Phosphorus supply and cycling at long-term forest monitoring sites in Germany

In this study, the supply and input–output balances of phosphorus (P) were investigated for a 10-year-period at 85 long-term monitoring sites in German forest ecosystems under the European Level II programme. These sites encompass 23 European beech (Fagus sylvatica L.) stands, 9 oak stands comprised of common oak (Quercus robur L.) and/or sessile oak (Quercus petraea Liebl.), 20 Scots pine (Pinus sylvestris L.) and 33 Norway spruce (Picea abies H.Karst.) stands. We quantified P concentrations in needles and leaves, P inputs from the atmosphere, P outputs through leaching and harvesting, and total P in the soil and humus layers. The P concentrations in European beech leaves from two sites (>1 mg P g⁻¹ dry weight), and in Norway spruce needles from four sites (>1.2 mg P g⁻¹ dry weight), were deficient over several years. In contrast, the oak and Scots pine sites were well supplied with P. When P removal through harvesting was disregarded, P balances were positive or stable (median 0.21 kg P ha⁻¹ a⁻¹). With harvesting, balances were mostly negative (median −0.35 kg P ha⁻¹ a⁻¹), with long-term P removal from the forest ecosystems.     

Über die Ursachen und Kriterien der Immissionsresistenz bei Fichte,Picea abies (L.) Karst

Causes and criteria of resistancë to air pollution in Norway spruce. 3. Tolerance of toxic materials (“internal” resistance). Comparing relatively resistant and not resistant Norway spruce clones, an attempt was made to find differences by investigating the relation of needle colour and absorption of sulfur, needle pH-value and buffer capacity, sulphur distribution within several of their needle components and within the total plant, the ratio of organically fixed sulfur to total sulfur content of noedles, and content of sulfhydril groups, protein and chlorophyll of needles. Significant differences were shown only in buffer capacity of unfumigated needles, and in thc propornon of organically fixed sulfur before and after SO₂ fumigation.     

Short-term impacts of harvesting and burning disturbances on physical and chemical characteristics of forest soils in western Newfoundland,Canada

The short-term impacts of prescribed burning were determined for the physical and chemical properties of a forest soil on a clear-cut in western Newfoundland, Canada. The experimental site was predominantly covered by black spruce trees (Picea mariana Mill.) that were harvested in 1996. In August 1998, prescribed burning removed most of the logging residues left on the site after harvesting, all above-ground surface vegetation, and parts of the upper humus layer. In October 1998, field samples were taken from four replicated burned plots and four replicated unburned plots. In each study plot, soil samples were taken from the organic layer (F+H) and from the top 10 cm of the mineral soil. In the burned treatments, mass of the humus layer (F+H) was reduced by 24% and, because of the accretion of basic ash materials, acidity of the humus layer was also reduced by up to 1 unit. In the organic layer and the mineral soil, total contents of Mg, Ca, and P, extractable Mg and Ca, available P, sum of NH₄Cl-extractable cations, and C/N ratios were increased by burning, while total C and N as well as total and extractable K remained unaffected by burning.     

The effect of nitrogen fertilization on fungistatic phenolic compounds in roots of beech (Fagus sylvatica) and Norway spruce (Picea abies)

The effect of nitrogen fertilization on fungistatic phenolic compounds in fine roots of beech and Norway spruce growing in afforestation plots was analysed. The plots were situated at two sites in Switzerland on acidic soil with low base saturation. For 9 years, the trees have been treated with dry ammonium nitrate to give 0, 10, 20, 40, 80, 160 kg N ha^?1 year^?1, respectively. The phenolic compounds responded differently to fertilization. Fine roots of beech showed a significant decrease of (?)‐epicatechin and piceatannol with increasing nitrogen fertilization. The concentration of protocatechuic acid was increased with fertilization. Roots of fertilized Norway spruce showed significantly decreased concentrations of 4‐hydroxyacetophenone and piceatannol. The mycelial growth of three isolates each of Heterobasidion annosum s.l. and Cylindrocarpon destructans was tested on agar media containing various phenolic compounds in concentrations found in fine roots of Norway spruce (Picea abies) and beech (Fagus sylvatica). All three H. annosum isolates were inhibited by p‐coumaric acid and (?)‐epicatechin. Two isolates were inhibited by another four phenolic compounds (p‐hydroxybenzoic acid, 4‐hydroxyacetophenone, piceatannol and protocatechuic acid), one by (+)‐catechin. Two of three C. destructans isolates were inhibited by all phenolic compounds except for (+)‐catechin which affected only one isolate, one isolate did not respond at all.     

Dissolved Al reduces Mg uptake in Norway spruce forest: Results from a long-term field manipulation experiment in Norway

Dissolved aluminium (Al) in soils, mobilized by acid deposition, is considered a threat to forest health through hampering root growth and nutrient uptake. Since the end of the 1980s dissolved Al in forest soil water plays a key role in the assessment of critical loads of acid deposition. So far, most evidence for toxicity of dissolved Al in forest soil water is based on nutrient solution studies and pot experiments. Here, we present results from one of the few in situ ecosystem-scale forest manipulation experiments to study the effect of Al on mature forest trees. A plotwise addition of dilute AlCl₃ was conducted during seven years in an even-aged spruce forest (Picea abies) in an area in Norway with low acid deposition. Soil solution concentrations of Al were increased to potentially toxic levels (up to 500 μmol L⁻¹) and base cation (Ca + Mg + K) to inorganic Al ratios in the soil solution in the root zone were mostly below 1 in the Al-addition treatments. In the control treatment (only water addition) Al concentrations did not exceed 15 μmol L⁻¹ and base cation to inorganic Al ratios were above 1. The toxic effects of Al on fine root growth and plant growth found in hydroponic studies and pot trials are not confirmed by this field manipulation. However, magnesium (Mg) contents in needles decreased significantly and persistently in plots with elevated Al concentrations, whereas the needle Ca content did not respond. The depletion of the Mg content in needles is suggested to be due to antagonistic effects of high Al concentrations at the root surface, consistent with observed reductions in Mg to Al ratio of inner bark. This study clearly supports a role for Al in critical load functions for forests as dissolved Al causes a decrease in uptake of Mg. However, other signs of reduced forest vitality were not observed. Soil base cation status may need to be included in risk evaluations of forest health under acid deposition.     

Analyses of the impact of changes in atmospheric deposition and climate on forest growth in European monitoring plots: A stand growth approach

During the last 15 years a number of studies have shown increasing forest growth in central Europe, rather than a decline as was expected due to negative effects of air pollution. We have here used data from intensive monitoring plots spread over Europe for a five year period in order to examine the influence of environmental factors on forest growth. Evaluations focussed on the influence of nitrogen, sulphur and acid deposition, temperatures, precipitation and on a drought index calculated as deviation from the long-term mean. The study included the main tree species Norway spruce, Scots pine, common beech as well as European and sessile oak and was based on data from 363 plots. As many other factors besides nitrogen and temperature influence tree growth, expected stem volume increments were modelled using site productivity, stand age and a stand density index. Relative volume increment was then calculated as actual increment in % of expected increment. The site productivity, assumed to be given by site conditions and past environmental conditions, was either taken from expert estimates or computed from site index curves from northern, central and southern Europe. The model explained between 18% and 39% of the variance with site productivity being positively related and age negatively related to actual increment. The various models and statistical approaches were fairly consistent, and indicated a fertilizing effect of nitrogen deposition, with slightly above one percent increase in volume increment per kg of nitrogen deposition per ha and year. This was most clear for spruce and pine, and most pronounced for plots having soil C/N ratios above 25. Also, we found a positive relationship between relative increment and summer temperature, i.e. May–August mean temperature deviation from the 1961–1990 means. The cause–effect relationship here is, however, less certain. Other influences were uncertain. Possibly, sulphur and acid deposition have effects on growth, but these effects are obscured by, and outweighed by the positive effect of nitrogen deposition, because of collinearity between these variables. Drought effects were uncertain also, and one reason for this might be large uncertainties in the precipitation data: precipitation measured on some 50% of the plots correlated poorly with the precipitation data obtained from Europe-wide databases. The major finding of this study was a positive relationship between higher than normal volume increment on one hand and nitrogen deposition on the other hand.     

Effects of the needle bladder rust (Chrysomyxa rhododendri) on Norway spruce: implications for subalpine forests

Norway spruce trees in the subalpine forests of the European Alps are frequently attacked by the needle rust Chrysomyxa rhododendri. The obligate parasite undergoes a complex life cycle with a host shift between rhododendrons (Rhododendron sp.) and Norway spruce [Picea abies (L.) Karsten] and causes a yellowing and defoliation of the current-year needles in summer. Infected trees show several anatomical, morphological and physiological modifications, including a decrease in pigment content and net photosynthesis of infected needles, lower biomass production and reduced radial and height growth. The consequences are diminished timber yield and cripple growth. Because of repeated heavy infections in recent years, forest managers report increasing difficulties in both natural regeneration and afforestation at high elevation sites, where rhododendrons occur. This review gives a summary of the present knowledge about the effects of C. rhododendri infections on Norway spruce, including so far unpublished findings and with particular attention to the phenomena of resistant trees. Implications for subalpine forests and counter strategies are discussed.     

Effects of slash retention and wood ash addition on fine root biomass and production and fungal mycelium in a Norway spruce stand in SW Sweden

In the study reported here we examined the short-term effects (1–3 years) of slash retention (SR) and the long-term effects (13–15 years) of wood-ash application (A) on fine roots and mycorrhizae in a 40-year-old Norway spruce forest in southwest Sweden. Soil cores were used to obtain estimates of the biomass (g m⁻²) of roots in three diameter classes (<0.5, 0.5–1 and 1–2 mm), root length density (RLD), specific root length (SRL) and mycorrhizal root tip density (RTD). Fine root (<1 mm) length production and mortality, and mycelium production, were estimated using minirhizotron and mesh bag techniques, respectively. Compared with the control plots (C), the biomass of fine roots in diameter classes <0.5 mm and 0.5–1 mm was significantly higher in A plots, but lower in SR plots. In addition, RLD was significantly lower in the humus layer of SR plots than in the humus layers of C and A plots, but not in the other layers. None of the treatments affected the SRL. In all soil layers, the SR treatment resulted in significant reductions in the number of ectomycorrhizal root tips, and the mycelia production of fungi in mesh bags, relative to the C treatment, but the C and A treatments induced no significant changes in these variables. Fine root length production in the C, A and SR plots amounted to 94, 87 and 70 mm tube⁻¹ during the 2003 growing season, respectively. Fine root mortality in treated plots did not change over the course of the study. We suggest that leaving logging residues on fertile sites may result in nitrogen mineralisation, which may in turn induce reductions in root biomass, and both root and mycelium production, and consequently affect nutrient uptake and the accumulation of organic carbon in soil derived from roots and mycorrhizae.     

The effects of nitrogen fertilization on the ultrastructure of mesophyll cells of conifer needles in northern Finland

Cytological injuries in needles of Scots pine (Pinus sylvestris L.) and Norway spruce [Picea abies (L.) Karst.] were studied at light and electron microscope level in northern Finland. Needle samples were taken from fertilized (mainly with nitrogen) and nonfertilized trees during winter 1980. The results are discussed in relation to the severe climate and growth response of the trees.     

Vergleichende Feinwurzeluntersuchungen an gesunden und erkrankten Altfichten auf zwei Standorten in Bayernein Beitrag zur Waldsterbensforschung

Fine root studies in two old growth spruce forests affected by “Wadsterben” in Bavaria. Fine roots of healthy and declining Norway spruce (Picea abies) were investigated. We observed fewer active myeorrhizac associated with affected spruce in two old growth Forests. It has been hypothesised that air pollution is dimaging the needles resulting in less photosynthate and that the reduction of active mycorrhizae is due to such an initial weakening of the tree.     

The impact of atmospheric deposition and climate on forest growth in European monitoring plots: An individual tree growth model

In the climate change discussion, the possibility of carbon sequestration of forests plays an important role. Therefore, research on the effects of environmental changes on net primary productivity is interesting. In this study we investigated the influence of changing temperature, precipitation and deposition of sulphur and nitrogen compounds on forest growth. The database consisted of 654 plots of the European intensive monitoring program (Level II plots) with 5-year growth data for the period 1994–1999. Among these 654 plots only 382 plots in 18 European countries met the requirements necessary to be used in our analysis. Our analysis was done for common beech (Fagus sylvatica), oak (Quercus petraea and Q. robur), Scots pine (Pinus sylvestris) and Norway spruce (Picea abies). We developed an individual tree growth model with measured basal area increment of each individual tree as responding growth factor and tree size (diameter at breast height), tree competition (basal area of larger trees and stand density index), site factors (soil C/N ratio, temperature), and environmental factors (temperature change compared to long-term average, nitrogen and sulphur deposition) as influencing parameters. Using a mixed model approach, all models for the tree species show a high goodness of fit with Pseudo-R² between 0.33 and 0.44. Diameter at breast height and basal area of larger trees were highly influential variables in all models. Increasing temperature shows a positive effect on growth for all species except Norway spruce. Nitrogen deposition shows a positive impact on growth for all four species. This influence was significant with p < 0.05 for all species except common beech. For beech the effect was nearly significant (p = 0.077). An increase of 1 kg N ha⁻¹ yr⁻¹ corresponds to an increase in basal area increment between 1.20% and 1.49% depending on species. Considering an average total carbon uptake for European forests near 1730 kg per hectare and year, this implies an estimated sequestration of approximately 21–26 kg carbon per kg nitrogen deposition.     

The nutrient status of Norway spruce in pure and in mixed-species stands

Atmospheric deposition of N and S appears to have caused nutrient imbalance in Norway spruce stands in southern Sweden. This calls for a change of forest management to procedures that promote nutrient balance. Studies have shown lower soil acidity in Norway spruce/deciduous mixed stands than in spruce monocultures, but the tree nutrient status in such mixtures has not been much investigated so far.

The nutrient status of Norway spruce foliage and top mineral soil chemistry in monocultures and in stands mixed with beech, birch, or oak was investigated through paired comparisons on 30 sites in southern Sweden (27 sites) and eastern Denmark (three sites). In total, 45 mixed stands and 34 pure stands were included in the study.

Spruce needles from mixed stands had higher concentrations and ratios to N of K, P, and Zn than needles from pure spruce stands. Among the mixed stands, the K status appeared to be positively correlated with the percentage of deciduous tree basal area. Soil samples from mixed stands had a higher Mg concentration, base saturation, and BC/Al ratio than soil samples from pure stands. The spruce needle nutrient status was comparable in pure stands on fertile sites and in mixed stands on poor sites. We did not detect any differences in spruce tree growth between pure and mixed stands.

This paper discusses possible reasons for a positive effect on the tree nutrient status in mixed-species stands and the possibility of using mixed-species stands as a forest management procedure to avoid nutrient imbalance.     

Importance of soil extractable phosphorus distribution for mature Norway spruce nutrition and productivity

Phosphorus is an essential nutrient for forest growth. In this study, we assessed the impact of soil extractable phosphorus using two simple extraction methods on nutrition and productivity of Norway spruce in sixteen mature forest stands on different bedrocks and soils in Bavaria, Southern Germany. Representative trees were sampled for needles, twigs, branches, stem bark, and stem wood. Total phosphorus content in the tree parts and soil phosphorus stock extractable with citric acid and sodium bicarbonate up to a soil depth of 80 cm were determined. We found that easily soil extractable phosphorus is a suitable indicator for estimating phosphorus uptake and stand productivity in Norway spruce. In contrast, organic layer phosphorus showed no significant correlation with aboveground biomass phosphorus contents. In the biomass, the highest phosphorus contents were measured in young needles and twigs, but the highest correlation with soil phosphorus was detected for phosphorus contents in needles and bark. The stock of phosphorus extracted by citric acid down to 40 cm soil depth revealed the best correlation with phosphorus in needles and bark. Therefore, as a supplemental or alternative method to needle analysis, our study suggests the use of phosphorus contents in stem bark to evaluate tree phosphorus nutrition. These results highlight the suitability of the citric acid soil extraction method to characterize plant available phosphorus in Norway spruce ecosystems.     

The influence of different forest organic matter on the growth of one-year old planted Norway spruce seedlings in a greenhouse experiment

A greenhouse trial was carried out to improve the knowledge of how forest organic matter could be utilized in site preparation and the choice of planting spot for Picea abies (L.) Karst. One-year old Norway spruce seedlings were grown for 26 weeks in pots containing pure mineral soil, forest organic matter in three different states of decomposition and combined treatments where the organic matter was mixed with or placed on the surface of the mineral soil (volume proportions of organic:mineral of 1:2). Watering was adapted to the water retention characteristics of each growing medium in order to keep the soil water potential between -4 and -5 kPa in all pots.In its pure form as well as combined with mineral soil, the moderately decomposed mor organic matter gave a higher seedling dry weight increment than the corresponding growing media containing the more decomposed humus. Adding mor or humus to mineral soil increased the seedling dry weight increment by 30–140% and 10–40%, respectively. The largest seedlings grew in the pure mor and humus. Mixing these forms of organic matter with mineral soil appeared to reduce seedling growth as compared to placing them on the mineral soil surface. The increment increase compared to plain mineral soil mostly consisted of proleptic growth and was probably due to the improved nutrient availability of the growing media. Fresh organic matter (chopped needles and twigs) seemed to impair seedling root function and reduced the dry weight increment by 30–50% as compared with pure mineral soil.The results suggest that as long as the water and temperature requirements are fulfilled planting Norway spruce seedlings without scarification and/or gathering extra mor and humus to the planting position should improve seedling growth as compared to the growth following scarification. Unmixed fresh needles and twigs or fresh needles and twigs combined with plain mineral soil should be avoided when planting Norway spruce seedlings.     

Some factors limiting denitrification in slurries of acid forest soils

Denitrifying enzyme activity (DEA) is defined as N₂O production rate over the first eight hours of anaerobic incubation of soil slurries at 15°C with added nitrate and acetylene. Organic fraction explained 81 % of the spatial variation in DEA among soil horizons of beech (Fagus), alder (Alnus), and spruce (Picea) forests, and spruce ciearcuts. Nitrate and pH limit denitrification in anaerobic O horizons at 15°C. Thick, poorly drained O horizons, such as in alder forest, have a high capability for denitrification. Failure to recover all added nitrate in O horizon incubations suggested both production of unmeasured NO and dissimilatory reduction of nitrate to ammonium. In mineral horizons, N₂O production was small and could not be increased by increasing nitrate, carbon, pH, and phosphorus; some other factor appears to prevent significant denitrifier activity in acid forest mineral horizons.     

Vegetation change and forest regeneration on the Kenai Peninsula,Alaska following a spruce beetle outbreak, 1987–2000

Forests of the Kenai Peninsula, Alaska experienced widespread spruce (Picea spp.) mortality during a massive spruce beetle (Dendroctonus rufipennis) infestation over a 15-year period. In 1987, and again in 2000, the U.S. Forest Service, Pacific Northwest Research Station, Forest Inventory and Analysis Program conducted initial and remeasurement inventories of forest vegetation to assess the broad-scale impacts of this infestation. Analysis of vegetation composition was conducted with indirect gradient analysis using nonmetric multidimensional scaling to determine the overall pattern of vegetation change resulting from the infestation and to evaluate the effect of vegetation change on forest regeneration. For the latter we specifically assessed the impact of the grass bluejoint (Calamagrostis canadensis) on white spruce (Picea glauca) and paper birch (Betula papyrifera) regeneration. Changes in vegetation composition varied both in magnitude and direction among geographic regions of the Kenai Peninsula. Forests of the southern Kenai Lowland showed the most marked change in composition indicated by relatively large distances between 1987 and 2000 measurements in ordination space. Specific changes included high white spruce mortality (87% reduction in basal area of white spruce >12.7 cm diameter-at-breast height (dbh)) and increased cover of early successional species such as bluejoint and fireweed (Chamerion angustifolium). Forests of the Kenai Mountains showed a different directional change in composition characterized by moderate white spruce mortality (46% reduction) and increased cover of late-successional mountain hemlock (Tsuga mertensiana). Forests of the Gulf Coast and northern Kenai Lowland had lower levels of spruce mortality (22% reduction of Sitka spruce (Picea sitchensis) and 28% reduction of white spruce, respectively) and did not show consistent directional changes in vegetation composition. Bluejoint increased by ≥10% in cover on 12 of 33 vegetation plots on the southern Kenai Lowland but did not increase by these amounts on the 82 plots sampled elsewhere on the Kenai Peninsula. Across the Kenai Lowland, however, regeneration of white spruce and paper birch did not change in response to the outbreak or related increases in bluejoint cover from 1987 to 2000. Although some infested areas will be slow to reforest owing to few trees and no seedlings, we found no evidence of widespread reductions in regeneration following the massive spruce beetle infestation.     

Dynamics of soil C,N and Ca in four Swedish forests after removal of tops,branches and stumps as predicted by the Q model

Abstract

We used the Q model to examine the dynamics of carbon (C), nitrogen (N) and calcium (Ca) in the litter/soil system in different scenarios of harvesting intensities, S (stems only), SSl (stems and slash, i.e. tops, and branches including needles) and SSlSt (stems, slash and stumps including coarse roots). Empirical data from long-term field experiments in Sweden, two sites with Norway spruce and two with Scots pine with different levels of productivity, were used to calibrate the model against the stem-only treatment. The highest initial reduction in soil C, N and Ca stores was predicted for SSlSt, and the reduction was more pronounced at low productive sites than at the high productive ones. Most of the decline in soil C and Ca stocks was offset by the litter production in the following forest stand. N showed an initial phase of immobilisation in stumps and coarse roots, while N was immediately released from tops and branches, which contained N-rich needles. Removal of stumps and coarse roots in combination with slash resulted in a similar load of inorganic soil N as for the S treatment, whereas the SSl treatment with stumps left in the soil initially reduced the inorganic soil N pool.