Substantial net N mineralization during the dormant season in temperate forest soils

In temperate forest soils, N net mineralization has been extensively investigated during the growing season, whereas N cycling during winter was barely addressed. Here, we quantified net ammonification and nitrification during the dormant season by in situ and laboratory incubations in soils of a temperate European beech and a Norway spruce forest. Further, we compared temperature dependency of N net mineralization in in situ field incubations with those from laboratory incubations at controlled temperatures. From November to April, in situ N net mineralization of the organic and upper mineral horizons amounted to 10.9 kg N (ha · 6 months)^–1 in the spruce soil and to 44.3 kg N (ha · 6 months)^–1 in the beech soil, representing 65% (beech) and 26% (spruce) of the annual above ground litterfall. N net mineralization was largest in the Oi/Oe horizon and lowest in the A and EA horizons. Net nitrification in the beech soil [1.5 kg N (ha · 6 months)^–1] was less than in the spruce soil [5.9 kg N (ha · 6 months)^–1]. In the range of soil temperatures observed in the field (0–8°C), the temperature dependency of N net mineralization was generally high for both soils and more pronounced in the laboratory incubations than in the in situ incubations. We suggest that homogenization of laboratory samples increased substrate availability and, thus, enhanced the temperature response of N net mineralization. In temperate forest soils, N net mineralization during the dormant season contributes substantially to the annual N cycling, especially in deciduous sites with large amounts of litterfall immediately before the dormant season. High Q₁₀ values of N net mineralization at low temperatures suggest a huge effect of future increasing winter temperature on the N cycle in temperate forests.     

Potential net soil N mineralization and decomposition of glycine-C in forest soils along an elevation gradient

The objective of this research was to better understand patterns of soil nitrogen (N) availability and soil organic matter (SOM) decomposition in forest soils across an elevation gradient (235-1670 m) in the southern Appalachian Mountains. Laboratory studies were used to determine the potential rate of net soil N mineralization and in situ studies of ¹³C-labelled glycine were used to infer differences in decomposition rates. Nitrogen stocks, surface soil (0-5 cm) N concentrations, and the pool of potentially mineralizable surface soil N tended to increase from low to high elevations. Rates of potential net soil N mineralization were not significantly correlated with elevation. Increasing soil N availability with elevation is primarily due to greater soil N stocks and lower substrate C-to-N ratios, rather than differences in potential net soil N mineralization rates. The loss rate of ¹³C from labelled soils (0-20 cm) was inversely related to study site elevation (r=−0.85; P<0.05) and directly related to mean annual temperature (+0.86; P<0.05). The results indicated different patterns of potential net soil N mineralization and ¹³C loss along the elevation gradient. The different patterns can be explained within a framework of climate, substrate chemistry, and coupled soil C and N stocks. Although less SOM decomposition is indicated at cool, high-elevation sites, low substrate C-to-N ratios in these N-rich systems result in more N release (N mineralization) for each unit of C converted to CO₂ by soil microorganisms.     

Fate of inorganic 15N in the profile of different coniferous forest soils

The fate of inorganic ¹⁵N added to different coniferous forest soils was traced throughout the soil profile (0–25 cm) in a laboratory experiment under controlled conditions of temperature and water content. Six soils with different chemical climates were compared. The sequestration of labelled N was significantly explained by the clay content but the correlation was improved when C and N content were included. The level of acidification, even in soil with a fine texture, reduced the immobilization. For a similar N input, sandy soils with low C content or high acidification showed a reduced N storage capacity, so that N excess would be able to pollute the ground-water.     

Bacterial and fungal response to nitrogen fertilization in three coniferous forest soils

Forest soil carbon (C) pools may act as sinks for, or sources of, atmospheric carbon dioxide, while nitrogen (N) fertilization may affect the net exchange of C in forest ecosystems. Since all major C and N processes in soil are driven by soil microorganisms, we evaluated the effects of N fertilization on biomass and bacterial and fungal activity in soils from three Norway spruce forests with different climatic and N availability conditions. N deposition and net N mineralization were higher at the sites in southern Sweden than at the site in northern Sweden. We also studied the extent to which N fertilization altered the nutrient(s) limiting bacterial growth in soil. We found that on average microbial biomass was reduced by ～40% and microbial activity by ～30% in fertilized plots. Bacterial growth rates were more negatively affected by fertilization than fungal growth rates, while fungal biomass (estimated using the phospholipid fatty acid (PLFA) 18:2ω6,9) decreased more than bacterial biomass as a consequence of fertilization. The microbial community structure (indicated by the PLFA pattern) was changed by fertilization, but not in the same way at the three sites. Soil bacteria were limited by a lack of carbon in all forests, with the carbon limitation becoming more evident in fertilized plots, especially in the forests that had previously been the most N-limited ones. This study thus showed that the effects of N fertilization differed depending on the conditions at the site prior to fertilization.     

Changes in net N mineralization rates and soil N and P pools in a pine forest wildfire chronosequence

The concern that climate change may increase fire frequency and intensity has recently heightened the interest in the effects of wildfires on ecosystem functioning. Although short-term fire effects on forest soils are well known, less information can be found on the long-term effects of wildfires on soil fertility. Our objective was to study the 17-year effect of wildfires on forest net mineralization rates and extractable inorganic nitrogen (N) and phosphorus (P) concentrations. We hypothesize that (1) burned forest stands should exhibit lower net mineralization rates than unburned ones; (2) these differences would be greatest during the growing season; (3) differences between soil variables might also be observed among plots from different years since the last fire; and (4) due to fire-resistant geochemical processes controlling P availability, this nutrient should recover faster than N. We used a wildfire chronosequence of natural and unmanaged Pinus canariensis forests in La Palma Island (Canary Islands). Soil samples were collected during winter and spring at 22 burned and unburned plots. We found significantly higher values for net N mineralization and extractable N pools in unburned plots. These differences were higher for the winter sampling date than for the spring sampling date. Unlike extractable N and N mineralization rates, extractable P levels of burned plots exhibited a gradual recovery over time after an initial decrease. These results demonstrate that P. canariensis forest soils showed low resilience after wildfires, especially for N, and that this disturbance might induce long-term changes in ecosystem functioning.     

Can δ15N profiles in forest soils predict {\rm{NO}}_3^ - loss and net N mineralization rates?

This paper studies changes of ¹⁵N signatures ('¹⁵N, ‰) and total N (TN, %) in soil profiles among forest stands with different NO₃^- {\rm NO}_3^ - losses within the same climatic zone. An additional aim was to investigate whether the change of '¹⁵N (('¹⁵N) within the 0-10, 10-20 and 20-30 cm depths of the mineral layer could be linked to measured potential net N mineralization rates. Soil samples were collected from five forest stands in Belgium: three mixed deciduous forests (G, AE, LD), a homogenous deciduous (SB) and a coniferous stand (CP). At the G site, five locations were sampled: one at the forest edge (GE), two deeper in the forest (GF1, GF2), one clear-cut spot (GO) and one in coppice wood (GC). The '¹⁵N and TN measurements were conducted for the litter layer, the fermentation + humus layer and the underlying mineral layers (0-30 cm, at 2 cm intervals). The '¹⁵N values increased with depth, ranging from -12‰ to -1‰ for the forest floor and from -7‰ to +15‰ for the mineral layers. The overall enrichment factor was greater for locations GE, AE and SB (-5.2‰, P <0.001, R² =0.86) than locations GF1, GF2, GO, GC, LD and CP (-2.4‰, P <0.001, R² =0.93), possibly indicating NO₃^- losses. A significant linear regression model could be calculated between ('¹⁵N and potential net N mineralization rates (y =0.04x), explaining 65% of the variability of ('¹⁵N. Thus, '¹⁵N profiles in forest soils might be useful as an indicator of NO₃^- {\rm NO}_3^ - loss and N mineralization behaviour, however, further research is needed to confirm our observations.     

Carbon and nitrogen mineralization in subarctic agricultural and forest soils

Summary C and N mineralization potentials were determined, in a 12-week laboratory incubation study, on soil samples obtained from recently cleared land which had been cropped to barley for 4 years (field soils) and from nearby undisturbed taiga (forest soils). Treatments for the cropped soils were conventional and no-tillage with and without crop residues removed. An average of about 3% of the total C was evolved as CO₂ from the field soils compared with > 10% and 4% for the upper (Oie) and lower (Oa) forest-floor horizons, respectively. Significantly more C was mineralized from the Ap of the no-till treatment with residue left on the surface than from the other field Ap horizons. Both forest-floor horizons showed rather long lag periods for net mineralization compared with the field soils, but at the end of the incubation, more mineral N was recovered from the forest Oie despite a rather wide C:N ratio, than from the field soils. After 12 weeks about 115, 200 and 20 g mineral N/g soil were recovered from the field Ap, the forest Oie and the forest Oa horizons, respectively. Very little C or N was mineralized from the B horizon of the forest or the field soils. Nitrification was rapid and virtually complete for the field soils but was negligible for both forest-floor O horizons.Paper no J-188 of the Journal Series of the Alaska Agricultural and Forestry Experiment Station     

Soil layer-specific variability in net nitrification and denitrification in an acid coniferous forest

 High spatial variation in nitrification potentials has been observed in forest soils, but explanations for this variability have remained speculative. In the present study we determined whether sample treatment, sample size, denitrification or small-scale variations in abiotic properties could explain spatial variation in nitrogen transformations in the organic horizon of a pine forest soil. Net nitrate production in homogenates of the organic horizon was extremely variable. Sample size (60–600 cm²) had no significant effect on nitrate production. In homogenised samples no increased nitrogen production was observed compared to intact incubated cores. High small-scale variation in nitrate production was observed in the litter (L) horizon. When this stratified L layer was subdivided, high net nitrate production was observed in moss (LM) and fragmented needles, whereas no net nitrate production was found in intact needles. The addition of acetylene, inhibiting nitrous oxide reductase, led to significant nitrous oxide production in the L layer. Low nitrous oxide production was found in the LM layer and none in the fragmentation layer. These results show that denitrification can explain part of the spatial variation and plays a major role in nitrogen transformations in the L layer. The relatively higher pH and the presence of fungi are suggested as factors responsible for high denitrification rates in the L layer. As a consequence homogenisation of the organic horizon could lead to highly variable nitrate production due to denitrifying activity from the needles being introduced into other layers. Received: 10 December 1999     

Bulk soil C to N ratio as a simple measure of net N mineralization from stabilized soil organic matter in sandy arable soils

Investigations of 23 northwestern German sandy Ap horizons (mean clay content 35 g kg⁻¹), that had higher organic matter (OM) levels than expected for sands, showed that the bulk soil C to N ratio reliably indicated the release of N from stabilized OM. Soils were incubated at 35 °C for 200 days under aerobic conditions. Cumulative N release curves were split into N released from fresh materials (N_fast) and N released from the larger pool of stabilized, older OM (N_slow rates, 0.06-0.77 μg N g⁻¹ soil d⁻¹, or 0.7-49 μg N g⁻¹ OM). Correlating the N_slow rates with total N contents of soils yielded no satisfactory relationships while their relationship with C to N ratios was very close (negative exponential, R²=0.88). Low rates of N release (N_slow) per unit of OM occurred if C to N exceeded 15. This was associated with historical factors like podzolization, calluna heathland, plaggen fertilization or a combination of these.     

Differential response of ectomycorrhizal and saprotrophic fungal mycelium from coniferous forest soils to selected monoterpenes

The mycelia of saprotrophic (SP) and ectomycorrhizal (ECM) fungi occur throughout the upper soil horizons in coniferous forests and could therefore be exposed to high concentrations of monoterpenes occurring in the needle litter of some tree species.Monoterpenes are mycotoxic and could potentially affect fungi that are exposed to them in the litter layers. In order to investigate whether monoterpenes typical of coniferous litters could influence fungal communities, we analysed the monoterpene content of freshly fallen needles of Pinus sylvestris, Picea abies and Picea sitchensis. The most abundant monoterpenes were found to be α-pinene, β-pinene and 3-carene. We evaluated the effects of these three monoterpene vapours on the biomass production of 23 SP isolates and 16 ECM isolates. Overall, 75% of ECM isolates and 26% of SP isolates were significantly inhibited by at least one of the monoterpene treatments and both intra- and inter-specific variations in response were observed.Monoterpene concentrations are highest in surface litters. The differential effects on fungal taxa may influence the spatial and temporal distribution of fungal community composition, indirectly affecting decomposition and nutrient cycling, the fundamental ecosystem processes in which fungi have a key role in coniferous forest soils.     

Clear-cutting affects the ammonia-oxidising community differently in limed and non-limed coniferous forest soils

The effects of clear-cutting on the ammonia-oxidising bacterial community were studied in the soil of limed and non-limed spruce forest plots located in the central part of southern Sweden. The communities were studied using denaturing gradient gel electrophoresis (DGGE) profiling after polymerase chain reaction (PCR) amplification from total DNA with primers reported to be specific for -subgroup ammonia-oxidising bacteria. The bands on the DGGE were sequenced and each unique sequence was interpreted as representing one ammonia-oxidising population. The relative abundance of each population was determined by measuring the fluorescence of the respective DGGE bands. In both limed and non-limed soil, the same two Nitrosospira populations were found, one belonging to cluster 2 (NScl2) and one to cluster 4 (NScl4). However, while NScl4 first appeared a year after the clear-cutting in the non-limed plot, it was present both before and after the cutting in the limed plot. Irrespective of previous liming, clear-cutting caused a shift in the ammonia-oxidiser community, from dominance by the NScl2 population to a community with approximately equal relative abundance of NScl2 and NScl4. In both plots the total size of the community increased after clear-cutting (based on increased DGGE band intensity), most likely due to increased NH₄⁺ availability, but the growth response was faster in the limed plot. Hence, the prior liming increased the responsiveness of the ammonia-oxidisers to the changes caused by cutting. This is the first study to report the effects of clear-cutting on the ammonia-oxidising community, and the results show a clear correlation between increased potential nitrification and a shift in the ammonia-oxidiser community.     

Experimental snowpack reduction alters organic matter and net N mineralization potential of soil macroaggregates in a northern hardwood forest

Climate change is predicted to reduce or delay annual wintertime snow pack formation in the forests of the northeastern US. Any delay in snowpack formation could increase soil freezing in winter and, thereby, alter soil characteristics and processes. We examined the hypothesis that delayed snowpack would disrupt soil structure and change organic matter bioavailability in an experimental snow removal study at the Hubbard Brook Experimental Forest (HBEF), NH, USA. Pairs of reference and snow removal treatment plots were studied in four different sites at HBEF. Snow was removed from November–January of two winters, inducing soil freezing throughout both winters. Size class distribution and organic matter concentration and content of aggregates, and carbon and nitrogen mineralization potential of size fractions were quantified for surface mineral soils in the spring of both years immediately after snowmelt. In the first year of sampling, the only significant effect of snow removal was an increase in the smallest (<53 μm) size fraction of mineral soil. In the second year, snow removal increased organic matter concentrations of macroaggregate (250–2,000 μm) and microaggregate (53–250 μm) size fractions. This change corresponded to an increase in net N mineralization potential and the ratio of N to C mineralized in the macroaggregate fraction, but there were no effects of snow removal on C mineralization. We propose that soil freezing increases the movement of organic matter from organic to mineral soil horizons and increases the N content of mineralizable substrates in mineral soil following years with delayed snowpack formation.     

The relationship between N mineralization or microbial biomass N with micromorphological properties in beech forest soils with different texture and pH

To test relationships between net N-mineralization, organic matter and soil organisms, we combined micromorphology with laboratory incubation experiments over a soil gradient. Microbial biomass N generally increased with pH, and from sandy to loamy soil, but net N-mineralization showed the opposite, and was highest in acid, sandy soil. Twenty-two micromorphological characteristics were analyzed with principal component analysis. PC1 had high eigenvalue (0.70), and clearly separated fungi from earthworms, microarthropods and bacteria. PC2 was less important (0.15). Organic layer and sand content clearly correlated with the fungi-end of PC1, but pH and C-content of the Ah with the opposite. Microbial N also correlated with the earthworm–bacteria end, but net N-mineralization did not. Efficiency of N-mineralization per unit microbe even correlated with the fungi end of PC1, in both organic layer and mineral topsoil. The results support the hypothesis that high (or low) litter turnover and biological activity can be counteracted by high (or low) microbial N-demand.     

Effects of acidification and liming on feeding groups of nematodes in coniferous forest soils

Summary Nematodes were sampled in untreated, acidified, and limed plots in a Norway spruce (Fexboda) and a Scots pine (Norrliden) stand. At Fexboda, the total number of nematodes was significantly reduced after the acidification. This reduction was probably due to a shock effect, because the samples were taken only 5 months after an application of 200 kg H₂SO₄ ha^-1 to the forest floor. However, the root/fungal-feeding Aphelenchoides was not reduced, probably because it is more tolerant of high acid concentrations than most other nematodes. At Norrliden, where the samples were taken 7 years after the last application of H₂SO₄, no significant differences were found between the acidified and untreated plots. If the treatment with H₂SO₄ caused similar effects as at Fexboda, the results indicate a recovery of the nematode populations. Decreased predation from lumbricids rather than a recovery of microfloral populations probably allowed this recovery. No marked effect of lime, spread 2 (Fexboda) and 12 years (Norrliden) before the sampling on the numbers of any of the nematode feeding groups was found. This correlated with almost no change in bacterial biomass after liming, while the active fraction of fungal hyphae was unaffected by liming at Fexboda and reduced by liming at Norrliden. A tendency for decreasing numbers of all nematode feeding groups in the limed plots at Norrliden coincided with increasing numbers of lumbricids.     

Effects of plants on net mineralization of nitrogen in forest soil microcosms

Summary The effects of plant roots on net N mineralization were examined by comparing soil microcosms with and without plants. Additionally, inorganic N amendments were used to test for competition for N between plants and microorganisms. Daily watering and the application of suction to microcosms eliminated the effects of transpiration on soil moisture content. Monthly litter collections reduced the influence of the aboveground portions of plants. Plants decreased net N mineralization by 23% during days 0–114 and then increased net mineralization by the same amount during days 144–124. Root-free soil collected from with-plant microcosms on day 244 evolved 24% more CO₂ in laboratory incubations than soil from without-plant microcosms. This indicates that plants had increased substrate availability to soil microorganisms. Inorganic N amendments had no significant effects on the microcosms or on laboratory soil incubations. Evidence is most consistent with the hypothesis that plant roots increased microbial activity due to the increased substrate availability. Different net N mineralization rates probably resulted from changes in the substrate C : N ratio.     

Factors controlling nitrogen mineralization and nitrification in forest ecosystems in New Mexico

Summary Forest floor and mineral soil from ponderosa-pine, Douglas-fir, aspen and spruce-fir ecosystems located along a rising gradient in New Mexico were tested with laboratory assays for factors controlling N mineralization and nitrification. We concluded that low pH in combination with factors associated with organic quality controlled N mineralization and almost completely limited nitrification in spruce-fir soils, while N mineralization in the forest floor of ponderosa-pine was limited by low nutrient availability (other than N). Organic quality of the substrate and temporal changes in organic quality appeared to control N-mineralization and nitrification processes in forest-floor and mineral soils from all other sites.     

Litter decomposition reduces either N2O or NO production in strongly acidic coniferous and broad-leaved forest soils under anaerobic conditions

Purpose

The beneficial effect to the environment of nitrate (NO₃ ^?) removal by denitrification depends on the partitioning of its end products into nitrous oxide (N₂O), nitric oxide (NO), and dinitrogen (N₂). However, in subtropical China, acidic forest mineral soils are characterized by negligible denitrification capacity and thus reactive forms of N could not be effectively converted to inert N₂, resulting in a negative environmental consequence. In this study, the influences of C input from litter decomposition on denitrification rate and its gaseous products under anoxic conditions in the acidic coniferous and broad-leaved forest soils in subtropical China were investigated using the acetylene (C₂H₂) blockage technique in the laboratory.

Materials and methods

The coniferous and broad-leaved forest soils with and without litter addition were incubated under anaerobic conditions for 244 h. There were three treatments for each forest soil including addition of 0.5 and 1% corresponding litter (gram of litter per gram of soil) and the control without addition of litter.

Results and discussion

The results showed that litter addition into the broad-leaved forest soil had no effect on average rates of denitrification (calculated as the sum of NO, N₂O, and N₂), whereas in the coniferous forest soil, the addition resulted in a significant increase in average denitrification rate. In the broad-leaved forest soil, both rates of litter addition decreased the production of NO but increased the production of N₂, and high rates of litter addition into the coniferous forest soil promoted the reduction of N₂O to N₂.

Conclusions

Increased decomposition of litter in the forest soils could effectively reduce N₂O and NO production through denitrification under anaerobic conditions.     

Microbial biomass and enzyme activities in coniferous forest soils as affected by lignite-derived deposition

A forest ecosystem study was conducted along a deposition gradient of air pollutants in old Scots pine stands located near the industrial belt around the city of Bitterfeld in northeast Germany from 1999 to 2000. In order to estimate the impact of different atmospheric deposition loads on microbial biomass and enzyme activities, samples were taken from the forest floor (L, F, and H horizon) and the mineral topsoil (0–10 cm). The emission-induced increases in ferromagnetic susceptibility, soil pH, concentrations of mobile (NH ₄NO ₃ extractable) Cr and Ni, effective cation exchange capacity, and base saturation in the humus layer along the 25-km long transect reflected that great portions of the past depositions were characterized by alkaline fly ash. Alkaline depositions significantly ( P <0.05) decreased the microbial biomass C and N contents, microbial biomass C-to-organic C ratios, and microbial respiration rates, but increased the metabolic quotient (qCO ₂) of the mineral topsoil and forest floor. Variations in microbial biomass and activity can mainly be predicted ( r ² =0.60) by the concentrations of Ca, Zn and Cd in these forest soils. The specific activities (activity kg ^-1 organic C) of l-asparaginase, l-glutaminase, arylsulfatase, and in part, acid phosphatase were significantly ( P <0.05) higher at forest sites receiving higher fly ash loads than those of the other sites, and thus followed the trend of the qCO ₂. In contrast, the specific activity of ß-glucosidase was significantly ( P <0.05) decreased at heavily affected sites compared to moderate and less affected sites, suggesting an inhibition of C mineralization in the forest floor of pine stands affected by predominantly alkaline emissions. A great portion ( r ²=0.91) of the variation in the specific enzyme activity data in forest soils in emission areas can be predicted from a linear combination of the variables total organic C and NH ₄Cl-extractable Ca, pH and effective cation exchange capacity.     

Effect of simulated acid precipitation on nitrogen mineralization and nitrification in forest soils

After exposure of samples of three forest soils (pH 3.4 to 3.9) from the Adirondacks region of New York to 60, 230, or 400 cm of simulated rain of pH 3.5 or 5.6 in 4, 14, or 24 weeks, respectively, the soil samples were separated into the 0 to 2 and 2 to 5 cm organic layers and further incubated. The rates of N mineralization in Woods soil exposed to the simulated precipitation were less for rain at pH 3.5 than at pH 5.6, but the inhibition decreased with increasing exposure of the 0 to 2 cm layer. In Panther soil, the rates of mineralization were usually not affected by the acidity of the simulated rain. In the upper layer of Sagamore soil, mineralization was not influenced by pH of the simulated rain, but the transformation was faster in the bottom layer of soil after prolonged exposure to simulated rain at pH 3.5 than at pH 5.6. The rate of nitrate formation in Panther and Woods soil amended with ammonium was inhibited by the more acid rain. Studies with ¹⁵NH₄ indicated that ammonium was oxidized to nitrate even though ammonium levels did not decline or declined only slightly after prolonged exposure of Panther or Woods soil to rain at pH 3.5. The growth of orchardgrass in Panther and Woods soil was inhibited by the more acid simulated rain.