Temporal evolution of biochar's impact on soil nitrogen processes – a 15N tracing study

Biochar addition to soils has been proposed as a means to increase soil fertility and carbon sequestration. However, its effect on soil nitrogen (N) cycling and N availability is poorly understood. To gain better insight into the temporal variability of the impact of biochar on gross soil N dynamics, two ¹⁵N tracing experiments, in combination with numerical data analysis, were conducted with soil from a biochar field trial, 1 day and 1 year after application of a woody biochar type. The results showed accelerated soil N cycling immediately following biochar addition, with increased gross N mineralization (+34%), nitrification (+13%) and ammonium (NH₄⁺) and nitrate (NO₃⁻) immobilization rates (+4500% and +511%, respectively). One year after biochar application, the biochar acted as an inert substance with respect to N cycling. In the short term, biochar's labile C fraction and a pH increase can explain stimulated microbial activity, while in the longer term, when the labile C fraction has been mineralized and the pH effect has faded, the accelerating effect of biochar on N cycling ceases. In conclusion, biochar accelerates soil N transformations in the short-term through stimulating soil microbial activity, thereby increasing N bio-availability. This effect is, however, temporary.     

Ten years of elevated atmospheric carbon dioxide alters soil nitrogen transformations in a sheep‐grazed pasture

The increasing concentration of atmospheric carbon dioxide (CO₂) is expected to lead to enhanced competition between plants and microorganisms for the available nitrogen (N) in soil. Here, we present novel results from a ¹⁵N tracing study conducted with a sheep‐grazed pasture soil that had been under 10 years of CO₂ enrichment. Our study aimed to investigate changes in process‐specific gross N transformations in a soil previously exposed to an elevated atmospheric CO₂ (eCO₂) concentration and to examine indicators for the occurrence of progressive nitrogen limitation (PNL). Our results show that the mineralization–immobilization turnover (MIT) was enhanced under eCO₂, which was driven by the mineralization of recalcitrant organic N. The retention of N in the grassland was enhanced by increased dissimilatory NO₃^? reduction to NH₄⁺ (DNRA) and decreased NH₄⁺ oxidation. Our results indicate that heterotrophic processes become more important under eCO₂. We conclude that higher MIT of recalcitrant organic N and enhanced N retention are mechanisms that may alleviate PNL in grazed temperate grassland.     

Nitrogen cycling in canopy soils of tropical montane forests responds rapidly to indirect N and P fertilization

Although the canopy can play an important role in forest nutrient cycles, canopy‐based processes are often overlooked in studies on nutrient deposition. In areas of nitrogen (N) and phosphorus (P) deposition, canopy soils may retain a significant proportion of atmospheric inputs, and also receive indirect enrichment through root uptake followed by throughfall or recycling of plant litter in the canopy. We measured net and gross rates of N cycling in canopy soils of tropical montane forests along an elevation gradient and assessed indirect effects of elevated nutrient inputs to the forest floor. Net N cycling rates were measured using the buried bag method. Gross N cycling rates were measured using ¹⁵N pool dilution techniques. Measurements took place in the field, in the wet and dry season, using intact cores of canopy soil from three elevations (1000, 2000 and 3000 m). The forest floor had been fertilized biannually with moderate amounts of N and P for 4 years; treatments included control, N, P, and N + P. In control plots, gross rates of NH₄⁺ transformations decreased with increasing elevation; gross rates of NO₃^? transformations did not exhibit a clear elevation trend, but were significantly affected by season. Nutrient‐addition effects were different at each elevation, but combined N + P generally increased N cycling rates at all elevations. Results showed that canopy soils could be a significant N source for epiphytes as well as contributing up to 23% of total (canopy + forest floor) mineral N production in our forests. In contrast to theories that canopy soils are decoupled from nutrient cycling in forest floor soil, N cycling in our canopy soils was sensitive to slight changes in forest floor nutrient availability. Long‐term atmospheric N and P deposition may lead to increased N cycling, but also increased mineral N losses from the canopy soil system.     

Soil nitrogen transformations under Populus tremuloides, Betula papyrifera and Acer saccharum following 3 years exposure to elevated CO2 and O3

Increases in atmospheric CO₂ and tropospheric O₃ may affect forest N cycling by altering plant litter production and the availability of substrates for microbial metabolism. Three years following the establishment of our free‐air CO₂–O₃ enrichment experiment, plant growth has been stimulated by elevated CO₂ resulting in greater substrate input to soil; elevated O₃ has counteracted this effect. We hypothesized that rates of soil N cycling would be enhanced by greater plant productivity under elevated CO₂, and that CO₂ effects would be dampened by O₃. We found that elevated CO₂ did not alter gross N transformation rates. Elevated O₃ significantly reduced gross N mineralization and microbial biomass N, and effects were consistent among species. We also observed significant interactions between CO₂ and O₃: (i) gross N mineralization was greater under elevated CO₂ (1.0 mg N kg^?1 day^?1) than in the presence of both CO₂ and O₃ (0.5 mg N kg^?1 day^?1) and (ii) gross NH₄⁺ immobilization was also greater under elevated CO₂ (0.8 mg N kg^?1 day^?1) than under CO₂ plus O₃ (0.4 mg N kg^?1 day^?1). We used a laboratory ¹⁵N tracer method to quantify transfer of inorganic N to organic pools. Elevated CO₂ led to greater recovery of NH₄⁺‐¹⁵N in microbial biomass and corresponding lower recovery in the extractable NO₃^? pool. Elevated CO₂ resulted in a substantial increase in NO₃^?‐¹⁵N recovery in soil organic matter. We observed no O₃ main effect and no CO₂ by O₃ interaction effect on ¹⁵N recovery in any soil pool. All of the above responses were most pronounced beneath Betula papyrifera and Populus tremuloides, which have grown more rapidly than Acer saccharum. Although elevated CO₂ has increased plant productivity, the resulting increase in plant litter production has yet to overcome the influence of the pre‐existing pool of soil organic matter on soil microbial activity and rates of N cycling. Ozone reduces plant litter inputs and also appears to affect the composition of plant litter in a way that reduces microbial biomass and activity.     

Effects of forest management on soil N cycling in beech forests stocking on calcareous soils

The effects of forest management (thinning) on gross and net N conversion, the balance of inorganic N production and consumption, inorganic N concentrations and on soil microbial biomass in the Ah layer were studied in situ during eight intensive field measuring campaigns in the years 2002–2004 at three beech (Fagus sylvatica L.) forest sites. At all sites adjacent thinning plots (“T”) and untreated control plots (“C”) were established. Since the sites are characterized either by cool-moist microclimate (NE site and NW site) or by warm-dry microclimate (SW site) and thinning took place in the year 1999 at the NE and SW sites and in the year 2003 at the NW site the experimental design allowed to evaluate (1) short-term effects (years 1–2) of thinning at the NW site and (2) medium-term effects (years 4–6) of thinning under different microclimate at the SW and NE site. Microbial biomass N was consistently higher at the thinning plots of all sites during most of the field campaigns and was overall significantly higher at the SWT and NWT plots as compared to the corresponding untreated control plots. The size of the microbial biomass N pool was found to correlate positively with both gross ammonification and gross nitrification as well as with extractable soil NO₃⁻ concentrations. At the SW site neither gross ammonification, gross nitrification, gross ammonium (NH₄⁺) immobilization and gross nitrate (NO₃⁻) immobilization nor net ammonification, net nitrification and extractable NH₄⁺ and NO₃⁻ contents were significantly different between control and thinning plot. At the NET plot lower gross ammonification and gross NH₄⁺ immobilization in conjunction with constant nitrification rates coincided with higher net nitrification and significantly higher extractable NO₃⁻ concentrations. Thus, the medium-term effects of thinning varied with different microclimate. The most striking thinning effects were found at the newly thinned NW site, where gross ammonification and gross NH₄⁺ immobilization were dramatically higher immediately after thinning. However, they subsequently tended to decrease in favor of gross nitrification, which was significantly higher at the NWT plot as compared to␣the␣NWC plot during all field campaigns after␣thinning except for April 2004. This increase␣in␣gross nitrification at the NWT plot (1.73 mg N kg⁻¹ sdw day⁻¹ versus 0.48 mg N kg⁻¹ sdw day⁻¹ at the NWC plot) coincided with significantly higher extractable NO₃⁻ concentrations (4.59 mg N kg⁻¹ sdw at the NWT plot versus 0.96 mg N kg⁻¹ sdw at the NWC plot). Pronounced differences in relative N retention (the ratio of gross NH₄⁺ immobilization + gross NO₃⁻ immobilization to gross ammonification + gross nitrification) were found across the six research plots investigated and could be positively correlated to the soil C/N ratio (R = 0.94; p = 0.005). In sum, the results obtained in this study show that (1) thinning can lead to a shift in the balance of microbial inorganic N production and consumption causing a clear decrease in the N retention capacity in the monitored forest soils especially in the first two years after thinning, (2)␣the resistance of the investigated forest ecosystems to disturbances of N cycling by thinning may vary with different soil C contents and C/N ratios, e. g. caused by differences in microclimate, (3) thinning effects tend to decline with the growth of understorey vegetation in the years 4–6 after thinning.     

Context-dependent tree species effects on soil nitrogen transformations and related microbial functional genes

Although it is generally accepted that tree species can influence nutrient cycling processes in soils, effects are not consistently found, nor are the mechanisms behind tree species effects well understood. Our objectives were to gain insights into the mechanism(s) underlying the effects of tree species on soil nitrogen cycling processes, and to determine the consistency of tree species effects across sites. We compared N cycling in soils beneath six tree species (ash, sycamore maple, lime, beech, pedunculate oak, Norway spruce) in common garden experiments planted 42 years earlier at three sites in Denmark with distinct land-use histories (forest and agriculture). We measured: (1) net and gross rates of N transformations using the ¹⁵N isotope pool-dilution method, (2) soil microbial community composition through qPCR of fungal ITS, bacterial and archaeal 16S, and (3) abundance of functional genes associated with N cycling processes—for nitrification the archaeal and bacterial ammonia-monooxygenase genes (amoA AOA and amoA AOB, respectively) and for denitrification, the nitrate reductase genes nirK and nirS. Carbon concentrations were higher in soils under spruce than under broadleaves, so N transformation rates were standardized per g soil C. Soil NH₄⁺ parameters (gross ammonification, gross NH₄⁺ consumption, net ammonification (net immobilization in this case), and NH₄⁺ concentrations, per g C) were all lowest in soils under spruce. Soils under spruce also had the lowest gene abundance of bacteria, bacterial:fungal ratio, denitrifying microorganisms, ammonia-oxidizing archaea and ammonia-oxidizing bacteria. Differences in N-cycling processes and organisms among the five broadleaf species were smaller. The ‘spruce effect’ on soil microbes and N transformations appeared to be driven by its acidifying effect on soil and tighter N cycling, which occurred at the previously forested sites but not at the previously agricultural site. We conclude that existing characteristics of soils, including those resulting from previous land use, mediate the effects of tree species on the soil microbial communities and activities that determine rates of N-cycling processes.     

Soil-mixing effects on inorganic nitrogen production and consumption in forest and shrubland soils

Soils that are physically disturbed are often reported to show net nitrification and NO₃⁻ loss. To investigate the response of soil N cycling rates to soil mixing, we assayed gross rates of mineralization, nitrification, NH₄⁺ consumption, and NO₃⁻ consumption in a suite of soils from eleven woody plant communities in Oregon, New Mexico, and Utah. Results suggest that the common response of net NO₃⁻ flux from disturbed soils is not a straightforward response of increased gross nitrification, but instead may be due to the balance of several factors. While mineralization and NH₄⁺ assimilation were higher in mixed than intact cores, NO₃⁻ consumption declined. Mean net nitrification was 0.12 mg N kg⁻¹ d⁻¹ in disturbed cores, which was significantly higher than in intact cores (−0.19 mg N kg⁻¹ d⁻¹). However, higher net nitrification rates in disturbed soils were due to the suppression of NO₃⁻ consumption, rather than an increase in nitrification. Our results suggest that at least in the short term, disturbance may significantly increase NO₃⁻ flux at the ecosystem level, and that N cycling rates measured in core studies employing mixed soils may not be representative of rates in undisturbed soils.     

Spring ephemeral herbs and nitrogen cycling in a northern hardwood forest: an experimental test of the vernal dam hypothesis

In the late 1970s R.N. Muller and F.H. Bormann posited their ”vernal dam” hypothesis, stating that spring-ephemeral herbs in deciduous forests serve as a temporary sink for N when overstory trees are dormant, and then release this N later, in the summer, when the trees are active. This hypothesis has gained wide acceptance, yet two of its critical assumptions have never been experimentally tested: (1) that N taken up by spring ephemerals would otherwise be lost from the ecosystem, and (2) that N from senesced ephemeral tissues contributes to increased rates of summertime N mineralization. To test these assumptions, I quantified patterns of N cycling and loss from a set of paired plots, half of which served as controls and from half of which all spring-ephemeral plants were removed. There were no significant differences in NO₃ ^– leaching between plots with and without spring ephemeral vegetation. These results are consistent with the relatively low rates of N uptake by the dominant spring ephemeral, Allium tricoccum, and its apparent preference for NH₄ ⁺, which is far less mobile in soil than NO₃ ^–. In addition, based on sequential sampling, I found that soil microorganisms took up 8 times as much N during the spring than did spring-ephemeral herbs (microbial uptake=3.19 vs. plant uptake=0.41 g N m^–2), suggesting that microbial immobilization of N is the dominant sink for N during this season. Removal of spring ephemeral vegetation also had no effect on summertime rates of net N mineralization. Furthermore, the addition of spring ephemeral litter to soil+forest floor microcosms did not significantly increase rates of N mineralization in a laboratory incubation. Instead, this experiment demonstrated the overwhelming influence of forest floor litter in controlling the release of mineral N from these soils. Overall, neither assumption of the vernal dam hypothesis holds true in this ecosystem, where patterns of N cycling and loss appear to be dominated by microbial decomposition of forest floor material and soil organic matter. Received: 24 August 1999 / Accepted: 23 March 2000     

Gross Nitrogen Turnover of Natural and Managed Tropical Ecosystems at Mt. Kilimanjaro,Tanzania

In our study at Mt. Kilimanjaro, East Africa, we quantified gross rates of ammonification, nitrification, nitrogen immobilization, and dissimilatory nitrate reduction to ammonium in soils across different land uses, climate zones (savanna, montane forest ecosystems, extensive agroforest homegarden, and intensively managed coffee plantation), and seasons (dry, wet, and transition from dry to wet season) to identify if and to what extent conversion of natural ecosystems to cultivated land has affected key soil microbial nitrogen turnover processes. Overall variation of gross soil nitrogen turnover rates across different ecosystems was more pronounced than seasonal variations, with the highest turnover rates occurring at the transition between dry and wet seasons. Nitrogen production and immobilization rates positively correlated with soil organic carbon and total nitrogen concentrations as well as substrate availability of dissolved organic carbon and nitrogen r > 0.67, P < 0.05), but did not correlate with soil ammonium and nitrate concentrations. Soil nitrogen turnover rates were highest in the montane Ocotea forest (ammonification 29.84, nitrification 12.67, NH₄ ⁺ immobilization 38.92, NO₃ ^? immobilization 10.74, and DNRA 1.54 µg N g^?1 SDW d^?1) and progressively decreased with decreasing annual rainfall and increasing land-use intensity. Using indicators of N retention and characteristics of soil nutrient status, we observed a grouping of faster, but tighter N cycling in the (semi-) natural savanna and Ocotea forest. This contrasted with a more open N cycle in managed systems (the homegarden and coffee plantation) where N was more prone to leaching or gaseous losses due to high nitrate production rates. The partly disturbed (selected logging) lower montane forest ranged between these two groups.     

Microbial immobilization drives nitrogen cycling differences among plant species

In many terrestrial ecosystems nitrogen (N) limits productivity and plant community composition is influenced by N availability. However, vegetation is not only controlled by N; plant species may influence ecosystem N dynamics through positive or negative effects on N cycling. We examined four potential mechanisms of plant species effects on nitrogen (N) cycling. We found no species differences in gross ammonification suggesting there are no changes in the ecosystem N cycling rate between the soil organic matter pool (SOM) and the plant/microbial pool. We also found weak differences among plant species in gross nitrification, thus plant species only marginally change the relative sizes of the NH₄⁺ and NO₃^? pools. Next, more than 90% of mineralized N was microbially immobilized, and microbial N immobilization was positively correlated with root biomass. Finally, while species differed in extractable soil NO3^? concentration, these differences were not related to root biomass suggesting that microbial immobilization drives net N mineralization and soil NO₃^? levels. Our results indicate that plant species do not cause feedbacks on the N cycling rate among the three major ecosystem N pools over nine years. However, plant carbon (C) inputs to the soil control microbial N immobilization and thereby change N partitioning between the plant and microbial N pools. Furthermore our results suggest that the SOM pool can act as a strong bottleneck for N cycling in these systems.     

Carbon and nitrogen turnover in adjacent grassland and cropland ecosystems

The effects of cultivation and soil texture on net and gross N mineralization, CO₂ evolution and C and N turnover were investigated using paired grassland and cropped sites on soils of three textures. Gross N mineralization and immobilization were measured using¹⁵N-isotope dilution. Grassland soils had high CO₂ evolution and gross N mineralization rates, and low net N mineralization rates. Cropland soils had low CO₂ evolution rates but had high net and gross N mineralization rates. Grassland soils thus had high immobilization rates and cropland soils had low immobilization rates. Cultivation increased N turnover but reduced C turnover. The data suggest that the microflora in grassland soils are N limited, while those of cropland soils are limited by C availability. Increasing clay content reduced N turnover. C turnover was less clearly related to texture. Differences in the immobilization potential of substrates help explain why agricultural soils have higher N losses than do grassland soils.     

Microbial biomass and nitrogen cycling responses to fertilization and litter removal in young northern hardwood forests

The influence of site fertility on soil microbial biomass and activity is not well understood but is likely to be complex because of interactions with plant responses to nutrient availability. We examined the effects of long-term (8 yr) fertilization and litter removal on forest floor microbial biomass and N and C transformations to test the hypothesis that higher soil resource availability stimulates microbial activity. Microbial biomass and respiration decreased by 20–30 % in response to fertilization. Microbial C averaged 3.8 mg C/g soil in fertilized, 5.8 mg C/g in control, and 5.5 mg C/g in litter removal plots. Microbial respiration was 200 µg CO₂-C g^–1 d^–1 in fertilized plots, compared to 270 µg CO₂-C g^–1 d^–1 in controls. Gross N mineralization and N immobilization did not differ among treatments, despite higher litter nutrient concentrations in fertilized plots and the removal of substantial quantities of C and N in litter removal plots. Net N mineralization was significantly reduced by fertilization. Gross nitrification and NO₃ ^– immobilization both were increased by fertilization. Nitrate thus became a more important part of microbial N cycling in fertilized plots even though NH₄ ⁺ availability was not stimulated by fertilization.Soil microorganisms did not mineralize more C or N in response to fertilization and higher litter quality; instead, results suggest a difference in the physiological status of microbial biomass in fertilized plots that influenced N transformations. Respiration quotients (qCO₂, respiration per unit biomass) were higher in fertilized plots (56 µg CO₂-C mg C^–1 d^–1) than control (48 µg CO₂-C mg C^–1 d ^–1) or litter removal (45 µg CO₂-C mg C^–1 d^–1), corresponding to higher microbial growth efficiency, higher proportions of gross mineralization immobilized, and lower net N mineralization in fertilized plots. While microbial biomass is an important labile nutrient pool, patterns of microbial growth and turnover were distinct from this pool and were more important to microbial function in nitrogen cycling.     

Pulse additions of soil carbon and nitrogen affect soil nitrogen dynamics in an arid Colorado Plateau shrubland

Biogeochemical cycles in arid and semi-arid ecosystems depend upon the ability of soil microbes to use pulses of resources. Brief periods of high activity generally occur after precipitation events that provide access to energy and nutrients (carbon and nitrogen) for soil organisms. To better understand pulse-driven dynamics of microbial soil nitrogen (N) cycling in an arid Colorado Plateau ecosystem, we simulated a pulsed addition of labile carbon (C) and N in the field under the canopies of the major plant species in plant interspaces. Soil microbial activity and N cycling responded positively to added C while NH₄⁺–N additions resulted in an accumulation of soil NO₃⁻. Increases in microbial activity were reflected in higher rates of respiration and N immobilization with C addition. When both C and N were added to soils, N losses via NH₃ volatilization decreased. There was no effect of soil C or N availability on microbial biomass N suggesting that the level of microbial activity (respiration) may be more important than population size (biomass) in controlling short-term dynamics of inorganic and labile organic N. The effects of C and N pulses on soil microbial function and pools of NH₄⁺–N and labile organic N were observed to last only for the duration of the moisture pulse created by treatment addition, while the effect on the NO₃⁻–N pool persisted after soils dried to pre-pulse moisture levels. We observed that increases in available C lead to greater ecosystem immobilization and retention of N in soil microbial biomass and also lowered rates of gaseous N loss. With the exception of trace gas N losses, the lack of interaction between available C and N on controlling N dynamics, and the subsequent reduction in plant available N with C addition has implications for the competitive relationships between plants species, plants and microbes, or both.     

The role of gross and net N transformation processes and NH4 + and NO3 − immobilization in controlling the mineral N pool of a temperate mixed deciduous forest soil

In many forests of Europe and north-eastern North America elevated N deposition has opened the forest N cycle, resulting in NO₃ ^? leaching. On the other hand, despite this elevated N deposition, the dominant fate of NO₃ ^? and NH₄ ⁺ in some of these forests is biotic or abiotic immobilization in the soil organic matter pool, preventing N losses. The environmental properties controlling mineral N immobilization and the variation and extent of mineral N immobilization in forest soils are not yet fully understood. In this study we investigated a temperate mixed deciduous forest, which is subjected to an average N deposition of 36.5 kg N ha^?1 yr^?1, but at the same time shows low NO₃ ^? concentrations in the groundwater. The aim of this study was to investigate whether the turnover rate of the mineral N pool could explain these low N leaching losses. A laboratory ¹⁵N pool dilution experiment was conducted to study gross and net N mineralization and nitrification and mineral N immobilization in the organic and uppermost (0–10 cm) mineral layer of the forest soil. Two locations, one at the forest edge (GE) and another one 145 m inside the forest (GF1), were selected. In the organic layers of GE and GF1, the gross N mineralization averaged 10.9 and 11.1 mg N kg^?1 d^?1, the net N mineralization averaged 6.1 and 6.8 mg N kg^?1 d^?1 and NH₄ ⁺ immobilization rates averaged 3.8 and 3.6 mg N kg^?1 d^?1. In the organic layer of GE and GF1, the average gross nitrification was 3.8 and 4.6 mg N kg^?1 d^?1, the average net nitrification was ?25.2 and ?31.3 mg N kg^?1 d^?1 and the NO₃ ^? immobilization rates averaged 29.0 and 35.9 mg N kg^?1 d^?1. For the mineral (0–10 cm) layer the same trend could be observed, but the N transformation rates were much lower for the NH₄ ⁺ pool and not significantly different from zero for the NO₃ ^? pool. Except for the turnover of the NH₄ ⁺ pool in the mineral layer, no significant differences were observed between location GE and GF1. The ratio of NH₄ ⁺ immobilization to gross N mineralization, gross N mineralization to gross nitrification, and NO₃ ^? immobilisation to gross nitrification led to the following observations. The NH₄ ⁺ pool of the forest soil was controlled by N mineralization and NO₃ ^? immobilization was importantly controlling the forest NO₃ ^? pool. Therefore it was concluded that this process is most probably responsible for the limited NO₃ ^? leaching from the forest ecosystem, despite the chronically high N deposition rates.     

Growth and Major Nutrient Concentrations in Brassica campestris Supplied with Different NH4^＋/NO3 Ratios

In the present study, we investigated whether growth and main nutrient ion concentrations of cabbage （Brassica campestris L.） could be increased when plants were subjected to different NH4^＋/NO3- ratios. Cabbage seedlings were grown in a greenhouse in nutrient solutions with five NH4^＋/NO3- ratios （1：0; 0.75：0.25; 0.5：0.5; 0.25：0.75; and 0：1）. The results showed that cabbage growth was reduced by 87% when the proportion of NH4^＋-N in the nutrient solution was more than 75% compared with a ratio NH4^＋/NO3- of 0.5：0.5 35 d after transplanting, suggesting a possible toxicity due to the accumulation of a large amount of free ammonia in the leaves. When the NH4＋/NO3- ratio was 0.5：0.5, fresh seedling weight, root length, and H2PO4- （P）, K^＋, Ca^2＋, and Mg^2＋ concentrations were all higher than those in plants grown under other NH4^＋/NO3- ratios. The nitrate concentration in the leaves was the lowest in plants grown at 0.5： 0.5 NH4^＋/NO3-. The present results indicate that an appropriate NH4^＋/NO3- ratio improves the absorption of other nutrients and maintains a suitable proportion of N assimilation and storage that should benefit plant growth and the quality of cabbage as a vegetable.     

Inorganic nitrogen immobilization in live and sterile soil of old-growth conifer and hardwood forests: implications for ecosystem nitrogen retention

Rapid immobilization of inorganic nitrogen (N) in soil contributes to ecosystem N accumulation, even in old-growth and chronically-fertilized forests once thought to have poor N retention capacity. In old-growth conifer and hardwood stands in Pennsylvania, we tested the hypotheses that biotic and abiotic N immobilization are regulated by N form and forest type. We added ¹⁵NH₄ ⁺, ¹⁵NO₂ ^?, and ¹⁵NO₃ ^? to sterile (γ-irradiated) and live organic-horizon soil and define N immobilization as the mass of added ¹⁵N remaining in soil following extractions conducted 15 min, 24 h, and 21 days later. Immobilization of NO₂ ^? (19–25% of added N) occurred in sterile soils within 15 min and was little changed thereafter. Tracer NO₃ ^? immobilization was not observed, although soils had been pretreated (refrigerated) so as to quantify the lower limit of immobilization potential. Immobilization of NH₄ ⁺ (27%) occurred in live conifer soils by 21 days but not in other treatments. In 21-day incubations, tracer N immobilization was greater in NO₃ ^?-poor and humic-rich soils. Immobilization was greater in sterile than in live soil, perhaps owing to artifacts of sterilization. Conifer stands exhibited more massive O-horizons, so NO₂ ^? immobilization per unit area was greater in conifer (1.46 mg N m^?2) than hardwood (0.43 mg N m^?2) stands, possibly accounting for lower N leaching from conifer forests. Areal immobilization rates appear to be fast enough to retain all N transformed to NO₂ ^?, so NO₂ ^? production may be a limiting step in soil N retention in old-growth ecosystems.     

Microbial dynamics and carbon and nitrogen cycling following re-wetting of soils beneath two semi-arid plant species

Sporadic summer rainfall in semi-arid ecosystems can provide enough soil moisture to drastically increase CO₂ efflux and rates of soil N cycling. The magnitudes of C and N pulses are highly variable, however, and the factors regulating these pulses are poorly understood. We examined changes in soil respiration, bacterial, fungal and microfaunal populations, and gross rates of N mineralization, nitrification, and NH₄⁺ and NO₃^– immobilization during the 10 days following wetting of dry soils collected from stands of big sagebrush (Artemisia tridentata) and cheatgrass (Bromus tectorum) in central Utah. Soil CO₂ production increased more than tenfold during the 17 h immediately following wetting. The labile organic C pool released by wetting was almost completely respired within 2–3 days, and was nearly three times as large in sagebrush soil as in cheatgrass. In spite of larger labile C pools beneath sagebrush, microbial and microfaunal populations were nearly equal in the two soils. Bacterial and fungal growth coincided with depletion of labile C, and populations peaked in both soils 2 days after wetting. Protozoan populations, whose biomass was nearly 3,000-fold lower than bacteria and fungi, peaked after 2–4 days. Gross N mineralization and nitrification rates were both faster in cheatgrass soil than in sagebrush, and caused greater nitrate accumulation in cheatgrass soil. Grazing of bacteria and fungi by protozoans and nematodes could explain neither temporal trends in N mineralization rates nor differences between soil types. However, a mass balance model indicated that the initial N pulse was associated with degradation of microbial substrates that were rich in N (C:N <8.3), and that microbes had shifted to substrates with lower N contents (C:N =15–25) by day 7 of the incubation. The model also suggested that the labile organic matter in cheatgrass soil had a lower C:N ratio than in sagebrush, and this promoted faster N cycling rates and greater N availability. This study provides evidence that the high N availability often associated with wetting of cheatgrass soils is a result of cheatgrass supplying substrates to microbes that are of high decomposability and N content.     

Microbial transformations of C and N in a boreal forest floor as affected by temperature

The effects of temperature on N mineralization were studied in two organic surface horizons (LF and H) of soil from a boreal forest. The soil was incubated at 5 °C and 15 °C after adding ¹⁵ N and gross N fluxes were calculated using a numerical simulation model. The model was calibrated on microbial C and N, basal respiration, and KCl-extractable NH₄ ⁺, NO₃ ⁻, ¹⁵NH₄ ⁺ and ¹⁵ NO₃ ⁻. In the LF layer, increased temperature resulted in a faster turnover of all N pools. In both layers net N mineralization did not increase at elevated temperature because both gross NH₄ ⁺ mineralization and NH₄ ⁺ immobilization increased. In the H layer, however, both gross NH₄ ⁺ mineralization and NH₄ ⁺ immobilization were lower at 15 °C than at 5 °C and the model predicted a decrease in microbial turnover rate at higher temperature although measured microbial activity was higher. The decrease in gross N fluxes in spite of increased microbial activity in the H layer at elevated temperature may have been caused by uptake of organic N. The model predicted a decrease in pool size of labile organic matter and microbial biomass at elevated temperature whereas the amount of refractory organic matter increased. Temperature averaged microbial C/N ratio was 14.7 in the LF layer suggesting a fungi-dominated decomposer community whereas it was 7.3 in the H layer, probably due to predominance of bacteria. Respiration and microbial C were difficult to fit using the model if the microbial C/N ratio was kept constant with time. A separate ¹⁵N-enrichment study with the addition of glucose showed that glucose was metabolized faster in the LF than in the H layer. In both layers, decomposition of organic matter appeared to be limited by C availability. This revised version was published online in June 2006 with corrections to the Cover Date.     

Enhanced foliar 15N enrichment with increasing nitrogen addition rates: Role of plant species and nitrogen compounds

Determining the abundance of N isotope (δ¹⁵N) in natural environments is a simple but powerful method for providing integrated information on the N cycling dynamics and status in an ecosystem under exogenous N inputs. However, whether the input of different N compounds could differently impact plant growth and their ¹⁵N signatures remains unclear. Here, the response of ¹⁵N signatures and growth of three dominant plants (Leymus chinensis, Carex duriuscula, and Thermopsis lanceolata) to the addition of three N compounds (NH₄HCO₃, urea, and NH₄NO₃) at multiple N addition rates were assessed in a meadow steppe in Inner Mongolia. The three plants showed different initial foliar δ¹⁵N values because of differences in their N acquisition strategies. Particularly, T. lanceolata (N₂-fixing species) showed significantly lower ¹⁵N signatures than L. chinensis (associated with arbuscular mycorrhizal fungi [AMF]) and C. duriuscula (associated with AMF). Moreover, the foliar δ¹⁵N of all three species increased with increasing N addition rates, with a sharp increase above an N addition rate of ~10 g N m⁻² year⁻¹. Foliar δ¹⁵N values were significantly higher when NH₄HCO₃ and urea were added than when NH₄NO₃ was added, suggesting that adding weakly acidifying N compounds could result in a more open N cycle. Overall, our results imply that assessing the N transformation processes in the context of increasing global N deposition necessitates the consideration of N deposition rates, forms of the deposited N compounds, and N utilization strategies of the co-existing plant species in the ecosystem.