Phosphorus and nitrogen fertilization of a Norway spruce forest-effects on needle concentrations and acid phosphatase activity in the humus layer

Increased atmospheric deposition of N might eventually lead to P deficiency. The relation between needle P concentration and acid phosphatase activity in the humus layer was studied during 1990–93 in a Norway spruce stand where the water and N and P supplies had been experimentally manipulated since 1988. Treatments included control (C), yearly application of ammonium sulphate (NS), N-free fertilizer (V), granulated wood ash (A), irrigation (I), drought (D) and water plus nutrients in an optimum combination (IF). We found indications of a feed-back mechanism for P, where low concentrations in the needles were associated with increased acid phosphatase activity in the humus layer. Acid phosphatase estimations made during moist soil conditions were much more informative than those made during dry conditions. We further argue that a site-specific base-line exists for acid phosphatase activity in the soil, mainly originating from enzymes immobilized in the field, but active in the assay. Increased phosphatase activity, above the base line, was generally found in the A, I and NS treatments, but in some cases also in C. Although P and N concentrations were significantly higher in the IF treatment as compared to the C and the D treatments, the P as fraction of N was 0.10 and thus balanced in all cases. In the A and I treatment P:N was around 0.09, while it was only 0.07 in the NS treatment, mainly due to high N concentrations. The latter treatment thus created an imbalanced situation where P additions most likely would have increased tree growth.     

Immobilization and mineralization of tracer nitrogen in forest raw humus

Summary An experiment was conducted to permit an assessment of the effect of the N-carrier on the interchange of fertilizer-N in humus maintained under waterlogged conditions. The experiment was carried out at 4°; 12°; and 20°C; and the incubation varied from 0 to 90 days. N¹⁵-enriched urea, ammonium chloride and potassium nitrate were used. The incorporation of fertilizer-N into a non KCl-extractable fraction occurred relatively fast in the urea treatment regardless of the temperature. The highest recovery of tracer-N in the mentioned fraction was attained at 20°C after 90 days' incubation. The urea was hydrolyzed at a high rate, and within 30 days of fertilizer application the inorganic N¹⁵-pool was almost stabilized at a level in the vicinity of 60% of the added N at 12° and 20°C. Nitrogen added as ammonium was incorporated into the non-extractable fraction at a lower rate than was the N added in the form of urea. The data reveal a continual N-immobilization, but at 20°C a substantial net re-mineralization took place during the last part of the experiment. The nitrate treatment was followed by a sizeable temporary net immobilization of tracer-N. The extremely great changes in the inorganic N¹⁵-pool and the net re-mineralization of tracer-N were positively correlated with temperature and incubation time. Within 30 days, about 95% of the added nitrate-N had disappeared from the inorganic pool at 12° and 20°C. Approximately 7.6% of the added NO₃-N was re-mineralized at 20°C after 90 days' incubation. The data reveal great losses of gaseous N from this waterlogged acid forest soil after treatment with nitrate. Contribution from the Forest Soil Fertilization Research Group, Vollebekk, Norway. This work was supported by the Agricultural Research Council of Norway.     

Immobilization and mineralization of tracer nitrogen in forest raw humus

Summary A series of laboratory incubation experiments was conducted to provide information about the effects of soil temperature on the nitrogen interchange in forest raw humus after addition of different forms of nitrogen fertilizers enriched with the N¹⁵-isotope.A positive correlation between temperature and immobilization of nitrogen (non-extractable inN KCl) added as urea-N¹⁵, was found at temperatures of 4°, 12°, and 20°C. When ammonium was added as the source of nitrogen the data indicated a gradual and continual immobilization of tracer nitrogen throughout the entire 90 days experimental period at temperatures of 4° and 12°C. At an incubation temperature of 20°C the amount of immobilized tracer nitrogen reached a relatively low but almost constant level within 10 days.A positive correlation between incubation temperature and the overall nitrogen turnover was found in the raw humus after the nitrate application. Within 3 days of incubation significant net re-mineralization of added tracer nitrate nitrogen took place at the highest temperature (20°C) used. The data show a negative correlation between incubation temperature and net accumulated nitrite-nitrate nitrogen after addition of urea, while no significant amount of nitrite-nitrate was detected in the humus after the ammonium application.At temperatures of 4° and 12°C the nitrogen added as ammonium as compared with nitrate was more rapidly used in supplying the needs of the micro-organisms decomposing forest raw humus. However, the results reveal that in spite of the presence of ammonium in the system, nitrate nitrogen is included in the pathway of the internal nitrogen cycle. Compared with the ammonium and the nitrate treatment the preferential utilization of nitrogen added as urea generally increased with increasing incubation time and temperature.Contribution from the Forest Soil Fertilization Research Group, Vollebekk, Norway. This work was supported by the Agricultural Research Council of Norway.     

Tracer studies on nitrogen immobilization-mineralization relationships in forest raw humus

Summary An investigation was conducted to study the effect of fertilizer-N, application rate, and incubation temperature on the immobilization-mineralization relationships of N in forest raw humus. Urea, ammonium chloride, and potassium nitrate enriched with N¹⁵ were used at rates of 0, 50, 100, 200, and 400 ppm N. During a 60 days' experimental period the soil was incubated at 12° and 20°C. The interchanges of N added to this highly acid humus material are to a great extent found to be governed by the N-carrier itself. The total recovery in the N¹⁵ inorganic pool at the end of the 60 days' period, increased with increasing fertilizer application rate regardless of the source of N added. With one exception only, the recovery of KCl-extractable N¹⁵ was larger in the ammonium treatment than in the urea-treated humus. The more striking differences between the interchanges of N¹⁵ in the urea and the ammonium treatments, as compared with the nitrate-treated humus, are revealed in the extremely high recovery of KCl-extractable N, and a concomitant low recovery of the added N in the non-extractable pool in the nitrate treatment. The immobilization and re-mineralization of N are found to be positively correlated with temperature. The total recovery of N¹⁵ was high in the nitrate treatment, and even higher in the ammonium-treated soil (96 to 104 per cent). However, the unaccounted-for losses reached a maximum of about 48 per cent of the added N at 20°C in the 400 ppm urea-N treatment. The results are consistent in showing a statistically highly significant effect of N-fertilizer, application rate, and temperature on the net mineralization of humus N. The release of soil N is discussed in relation to the highly significant two- and three-way interactions between the above-mentioned main variables. Contribution from the Forest Soil Fertilization Research Group, Vollebekk, Norway. This work was supported by the Agricultural Research Council of Norway.     

The use of different soil nitrogen sources by young Norway spruce plants

 Three-year-old Norway spruce trees were planted into a low-nitrogen mineral forest soil and supplied either with two different levels of mineral nitrogen (NH₄NO₃) or with a slow-release form of organic nitrogen (keratin). Supply of mineral nitrogen increased the concentrations of ammonium and nitrate in the soil solution and in CaCl₂-extracts of the rhizosphere and bulk soil. In the soil solution, in all treatments nitrate concentrations were higher than ammonium concentrations, while in the soil extracts ammonium concentrations were often higher than nitrate concentrations. After 7 months of growth, ¹⁵N labelled ammonium or nitrate was added to the soil. Plants were harvested 2 weeks later. Keratin supply to the soil did not affect growth and nitrogen accumulation of the trees. In contrast, supply of mineral nitrogen increased shoot growth and increased the ratio of above-ground to below-ground growth. The proportion of needle biomass to total above-ground biomass was not increased by mineral N supply. The atom-% ¹⁵N was higher in younger needles than in older needles, and in younger needles higher in plants supplied with ¹⁵N-nitrate than in plants supplied with ¹⁵N-ammonium. The present data show that young Norway spruce plants take up nitrate even under conditions of high plant internal N levels. Received: 1 April 1998 / Accepted: 9 October 1998     

Temperature sensitivity of nitrogen productivity for Scots pine and Norway spruce

Environmental conditions control physiological processes in plants and thus their growth. The predicted global warming is expected to accelerate tree growth. However, the growth response is a complex function of several processes with both direct and indirect effects. To analyse this problem we have used needle nitrogen productivity, which is an aggregate parameter for production of new foliage. Data on needle dry matter, production, and nitrogen content in needles of Scots pine ( Pinus sylvestris) and Norway spruce ( Picea abies) from a wide range of climatic conditions were collected and needle nitrogen productivities, defined as dry matter production of needles per unit of nitrogen in the needle biomass, were calculated. Our results show that the nitrogen productivity for spruce is insensitive to temperature. However, for pine, temperature affects both the magnitude of nitrogen productivity at low needle biomass and the response to self-shading but the temperature response is small at the high end of needle biomass. For practical applications it may be sufficient to use a species-specific nitrogen productivity that is independent of temperature. Because temperature affects tree growth indirectly as well as through soil processes, the effects of temperature change on tree growth and ecosystem carbon storage should mainly be derived from effects on nitrogen availability through changes in nitrogen mineralization. In addition, this paper summarises data on dry matter, production and nitrogen content of needles of conifers along a temperature gradient.     

Nitrogen transformations in limed and nitrogen fertilized soil in Norway spruce stands

Nitrogen transformations in the soil, and the resulting changes in carbon and nitrogen compounds in soil percolate water, were studied in two stands of Norway spruce (Picea abies L.). Over the last 30 years the stands were repeatedly limed (total 6000 kg ha^–1), fertilized with nitrogen (total about 900 kg ha^–1), or both treatments together. Both aerobic incubations of soil samples in the laboratory, and intact soil core incubations in the field showed that in control plots ammonification widely predominated over nitrification. In both experiments nitrogen addition increased the formation of mineral-N. In one experiment separate lime and nitrogen treatments increased nitrification, in the other, only lime and nitrogen addition together had this effect. In one experiment immobilization of nitrogen to soil microbial biomass was lower in soil only treated with nitrogen. Soil percolate water was collected by means of lysimeters placed under the humus layer and 10 cm below in the mineral soil. Total N, NH₄-N and NO₃-N were measured, and dissolved organic nitrogen was fractioned according to molecular weight. NO₃-N concentrations in percolate water, collected under the humus layer, were higher in plots treated with N-fertilizer, especially when lime was also added. The treatments had no effect on the N concentrations in mineral soil. A considerable proportion of nitrogen was leached in organic form.     

Carbon and nitrogen flow in silver birch and Norway spruce connected by a common mycorrhizal mycelium

 Spruce and birch seedlings were grown together in boxes filled with unsterile peat. Both seedlings were colonized by the ectomycorrhizal fungus Scleroderma citrinum. The two plants thus shared a common external mycelium. ¹⁵N-labelled ammonium was supplied exclusively to the fungus, while the birch or the spruce plant was continuously fed with ¹³C-labelled CO₂ for 72 h. The carbon and nitrogen transfer rates were strikingly different for birch and spruce seedlings. The mycorrhizal mycelium received carbohydrates mainly from the birch plant and the nitrogen transfer by the fungus to the plants was largely directed towards the birch. Carbon assimilates were also transferred in both directions between birch and spruce; however, there was no conclusive evidence for a net transfer of carbon between the plants. Accepted: 20 September 1996     

Karyotype of Norway spruce by multicolor FISH

The chromosomes (2n = 2x = 24) of Norway spruce are very large since their size reflects the huge amount of genomic DNA (2C = 30 × 10⁹ bp). However, the identification of homologous pairs is hampered by their high degree of similarity at the morphological level. Data so far presented in the literature were not sufficient to solve all the ambiguities in chromosome identification. Several genomic Norway spruce DNA clones containing highly repetitive sequences have been identified and characterised in our laboratory. Three of them were selected for fluorescent in situ hybridization (FISH) experiments because of their strong signals and suitability for chromosome identification: PATR140 hybridized at the centromeric site of three chromosome pairs; PAF1 hybridized in six subtelomeric and two centromeric sites; 1PABCD6 co-localized with the subtelomeric sites identified by PAF1. The statistical analysis of microscopic measurements of chromosomes in combination with the FISH signals of these probes allowed the unambigous construction of Norway spruce karyotype. We also compared the karyotype of Norway spruce with that of other spruce species to infer the number and kind of rearrangements that have occurred during the evolution of these species.Communicated by D.B. Neale     

Accumulation of ammonium in Norway spruce (Picea abies) seedlings measured by in vivo 14N-NMR

(14)N-NMR and (31)P-NMR have been used to monitor the in vivo pH in roots, stems, and needles from seedlings of Norway spruce, a typical ammonium-tolerant plant. The vacuolar and cytoplasmic pH measured by (31)P-NMR was found to be c. pH 4.8 and 7.0, respectively, with no significant difference between plants growing with ammonium or nitrate as the N-source. The (1)H-coupled (14) NH 4+ resonance is pH-sensitive: at alkaline pH it is a narrow singlet line and below pH 4 it is an increasing multiplet line with five signals. The pH values in ammonium-containing compartments measured by (14)N-NMR ranged from 3.7 to 3.9, notably lower than the estimated pH values of the P(i) pools. This suggests that, in seedlings of Norway spruce, ammonium is stored in vacuoles with low pH possibly to protect the seedlings against the toxic effects of ammonium ( NH 4+) or ammonia (NH3). It was also found that concentrations of malate were 3-6 times higher in stems than in roots and needles, with nitrate-grown plants containing more malate than plants grown with ammonium.     

Peroxisome proliferation in Norway spruce induced by ozone

Summary Catalase (EC 1.11.1.6) activity (both total and specific activity) of particulate fractions of needles of Norway spruce [Picea abies (L.) Karst.] was elevated approximately 2-fold following exposure of trees to 60–70 g/m³ of ozone during the growing season compared to trees receiving charcoal filtered air (about 15 g/m³ ozone). Measurements were from homogenates fractionated into particulate and soluble (supernatent) activities. In contrast, the catalase activity of the supernatant was unchanged in response to ozone treatment. Catalase activity declined as the needles aged comparing current, 1-, and 2-year needles but the ozone-induced increment remained constant. Electron microscope cytochemistry using peroxidatic coupling with 3,3-diaminobenzidine carried out in parallel, revealed catalase-containing peroxisomes both in situ and in the particulate fractions analyzed for catalase activity. The tissue volume occupied by peroxisomes in response to needle age and ozone appeared to vary approximately in proportion to the measured catalase activity. Overall cytochemical reactivity for catalase declined with needle age, but, for all years, was greater in needles of trees receiving air supplemented with ozone compared to those of trees receiving charcoal filtered air.Abbreviations DAB 3,3-diaminobenzidine tetrahydrochloride     

Biological activity of humus acids isolated from spruce wood decomposed by fungi

Biological activity of humic acid, hymatomelanic acid, lignofulvic acid, and fulvic acids isolated from peat, from spruce wood decomposed by the fungusFomes marginatus (Fr.) Gillet, and from spruce wood decomposed by the fungusPoria cinerascens Bres. was tested on cress seedlings (Lepidium sativum L.). The results evaluated by the analysis of variance indicate that humus acids show a high degree of affinity in their biological activity which is a condition for the applicability of the decomposition of wood by fungi as a model of the humus formation. Humic and hymatomelanic acids, products of an advanced step of humification, have a predominantly stimulatory effect especially on the hypocotyl. Lignofulvic acids and fulvic acids exert a rather inhibitory effect with the root being more affected.     

Influences of different nitrate to ammonium ratios on chlorosis,cation concentrations and the binding forms of Mg and Ca in needles of Mg-deficient Norway spruce

Effects of Mg deficiency and variations of the NO₃ ^-/NH₄ ⁺-ratio on chlorophyll, mineral nutrient concentrations and the binding forms of Mg and Ca were investigated in current-year, one- and two-year-old needles of clonal Norway spruce (Picea abies [L.] Karst.). Six-year-old spruce plants were grown for one year in sand culture with circulating nutrient solutions containing sufficient (0.2 mt M) or limiting (0.04 mt M) concentrations of Mg. The NO₃ ^-/NH₄ ⁺-ratio in the nutrient solutions administered to the experimental trees was adjusted to 0.76 in the Mg-sufficient treatment and to 1.86, 0.76 and 0.035 in Mg-limited treatments. Mg and chlorophyll concentrations, were strongly influenced by the applied nitrogen source in current-year needles and - to a less extent - also in one-year-old needles. NH₄ ⁺-dominated nutrition resulted in decreased height growth and significantly lower Mg and chloropyhll concentrations in current-year and one-year-old needles compared to NO₃ ^--dominated nutrition. Decreases in total Mg were linearly correlated to reductions of water-soluble Mg and water-unsoluble Mg not bound to chlorophyll. Mg bound to chlorophyll, however, was only reduced, when total Mg decreased below a physiological threshold value of 2% of the total nitrogen concentration in the respective needles. Total Ca concentrations in the needles, which were reduced by Mg deficiency especially when nutrition was NH₄ ⁺-dominated, were strongly correlated to the portion of Ca bound to oxalate. The amount of water-soluble Ca and pectate-bound Ca remained nearly constant, independent from changes of total Ca concentrations. Negative effects of increasing NH₄ ⁺ supply on concentrations of Mg and other cations in the needles can be attributed to an inhibition of cation uptake induced by ion antagonism and/or reduced root growth.     

Nitrogen distribution in young Norway spruce (Picea abies) trees as affected by pedospheric nitrogen supply

In numerous locations in Europe spruce trees are exposed to high loads of nitrogen. The present study was performed to characterize the distribution of nitrogen compounds under these conditions. For this purpose Norway spruce ( Picea abies [L.] Karst.) trees were cultivated under close-to-natural conditions of a forest understory in soil from an apparently nitrogen-limited field site in the Black Forest either with, or without supplementation of nitrogen as ammonium nitrate. After 11 and 20 months, growth, total nitrogen contents of the biomass, and total soluble non-proteinogenic nitrogen compounds (TSNN, i.e. nitrate, ammonium, soluble proteinogenic and non-proteinogenic amino compounds) in needles, xylem sap and phloem exudate were analysed. After 20 months of growth, N-fertilization had slightly enhanced the biomass of current-, but not of 1-year-old shoots. At both harvests, total N-content of 1-year-old needles was increased by N-fertilization, whereas current-year needles were not significantly affected. By contrast, TSNN was elevated by N-fertilization in both current-year and 1-year-old needles. The increase in TSNN was mainly attributed to an accumulation of arginine. Xylem sap analysis showed that the increase in TSNN of the needles was a consequence of enhanced nitrogen assimilation of the roots rather than the shoot. Since also TSNN in phloem exudates was enhanced, it appears that N-fertilization elevates the cycling pool of amino compounds in young Norway spruce trees. However, this pool seems to be subject to metabolic interconversion, since mainly glutamine and aspartate are transported in the xylem from the roots to the shoot, but arginine accumulated in the needles and the phloem.     

Effect of exposure to fluoride,nitrogen compounds and SO2 on the numbers of spruce shoot aphids on Norway spruce seedlings

Summary Development of spruce shoot aphid (Cinara pilicornis Hartig) populations was monitored in natural and artificial infestations of Norway spruce (Picea abies Karst.) seedlings, exposed to air pollutants in an experimental field. The pollutants, applied both singly and in mixtures, were gaseous sulphur dioxide, NaF (30 mg l^-1 F) and Ca(NO₃)₂ or (NH₄)₂SO₄ in aqueous solutions (200 mg l^-1 N). Aphid numbers on 10 seedlings in each treatment and two control plots were counted at 2-week intervals. At the beginning of the experiment aphid numbers did not differ between treatments. Aphid populations peaked in late June and early July. All the pollutants and their combinations significantly increased the numbers of aphids per seedling. Four apterous females were transferred to spruce seedlings which were growing in containers in the same plots. After 4–5 weeks aphid numbers were significantly higher in the fluoride treatment and in the combined treatment of fluoride, nitrogen and SO₂. The pollution treatments did not have a significant effect on shoot growth. Concentrations of F and S in needles were higher in treatments involving these pollutants. There were no significant differences in the concentrations of free amino acids in shoot stems between control and fluoride treatment. However, the relatively low concentration of arginine in the F treatment at the end of the growing season might indicate disturbances in the nitrogen metabolism of spruce seedlings.     

Fate of ammonium 15N in a Norway spruce forest under long-term reduction in atmospheric N deposition

In the last decades, in particular forest ecosystems became increasingly N saturated due to elevated atmospheric N deposition, resulting from anthropogenic N emission. This led to serious consequences for the environment such as N leaching to the groundwater. Recent efforts to reduce N emissions raise the question if, and over what timescale, ecosystems recover to previous conditions. In order to study the effects on N distribution and N transformation processes under the lowered N deposition treatment, we investigated the fate of deposited NH₄ ⁺-¹⁵N in soil of a N-saturated Norway spruce forest (current N deposition: 34 kg ha^?1 year^?1; critical N load: 14 kg ha^?1 year^?1), where N deposition has been reduced to 11.5 kg ha^?1 year^?1 since 14.5 years. We traced the deposited ¹⁵N in needle litter, bulk soil, and amino acids, microbial biomass and inorganic N in soil. Under reduced N deposition, 123 ± 23% of the deposited N was retained in bulk soil, while this was only 72 ± 15% under ambient deposition. We presume that with reduced deposition the amount of deposited N was small enough to become completely immobilized in plant and soil and no leaching losses occurred. Trees receiving reduced N deposition showed a decline in N content as well as in ¹⁵N incorporation into needle litter, indicating reduced N plant uptake. In contrast, the distribution of ¹⁵N within the soil over active microbial biomass, microbial residues and inorganic N was not affected by the reduced N deposition. We conclude that the reduction in N deposition impacted only plant uptake and drainage losses, while microbial N transformation processes were not influenced. We assume changes in the biological N turnover to start with the onset of the decomposition of the new, N-depleted litter.     

Long-term dynamics of aboveground fungal communities in a subalpine Norway spruce forest under elevated nitrogen input

As anthropogenic N deposition has been suspected to be the main reason for the decline of macromycetous sporocarp production in forest ecosystems, various N-fertilization experiments were started in the mid 1990s. The dynamics of ectomycorrhizal (root-inhabiting) and terricolous saprobic (litter-inhabiting) fungal communities were studied by exhaustive sporocarp inventories in a substitution Norway spruce (Picea abies) forest in two 256-m² plots sampled for periods of 1 week at 1-m² resolution between 1994 and 2007. N was added to the soil twice per year in one plot from the fourth year onwards. The effects of N input and time on aboveground fungal communities were assessed using redundancy analysis, principal response curves and non-parametric multivariate ANOVA. Results of this long-term experiment revealed that both ectomycorrhizal and saprobic fungal communities responded to an increase in soil N input. The ectomycorrhizal community reacted by a fast decrease in sporocarp production and in species richness, whereas the saprobic community was less affected. The response was highly species specific, especially for the saprobic community. The difference in species composition between control and fertilized plots was significant after 1 year of N addition for ectomycorrhizal fungi and only after 3 years for saprobic fungi. An aging effect affected sporocarp production in the whole area. For both communities, this unidirectional drift in species composition was as important as the treatment effect. This result highlights the importance of considering the respective role of treatment and year effects in long-term field experiments on fungal communities.     

Compartmentation and flux characteristics of ammonium in spruce

Using ¹³NH ₄ ⁺ as a tracer, compartmental analyses for NH ₄ ⁺ were performed in non-mycorrhizal roots of intact Picea glauca (Moench) Voss. seedlings at four different concentration regimes of external NH ₄ ⁺ ([NH ₄ ⁺ ]_o), i.e. 0, 10, 100, and 1500 M. Three kinetically distinct compartments were identified, with half-lives of exchange of approximately 2 s, 30 s, and 14 min, assumed to represent surface adsorption, Donnan free space, and cytoplasm, respectively. No significant differences were found in half-lives of exchange with changes in [NH ₄ ⁺ ]_o. Influx was calculated to be 0.96 mol·g^–1·h^–1 in N-deprived plants (measured at 10 M [NH ₄ ⁺ ]_o), while under steady-state conditions it was 0.21 mol·g^–1h^–1 at 10 M [NH ₄ ⁺ ]_o, 1.96 mol·g^–1h·^–1 at 100 M [NH ₄ ⁺ ]_o, and 6.45 mol·g^–1·h^–1 at 1.5 mM [NH ₄ ⁺ ]_o. Efflux measured over the same range constituted approximately 9% of influx in N-deprived plants, 10% at 10 M, 28% at 100 M, and 35% at 1.5 mM [NH ₄ ⁺ ]_o. Cytoplasmic [NH ₄ ⁺ ] was estimated at 6 m M in N-deprived plants, 2 mM at 10 M [NH ₄ ⁺ ]_o, 14 mM at 100 M, and 33 mM at 1.5 mM. Free-space [NH ₄ ⁺ ] was 84 M, 50 M, 700 M, and 8 mM, respectively. In comparison with previously published data on fluxes and compartmentation of NO ₃ ^– in white-spruce seedlings, results of this study identify a pronounced physiological preference of this species for NH ₄ ⁺ over NO ₃ ^– as an inorganic N source in terms of uptake and intracellular accumulation. The significant ecological importance of this N-source preference is discussed.The research was supported by a Natural Sciences and Engineering Research Council, Canada, grant to Dr. A.D.M. Glass and a University of British Columbia Graduate Fellowship to Herbert J. Kronzucker. Our thanks go to Dr. M. Adam and Mr. P. Culbert at the particle accelerator facility TRIUMF on the University of British Columbia campus for providing ¹³N, to Drs. R.D. Guy and S. Silim for providing plant material, and to Dr. M.Y. Wang, Mr. J. Bailey, Mr. J. Mehroke and Mr. P. Poon for essential assistance in experiments.     

Nitrogen uptake in a Norway spruce stand following ammonium sulphate application,fertigation, irrigation,drought and nitrogen-free-fertilisation

The above-ground accumulation of N,N uptake and litter quality resulting from improved or deteriorated availability of water and nutrients in a 25 year old Norway spruce stand in SW Sweden (as part of the Skogaby project) is presented. Treatment include irrigation; artificial drought; ammonium sulphate addition; N-free-fertilisation and irrigation with liquid fertilisers including a complete set of nutrients according to the Ingested principle (fertigation). At start of the experiment the stand contained 86.5 t dry mass and 352 kg N ha⁻¹. The following three years the annual N uptake in untreated trees was 32 kg N ha⁻¹ to be compared with the annual N throughfall of 17 kg ha⁻¹. Simultaneously, the treatment with ammonium sulphate and liquid fertilisation resulted in 48 and 56 kg ha⁻¹ y⁻¹, respectively, in treatment specific N-uptake following an application of 100 kg N ha⁻¹ y⁻¹. Addition of a N-free fertiliser resulted in improved N-uptake by 19 kg N ha⁻¹ y⁻¹ and irrigation by 10 kg N ha⁻¹ y⁻¹, compared to control. A linear relation between total above-ground dry mass production and N-uptake was found for trees growing with similar water availability. Dry mass production increased with increased water availability given the same N-uptake. It is concluded that the studied stand this far is not N saturated', as N fertilisation resulted in both increased N uptake and increased growth. Addition of a N-free-fertiliser resulted in increased uptake of N compared to the control, indicating an increased mineralisation rate or uptake capacity of the root system. The linear relation between N uptake and biomass production shows that at this study site N is a highly limiting factor for growth.