Leaf-cutting ant (Atta Sexdens) and nutrient cycling: deep soil inorganic nitrogen stocks, mineralization, and nitrification in Eastern Amazonia

Nest excavation and agricultural activities of the leaf-cutting ant Atta sexdens create complex belowground heterogeneity in secondary forests of Eastern Amazonia. We examined the effects of this heterogeneity on inorganic-N stocks, net mineralization, and net nitrification to test the hypothesis that the bulk soil of the nests has higher net rates of mineralization and nitrification than soil that was not affected by the influences of ant nests, throughout the profile. This study was conducted in a secondary forest at Fazenda Vitoria, near Paragominas in the Eastern Brazilian Amazon, where a previous study showed that the bulk soil of ant nests had elevated NO₃⁻. The results of the inorganic-N measurements were consistent with the previous study, showing elevated NO₃⁻ deep in the soil profile of the nests. However, neither net mineralization nor net nitrification were significantly greater at depth in the mineral soil of the nests compared to soil that was not influenced by nests (P=0.05), although variability was higher in the nest soil. These results suggest that the NO₃⁻ may have diffused into the surrounding mineral from the N-rich organic matter buried by the ants in chambers within the deep soil.     

Impact of land-use types on soil nitrogen net mineralization in the sandstorm and water source area of Beijing,China

Changes of land-use type (LUT) can affect soil nutrient pools and cycling processes that relate long-term sustainability of ecosystem, and can also affect atmospheric CO₂ concentrations and global warming through soil respiration. We conducted a comparative study to determine NH₄⁺ and NO₃⁻ concentrations in soil profiles (0–200 cm) and examined the net nitrogen (N) mineralization and net nitrification in soil surface (0–20 cm) of adjacent naturally regenerated secondary forests (NSF), man-made forests (MMF), grasslands and cropland soils from the windy arid and semi-arid Hebei plateau, the sandstorm and water source area of Beijing, China. Cropland and grassland soils showed significantly higher inorganic N concentrations than forest soils. NO₃⁻-N accounted for 50–90% of inorganic N in cropland and grassland soils, while NH₄⁺-N was the main form of inorganic N in NSF and MMF soils. Average net N-mineralization rates (mg kg⁻¹ d⁻¹) were much higher in native ecosystems (1.51 for NSF soils and 1.24 for grassland soils) than in human disturbed LUT (0.15 for cropland soils and 0.85 for MMF soils). Net ammonification was low in all the LUT while net nitrification was the major process of net N mineralization. For more insight in urea transformation, the increase in NH₄⁺ and, NO₃⁻ concentrations as well as C mineralization after urea addition was analyzed on whole soils. Urea application stimulated the net soil C mineralization and urea transformation pattern was consistent with net soil N mineralization, except that the rate was slightly slower. Land-use conversion from NSF to MMF, or from grassland to cropland decreased soil net N mineralization, but increased net nitrification after 40 years or 70 years, respectively. The observed higher rates of net nitrification suggested that land-use conversions in the Hebei plateau might lead to N losses in the form of nitrate.     

Seasonal,annual, and treatment-induced variation in available nitrogen pools and nitrogen-cycling processes in soils of two Douglas-fir stands

Summary Forest-floor and 0–10 cm depth mineral soil horizons in two stands of Douglas fir were sampled for available NH₄ ⁺-N and NO₃ ^–-N, N-mineralization potentials, and nitrification potentials for 2 years. The plots in each stand were sampled for 1 year, treated with either ammonium sulfate, carbohydrate (sawdust-sucrose), irrigation, carbohydrate plus irrigation, or no treatment (control), and then sampled for 1 year following treatment. In general, the direction of change following the treatments was the same for both the forest-floor and the mineral soils. Fertilization increased the NH₄ ⁺-N and NO₃ ^–-N pools, nitrification potential, and N-mineralization potential, while treatment with carbohydrate decreased all of these characteristics. Irrigation generally increased NH₄ ⁺-N pools, nitrification potential, and N-mineralization potential, but decreased these characteristics in the soil at one site. Irrigation plus carbohydrate gave similar results to those of carbohydrate alone. Treatments altered pool sizes and/or potentials, but did not reduce within-year variance in any of these characteristics. Distinct seasonal patterns occurred in all measurements, suggesting that control of short-term variation in N-transformation processes is by factors which are dynamic in nature.     

Biological and physical effects of non-native earthworms on nitrogen cycling in riparian soils

Global nitrogen cycling is being altered by anthropogenic disturbances including invasion by non-native species. European and Asian earthworms have invaded northern temperate forests in North America with dramatic consequences for litter thickness, forest floor plant diversity, and soil nitrogen cycling. Invasive earthworms present at the boundary of terrestrial and aquatic ecosystems (i.e., riparian zones) may alter the flux of nitrogen into adjacent aquatic ecosystems. We examined how nitrogen cycling in riparian soil responds to amendments of invasive earthworms or artificial earthworm burrows. In earthworm-free riparian plots (0.25 m²), we established treatments of invasive earthworms (60 g fresh mass·m⁻²), artificial burrows (120 m⁻²), or control plots and sampled the plots after 30 days. Before and after treatment application we measured major soil characteristics (water-filled pore space, organic matter, and pH), nitrogen pools (exchangeable NH₄⁺ and NO₃⁻), and nitrogen transformation rates (net N-mineralization, net nitrification, and denitrification). Exchangeable NH₄⁺ and NO₃⁻ changed through time but did not differ among treatments. Net N-mineralization and net nitrification rates did not change through time and were similar across all treatments. However, denitrification rates in plots with added earthworms were 4 times greater than rates in control and burrow-only plots, which represents a large rapid increase in gaseous nitrogen flux out of these riparian soils. For all response variables, artificial burrows responded similarly to control plots, suggesting that earthworm biological activity (i.e., feeding, excretion, and mucus production) rather than physical effects (i.e., burrowing and soil aeration) drove the changes in nitrogen cycling. Examination of soil nitrogen pool and flux measurements suggest that this increase in denitrification was coupled with NH₄⁺ consumption by nitrifying bacteria, but future studies are needed to confirm this hypothesis. We conclude that the activity of invasive earthworms in riparian zones can increase the flux of N out of riparian zones, but the hydrologic context of the riparian zone (e.g., pore-water residence time) ultimately controls whether denitrification or nitrate leaching is the dominant flux of N.     

Initial recovery of soil carbon and nitrogen pools and dynamics following disturbance in jack pine forests: A comparison of wildfire and clearcut harvesting

Forests naturally maintained by stand-replacing wildfires are often managed with clearcut harvesting, yet we know little about how replacing wildfire with clearcutting affects soil processes and properties. We compared the initial recovery of carbon (C) and nitrogen (N) pools and dynamics following disturbance in jack pine (Pinus banksiana) stands in northern Lower Michigan, USA, by sampling soils (Oa+A horizons) from three “treatments”: 3-6-year-old harvest-regenerated stands, 3-6-year-old wildfire-regenerated stands and 40-55-year-old intact, mature stands (n=4 stands per treatment). We measured total C and N; microbial biomass and potentially mineralizable C and N; net nitrification; and gross rates of N mineralization and nitrification. Burned stands exhibited reduced soil N but not C, whereas clearcut and mature stands had similar quantities of soil organic matter. Both disturbance types reduced microbial biomass C compared to mature stands; however, microbial biomass N was reduced in burned stands but not in clearcut stands. The experimental C and N mineralization values were fit to a first-order rate equation to estimate potentially mineralizable pool size (C₀ and N₀) and rate parameters. Values for C₀ in burned and clearcut stands were approximately half that of the mature treatment, with no difference between disturbance types. In contrast, N₀ was lowest in the wildfire stands (170.2 μg N g⁻¹), intermediate in the clearcuts (215.4 μg N g⁻¹) and highest in the mature stands (244.6 μg N g⁻¹). The most pronounced difference between disturbance types was for net nitrification. These data were fit to a sigmoidal growth equation to estimate potential NO₃⁻ accumulation (Nit_max) and kinetic parameters. Values of Nit_max in clearcut soils exceeded that of wildfire and mature soils (149.2 vs. 83.5 vs. 96.5 μg NO₃⁻-N g⁻¹, respectively). Moreover, the clearcut treatment exhibited no lag period for net NO₃⁻ production, whereas the burned and mature treatments exhibited an approximate 8-week lag period before producing appreciable quantities of NO₃⁻. There were no differences between disturbances in gross rates of mineralization or nitrification; rather, lower NO₃⁻ immobilization rates in the clearcut soils, 0.20 μg NO₃⁻ g⁻¹ d⁻¹ compared to 0.65 in the burned soils, explained the difference in net nitrification. Because the mobility of NO₃⁻ and NH₄⁺ differs markedly in soil, our results suggest that differences in nitrification between wildfire and clearcutting could have important consequences for plant nutrition and leaching losses following disturbance.     

Soil microbial biomass and nitrogen transformations among five tree species of the Catskill Mountains, New York, USA

We measured soil microbial biomass nitrogen (MBN), microbial uptake of ¹⁵N, potential net mineralization and net nitrification in the laboratory to determine the influence of tree species on nitrogen (N) transformations in soils of the Catskills Mountains, New York, USA. Organic horizon soils were taken from single species plots of beech (Fagus grandifolia), hemlock (Tsuga canadensis), red oak (Quercus rubra), sugar maple (Acer saccharum) and yellow birch (Betula alleghaniensis). ¹⁵NH₄Cl was added to the soils and N pools were sampled at 1, 3, 10 and 28 days to examine microbial uptake of ¹⁵N over time. Soil MBN was about 60% lower in red oak and sugar maple soils than in the other three species. Soil pools of NO₃⁻ and rates of net nitrification were significantly greater in soils associated with sugar maple than hemlock, red oak and yellow birch. With the exception of sugar maple soils, microbial recovery of ¹⁵N was significantly greater after 10 and 28 days compared to 60 min and 1 day following ¹⁵N tracer addition. Microbial ¹⁵N recovery declined significantly within sugar maple stands within the first 3 days of incubation. Soil carbon to nitrogen ratio (C:N) was lowest in sugar maple soils and highest in red oak soils. However, correlations between soil C:N and MBN or rates of net mineralization and nitrification were not significant. Soil moisture could account for 22% of the variation in MBN and 36% of the variation in net mineralization. Soil microbial transformations of N vary among tree species stands and may have consequences for forest N retention and loss.     

The N2O product ratio of nitrification and its dependence on long-term changes in soil pH

The contribution of nitrification to the emission of nitrous oxide (N₂O) from soils may be large, but its regulation is not well understood. The soil pH appears to play a central role for controlling N₂O emissions from soil, partly by affecting the N₂O product ratios of both denitrification (N₂O/(N₂+N₂O)) and nitrification (N₂O/(NO₂⁻+NO₃⁻). Mechanisms responsible for apparently high N₂O product ratios of nitrification in acid soils are uncertain. We have investigated the pH regulation of the N₂O product ratio of nitrification in a series of experiments with slurries of soils from long-term liming experiments, spanning a pH range from 4.1 to 7.8. ¹⁵N labelled nitrate (NO₃⁻) was added to assess nitrification rates by pool dilution and to distinguish between N₂O from NO₃⁻ reduction and NH₃ oxidation. Sterilized soil slurries were used to determine the rates of chemodenitrification (i.e. the production of nitric oxide (NO) and N₂O from the chemical decomposition of nitrite (NO₂⁻)) as a function of NO₂⁻ concentrations. Additions of NO₂⁻ to aerobic soil slurries (with ¹⁵N labelled NO₃⁻ added) were used to assess its potential for inducing denitrification at aerobic conditions. For soils with pH?5, we found that the N₂O product ratios for nitrification were low (0.2-0.9‰) and comparable to values found in pure cultures of ammonia-oxidizing bacteria. In mineral soils we found only a minor increase in the N₂O product ratio with increasing soil pH, but the effect was so weak that it justifies a constant N₂O product ratio of nitrification for N₂O emission models. For the soils with pH 4.1 and 4.2, the apparent N₂O product ratio of nitrification was 2 orders of magnitude higher than above pH 5 (76‰ and 14‰). This could partly be accounted for by the rates of chemodenitrification of NO₂⁻. We further found convincing evidence for NO₂⁻-induction of aerobic denitrification in acid soils. The study underlines the role of NO₂⁻, both for regulating denitrification and for the apparent nitrifier-derived N₂O emission.     

Concurrent measurements of net mineralization, nitrification, denitrification and leaching from field incubated soil cores

An improved method is described for incubating intact soil cores in the field, which permits concurrent measurement of net mineralization, nitrification, denitrification and leaching. Cores were enclosed in PVC tubes with minimal disturbance to the physical state or to the natural cycles of wetting/drying, soil temperature and aeration during an incubation lasting 4–5 days. An example of the application of the method is given in which soils with contrasting drainage characteristics were compared. Over a 64-day experimental period, 58% of the mineralized nitrogen (N) in a freely drained soil was nitrified and 36% of the nitrate-N (NO₃ ^–-N) was denitrified. In a poorly drained soil, 72% of the mineralized N was nitrified and 63% of the NO₃ ^–-N was denitrified. In both soil types, 18% of the remaining NO₃ ^–-N was leached. Rates of nitrification were significantly correlated with net mineralization (r ²=0.41 and 0.52) and also closely correlated with denitrification (r ²=0.67 and 0.68) in the freely and poorly drained soils, respectively. Independent measurements of these processes, using alternative techniques (for the same period), compared favourably with measurements obtained with the improved incubation method. Adoption of this method has a number of advantages with respect to field net N mineralization, and also allows interpretation of the impact this may have on other N transformation processes. Received: 18 June 1997     

Soil biochemical and chemical changes in relation to mature spruce (Picea abies) forest conversion and regeneration

To investigate soil changes from forest conversion and regeneration, soil net N mineralization, potential nitrification, microbial biomass N, L‐asparaginase, L‐glutaminase, and other chemical and biological properties were examined in three adjacent stands: mature pure and dense Norway spruce (Picea abies (L.) Karst) (110 yr) (stand I), mature Norway spruce mixed with young beech (Fagus sylvatica) (5 yr) (stand II), and young Norway spruce (16 yr) (stand III). The latter two stands were converted or regenerated from the mature Norway spruce stand as former. The studied soils were characterized as having a very low pH value (2.9 – 3.5 in 0.01 M CaCl₂), a high total N content (1.06 – 1.94 %), a high metabolic quotient (qCO₂) (6.7 – 16.9 g CO₂ kg^–1 h^–1), a low microbial biomass N (1.1 – 3.3 % of total N, except LOf₁ at stand III), and a relatively high net N mineralization (175 – 1213 mg N kg^–1 in LOf₁ and Of₂, 4 weeks incubation). In the converted forest (stand II), C : N ratio and qCO₂ values in the LOf₁ layer decreased significantly, and base saturation and exchangeable Ca showed a somewhat increment in mineral soil. In the regenerated forest (stand III), the total N storage in the surface layers decreased by 30 %. The surface organic layers (LOf₁, Of₂) possessed a very high net N mineralization (1.5 – 3 times higher than those in other two stands), high microbial biomass (C, N), and high basal respiration and qCO₂ values. Meanwhile, in the Oh layer, the base saturation and the exchangeable Ca decreased. All studied substrates showed little net nitrification after the first period of incubation (2 weeks). In the later period of incubation (7 – 11 weeks), a considerable amount of NO₃‐N accumulated (20 – 100 % of total cumulative mineral N) in the soils from the two pure spruce stands (I, III). In contrast, there was almost no net NO₃‐N accumulation in the soils from the converted mixed stand (II) indicating that there was a difference in microorganisms in the two types of forest ecosystems. Soil microbial biomass N, mineral N, net N mineralization, L‐asparaginase, and L‐glutaminase were correlated and associated with forest management.     

Nitrogen mineralisation in the soil of indigenous oak and pine plantation forests in a Mediterranean environment

Nitrogen mineralisation in soils of various forest sites (pine plantation, natural and thinned oak) at Uluda? University campus in Bursa, Turkey was investigated continuously over a year by the field incubation method. Net nitrogen mineralisation and nitrification rates varied depending on sampling dates. Although nitrogen mineralisation and nitrification rates increased in the spring and summer months, there was no seasonal variation in the soils of the examined forests. Annual net nitrate (NO₃^?–N) accumulation in the upper soil layer (0–5 cm) was higher in Oak I and Oak II (14 kg ha y^?1 and 12 kg ha y^?1) than in the pine plantation (8 kg ha y^?1). While annual net NO₃^?–N accumulation (0–5 cm) varied between the oak forests (possibly due to forest management practices), annual net N_min values were similar in these forests. No significant correlation was found between the examined soil parameters and net nitrification and mineralisation rates in the soils (P > 0.05). These results indicate that tree species and forest management practices play important roles in N cycling in forest ecosystems.     

Natural N abundances of inorganic nitrogen in soil treated with fertilizer and compost under changing soil moisture regimes

This study was conducted to examine whether the applications of N-inputs (compost and fertilizer) having different N isotopic compositions (δ¹⁵N) produce isotopically different inorganic-N and to investigate the effect of soil moisture regimes on the temporal variations in the δ¹⁵N of inorganic-N in soils. To do so, the temporal variations in the concentrations and the δ¹⁵N of NH₄⁺ and NO₃⁻ in soils treated with two levels (0 and 150 mg N kg⁻¹) of ammonium sulfate (δ¹⁵N=−2.3‰) and compost (+13.9‰) during a 10-week incubation were compared by changing soil moisture regime after 6 weeks either from saturated to unsaturated conditions or vice versa. Another incubation study using ¹⁵N-labeled ammonium sulfate (3.05 ¹⁵N atom%) was conducted to estimate the rates of nitrification and denitrification with a numerical model FLUAZ. The δ¹⁵N values of NH₄⁺ and NO₃⁻ were greatly affected by the availability of substrate for each of the nitrification and denitrification processes and the soil moisture status that affects the relative predominance between the two processes. Under saturated conditions for 6 weeks, the δ¹⁵N of NH₄⁺ in soils treated with fertilizer progressively increased from +2.9‰ at 0.5 week to +18.9‰ at 6 weeks due to nitrification. During the same period, NO₃⁻ concentrations were consistently low and the corresponding δ¹⁵N increased from +16.3 to +39.2‰ through denitrification. Under subsequent water-unsaturated conditions, the NO₃⁻ concentrations increased through nitrification, which resulted in the decrease in the δ¹⁵N of NO₃⁻. In soils, which were unsaturated for the first 6-weeks incubation, the δ¹⁵N of NH₄⁺ increased sharply at 0.5 week due to fast nitrification. On the other hand, the δ¹⁵N of NO₃⁻ showed the lowest value at 0.5 week due to incomplete nitrification, but after a subsequence increase, they remained stable while nitrification and denitrification were negligible between 1 and 6 weeks. Changing to saturated conditions after the initial 6-weeks incubation, however, increased the δ¹⁵N of NO₃⁻ progressively with a concurrent decrease in NO₃⁻ concentration through denitrification. The differences in δ¹⁵N of NO⁻₃ between compost and fertilizer treatments were consistent throughout the incubation period. The δ¹⁵N of NO₃⁻ increased with the addition of compost (range: +13.0 to +35.4‰), but decreased with the addition of fertilizer (−10.8 to +11.4‰), thus resulting in intermediate values in soils receiving both fertilizer and compost (−3.5 to +20.3‰). Therefore, such differences in δ¹⁵N of NO₃⁻ observed in this study suggest a possibility that the δ¹⁵N of upland-grown plants receiving compost would be higher than those treated with fertilizer because NO₃⁻ is the most abundant N for plant uptake in upland soils.     

Controls on nitrification in a water-limited ecosystem: experimental inhibition of ammonia-oxidising bacteria in the Patagonian steppe

We studied controls on nitrification in an undisturbed water-limited ecosystem by inhibiting autotrophic nitrifying bacteria in soils with varying levels of vegetative cover. The activity of nitrifying bacteria was disrupted using nitrapyrin, 2-chloro-6-(trichloromethyl)-pyridine, under field conditions in three microenvironments (underneath shrubs, next to grasses and in bare soil). Ammonia-oxidising bacteria were detected by PCR analysis of DNA in soils. The inhibition of nitrification changed the concentrations of NO₃⁻ and NH₄⁺ in the soil, while the microenvironment was most important in determining the response of bacteria to the inhibitor. Nitrapyrin application resulted in a significant (p<0.05) reduction in soil NO₃⁻ concentration (39%) and a significant increase (p<0.001) in soil NH₄⁺ concentration (41%). Untreated bare-soil microenvironments had the lowest concentrations of NH₄⁺ (1.57 μg/g of dry soil) and NO₃⁻ (0.49 μg/g of dry soil) when compared to the other microenvironments, and showed the highest impacts of nitrification inhibition. For example, NH₄⁺ concentrations increased 288% and NO₃⁻ concentrations decreased 60% in inhibited bare-soil microenvironments. In contrast, untreated microenvironments underneath shrubs had the highest levels of NH₄⁺ (10.01 μg/g of dry soil) and NO₃⁻ (0.69 μg/g of dry soil), but showed no significant effects of inhibition of nitrification on soil nitrogen concentrations.     

Nitrogen transformations in cold and frozen agricultural soils following organic amendments

Though microbial activity is known to occur in frozen soils, little is known about the fate of animal manure N applied in the fall to agricultural soils located in areas with prolonged winter periods. Our objective was to examine transformations of soil and pig slurry N at low temperatures. Loamy and clay soils were either unamended (Control), amended with ¹⁵NH₄-labeled pig slurry, or amended with the pig slurry and wheat straw. Soils were incubated at −6, −2, 2, 6, and 10 °C. The amounts of NH₄, NO₃ and microbial biomass N (MBN), and the presence of ¹⁵N in these pools were monitored. Total mineral N, NO₃ and ¹⁵NO₃ increased at temperature down to −2 °C in the loam soil and −6 °C in the clay soil, indicating that nitrification and mineralization proceeded in frozen soils. Nitrification and mineralization rates were 1.8-4.9 times higher in the clay than in the loamy soil, especially below freezing point (3.2-4.9), possibly because more unfrozen water remained in the clay than in the loamy soil. Slurry addition increased nitrification rates by 3-14 times at all temperatures, indicating that this process was N-limited even in frozen soils. Straw incorporation caused significant net N immobilization only at temperatures ≥2 °C in both soils; the rates were 1.4-3.4 higher in the loam than in the clay soil. Nevertheless, up to 30% of the applied ¹⁵N was present in MBN at all temperatures. These findings indicate that microbial N immobilization occurred in frozen soils, but was not strong enough to induce net immobilization below the freezing point, even in the presence of straw. The Q₁₀ values for estimated mineralization and nitrification rates were one to two orders-of-magnitude larger below 2 °C than above this temperature (13-208 versus 1.5-6.9, respectively), indicating that these processes are highly sensitive to a small increase in soil temperature around the freezing point of water. This study confirms that net mineralization and nitrification can occur at potentially significant rates in frozen agricultural soils, especially in the presence of organic amendments. In contrast, net N immobilization could be detected essentially above the freezing point. Our results imply that fall-applied N could be at risk of overwinter losses, particularly in fine-textured soils.     

Differences in isotopic fractionation of nitrogen in water-saturated and unsaturated soils

Temporal variations in δ¹⁵N of NH₄⁺ and NO₃⁻ in water-saturated and unsaturated soils were examined in a laboratory incubation study. Ammonium sulfate (δ¹⁵N=−2.6‰) was added to 25 g samples of soil at concentrations of 160 mg N kg⁻¹. Soils were then incubated under unsaturated (50% of water holding capacity at saturation, WHC) or saturated (100% of WHC) water conditions for 7 and 36 d, respectively. During 7 d incubation of unsaturated soil, the NH₄⁺-N concentration decreased from 164.8 to 34.4 mg kg⁻¹, and the δ¹⁵N of NH₄⁺ increased from −0.4 to +57.2‰ through nitrification, as evidenced by corresponding increase in NO₃⁻-N concentration and lower δ¹⁵N of NO₃⁻ (product) than that of NH₄⁺ (substrate) at each sampling time. In saturated soil, the concentration of NH₄⁺-N decreased gradually from 162.4 to 24.2 mg kg⁻¹, and the δ¹⁵N values increased from +0.8 to +21.0‰ during 36 d incubation. However, increase in NO₃⁻ concentration was not observed due to loss of NO₃⁻ through concurrent denitrification in anaerobic sites. The apparent isotopic fractionation factors (α_s/p) associated with decrease in NH₄⁺ concentration were 1.04 and 1.01 in unsaturated and saturated soils, respectively. Since nitrification is likely to introduce greater isotope fractionation than microbial immobilization, the higher value for unsaturated soil probably reflected faster nitrification under aerobic conditions. The lower value for saturated soil suggests that immobilization and subsequent remineralization of NH₄⁺ were relatively more dominant than nitrification under the anaerobic conditions.     

Modelling forest carbon balances considering tree mortality and removal

The determination of ecosystem carbon balances is a major issue in environmental research. Forest inventories and - more recently - Eddy covariance measurements have been set up to guide sustainability assessments as well as carbon accounting. A differentiation between ecosystem compartments of carbon such as soil and vegetation, or above- and belowground storages nevertheless requires further empirical assumptions or model simulations. However, models to estimate carbon balances often do not account for carbon export by management and the direct and indirect impacts of forest management. To overcome this obstacle, we complemented a physiologically based process model (MoBiLE-PSIM) with routines for dimensional tree growth and mortality and evaluated the full model with measurements of water availability, primary production, respiration fluxes and forest development (tree dimensions and numbers per hectare).The model is applied to three forests representing different physiological types and climatic environments: Norway spruce, European beech and Mediterranean holm oak. Simulated carbon balances are presented on a daily, annual and decadal time scale throughout the years 1998-2008 for all three stands. On average, gross primary production is 2.0, 1.7, and 1.4 and net ecosystem production 0.6, 0.6, and 0.3 kg C m⁻² a⁻¹. Export of carbon by thinning is highest in the middle-aged beech stand (0.24 kg C m⁻² a⁻¹) which decreases net ecosystem production by 15% compared with an unthinned stand. Between 46 (spruce) and 72 (oak) % of carbon gained by net ecosystem production is sequestered below ground (incl. roots) - a share that is decreased if a part of the carbon is exported as timber. The role of further impacts, in particular carry-over effects in years that follow intense drought periods, is highlighted and the usefulness of the approach for highly resolved environmental change studies is discussed.     

CH4 oxidation and emissions of CH4 and N2O from Lolium perenne swards under elevated atmospheric CO2

Emissions of N₂O and CH₄ and CH₄ oxidation rates were measured from Lolium perenne swards in a short-term study under ambient (36 Pa) and elevated (60 Pa) atmospheric CO₂ at the Free Air Carbon dioxide Enrichment experiment, Eschikon, Switzerland. Elevated pCO₂ increased (P<0.05) N₂O emissions from high N fertilised (11.2 g N m⁻²) swards by 69%, but had no significant effect on net emissions of CH₄. Application of ¹³C-CH₄ (11 μl l⁻¹; 11 at.% excess ¹³C) to closed chamber headspaces in microplots enabled determination of rates of ¹³C-CH₄ oxidation even when net CH₄ fluxes from main plots were positive. We found a significant interaction between fertiliser application rate and atmospheric pCO₂ on ¹³C-CH₄ oxidation rates that was attributed to differences in gross nitrification rates and C and N availability. CH₄ oxidation was slower and thought to be temporarily inhibited in the high N ambient pCO₂ sward. The most rapid CH₄ oxidation of 14.6 μg ¹³C-CH₄ m⁻² h⁻¹ was measured in the high fertilised elevated pCO₂ sward, and we concluded that either elevated pCO₂ had a stimulatory effect on CH₄ oxidation or inhibition of oxidation following fertiliser application was lowered under elevated pCO₂. Application of ¹⁴NH₄¹⁵NO₃ and ¹⁵NH₄¹⁵NO₃ (10 at.% excess ¹⁵N) to different replicates enabled determination of the respective contributions of nitrification and denitrification to N₂O emissions. Inhibition of CH₄ oxidation in the high fertilised ambient pCO₂ sward, due to competition between NH₃ and CH₄ for methane monooxygenase enzymes or toxic effects of NH₂OH or NO₂⁻ produced during nitrification, was hypothesised to increase gross nitrification (12.0 mg N kg dry soil⁻¹) and N₂O emissions during nitrification (327 mg ¹⁵N-N₂O m⁻² over 11 d). Our results indicate that increasing atmospheric concentrations of CO₂ may increase emissions of N₂O by denitrification, lower nitrification rates and either increase or decrease the ability of soil to act as a sink for atmospheric CH₄ depending on fertiliser management.     

Evidence of carbon stimulated N transformations in grassland soil after slurry application

High nitrification rates which convert ammonium (NH₄⁺) to the mobile ions NO₂⁻ and NO₃⁻ are of high ecological significance because they increase the potential for N losses via leaching and denitrification. Nitrification can be performed by chemoautotrophic or heterotrophic organisms and heterotrophic nitrifiers can oxidise either mineral (NH₄⁺) or organic N. Selective nitrification inhibitors and ¹⁵N tracer studies have been used in an attempt to separate heterotrophic and autotrophic nitrification. In a laboratory study we determined the effect of cattle slurry on the oxidation of mineral NH₄⁺-N and organic-N by labelling the NH₄⁺ or NO₃⁻ pools separately or both together with ¹⁵N. The size and enrichment of the mineral N pools were determined at intervals. To calculate gross N transformation rates a ¹⁵N tracing model was developed. This model consists of the three N-pools NH₄⁺, NO₃⁻ and organic N. Sub-models for decomposition of degradable carbon in the soil and the slurry were added to the model and linked to the N transformation rates. The model was set up in the software ModelMaker which contains non-linear optimization routines to determine model parameters. The application of cattle slurry increased the rate of nitrifcation by a factor of 20 compared with the control. The size and enrichment of the mineral N pools provided evidence that nitrification was due to the conversion of NH₄⁺ to NO₃⁻ and not the conversion of organic N to NO₃⁻. There was evidence that slurry-enhanced oxidation of NH₄⁺ to NO₃⁻ was due to a combination of autotrophic and heterotrophic transformations. Slurry application increased the mineralisation rate by approximately a factor of two compared with the control and the rate of immobilisation of NH₄⁺ by approximately a factor of three.     

Microbial immobilization of ammonium and nitrate in relation to ammonification and nitrification rates in organic and conventional cropping systems

Agricultural systems that receive high or low organic matter (OM) inputs would be expected to differ in soil nitrogen (N) transformation rates and fates of ammonium (NH₄⁺) and nitrate (NO₃⁻). To compare NH₄⁺ availability, competition between nitrifiers and heterotrophic microorganisms for NH₄⁺, and microbial NO₃⁻ assimilation in an organic vs. a conventional irrigated cropping system in the California Central Valley, chemical and biological soil assays, ¹⁵N isotope pool dilution and ¹⁵N tracer techniques were used. Potentially mineralizable N (PMN) and hot minus cold KCl-extracted NH₄⁺ as indicators of soil N supplying capacity were measured five times during the tomato growing season. At mid-season, rates of gross ammonification and gross nitrification after rewetting dry soil were measured in microcosms. Microbial immobilization of NO₃⁻ and NH₄⁺ was estimated based on the uptake of ¹⁵N and gross consumption rates. Gross ammonification, PMN, and hot minus cold KCl-extracted NH₄⁺ were approximately twice as high in the organically than the conventionally managed soil. Net estimated microbial NO₃⁻ assimilation rates were between 32 and 35% of gross nitrification rates in the conventional and between 37 and 46% in the organic system. In both soils, microbes assimilated more NO₃⁻ than NH₄⁺. Heterotrophic microbes assimilated less NH₄⁺ than NO₃⁻ probably because NH₄⁺ concentrations were low and competition by nitrifiers was apparently strong. The high OM input organic system released NH₄⁺ in a gradual manner and, compared to the low OM input conventional system, supported a more active microbial biomass with greater N demand that was met mainly by NO₃⁻ immobilization.     

Effects of repeated fertilizer and cattle slurry applications over 38 years on N dynamics in a temperate grassland soil

The effects of repeated synthetic fertilizer or cattle slurry applications at annual rates of 50, 100 or 200 m³ ha⁻¹ yr⁻¹ over a 38 year period were investigated with respect to herbage yield, N uptake and gross soil N dynamics at a permanent grassland site. While synthetic fertilizer had a sustained and constant effect on herbage yield and N uptake, increasing cattle slurry application rates increased the herbage yield and N uptake linearly over the entire observation period. Cattle slurry applications, two and four times the recommended rate (50 m³ ha⁻¹ yr⁻¹, 170 kg N ha⁻¹), increased N uptake by 46 and 78%, respectively after 38 years. To explain the long-term effect, a ¹⁵N tracing study was carried out to identify the potential change in N dynamics under the various treatments. The analysis model evaluated process-specific rates, such as mineralization, from two organic-N pools, as well as nitrification from NH₄⁺ and organic-N oxidation. Total mineralization was similar in all treatments. However, while in an unfertilized control treatment more than 90% of NH₄⁺ production was related to mineralization of recalcitrant organic-N, a shift occurred toward a predominance of mineralization from labile organic-N in the cattle slurry treatments and this proportion increased with the increase in slurry application rate. Furthermore, the oxidation of recalcitrant organic-N shifted from a predominant NH₄⁺ production in the control treatment, toward a predominant NO₃⁻ production (heterotrophic nitrification) in the cattle slurry treatments. The concomitant increase in heterotrophic nitrification and NH₄⁺ oxidation with increasing cattle slurry application rate was mainly responsible for the increase in net NO₃⁻ production rate. Thus the increase in N uptake and herbage yield on the cattle slurry treatments could be related to NO₃⁻ rather than NH₄⁺ production. The ¹⁵N tracing study was successful in revealing process-specific changes in the N cycle in relationship to long-term repeated amendments.     

Potential importance of bacteria and fungi in nitrate assimilation in soil

Soil microorganisms can use a wide range of N compounds but are thought to prefer NH₄⁺. Nevertheless, ¹⁵N isotope dilution studies have shown that microbial immobilization of NO₃⁻ can be an important process in many soils, particularly relatively undisturbed soils. Our objective was to develop a method for measuring NO₃⁻ immobilization potential so that the relative contributions of bacteria and fungi could be determined. We modified and optimized a soil slurry method that included amendments of KNO₃, glucose, and methionine sulfoximine (an inhibitor of N assimilation) in the presence of two protein synthesis inhibitors: chloramphenicol, which inhibits bacteria, or cycloheximide, which inhibits fungi. By adding ¹⁵N-labeled KNO₃, we were able to measure gross rates of NO₃⁻ production (i.e., gross nitrification) and consumption (i.e., gross NO₃⁻ immobilization). We found that bacteria, not fungi, had the greatest potential for assimilating, or immobilizing, NO₃⁻ in these soils. This is consistent with their growth habit and distribution in the heterogeneous soil matrix.