15N natural abundances and N use by tundra plants

Plant species collected from tundra ecosystems located along a north-south transect from central Alaska to the north coast of Alaska showed large and consistent differences in ¹⁵N natural abundances. Foliar ¹⁵N values varied by about 10% among species within each of two moist tussock tundra sites. Differences in ¹⁵N contents among species or plant groups were consistent across moist tussock tundra at several other sites and across five other tundra types at a single site. Ericaceous species had the lowest ¹⁵N values, ranging between about –8 to –6. Foliar ¹⁵N contents increased progressively in birch, willows and sedges to maximum ¹⁵N values of about +2 in sedges. Soil ¹⁵N contents in tundra ecosystems at our two most intensively studied sites increased with depth and ¹⁵N values were usually higher for soils than for plants. Isotopic fractionations during soil N transformations and possibly during plant N uptake could lead to observed differences in ¹⁵N contents among plant species and between plants and soils. Patterns of variation in ¹⁵N content among species indicate that tundra plants acquire nitrogen in extremely nutrient-poor environments by competitive partitioning of the overall N pool. Differences in plant N sources, rooting depth, mycorrhizal associations, forms of N taken up, and other factors controlling plant N uptake are possible causes of variations in ¹⁵N values of tundra plant species.     

Nitrogen Transfer Between Plants: A 15N Natural Abundance Study with Crop and Weed Species

An increasing amount of evidence indicates that N can be transferred between plants. Nonetheless, a number of fundamental questions remain. A series of experiments was initiated in the field to examine N transfer between N₂-fixing soybean (Glycine max [L.] Merr.) varieties and a non-nodulating soybean, and between N₂-fixing peanut (Arachis hypogaea L.) or soybean and neighboring weed species. The experiments were conducted in soils with low N fertilities and used differences in N accumulation and/or ¹⁵N natural abundance to estimate N transfer. Mixtures of N₂-fixing and non-nod soybean indicated that substantial inter-plant N transfer occurred. Amounts were variable, ranging from negligible levels to 48% of the N found in the non-nod at maturity. Transfer did not appear to strongly penalize the N₂-fixing donor plants. But, in cases where high amounts of N were transferred, N content of donors was noticeably lowered. Differences were evident in the amount of N transferred from different N₂-fixing donor genotypes. Results of experiments with N₂-fixing crops and the weed species prickly sida (Sida spinosa L.) and sicklepod (Senna obtusifolia [L.] Irwin & Barneby) also indicated substantial N transfer occurred over a 60-day period, with amounts accounting for 30–80% of the N present in the weeds. Transfer of N, however, was generally very low in weed species that are known to be non-hosts for arbuscular mycorrhizae (yellow nutsedge, Cyperus esculentus L. and Palmer amaranth, Amaranthus palmeri [S.] Watson). The results are consistent with the view that N transfer occurs primarily through mycorrhizal hyphal networks, and they reveal that N transfer may be a contributing factor to weed problems in N₂-fixing crops in low N fertility conditions.     

Natural abundance of 15N in actinorhizal plants and nodules

Substantial enrichment of some plant parts in ¹⁵N relative to the rest of the plant is unusual, but is found in the nitrogen-fixing nodules of many legumes. A range of actinorhizal plants was surveyed to determine whether the nodules of any of them are also substantially enriched in ¹⁵N. The nonlegume Parasponia, nodulated by a rhizobium, was also included. Four of the actinorhizal genera and Parasponia were grown in N-free culture, and three actinorhizal genera were collected from the field. Nodules of Parasponia, Casuarina and Alnus were¹⁵N enriched relative to other plant parts, but only Parasponia approached the degree of enrichment found in some legume nodules. The nodules of Datisca, Myrica, Elaeagnus, Shepherdia, and Coriaria were depleted in ¹⁵N. Thus many actinorhizal nodules are depleted in ¹⁵N compared to other plant parts and enrichment is modest when it does occur. Whole plant ¹⁵N content (¹⁵N) in four actinorhizal plants and Parasponia showed a relatively narrow range of –1.41 to –1.90. Hence regardless of the degree of nodule enrichment or depletion, whole plant ¹⁵N content appears to vary little in plants grown in N-free culture.     

15N natural abundance of foliage and soil across boreal forests of Finland

This study presents the latitudinal variation (from 60° 30′ N to 68° 2′ N latitude) of natural abundances of ¹⁵N in the foliage, humus and soils of boreal forests of Finland. Our results clearly showed that N concentration of the foliage did not change significantly with latitudes but their ¹⁵N values were significantly higher in higher latitude sites relative to that of the mid and lower latitude sites, indicating the different forms of N uptake at the higher latitudes compared to the lower latitudes. We assume that the higher foliage δ¹⁵N values of the higher latitudes trees might be due to either more openness of N cycle (greater proportional N losses) in these latitudes compared to the sites of southern latitudes (lower N losses) or the differences in their mycorrhizal associations. Regression analysis showed that the temperature was the main factor influencing the ¹⁵N natural abundance of humus and soil of all forest ecosystems, both before and after clear-cut, whereas rainfall was the main controlling factor to the foliage ¹⁵N. Possible reasons behind the increasing δ¹⁵N natural abundances of plants and soils with increasing latitudes are discussed in this paper. The clear-cut did not show any specific trend on the ¹⁵N fractionation in humus and soil, i.e. both ¹⁵N-enrichment and -depletion occurred after clear-cut.     

Agronomic assessment and 15N recovery of urea amended with the urease inhibitor nBTPT (N-(n-butyl) thiophosphoric triamide) for temperate grassland

Three field experiments were undertaken concurrently at one site to evaluate a range of surface-applied nBTPT-amended urea products (0.01, 0.05, 0.1, 0.25 and 0.5% nBTPT w/w) on NH₃ volatilization, grass yield and ¹⁵N recovery in the plant-soil system. Each experiment was repeated on five separate occasions over the 1992 growing season to cover a range of weather conditions. Total NH₃ loss from unamended-urea ranged from 5.5% in early May to 20.8% in June. The inhibitor was highly effective in reducing ammonia volatilization and delaying the time at which maximum rate of NH₃ loss occurred. Over all time periods the % inhibition was 50.4, 82.8, 89.0, 96.5 and 97.0% at the 0.01, 0.05, 0.1, 0.25 and 0.5% nBTPT levels respectively. There was no significant difference in the overall % inhibition in ammonia loss at different times suggesting that the effectiveness of the inhibitor was not dependent on climatic conditions.Over all times incorporation of nBTPT at the 0.05% level increased dry-matter yield by 9% compared to urea alone and increased the shoot recovery of N from 66.7% to 80.9%. Nitrogen saved from volatilization was taken up by the plant, however, the subsequent translation into dry-matter yield appeared to be adversely affected at the high inhibitor rates.There was no significant effect of inhibitor on ¹⁵N recovery in soil at any depth down to 15 cms. nBTPT significantly increased (p < 0.001) the % N derived from fertilizer (% N dff) in the shoot compared to unamended-urea and increased (p < 0.01) the shoot recovery of ¹⁵N from 32% up to 39%. Total ¹⁵N recovery in the soil-plant system was increased by up to 17% by amending urea with nBTPT. This urease inhibitor has been shown to improve the efficiency of urea however, its potential for the European market will be dependent on economic factors.Faculty of Agriculture and Food Science, The Queen's University of Belfast     

Increased soil moisture content increases plant N uptake and the abundance of 15N in plant biomass

The natural abundance of ¹⁵N in plant biomass has been used to infer how N dynamics change with elevated atmospheric CO₂ and changing water availability. However, it remains unclear if atmospheric CO₂ effects on plant biomass ¹⁵N are driven by CO₂-induced changes in soil moisture. We tested whether ¹⁵N abundance (expressed as δ¹⁵N) in plant biomass would increase with increasing soil moisture content at two atmospheric CO₂ levels. In a greenhouse experiment we grew sunflower (Helianthus annuus) at ambient and elevated CO₂ (760 ppm) with three soil moisture levels maintained at 45, 65, and 85% of field capacity, thereby eliminating potential CO₂-induced soil moisture effects. The δ¹⁵N value of total plant biomass increased significantly with increased soil moisture content at both CO₂ levels, possibly due to increased uptake of ¹⁵N-rich organic N. Although not adequately replicated, plant biomass δ¹⁵N was lower under elevated than under ambient CO₂ after adjusting for plant N uptake effects. Thus, increases in soil moisture can increase plant biomass δ¹⁵N, while elevated CO₂ can decrease plant biomass δ¹⁵N other than by modifying soil moisture.     

Natural abundance of 15N in tropical plants with emphasis on tree legumes

Natural abundance of ¹⁵N ( ¹⁵N) of leaves harvested from tropical plants in Brazil and Thailand was analyzed. The ¹⁵N values of non-N₂-fixing trees in Brazil were +4.5±1.9, which is lower than those of soil nitrogen (+8.0±2.2). In contrast, mimosa and kudzu had very low ¹⁵N values (–1.4+0.5). The ¹⁵N values of Panicum maximum and leguminous trees, except Leucaena leucocephala, were similar to those of non-N₂-fixing trees, suggesting that the contribution of fixed N in these plants is negligible. The ¹⁵N values of non-N₂-fixing trees in Thailand were +4.9±2.0. Leucaena leucocephala, Sesbania grandiflora, Casuarina spp. and Cycas spp. had low ¹⁵N values, close to the value of atmospheric N₂ (0), pointing to a major contribution of N₂ fixation in these plants. Cassia spp. and Tamarindus indica had high ¹⁵N values, which confirms that these species are non-nodulating legumes. The ¹⁵N values of Acacia spp. and Gliricidia sepium and other potentially nodulating tree legumes were, on average, slightly lower than those of non-N₂-fixing trees, indicating a small contribution of N₂ fixation in these legumes.     

Estimation of symbiotically fixed nitrogen in field grown soybeans: An application of natural15N/14N abundance and a low level15N-tracer technique

Summary Plants from agricultural and natural upland ecosystem were investigated for¹⁵N content to evaluate the role of symbiotic N₂-fixation in the nitrogen nutrition of soybean. Increased yields and lower δ¹⁵N values of nodulating soybeansvs, non-nodulating isolines gave semi-quantitative estimates of N₂ fixation. A fairly large discrepancy was found between estimations by δ¹⁵N and by N yield at 0 kg N/ha of fertilizer. More precise estimates were made by following changes in plant δ¹⁵N when fertilizer δ¹⁵N was varied near¹⁵N natural abundance level. Clearcut linear relationships between δ¹⁵N values of whole plants and of fertilizer were obtained at 30 kg N/ha of fertilizer for three kinds of soils. In experimental field plots, nodulating soybeans obtained 13±1% of their nitrogen from fertilizer, 66±8% from N₂ fixation and 21±10% from soil nitrogen in Andosol brown soil; 30%, 16% and 54% in Andosol black soil; 7%, 77% and 16% in Alluvial soil, respectively. These values for N₂ fixation coincided with each corresponding estimation by N yield method. Other results include: 1)¹⁵N content in upland soils and plants was variable, and may reflect differences in the mode of mineralization of soil organics, and 2) nitrogen isotopic discrimination during fertilizer uptake (δ¹⁵N of plant minus fertilizer) ranged from −2.2 to +4.9‰ at 0–30 kg N/ha of fertilizer, depending on soil type and plant species. The proposed method can accurately and relatively simply establish the importance of symbiotic nitrogen fixation for soybeans growing in agricultural settings.     

Evaluation of N2-fixation and nitrogen economy of a maize/cowpea intercrop system using15N dilution methods

Yields of above ground biomass and total N were determined in summer-grown maize and cowpea as sole crops or intercrops, with or without supplementary N fertilizer (25 kg N ha⁻¹, urea) at an irrigated site in Waroona, Western Australia over the period 1982–1985. Good agreement was obtained between estimates of N₂ fixation of sole or intercrop cowpea (1984/85 season) based on the¹⁵N natural abundance and¹⁵N fertilizer dilution techniques, both in the field and in a glasshouse pot study. Field-grown cowpea was estimated to have received 53–69% of its N supply from N₂-fixation, with N₂-fixation onlyslightly affected by intercropping or N fertilizer application. Proportional reliance on N₂-fixation of cowpea in glasshouse culture was lower (36–66%) than in the field study and more affected by applied N. Budgets for N were drawn up for the field intercrops, based on above-ground seed yields, return of crop residues, inputs of fixed N and fertilizer N. No account was taken of possible losses of N through volatilization, denitrification and leaching or gains of N in the soil from root biomass. N₂-fixation was estimated tobe 59 kg N ha⁻¹ in the plots receiving no fertilizer N, and 73 kg N ha⁻¹ in plots receiving 25 kg N ha⁻¹ as urea. Comparable fixation by sole cowpea was higher (87 and 82 kg N ha⁻¹ respectively) but this advantage was outweighed by greater land use efficiency by the intercrop than sole crops.     

The fate of labelled15N urea and ammonium nitrate applied to a winter wheat crop

Labelled urea or ammonium nitrate was applied to winter wheat growing on a loamy soil in Northern France. Two applications of fertilizer were given: 50 kg N ha^–1 at tillering (early March) and 110 kg N ha^–1 at the beginning of stem elongation (mid-April). The kinetics of urea hydrolysis, nitrification of ammonium and the disappearance of inorganic nitrogen were followed at frequent intervals. Inorganic nitrogen soon disappeared, mainly immobilized by soil microflora and absorbed by the crop. Net immobilization of fertilizer N occured at a very similar rate for urea and ammonium nitrate. Maximum immobilization (16 kg N ha¹) was found at harvest for the first dressing and at anthesis for the second dressing (23 kg N ha¹). During the nitrification period, the labelled ammonium pool was immobilized two to three times faster than the labelled nitrate pool. No significant net¹⁵N remineralization was found during the growth cycle.The actual denitrification and volatilization losses were probably more important than indicated from calculations made by extrapolation of fluxes measured over short intervals. However microbial immobilization was the most important of the processes which compete with plant uptake for nitrogen.     

The use of mean pool abundances to interpret 15N tracer experiments

A pulse dilution ¹⁵N technique was used in the field to determine the effect of the ammonium to nitrate ratio in a fertilizer application on the uptake of ammonium and nitrate by ryegrass and on gross rates of mineralization and nitrification. Two experiments were performed, corresponding approximately to the first and second cuts of grass. Where no substantial recent immobilization of inorganic nitrogen had occurred, mineralization was insensitive to the form of nitrogen applied, ranging from 2.1–2.6 kg N ha^-1 d^-1. The immobilization of ammonium increased as the proportion of ammonium in the application increased. In the second experiment there was evidence that high rates of immobilization in the first experiment were associated with high rates of mineralization in the second. The implication was that some nitrogen immobilized in the first experiment was re-mineralized during the second. Whether this was nitrogen taken up, stored in roots and released following defoliation was not clear. Nitrification rates in this soil were low (0.1–0.63 kg N ha^-1 d^-1), and as a result, varying the ratio of ammonium to nitrate applied markedly altered the relative uptake of ammonium and nitrate. In the first experiment, where temperatures were low, preferential uptake of ammonium occurred, but where >90% of the uptake was as ammonium, a reduction in yield and nitrogen uptake was observed. In the second experiment, where temperatures and growth rates were higher, the proportion of ammonium to nitrate taken up had no effect on yield or nitrogen uptake.     

Effects of N-deficiency and timing of N supply on the recovery and distribution of labeled 15N in contrasting maize hybrids

Little information exists on the pattern of nitrogen (N) uptake, remobilization and N use efficiency (NUE) in Leafy and stay-green (SG) maize (Zea mays L.) genotypes. A pot experiment was conducted under controlled nutrition and growing conditions to determine the response of Leafy and SG maize genotypes to different levels of N-deficiency and timing of N supply. Three contrasting maize hybrids, Pioneer 3905 (a conventional hybrid with moderate SG characteristics), Pioneer 39F06 Bt (with a high score of SG trait) and Maizex LF850-RR (with a Leafy trait) were grown in 6 L plastic pots. Five different N treatments [no supply of N until V8 (N₁), no supply of N after V8 (N₂), no supply of N after silking (N₃), no supply of N beyond 3 weeks after silking (N₄), and continuous N supply from emergence to physiological maturity (N₅; standard check)] were imposed through modified Hoagland solution applied manually. Labeled ¹⁵N of 5% ¹⁵N₂–NH₄NO₃fertilizer was applied at 3 g per pot at the start of each schedule N treatment. Total amounts of N applied in different treatments were 3.13, 1.32, 1.90, 2.63 and 3.40 g, respectively in N₁, N₂, N₃, N₄ and N₅. Dry matter, N concentration, ¹⁵N (atom% enrichment) and NUE were determined in roots, stalk, leaves and grains at crop maturity. The three contrasting hybrids did not differ in grain yield, total N acquisition, partitioning of ¹⁵N and NUE. Restriction of N supply until V8, and from V8 to physiological maturity significantly reduced grain yield and N-uptake in all hybrids. Irrespective of the level of N-deficiency in plant and timing when the labeled fertilizer was applied, the amount of ¹⁵N recovered in the matured plant was the same in all N treatments. It has been evident that maize continued to take up N beyond 3 weeks after silking and the later N was applied during the development, the higher proportion of it was partitioned to grains. Of the total ¹⁵N uptake, 78% was partitioned to kernels in the N₄ treatment compared to only 61% in the control. Our data showed no evidence of differential N uptake, remobilization and NUE in the SG or Leafy hybrids tested, but the timing of N application and level of N-deficiency in plant significantly influenced N uptake, remobilization and N-dynamics in maize.     

Below-ground nitrogen transfer between different grassland species: Direct quantification by 15N leaf feeding compared with indirect dilution of soil 15N

Nitrogen (N) transfer from one species to another is important for the N cycling in low-input grassland. In the present work, estimates obtained by an indirect ¹⁵N dilution technique were compared with estimates obtained by a direct ¹⁵N leaf feeding technique over two complete growing seasons in red clover-ryegrass and white clover-ryegrass mixtures under field conditions. The direct technique confirmed that N transfer between clovers and ryegrass is a bi-directional process. The transfer of N from both clovers to ryegrass occurred within 25 days upon the first labelling event. A very high N transfer occurred from white clover to the associated ryegrass, 4.5 and 7.5 g m⁻² in the 1st and 2nd production year, respectively. The corresponding values for transfer from red clover to the associated ryegrass were 1.7 and 3.6 g m⁻². Quantified relatively to the total above-ground N content of white clover- ryegrass and red clover-ryegrass mixtures, the N transfer exceeded 50% and 10%, respectively, in three out of seven harvests. The N transfer from ¹⁵N labelled grass to associated clovers constituted a relatively constant proportion of approx. 8% of the above-ground N content of the mixtures. Estimates based on the soil ¹⁵N dilution technique generally underestimated the net N transfer by more than 50% compared to the direct ¹⁵N labelling technique. Furthermore, the indirect ¹⁵N dilution technique estimated only marginal differences between red and white clover in the quantities of N transferred, whereas the direct ¹⁵N labelling technique showed the N transfer from white clover to the associated ryegrass to be significantly higher than that involving red clover. It is concluded that N transfer is a much more dynamic and quantitatively important process in grassland than previously recognised. This revised version was published online in June 2006 with corrections to the Cover Date.     

Nitrogen uptake and recovery from urea and green manure in lowland rice measured by15N and non-isotope techniques

In the recent past considerable attention is paid to minimize dependence on purchased inputs such as inorganic nitrogen fertilizer. Green manure in the form of flood-tolerant, stem-nodulatingSesbania rostrata andAeschynomene afraspera is an alternative N source for rice, which may also increase N use efficiency. Therefore research was conducted to determine the fate of N applied to lowland rice (Oryza sativa L.) in the form ofSesbania rostrata andAeschynomene afraspera green manure and urea in two field experiments using¹⁵N labeled materials.¹⁵N in the soil and rice plant was determined, and¹⁵N balances established. Apparent N recoveries were determined by non-tracer method. ¹⁵N recoveries averaged 90 and 65% of N applied for green manure and urea treatments, respectively. High partial pressures of NH₃ in the floodwater, and high pH probably resulted from urea application and favoured losses of N from the urea treatment. Results show that green manure N can supply a substantial proportion of the N requirements of lowland rice. Nitrogen released fromSesbania rostrata andAeschynomene afraspera green manure was in synchrony with the demand of the rice plant. The effect of combined application of green manure and urea on N losses from urea fertilizer were also investigated. Green manure reduced the N losses from¹⁵N labeled urea possibly due to a reduction in pH of the floodwater. Positive added N interactions (ANIs) were observed. At harvest, an average of 45 and 25% of N applied remained in the soil for green manure and urea, respectively.Contribution from IRRI, Los Baños, Philippines and Justus-Liebig-University, Giessen, GermanyContribution from IRRI, Los Baños, Philippines and Justus-Liebig-University, Giessen, Germany     

Symbiotic N2 fixation by soybean in organic and conventional cropping systems estimated by 15N dilution and 15N natural abundance

Nitrogen (N) is often the most limiting nutrient in organic cropping systems. N₂ fixing crops present an important option to improve N supply and to maintain soil fertility. In a field experiment, we investigated whether the lower N fertilization level and higher soil microbial activity in organic than conventional systems affected symbiotic N₂ fixation by soybean (Glycine max, var. Maple Arrow) growing in 2004 in plots that were since 1978 under the following systems: bio-dynamic (DYN); bio-organic (ORG); conventional with organic and mineral fertilizers (CON); CON with exclusively mineral fertilizers (MIN); non-fertilized control (NON). We estimated the percentage of legume N derived from the atmosphere (%Ndfa) by the natural abundance (NA) method. For ORG and MIN we additionally applied the enriched ¹⁵N isotope dilution method (ID) based on residual mineral and organic ¹⁵N labeled fertilizers that were applied in 2003 in microplots installed in ORG and MIN plots. These different enrichment treatments resulted in equal %Ndfa values. The %Ndfa obtained by NA for ORG and MIN was confirmed by the ID method, with similar variation. However, as plant growth was restricted by the microplot frames the NA technique provided more accurate estimates of the quantities of symbiotically fixed N₂ (Nfix). At maturity of soybean the %Ndfa ranged from 24 to 54%. It decreased in the order ORG > CON > DYN > NON > MIN, with significantly lowest value for MIN. Corresponding Nfix in above ground plant material ranged from 15 to 26 g N m^-2, with a decreasing trend in the order DYN = ORG > CON > MIN > NON. For all treatments, the N withdrawal by harvested grains was greater than Nfix. This shows that at the low to medium %Ndfa, soybeans did not improve the N supply to any system but removed significant amounts of soil N. High-soil N mineralization and/or low-soil P availability may have limited symbiotic N₂ fixation.     

Regional Assessment of N Saturation using Foliar and Root \varvec {\delta}^{\bf 15}{\bf N}

N saturation induced by atmospheric N deposition can have serious consequences for forest health in many regions. In order to evaluate whether foliar may be a robust, regional-scale measure of the onset of N saturation in forest ecosystems, we assembled a large dataset on atmospheric N deposition, foliar and root and N concentration, soil C:N, mineralization and nitrification. The dataset included sites in northeastern North America, Colorado, Alaska, southern Chile and Europe. Local drivers of N cycling (net nitrification and mineralization, and forest floor and soil C:N) were more closely coupled with foliar than the regional driver of N deposition. Foliar increased non-linearly with nitrification:mineralization ratio and decreased with forest floor C:N. Foliar was more strongly related to nitrification rates than was foliar N concentration, but concentration was more strongly correlated with N deposition. Root was more tightly coupled to forest floor properties than was foliar . We observed a pattern of decreasing foliar values across the following species: American beech>yellow birch>sugar maple. Other factors that affected foliar included species composition and climate. Relationships between foliar and soil variables were stronger when analyzed on a species by species basis than when many species were lumped. European sites showed distinct patterns of lower foliar , due to the importance of ammonium deposition in this region. Our results suggest that examining values of foliage may improve understanding of how forests respond to the cascading effects of N deposition.     

Nitrogen fixation in legumes and actinorhizal plants in natural ecosystems: values obtained using 15N natural abundance

Background: Nitrogen fixation has been quantified for a range of crop legumes and actinorhizal plants under different agricultural/agroforestry conditions, but much less is known of legume and actinorhizal plant N₂ fixation in natural ecosystems.

Aims: To assess the proportion of total plant N derived from the atmosphere via the process of N₂ fixation (%Ndfa) by actinorhizal and legume plants in natural ecosystems and their N input into these ecosystems as indicated by their ¹⁵N natural abundance.

Methods: A comprehensive collation of published values of %Ndfa for legumes and actinorhizal plants in natural ecosystems and their N input into these ecosystems as estimated by their ¹⁵N natural abundance was carried out by searching the ISI Web of Science database using relevant key words.

Results: The %Ndfa was consistently large for actinorhizal plants but very variable for legumes in natural ecosystems, and the average value for %Ndfa was substantially greater for actinorhizal plants. High soil N, in particular, but also low soil P and water content were correlated with low legume N₂ fixation. N input into ecosystems from N₂ fixation was very variable for actinorhizal and legume plants and greatly dependent on their biomass within the system.

Conclusions: Measurement of ¹⁵N natural abundance has given greater understanding of where legume and actinorhizal plant N₂ fixation is important in natural ecosystems. Across studies, the average value for %Ndfa was substantially greater for actinorhizal plants than for legumes, and the relative abilities of the two groups of plants to utilise mineral N requires further study.     

Dynamics of soil microbial biomass N under zero and shallow tillage for spring wheat,using15N urea

Summary Field studies to determine the effect of zero and shallow (10 cm) cultivation on microbial biomass were conducted on several Chernozemic soils in western Canada. Using the CHCl₃ fumigation method, the distribution of microbial biomass N and the immobilization and subsequent release of added¹⁵N (¹⁵N-urea) from the microbial biomass were determined in the A horizon, at the 0 to 5 and 5 to 10 cm depth, during the growing season for spring wheat.Temporal variation in microbial biomass N, associated with the development of the rhizosphere, was characterized by an increase between Feekes stage 1 and 5 or 10 and decrease at Feekes stage 11.4. Over the long term, the variation in biomass N between tillage systems corresponded with crop residue distribution. Immobilization of fertilizer N was related to the increase in biomass N from Feekes stage 1, which in turn, was associated with the incorporation of recent crop residues or levels of labile organic matter in the surface soil. The study demonstrated the relatively rapid remineralization of immobilized fertilizer N under field conditions and emphasized the role of the microbial biomass N as both a sink and source of mineral N.     

Natural 15N abundance of soil N pools and N2O reflect the nitrogen dynamics of forest soils

Natural ¹⁵N abundance measurements of ecosystem nitrogen (N) pools and ¹⁵N pool dilution assays of gross N transformation rates were applied to investigate the potential of δ¹⁵N signatures of soil N pools to reflect the dynamics in the forest soil N cycle. Intact soil cores were collected from pure spruce (Picea abies (L.) Karst.) and mixed spruce-beech (Fagus sylvatica L.) stands on stagnic gleysol in Austria. Soil δ¹⁵N values of both forest sites increased with depth to 50 cm, but then decreased below this zone. δ¹⁵N values of microbial biomass (mixed stand: 4.7 ± 0.8‰, spruce stand: 5.9 ± 0.9‰) and of dissolved organic N (DON; mixed stand: 5.3 ± 1.7‰, spruce stand: 2.6 ± 3.3‰) were not significantly different; these pools were most enriched in ¹⁵N of all soil N pools. Denitrification represented the main N₂O-producing process in the mixed forest stand as we detected a significant ¹⁵N enrichment of its substrate NO₃⁻ (3.6 ± 4.5‰) compared to NH₄⁺ (−4.6 ± 2.6‰) and its product N₂O (−11.8 ± 3.2‰). In a ¹⁵N-labelling experiment in the spruce stand, nitrification contributed more to N₂O production than denitrification. Moreover, in natural abundance measurements the NH₄⁺ pool was slightly ¹⁵N-enriched (−0.4 ± 2.0 ‰) compared to NO₃⁻ (−3.0 ± 0.6 ‰) and N₂O (−2.1 ± 1.1 ‰) in the spruce stand, indicating nitrification and denitrification operated in parallel to produce N₂O. The more positive δ¹⁵N values of N₂O in the spruce stand than in the mixed stand point to extensive microbial N₂O reduction in the spruce stand. Combining natural ¹⁵N abundance and ¹⁵N tracer experiments provided a more complete picture of soil N dynamics than possible with either measurement done separately.     

Measurements of abundances of 15N and 13C as tools in retrospective studies of N balances and water stress in forests: A discussion of preliminary results

Preliminary attempts to make retrospective studies of N balances and water stress in forest fertilization experiments by analyzing changes in the abundances of ¹⁵N and ¹³C, respectively, are discussed. Most evidence is from the Swedish Forest Optimum Nutrition Experiments, which have been running for two decades. Annual additions of N have been given either alone or in combination with other elements, notably P and K, every third year. Processes leading to loss of N, e.g. volatilization of ammonia, nitrification followed by leaching or denitrification, and denitrification alone, discriminate against the heavy isotope ¹⁵N. A correlation was found between fractional losses of added N and the change in ¹⁵N () during 19 years in current needles in a Scots pine forest, irrespective of source of N. Isotope effects were larger on urea than on ammonium nitrate plots (2 as compared to 9 ¹⁵N ()) because of ammonia volatilization and higher rates of nitrification. They developed gradually over time, which opens possibilities to analyse the development of N saturation. However, the analysis may be confounded by shifts in ¹⁵N abundance of fertilizer N. In another trial, N isotope effects could be seen in both plants and soils 10 years after the last fertilization; they were smaller in soils because of a large pretreatment memory effect, but we expect them to persist there for decades.The enzyme RuBisCo discriminates strongly against the heavy isotope ¹³C during photosynthesis, but this effect becomes less expressed as stomata close because of water stress. The supply of N may also affect the ¹³C () via effects on rates of photosynthesis, and the source of N may have an influence directly via non-RubisCo carboxylations, and indirectly via effects on water use efficiency. In a trial with Norway spruce, the effect of N fertilization on the ¹³C () of current needles was strongly correlated with production and weakly so with foliar biomass a dry year, but not a wet year. This suggested that these variations are primarily related to induced differences in the balance between supply and demand for water. Hence, studies of {au13}C abundance can disentangle the role of water as such from its effects on mineralization of N and flow of N.