulation of extractable inorganic N (NH₄−N + NO₃−N) on ion exchange resins deployed in the 0–10 cm mineral soil layer and higher soil extractable N in the subsurface (30–40 cm) mineral soil than plots exposed to ambient N. Chaparral and CSS shrubs exposed to added N also exhibited a significant increase in tissue N concentration and a decline in the tissue C:N ratio, and added N significantly altered the shrub tissue δ ¹⁵N natural abundance. Leaching of inorganic N to 1 m below the soil surface was on average 2–3 times higher in the added N plots, but large within treatment variability cause these differences to be statistically insignificant. Although a large fraction of the added N could not be accounted for in the shrub and soil N pools investigated, these observations suggest that dry season N inputs can significantly and rapidly alter N availability and shrub tissue chemistry in Mediterranean-type chaparral and CSS shrublands of southern California.     

Rhizosphere effects on ion concentrations near different root zones of Norway spruce (Picea abies (L.) Karst.) and root types of Douglas-fir (Pseudotsuga menziesii L.) seedlings

In many temperate ecosystems, rates of atmospheric nitrogen deposition remain high over winter despite decreased agricultural activity over this season. The extent to which this nitrogen is accessible for plant growth over the following growing season may depend strongly on uptake by plants and soil microorganisms from late fall through early spring, when the majority of aboveground plant tissue has senesced. We added Ca(¹⁵NO₃)₂ (5 atom %¹⁵N) at a rate of 2 g m^?2 of N (corresponding to 100 mg ¹⁵N m^?2) to the surface of plots in a temperate old field during either late fall, winter, spring melt or early spring. We quantified the recovery of excess ¹⁵N in the soil microbial biomass and soil extracts following spring melt and in aboveground plant tissue at the peak of the plant growing season. Nitrate additions had no significant effect on total aboveground plant biomass, relative species abundance or percent tissue nitrogen. However, mean excess ¹⁵N in aboveground plant tissue varied significantly among treatments, with values of 8.1, 2.6, 0.3 and 7.3 mg m^?2 for late fall, winter, spring melt and early spring addition plots, respectively. Corresponding values of excess ¹⁵N were 3.1, 1.4 and 0.2 mg m^?2 in microbial biomass, and 0.17, 0.07 and 0.03 mg m^?2 in soil extracts, for late fall, winter and spring melt addition plots, respectively. Overall, these results indicate that nitrogen retention from late fall through early spring may depend highly on plant uptake in this system, and that only a small fraction of the nitrogen that accumulates in the winter snow pack may be available to plants.     

Additive Effects of Warming and Increased Nitrogen Deposition in a Temperate Old Field: Plant Productivity and the Importance of Winter

Both climate warming and atmospheric nitrogen (N) deposition are predicted to alter plant productivity and species composition over the next century. However, the extent to which their effects may interact is unclear. For example, over winter, the effects of warming on soil freezing dynamics may promote ecosystem N losses, thereby limiting increases in productivity in response to warming, yet these losses may be compensated for by increased N deposition. We measured plant production and species composition in response to warming (winter-only or year-round) and N addition in a temperate old field. We used shoot allometric relationships to estimate aboveground production non-destructively and sampled root biomass destructively throughout two growing seasons. We also used spectral data (normalized difference vegetation index—NDVI) to examine the treatment effects on the timing of plant green-up and senescence. In 2007, which featured an exceptionally dry summer, there were no treatment effects on plant growth. However, in 2008, warming (both winter-only and year-round) and N addition combined approximately doubled aboveground productivity, and these effects were additive. Warming increased root biomass, but no N effect was evident. Conversely, N addition increased NDVI, but NDVI was unresponsive to warming. Overall, our results do not support the hypothesis that warming-induced changes to soil freezing dynamics limit plant productivity in our system. On the contrary, they demonstrate that winter warming alone can increase primary productivity to the same extent as year-round warming, and that this effect may interact very strongly with interannual variation in precipitation.     

N uptake and growth responses to sub-lethal freezing in the grass Poa pratensis L.

Background and aims

Climate warming has the potential to increase both the exposure and vulnerability of grass roots to frost in temperate regions by reducing snow cover and altering the timing of cold acclimation. Despite a strong research focus on the direct effects of freezing on grass mortality, the direct sub-lethal effects of freezing on grass performance have not been well-characterized. We examined sub-lethal responses of the grass Poa pratensis to variation in the timing, severity, rate and length of freezing.

Methods

We assessed short term root functional responses (¹⁵N uptake) and longer term plant growth responses to freezing administered both under controlled conditions in a refrigerated incubator, and in the field by manipulating snow and litter cover.

Results

In fall and spring, ¹⁵N uptake declined in response to 1?day of freezing down to ?10?°C or to 3?days of freezing at ?5?°C, whereas in winter, ¹⁵N uptake was insensitive to freezing. Long term growth responses were similar, with reduced growth only occurring for grasses frozen for 3?days at ?5?°C in spring, but not for grasses frozen in fall or winter. Snow and litter removal intensified soil freezing over winter, but did not significantly affect plant growth.

Conclusions

Our results demonstrate that while P. pratensis is relatively tolerant to frost damage over winter, it may be vulnerable to sub-lethal frost effects in fall, and particularly in spring. These sub-lethal effects occur at temperatures approximately 15–20?°C warmer than the published LT₅₀ values for this species.     

Fate of ammonium 15N in a Norway spruce forest under long-term reduction in atmospheric N deposition

In the last decades, in particular forest ecosystems became increasingly N saturated due to elevated atmospheric N deposition, resulting from anthropogenic N emission. This led to serious consequences for the environment such as N leaching to the groundwater. Recent efforts to reduce N emissions raise the question if, and over what timescale, ecosystems recover to previous conditions. In order to study the effects on N distribution and N transformation processes under the lowered N deposition treatment, we investigated the fate of deposited NH₄ ⁺-¹⁵N in soil of a N-saturated Norway spruce forest (current N deposition: 34 kg ha^?1 year^?1; critical N load: 14 kg ha^?1 year^?1), where N deposition has been reduced to 11.5 kg ha^?1 year^?1 since 14.5 years. We traced the deposited ¹⁵N in needle litter, bulk soil, and amino acids, microbial biomass and inorganic N in soil. Under reduced N deposition, 123 ± 23% of the deposited N was retained in bulk soil, while this was only 72 ± 15% under ambient deposition. We presume that with reduced deposition the amount of deposited N was small enough to become completely immobilized in plant and soil and no leaching losses occurred. Trees receiving reduced N deposition showed a decline in N content as well as in ¹⁵N incorporation into needle litter, indicating reduced N plant uptake. In contrast, the distribution of ¹⁵N within the soil over active microbial biomass, microbial residues and inorganic N was not affected by the reduced N deposition. We conclude that the reduction in N deposition impacted only plant uptake and drainage losses, while microbial N transformation processes were not influenced. We assume changes in the biological N turnover to start with the onset of the decomposition of the new, N-depleted litter.     

Interactive Effects of Nitrogen and Phosphorus on Soil Microbial Communities in a Tropical Forest

Elevated nitrogen (N) deposition in humid tropical regions may exacerbate phosphorus (P) deficiency in forests on highly weathered soils. However, it is not clear how P availability affects soil microbes and soil carbon (C), or how P processes interact with N deposition in tropical forests. We examined the effects of N and P additions on soil microbes and soil C pools in a N-saturated old-growth tropical forest in southern China to test the hypotheses that (1) N and P addition will have opposing effects on soil microbial biomass and activity, (2) N and P addition will alter the composition of the microbial community, (3) the addition of N and P will have interactive effects on soil microbes and (4) addition-mediated changes in microbial communities would feed back on soil C pools. Phospholipid fatty acid (PLFA) analysis was used to quantify the soil microbial community following four treatments: Control, N addition (15 g N m⁻² yr⁻¹), P addition (15 g P m⁻² yr⁻¹), and N&P addition (15 g N m⁻² yr⁻¹ plus 15 g P m⁻² yr⁻¹). These were applied from 2007 to 2011. Whereas additions of P increased soil microbial biomass, additions of N reduced soil microbial biomass. These effects, however, were transient, disappearing over longer periods. Moreover, N additions significantly increased relative abundance of fungal PLFAs and P additions significantly increased relative abundance of arbuscular mycorrhizal (AM) fungi PLFAs. Nitrogen addition had a negative effect on light fraction C, but no effect on heavy fraction C and total soil C. In contrast, P addition significantly decreased both light fraction C and total soil C. However, there were no interactions between N addition and P addition on soil microbes. Our results suggest that these nutrients are not co-limiting, and that P rather than N is limiting in this tropical forest.     

Minor contribution of leaf litter to N nutrition of beech (Fagus sylvatica) seedlings in a mountainous beech forest of Southern Germany

Aims

Our aims were to characterize the fate of leaf-litter-derived nitrogen in the plant-soil-microbe system of a temperate beech forest of Southern Germany and to identify its importance for N nutrition of beech seedlings.

Methods

¹⁵N-labelled leaf litter was traced in situ into abiotic and biotic N pools in mineral soil as well as into beech seedlings and mycorrhizal root tips over three growing seasons.

Results

There was a rapid transfer of ¹⁵N into the mineral soil already 21 days after tracer application with soil microbial biomass initially representing the dominant litter-N sink. However, ¹⁵N recovery in non-extractable soil N pools strongly increased over time and subsequently became the dominant ¹⁵N sink. Recovery in plant biomass accounted for only 0.025 % of ¹⁵N excess after 876 days. After three growing seasons, ¹⁵N excess recovery was characterized by the following sequence: non-extractable soil N?>>?extractable soil N including microbial biomass?>>?plant biomass?>?ectomycorrhizal root tips.

Conclusions

After quick vertical dislocation and cycling through microbial N pools, there was a rapid stabilization of leaf-litter-derived N in non-extractable N pools of the mineral soil. Very low ¹⁵N recovery in beech seedlings suggests a high importance of other N sources such as root litter for N nutrition of beech understorey.     

Snow depth, soil freezing and nitrogen cycling in a northern hardwood forest landscape

Increases in soil freezing associated with decreases in snow cover have been identified as a significant disturbance to nitrogen (N) cycling in northern hardwood forests. We created a range of soil freezing intensity through snow manipulation experiments along an elevation gradient at the Hubbard Brook Experimental Forest (HBEF) in the White Mountains, NH USA in order to improve understanding of the factors regulating freeze effects on nitrate (NO₃ ^?) leaching, nitrous oxide (N₂O) flux, potential and in situ net N mineralization and nitrification, microbial biomass carbon (C) and N content and respiration, and denitrification. While the snow manipulation treatment produced deep and persistent soil freezing at all sites, effects on hydrologic and gaseous losses of N were less than expected and less than values observed in previous studies at the HBEF. There was no relationship between frost depth, frost heaving and NO₃ ^? leaching, and a weak relationship between frost depth and winter N₂O flux. There was a significant positive relationship between dissolved organic carbon (DOC) and NO₃ ^? concentrations in treatment plots but not in reference plots, suggesting that the snow manipulation treatment mobilized available C, which may have stimulated retention of N and prevented treatment effects on N losses. While the results support the hypothesis that climate change resulting in less snow and more soil freezing will increase N losses from northern hardwood forests, they also suggest that ecosystem response to soil freezing disturbance is affected by multiple factors that must be reconciled in future research.     

Biogeochemical pools and fluxes of carbon and nitrogen in a maritime tundra near penguin colonies along the Antarctic Peninsula

There is limited information regarding biogeochemical pools and fluxes in maritime tundra ecosystems along the Antarctic Peninsula. To collect baseline information on biogeochemical processes in a tundra ecosystem dominated by two vascular plant species (Colobanthus quitensis and Deschampsia antarctica) at Biscoe Point off the coast of Anvers Island, we measured pools and fluxes of C and N in transplanted tundra microcosm cores, complemented with sampling of precipitation and surface runoff. Snow and snowmelt from the tundra collection site and soil leachates from the cores were enriched with N and dissolved organic carbon compared to precipitation and snowmelt samples collected at Palmer Station, indicating high loading of N and organic matter from the penguin colonies adjacent to the tundra site. Relatively high values of δ¹⁵N in the live and dead biomass of D. antarctica and C. quitensis (5.6–25.1‰) indicated an enrichment of N in this tundra ecosystem, possibly through N inputs from adjacent penguin colonies. Stepwise multiple linear regressions found that ecosystem respiration and gross primary production were best predicted by live biomass of D. antarctica, suggesting a disproportionately high contribution of D. antarctica to CO₂ fluxes. The cores with higher δ¹⁵N and lower δ¹³C in the soil organic horizon exhibited higher CO₂ fluxes. The results suggest that abundant N inputs from penguin colonies and the competitive balance between plant species might play a critical role in the response of tundra ecosystems along the Antarctic Peninsula to projected climate change.     

The fate of15N enriched throughfall in two coniferous forest stands at different nitrogen deposition levels

The stable isotope¹⁵N was added as (¹⁵NH₄)₂SO₄ to throughfall water for one year, to study the fate of the deposited nitrogen at different levels of N deposition in two N saturated coniferous forests ecosystems in the Netherlands. The fate of the¹⁵N was followed at high-N (44–55 kg N ha^–1 yr^–1) 1) and low-N (4–6 kg N ha^–1 yr^–1) deposition in plots established under transparent roofs build under the canopy in a Douglas fir (Pseudotsuga menziesii (Mirb.) Franco.) and Scots pine (Pinus sylvestris L.) forest.The applied¹⁵N was detectable in needles and twigs, the soil and soil water leaching below the rooting zone (90 cm depth). Total¹⁵N recovery in major ecosystem compartments was 71–100% during two successive growing seasons after the start of a year-round¹⁵N application to throughfall-N. Nine months after the year-round¹⁵N application, the¹⁵N assimilated into tree biomass was 29–33% of the¹⁵N added in the Douglas fir stand and less than 17% in the Scots pine stand. At the same time total¹⁵N retention in the soil (down to 70 cm) of the high-N plots was about 37% of the deposited¹⁵NH₄-N, whereas 46% and 65% of the¹⁵N was found in the soil of the low-N deposition plots at the Douglas fir and Scots pine stand, respectively. The organic layers accounted for 60% of the¹⁵N retained in the soil. The total N deposition exceeded the demand of the vegetation and microbial immobilization. Total¹⁵N leaching losses within a year (below 90 cm) were 10–20% in the high-N deposition plots in comparison to 2–6% in the lowered nitrogen input plots. Relative retention in the soil and vegetation increased at lower N-input levels.Species differences in uptake and tree health seem to contribute to lower¹⁵N recoveries in the Scots pine trees compared to the Douglas fir trees. The excessive N deposition and resulting N saturation lead to conditions were the health and functioning of biota were negatively influenced. At decreased N deposition, lower leaching losses together with increased soil and plant retention indicated a change in the fate of the¹⁵N deposited. This may have resulted from changes in ecosystem processes, and thus a shift along the continuum of N saturation to N limitation.     

Nitrogen deposition and carbon sequestration in alpine meadows

Nitrogen deposition experiments were carried out in alpine meadow ecosystems in Qinghai-Xizang Plateau in China, in order to explore the contribution of nitrogen deposition to carbon sequestration in alpine meadows. Two methods were used in this respect. First, we used the allocation of ¹⁵N tracer to soil and plant pools. Second, we used increased root biomass observed in the nitrogen-amended plots. Calculating enhanced carbon storage, we considered the net soil CO₂ emissions exposed to nitrogen deposition in alpine meadows. Our results show that nitrogen deposition can enhance the net soil CO₂ emissions, and thus offset part of carbon uptake by vegetation and soils. It means that we have to be cautious to draw a conclusion when we estimate the contribution of nitrogen deposition to carbon sequestration based on the partitioning of ¹⁵N tracer in terrestrial ecosystems, in particular in N-limited ecosystems. Even if we assess the contribution of nitrogen deposition to carbon sequestration based on increased biomass exposed to nitrogen deposition in terrestrial ecosystems, likewise, we have to consider the effects of nitrogen deposition on the soil CO₂ emissions.     

Multiyear fate of a 15N tracer in a mixed deciduous forest: retention,redistribution, and differences by mycorrhizal association

The impact of atmospheric nitrogen deposition on forest ecosystems depends in large part on its fate. Past tracer studies show that litter and soils dominate the short‐term fate of added ¹⁵N, yet few have examined its longer term dynamics or differences among forest types. This study examined the fate of a ¹⁵N‐ tracer over 5–6 years in a mixed deciduous stand that was evenly composed of trees with ectomycorrhizal and arbuscular mycorrhizal associations. The tracer was expected to slowly mineralize from its main initial fate in litter and surface soil, with some ¹⁵N moving to trees, some to deeper soil, and some net losses. Recovery of added ¹⁵N in trees and litterfall totaled 11.3% both 1 and 5–6 years after the tracer addition, as ¹⁵N redistributed from fine and especially coarse roots into cumulative litterfall and small accumulations in woody tissues. Estimates of potential carbon sequestration from tree ¹⁵N recovery amounted to 12–14 kg C per kg of N deposition. Tree ¹⁵N acquisition occurred within the first year after the tracer addition, with no subsequent additional net transfer of ¹⁵N from detrital to plant pools. In both years, ectomycorrhizal trees gained 50% more of the tracer than did trees with arbuscular mycorrhizae. Much of the ¹⁵N recovered in wood occurred in tree rings formed prior to the ¹⁵N addition, demonstrating the mobility of N in wood. Tracer recovery rapidly decreased over time in surface litter material and accumulated in both shallow and deep soil, perhaps through mixing by earthworms. Overall, results showed redistribution of tracer ¹⁵N through trees and surface soils without any losses, as whole‐ecosystem recovery remained constant between 1 and 5–6 years at 70% of the ¹⁵N addition. These results demonstrate the persistent ecosystem retention of N deposition even as it redistributes, without additional plant uptake over this timescale.     

Dynamics of mineral N availability in grassland ecosystems under increased [CO2]: hypotheses evaluated using the Hurley Pasture Model

The following arguments are outlined and then illustrated by the response of the Hurley Pasture Model to [CO₂] doubling in the climate of southern Britain. 1. The growth of N-limited vegetation is determined by the concentration of N in the soil mineral N pools and high turnover rates of these pools (i.e., large input and output fluxes) contribute positively to growth. 2. The size and turnover rates of the soil mineral N pools are determined overwhelmingly by N cycling into all forms of organic matter (plants, animals, soil biomass and soil organic matter — `immobilisation' in a broad sense) and back again by mineralisation. Annual system N gains (by N₂ fixation and atmospheric deposition) and losses (by leaching, volatilisation, nitrification and denitrification) are small by comparison. 3. Elevated [CO₂] enriches the organic matter in plants and soils with C, which leads directly to increased removal of N from the soil mineral N pools into plant biomass, soil biomass and soil organic matter (SOM). ‘Immobilisation’ in the broad sense then exceeds mineralisation. This is a transient state and as long as it exists the soil mineral N pools are depleted, N gaseous and leaching losses are reduced and the ecosystem gains N. Thus, net immobilisation gradually increases the N status of the ecosystem. 4. At the same time, elevated [CO₂] increases symbiotic and non-symbiotic N₂ fixation. Thus, more N is gained each year as well as less lost. Effectively, the extra C fixed in elevated [CO₂] is used to capture and retain more N and so the N cycle tracks the C cycle. 5. However, the amount of extra N fixed and retained by the ecosystem each year will always be small (ca. 5–10 kg N ha^-1 yr^-1) compared with amount of N in the immobilisation-mineralisation cycle (ca. 1000 kg N ha^-1 yr^-1). Consequently, the ecosystem can take decades to centuries to gear up to a new equilibrium higher-N state. 6. The extent and timescale of the depletion of the mineral N pools in elevated [CO₂] depends on the N status of the system and the magnitude of the overall system N gains and losses. Small changes in the large immobilisation—mineralisation cycle have large effects on the small mineral N pools. Consequently, it is possible to obtain a variety of growth responses within 1–10 year experiments. Ironically, ecosystem models — artificial constructs — may be the best or only way of determining what is happening in the real world. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of nitrogen additions on above- and belowground carbon dynamics in two tropical forests

Anthropogenic nitrogen (N) deposition is increasing rapidly in tropical regions, adding N to ecosystems that often have high background N availability. Tropical forests play an important role in the global carbon (C) cycle, yet the effects of N deposition on C cycling in these ecosystems are poorly understood. We used a field N-fertilization experiment in lower and upper elevation tropical rain forests in Puerto Rico to explore the responses of above- and belowground C pools to N addition. As expected, tree stem growth and litterfall productivity did not respond to N fertilization in either of these N-rich forests, indicating a lack of N limitation to net primary productivity (NPP). In contrast, soil C concentrations increased significantly with N fertilization in both forests, leading to larger C stocks in fertilized plots. However, different soil C pools responded to N fertilization differently. Labile (low density) soil C fractions and live fine roots declined with fertilization, while mineral-associated soil C increased in both forests. Decreased soil CO₂ fluxes in fertilized plots were correlated with smaller labile soil C pools in the lower elevation forest (R² = 0.65, p < 0.05), and with lower live fine root biomass in the upper elevation forest (R² = 0.90, p < 0.05). Our results indicate that soil C storage is sensitive to N deposition in tropical forests, even where plant productivity is not N-limited. The mineral-associated soil C pool has the potential to respond relatively quickly to N additions, and can drive increases in bulk soil C stocks in tropical forests.     

Turnover of residual 15N-labelled fertilizer N in soil following harvest of oilseed rape (t Brassica napus L.)

We studied the fate of ¹⁵N-labelled fertilizer nitrogen in a sandy loam soil after harvest of winter oilseed rape (Brassica napus L. cv. Ceres) given 100 or 200 kg N ha^-1 in spring, with or without irrigation. Our main objective was to quantify the temporal variations of the soil mineral N, the extractable soil organic N and soil microbial biomass N, and fertilizer derived N in these pools during autumn and winter. Nitrogen use efficiency of the oilseed rape crop varied from 47% of applied N in the 100N, irrigated treatment to 34% in the 200N, non-irrigated treatment. However, only in the latter treatment did we find significantly higher fertilizer derived soil mineral N than in the three other treatments which all had low soil mineral N contents at the first sampling after harvest (8 days after stubble tillage). Between 31% and 42% of the applied N could not be accounted for in the harvested plants or 0-15 cm soil layer at this first sampling. Over the following autumn and winter none of the remaining fertilizer derived soil N was lost from the 0–5 cm depth, but from the 5–15 cm depth a marked proportion of N derived from fertilizer was lost, probably by leaching. Negligible amounts of fertilizer derived extractable soil organic and mineral N (<1 kg N ha^-1, 0-15 cm) were found in all treatments after the first sampling.Soil microbial biomass N was not significantly affected by treatments and showed only small temporal variability (±11% of the mean 76 kg N ha^-1, 0- 15 cm depth). Surprisingly, the average amount of soil microbial biomass N derived from fertilizer was significantly affected by the treatments, with the extremes being 5.5 and 3.1 kg N ha^-1 in the 200N, non-irrigated and 100N, irrigated treatments, respectively. Also, the estimated exponential decay rate of microbial biomass N derived from fertilizer, differed greatly (2 fold) between these two treatments, indicating highly different microbial turnover rates in spite of the similar total microbial biomass N values. In studies utilising ¹⁵N labelling to estimate turnover rates of different soil organic matter pools this finding is of great importance, because it may question the assumption that turnover rates are not affected by the insertion of the label.