The effect of harvesting intensity on the fate of applied 15N-ammonium to the organic horizons of a coniferous forest in N. Wales

¹⁵N-ammonium sulphate equivalent to 0.5 kg N/ha was added as a tracer to lysimeters containing the organic horizons of an acid forest soil. The effect of logging debris (brash), vegetation and second rotationPicea sitchensis seedlings on the amount of the¹⁵N found in various soil, vegetation and leachate pools was followed over a period of 60 days. Transformation of¹⁵N-ammonium to nitrate occurred within 24 hours. Although total nitrate leachate losses were high, tracer-derived nitrate represented only 0.4%–4.2% of the applied¹⁵N-ammonium. The atom % excess of the KCI-extractable organic-N pool was initially lower than for the inorganic species but due to the large pool size, consistently represented 3–6% of the applied¹⁵N-ammonium. The similarity of the atom % excess of the ammonium and nitrate pools indicated an autotrophic nitrification pathway.A significant proportion of the¹⁵N-ammonium passed through the microbial biomass which contained between 16 and 48% of the¹⁵N-ammonium 2 days after addition of the¹⁵N-ammonium. This nitrogen was in a readily available form or short-term pool for the first two weeks (with no change in the overall biomass pool), after which the nitrogen appeared to become transformed into more stable compounds representing a long-term pool. Total recovery of the¹⁵N was between 68% and 99% for the different treatments. The presence of brash reduced microbial immobilisation of the¹⁵N-ammonium and total retention in the organic matter. This is suggested to be a consequence of greater nitrification and denitrificatiion rate in organic horizons beneath a brash covering due to different microclimatic conditions.     

Turnover of residual 15N-labelled fertilizer N in soil following harvest of oilseed rape (t Brassica napus L.)

We studied the fate of ¹⁵N-labelled fertilizer nitrogen in a sandy loam soil after harvest of winter oilseed rape (Brassica napus L. cv. Ceres) given 100 or 200 kg N ha^-1 in spring, with or without irrigation. Our main objective was to quantify the temporal variations of the soil mineral N, the extractable soil organic N and soil microbial biomass N, and fertilizer derived N in these pools during autumn and winter. Nitrogen use efficiency of the oilseed rape crop varied from 47% of applied N in the 100N, irrigated treatment to 34% in the 200N, non-irrigated treatment. However, only in the latter treatment did we find significantly higher fertilizer derived soil mineral N than in the three other treatments which all had low soil mineral N contents at the first sampling after harvest (8 days after stubble tillage). Between 31% and 42% of the applied N could not be accounted for in the harvested plants or 0-15 cm soil layer at this first sampling. Over the following autumn and winter none of the remaining fertilizer derived soil N was lost from the 0–5 cm depth, but from the 5–15 cm depth a marked proportion of N derived from fertilizer was lost, probably by leaching. Negligible amounts of fertilizer derived extractable soil organic and mineral N (<1 kg N ha^-1, 0-15 cm) were found in all treatments after the first sampling.Soil microbial biomass N was not significantly affected by treatments and showed only small temporal variability (±11% of the mean 76 kg N ha^-1, 0- 15 cm depth). Surprisingly, the average amount of soil microbial biomass N derived from fertilizer was significantly affected by the treatments, with the extremes being 5.5 and 3.1 kg N ha^-1 in the 200N, non-irrigated and 100N, irrigated treatments, respectively. Also, the estimated exponential decay rate of microbial biomass N derived from fertilizer, differed greatly (2 fold) between these two treatments, indicating highly different microbial turnover rates in spite of the similar total microbial biomass N values. In studies utilising ¹⁵N labelling to estimate turnover rates of different soil organic matter pools this finding is of great importance, because it may question the assumption that turnover rates are not affected by the insertion of the label.     

Effects of atmospheric CO2 enrichment on δ 13C, δ 15N values and turnover times of soil organic matter pools isolated by thermal techniques

CO₂ applied for Free-Air CO₂ Enrichment (FACE) experiments is strongly depleted in ¹³C and thus provides an opportunity to study C turnover in soil organic matter (SOM) based on its δ ¹³C value. Simultaneous use of ¹⁵N labeled fertilizers allows N turnover to be studied. Various SOM fractionation approaches (fractionation by density, particle size, chemical extractability etc.) have been applied to estimate C and N turnover rates in SOM pools. The thermal stability of SOM coupled with C and N isotopic analyses has never been studied in experiments with FACE. We tested the hypothesis that the mean residence time (MRT) of SOM pools is inversely proportional to its thermal stability. Soil samples from FACE plots under ambient (380 ppm) and elevated CO₂ (540 ppm; for 3 years) treatments were analyzed by thermogravimetry coupled with differential scanning calorimetry (TG-DSC). Based on differential weight losses (TG) and energy release or consumption (DSC), five SOM pools were distinguished. Soil samples were heated up to the respective temperature and the remaining soil was analyzed for δ ¹³C and δ ¹⁵N by IRMS. Energy consumption and mass losses in the temperature range 20–200°C were mainly connected with water volatilization. The maximum weight losses occurred from 200–310°C. This pool contained the largest amount of carbon: 61% of the total soil organic carbon in soil under ambient treatment and 63% in soil under elevated CO₂, respectively. δ ¹³C values of SOM pools under elevated CO₂ treatment showed an increase from −34.3‰ of the pool decomposed between 20–200°C to −18.1‰ above 480°C. The incorporation of new C and N into SOM pools was not inversely proportional to its thermal stability. SOM pools that decomposed between 20–200 and 200–310°C contained 2 and 3% of the new C, with a MRT of 149 and 92 years, respectively. The pool decomposed between 310–400°C contained the largest proportion of new C (22%), with a MRT of 12 years. The amount of fertilizer-derived N after 2 years of application in ambient and elevated CO₂ treatments was not significantly different in SOM pools decomposed up to 480°C having MRT of about 60 years. In contrast, the pool decomposed above 480°C contained only 0.5% of new N, with a MRT of more than 400 years in soils under both treatments. Thus, the separation of SOM based on its thermal stability was not sufficient to reveal pools with contrasting turnover rates of C and N. Responsible Editor: Bernard Nicolardot.     

Light affects competition for inorganic and organic nitrogen between maize and rhizosphere microorganisms

Effects of light on the short term competition for organic and inorganic nitrogen between maize and rhizosphere microorganisms were investigated using a mixture of amino acid, ammonium and nitrate under controlled conditions. The amount and forms of N added in the three treatments was identical, but only one of the three N forms was labeled with ¹⁵N. Glycine was additionally labeled with ¹⁴C to prove its uptake by maize and incorporation into microbial biomass in an intact form. Maize out-competed microorganisms for during the whole experiment under low and high light intensity. Microbial uptake of ¹⁵N and ¹⁴C was not directly influenced by the light intensity, but was indirectly related to the impact the light intensity had on the plant. More was recovered in microbial biomass than in plants in the initial 4 h under the two light intensities, although more ¹⁵N-glycine was incorporated into microbial biomass than in plants in the initial 4 h under low light intensity. Light had a significant effect on uptake by maize, but no significant effects on the uptake of or ¹⁵N-glycine. High light intensity significantly increased plant uptake of and glycine ¹⁴C. Based on ¹⁴C to ¹⁵N recovery ratios of plants, intact glycine contributed at least 13% to glycine-derived nitrogen 4 h after tracer additions, but it contributed only 0.5% to total nitrogen uptake. These findings suggest that light intensity alters the competitive relationship between maize roots and rhizosphere microorganisms and that C4 cereals such as maize are able to access small amounts of intact glycine. We conclude that roots were stronger competitor than microorganisms for inorganic N, but microorganisms out competed plants during a short period for organic N, which was mineralized into inorganic N within a few hours of application to the soil and was thereafter available for root uptake.     

Recruitment preferences of blue mussel spat (Mytilus edulis) for different substrata and microhabitats in the White Sea (Russia)

We adapted the chloroform fumigation method to determine microbial nitrogen (N) and microbial incorporation of ¹⁵N on three common substrates [leaves, wood and fine benthic organic matter (FBOM)] in three forest streams. We compared microbial N and ¹⁵N content of samples collected during a 6-week ¹⁵N–NH₄ tracer addition in each stream. The ¹⁵N was added during late autumn to Upper Ball Creek, a second-order stream at the Coweeta Hydrologic Lab, North Carolina, U.S.A.; during spring to Walker Branch, a first-order stream on DOE's Oak Ridge National Environmental Research Park, Tennessee; and during summer to Bear Brook, a first-order stream in the Hubbard Brook Experimental Forest, New Hampshire. FBOM was the largest component of organic matter and N standing stock in all streams. Microbial N represented the highest proportion of total N in leaves and least in FBOM in Walker Branch and Bear Brook. In Upper Ball Creek, the proportion of microbial N was higher in FBOM than in used biofilm or on leaves. Standing stock of microbial N on leaves and in FBOM ranged from 37 mg N m^–2 in Bear Brook to 301 mg N m^–2 in Walker Branch. Percent of detrital N in living microbial cells was directly related to total microbial biomass (fungal and bacterial biomass) determined from microscopic counts. ¹⁵N values for microbes were generally higher than for bulk detritus, which would result in higher ¹⁵N values for animals preferentially consuming or assimilating microbial cells. The proportion of ¹⁵N taken up by detritus during the ¹⁵N experiments that remained in microbial cells by the end of the experiments was highest for wood biofilm in Upper Ball Creek (69%), leaves in Walker Branch (65%) and FBOM in Upper Ball Creek (31%). Lower retention proportions (<1–25%) were observed for other substrates. Our results suggest that microbial cells associated with leaves and wood biofilm were most active in ¹⁵N–NH₄ immobilization, whereas microbial cells associated with FBOM immobilized little ¹⁵N from stream water.     

Root contribution to nitrate reduction in barley seedlings (Hordeum vulgare L.)

Summary The leaf and root nitrate reductase activities were measured in 7 day-old barley seedlings by anoxic nitrite accumulation in darkness, during 48h after the transfer from a N-starved medium to a 1.5 mM K¹⁵NO₃ medium. Thisin situ nitrate reduction was compared with the¹⁵N incorporation in the reduced N fraction of the whole seedlings.The nitrate reduction integrated fromin situ measurements was lower than the reduced¹⁵N accumulation. The rootin situ nitrate reductase activity seemed to account for only the third of the real root nitrate reduction, which may have been responsible for the overall underestimation. This discrepancy was partly explained by the ability of the root to reduce nitrite in an anoxic environment.These results suggest that, after correction of thein situ estimation of the nitrate reduction. the roots contribute to about 50% of the total assimilation.     

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.     

Ammonia Assimilation in the Roots of Nitrate- and Ammonia-Grown Hordeum Vulgare (cv Golden Promise)

¹⁵N kinetic labeling studies were performed on seedlings of Hordeum vulgare L. var. Golden Promise growing under steady state conditions. Patterns of label incorporation in the pools of nitrogen compounds of roots fed [¹⁵N]ammonium were compared with computer-simulated labeling curves. The data were found to be quantitatively consistent with a three-compartment model in which ammonium is assimilated solely into the amide-N of glutamine. Labeling data from roots fed [¹⁵N]nitrate were also found to be at least qualitatively consistent with the assimilation of ammonia into glutamine. Methionine sulfoximine almost completely blocked the incorporation of ¹⁵N label into the amino acid pools of barley roots fed [¹⁵N]nitrate. These observations suggest that ammonia assimilation occurs solely via the glutamine synthetase/glutamate synthase pathway in both nitrate- and ammonia-grown barley roots.     

Nitrate uptake and storage in the seaweed Ulva rigida C. Agardh in relation to nitrate availability and thallus nitrate content in a eutrophic coastal lagoon (Sacca di Goro, Po River Delta, Italy)

The seasonal cycle of biomass and tissue composition of Ulva rigida C. Agardh, in relation to nitrogen availability in the water column, was studied in 1991-1992 in the Sacca di Goro, a highly eutrophic lagoon in the Po River Delta (Italy). Nitrate uptake rates and storage capacity were also determined in laboratory experiments. The seasonal growth of U. rigida was related to the seasonal trend of nitrogen concentration in the water column. U. rigida biomass increased exponentially during spring and attained peaks of about 300-400 g dry mass (DM) m⁻² in June. As biomass increased, U. rigida depleted nitrate in the water column. Thallus nitrate reserves also declined from 100 μmol N (g DM)⁻¹ to almost undetectable levels, and total thallus nitrogen declined from 4% to 2.5% DM and 1.25% DM in 1991 and 1992, respectively. During summer, U. rigida decomposition increased, and organic nitrogen concentrations in the water column increased. The uptake experiments demonstrated an inverse relationship between thallus nitrate content and nitrate uptake rates. A modified Michaelis-Menten equation that accounts for thallus nitrate fit the uptake data well. U. rigida can accumulate up to about 400-500 μmol nitrate (g DM)⁻¹ in cellular reserves. U. rigida in the Sacca di Goro has higher K_m and lower V_max/K_m ratios for nitrate uptake than other chlorophycean species, indicating a low efficiency of uptake at low nitrate concentrations. This low uptake efficiency, and the ability to exploit N availability by storing cellular nitrate pools in excess of immediate growth needs, may represent a physiological response to an eutrophic environment where nitrate is in large supply for most of the year.     

天山林区不同类型群落土壤氮素对冻融过程的动态响应

季节性冻融过程对北方温带森林土壤氮素的转化与流失具有重要影响,但不同类型群落对冻融过程响应的差异尚不明确。通过在林地、草地、灌丛上设置系列监测样地,采用原位培养的方法,利用林冠遮挡形成的自然雪被厚度差异,监测分析了冻融期天山林区不同群落表层土壤(0—15 cm)的氮素动态及净氮矿化速率间的差异。结果表明:(1)不同类型群落土壤的铵态氮(NH+4-N)含量、微生物量氮(MBN)含量基本与土壤(5 cm)温度呈正相关,深冻期林地土壤铵态氮含量低于其他群落类型而硝态氮含量高于其他群落类型;(2)硝态氮(NO-3-N)为天山林区季节性冻融期间土壤矿质氮的主体,占比达78.4%。灌丛土壤硝态氮流失风险较大,融化末期较融化初期灌丛土壤硝态氮含量下降了64.6%;(3)冻融时期对整体氮素矿化速率影响显著,群落类型对氨化速率影响显著;(4)天山林区土壤氮素在冻结期主要以氮固持为主。通过揭示不同类型群落土壤氮素对冻融格局的响应,能够助益于对北方林区冬季土壤氮素循环的认识。     

Dilution of 15N in Dunaliella tertiolecta by uptake of an unidentified compound following nitrate exhaustion

Nitrate (about 20 μM) was added as ¹⁵NO₃ to a nitrate-limited continuous culture of Dunaliella tertiolecta at steady-state. Nitrate uptake was then estimated from the decrease in nitrate in the medium, the incorporation of ¹⁵N into cells, and the increase in cellular nitrogen. Although the overall nitrogen budget over 5 h was balanced, there were large differences in estimates (up to a factor of five) of nitrate assimilation by the three methods on shorter time scale. After nitrate was exhausted from the medium, cellular nitrogen continued to increase while the ¹⁵N content of the particulate matter decreased over the next 1.5 h. This indicated that an unidentified, unlabelled nitrogen form, which was neither nitrite, ammonium nor dissolved free amino acids, was being taken up by the cells, at rates comparable to those of nitrate. This phenomenon leads to an underestimation of new biomass production when assessed through ¹⁵N incorporation into cells.     

Partitioning of 15N-labeled ammonium and nitrate among soil, litter, below- and above-ground biomass of trees and understory in a 15-year-old Picea abies plantation

The partitioning of nitrogen deposition among soil, litter, below- and above-ground biomass of trees and understory vegetation was investigated in a 15-year-old Picea abies (L.) Karst. plantation in the Fichtelgebirge, Germany, by labeling with 62 mg of¹⁵N tracer per square meter in March 1991. Ammonium and nitrate depositions were simulated on five plots each, by labeling with either¹⁵N-NH₄ ⁺ or¹⁵N-NO₃ ^–, and the¹⁵N pulse was followed during two successive growing seasons (1991 and 1992). Total recovery rates of the¹⁵N tracer in the entire stand ranged between 93 and 102% for both nitrogen forms in 1991, and 82% in June 1992. ⁵ N ratios increased rapidly in all compartments of the ecosystem. Roots and soils (to 65 cm depth) showed significant¹⁵N enrichments for both¹⁵N-treatments compared to reference plots. Newly grown spruce tissues were more enriched than older ones, but the most enriched ¹⁵N values were found in the understory vegetation. Although spruce trees were a much larger pool (1860 g biomass/m²) than understory vegetation (Vaccinium myrtillus 333 g/m², Calluna vulgaris 142 g/m², Deschampsia flexuosa 22 g/m²), the ericaceous shrubs and the perennial grass were a much greater sink for the¹⁵N label. Eight months after labeling, 9% of the ammonium and 15% of the nitrate label were found in the understory. P.abies retained only 3% of the¹⁵N-ammonium and 7% of the¹⁵N-nitrate. The main sink for both¹⁵N tracers was the soil, where 87% of the ammonium and 79% of the nitrate tracer were found. The organic soil horizon (5-0 cm depth) contained 63% of the¹⁵N-ammonium and 46% of the¹⁵N -nitrate suggesting strong immobilization by microorganisms of both N forms. Eight months after tracer application, about 16% of both¹⁵N-tracers was found below 25 cm soil depth. This 16% corresponds well to a 20% decrease in the recovery of both¹⁵N tracers after 15 months and indicates a total loss out of the ecosystem. Highly enriched ¹⁵N values were found in fruit bodies of fungi growing in reference lots (no¹⁵N addition), although soils did not show increased ¹⁵N ratios. No transfer of¹⁵N-tracer between fungi and spruce or understory vegetation was apparent yet.     

Nitrogen form, availability, and mycorrhizal colonization affect biomass and nitrogen isotope patterns in Pinus sylvestris

Nitrogen (N) isotope patterns are useful for understanding carbon and nitrogen dynamics in mycorrhizal systems but questions remain about how different N forms, fungal symbionts, and N availabilities influence δ¹⁵N signatures. Here, we studied how biomass allocation and δ¹⁵N patterns in Pinus sylvestris L. cultures were affected by nitrogen supply rate (3% per day or 4% per day relative to the nitrogen already present), nitrogen form (ammonium versus nitrate), and mycorrhizal colonization by fungi with a greater (Laccaria laccata) or lesser (Suillus bovinus) ability to assimilate nitrate. Mycorrhizal (fungal) biomass was greater with ammonium than with nitrate nutrition for Suillus cultures but similar for Laccaria cultures. Total biomass was less with nitrate nutrition than with ammonium nutrition for nonmycorrhizal cultures and was less in mycorrhizal cultures than in nonmycorrhizal cultures. The sequestration of available N by mycorrhizal fungi limited plant N supply. This limitation and the higher energetic cost of nitrate reduction than ammonium assimilation appeared to control plant biomass accumulation. Colonization decreased foliar δ¹⁵N by 0.5 to 2.2‰ (nitrate) or 1.7 to 3.5‰ (ammonium) and increased root tip δ¹⁵N by 0 to 1‰ (nitrate) or 0.6 to 2.3‰ (ammonium). Root tip δ¹⁵N and fungal biomass on root tips were positively correlated in ammonium treatments (r ²?=?0.52) but not in nitrate treatments (r ²?=?0.00). Fungal biomass on root tips was enriched in ¹⁵N an estimated 6–8‰ relative to plant biomass in ammonium treatments. At high nitrate availability, Suillus colonization did not reduce plant δ¹⁵N. We conclude that: (1) transfer of ¹⁵N-depleted N from mycorrhizal fungi to plants produces low plant δ¹⁵N signatures and high root tip and fungal δ¹⁵N signatures; (2) limited nitrate reduction in fungi restricted transfer of ¹⁵N-depleted N to plants when nitrate is supplied and may account for many field observations of high plant δ¹⁵N under such conditions; (3) plants could transfer assimilated nitrogen to fungi at high nitrate supply but such transfer was without ¹⁵N fractionation. These factors probably control plant δ¹⁵N patterns across N availability gradients and were here incorporated into analytical equations for interpreting nitrogen isotope patterns in mycorrhizal fungi and plants.     

Field measurement of lupin belowground nitrogen accumulation and recovery in the subsequent cereal-soil system in a semi-arid Mediterranean-type climate

In situ ¹⁵N-labelling was used to provide a quantitative assessment of the total contribution of lupin (Lupinus angustifolius) to below-ground (BG) N accumulation during a growing season under field conditions, and to directly trace the fate of the lupin BG N in the next season, including quantifying the N benefit from lupin to a following wheat (Triticum aestivum) crop. The experiments were conducted at two sites, both experiencing a semi-arid Mediterranean-type climate in the wheat-growing region of Western Australia but with differing soil types, a deep sand (Moora) and a sand-over-clay shallow duplex soil (East Beverley, EB). Lupin shoot and root dry matter and total plant N accumulation, proportional dependence on nitrogen fixation and grain yield were greater at the deep sand site than the duplex soil site, although there was a similar proportion of shoot N to estimated total BG N at both sites. The proportion of total plant BG N decreased from the vegetative stage (42–51%) to peak biomass (25–39%) and maturity (23–34%). From 56–67% of BG N on the deep sand and 74–86% on the duplex soil was not recovered in coarse roots (>2 mm) or as soluble N, but was present in the insoluble organic N fraction. There was evidence for cycling of lupin root-derived N into soil microbial biomass and soluble organic N during lupin growth (by the late vegetative stage), but no evidence for leaching of legume derived BG N during the lupin season. Estimates of fixed N input BG were at least four times greater if based on total lupin BG N rather than on N recovered in coarse roots (>2 mm). There were no apparent losses of lupin BG N during the summer fallow period subsequent to lupin harvest at either site. Also, immediately prior to sowing of wheat there were similar proportions of lupin BG N in the inorganic (20–25%) and microbial biomass (6–9%) pools at both sites, with the majority of BG N detected in the <2 mm fraction of the soil column. However, the proportion of residual lupin BG N estimated to benefit the aboveground wheat biomass was relatively low, 10% on the deep sand and only 3% on the shallow duplex. Some (14%) residual lupin BG N was leached as nitrate to 1 m on the deep sand compared to 8% of residual lupin BG N leached to the clay layer (0.3 m) on the shallow duplex. About 27% of the residual lupin BG N on the deep sand at Moora had apparently mineralised by the end of the succeeding wheat season (i.e. recovered either in the wheat shoots, as inorganic N in the soil profile or as leached nitrate) compared to only 12% at EB. There was an unaccounted for large loss of residual lupin BG N (50%) from the duplex soil at EB during the wheat season, postulated to be chiefly via denitrification. At both sites after the wheat season a substantial proportion (32–55%) of legume derived BG N was still present as residual insoluble organic N, considered to be an important contribution to structural and nutritional long-term sustainability of these soils.     

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.     

15N-ammonium and 15N-nitrate uptake of a 15-year-old Picea abies plantation

Throughfall nitrogen of a 15-year-old Picea abies (L.) Karst. (Norway spruce) stand in the Fichtelgebirge, Germany, was labeled with either ¹⁵N-ammonium or ¹⁵N-nitrate and uptake of these two tracers was followed during two successive growing seasons (1991 and 1992). ¹⁵N-labeling (62 mg ¹⁵N m^-2 under conditions of 1.5 g N m^-2 atmospheric nitrogen deposition) did not increase N concentrations in plant tissues. The ¹⁵N recovery within the entire stand (including soils) was 94%±6% of the applied ¹⁵N-ammonium tracer and 100%±6% of the applied ¹⁵N-nitrate tracer during the 1st year of investigation. This decreased to 80%±24% and 83%±20%, respectively, during the 2nd year. After 11 days, the ¹⁵N tracer was detectable in 1-year-old spruce needles and leaves of understory species. After 1 month, tracer was detectable in needle litter fall. At the end of the first growing season, more than 50% of the ¹⁵N taken up by spruce was assimilated in needles, and more than 20% in twigs. The relative distribution of recovered tracer of both ¹⁵N-ammonium and ¹⁵N-nitrate was similar within the different foliage age classes (recent to 11-year-old) and other compartments of the trees. ¹⁵N enrichment generally decreased with increasing tissue age. Roots accounted for up to 20% of the recovered ¹⁵N in spruce; no enrichment could be detected in stem wood. Although ¹⁵N-ammonium and ¹⁵N-nitrate were applied in the same molar quantities (¹⁵NH ₄ ⁺ : ¹⁵NO ₃ ^- =1:1), the tracers were diluted differently in the inorganic soil N pools (¹⁵NH ₄ ⁺ /NH ₄ ⁺ : ¹⁵NO ₃ ^- /NO ₃ ^- =1:9). Therefore the measured ¹⁵N amounts retained by the vegetation do not represent the actual fluxes of ammonium and nitrate in the soil solution. Use of the molar ammonium-to-nitrate ratio of 9:1 in the soil water extract to estimate ¹⁵N uptake from inorganic N pools resulted in a 2–4 times higher ammonium than nitrate uptake by P. abies.     

Unusual seasonal patterns and inferred processes of nitrogen retention in forested headwaters of the Upper Susquehanna River

Atmospheric deposition contributes a large fraction of the annual nitrogen (N) input to the basin of the Susquehanna River, a river that provides two-thirds of the annual N load to the Chesapeake Bay. Yet, there are few measurements of the retention of atmospheric N in the Upper Susquehanna’s forested headwaters. We characterized the amount, form (nitrate, ammonium, and dissolved organic nitrogen), isotopic composition (δ¹⁵N- and δ¹⁸O-nitrate), and seasonality of stream N over 2 years for 7–13 catchments. We expected high rates of N retention and seasonal nitrate patterns typical of other seasonally snow-covered catchments: dormant season maxima and growing season minima. Coarse estimates of N export indicated high rates of inorganic N retention (>95%), yet streams had unexpected seasonal nitrate patterns, with summer peaks (14–96 μmol L⁻¹), October crashes (<1 μmol L⁻¹), and modest rebounds during the dormant season (<1–20 μmol L⁻¹). Stream δ¹⁸O-nitrate values indicated microbial nitrification as the primary source of stream nitrate, although snowmelt or other atmospheric source contributed up to 47% of stream nitrate in some March samples. The autumn nitrate crash coincided with leaffall, likely due to in-stream heterotrophic uptake of N. Hypothesized sources of the summer nitrate peaks include: delayed release of nitrate previously flushed to groundwater, weathering of geologic N, and summer increases in net nitrate production. Measurements of shale δ¹⁵N and soil-, well-, and streamwater nitrate within one catchment point toward a summer increase in soil net nitrification as the driver of this pattern. Rather than seasonal plant demand, processes governing the seasonal production, retention, and transport of nitrate in soils may drive nitrate seasonality in this and many other systems.     

Long-Term Nitrogen Additions and Nitrogen Saturation in Two Temperate Forests

This article reports responses of two different forest ecosystems to 9 years (1988–96) of chronic nitrogen (N) additions at the Harvard Forest, Petersham, Massachusetts. Ammonium nitrate (NH₄NO₃) was applied to a pine plantation and a native deciduous broad-leaved (hardwood) forest in six equal monthly doses (May–September) at four rates: control (no fertilizer addition), low N (5 g N m^-2 y^-1), high N (15 g N m^-2 y^-1), and low N + sulfur (5 g N m^-2 y^-1 plus 7.4 g S m^-2 y^-1). Measurements were made of net N mineralization, net nitrification, N retention, wood production, foliar N content and litter production, soil C and N content, and concentrations of dissolved organic carbon (DOC) and nitrogen (DON) in soil water. In the pine stand, nitrate losses were measured after the first year of additions (1989) in the high N plot and increased again in 1995 and 1996. The hardwood stand showed no significant increases in nitrate leaching until 1995 (high N only), with further increases in 1996. Overall N retention efficiency (percentage of added N retained) over the 9-year period was 97–100% in the control and low N plots of both stands, 96% in the hardwood high N plot, and 85% in the pine high N plot. Storage in aboveground biomass, fine roots, and soil extractable pools accounted for only 16–32% of the added N retained in the amended plots, suggesting that the one major unmeasured pool, soil organic matter, contains the remaining 68–84%. Short-term redistribution of ¹⁵N tracer at natural abundance levels showed similar division between plant and soil pools. Direct measurements of changes in total soil C and N pools were inconclusive due to high variation in both stands. Woody biomass production increased in the hardwood high N plot but was significantly reduced in the pine high N plot, relative to controls. A drought-induced increase in foliar litterfall in the pine stand in 1995 is one possible factor leading to a measured increase in N mineralization, nitrification, and nitrate loss in the pine high N plot in 1996. Received 2 April 1999; Accepted 29 October 1999.     

Influence of light intensity on the distribution of carbon and consequent effects on mineralization of soil nitrogen in a barley (Hordeum vulgare L.)-soil system

A pot experiment was conducted in a ¹⁴C-labelled atmosphere to study the influence of living plants on organic-N mineralization. The soil organic matter had been labelled, by means of a 200-days incubation, with ¹⁵N. The influence of the carbon input from the roots on the formation of microbial biomass was evaluated by using two different light intensities (I). Mineralization of ¹⁵N-labelled soil N was examined by following its fate in both the soil biomass and the plants. Less dry matter accumulated in shoots and roots at the lower light intensity. Furthermore, in all the plant-soil compartments examined, with the exception of rhizosphere respiration, the proportion of net assimilated ¹⁴C was lower in the low-I treatment than in the high-I treatment. The lower rates of ¹⁴C and ¹⁵N incorporation into the soil biomass were associated with less root-derived ¹⁴C. During the chamber period (¹⁴CO₂-atmosphere), mineralized amounts of ¹⁵N (measured as plant uptake of ¹⁵N) were small and represented about 6.8 to 7.8% of the initial amount of organic ¹⁵N in the soil. Amounts of unlabelled N found in the plants, as a percentage of total soil N, were 2.5 to 3.3%. The low availability of labelled N to microorganisms was the result of its stabilization during the 210 days of soil incubation. Differences in carbon supply resulted in different rates of N mineralization which is consistent with the hypothesis that roots induce N mineralization. N mineralization was higher in the high-I treatment. On the other hand, the rate of mineralization of unlabelled stable soil N was lower than labelled soil ¹⁵N which was stabilized. The amounts of ¹⁵N mineralized in planted soil during the chamber period (43 days) which were comparable with those mineralized in unplanted soil incubated for 210 days, also suggested that living plants increased the turnover rate of soil organic matter.     

Protein degradation in different feedstuffs labelled with 15N by using the nylon bag technique

The nylon bag technique was used to determine the Nitrogen (N) and ¹⁵N degradation of ¹⁵N labelled feedstuffs in the rumen. The N and ¹⁵N degradation values were calculated according to ?rskov and McDonald (1979) and ranged from 46.8 to 92.0 and from 61.8 to 93.6%, respectively. The differences between N and ¹⁵N degradation values of high fibre content feedstuffs are the highest, thus the measuring errors were greatest here. But differences also existed in concentrates. This study indicated that especially barley had a higher proportion of microbial N in the bag residues after the washing than the other concentrates. Therefore it is necessary to correct the N degradation values not only in cases of high fibre content but also in cases of low nitrogen content of feedstuffs. The calculation of the N degradation values could be possible on the basis of crude fibre and crude protein contents of feedstuffs. But experiments with a much larger number of ¹⁵N labelled feedstuffs have to be realized to give an accurate prediction of N degradation.