Soil acidification as caused by the nitrogen uptake pattern of Scots pine (Pinus sylvestris)

Three-year-old Scots pine (Pinus sylvestris) trees were grown on a sandy forest soil in pots, with the objective to determine their NH₄/NO₃ uptake ratio and proton efflux. N was supplied in three NH₄-N/NO₃-N ratios, 3:1, 1:1 and 1:3, either as ¹⁵NH₄+¹⁴NO₃ or as ¹⁴NH₄+¹⁵NO₃. Total N and ¹⁵N acquisition of different plant parts were measured. Averaged over the whole tree, the NH₄/NO₃ uptake ratios throughout the growing season were found to be 4.2, 2.5, and 1.5 for the three application ratios, respectively. The excess cation-over-anion uptake value (C_a-A_a) appeared to be linearly related to the natural logarithm of the NH₄/NO₃ uptake ratio. Further, this uptake ratio was related to the NH₄/NO₃ ratio of the soil solution. From these relationship it was estimated that Scots pine exhibits an acidifying uptake pattern as long as the contribution of nitrate to the N nutrition is lower than 70%. Under field circumstances root uptake may cause soil acidification in the topsoil, containing the largest part of the root system, and soil alkalization in deeper soil layers.     

Cyanobacterial inoculation and nitrogen fertilization in rice

During three rice-growing seasons in Uruguay, field experiments were conducted to study the contribution of cyanobacterial inoculation and chemical N fertilization to rice production. Neither grain yield nor fertilizer recovery by the plant were affected by inoculation with native cyanobacterial isolates. A low fertilizer use efficiency (around 20%) was observed when labelled (NH₄)₂SO₄ was applied at sowing. Recovery of applied ¹⁵N by the soil–plant system was 50%. Inoculation did not modify ¹⁵N uptake by the plant when the fertilizer was three-split applied either. The total N-fertilizer recovery was higher when the fertilizer was split than when applied in a single dose. Plant N-fertilizer uptake was higher when the fertilizer was applied at tillering. Uptake of ¹⁵N from cyanobacteria by rice was studied in a greenhouse pots experiment without chemical nitrogen addition. Recovery of ¹⁵N from labelled cyanobacteria by rice in greenhouse growth conditions was similar to that of partial recovery of (NH₄)₂SO₄ applied at sowing in the field. Cyanobacterial N mineralization under controlled conditions was fast as cyanobacterial N was detected in plants after 25 days. Moreover 40 days after inoculation non-planted and inoculated soil had more inorganic N than the non-inoculated one.     

Crop uptake and leaching of 15N applied in ruminant slurry with selectively labelled faeces and urine fractions

Two animal slurries either labelled with ¹⁵N in the urine or in the faeces fraction, were produced by feeding a sheep with unlabelled and ¹⁵N-labelled hay and collecting faeces and urine separately. The slurries were applied (12 g total N ^-2) to a coarse sand and a sandy loam soil confined in lysimeters and growing spring barley (Hordeum vulgare L). Reference lysimeters without slurry were supplied with¹⁵ NH₄ ¹⁵NO³ corresponding to the inorganic N applied with the slurries (6 g N m^-2). In the second year, all lysimeters received unlabelled mineral fertilizer (6 g N m^-2) and grew spring barley. N harvested in the two crops (grain + straw) and the loss of nitrate by leaching were determined. ¹⁵N in the urine fraction was less available for crop uptake than mineral fertilizer ¹⁵N. The first barley crop on the sandy loam removed 49% of the ¹⁵N applied in mineral fertilizer and 36% of that applied with urine. The availability of fertilizer ¹⁵N (36%) and urine¹⁵ N (32%) differed less on the coarse sand. Of the¹⁵ N added with the faeces fraction, 12–14% was taken up by the barley crop on the two soils. N mineralized from faeces compensated for the reduced availability of urine N providing a similar or higher crop N uptake in manured lysimeters compared with mineral fertilized ones.About half of the total N uptake in the first crop originated from the N applied either as slurry or mineral fertilizer. The remaining N was derived from the soil N pool. Substantially smaller but similar proportions of¹⁵ N from faeces, urine and fertilizer were found in the second crop. The similar recoveries indicated a slow mineralization rate of the residual faeces N since more faeces was left in the soil after the first crop.More N was lost by leaching from manured lysimeters but as a percentage of N applied, losses were similar to those from mineral fertilizer. During the first and second winter, 3–5% and 1–3%, respectively, of the ¹⁵N in slurry and mineral fertilizer was leached as nitrate. Thus slurry N applied in spring just before sowing did not appear to be more prone to loss by nitrate leaching than N given in mineral fertilizer. Slurry N accounted for a higher proportion of the N leached, however, because more N was added in this treatment.     

Effects of calcium and aluminium on nodulation,nitrogen fixation and growth of groundnut in solution culture

Uptake of NH₄ and NO₃ by above ground parts of beech trees was studied by spraying young trees with varying concentrations of ¹⁵N labeled solutions, different N-forms, and spray regimes over four months. Following treatment, the trees were harvested and analyzed for ¹⁵N and major element content. Throughfall was collected and analyzed in addition in order to study the interaction between nitrogen uptake and cation leaching. Significant amounts of N were taken up by the above ground plant parts in all treatments as indicated by ¹⁵N analysis of the trees as well as by throughfall measurements. NH₄ uptake exceeded the uptake of NO₃ if applied in the same concentration. Uptake of N increased linearly with increasing concentration in the spray solution and with spray intensity. The uptaken N was translocated within the plant. The contribution of N from uptake by above ground parts to the total N content of tissues differed and reached a maximum level of 6% in leaves. No effect of above ground N uptake on the total N content of tissues was found. Calculating atmospheric N inputs to forest ecosystems by throughfall measurements may underestimate the actual N input.     

Nitrogen losses in puddled soils as affected by timing of water deficit and nitrogen fertilization

Erratic rainfall in rainfed lowlands and inadequate water supply in irrigated lowlands can results in alternate soil drying and flooding during a rice (Oryza sativa L.) cropping period. Effects of alternate soil drying and flooding on N loss by nitrification-denitrification have been inconsistent in previous field research. To determine the effects of water deficit and urea timing on soil NO₃ and NH₄, floodwater NO₃, and N loss from added ¹⁵N-labeled urea, a field experiment was conducted for 2 yr on an Andaqueptic Haplaquoll in the Philippines. Water regimes were continuously flooded, not irrigated from 15 to 35 d after transplanting (DT), or not irrigated from 41 to 63 DT. The nitrogen treatments in factorial combination with water regimes were no applied N and 80 kg urea-N ha^–1, either applied half basally and half at 37 DT or half at 11 DT and half at 65 DT. Water deficit at 15 to 35 DT and 41 to 63 DT, compared with continuous soil flooding, significantly reduced extractable NH₄ in the top 30-cm soil layer and resulted in significant but small (<1.0 kg N ha^–1) soil NO₃ accumulations. Soil NO₃, which accumulated during the water deficit, rapidly disappeared after reflooding. Water deficit at 15 to 35 DT, unlike that at 41 to 63 DT, increased the gaseous loss of added urea N as determined from unrecovered ¹⁵N in ¹⁵N balances. The results indicate that application of urea to young rice in saturated or flooded soil results in large, rapid losses of N (mean = 35% of applied N), presumably by NH₃ volatilization. Subsequent soil drying and flooding during the vegetative growth phase can result in additional N loss (mean = 14% of applied N), presumably by nitrification-denitrification. This additional N loss due to soil drying and flooding decreases with increasing crop age, apparently because of increased competition by rice with soil microorganisms for NH₄ and NO₃.     

Fertilization rates and clay fixed ammonium in two Quebec soils

Clay fixed NH₄ ⁺ can provide a significant sink for fertilizer N, as well as a source of N for plant uptake. Knowledge or soil NH₄ ⁺ fixing capacity and release for crops is necessary to develop long-term fertilizer programs. Field experiments with corn (Zea mays L.) were carried out to investigate soil NH₄ ⁺ fixing capacity and subsequent release as influenced by fertilizer rates using ¹⁵N in a Ste. Rosalie clay (fine, mixed, frigid, Typic Humaquept) and a Chicot sandy clay loam (fine-loamy, mixed, frigid, Typic Hapludalf). With high N rates increased NH₄ ⁺ fixation occurred only in the Ste. Rosalie soil. At the end of the first growing season, fertilizer N recovery as clay fixed NH₄ ⁺ for high and normal rates of fertilizer in the Ste. Rosalie soil was 17.8% and 28.7%, respectively and the recovery for the high and normal rates in the Chicot soil was 4.6 and 10.5%, respectively. Significant amounts of clay fixed NH₄ ⁺-N were released in the soil profile in the second year after ¹⁵N application on the Chicot soil. Recently clay fixed fertilizer NH₄ ⁺N was released more rapidly than that of the native fixed NH₄ ⁺, from the surface layer of the Ste. Rosalie soil. The fertilizer fixed NH₄ ⁺ seems to be in a more labile N pool than the native fixed NH₄ ⁺-N in the Chicot soil.     

Isotopic fractionation accompanying fertilizer nitrogen transformations in soil and trees of a Scots spine ecosystem

Foliage from a mature stand of Scots pine (Pinus sylvestris L.) receiving increasing doses of ammonium nitrate and urea nitrogen was assayed during the five subsequent growing seasons for total N concentration and ¹⁵N abundance. The aim of the study was to examine the potential of the ¹⁵N technique to provide estimates on fertilizer N recovery and its fate in the ecosystem. The ¹⁵N abundance in the foliage increased in proportion to the dose of fertilizer application. This was generally owing to the fact that the ¹⁵N of the fertilizer N was significantly higher than that in the soil inorganic-N pool, as well as in the needle biomass of the Scots pine trees on the nonfertilized plots. Due to ¹⁵N isotope discrimination occurring during N transformations in soil the relationship was however not very close. Calculations based on the principle of isotope dilution yielded only rough and, in some cases, even misleading estimates of the fraction of the fertilizer-derived nitrogen (N_dff) in the needles. This was especially the case for the urea-N, which undergoes significant isotopic fractionation during the process of ammonia volatilization and possibly microbial NH₄ ⁺ assimilation in soil. Over five growing seasons, foliar total N concentration peaked at the end of the second season while the ¹⁵N abundance continued to increase. Although large methodological errors may be involved when interpreting natural ¹⁵N abundance, the measurement of ¹⁵N seems to provide semi-quantitative information about fertilizer N accumulation and transformation processes in coniferous ecosystems. A better understanding of the tree and soil processes causing isotopic fractionation is a prerequisite for correct interpretation of ¹⁵N data.     

The role of below-ground competition during early stages of secondary succession: the case of 3-year-old Scots pine (Pinus sylvestris L.) seedlings in an abandoned grassland

In abandoned or extensively managed grasslands, the mechanisms involved in pioneer tree species success are not fully explained. Resource competition among plants and microclimate modifications have been emphasised as possible mechanisms to explain variation of survivorship and growth. In this study, we evaluated a number of mechanisms that may lead to successful survival and growth of seedlings of a pioneer tree species (Pinus sylvestris) in a grass-dominated grassland. Three-year-old Scots pines were planted in an extensively managed grassland of the French Massif Central and for 2 years were either maintained in bare soil or subjected to aerial and below-ground interactions induced by grass vegetation. Soil temperatures were slightly higher in bare soil than under the grass vegetation, but not to an extent explaining pine growth differences. The tall grass canopy reduced light transmission by 77% at ground level and by 20% in the upper part of Scots pine seedlings. Grass vegetation presence also significantly decreased soil volumetric water content (Hv) and soil nitrate in spring and in summer. In these conditions, the average tree height was reduced by 5% compared to trees grown in bare soil, and plant biomass was reduced by 85%. Scots pine intrinsic water-use efficiency (A/g), measured by leaf gas-exchange, increased when Hv decreased owing to a rapid decline of stomatal conductance (g). This result was also confirmed by δ ¹³C analyses of needles. A summer ¹⁵N labelling of seedlings and grass vegetation confirmed the higher NO₃ capture capacity of grass vegetation in comparison with Scots pine seedlings. Our results provide evidence that the seedlings' success was linked to tolerance of below-ground resource depletion (particularly water) induced by grass vegetation based on morphological and physiological plasticity as well as to resource conservation.     

The partitioning of fertilizer-N between soil and crop: Comparison of ammonium and nitrate applications

Field experiments were carried out in 1987 on winter wheat crops grown on three types of soil. ¹⁵N-labelled urea, ¹⁵NH₄NO₃ or NH₄ ¹⁵NO₃ (80 kg N ha^-1) was applied at tillering. The soils (chalky soil, hydromorphic loamy soil, sandy clay soil) were chosen to obtain a range of nitrogen dynamics, particularly nitrification. Soil microbial N immobilization and crop N uptake were measured at five dates. Shortly after fertilizer application (0–26 days), the amount of N immobilized in soil were markedly higher with labelled urea or ammonium than that with nitrate in all soils. During the same period, crop ¹⁵N uptake occurred preferentially at the expense of nitrate. Nitrification differed little between soils, the rates were 2.0 to 4.7 kg N ha^-1 day^-1 at 9°C daily mean temperature. The differences in immobilization and uptake had almost disappeared at flowering and harvest. ¹⁵N recovery in soil and crop varied between 50 and 100%. Gaseous losses probably occurred by volatilization in the chalky soil and denitrification in the hydromorphic loamy soil. These losses affected the NH₄ ⁺ and NO₃ ^- pools differently and determined the partitioning of fertilizer-N between immobilization and absorption.     

Fertilizer and soil nitrogen accumulation by irrigated barley grown on a red-brown earth in south-eastern Australia

¹⁵N labelled (NH₄)₂SO₄ was applied to barley at 5 g N m⁻² (50 kg N ha⁻¹) in microplots at sowing to study the timing of the N losses and the contribution of soil and fertilizer N to the plant. Water treatments included rainfed and irrigation at 45–50 mm deficit beginning in the spring. Recovery of¹⁵N in the plant increased to a maximum of about 20% within 91 days after sowing (DAS 91) and then remained constant. Approximately 16% (0.8 g N m⁻²) of the fertilizer was in the stem and leaves at DAS 91 and this N was subsequently redistributed to the head. At maturity, approximately 75% of the¹⁵N assimilated by the tops was recovered in the grain. Soil N contributed 3.6 g N m⁻² to the head; 2.2 g N m⁻² was remobilized from the stem and leaves, and the balance, approximately 1.4 g N m⁻², was taken up from the soil between DAS 69 to 91. Effects of irrigation treatments on N accumulation were not significant. Residual¹⁵N fertilizer in the soil decreased with time from sowing, and at maturity 40% of the applied N was recovered in the surface 0.15 m.¹⁵N movement to depth was limited and less than 5% of the fertilizer was recovered below 0.15 m. Irrigation had no effect on the¹⁵N recovery at depth. Total recovery of the¹⁵N varied between 60 and 67% and implies that 33–40% was lost from the soil-plant system. The total recovery in the soil and plant was not affected by time or irrigation in the interval DAS 39 to 134. Losses occurred before DAS 39 when crop uptake of N was small and soil mineral N content was high. There was an apparent loss of 1.9 g fertilizer N m⁻² (i.e. 38% of that applied) between DAS 1 and 15. This loss occurred before crop emergence when rainfall provided conditions suitable for denitrification.     

Fertilizer nitrogen budget of Na15NO3 and (15NH4)2SO4 split-applied to winter wheat in microplots on a loam soil

Labelled fertilizer N applied to winter wheat as Na¹⁵NO₃ and (¹⁵NH₄)₂SO₄ at a total N dressing of 100kg ha⁻¹ was used in a microplot balance study to investigate the fate of each split fraction at three growth stages: end of tillering, heading and beginning of flowering. Results indicated that while the percentage utilization of the applied N by the grain and total crop increased considerably from the first to the third split application, these values diminished steadily in the straw. Grain recovery values for the first, second and third split applications were 34.2%, 51.5% and 55.7% for the NO₃ and 32.3%, 48.4% and 52.5% for the NH₄ carrier, respectively. The corresponding recovery values for the whole plant were 54.6%, 67.8% and 69.9% for the NO₃ and 51.7%, 63.5% and 66.1% for the NH₄ carrier. A greater proportion of the fertilizer N applied at the end of tillering stage was found in the vegetative plant components as compared with the grain. The reverse occurred for the N applied at the heading and at the beginning of the flowering stages. The residual fertilizer N found in the soil amounted to 18.0%, 10.4% and 11.6% of the applied NO₃−N and to 22.5%, 12.7% and 15.2% of the applied NH₄−N for the respective split applications. No differences were found for each split application between the two carriers as far as the unaccounted fertilizer N was concerned. The losses were 26.6%, 22.3% and 18.6% of the applied N for the three split applications, respectively. The application of fertilizer N did not lead to any increase in soil N uptake by the crop.     

Mineralization-immobilization and plant uptake of nitrogen as influenced by the spatial distribution of cattle slurry in soils of different texture

The effect of incorporating cattle slurry in soil, either by mixing or by simulated injection into a hollow in soil, on the ryegrass uptake of total N and ¹⁵NH₄ ⁺-N was determined in three soils of different texture. The N accumulation in Italian ryegrass (Lolium multiflorum L.) from slurry N and from an equivalent amount of NH₄ ⁺-N in (¹⁵NH₄) SO₄ (control) was measured during 6 months of growth in pots. After this period the total recovery of labelled N in the top soil plus herbage was similar in the slurry and the control treatments. This indicated that gaseous losses from slurry NH₄ ⁺-N were insignificant. Consequently, the availability of slurry N to plants was mainly influenced by the mineralization-immobilization processes. The apparent utilization of slurry NH₄ ⁺-N mixed into soil was 7%, 14% and 24% lower than the utilization of (NH₄)₂SO₄-N in a sand soil, a sandy loam soil and a loam soil, respectively. Thus, the net immobilization of N due to slurry application increased with increasing soil clay content, whereas the recovery in plants of ¹⁵N-labelled NH₄ ⁺-N from slurry was similar on the three soils. A parallel incubation experiment showed that the immobilization of slurry N occurred within the first week after slurry application. The incorporation of slurry N by simulated injection increased the plant uptake of both total and labelled N compared to mixing the slurry into the soil. The apparent utilization of injected slurry NH₄ ⁺-N was 7% higher, 8% lower and 4% higher than the utilization of (NH₄)₂SO₄-N in the sand, the sandy loam and the loam soil, respectively. It is concluded that the spatial distribution of slurry in soil influenced the net mineralization of N to the same degree as did the soil type.     

Effects of varied soil nitrogen supply on Norway spruce (Picea abies [L.] Karst.)

During a seven-month period the effect of different nitrogen (N) availability in soil on growth and nutrient uptake was studied in three-year-old Norway spruce (Picea abies [L.] Karst.) trees. The plants were grown in pots on N-poor forest soil supplied with various amounts and forms (inorganic and organic) of N. Increasing supply of inorganic N (as NH₄NO₃) increased the formation of new shoots and shoot dry weight. The root/shoot dry weight ratio of new growth was drastically decreased from 1.6 in plants without N supply to 0.5 in plants supplied with high levels of NH₄NO₃. This decrease in root/shoot dry weight ratio was associated with distinct changes in root morphology in favour of shorter and thicker roots. The addition of keratin as organic N source did neither affect growth nor root morphology of the trees. The amount of N taken up by plants was closely related to the supply of inorganic N, and trees supplied with highest levels of NH₄NO₃ also had the highest N contents in the dry matter of needles and roots. In contrast, N contents in needles of trees grown without additional N, or with keratin supply, were in the deficiency range. Supply of NH₄NO₃ decreased the contents of phosphate (P) and potassium (K) and therefore markedly increased N/P and N/K ratios in the needles. On the other hand, the contents of calcium (Ca), magnesium (Mg), and manganese (Mn) in the needles were increased in the plants supplied with inorganic N, suggesting high soil availability and promotion of uptake of these divalent cations by high nitrate uptake. The observed effects on root/shoot dry weight ratio, root morphology, and mineral nutrient composition of the needles indicated that high inorganic N supply may increase above-ground productivity but at the same time decrease the tolerance of trees against soil-borne (e.g. deficiency of other mineral nutrients) stress factors. Deceased 21 September 1996 Deceased 21 September 1996     

Functional role of DNRA and nitrite reduction in a pristine south Chilean <Emphasis Type="Italic">Nothofagus</Emphasis> forest

Nitrite (NO₂ ⁻) is an intermediate in a variety of soil N cycling processes. However, NO₂ ⁻ dynamics are often not included in studies that explore the N cycle in soil. Within the presented study, nitrite dynamics were investigated in a Nothofagus betuloides forest on an Andisol in southern Chile. We carried out a ¹⁵N tracing study with six ¹⁵N labeling treatments, including combinations of NO₃ ⁻, NH₄ ⁺ and NO₂ ⁻. Gross N transformation rates were quantified with a ¹⁵N tracing model in combination with a Markov chain Monte Carlo optimization routine. Our results indicate the occurrence of functional links between (1) NH₄ ⁺ oxidation, the main process for NO₂ ⁻ production (nitritation), and NO₂ ⁻ reduction, and (2) oxidation of soil organic N, the dominant NO₃ ⁻ production process in this soil, and dissimilatory NO₃ ⁻ reduction to NH₄ ⁺ (DNRA). The production of NH₄ ⁺ via DNRA was approximately ten times higher than direct mineralization from recalcitrant soil organic matter. Moreover, the rate of DNRA was several magnitudes higher than the rate of other NO₃ ⁻ reducing processes, indicating that DNRA is able to outcompete denitrification, which is most likely not an important process in this ecosystem. These functional links are most likely adaptations of the microbial community to the prevailing pedo-climatic conditions of this Nothofagus ecosystem.     

Land-use change effects on soil C and N transformations in soils of high N status: comparisons under indigenous forest, pasture and pine plantation

Globally, land-use change is occurring rapidly, and impacts on biogeochemical cycling may be influenced by previous land uses. We examined differences in soil C and N cycling during long-term laboratory incubations for the following land-use sequence: indigenous forest (soil age = 1800 yr); 70-year-old pasture planted after forest clearance; 22-year-old pine (Pinus radiata) planted into pasture. No N fertilizer had been applied but the pasture contained N-fixing legumes. The sites were adjacent and received 3–6 kg ha^–1 yr^–1volcanic N in rain; NO₃ ^–-N leaching losses to streamwater were 5–21 kg ha^–1 yr^–1, and followed the order forest < pasture = pine. Soil C concentration in 0–10 cm mineral soil followed the order: pasture > pine = forest, and total N: pasture > pine > forest. Nitrogen mineralization followed the order: pasture > pine > forest for mineral soil, and was weakly related to C mineralization. Based on radiocarbon data, the indigenous forest 0–10 cm soil contained more pre-bomb C than the other soils, partly as a result of microbial processing of recent C in the surface litter layer. Heterotrophic activity appeared to be somewhat N limited in the indigenous forest soil, and gross nitrification was delayed. In contrast, the pasture soil was rich in labile N arising from N fixation by clover, and net nitrification occurred readily. Gross N cycling rates in the pine mineral soil (per unit N) were similar to those under pasture, reflecting the legacy of N inputs by the previous pasture. Change in land use from indigenous forest to pasture and pine resulted in increased gross nitrification, net nitrification and thence leaching of NO₃ ^–-N.     

Site-dependent N uptake from N-form mixtures by arctic plants,soil microbes and ectomycorrhizal fungi

Soil microbes constitute an important control on nitrogen (N) turnover and retention in arctic ecosystems where N availability is the main constraint on primary production. Ectomycorrhizal (ECM) symbioses may facilitate plant competition for the specific N pools available in various arctic ecosystems. We report here our study on the N uptake patterns of coexisting plants and microbes at two tundra sites with contrasting dominance of the circumpolar ECM shrub Betula nana. We added equimolar mixtures of glycine-N, NH₄⁺–N and NO₃⁻–N, with one N form labelled with ¹⁵N at a time, and in the case of glycine, also labelled with ¹³C, either directly to the soil or to ECM fungal ingrowth bags. After 2 days, the vegetation contained 5.6, 7.7 and 9.1% (heath tundra) and 7.1, 14.3 and 12.5% (shrub tundra) of the glycine-, NH₄⁺- and NO₃⁻–¹⁵N, respectively, recovered in the plant–soil system, and the major part of ¹⁵N in the soil was immobilized by microbes (chloroform fumigation-extraction). In the subsequent 24 days, microbial N turnover transferred about half of the immobilized ¹⁵N to the non-extractable soil organic N pool, demonstrating that soil microbes played a major role in N turnover and retention in both tundra types. The ECM mycelial communities at the two tundras differed in N-form preferences, with a higher contribution of glycine to total N uptake at the heath tundra; however, the ECM mycelial communities at both sites strongly discriminated against NO₃⁻. Betula nana did not directly reflect ECM mycelial N uptake, and we conclude that N uptake by ECM plants is modulated by the N uptake patterns of both fungal and plant components of the symbiosis and by competitive interactions in the soil. Our field study furthermore showed that intact free amino acids are potentially important N sources for arctic ECM fungi and plants as well as for soil microorganisms. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Changes in Canopy Processes Following Whole-Forest Canopy Nitrogen Fertilization of a Mature Spruce-Hemlock Forest

Abstract Most experimental additions of nitrogen to forest ecosystems apply the N to the forest floor, bypassing important processes taking place in the canopy, including canopy retention of N and/or conversion of N from one form to another. To quantify these processes, we carried out a large-scale experiment and determined the fate of nitrogen applied directly to a mature coniferous forest canopy in central Maine (18–20 kg N ha⁻¹ y⁻¹ as NH₄NO₃ applied as a mist using a helicopter). In 2003 and 2004 we measured NO₃ ⁻, NH₄ ⁺, and total dissolved N (TDN) in canopy throughfall (TF) and stemflow (SF) events after each of two growing season applications. Dissolved organic N (DON) was greater than 80% of the TDN under ambient inputs; however NO₃ ⁻ accounted for more than 50% of TF N in the treated plots, followed by NH₄ ⁺ (35%) and DON (15%). Although NO₃ ⁻ was slightly more efficiently retained by the canopy under ambient inputs, canopy retention of NH₄ ⁺as a percent of inputs increased markedly under fertilization. Recovery of less than 30% of the fertilizer N in TF suggested that the forest canopy retained more than 70% of the applied N (>80% when corrected for N which bypassed tree surfaces at the time of fertilizer addition). Results from plots receiving ¹⁵N enriched NO₃ ⁻ and NH₄ ⁺ confirmed bulk N estimations that more NO₃ ⁻ than NH₄ ⁺ was washed from the canopy by wet deposition. The isotope data did not show evidence of canopy nitrification, as has been reported in other spruce forests receiving much higher N inputs. Conversions of fertilizer-N to DON were observed in TF for both ¹⁵NH₄ ⁺ and ¹⁵NO₃ ⁻ additions, and occurred within days of the application. Subsequent rain events were not significantly enriched in ¹⁵N, suggesting that canopy DON formation was a rapid process related to recent N inputs to the canopy. We speculate that DON may arise from lichen and/or microbial N cycling rather than assimilation and re-release by tree tissues in this forest. Canopy retention of experimentally added N may meet and exceed calculated annual forest tree demand, although we do not know what fraction of retained N was actually physiologically assimilated by the plants. The observed retention and transformation of DIN within the canopy demonstrate that the fate and ecosystem consequences of N inputs from atmospheric deposition are likely influenced by forest canopy processes, which should be considered in N addition studies. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Fate of nitrogen fertilizer applied on two main arables crops,winter wheat (Triticum aestivum) and sugar beet (Beta vulgaris) in the loam region of Belgium

Since 1986, the fate of fertilizer N (NH₄NO₃ or NaNO₃) applied in field conditions on two main arable crops, winter wheat (Triticum aestivum) and sugar beet (Beta vulgaris), has been studied using ¹⁵N. Up to a rate of 200 kg ha^-1 of N, mean recovery of fertilizer by winter wheat was 70%, provided it had been split applied. Single application (with or without dicyandiamid) was less effective. For sugar beet, in 1990, 1991 and 1992, 40% of fertilizer N was found in the crop at harvest when NH₄NO₃ had been broadcast at 100 to 160 kg N ha^-1 at sowing time. For the same N rate, recovery was 50% when row applied near the seeds and 60% for 80 kg N ha^-1. For the two experimental crops, residual fertilizer N in soil was exclusively organic. It ranged from 15 to 30% of applied N and was located in the 30 cm upper layer. Losses were generally lower with winter wheat (12%) than with sugar beet (20–40%) and could be ascribed to volatilization and denitrification. Soil derived N taken up by the plant was site and year dependent.     

Abiotic immobilization of nitrate in two soils of relic <Emphasis Type="Italic">Abies pinsapo</Emphasis>-fir forests under Mediterranean climate

Evidence for abiotic immobilization of nitrogen (N) in soil is accumulating, but remains controversial. Identifying the fate of N from atmospheric deposition is important for understanding the N cycle of forest ecosystems. We studied soils of two Abies pinsapo fir forests under Mediterranean climate seasonality in southern Spain—one with low N availability and the other with symptoms of N saturation. We hypothesized that biotic and abiotic immobilization of nitrate (NO₃ ⁻) would be lower in soils under these forests compared to more mesic temperate forests, and that the N saturated stand would have the lowest rates of NO₃ ⁻ immobilization. Live and autoclaved soils were incubated with added ¹⁵NO₃ ⁻ (10 μg N g⁻¹ dry soil; 99% enriched) for 24 h, and the label was recovered as total dissolved-N, NO₃ ⁻, ammonium (NH₄ ⁺), or dissolved organic-N (DON). To evaluate concerns about possible iron interference in analysis of NO₃ ⁻ concentrations, both flow injection analysis (FIA) and ion chromatography (IC) were applied to water extracts, soluble iron was measured in both water and salt extracts, and standard additions of NO₃ ⁻ to salt extracts were analyzed. Good agreement between FIA and IC analysis, low concentrations of soluble Fe, and 100% (±3%) recovery of NO₃ ⁻ standard additions all pointed to absence of an interference problem for NO₃ ⁻ quantification. On average, 85% of the added ¹⁵NO₃ ⁻ label was recovered as ¹⁵NO₃ ⁻, which supports our hypothesis that rates of immobilization were generally low in these soils. A small amount (mean = 0.06 μg N g⁻¹ dry soil) was recovered as ¹⁵NH₄ ⁺ in live soils and none in sterilized soils. Mean recovery as DO¹⁵N ranged from 0.6 to 1.5 μg N g⁻¹ dry soil, with no statistically significant effect of sterilization or soil type, indicating that this was an abiotic process that occurred at similar rates in both soils. These results demonstrate a detectable, but modest rate of abiotic immobilization of NO₃ ⁻ to DON, supporting our first hypothesis. These mineral soils may not have adequate carbon availability to support the regeneration of reducing microsites needed for high rates of NO₃ ⁻ reduction. Our second hypothesis regarding lower expected abiotic immobilization in soils from the N-saturated site was not supported. The rates of N deposition in this region may not be high enough to have swamped the capacity for soil NO₃ ⁻ immobilization, even in the stand showing some symptoms of N saturation. A growing body of evidence suggests that soil abiotic NO₃ ⁻ immobilization is common, but that rates are influenced by a combination of factors, including the presence of plentiful available carbon, reduced minerals in anaerobic microsites and adequate NO₃ ⁻ supply.     

Pertilization and nutrition experiments with conifer seedlings in pots

Summary In conifer fertilization and nutrition experimentsPinus halepensis, P. radiata andP. maritima seedlings were grown in pots, filled with soil derived from mica schist and siliceous tertiary deposits and also in peat substrate in paperpots.Fertilization with P ofP. radiata andP. maritima seedlings growing in soil low in available P and N improved seedling height only in combination with N fertilization and fertilization with alone induced P deficiency symptoms. N fertilization with from 100 to 150 ppm (2.4 to 3.2 g N/kg, respectively) in the soil regardless of the form of N (NH₄ ⁺ or NO₃ ^–) applied in the summer or autumn together with application of 20 ppm P before sowing was the fertilization regime which produced the best seedlings.Fertilization of peat before sowingP. halepensis, P. radiata andP. maritima with omission of one of the nutrients N, P and K resulted in visible symptoms of N, P and K deficiency, respectively, in the seedlings. Comparative chemical analysis of needles from the three kinds of conifer seedlings with deficiency symptoms and healthy ones verified the visual symptoms of N and P deficiency but not so convincingly the K deficiency symptoms.