Measured and modelled differences in nutrient concentrations between rhizosphere and bulk soil in a Norway spruce stand

The gradient in soil characteristics from the bulk soil to the root surface is important to roots and to the organisms that live in the rhizosphere. Our ability to measure ion concentrations at the root surface is extremely limited, and models are largely untested. We used data from a well studied Norway spruce stand in SW Sweden to compare the measured difference in nutrient concentrations between rhizosphere and bulk soil with the difference predicted by a steady-state simulation model based on ecosystem budgets of nutrient uptake. The simulation model predicted depletion of NH₄, Ca, Mg, K in the rhizosphere, which shows that budgeted uptake rates were greater than the mass flow of bulk solution towards the root. In plots treated with ammonium sulphate, the model predicted an accumulation of S in the rhizosphere. In contrast, the observed rhizosphere concentrations were generally enriched in nutrients, relative to bulk soil. Collecting rhizosphere soil adhering to root surfaces may not be an appropriate method for describing the concentration gradient around the root. In addition, the simulation model omits some processes affecting conditions in the rhizosphere that are important to explaining nutrient uptake.     

Fine-root vitality in a Norway spruce stand subjected to various nutrient supplies

The vitality of fine roots in a Norway spruce stand subjected to application of ammonium sulphate (NS), wood ash (A) and nitrogen-free fertilizer (V) respectively, was investigated using i) vitality classification of fine roots based on morphological characteristics and ii) the triphenyl tetrazolium chloride (TTC) method of estimating dehydrogenase activity.Although the NS-treated areas showed a 30% increase in above-ground production in response to the NS-application, the vitality of the fine-root system in the humus layer appeared to be in a state of deterioration, as indicated by a decrease in fine-root biomass, an increased amount of dead fine (0–1 mm) and small (1–2 mm) roots, a decreased specific root length (SRL = fine root length/fine root dry weight) and an increased dehydrogenase activity. The impact of the the A and V treatments was reflected in a decrease in fine-root biomass and an increase in SRL. The results make it clear that in order to study the vitality of forest trees, both fine-root studies and studies of above-ground tree parts are necessary.     

Effects of nutrition and soil warming on stemwood production in a boreal Norway spruce stand

The boreal forest is expected to experience the greatest warming of all forest biomes. The extent of the boreal forest, the large amount of carbon contained in the soil, and the expected climate warming, make the boreal forest a key biome to understand and represent correctly in global carbon models. It has been suggested that an increase in temperature could stimulate the release of CO₂ caused by an increased decomposition rate, more than biomass production, which could convert current carbon sinks into carbon sources. Most boreal forests are currently carbon sinks, but it is unclear for how long in the future the carbon sink capacity of the boreal forest is likely to be maintained. The impact of soil warming on stem volume growth was studied during 6 years, in irrigated (I) and irrigated‐fertilized (IL) stands of 40‐year‐old Norway spruce in Northern Sweden. From May to October heating cables were used to maintain the soil temperature on heated‐irrigated plots (Ih and ILh) 5 °C above that on unheated control plots (Ic and ILc). After six seasons' warming, stem volume production (m³ ha^?1 a^?1) was 115% higher on Ih than on unheated (Ic) plots, and on heated and irrigated‐fertilized plots (ILh) it was 57% higher than on unheated plots (ILc). The results indicate that in a future warmer climate, an increased availability of nitrogen, combined with a longer growing season, may increase biomass production substantially, on both low‐ and high‐fertility sites. It is, however, too early to decide whether the observed responses are transitory or long lasting. It is therefore crucial to gain a better understanding of the responses of boreal forest ecosystems to climate change, and to provide data to test and validate models used in predicting the impact of climate change.     

Spatial and temporal variation of seepage water chemistry after femel and small scale clear-cutting in a N-saturated Norway spruce stand

The chemistry of seepage water was studied before and after small scale clear-cutting and femel cutting (removing 20% of the trees) between 1999 and 2002 at the H?glwald site in southern Bavaria. The interventions were performed in February 2000 on mature, N-saturated Norway spruce (Picea abies (L.) Karst.) stands with high NO ₃ ⁻ concentrations before felling. Seepage water was collected with suction cups at 40 cm soil depth in the following treatments: (I) a mature stand (control), (II) a femel-cut, and (III) a clear-cut. In the femel cut subvariants were created with suction cups (plots) at varying distances from pre-selected spruce, which were later removed. The femel treatment was replanted with beech (Fagus sylvatica L.) saplings. On the clear-cut, subvariants of planted beech (close to the stem, interstem area), planted spruce (interstem), or natural spruce regeneration were investigated. Clear-cutting caused high NO ₃ ⁻ peaks (average values up to 2750 μM) during 2000 and 2001 in all planted tree subvariants during times of comparatively low water fluxes. In contrast to peak concentrations, flux weighted yearly average concentrations showed different trends. In 2000, flux weighted yearly average NO ₃ ⁻ concentrations were significantly elevated, but only on the subvariants of the interstem area, which covered in the clear-cut plot ca. 62% of the area. However, the subvariant close to the stem (31% of clear-cut area), or the natural spruce regeneration subvariant (6% of clear-cut area) exhibited no significant felling effect. With respect to the whole treatment area, this resulted in no significant felling effect as compared with the control. In the next year (2001), flux weighted yearly average NO ₃ ⁻ concentrations were not significantly affected by clear-cutting, while the concentrations were even reduced for all of the clear-cut subvariants in 2002. On the subvariant natural spruce regeneration, NO ₃ ⁻ concentrations remained below the European limit of drinking water (806 μM) during almost the whole investigation period. Selective cutting resulted in slightly reduced NO ₃ ⁻ concentrations in 2000 and 2001 on the femel treatment. However, no significant effect could be detected for any subvariant in the femel-cut, even not for the subvariant with suction cups closest to the felled spruce. In contrast to many other investigations, clear-cutting did not increase the NO ₃ ⁻ problem on the treatment to a relevant extend. Quite contrary, a decline in NO ₃ ⁻ concentrations to values below the EU level for drinking water and levels below the control and femel treatment just 2 years after cutting were observed. Al³⁺ concentrations showed nearly the same trend as NO ₃ ⁻ , while Ca²⁺, Mg²⁺, and K⁺ concentrations were affected to a lesser degree. Only in 2002 was Ca²⁺ significantly lower on the clear-cut as compared to the femel treatment, but not compared to the control. Mg²⁺ increased in 2000 on the clear-cut subvariants in the interstem area, but decreased in the years 2001 and 2002. Changes could be observed for K⁺ only periodically on some subvariants, but not for the whole treatment area. Concentrations of SO ₄ ²⁻ , Na⁺, and Cl⁻ were reduced after clear-cutting and remained nearly unchanged after femel cutting.     

Short-term impacts of energy wood harvesting on ectomycorrhizal fungal communities of Norway spruce saplings

The increased demand for harvesting energy wood raises questions about its effects on the functioning of the forest ecosystems, soil processes and biodiversity. Impacts of tree stump removal on ectomycorrhizal fungal (EMF) communities of Norway spruce saplings were studied with 454-pyrosequencing in a 3-year field experiment replicated in 3 geographical areas. This is possibly the most thorough investigation of EMF communities associated with saplings grown on sites subjected to energy wood harvesting. To separate impacts of tree stump and logging residue removal on EMF and plant variables, we used three harvesting treatments with increasing complexity from patch mounding alone (P) to patch mounding combined with logging residue removal (RP), and patch mounding combined with both logging residue and stump removal (SRP). Saplings grown in uncut forests (F) served as references for harvesting treatments. A majority of sequences (>92%) and operational taxonomic units (OTUs, 55%) were assigned as EMF. EMF OTU richness, fungal community composition or sapling growth did not differ between harvesting treatments (P, RP and SRP), while EMF OTU richness, diversity and evenness were highest and sapling growth lowest in the undisturbed reference forests (F). The short study period may partially explain the similarities in fungal and sapling variables in different harvesting treatments. In conclusion, our results indicate that neither stump removal nor logging residue removal have significant additional negative impacts on EMF communities or growth of Norway spruce saplings in the short-term compared with the impacts of more conventional harvesting methods, including clear cutting and patch mounding.     

Interactive effects of soil warming and fertilization on root production, mortality, and longevity in a Norway spruce stand in Northern Sweden

The effects of soil warming and nitrogen availability on root production, longevity and mortality were studied using minirhizotrons in irrigation (C), fertilized (F), heated (H), and heated‐fertilized (HF) plots in a Norway spruce stand in northern Sweden from October 1996 to October 1997. Irrigation was included in all treatment plots. Heating cables were used to maintain the soil temperature in heated plots at 5°C above that in unheated plots during the growing season. A Kaplan–Meier approach was used to estimate the longevity of fine roots and Cox proportional hazards regression to analyze the effects of the H, F, and HF treatments on the risk of root mortality. The proportion of annual root length production contributed by winter–spring production amounted to 52% and 49% in heated plots and heated‐fertilized plots, respectively. The annual root length production in C plots was significantly higher than in other treatments, while the HF treatment gave significantly greater production compared with the F treatment. The risk of mortality (hazard ratio) relative to C plots was higher in H plots (358%) and F plots (191%). The interaction between heating and fertilizing was strongly significant. The increase in the risk of root mortality in combined fertilization and heating (103%) was lower than that in the H or F plots. The results show that nitrogen addition combined with warmer temperatures decreases the risk of root mortality, and fine root production is a function of the length of the growing season. In the future, fertilization combined with the warmer temperatures expected to follow predicted climatic change may increase root production in boreal forests at low fertility sites.     

Needle chemistry in young Norway spruce stands after application of crushed wood ash

Nutrient concentrations in current and 1-year-old needles were analyzed annually for 5 years after application of hardened wood ash in 1–4-year-old Norway spruce (Picea abies (L.) Karst.) stands within a range of climate and fertility gradients. At each site, 3000 kg ha^–1 hardened wood ash of two types, Nymölla and Perstorp, was applied in a randomized block design. Wood ash Nymölla contained 12 kg ha^–1 P, 30 kg ha^–1 K, 891 kg ha^–1 Ca, 72 kg ha^–1 Mg and wood ash Perstorp contained 12 kg ha^–1 P, 60 kg ha^–1 K, 486 kg ha^–1 Ca, and 60 kg ha^–1 Mg. The ash was intended to compensate for nutrients removed at the preceding harvest when logging residues were collected and removed from the site (whole-tree harvesting). The climate gradient included four climate zones throughout Sweden and each of these included a fertility gradient of three sites classified according to their ground vegetation type. There were no effects on nutrient concentrations in the needles 1 year after the application of wood ash. Five years after ash application, the concentrations of P, K and Ca in current and 1-year-old needles were higher than in the control plots. The results were consistent over all stands, irrespective of climate zone and fertility status. P and K concentrations were higher in spruce needles from plots treated with Perstorp wood ash, whereas Ca concentrations were higher in those of Nymölla treated plots. Analyses across all study sites revealed a treatment effect in terms of increased ratios of P:N, K:N and Ca:N in 1-year-old needles. The ratio P:N tended to increase with time in the Perstorp wood ash treatment compared with the control. The needle concentrations of Mg and S were not affected by the ash applications. The increase in needle nutrient concentrations after application of hardened wood ash suggests that wood ash recycling could be used in order to replace nutrients removed at whole-tree harvesting.     

Nutrient status in needles of Norway spruce and Scots pine following harvesting of logging residues

Nutrient concentrations in current and 1-year old needles from two Picea abies (L.) Karst and two Pinus sylvestris L. stands in Sweden were determined 8–10, 16–18 and 22–24 years after clear-felling and experimental manipulation of harvesting intensity. On all sites, three levels of harvest intensity had been applied in a randomized block design (n=4); (i) conventional stem-only harvesting, where all logging residues (i.e. tops, branches and needles) were evenly distributed on the ground, (ii) harvesting all above-ground tree parts except needles and (iii) above-ground whole-tree harvesting (no residues left on site). At stand age 8–10 years, nitrogen concentrations in the current year needles in plots where all residues or needles only were retained were higher than in whole-tree harvested plots, whereas concentrations of K, Ca and Mg were lower. The latter response was interpreted as a dilution effect. P:N, K:N, Ca:N, Mg:N, Mn:N and Zn:N were in general higher after whole-tree harvesting treatments than after the treatments where all residues or only needles had been left on site. At stand age 16–18 years, no significant differences in nitrogen concentrations were observed between treatments, but the levels of Ca, Mg and Mn in both current and 1-year-old needles were lower after whole-tree harvesting than after the treatments where logging residues remained on site. By contrast, potassium levels in the foliage were highest in treatments where only the needles were left on site, whereas the lowest levels were observed for treatments where all residues was left. At stand age 22–24 years, the treatment effects had diminished, except for the effects on Ca and K on the southern Norway spruce stand. It is concluded that the nutrient release from logging residues enhances nutrient uptake in trees of the succeeding forest generation, but this effect does not occur simultaneously for all elements. This revised version was published online in June 2006 with corrections to the Cover Date.     

Influence of Norway spruce seedlings on the nutrient availability in mineral soil and forest floor material

A greenhouse experiment was performed to evaluate the effect of Norway spruce (Picea abies (L.) Karst.) seedlings on net nutrient availability in five different growing media containing F- or H-layer and mineral soil originating from a haplic podzol in northern Sweden. The initial total amounts of eight nutrient elements (N, K, P, Ca, Mg, Mn, Fe, Zn) and exchangeable amounts of the same elements were analyzed in pots with or without spruce seedlings. In the planted pots seedling nutrient uptake was also estimated. After 26 weeks, higher net nutrient availability with seedlings was found in 25 out of the 40 (62%) growing media and nutrient element combinations. A positive seedling effect on net nutrient availability might be explained by rhizodeposition stimulating the soil microorganism activity and accelerating the weathering of minerals or by seedling roots promoting the nutrient providing processes through changes in soil chemical and physical properties. Nitrogen availability was primarily affected by what part of the forest floor the growing medium contained although the positive response to seedling presence was apparent. The positive net availability response of P, Ca, Mg, Mn, Fe and Zn to seedling presence was on the other hand relatively strong. In the case of P, K, and Zn the growing medium composition (if the F- and H-layer was pure or mixed with mineral soil) was also an important factor for the estimated net availability. Pure F-and H-layer provided greater P- and K-availability while the availability of Zn increased when mineral soil was added. The influence of growing plants ought to be considered when soil samples are used for assessing the nutrient availability.     

Spatial variability of throughfall chemistry and selected soil properties as influenced by stem distance in a mature Norway spruce (Picea abies,Karst.) stand

Our aims were to investigate the spatial variability of throughfall chemistry and soil parameters as influenced by stem distance and to evaluate the implication of the observed systematic and random patterns for the sampling strategy.One hundred throughfall samplers with a sampling area of 106 cm² each were established in a systematic grid around 5 trees in a mature Norway spruce; site of the Fichtelgebirge (Germany). One hundred soil cores were taken with an auger of 50 cm² next to the throughfall samplers. Soil samples were stratified according to genetic soil horizons and analysed for pH, exchangeable NH₄ ⁺, SO₄ ^2– and total-S. Throughfall samples were collected over a period of 6 months. For each sampler an aliquod sample was mixed over the observation period and analysed for major ions.The spatial variability of the element concentrations in throughfall, expressed by the coefficient of variance, was 21–164%, depending on the element considered. For precipitation volume, the coefficient of variance was only 3%. The distance to the stem influenced most element concentrations in throughfall with increasing concentrations approaching the stem. Steepest gradients were observed in case of SO₄ ^2– and H⁺.The spatial variability of the investigated soil parameters was also very high with the exception of pH. The SO₄ ^2– content of the forest floor reflected the gradients observed in throughfall, while for the other investigated soil parameters and soil horizons no significant relations to stem distance were found.To determine site representative throughfall concentrations and soil properties with the sample volumes and time intervals we used, the number of samples required to get a statistical error of less than 10% (with 95% probability) can be very high. In case of throughfall, more than 100, and in case of the soil parameters, more than 300 replicates would be required.     

Experimental manipulations of water and nutrient input to a Norway spruce plantation at Klosterhede,Denmark

At Klosterhede, Denmark experimental manipulations of water and nutrient inputs to a forest stand were carried out under a 1200 m² roof simulating i) summer drought, ii) removal of N and S input, and iii) optimal nutrition and water availability. In practise, manipulation of element fluxes in a complex system like a forest may cause unintended changes such as climatic effects from the roof which may interfere with the results of the intended manipulation. This paper illustrates the extent and effects of such unintended changes.The most important climatic change under the roof was a 50% reduction of photosynthetic light which caused a decrease in moss cover on the ground floor. Soil temperatures under the roof were decreased 0.3°C during summer and increased 0.2°C during winter. Air humidity was not changed. Throughfall water amount decreased close to the stems whereas element concentrations in throughfall increased close to the stems and with tree size on the ambient plots. This distinct spatial pattern was changed under the roof to a more variable water distribution from the sprinkling system and to constant element concentrations in the sprinkling water. This loss/change of spatial variability in the input was reflected in the soil solution. The concentration patterns found in soil water (increasing close to the stem and with tree size) in the ambient plots were no longer observed under the roof.Exclusion of throughfall by the roof disturbed the internal cycle of nutrients leached from the canopy. The removal of canopy leached Ca and K by the roof caused a decline in soil water concentrations, even on the drought plot where natural throughfall was only cut off during two summer months. The internal cycle of Ca and K had to be restored by addition of Ca and K under the roof. Further, leachable elements and soluble compounds may be washed out from litterfall during the collection period on the roof, and thus lost from the internal cycle by by exclusion of throughfall.It is important to consider these problems in the design of new manipulation experiments and in the interpretation of the results. Some recommendations are given.     

Experimental manipulation of water and nutrient input to a Norway spruce plantation at Klosterhede,Denmark

Water and nutrient supply to forest trees are major factors controlling tree growth and forest vitality. Therefore, changes in the supply of water and nutrients to the trees may be important contributing reasons to the forest damages observed in Europe. Such changes may be caused by several factors, e.g. air pollution, soil acidification and climate change. The present study investigates possible effects on tree growth, growth related parameters and nutrition related to changes in the water and nutrient supply. Water and nutrient supply to the forest soil was manipulated in three roof covered plots. The treatments consisted of 1) summer drought, 2) irrigation and 3) combined irrigation and fertilizer application (fertigation). The results from the roof covered treatment plots were compared to a control plot without roof. Increased supply of water during the spring and early summer increased the diameter growth, whereas application of nutrients in addition to irrigation had no additional effect on tree growth. Addition of nutrients increased the needle content of P, Mn, Ca and Mg. Extended summer drought for 2 months had no effect on the tree growth or other growth parameters, but drought reduced the root development in the upper soil layer. Furthermore, drought induced significant stress symptoms by increasing the cone shredding substantially.     

The impact of spruce aphids on nutrient flows in the canopy of Norway spruce

1 This study investigated the effects of honeydew from aphids in the canopy of Norway spruce (Picea abies (L.) Karst.) on the nitrogen chemistry of throughfall using a rainfall simulation experiment. Throughfall collected beneath infested trees was compared with that from beneath uninfested trees, while standardizing the quality and quantity of the precipitation and plant age. 2 Honeydew excreted by Cinara pilicornis (Hartig) and C. costata (Zett.) significantly increased the concentrations of dissolved organic carbon (DOC) and hexose-C in throughfall. The average concentrations of nitrogenous compounds (NH₄-N, NO₃-N) in throughfall collected beneath infested trees were significantly lower than beneath uninfested trees. 3 Multiple regression analysis indicated that the amount of rain and NH₄-N concentrations were the best predictors of the concentrations of dissolved organic nitrogen (DON) in throughfall. Parameters that were closely associated with the level of infestation (DOC, hexose-C concentrations) did not have a direct relationship with DON. About 40% of the reduction in the concentration of DON in the throughfall was attributed to aphid–micro-organism interactions. 4 Particle amino nitrogen (PAN)-concentrations were highest under infested trees in July after aphid numbers had declined, indicating a concomitant decline in microbial biomass after honeydew becomes a limiting resource. 5 The comparison of the concentrations of different nitrogen compounds in throughfall of infested and uninfested trees indicated that aphids affect the carbon and nitrogen cycles in the phyllosphere by providing energy that fuels the metabolism of the micro-organisms. These processes seem to occur very rapidly. 6 We discuss the significance of the results and the prospects of linking the ecology of micro-organisms and herbivores with flows of nutrients through the canopy of trees.     

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     

Effects of long-term CO2 enrichment and nutrient availability in Norway spruce. II. Foliar chemistry

 Branches on 30-year-old Norway spruce trees [Picea abies (L.) Karst.] were exposed to ambient (AMB) or ambient plus 350 μmol CO₂ mol^–1 (EL) for 4 years (except winters), using the branch bag technique (BB). The trees were growing on plots with low (control) and high (irrigated-fertilised) availability of soil nutrients. The seasonal variation in foliar macronutrients and non-structural carbohydrates in current and 1-year-old shoots was monitored throughout the treatment period. When the branches were harvested at the end of treatment, macronutrients were analysed in five age classes of foliage. The concentration of all elements, except magnesium, generally increased in AMB, i.e. a ’bag effect’, but decreased as an effect of EL, i.e. a ’CO₂ effect’. At the final harvest K, P, N and S were reduced in young needles by EL, whereas Mg was reduced in older needles on both plots. A change in needle morphology by EL possibly caused a dilution effect in irrigated-fertilised needles, but not in control needles. Reductions in K and Mg are suggested to be an effect of increased phloem transport from the branch, in consequence of higher rates of carbon fixation in EL. Foliage in BBs had higher concentration of Ca, but there was no significant effect of the EL-treatment, indicating that elevated CO₂ had no effect on stomatal conductance. Quinic acid concentration decreased, but shikimic acid concentration increased in BBs, independently of CO₂ treatment. Concentrations of starch and sugars increased in the EL-treatment, but pinitol decreased. Received: 23 October 1998 / Accepted: 1 December 1998     

Short-term effects of manipulated increase in acid deposition on soil, soil solution chemistry and fine roots in Scots pine (Pinus sylvestris) stand on a podzol

A manipulated increase in acid deposition (15 kg S ha⁻¹), carried out for three months in a mature Scots pine (Pinus sylvestris) stand on a podzol, acidified the soil and raised dissolved Al at concentrations above the critical level of 5 mg l⁻¹ previously determined in a controlled experiment with Scots pine seedlings. The induced soil acidification reduced tree fine root density and biomass significantly in the top 15 cm of soil in the field. The results suggested that the reduction in fine root growth was a response not simply to high Al in solution but to the depletion of exchangeable Ca and Mg in the organic layer, K deficiency, the increase in NH₄:NO₃ ratio in solution and the high proton input to the soil by the acid manipulation. The results from this study could not justify the hypothesis of Al-induced root damage under field conditions, at least not in the short term. However, the study suggests that a short exposure to soil acidity may affect the fine root growth of mature Scots pine.     

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.     

Effects of wood ash and nitrogen fertilization on fine root biomass and soil and foliage nutrients in a Norway spruce stand in Finland

We determined the effects of wood ash fertilization, given together with nitrogen (WAN), and nitrogen given together with P, B and Cu (SSF), on soil and foliage nutrients and fine root biomass in a 45-year-old Norway spruce stand in southern Finland. Fine roots were sampled 9 years, and the soil 10 years after ash (3 t/ha) and nitrogen (150 kg/ha) application. Fine root biomass tended to be lower, the necromass higher, and the fine root distribution relatively deeper on the WAN than on the control and SSF plots. The response of fine root biomass to WAN was probably related to changes in soil acidity. pH, base saturation, total and extractable concentrations of Ca, K, Mg and P, and total B, Cd, Mn, Ni and Zn concentrations in the organic layer were significantly higher on the WAN plots than on the SSF and the control plots with no ash and nutrient addition. On the WAN plots, the pH was 1.2 pH-units higher, the exchangeable Ca concentrations fourfold and those of Mg over twofold compared to the control plots. WAN increased the concentrations of K but decreased those of Mn and Ni in the needles compared to the control and SSF treatment. Even though ash and nitrogen fertilisation tended to decrease the fine root biomass, this decrease was not likely to affect tree growth during a 10-year period.     

Effects of harvesting on nutrient leaching in a Norway spruce (Picea abies Karst.) ecosystem on a Lithic Leptosol in the Northern Limestone Alps

On a heavily karstified site in the Northern Limestone Alps (Austria), nutrient budgets and leaching in Norway spruce stands were investigated along a chronosequence (clearcut, 10-year-old plantation (25% cover of planted and naturally regenerated spruce and larch, 75% weed cover) and mature stand). The soils were Lithic Leptosols on very pure limestone. Nutrient fluxes were studied during three growth periods (4–5 months each). Despite of inorganic nitrogen inputs from precipitation between 5 and 10 kg ha^–1, inorganic nitrogen output with seepage water from the mature stand and the regeneration plot was only 0.5–1.2 kg ha^–1 during these periods. In the first and second growth periods after clearcut, inorganic N fluxes with seepage increased to 20 and 30 kg ha^–1, respectively, declining in the third growth period to 8 kg ha^–1. DON output during the growth period was between 3 and 6 kg ha^–1 in the mature stand and 7 and 11 kg ha^–1 in the clearcut as well as in the regeneration plot. K output rates achieved 30 kg ha^–1 in the first, 20 kg ha^–1 in the second and 9 kg ha^–1 in the third growth period after clear-cutting while output rates during the growth periods were less than 2 kg ha^–1 in the mature stand and in the regeneration plot. K pools in the humus layer were only 150–210 kg ha^–1, total K pools including above and below ground biomass in the mature stand were 360 kg ha^–1. Thus, post-harvest hydrological losses comprise a substantial depletion of K for this specific ecosystem. Since precipitation is high in this area (1400 mm a^–1), forest growth is limited by nutrient rather than by water supply. Needle analyses already indicate a deficient potassium supply. Harvesting and post-harvesting losses of K in combination with elevated nitrogen deposition may have negative influences on the stability of forest stands on the studied sites.