Effects of carbon and phosphorus addition on microbial respiration,N<Subscript>2</Subscript>O emission,and gross nitrogen mineralization in a phosphorus-limited grassland soil

Soil microbes are frequently limited by carbon (C), but also have a high phosphorus (P) requirement. Little is known about the effect of P availability relative to the availability of C on soil microbial activity. In two separate experiments, we assessed the effect of P addition (20 mg P kg^?1 soil) with and without glucose addition (500 mg C kg^?1 soil) on gross nitrogen (N) mineralization (¹⁵N pool dilution method), microbial respiration, and nitrous oxide (N₂O) emission in a grassland soil. In the first experiment, soils were incubated for 13 days at 90% water holding capacity (WHC) with addition of NO₃^? (99 mg N kg^?1 soil) to support denitrification. Addition of C and P had no effect on gross N mineralization. Initially, N₂O emission significantly increased with glucose, but it decreased at later stages of the incubation, suggesting a shift from C to NO₃^? limitation of denitrifiers. P addition increased the N₂O/CO₂ ratio without glucose but decreased it with glucose addition. Furthermore, the ¹⁵N recovery was lowest with glucose and without P addition, suggesting a glucose by P interaction on the denitrifying community. In the second experiment, soils were incubated for 2 days at 75% WHC without N addition. Glucose addition increased soil ¹⁵N recovery, but had no effect on gross N mineralization. Possibly, glucose addition increased short-term microbial N immobilization, thereby reducing N-substrates for nitrification and denitrification under more aerobic conditions. Our results indicate that both C and P affect N transformations in this grassland soil.     

Ecosystem partitioning of N-glycine after long-term climate and nutrient manipulations, plant clipping and addition of labile carbon in a subarctic heath tundra

Low temperatures and high soil moisture restrict cycling of organic matter in arctic soils, but also substrate quality, i.e. labile carbon (C) availability, exerts control on microbial activity. Plant exudation of labile C may facilitate microbial growth and enhance microbial immobilization of nitrogen (N). Here, we studied ¹⁵N label incorporation into microbes, plants and soil N pools after both long-term (12 years) climate manipulation and nutrient addition, plant clipping and a pulse-addition of labile C to the soil, in order to gain information on interactions among soil N and C pools, microorganisms and plants. There were few effects of long-term warming and fertilization on soil and plant pools. However, fertilization increased soil and plant N pools and increased pool dilution of the added ¹⁵N label. In all treatments, microbes immobilized a major part of the added ¹⁵N shortly after label addition. However, plants exerted control on the soil inorganic N concentrations and recovery of total dissolved ¹⁵N (TD¹⁵N), and likewise the microbes reduced these soil pools, but only when fed with labile C. Soil microbes in clipped plots were primarily C limited, and the findings of reduced N availability, both in the presence of plants and with the combined treatment of plant clipping and addition of sugar, suggest that the plant control of soil N pools was not solely due to plant uptake of soil N, but also partially caused by plants feeding labile C to the soil microbes, which enhanced their immobilization power. Hence, the cycling of N in subarctic heath tundra is strongly influenced by alternating release and immobilization by microorganisms, which on the other hand seems to be less affected by long-term warming than by addition or removal of sources of labile C.     

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.     

Effect of P stoichiometry on the abundance of nitrogen-cycle genes in phosphorus-limited paddy soil

Previous studies have shown that phosphorus addition to P-limited soils increases gaseous N loss. A possible explanation for this phenomenon is element stoichiometry (specifically of C:N:P) modifying linked nutrient cycling, leading to enhanced nitrification and denitrification. In this study, we investigated how P stoichiometry influenced the dynamics of soil N-cycle functional genes. Rice seedlings were planted in P-poor soils and incubated with or without P application. Quantitative PCR was then applied to analyze the abundance of ammonia-oxidizing (amoA) and denitrifying (narG nirK, nirS, nosZ) genes in soil. P addition reduced bacterial amoA abundance but increased denitrifying gene abundance. We suggest this outcome is due to P-induced shifts in soil C:P and N:P ratios that limited ammonia oxidization while enhancing P availability for denitrification. Under P application, the rhizosphere effect raised ammonia-oxidizing bacterial abundance (amoA gene) and reduced nirK, nirS, and nosZ in rhizosphere soils. The change likely occurred through greater C input and O₂ release from roots, thus altering C availability and redox conditions for microbes. Our results show that P application enhances gaseous N loss potential in paddy fields mainly through stimulating denitrifier growth. We conclude that nutrient availability and elemental stoichiometry are important in regulating microbial gene responses, thereby influencing key ecosystem processes such as denitrification.

Open image in new window

Graphical abstract ?

     

Short-term competition between crop plants and soil microbes for inorganic N fertilizer

Agricultural systems that receive high amounts of inorganic nitrogen (N) fertilizer in the form of either ammonium (NH₄⁺), nitrate (NO₃⁻) or a combination thereof are expected to differ in soil N transformation rates and fates of NH₄⁺ and NO₃⁻. Using ¹⁵N tracer techniques this study examines how crop plants and soil microbes vary in their ability to take up and compete for fertilizer N on a short time scale (hours to days). Single plants of barley (Hordeum vulgare L. cv. Morex) were grown on two agricultural soils in microcosms which received either NH₄⁺, NO₃⁻ or NH₄NO₃. Within each fertilizer treatment traces of ¹⁵NH₄⁺ and ¹⁵NO₃⁻ were added separately. During 8 days of fertilization the fate of fertilizer ¹⁵N into plants, microbial biomass and inorganic soil N pools as well as changes in gross N transformation rates were investigated. One week after fertilization 45-80% of initially applied ¹⁵N was recovered in crop plants compared to only 1-10% in soil microbes, proving that plants were the strongest competitors for fertilizer N. In terms of N uptake soil microbes out-competed plants only during the first 4 h of N application independent of soil and fertilizer N form. Within one day microbial N uptake declined substantially, probably due to carbon limitation. In both soils, plants and soil microbes took up more NO₃⁻ than NH₄⁺ independent of initially applied N form. Surprisingly, no inhibitory effect of NH₄⁺ on the uptake and assimilation of nitrate in both, plants and microbes, was observed, probably because fast nitrification rates led to a swift depletion of the ammonium pool. Compared to plant and microbial NH₄⁺ uptake rates, gross nitrification rates were 3-75-fold higher, indicating that nitrifiers were the strongest competitors for NH₄⁺ in both soils. The rapid conversion of NH₄⁺ to NO₃⁻ and preferential use of NO₃⁻ by soil microbes suggest that in agricultural systems with high inorganic N fertilizer inputs the soil microbial community could adapt to high concentrations of NO₃⁻ and shift towards enhanced reliance on NO₃⁻ for their N supply.     

Nitrification activity in submerged soils and its relation to denitrification loss

Summary Nitrification activity (formation of NO ₂ ^– + NO ₃ ^– per unit soil weight) was measured in the surface layer of 15 presubmerged soils incubated in petri dishes under flooded but aerobic conditions. soils with pH above 5 nitrified quickly, whereas soils with pH below this level did not nitrify or nitrified slowly. The pH values between 7 and 8.5 were optimal for nitrification. Organic-matter levels in the 15 soils of our study did not influence their nitrification activities. In a follow-up greenhouse pot study, after a period of 3 weeks, ¹⁵N-balance measurements showed that the loss of N through apparent denitrification did not follow the nitrification patterns of the soils observed in the petri dishes. Apparent denitrification accounted for 16.8% and 18.9% loss of ¹⁵N from a soil with insignificant nitrification activity and a soil with high nitrification activity, respectively. These results, thus, indicate a lack of correspondence between the nitrification activities of soil and the denitrification loss of N when the former was measured in the dark and the latter was estimated in the light. Soils that nitrified in the darkness of the incubator did not nitrify in the daylight in the greenhouse.     

Combined effects of soil waterlogging and compaction on rice (Oryza sativa L.) growth, soil aeration, soil N transformations and 15N discrimination

 The combined effects of soil compaction and soil waterlogging on the growth of two rice cultivars (Oryza sativa L., cultivars Kanto 168 and Koshihikari) and soil N transformations were studied in pots. Although waterlogging eliminated initial differences in mechanical resistance between compacted and loose soils, Kanto 168 and Koshihikari roots had, respectively, less biomass and a lower porosity if soil was compacted prior to waterlogging. The cause for this was probably established before waterlogging. Redox values showed that upland soils were well aerated. Loose waterlogged soils contained oxic sites, but compacted waterlogged soils did not. Potential denitrification was stimulated by waterlogging and, to a larger extent, by plant presence. Waterlogging lowered potential nitrifying capacities, by competition between plants and micro-organisms for NH₄ ⁺ rather than by oxygen shortage. Compaction prior to waterlogging benefited the potential nitrifying capacity of soils with either cultivar and the potential denitrifying capacity for soils with Koshihikari. Compaction had no effect on nitrification or denitrification in upland soils. N recoveries were low, especially in pots without plants, as a result from sampling strategy and N loss. On day 42/43 after potting, total δ¹⁵N values of waterlogged pots were positive, whereas after 22 days all pots had negative total δ¹⁵N values. Final δ¹⁵N values of plant parts from waterlogged and upland soils were positive and negative, respectively. Although the δ¹⁵N values generally accorded well with the other results, they did not support higher N losses from compacted waterlogged soils than from loose waterlogged soils with plants, as suggested by potential denitrifying activities. Received: 4 February 2000     

Mineralization and denitrification in upland, nearly saturated and flooded subtropical soil I. Effect of nitrate and ammoniacal nitrogen

 The influence of fertilizer N applied through nitrate and ammoniacal sources on the availability of nitrate, supply of C, and gaseous N losses via denitrification (using acetylene inhibition technique) in a semiarid subtropical soil (Typic Ustochrepts) was investigated in a growth chamber simulating upland [60% water-filled pore space (WFPS)], nearly saturated (90% WFPS), and flooded (120% WFPS) conditions. The rate of denitrification was very low in the upland soil conditions, irrespective of fertilizer N treatments. Increasing water content to nearly saturated and flooded conditions resulted in four- to sixfold higher rates of denitrification within 2 days, suggesting that the denitrifying activity commences quickly. Results of this study reveal that (1) under restricted aeration, these soils could support high rates of denitrification (∼6 mg N kg^–1 day^–1) for short periods when nitrate is present; (2) application of fertilizer N as nitrate enhances N losses via denitrification (∼10 mg N kg^–1 day^–1) – however, the supply of available C determines the intensity and duration of denitrification; (3) when fertilizer N is applied as an ammoniacal form, nitrification proceeds slowly and nitrate availability limits denitrification in flooded soil; (4) the nearly saturated soil, being partially aerobic, supported greater nitrification of applied ammoniacal fertilizer N than flooded soil resulting in higher relative rates of denitrification; and (5) under aerobic soil conditions, 26 mg mineral N kg^–1 accumulated in control soil over a 16-day period, demonstrating a modest capacity of such semiarid subtropical soils, low in organic matter, to supply N to growing plants. Received: 7 June 1999     

Soil warming and liming impacts on the recovery of 15N in an acidic soil under soybean cropping

Liming has important implications for N dynamics in acidic soils planted with legumes that are not fully understood. We used a ¹⁵N tracer (K¹⁵NO₃) to examine N dynamics in a Chromic Luvisol planted with soybean with and without lime in environmentally‐controlled chambers set at 20°C and 30°C (full factorial design). Liming increased total N and ¹⁵N recovery in soybean, but had no effect on microbial recovery. Elevated temperature, increased total plant N, decreased ¹⁵N recovery in soybean and microbes, and increased loss of N through leaching. Our results show enhanced uptake of soil mineral N by soybean with liming, thereby reducing N loss from soil, while an increase in temperature from 20°C to 30°C may enhance N loss in these systems.     

Clay Addition to Sandy Soil Reduces Nutrient Leaching—Effect of Clay Concentration and Ped Size

Leaching of nutrients, particularly in sandy soil with low nutrient and water holding capacity (WHC), is a major threat to marine and fresh water pollution. Addition of clay soil to sandy soil could be an option to increase water and nutrient holding capacity of sandy soils, but the effect of clay soil addition may depend on the form in which the clay soil is added and the addition rate. Clay soil was added to sandy soil at rate of 10 or 20% (w/w) finely ground (<2 mm) or 2 and 5 mm peds with and without nitrogen (N) and phosphorus (P) fertilizer equivalent to 60 kg N ha^?1 and 15 kg P ha^?1. The clay sand mixture for each treatment was weighed (30 g) in cores with nylon mesh at the bottom. The soils were incubated at 80% WHC for 7 weeks. To obtain leachate, 20 mL reverse osmosis (RO) water was added every week to each core. Leachate was analysed for inorganic N, P, and pH. Soil was analyzed for N, P, and pH before and after the leaching. Clay addition significantly reduced the leaching of N and P compared to sandy soil alone, with greatest reduction by finely ground clay soil and least with 5 mm peds. Compared to sandy soil alone, 83% more N was retained in clay-amended soil and P retention was doubled. This study showed that addition of finely ground clay soil can substantially reduce N and P leaching and thereby increase fertilizer retention compared to sandy soil alone.     

Influence of soil phosphorus status and nitrogen addition on carbon mineralization from 14C-labelled glucose in pasture soils

 This study examines the effect of soil P status and N addition on the decomposition of ¹⁴C-labelled glucose to assess the consequences of reduced fertilizer inputs on the functioning of pastoral systems. A contrast in soil P fertility was obtained by selecting two hill pasture soils with different fertilizer history. At the two selected sites, representing low (LF) and high (HF) fertility status, total P concentrations were 640 and 820 mg kg^–1 and annual pasture production was 4,868 and 14,120 kg DM ha^–1 respectively. Soils were amended with ¹⁴C-labelled glucose (2,076 mg C kg^–1 soil), with and without the addition of N (207 mg kg^–1 soil), and incubated for 168 days. During incubation, the amounts of ¹⁴CO₂ respired, microbial biomass C and ¹⁴C, microbial biomass P, extractable inorganic P (P_i) and net N mineralization were determined periodically. Carbon turnover was greatly influenced by nutrient P availability. The amount of glucose-derived ¹⁴CO₂ production was high (72%) in the HF and low (67%) in the LF soil, as were microbial biomass C and P concentrations. The ¹⁴C that remained in the microbial biomass at the end of the 6-month incubation was higher in the LF soil (15%) than in the HF soil (11%). Fluctuations in P_i in the LF soil during incubation were small compared with those in HF soil, suggesting that P was cycling through microbial biomass. The concentrations of P_i were significantly greater in the HF samples throughout the incubation than in the LF samples. Net N mineralization and nitrification rates were also low in the LF soils, indicating a slow turnover of microorganisms under limited nutrient supply. Addition of N had little effect on biomass ¹⁴C and glucose utilization. This suggests that, at limiting P fertility, C turnover is retarded because microbial biomass becomes less efficient in the utilization of substrates. Received: 18 October 1999     

Plant and microbial uptake of nitrogen and phosphorus affected by drought using 15N and 32P tracers

Competition for nutrients between plants and microbes is an important determinant for plant growth, biodiversity and carbon cycling. Perturbations such as drought affect the availability of nitrogen (N) and phosphorus (P), and may cause shifts in uptake of N and P between plants and microbes. Competitiveness for these nutrients may depend on how flexible plants and microbes are in taking up N and P. We used a novel dual isotope labelling technique (¹⁵N and ³²P) to assess short-term uptake of N and P by plants and microbes affected by drought in two different plant–soil systems. Mesocosms were extracted from two grassland sites differing in soil nutrient availability and plant species. Half of the mesocosms were subjected to drought one week prior to injection of ¹⁵N (as KNO₃) and ³²P (as H₃PO₄) tracers. Uptake rates of ${{\text{NO}}_3}^{-}$ and P in plants and microbes were estimated based on average source pool enrichment during the labelling period and on plant and microbial recovery of ¹⁵N and ³²P measured after 4 days of labelling. Overall competition for N and P was reduced with drought as less ${{\text{NO}}_3}^{-}$ and P was taken up in plants and microbes. However, plant uptake of ${{\text{NO}}_3}^{-}$ was more sensitive to drought than microbial ${{\text{NO}}_3}^{-}$ uptake, while microbial P uptake was more sensitive than plant P uptake. These different sensitivities to drought by plants and microbes may decouple the N and P cycle with increased drought conditions.     

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

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

Phosphorus addition enhances gross microbial N cycling in phosphorus-poor soils: a <Superscript>15</Superscript>N study from two long-term fertilization experiments

The tight coupling between nitrogen (N) and phosphorus (P) suggests that P availability may affect soil microbial N dynamics in terrestrial ecosystems. However, how P addition affects the internal N transformations in P-deficient agricultural soil remains poorly understood. We hypothesized that an increase in gross microbial N rates in P-deficient soil should occur after long-term P inputs in agricultural soils. We thus conducted a ¹⁵N pool dilution experiment to quantify the gross microbial N transformation rates after long-term mineral fertilizer applications in an upland fluvo-aquic soil (from Fengqiu with pH 8.55) and upland red soil (from Qiyang with pH 5.49) in China. We found that P addition significantly enhanced the gross N mineralization and immobilization rates when N and K were also applied, probably due to the increased soil total C and N concentrations at both soils. Also, gross nitrification rate was stimulated by P addition, perhaps because of enhanced gross N mineralization rates and associated NH₄⁺ substrate availability. Our results showed that long-term P addition may stimulate soil gross N dynamics and hence increase overall N availability for crops in P-deficient agricultural soils.     

Effects of soil moisture on gross N transformations and N2O emission in acid subtropical forest soils

Soil moisture changes, arising from seasonal variation or from global climate changes, could influence soil nitrogen (N) transformation rates and N availability in unfertilized subtropical forests. A ¹⁵?N dilution study was carried out to investigate the effects of soil moisture change (30–90 % water-holding capacity (WHC)) on potential gross N transformation rates and N₂O and NO emissions in two contrasting (broad-leaved vs. coniferous) subtropical forest soils. Gross N mineralization rates were more sensitive to soil moisture change than gross NH₄ ⁺ immobilization rates for both forest soils. Gross nitrification rates gradually increased with increasing soil moisture in both forest soils. Thus, enhanced N availability at higher soil moisture values was attributed to increasing gross N mineralization and nitrification rates over the immobilization rate. The natural N enrichment in humid subtropical forest soils may partially be due to fast N mineralization and nitrification under relatively higher soil moisture. In broad-leaved forest soil, the high N₂O and NO emissions occurred at 30 % WHC, while the reverse was true in coniferous forest soil. Therefore, we propose that there are different mechanisms regulating N₂O and NO emissions between broad-leaved and coniferous forest soils. In coniferous forest soil, nitrification may be the primary process responsible for N₂O and NO emissions, while in broad-leaved forest soil, N₂O and NO emissions may originate from the denitrification process.     

Nitrification is the key process determining N use efficiency in paddy soils

Nitrogen (N) fertilizer use efficiency (NUE) in flooded paddy fields is relatively low. Many N fertilizer management options have been proposed to enhance NUE and minimize environmental damage. However, few investigations are focusing on the role of the characteristics of soil N transformations in regulating NUE and N losses in paddy fields. In this study, we test the role of soil N transformations on NUE and N losses under rice growth conditions in two paddy soils collected from Jiangxi (JX) and Sichuan (SC) in China. The N recoveries of applied ¹⁵N either as nitrate or ammonium in plant and soil, and N losses estimated by ¹⁵N balance were investigated in rice pot experiments using a ¹⁵N tracing technique. The results showed that gross nitrification rates in soil collected from JX were much lower than those in soils collected from SC either at 60% water holding capacity (WHC) or rice growth (flooding) conditions, which could be due to the difference in soil pH. The ‐N concentration in soil solution was maintained at a relatively high level for a long time period after N fertilizer application in the JX soil (41 d) compared to the SC soil (26 d), caused by different nitrification rates owing to different soil pH. The ¹⁵N uptake by rice in the JX soil (29–78%) was always significantly higher than that in the SC soil (22–54%), while N losses from the plant–soil system in the JX soil (17–21%) were always significantly lower than those from the SC soil (20–34%) at the same rice growth stage in the labeled ¹⁵N ammonium treatment. However, there were no significant differences in ¹⁵N uptake by rice and N losses in applied treatment between the two studied soils. These results indicate that nitrification, not denitrification, was the key process determining NUE and N losses in paddy soils. The results of the N application gradient experiment also indicated that higher amounts of N fertilizer should be applied for the same amount of N uptake, however, this caused higher N losses, in soils characterized by high nitrification rate (e.g ., the alkaline soil). Results highlighted that soil N transformations in particular nitrification rate provided a very good guideline for an optimized N management.     

Evidence for nitrification ability controlling nitrogen use efficiency and N losses via denitrification in paddy soils

For understanding the effects of nitrification ability on nitrogen (N) use efficiency and N losses via denitrification in paddy soils under flooding conditions, six paddy soils with different nitrification activities were sampled from various sites of China and a pot experiment was conducted. Rice plants at tillering stage were transplanted into pots and harvested 7.5 days after transplanting, ¹⁵N-(NH₄)₂SO₄ was applied 2.5 days after rice transplanting under continuously flooding conditions. The N losses by denitrification were determined by the unrecovered ¹⁵N applied as ¹⁵NH₄ ⁺ and the N use efficiency (NUE) was calculated by ¹⁵N taken up by rice plants. Plant height (from 33.8 to 37.3 cm) and biomass (from 1.07 g pot^?1 to 1.52 g pot^?1) increased significantly with the native NH₄ ⁺ concentration in the studied soils (P < 0.01). The NUE decreased, whereas the N losses via denitrification increased due to the increase in the nitrification rate of soils determined at 60% water holding capacity (P < 0.05). The results implied that the nitrification activity of paddy soils is a key factor in controlling NUE and N losses via denitrification.     

Plant‐available N:P alters root litter N recycling in a Mediterranean tree–grass ecosystem

Background: Nitrogen deposition can cause an ecosystem‐level shift in available N (nitrogen) to P (phosphorus) availability. However, most plant N nutrition is from edaphic sources rather than deposition and in seasonally dry grassland systems, root litter is the predominant nutrient source. Aims: We were interested how litter turnover and altered nutrient recycling from dead biomass can compensate for these shifts in ecosystem stoichiometry. Methods: We studied a Mediterranean savanna amended with N or NP treatments three years prior. We measured root and plant‐available soil N:P stoichiometry in two micro‐habitats: open pasture and beneath oak canopies. ¹⁵N‐labelled root litter incubated in topsoils without litterbags was used to trace uptake of litter N by herbaceous strata roots. Results: Since fertilization, NP added sites have become relatively P enriched, resulting in lower N:P ratios in living roots than either when N was added alone or control sites. Total litter‐derived ¹⁵N uptake by roots was proportional to root ingrowth response but higher in the NP than N treatment, indicating a higher N demand when N and P were added together. We observed more ¹⁵N uptake by plants under tree canopies, indicating a tighter nutrient recycling loop in these micro‐habitats in contrast to treatment level ‘fertility' trends. Conclusions: Root stoichiometry responded to manipulated soil nutrient availability and N uptake was altered as plants attempted to compensate for nutrient availability imbalances, indicating that these ecosystem perturbations have long term effects on nutrient cycling which can propagate to whole system function. This was also related to functional community‐level adaptions between micro‐habitats with under canopy communities more able to take advantage of the litter nutrient source.     

Nitrogen losses from two grassland soils with different fungal biomass

Nitrogen losses from agricultural grasslands cause eutrophication of ground- and surface water and contribute to global warming and atmospheric pollution. It is widely assumed that soils with a higher fungal biomass have lower N losses, but this relationship has never been experimentally confirmed. With the increased interest in soil-based ecosystem services and sustainable management of soils, such a relationship would be relevant for agricultural management. Here we present a first attempt to test this relationship experimentally. We used intact soil columns from two plots from a field experiment that had consistent differences in fungal biomass (68 ± 8 vs. 111 ± 9 μg C g⁻¹) as a result of different fertilizer history (80 vs. 40 kg N ha⁻¹ y⁻¹ as farm yard manure), while other soil properties were very similar. We performed two greenhouse experiments: in the main experiment the columns received either mineral fertilizer N or no N (control). We measured N leaching, N₂O emission and denitrification from the columns during 4 weeks, after which we analyzed fungal and bacterial biomass and soil N pools. In the additional ¹⁵N experiment we traced added N in leachates, soil, plants and microbial biomass. We found that in the main experiment, N₂O emission and denitrification were lower in the high fungal biomass soil, irrespective of the addition of fertilizer N. Higher ¹⁵N recovery in the high fungal biomass soil also indicated lower N losses through dentrification. In the main experiment, N leaching after fertilizer addition showed a 3-fold increase compared to the control in low fungal biomass soil (11.9 ± 1.0 and 3.9 ± 1.0 kg N ha⁻¹, respectively), but did not increase in high fungal biomass soil (6.4 ± 0.9 after N addition vs. 4.5 ± 0.8 kg N ha⁻¹ in the control). Thus, in the high fungal biomass soil more N was immobilized. However, the ¹⁵N experiment did not confirm these results; N leaching was higher in high fungal biomass soil, even though this soil showed higher immobilization of ¹⁵N into microbial biomass. However, only 3% of total ¹⁵N was found in the microbial biomass 2 weeks after the mineral fertilization. Most of the recovered ¹⁵N was found in plants (approximately 25%) and soil organic matter (approximately 15%), and these amounts did not differ between the high and the low fungal biomass soil. Our main experiment confirmed the assumption of lower N losses in a soil with higher fungal biomass. The additional ¹⁵N experiment showed that higher fungal biomass is probably not the direct cause of higher N retention, but rather the result of low nitrogen availability. Both experiments confirmed that higher fungal biomass can be considered as an indicator of higher nitrogen retention in soils.     

Mineralization of soil organic nitrogen solubilized by aqua ammonia fertilizer

Organic N solubilized by NH_3(aq) was extracted from ¹⁵N-labelled or unlabelled soil, concentrated and added to non-extracted soil, which was incubated under aerobic conditions at 27±1°C. Gross N mineralization, gross N immobilization, and nitrification in soils with or without addition of unlabelled soluble organic N were estimated by models based on the dilution of the NH ₄ ⁺ or NO _inf3 ^sup- pools, which were labelled with ¹⁵N at the beginning of incubation. Mineralization of labelled organic N was measured by the appearance of label in the mineral N pool. Although gross N mineralization and gross N immobilization were increased in two soils between day 0 and day 7 following addition of unlabelled organic N solubilized by NH_3(aq), there was no increase in net N mineralization. Solubilization of ¹⁵N-labelled organic N increased and the ¹⁵N enrichment of the soluble organic N decereased as the concentration of NH_3(aq) added increased. A constant proportion of approximately one-quarter of the labelled organic N added at different rates to non-extracted soil was recovered in the mineral N pool after an incubation period of 14 days, and the availability ratios calculated from net N mineralization data were 1.1:1 and 2.1:1 for 111 and 186 mg added organic-N kg^-1 soil, respectively, indicating that the mineralization of organic N was increased by solubilization.