Factors controlling spatial distribution of soil acidification and Al forms in forest soils

Soil acidification and Al release in forest soils is controlled by a number of factors, like acid deposition, forest type, parent rock, altitude, etc. This paper studies the principal stand factors affecting spatial distribution of the content of KCl-extractable Al (Al(KCl), mainly exchangeable), Na4P2O7-extractable Al (Al(Na4P2O7), mainly organically bound), and other soil characteristics related to acidification in surface organic (O) and subsurface mineral (B) horizons in the Jizera Mountains region. Geostatistical methods were exploited. The highest Al(KCl) contents in the O horizons were related to high S and N content, low pH and low Ca and Mg content in soil. Liming decreased Al(KCl) contents in the O horizons. Al(Na4P2O7) in the O horizons was more abundant under spruce than under beech; in both horizons it was increased on the immission clear-cut areas populated by grass. Surface horizons are more sensitive to external influence (acid deposition, liming) and their spatial variation is stronger. In the mineral horizons, the effect of pedogenetic processes is more important. The effect of stand factors on Al behaviour is complex and often indirect, mediated for example by organic matter or soil reaction. It is difficult to clearly distinguish the effects of the particular factors.     

Assessment of soil aluminium pools along three mountainous elevation gradients

Anthropogenic soil acidification in mountain forests and consequent Al release still present a significant problem in many regions. The effect of deposition may differ according to stand conditions, including altitude. This contribution is focused on three elevation transects, two in the Jizera Mountains strongly influenced by acid deposition, one in the less affected Novohradske Mountains. Quantification of pools of different Al forms and related soil characteristics (organic carbon, exchangeable hydrogen cations, sorption characteristics, etc.) is evaluated. In the Novohradske Mountains, the pool of both organically bound and water-soluble Al increases with increasing altitudes. In the Jizera Mountains, the distribution is more complicated; it is strongly affected by different forest type (beech vs. spruce), deforestation, and other local differences. Higher amounts of Al are bound in the mineral horizons compared to the surface organic horizons, even in the case of organically bound Al pools. Further differences between different altitudes and between soil horizons in Al distribution were revealed by detailed Al speciation using HPLC/IC method.     

Time change of aluminium toxicity in the acid bulk soil and the rhizosphere in Norway spruce (Picea abies (L.) Karst.) and beech (Fagus sylvatica L.) stands

Context

In acidic forest soils, aluminium can alter tree health due to its potential toxicity. Aluminium phytotoxicity is mainly influenced by its chemical form and its availability.

Methods

As physical-chemical indicators of Al toxicity in soil, Al speciation in soil solutions and in the exchange complex was measured in the rhizosphere and the bulk soil of two tree species (Norway spruce (Picea abies (L.) Karst.) and European Beech (Fagus sylvatica L.) in an acidic soil and in 4 months (November, February, May and August) representing the four seasons in a year.

Results

In the bulk soil, Al toxicity was generally higher under Norway spruce than under beech. Furthermore, temporal changes in Al behaviour were identified under Norway spruce but not under beech. The monomeric Al in the soil solutions and the exchangeable Al in the solid soil increased significantly in February under Norway spruce and were positively correlated with nitrate concentration, suggesting that nitrate influence Al speciation and mobility under Norway spruce. In the rhizosphere, Al toxicity was restricted through Al complexation by organic compounds and by nutrient contents independently from the season. The ecological importance of the rhizosphere in Al detoxification is discussed.

Conclusions

This study suggests that plant specific differences as well as seasonal changes in plant physiology, microbial activity and microclimatology influence aluminum toxicity in acid forest soils.     

Effects of airborne sea salts on soil water acidification and leaching of aluminium in different forest ecosystems in Denmark

Studies on biogeochemical cycling of elements are performed in even aged forest ecosystems of Norway spruce, Sitka spruce and beech at several locations in Denmark. Episodes with high sea salt deposition and unusual low water surplus percolation caused extreme salt concentrations in the soil solutions. The changes were smallest beneath beech stands and most pronounced at the most salt-affected Sitka spruce stands. Contemporary, the pH drops and the Al³⁺ concentration increased above 20 mg L^-1 owing to cation exchange with Na⁺. The Cl^- and Al³⁺ concentrations reached levels reported as toxic for Norway spruce, but not for Sitka spruce and beech. The changes in the soil water chemistry must be considered as important factors for the reduced vitality in Norway spruce ecosystems in Denmark.     

Soil Base Saturation Combines with Beech Bark Disease to Influence Composition and Structure of Sugar Maple-Beech Forests in an Acid Rain-Impacted Region

Sugar maple, an abundant and highly valued tree species in eastern North America, has experienced decline from soil calcium (Ca) depletion by acidic deposition, while beech, which often coexists with sugar maple, has been afflicted with beech bark disease (BBD) over the same period. To investigate how variations in soil base saturation combine with effects of BBD in influencing stand composition and structure, measurements of soils, canopy, subcanopy, and seedlings were taken in 21 watersheds in the Adirondack region of NY (USA), where sugar maple and beech were the predominant canopy species and base saturation of the upper B horizon ranged from 4.4 to 67%. The base saturation value corresponding to the threshold for Al mobilization (16.8%) helped to define the species composition of canopy trees and seedlings. Canopy vigor and diameter at breast height (DBH) were positively correlated (P < 0.05) with base saturation for sugar maple, but unrelated for beech. However, beech occupied lower canopy positions than sugar maple, and as base saturation increased, the average canopy position of beech decreased relative to sugar maple (P < 0.10). In low-base saturation soils, soil-Ca depletion and BBD may have created opportunities for gap-exploiting species such as red maple and black cherry, whereas in high-base saturation soils, sugar maple dominated the canopy. Where soils were beginning to recover from acidic deposition effects, sugar maple DBH and basal area increased progressively from 2000 to 2015, whereas for beech, average DBH did not change and basal area did not increase after 2010.     

Aluminium effects on spruce and beech seedlings

Summary Natural spruce seedlings sampled on acid brown soil of the Belgian Ardennes, either on mineral soil, or on the litter layer, show higher Al content in the latter case, where substrate Al content is lower. Absorbing power of the root for Al organic chelates is probably higher than for Al⁺³ ions.Most soluble Al is chelated, especially in the holorganic and hemiorganic layers, under spruce and beech. In the mineral layers pH is above 3.8, and total Al in soil solutions is always found to be less than 2 eq/ml.     

Long-term changes of ecosystem services at Solling,Germany: Recovery from acidification,but increasing nitrogen saturation?

Time series of values of ingenious parameters indicating ecosystem services from European beech and Norway spruce ecosystems at Solling, Germany, were evaluated with respect to resilient or adaptive behaviour. Studied indicators comprise the use of monitoring data with up to more than 40 years of observation on deposition of potential acidity, sulphate (SO₄²⁻) budgets, exchangeable base cation pools, Bc/Al ratio in soil solution, nitrogen (N) budgets, foliar nutrition as indicated by the foliar Bc/N ratio, and defoliation. Deposition of potential acidity decreased considerably at both ecosystems. SO₄²⁻ budgets reveal retention of sulphur in the soils affecting acid/base budgets. Exchangeable base cation pools decreased at both ecosystems by about 60%. Bc/Al ratio in soil solution in the mineral soil was mostly below critical limits indicating potential toxic stress to tree roots. N retention in the soils decreased from about 40 kg ha⁻¹ yr⁻¹ in the 1970s to currently very low rates of 0–20 kg ha⁻¹ yr⁻¹ indicating increasing N saturation. Foliar Bc/N ratio decreased at the spruce ecosystem indicating possible nutrient imbalances. Defoliation at both Solling ecosystems is on a high level compared to other forests in Germany, but reveals no distinct relation to soil acidification or N saturation. From the selected indicators, SO₄²⁻ and N budgets reveal resilient behaviour, whereas indicators related to the acid/base status tend to adaptive behaviour.     

The effect of stand age on throughfall chemistry in spruce stands in the Potok Dupniañski catchment in the Silesian Beskid Mountains, southern Poland

The chemical composition of throughfall depends on the age of the Norway spruce (Picea abies Karst) stands and season of the year. The pH of throughfall decreased and the amount of hydrogen ion in throughfall deposited to the soil increased with increasing age of spruce stands, especially in the winter season. Concentrations of K+, H+, SO4(2-), Mn2+, and NH4(+) in throughfall were higher than bulk precipitation for the whole year and K+, H+, and Mn2+ concentrations were higher in throughfall in winter and the growing season. This indicates that these ions were washed out or washed from the surface of needles and/or the bark, and that NO3(-), NH4(+), Ca2+, Mg2+, Fe2+, and Zn2+ were absorbed in the canopy. The effect of high nitrogen deposition, above critical loads, and an increase in the amount of sulfur and in the sum of the strong acids (S-SO4(2-) and N-NO3(-)) that reached the soil with throughfall may have implications for the vitality of spruce stands, especially in older age classes. The application of Principal Component Analysis (PCA) has led to identification of five factors responsible for the data structure ("mineral dust", "acidic emissions", "heavy metals-dust particles", "ammonium [NH4(+)]", and "H+"). They explain more than 60% of the total variance system. The strong positive correlation between stand age class and ionic concentrations in throughfall occurs for all year and the winter period for ions within the following categories: "acidic emissions", SO4(2-) + NO3(-); "heavy metals-dust particles", Fe2+ + Mn2+ + Zn2+; "mineral dust", Na+ + K+ + Ca2+ + Mg2+; "NH4(+)"; and "H+". The strength of the relationship decreases in the growing period, probably due to processes occurring in the canopy (adsorption, leaching, etc.).     

Forest soil solutions: Acid/base chemistry and response to calcite treatment

Soil solution chemistry was investigated at a forested watershed draining into Woods Lake. N.Y. as part of the Experimental Watershed Liming Study (EWLS). The objective of this study was to assess the response of soil water to watershed treatment of calcite (CaCO₃). This material was applied in an effort to mitigate the effects of acidic atmospheric deposition. Soil solutions draining Oa and Bs horizons in reference subcatchments were characterized by low pH and acid neutralizing capacity (ANC) due to elevated concentrations of SO ₄ ^2– , NO ₃ ^– and organic anions relative to the sum of base cation (C_B Ca²⁺, Mg²⁺, Na⁺, K⁺) concentrations. Seasonal and spatial variation of pH andANC in soil solutions appeared to belargely controlled by variations in the concentrations of dissolved organic acids which, in turn, were regulated by reactions of Al with soil organic matter. Nitrate was positively correlated and SO²⁺ was negatively correlated with Ca²⁺ and Al concentrations in reference soil solutions, indicating that changes in NO ₃ ^– influences spatial and seasonal variations in Ca²⁺ and Al concentrations. On this basis, NO ₃ ^– appears to be important in soil acidification and the dynamics of drainage water acidity. Comparison of our results with historical data for the site showed declines in concentrations of SO ₄ ^2– , which are consistent with decreases in emissions of SO₄, in the eastern U.S. and atmospheric deposition of SO ₄ ^2– , to the Adirondack region. Mineral soil solutions have shown large increases in concentrations of NO ₃ ^– . Declines in concentrations of C_B and increases in concentrations of Al have occurred over the last ten years, suggesting depletion of soil pools of exchangeable basic cations and increased sensitivity to acidic deposition. Calcite (CaCO₃) treatment of 6.89 Mg/ha resulted in a significant increase of Ca²⁺, ANC and pH in both Oa and Bs horizon soil solutions. Soil water response to CaCO₃ addition was most evident during the first year after treatment, apparently due to macropore transport of particulate and dissolved CaCO₃ However, increases in ANC and pH in the mineral soil waters were not sustained and appeared insufficient to result in substantial improvement in surface water quality over the 43 month study period.     

Plant-soil feedback in spruce (Picea abies) and mixed spruce-beech (Fagus sylvatica) stands as indicated by dendrochemistry

Ten pairs of secondary pure spruce (Picea abies) and adjacent mixed spruce-beech (Fagus sylvatica) stands on comparable sites were selected on two different bedrocks for soil formation (Flysch: nutrient rich and high soil pH; Molasse: poor nutrient supply and acidic) to study how an admixture of beech to spruce stands affects nutrient cycling and consequently soil chemistry. Soil analyses indicated accumulation of Ca under the mixed stands while the top soil under pure spruce was acidified. It was hypothesized that changes of soil chemical properties due to species composition over the last six decades are reflected in the stem wood of spruce. Three healthy dominant spruce trees per plot were selected for coring. Cores were crossdated and half-decadal samples were analyzed for Ca, Mg, Mn and Al. Calcium and Mg concentrations in stem wood of spruce were significantly higher for the pure spruce than for the mixed stands in spite of lower Ca and Mg stores in the soil. We assume that acidification caused by pure spruce mobilized these cations temporarily, increasing soil solution contents and consequently stem wood concentrations. It was possible to reconstruct soil pH from the element ratios Ca/Al (pure stands) and Ca/Mg (mixed stands), since these ratios in the stem wood of the last half-decade did correlate with soil pH for selected soil depths. Reconstructed soil pH showed a decline over the last 60 years under both species compositions due to accumulation of base cations in the increasing biomass. Comparisons of reconstructed soil pH in 0–5 and 10–20 cm soil depth indicated more pronounced top soil acidification (lower soil pH in 0–5 cm) by spruce on the nutrient rich soil (Flysch) than on the acidic soil (Molasse). However, admixture of beech caused higher pH values in 0–5 cm than in 10–20 cm soil depth on Flysch due to the observed Ca-pump effect of beech (uptake of Ca from deeper soil horizons).     

The soil fauna community in pure and mixed stands of beech and spruce of different age: trophic structure and structuring forces

This study investigates the response of the soil fauna community to replacement of beech by spruce or by mixed stands of beech and spruce. Stands of different age were investigated in a factorial design with the factors tree species (beech and spruce) and stand age (30 and 120 yr). The input of leaf/needle litter did not differ significantly between the study sites. By contrast, the amount of organic matter in upper soil layers (L/F, H/Ah) of spruce forests strongly exceeded that of beech forests particularly in mature stands. The increase in organic matter in spruce stands was not associated by an increase in the amount of microbial biomass. Biomass of eight (bacterivorous, fungivorous and omnivorous nematodes, enchytraeids, earthworms, isopodes, mycetophilid and cecidomyiid Diptera) of the twelve microbi‐detritivorous soil animal groups studied was significantly increased in beech forests; only that of one group (elaterid beetles) was increased in spruce forests and three groups did not respond significantly (collembolans, oribatid mites, sciarid Diptera). This indicates that in the forests studied neither habitat space (amount of organic matter in L/F and H/Ah layers) nor the amount of microbial biomass controlled microbi‐detritivores. Rather, the quality of litter materials and the concentration of microbial biomass therein appeared to be most important. Herbivores and predators also were favoured by beech: the biomass of one (rhizophagous nematodes) of the three herbivorous groups studied were significantly increased in beech stands and none in spruce stands; the biomass of four (predatory nematodes, centipedes, carabid and cantharid beetles) of the seven carnivorous groups studied were increased in beech stands, none in spruce stands. Generally, the biomass ratio between prey and predators was at a minimum in mature beech and mixed stands indicating more intense top‐down control in these forests. Overall, the study documents that replacement of beech by spruce strongly alters the soil food web. Mixed stands were more similar to spruce stands in respect to the biomass of soil animal groups but predator–prey interactions appeared to be more similar in mature beech and mixed stands. Differences between tree species usually were more pronounced in 120 compared to 30 yr old stands indicating that the development of stand characteristics is slow.     

Does soil acidity reduce subsoil rooting in Norway spruce (Picea abies)?

Increasing evidence suggests that forest soils in central and northern Europe as well as in North America have been significantly acidified by acid deposition during the last decades. The present investigation was undertaken to examine the effect of soil acidity on rooting patterns of 40-year-old Norway spruce trees by comparing fine and coarse roots among four stands which differed in soil acidity and Mg (and Ca) nutrition. The coarse root systems of four to five 40-year-old Norway spruce trees per stand were manually excavated. The sum of cross sectional area (CSA) at 60 cm soil depth and below of all vertical coarse roots, as a measure of vertical rooting intensity, was strongly reduced with increasing subsoil acidity of the stands. This pattern was confirmed when 5 additional acidic sites were included in the analysis. Fine root biomass in the mineral soil estimated by repeated soil coring was strongly reduced in the heavily acidified stands, but increased in the humic layer. Using ingrowth cores and a screen technique, we showed that the higher root biomass in the humic layer of the more acidic stands was a result of higher root production. Thus, reduced fine root biomass and coarse root CSA in deeper soil layers coincided with increased root growth in the humic layer. Root mineral analysis showed Ca/Al ratios decreased with decreasing base saturation in the deeper mineral soil (20–40 cm). In the top mineral soil, only minor differences were observed among stands. In general, low Ca/Al ratios coincided with low fine root biomass. Calcium/aluminum ratios determined in cortical cell walls using X-ray microanalysis showed a similar pattern as Ca/Al ratios based on analysis of whole fine roots, although the amplitude of changes among the stands was much greater. Aluminum concentrations and Ca/Al ratios in cortical cell walls were at levels found to inhibit root growth of spruce seedlings in laboratory experiments. The data support the idea that Al (or Ca/Al ratios) and acid deposition-induced Mg (and possibly Ca) deficiency are important factors influencing root growth and distribution in acidic forest soils. Changes in carbon partitioning within the root system may contribute to a reduction in deep root growth.     

Deposition of ammonia to the forest floor under spruce and beech at the Höglwald site

Laboratory and field measurements of the flux of ammonia to forest floor canopies of spruce and beech stands at the Höglwald site in southern Bavaria are reported. Measurements were performed with an open chamber method. A linearity between ammonia concentration and ammonia flux from the atmosphere to the ground floor canopy was detected. Deposition of ammonia showed no saturation even at air concentrations up to 50 g NH₃ m^–3 air. Temperature, water content and the moss layer of the ground floor canopy had a minor influence on the deposition velocity in laboratory experiments. Deposition velocity of ammonia was higher to the spruce (1.3 cm s^–1), and limed spruce ground floor canopy (1.17 cm s^–1) compared to the beech stand (0.79 cm s^–1). In field studies, a diurnal course of the deposition velocity was detected with highest velocities in midday and minor during night times, but not in the climatic chamber. The flux of ammonia to the ground floor canopy was estimated of app. 10 kg N ha^–1 yr^–1 for the soil under spruce, 9 kg N ha^–1 yr^–1 for the limed spruce and 6 kg N ha^–1yr^–1 for the soil under beech. The fluxes are interpreted as fluxes from the atmosphere to the ground canopies of the stands.     

Chemical composition of soil solutions extracted from New Zealand beech forests and West German beech and spruce forests

Concentrations of ions were measured in soil solutions from beech (Nothofagus) forests in remote areas of New Zealand and in solutions from beech (Fagus sylvatica) and Norway spruce (Picea abies) forests in North-East Bavaria, West Germany, to compare the chemistry of soil solutions which are unaffected by acid deposition (New Zealand) with those that are affected (West Germany). In New Zealand, soil solution SO₄ ^2– concentrations ranged between <2 and 58 mol L^–1, and NO₃ ^– concentrations ranged between <1 and 3 mol L^–1. In West Germany, SO₄ ^2– concentrations ranged between 80 and 700 mol L^–1, and NO₃ ^– concentrations at three of six sites ranged between 39 and 3750 mol L^–1, but was not detected at the remaining three sites. At all sites in New Zealand, and at sites where the soil base status was moderately high in West Germany, pH levels increased, and total Al (Al_t) and inorganic monomeric Al (Al_i) levels decreased rapidly with increasing soil depth. In contrast, at sites on soils of low base status in West Germany, pH levels increased only slightly, and Al levels did not decline with increasing soil depth.Under a high-elevation Norway spruce stand showing severe Mg deficiency and dieback symptoms in West Germany, soil solution Mg²⁺ levels ranged between 20 and 60 mol L^–, and were only half those under a healthy stand. Al_t and Al_i levels were substantially higher the healthy stand than under the unhealthy stand, indicating that Al toxicity was not the main cause of spruce decline.     

Long-term changes in aluminum fractions of drainage waters in two forest catchments with contrasting lithology

Aluminum (Al) chemistry was studied in soils and waters of two catchments covered by spruce (Picea abies) monocultures in the Czech Republic that represent geochemical end-members of terrestrial and aquatic sensitivity to acidic deposition. The acid-sensitive Lysina catchment, underlain by granite, was compared to the acid-resistant Pluhův Bor catchment on serpentine. Organically-bound Al was the largest pool of reactive soil Al at both sites. Very high median total Al (Alt) concentrations (40 μmol L⁻¹) and inorganic monomeric Al (Ali) concentrations (27 μmol L⁻¹) were observed in acidic (pH 4.0) stream water at Lysina in the 1990s and these concentrations decreased to 32 μmol L⁻¹ (Alt) and 13 μmol L⁻¹ (Ali) in the 2000s. The potentially toxic Ali fraction decreased in response to long-term decreases in acidic deposition, but Ali remained the largest fraction. However, the organic monomeric (Alo) and particulate (Alp) fractions increased in the 2000s at Lysina. In contrast to Lysina, marked increases of Alt concentrations in circum-neutral waters at Pluhův Bor were observed in the 2000s in comparison with the 1990s. These increases were entirely due to the Alp fraction, which increased more than 3-fold in stream water and up to 8-fold in soil water in the A horizon. Increase of Alp coincided with dissolved organic carbon (DOC) increases. Acidification recovery may have increased the content of colloidal Al though the coagulation of monomeric Al.     

Ecophysiological responses of Fagus sylvatica seedlings to changing light conditions. II. The interaction of light environment and soil fertility on seedling physiology

The survival and growth of natural beech regeneration after canopy removal is variable and little is known about ecophysiological mechanisms of these responses. Biomass, nonstructural carbohydrate levels and nitrogen concentrations were measured in an Italian population of European beech seedlings. Seedlings were container-grown in two types of soil, organic and mineral, collected at the study site. The seedlings were grown under three light treatments: under full beech canopy (understory), exposed to full sun only during midday (gap) and under full sun (clearing). Leaf gas exchange and chlorophyll a fluorescence parameters were measured and then foliar analyses were conducted for chlorophyll, phenolic and tannin levels. Biomass and allocation were significantly affected by light and soil treatments. The clearing seedlings and those in organic soil were larger than seedlings in the other light treatments or soil type. Total nonstructural carbohydrate concentrations were lower in the understory seedlings and significant differences between soil types were present in the gap and clearing seedlings. Nitrogen concentrations were higher in the understory seedlings and those growing in the organic soil compared to the other treatments. Gas exchange rates were highest in clearing and the organic soil seedlings. Gap seedlings exhibited photosynthetic acclimation that allowed them to utilize high light of midday and any sunflecks during the morning and afternoon. Relative fluorescence was significantly influenced by both light treatment and soil type, with the highest values observed in the gap seedlings. Light response curves showed decreasing apparent maximum quantum efficiency from the understory to clearing, while maximum photosynthetic rate was highest in the gap seedlings. Chlorophyll concentration was highest in understory seedlings and those growing in organic soil and higher in seedlings growing in organic than in mineral soil. Both foliar tannin and phenolic levels were highest in clearing seedlings, and only tannin concentrations were affected by soil type. Understory seedlings had the highest mortality and insect herbivory; the latter was found to be inversely related to tannin concentration. Overall, growth and photosynthesis in beech seedlings responded positively to high light associated with small canopy gaps. Organic soil increased seedling size, particularly in the gap and clearing environments. We conclude that forest gaps are favorable for photosynthesis and growth of European beech seedlings.     

Consequences of elevated CO2, augmented nitrogen-deposition and soil type on the soluble nitrogen and sulphur in the phloem of beech (Fagus sylvatica) and spruce (Picea abies) in a competitive situation

Mixed spruce-beech plantations grown in large open-top chambers (OTC) were used to study consequences of elevated CO2, nitrogen-deposition and soil type on plant internal nitrogen and sulphur cycling of juvenile beech (Fagus sylvatica L.) and spruce (Picea abies Karst.) in a competitive situation. Processes of re-cycling as a consequence of protein turnover during leaf senescence in autumn were of further interest. For this purpose, phloem sap was collected in September 1998 and analysed for the composition and concentrations of organic and inorganic nitrogen and sulphur compounds. The phloem exudate of spruce showed higher total soluble non-protein nitrogen (TSNN) concentration on calcareous soil than on acidic soil, independent of the treatment. N-fertilization increased the N-concentration of phloem exudate significantly on both soil types, mainly by an increase of Arg and Gln concentrations. Elevated CO2 slightly increased TSNN on calcareous, but not on acidic soil. The combination of elevated CO2 and augmented N-deposition induced a further increase of TSNN on calcareous soil, but caused a lower N-effect on TSNN on acidic soil. Arg, the main TSNN component in phloem exudate, mediated this effect. Since Arg is considered to be a major nitrogen storage compound, it is concluded that in autumn elevated CO2 and augmented N-deposition, influence storage of N rather than N-supply of spruce. An effect of elevated CO2 and augmented N-deposition on GSH and sulphate concentrations in phloem exudate of spruce was not observed on acidic soil. On calcareous soil augmented N-deposition enhanced, elevated CO2 decreased phloem exudate GSH contents. In combination, elevated CO2 compensated the positive effect of N-deposition. The effects of elevated CO2 and augmented N-deposition on phloem sap N- and S-contents described above were not observed for beech trees. Apparently, elevated CO2 and augmented N-deposition did not affect plants internal S and N cycling of beech grown in spruce-beech plantations.     

Biomass, morphology and nutrient contents of fine roots in four Norway spruce stands

Fine root systems may respond to soil chemical conditions, but contrasting results have been obtained from field studies in non-manipulated forests with distinct soil chemical properties. We investigated biomass, necromass, live/dead ratios, morphology and nutrient concentrations of fine roots (<2 mm) in four mature Norway spruce (Picea abies [L.] Karst.) stands of south-east Germany, encompassing variations in soil chemical properties and climate. All stands were established on acidic soils (pH (CaCl₂) range 2.8–3.8 in the humus layer), two of the four stands had molar ratios in soil solution below 1 and one of the four stands had received a liming treatment 22 years before the study. Soil cores down to 40 cm mineral soil depth were taken in autumn and separated into four fractions: humus layer, 0–10 cm, 10–20 cm and 20–40 cm. We found no indications of negative effects of N availability on fine root properties despite large variations in inorganic N seepage fluxes (4–34 kg N ha⁻¹ yr⁻¹), suggesting that the variation in N deposition between 17 and 26 kg N ha⁻¹ yr⁻¹ does not affect the fine root system of Norway spruce. Fine root biomass was largest in the humus layer and increased with the amount of organic matter stored in the humus layer, indicating that the vertical pattern of fine roots is largely affected by the thickness of this horizon. Only two stands showed significant differences in fine root biomass of the mineral soil which can be explained by differences in soil chemical conditions. The stand with the lowest total biomass had the lowest Ca/Al ratio of 0.1 in seepage, however, Al, Ca, Mg and K concentrations of fine roots were not different among the stands. The Ca/Al ratio in seepage might be a less reliable stress parameter because another stand also had Ca/Al ratios in seepage far below the critical value of 1.0 without any signs of fine root damages. Large differences in the live/dead ratio were positively correlated with the Mn concentration of live fine roots from the mineral soil. This relationship was attributed to faster decay of dead fine roots because Mn is known as an essential element of lignin degrading enzymes. It is questionable if the live/dead ratio can be used as a vitality parameter of fine roots since both longevity of fine roots and decay of root litter may affect this parameter. Morphological properties were different in the humus layer of one stand that was limed in 1983, indicating that a single lime dose of 3–4 Mg ha⁻¹ has a long-lasting effect on fine root architecture of Norway spruce. Almost no differences were found in morphological properties in the mineral soil among the stands, but vertical patterns were apparently different. Two stands with high base saturation in the subsoil showed a vertical decrease in specific root length and specific root tip density whereas the other two stands showed an opposite pattern or no effect. Our results suggest that proliferation of fine roots increased with decreasing base saturation in the subsoil of Norway spruce stands.     

Ecophysiological responses of Japanese forest tree species to ozone, simulated acid rain and soil acidification

In this review, I summarized the results obtained from experimental studies on the ecophysiological responses of Japanese forest tree species to O₃, simulated acid rain and soil acidification. Based on the studies conducted in Japan, exposure to ambient levels of O₃ below 100 nl·l⁻¹ (ppb) for several months is sufficient to inhibit dry matter production and net photosynthesis of sensitive Japanese forest tree species such as Siebold's beech and Japanese zelkova. On the other hand, exposure to simulated acid rain with a pH of 4.0 or above for several months cannot induce any adverse effects on dry matter production and physiological functions of Japanese forest tree species. However, when the pH of simulated rain or fog is lowered below 4.0, negative effects appear on dry matter production and physiological functions such as transpiration in several sensitive Japanese forest tree species such as Japanese fir and Nikko fir. Based on limited information, it may be concluded that (1) Al dissolved into soil solution is the most important limiting factor for dry matter production, physiological functions and nutrient status of Japanese forest tree species grown in acidic soil, (2) the (Ca+Mg+K)/Al molar ratio in soil solution is a useful indicator to evaluate and predict the effects of soil acidification due to acid deposition on whole-plant dry matter production of Japanese forest tree species at the present time and in the future, and (3) Japanese coniferous tree species such as Japanese cedar and red pine are relatively sensitive to a reduction in (Ca+Mg+K)/Al molar ratio in soil solution compared with European forest tree species such as Norway spruce.     

Effects of changes in tree species composition on water flow dynamics – Model applications and their limitations

Water-plant relations play a key role in the water cycling in terrestrial ecosystems. Consequently, changes in tree species composition may have distinct effects on the water retention capacity as well as on the pattern of streamflow generation. Such changes may result from modified interception properties and transpiration related to differences in canopy properties and root distribution. In order to evaluate the potential hydrological effects of the current silvicultural conversion from monocultural conifer stands into mixed or pure deciduous stands the hydrological model BROOK90 was applied to two forested upland catchments in Germany. The Rotherdbach catchment (9.4 ha, 93 yr-old Norway spruce) is situated in the Eastern Ore Mountains. The Schluchsee catchment (11 ha, 55-yr-old Norway spruce) is located in the higher altitudes of the Black Forest. The calibrated model is capable to describe rather well the temporal variation of streamflow but also the portions of the individual flow components. Data for a beech scenario were adapted for each site using a standard parameter set for deciduous trees provided by BROOK90. The annual discharge in the fictional beech stand at Rotherdbach is 30 to 50% higher compared to spruce with an increase of soil moisture and especially the slow streamflow components. This mainly results from low interception rates during winter time. In contrast, the spruce stand has a permanently higher interception rate. Effects of tree species conversion are moderate at Schluchsee. The annual discharge of a fictional beech stand at Schluchsee is 7 to 14% higher compared to spruce. There in contrast to Rotherdbach, effects of tree species conversion on soil moisture dynamics are small since vertical percolation in the highly permeable soil dominates and precipitation is abundant. Practical forestry will favorably establish mixed beech–spruce rather than pure beech stands. However, it is critical to simulate mixed stands with BROOK90. Therefore, a simple summation of model results from spruce and beech according to their respective area in a fictional mixed stand can only be a first approximation. Advanced hydrological simulation of mixed stand conditions should regard interactions of tree species and spatial parameter distribution. However, this is not yet feasible due to a distinct lack of information. As a consequence, there is a strong need to collect relevant hydrological and ecophysiological data in mixed stands in the future.