Microbial immobilization and mineralization of dissolved organic nitrogen from forest floors

Dissolved organic nitrogen (DON) plays a key role in the N cycle of many ecosystems, as DON availability and biodegradation are important for plant growth, microbial metabolism and N transport in soils. However, biodegradation of DON (defined as the sum of mineralization and microbial immobilization) is only poorly understood. In laboratory incubations, biodegradation of DON and dissolved organic carbon (DOC) from Oi and Oa horizons of spruce, beech and cypress forests ranged from 6 to 72%. Biodegradation of DON and DOC was similar in most samples, and mineralization of DON was more important than microbial immobilization. Nitrate additions (0-10 mg N L⁻¹) never influenced either DON immobilization by microorganisms or mineralization. We conclude that soil microorganisms do not necessarily prefer mineral N over DON for meeting their N demand, and that biodegradation of DON seems to be driven by the microbial demand for C rather than N. Quantifying the dynamics of DON in soils should include consideration of both C and N demands by microbes.     

Nitrogen Saturation and Soil N Availability in a High-Elevation Spruce and Fir Forest

A field study was conducted during the summer of 1995 to gain abetter understanding of the causes of nitrate (NO₃-N)leaching and ongoing changes in soil nitrogen (N) availabilityin high-elevation (1524–2000 m) spruce (Picea rubens) andfir (Abies fraseri) forests of the Great Smoky MountainsNational Park, Tennessee and North Carolina, U.S.A. Indicatorsof soil N availability (total soil N concentrations,extractable NH₄-N, extractable NO₃-N, and C/N ratios)were measured in Oa and A horizons at 33 study plots. Dynamicmeasures included potential net soil N mineralization determinedin 12-week aerobic laboratory incubations at 22 °C.Potential net nitrification in the A horizon was correlated (r =+0.83, P < 0.001) with total soil N concentrations. Mostmeasures of soil N availability did not exhibit significanttrends with elevation, but there were topographic differences.Potential net soil N mineralization and net nitrification in theA horizon were higher in coves than on ridges. Relative amountsof particulate and organomineral soil organic matter influencedpotential net N mineralization and nitrification in the Ahorizon. Calculations indicate that soil N availability andNO₃-N leaching in high-elevation spruce and fir forests ofthe Great Smoky Mountains National Park will increase inresponse to regional warming.     

Phosphate additions have no effect on microbial biomass and activity in a northern hardwood forest

High rates of atmospheric nitrogen (N) deposition have raised questions about shifting patterns of nutrient limitation in northern hardwood forests. Of particular interest is the idea that increased supply of N may induce phosphorus (P) limitation of plant and microbial processes, especially in acid soils where P sorption by Al is high. In this study, we established field plots and plant-free laboratory mesocosms with P and Ca additions to test the hypotheses that 1) microbial biomass and activity are limited by P in the northern hardwood forest soils at the Hubbard Brook Experimental Forest in NH USA; 2) elevated Ca increases inherent P availability and therefore reduces any effects of added P and 3) P effects are more marked in the more carbon (C) rich Oie compared to the Oa horizon. Treatments included P addition (50 kg P ha⁻¹), Ca addition (850 kg Ca ha⁻¹) and Ca + P addition (850 kg Ca ha⁻¹ and 50 kg P ha⁻¹). The P treatments increased resin-available P levels and reduced phosphatase activity, but had no effect on microbial biomass C, microbial respiration, C metabolizing enzymes, potential net N mineralization and nitrification in the Oie or Oa horizon of either field plots or plant free mesocosms, in either the presence or absence of Ca. Total, prokaryote, and eukaryote PLFA were reduced by P addition, possibly due to reductions in mycorrhizal fungal biomass. These results suggest that increased N deposition and acidification have not created P limitation of microbial biomass and activity in these soils.     

Minor response of gross N turnover and N leaching to drying,rewetting and irrigation in the topsoil of a Norway spruce forest

Forest floors in the temperate climate zone are frequently subjected to strong changes in soil moisture, but the consequences for the soil N cycle are poorly known. In a field experiment we tested the hypotheses that soil drying leads to a decrease of gross N turnover and that natural rewetting causes a pulse of gross N turnover and an increase of N leaching from the forest floor. A further hypothesis was that optimal water availability induced by irrigation causes maximum N turnover and N leaching. Replicated control, throughfall exclusion and irrigation plots were established in a Norway spruce forest to simulate different precipitation patterns during a growing season. Gross N turnover rates were determined in undisturbed soil cores from Oi + Oe and Oa + EA horizons by the ¹⁵N pool dilution technique. Forest floor percolates were periodically collected by suction plates. After 142 mm throughfall was excluded, the median soil water potential at the throughfall exclusion plots increased from pF 1.9 to 4.5 in the Oi + Oe horizon and from pF 1.8 to 3.8 in the Oa + EA horizon. Gross ammonification ranged from 14 to 45 mg N kg⁻¹ soil day⁻¹ in the Oi + Oe horizon and from 4.6 to 11.4 mg N kg⁻¹ soil day⁻¹ in the Oa + EA horizon. Gross ammonification of both horizons was smallest in the throughfall exclusion plots during the manipulation, but the differences between all treatments were not statistically significant. Gross nitrification in both horizons was very small, ranging from 1.6 to 11.1 mg N kg⁻¹ soil day⁻¹. No effects of decreasing water potential and rewetting on gross nitrification rates were observed because of the small rates and huge spatial variations. Irrigation had no effect as the differences from the control in soil water potential remained small. N leaching from the forest floor was not affected by the treatments. Our findings suggest that ammonification in forest floors continues at considerable rates even at small water potentials. The hypotheses of increased N turnover and N leaching following rewetting of dry forest floor or irrigation were not confirmed.     

Phosphate additions have no effect on microbial biomass and activity in a northern hardwood forest

High rates of atmospheric nitrogen (N) deposition have raised questions about shifting patterns of nutrient limitation in northern hardwood forests. Of particular interest is the idea that increased supply of N may induce phosphorus (P) limitation of plant and microbial processes, especially in acid soils where P sorption by Al is high. In this study, we established field plots and plant-free laboratory mesocosms with P and Ca additions to test the hypotheses that 1) microbial biomass and activity are limited by P in the northern hardwood forest soils at the Hubbard Brook Experimental Forest in NH USA; 2) elevated Ca increases inherent P availability and therefore reduces any effects of added P and 3) P effects are more marked in the more carbon (C) rich Oie compared to the Oa horizon. Treatments included P addition (50 kg P ha⁻¹), Ca addition (850 kg Ca ha⁻¹) and Ca + P addition (850 kg Ca ha⁻¹ and 50 kg P ha⁻¹). The P treatments increased resin-available P levels and reduced phosphatase activity, but had no effect on microbial biomass C, microbial respiration, C metabolizing enzymes, potential net N mineralization and nitrification in the Oie or Oa horizon of either field plots or plant free mesocosms, in either the presence or absence of Ca. Total, prokaryote, and eukaryote PLFA were reduced by P addition, possibly due to reductions in mycorrhizal fungal biomass. These results suggest that increased N deposition and acidification have not created P limitation of microbial biomass and activity in these soils.     

Carbon mineralization is promoted by phosphorus and reduced by nitrogen addition in the organic horizon of northern hardwood forests

Limitations to the respiratory activity of heterotrophic soil microorganisms exert important controls of CO₂ efflux from soils. In the northeastern US, ecosystem nutrient status varies across the landscape and changes with forest succession following disturbance, likely impacting soil microbial processes regulating the transformation and emission of carbon (C). We tested whether nitrogen (N) or phosphorus (P) limit the mineralization of soil organic C (SOC) or that of added C sources in the Oe horizon of successional and mature northern hardwood forests in three locations in central New Hampshire, USA. Added N reduced mineralization of C from SOC and from added leaf litter and cellulose. Added P did not affect mineralization from SOC; however, it did enhance mineralization of litter- and cellulose- C in organic horizons from all forest locations. Added N increased microbial biomass N and K₂SO₄-extractable DON pools, but added P had no effect. Microbial biomass C increased with litter addition but did not respond to either nutrient. The direction of responses to added nutrients was consistent among sites and between forest ages. We conclude that in these organic horizons limitation by N promotes mineralization of C from SOC, whereas limitation by P constrains mineralization of C from new organic inputs. We also suggest that N suppresses respiration in these organic horizons either by relieving the N limitation of microbial biomass synthesis, or by slowing turnover of C through the microbial pool; concurrent measures of microbial growth and turnover are needed to resolve this question.     

Phosphorus mineralization in subarctic agricultural and forest soils

Summary Potential P and C mineralization rates were determined in a 12-week laboratory incubation study on subarctic forest and agricultural soil samples with and without N fertilizer added. There was no significant difference in net inorganic P produced between N fertilized and unfertilized soils. The forest soil surface horizons had the highest net inorganic P mineralized, 32 mg P kg^-1 soil for the Oie and 17 mg P kg^-1 soil for the Oa. In the cropped soils net inorganic P immobilization started after 4 weeks and lasted through 12 weeks of incubation. Cumulative CO₂–C evolution rates differed significantly among soils, and between fertilizer treatments, with the N-fertilized soils evolving lower rates of CO₂–C than the unfertilized soils. Soils from the surface horizons in the forest evolved the highest rates of CO₂–C (127.6 and 89.4 mg g^-1 soil for the Oie and Oa horizons, respectively) followed by the cleared uncropped soil (42.8 mg g^-1 soil C), and the cropped soils (25.4 and 29.0 mg g^-1 soil C). In vitro soil respiration rates, or potential soil organic matter decomposition rates, decreased with increasing time after clearing and in accord with the degree of disturbance. Only soils with high potential C mineralization rates and high organic P to total P ratios, mineralized P by the end of the study. Mineralizable P appeared to be associated with readily mineralizable organic C.     

Stock, turnover time and accumulation of organic matter in bulk and density fractions of a Podzol soil

Temperate forest soils store large amounts of organic matter and are considered as net sinks for atmospheric carbon dioxide. Information about the sink strength and the turnover time of soil organic carbon (SOC) is required to assess the potential response of soils to climate change. Here we report on stocks, turnover times (TT) and accumulation of SOC in bulk soil and density fractions from genetic horizons of a Podzol in the Fichtelgebirge, Germany. Stocks of SOC, total nitrogen and exchangeable cations determined in nine quantitative soil pits strongly varied with stone content and thickness of horizons in both the organic layer and the mineral soil. On the basis of radiocarbon signatures, mean turnover times of 4, 9 and 133 years, respectively, were calculated for Oi, Oe and Oa horizons from three soil pits, using a non-steady-state model. The Oa horizons accumulated 4–8 g C m⁻² year⁻¹ whereas the Oi and Oe horizons were close to steady-state during the past decade. Free particulate organic matter (FPOM) was the most abundant fraction in the Oa and EA horizons with TT of 70–480 years. In the B horizons, mineral associated organic matter (MAOM) dominated with over 40% of total SOC and had TT of 390–2170 years. In contrast to other horizons, MAOM in the Bsh and Bs horizon had generally faster TT than occluded particulate organic matter (OPOM), possibly because of sorption of dissolved organic carbon by iron and aluminium oxides/hydroxides. Our results suggest that organic horizons with relatively short turnover times could be particularly vulnerable to changes in climate or other disturbances.     

Carbon additions increase nitrogen availability in northern hardwood forest soils

 The effects of acetate additions to northern hardwood forest soils on microbial biomass carbon (C) and nitrogen (N) content, soil inorganic N levels, respirable C and potential net N mineralization and nitrification were evaluated. The experiment was relevant to a potential watershed-scale calcium (Ca) addition that aims to replace Ca depleted by long-term exposure to acid rain. One option for this addition is to use calcium-magnesium (Mg) acetate, a compound that is inexpensive and much more readily soluble than the Ca carbonate that is generally used for large-scale liming. Field plots were treated with sodium (NA) acetate, Na bicarbonate or water (control) and were sampled (forest floor – Oe and Oa combined) 2, 10 and 58 days following application. It was expected that the addition of C would lead to an increase in biomass C and N and a decrease in inorganic N. Instead, we observed no effect on biomass C, a decline in biomass N and an increase in N availability. One possible explanation for our surprising results is that the C addition stimulated microbial activity but not growth. A second, and more likely, explanation for our results is that the C addition did stimulate microbial growth and activity, but there was no increase in microbial biomass due to predation of the new biomass by soil fauna. The results confirm the emerging realization that the effects of increases in the flow of C to soils, either by deliberate addition or from changes in atmospheric CO₂, are more complex than would be expected from a simple C : N ratio analysis. Evaluations of large-scale manipulations of forest soils to ameliorate effects of atmospheric deposition or to dispose of wastes should consider microbial and faunal dynamics in considerable detail. Received: 13 March 1998     

Clear-cutting of a Norway spruce stand: implications for controls on the dynamics of dissolved organic matter in the forest floor

Clear‐cutting of forest provides a unique opportunity to study the response of dynamic controls on dissolved organic matter. We examined differences in concentrations, fluxes and properties of dissolved organic matter from a control and a clear‐cut stand to reveal controlling factors on its dynamics. We measured dissolved organic C and N concentrations and fluxes in the Oi, Oe and Oa horizons of a Norway spruce stand and an adjacent clear‐cutting over 3 years. Aromaticity and complexity of organic molecules were determined by UV and fluorescence spectroscopy, and we measured δ¹³C ratios over 1 year. Annual fluxes of dissolved organic C and N remained unchanged in the thin Oi horizon (～ 260 kg C ha^?1, ～ 8.5 kg N ha^?1), despite the large reduction in fresh organic matter inputs after clear‐cutting. We conclude that production of dissolved organic matter is not limited by lack of resource. Gross fluxes of dissolved organic C and N increased by about 60% in the Oe and 40% in the Oa horizon upon clear‐cutting. Increasing organic C and N concentrations and increasing water fluxes resulted in 380 kg C ha^?1 year^?1 and 10.5 kg N ha^?1 year^?1 entering the mineral soil of the clear‐cut plots. We found numerous indications that the greater microbial activity induced by an increased temperature of 1.5°C in the forest floor is the major factor controlling the enhanced production of dissolved organic matter. Increasing aromaticity and complexity of organic molecules and depletion of ¹³C pointed to an accelerated processing of more strongly decomposed parts of the forest floor resulting in increased release of lignin‐derived molecules after clear‐cutting. The largest net fluxes of dissolved organic C and N were in the Oi horizon, yet dissolved organic matter sampled in the Oa horizon did not originate mainly from the Oi horizon. Largest gross fluxes in the Oa horizon (control 282 kg C ha^?1) and increased aromaticity and complexity of the molecules with increasing depth suggested that dissolved organic matter was derived mainly from decomposition, transformation and leaching of more decomposed material of the forest floor. Our results imply that clear‐cutting releases additional dissolved organic matter which is sequestered in the mineral soil where it has greater resistance to microbial decay.     

Influence of balsam poplar tannin fractions on carbon and nitrogen dynamics in Alaskan taiga floodplain soils

The feedbacks between plant and soil processes play an important role in driving forest succession. One poorly understood feedback mechanism is the interaction between plant secondary chemicals and soil microbes. In the Alaskan taiga, changes in nutrient cycling caused by balsam poplar (Populus balsamifera) secondary chemicals may affect the transition from alder (Alnus tenuifolia) to balsam poplar on river floodplains. We examined the effects of four poplar condensed tannin fractions on N cycling in alder and poplar soils. Tannins were added to forest floor samples from both poplar and alder sites. Samples were incubated for 1 month in the laboratory with soil respiration rates measured over the course of the incubation. At the end of the incubation we measured both net and gross nitrogen mineralization and nitrification, microbial biomass C and N, and the activity of various exoenzymes. In all soils, tannin additions reduced N availability, however, the mechanisms differed depending on the molecular weight of the tannin and the native soil microbial community. Low molecular weight tannin fractions served as a labile C source in poplar Oi, poplar Oe, and alder Oe horizons but were toxic to microbes in alder Oi. High molecular weight tannin fractions appeared to act primarily by binding extracellular substrates and thus limiting C and N mineralization, with the strongest effects observed in the alder soils.     

Drying–rewetting cycles release phosphorus from forest soils

Drying–rewetting cycles (D/W) occur frequently in topsoils and may mobilize phosphorus (P). We investigated the effect of repeated D/W on the release of dissolved inorganic (DIP) and organic P (DOP) from forest floors and A horizons. Samples were taken from 3 European beech sites and from 3 Norway spruce sites. Soils were desiccated up to pF 6 (–100 MPa) in three D/W cycles in the laboratory, while the controls were kept permanently at 50% water holding capacity. After each drying, P was extracted from the soils in water. D/W caused the release of DIP and DOP especially from O layers. There was no general difference in response to D/W between samples from beech and spruce. The net release of DIP after D/W was largest from the Oe horizons (average 50–60 mg P kg^?1) for both beech and spruce forest soils. The net release of DIP from Oi layers was on average 7.8 mg P kg^?1 and from spruce Oa layers 21.1 mg P kg^?1. In the A horizons, net DIP release was similar in beech and spruce soils with 0.4 mg P kg^?1. The release of DOP was less than the release of DIP except for the A horizons. Repeated cycles did not increase the release of DIP and DOP. The release of DIP and DOP was positively correlated with the microbial biomass in Oe and Oa layers but not in Oi layers. Our results suggest that D/W may significantly influence the short term availability of dissolved P in both beech and spruce forest soils.     

Effects of decreasing water potential on gross ammonification and nitrification in an acid coniferous forest soil

Changes in the soil water regime, predicted as a consequence of global climate change, might influence the N cycle in temperate forest soils. We investigated the effect of decreasing soil water potentials on gross ammonification and nitrification in different soil horizons of a Norway spruce forest and tested the hypotheses that i) gross rates are more sensitive to desiccation in the Oa and EA horizon as compared to the uppermost Oi/Oe horizon and ii) that gross nitrification is more sensitive than gross ammonification. Soil samples were adjusted by air drying to water potentials from about field capacity to around −1.0 MPa, a range that is often observed under field conditions at our site. Gross rates were measured using the ¹⁵N pool dilution technique. To ensure that the addition of solute label to dry soils and the local rewetting does not affect the results by re-mineralization or preferential consumption of ¹⁵N, we compared different extraction and incubation times.T₀ times ranging from 10 to 300 min and incubation times of 48 h and 72 h did not influence the rates of gross ammonification and nitrification. Even small changes of water potential decreased gross ammonification and nitrification in the O horizon. In the EA horizon, gross nitrification was below detection limit and the response of the generally low rates of gross ammonification to decreasing water potentials was minor. In the Oi/Oe horizon gross ammonification and nitrification decreased from 37.5 to 18.3 mg N kg⁻¹ soil d⁻¹ and from 15.4 to 5.6 mg N kg⁻¹ soil d⁻¹ when the water potential decreased from field capacity to −0.8 MPa. In the Oa horizon gross ammonification decreased from 7.4 to 4.0 mg N kg⁻¹ soil d⁻¹ when the water potential reached −0.6 MPa. At such water potential nitrification almost ceased, while in the Oi/Oe horizon nitrification continued at a rather high level. Hence, only in the Oa horizon nitrification was more sensitive to desiccation than ammonification. Extended drought periods that might result from climate change will cause a reduction in gross N turnover rates in forest soils even at moderate levels of soil desiccation.     

Microbially available carbon in buried riparian soils in a glaciated landscape

Buried horizons and lenses in riparian soil profiles harbor large amounts of carbon relative to the surrounding soil horizons. Because these buried soil horizons, as well as deep surface horizons, frequently lie beneath the water table, their impact on nitrogen transport across the terrestrial–aquatic interface depends upon their frequency and spatial distribution, and upon the lability of associated organic matter. We collected samples of 51 soil horizons from 14 riparian zones Rhode Island, USA, where soil profiles are characterized by glacial outwash and alluvial deposits. These soil samples came from as deep as 2 m and ranged in carbon content from <1% to 44% in a buried O horizon 54–74 cm deep. We used these samples to: (1) determine the extent to which carbon in buried horizons, and deep surface horizons, is potentially microbially available; (2) identify spatial patterns of carbon mineralization associated with surface and buried horizons; and (3) evaluate likely relationships between soil horizon types, chemical characteristics and carbon mineralization. Carbon mineralization rates associated with buried horizons during anaerobic incubations ranged from 0.0001 to 0.0175 μmol C kg soil^?1 s^?1 and correlated positively with microbial biomass (R=0.89, P<0.0001, n=21). Excluding surface O horizons from the analysis, carbon mineralization varied systematically with horizon type (surface A, buried A, buried O, lenses, A/C, B, C) (P<0.05) but not with depth or depth x horizon interaction (overall R²=0.59, P<0.0005, n=47). In contrast to this result and to most published data sets, ¹³C-to-¹²C and ¹⁵N-to-¹⁴N ratios of organic matter declined with depth (¹³C?26.9 to ?29.3 per mil, ¹⁵N+5.6 to ?0.8 per mil). The absence of a relationship between horizon depth and C availability suggests that carbon availability in these buried horizons may be determined by the abundance and quality of organic matter at the time of horizon formation or burial, rather than by duration since burial, and implies that subsurface microbial activity is largely disconnected from surface ecosystems. Our results contribute to the emerging view that buried horizons harbor microbially available C in quantities relevant to ecosystem processes, and suggest that buried C-rich soil horizons need to be incorporated into assessments of the depth of the biologically active zone in near-stream subsurface soils.     

Buried organic horizons represent amino acid reservoirs in boreal forest soils

We examined the composition and concentration of amino acids by soil horizon and depth on the Tanana River floodplain in interior Alaska. Soils from mid-successional stages of balsam poplar and white spruce were separated into successive forest floor (Oe/Oa), buried organic horizons (BOHs), and mineral horizons; and water-extractable amino acid composition and concentration were determined by HPLC. The number, depth, and thickness of BOHs were highly variable across the landscape and among replicates of the same stand type, reflecting differences in terrace age, flood frequency, flood intensity, river channel position, vegetation inputs, and decomposition. BOHs generally had lower pH and bulk density, higher moisture content, and greater concentrations of carbon, nitrogen, and roots than the surrounding mineral horizons. In each horizon of both successional stages, the soil amino acid pool was dominated by glutamic acid, glutamine, alanine, asparagine, aspartic acid, and histidine, which together accounted for approximately 80% of the total amino acids found. Despite the similar overall amino acid composition among the horizons, proportions of glutamine generally increased with depth and were generally greater in the mineral horizons than in the BOHs, suggesting root exudation or fine root turnover as an amino acid source. In both successional stages, amino acid concentrations were nearly always highest in the Oe/Oa horizon and rapidly decreased with depth. BOHs generally had greater amino acid concentrations than the surrounding mineral horizons in both successional stages, but amino acid concentrations in successive BOHs declined with depth in the soil profile, suggesting that although BOHs do remain as biological hot spots and potential nutrient reservoirs as far down as 60 cm depth, their importance declines over time.     

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     

Microbial processes controlling P availability in forest spodosols as affected by soil depth and soil properties

Because carbon dioxide (CO₂) concentration is rising, increases in plant biomass and productivity of terrestrial ecosystems are expected. However, phosphorus (P) unavailability may disable any potential enhanced growth of plants in forest ecosystems. In response to P scarcity under elevated CO₂, trees may mine deeper the soil to take up more nutrients. In this scope, the ability of deep horizons of forest soils to supply available P to the trees has to be evaluated. The main objective of the present study was to quantify the relative contribution of topsoil horizons and deep horizons to P availability through processes governed by the activity of soil micro-organisms. Since soil properties vary with soil depth, one can therefore assume that the role of microbial processes governing P availability differs between soil layers. More specifically, our initial hypothesis was that deeper soil horizons could substantially contribute to total plant available P in forested ecosystems and that such contribution of deep horizons differs among sites (due to contrasting soil properties). To test this hypothesis, we quantified microbial P and mineralization of P in ‘dead’ soil organic matter to a depth of 120 cm in forest soils contrasting in soil organic matter, soil moisture and aluminum (Al) and iron (Fe) oxides. We also quantified microbiological activity and acid phosphomonoesterase activity. Results showed that the role of microbial processes generally decreases with increasing soil depth. However, the relative contribution of surface (litter and 0–30 cm) and deep (30–120 cm) soil layers to the stocks of available P through microbial processes (51–62 kg P ha^?1) are affected by several soil properties, and the contribution of deep soil layers to these stocks vary between sites (from 29 to 59%). This shows that subsoils should be taken into account when studying the microbial processes governing P availability in forest ecosystems. For the studied soils, microbial P and mineralization of P in ‘dead’ soil organic matter particularly depended on soil organic matter content, soil moisture and, to a minor extent, Al oxides. High Al oxide contents in some sites or in deep soil layers probably result in the stabilization of soil organic compounds thus reducing microbiological activity and mineralization rates. The mineralization process in the litter also appeared to be P-limited and depended on the C:P ratio of soil organic matter. Thus, this study highlighted the effects of soil depth and soil properties on the microbial processes governing P availability in the forest spodosols.     

Spatial and Temporal Variation in Calcium and Aluminum in Northern Hardwood Forest Floors

Acid rain results in losses of exchangeable base cations from soils, but the mechanism of base cation displacement from the forest floor is not clear, and has been hypothesized to involve mobilization of aluminum from the mineral soil. We attempted to test the hypothesis that losses of calcium from the forest floor were balanced by increases in Al in NewHampshire northern hardwoods. We measured exchangeable (six stands) and acid extractable (13 stands) Ca and Al in horizons of the forest floor over an interval of 15 years. Our sampling scheme was quite intensive, involving 50 or 60 blocks per stand, composited in groups of 10 for chemical analysis. Even at this level of effort, few stands exhibited changes large enough to be significant. Because of high spatial variability, differences would have had to be greater than about 50% to be statistically detectable. Differences in Ca and Al concentrations between Oi, Oe, Oa, and A horizons, however, were readily detected. Acid-extractable Al increased with depth, while Ca concentrations decreased; Ca-to-Al ratios decreased from 8.3 (charge basis) in the Oi to 0.2 in the A horizon. Therefore, a small change in sampling depth, or the inclusion of more or less A horizon material in the forest floor, could cause large differences in measured Ca and Al concentrations. To detect small changes in exchangeable cations over time would require sampling very intensively with careful control for comparability of horizons.     

Simulated grazer effects on microbial respiration in a subarctic meadow: Implications for nutrient competition between plants and soil microorganisms

Plant-microbial competition for nutrients is considered to be a strong mechanism affecting nutrient distribution in subarctic ecosystems, but the role of grazers on the distribution of nutrients between the plants and soil microorganisms remain poorly understood. We designed a factorial fertilization and clipping experiment to study the potential competition between plants and soil microorganisms for soil nitrogen in an ecosystem under grazing. We assumed that clipping reduces plant photosynthetic capacity and C flux to the soil, which ultimately results in lower microbial substrate availability and reduced potential for N immobilization. In concurrence with microbial substrate availability, increased nutrient availability through fertilization was expected to enhance microbial N in the unclipped but not in the clipped treatment.Clipping significantly reduced microbial respiration, suggesting that grazing reduces the labile C available for soil microbes in the system. Clipping had no effect on microbial C and N and the amount of NH₄-N captured in ion exchange resin bags, which was used as an index of net N mineralization. Microbial potential for N immobilization thus seemed insensitive to grazer-mediated changes in microbial availability of labile substrates. Fertilization had no effects or interactions with clipping on microbial C and N. By contrast, we found a close negative correlation between the plant root biomass and microbial N, indicating that plants had a negative impact on the microbial nutrient acquisition. The subarctic grassland vegetation seemed superior to the soil microorganisms in the competition for nutrients even when the plants were subjected to artificial grazing. We suggest that nutrient competition by higher plants constrained the microbial N immobilization in the system, which could explain why the reduction in microbial C availability by clipping had little effects on microbial N acquisition. In this subarctic system, grazing has significant influences on soil C cycling, but due to plant predominance in the competition for nutrients, does not affect N allocation between the plants and the soil microorganisms.     

Soil microbial biomass C:N:P stoichiometry and microbial use of organic phosphorus

Microbial mineralization and immobilization of nutrients strongly influence soil fertility. We studied microbial biomass stoichiometry, microbial community composition, and microbial use of carbon (C) and phosphorus (P) derived from glucose-6-phosphate in the A and B horizons of two temperate Cambisols with contrasting P availability. In a first incubation experiment, C, nitrogen (N) and P were added to the soils in a full factorial design. Microbial biomass C, N and P concentrations were analyzed by the fumigation-extraction method and microbial community composition was analyzed by a community fingerprinting method (automated ribosomal intergenic spacer analysis, ARISA). In a second experiment, we compared microbial use of C and P from glucose-6-phosphate by adding ¹⁴C or ³³P labeled glucose-6-phosphate to soil. In the first incubation experiment, the microbial biomass increased up to 30-fold due to addition of C, indicating that microbial growth was mainly C limited. Microbial biomass C:N:P stoichiometry changed more strongly due to element addition in the P-poor soils, than in the P-rich soils. The microbial community composition analysis showed that element additions led to stronger changes in the microbial community in the P-poor than in the P-rich soils. Therefore, the changed microbial biomass stoichiometry in the P-poor soils was likely caused by a shift in the microbial community composition. The total recovery of ¹⁴C derived from glucose-6-phosphate in the soil microbial biomass and in the respired CO₂ ranged between 28.2 and 37.1% 66 h after addition of the tracer, while the recovery of ³³P in the soil microbial biomass was 1.4–6.1%. This indicates that even in the P-poor soils microorganisms mineralized organic P and took up more C than P from the organic compound. Thus, microbial mineralization of organic P was driven by microbial need for C rather than for P. In conclusion, our experiments showed that (i) the microbial biomass stoichiometry in the P-poor soils was more susceptible to additions of C, N and P than in the P-rich soils and that (ii) even in the P-poor soils, microorganisms were C-limited and the mineralization of organic P was mainly driven by microbial C demand.