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.     

Effects of forest liming on soil processes

On the basis of a field experiment in Norway spruce with acid irrigation and compensatory liming of the soil surface (Höglwald, S-Bavaria), liming effects are described as lime dissolution rate, transformation of carbonate buffer to exchange buffer, time required for deacidification of soil and drainage water, mobilization of Cu and Pb, changes in soil organisms, humus decomposition, and nitrogen turnover. It was shown that lime dissolution followed an exponentially decreasing curve. 4 t ha^-1 dolomitic lime were dissolved within 6 years. Additional acid irrigation of 4 kmol H⁺ ha^-1 yr^-1 as sulphuric acid speeded up the lime dissolution to about 4 years. After dissolution of lime about 70% of Ca and about 30% of Mg, both originating from lime dissolution, are retained in the surface humus layer, loading the exchange buffer capacity there. Liming acted as a protection against acid irrigation but the extension of soil deacidification downwards proceeded slowly due to the high base neutralizing capacity of protonated functional groups of the organic matter. The main depth effect is caused by Mg translocation. A significant increase of organic Cu complexes occurred due to mobilization of water soluble humus decomposition products. The effect of liming on litter decomposing organisms is demonstrated with microorganisms, collembolae and earthworms regarding the abundance and the structure of dominance. It was shown that liming may induce unusually large changes in biocenoses of forest soils. The decay of surface humus accounted for 7.2 t ha^-1 or 23% of the store within 7 years. Within the same time span, liming caused a loss of about 170 kg N ha^-1 or 14% of the store of the surface humus layer. The nitrate concentration in the drainage water thus increased by about 50 to 60 mg NO₃ ^- L^-1. Site-specific conditions are discussed, which produce such negative liming effects as increased nitrate concentration of seepage, humus decay and heavy metal mobilization. Redistribution of tree roots, induction of boron deficiency and root rot are also considered. It is indicated that liming may aggravate the increasing problem of nitrate contamination of forest ground water resources which is associated with deposition of atmogenous nitrogen compounds. Some recommendations are given regarding forest practice.     

Nutrient pools and fluxes of the ground vegetation in coniferous forests due to fertilizing,liming and amelioration

The effects of fertilization and amelioration treatments on some nutrient pools and fluxes of ground vegetation in mature pine and spruce stands on acid soils in South Germany are described. In N-limited pine forests with moderate canopy density and with Deschampsia flexuosa an additional N-accumulation in biomass of 20–40 kg ha^-1 occurred 3 years after pure N-fertilization. The N, P, K-cycling through ground vegetation was stimulated more than 10 years by a combined N + CaCO₃ + P treatment leading toa shift in dominance from cryptogams and Ericaceae towards Deschampsia flexuosa and ruderal species like Epilobium angustifolium. The effect of a lupine treatment (combined with initial soil preparation, liming and P supply) was far stronger than the effect of the other experimental procedures. But the fertilizer and amelioration effects on the herb layer of pine forests tended to decline after two decades for different reasons.The shade-tolerant ground vegetation in a nitrogen-saturated spruce forest was not able to prevent heavy additional nitrate losses from upper mineral soil after dolomitic liming. But the Ca, Mg and K fluxes through ground vegetation were strongly elevated in the third year after treatment.     

Effects of liming and boron fertilization on boron uptake of Picea abies

The effects of liming on concentrations of boron and other elements in Norway spruce [Picea abies (L) Karst.] needles and in the mor humus layer were studied in long-term field experiments with and without B fertilizer on podzolic soils in Finland. Liming (2000+4000 kg ha^-1 last applied 12 years before sampling) decreased needle B concentrations in the four youngest needle age classes from 6–10 mg kg^-1 to 5 mg kg^-1. In boron fertilized plots the corresponding concentrations were 23–35 mg kg^-1 in control plots and 21–29 mg kg^-1 in limed plots. Both liming and B fertilizer decreased the Mn concentrations of needles. In the humus layer, total B concentration was increased by both lime and B fertilizer, and Ca and Mg concentrations and pH were still considerably higher in the limed plots than controls. Liming decreased the organic matter concentration in humus layer, whilst B fertilizer increased it.The results about B uptake were confirmed in a pot experiment, in which additionally the roles of increased soil pH and increased soil Ca concentration were separated by means of comparing the effects of CaCO₃ and CaSO₄. Two-year-old bare-rooted Norway spruce seedlings were grown in mor humus during the extension growth of the new shoot. The two doses of lime increased the pH of soil from 4.1 to 5.6 to 6.1, and correspondingly decreased the B concentrations in new needles from 22 to 12 to 9 mg kg^-1. However, CaSO₄ did not affect the pH of the soil or needle B concentrations. Hence the liming effect on boron availability in these soils appeared to be caused by the increased pH rather than increased calcium concentration.     

Effects of Mg(OH)2-fertilization on nutrient cycling in a heavily damaged Norway spruce ecosystem (NE Bavaria/FRG)

Main objective of this study was to test the effects of Mg(OH)₂-fertilization in a Norway spruce ecosystem showing severe symptoms of Mg-deficiency.The site is characterized by high atmospheric inputs with deposition rates of 1.25 kg H, 42 kg S, and 32 kg N per ha and year. The typic Dystrochrept derived from granite is acidified down to greater depths. The pH-values in soil solution of the organic surface layer and the upper mineral soil are around 3.5. Concentrations of Al, SO₄ ^2-, and especially NO₃ ^- and DOC are very high. The element balance indicates a significant influence of N-inputs and processes of N-turnover on the chemical status of the soil and probably on tree nutrition. Nitrification in the upper mineral soil leads to a transformation of a major part of NH₄ ⁺ into NO₃ ^-, which is quantitatively leached, resulting in an ecosystem-internal H⁺-production of 1.8 keq ha^-1yr^-1. NO₃ ^- and SO₄ ^2- govern the seepage output from the ecosystem.Mg(OH)₂ fertilization resulted in manifold increased Mg²⁺ concentrations in soil solution down to 70 cm soil depth and to a significant increase of pH down to 25 cm mineral soil depth. Nitrate concentrations were elevated after fertilization, but decreased within 15 months below the level of the control plot. As a mean over the whole experimental period, N-output was not increased by fertilization. Despite an elevated internal proton production due to nitrification, acid buffering in the soil was clearly increased, but enhanced Al-mobilization was not observed. Mg/Al- and Ca/H-ratios in soil solution indicate much more favourable conditions for fine root growth. Fertilization also increased the amount of exchangeable Mg down to 40cm mineral soil depth. Mg contents in current-year needles increased after three vegetation periods. Thirty months after application, only 10% and 4% of the fertilized Mg had left the organic surface layer and the mineral soil with seepage water output, respectively.     

Effect of artificial soil acidification and liming on growth and nutrient status of mycorrhizal roots of Norway spruce (Picea abies [L.] Karst.)

Effects of soil acidification and liming on biomass responses and free Al, Ca, K, Mg, Mn and P contents of mycorrhizal roots of mature Norway spruce (Picea abies [L.] Karst.) were studied at Höglwald Forest in Southern Germany.At the untreated site, mycorrhizal root biomass was lower in the acid humus (pH = 3.3) than in the less acid upper (0–5 cm) mineral soil (pH 4.1). Mycorrhizal roots from the humus contained 10% of the level free Al in mycorrhizal roots from the upper mineral soil. During seven years of soil acidification the quantity of mycorrhizal roots remained unaffected in the humus and the upper mineral soil, perhaps due to the high buffering capacity of the humus which prevented a significant alteration of the nutrient status of the roots. However, two years after soil acidification had been terminated, the percentage of mycorrhizal roots in the humus decreased, possibly because the free root concentrations of K had decreased.On the other hand, six years after liming, there was a two-fold increase of the annual mean quantity of mycorrhizal roots in the humus. Compensatory liming (acid irrigation after liming) had a similar effect on mycorrhizal root production in the humus. However, two years after acid irrigation had been terminated a decrease of mycorrhizal roots in the upper mineral soil (0–5 cm) was observed. Since the total amount of mycorrhizal roots in the humus and upper mineral soil remained constant, compensatory liming produced a shift in fine roots to the humus layer.The higher mass of living mycorrhizal roots in the upper mineral soil (0–5 cm) as compared to the humus of the untreated plot as well as the increased mass of mycorrhizal roots in the humus after liming or compensatory liming are both attributed to an increase in pH to 4.5 rather than alleviation of Al toxicity.     

Effect of N,K, and P fertilization,N source,and clipping on potential tetany hazard of bromegrass

Summary The objective of this field study was to determine early-season effects of N source, N, K, and P fertilization, and clipping (to simulate grazing) on potential tetany hazard of bromegrass (Bromus inermis L.) as indicated by the chemical composition of its forage. Tetany is a metabolic disorder of ruminants resulting from forage with low Mg availability. Chemical components considered in the forage were inorganic cations, organic acids, aconitate, and per cent total N/per cent total water soluble carbohydrate (N/TWSC). Differences between the sum (in meq/kg) of inorganic cations (Mg, Ca, K, and Na) and inorganic anions (Cl, NO₃, H₂PO₄, and SO₄) in forage were defined as the concentration of organic acids (C-A). Soil was Parshall fsl, a Pachic Haploboroll. Yields and chemical composition of oven-dried forage from previously unclipped and reclipped plots were determined at 3-week intervals beginning May 22 and June 12, respectively. A water budget was determined using soil-water and rainfall data.Forage yields were increased 2- to 3-fold by N fertilization with the NO₃-N source generally outyielding the NH₄-N source. A slight additional yield response to that obtained with N alone was obtained with K+P fertilization but not with K or P alone with or without N. Much less total forage was removed from reclipped plots than from unclipped plots. Forage Mg content was decreased only slightly by K or NH₄-N fertilization. Soil analysis indicated that high NH₄-N levels were present at the May 22 harvest. Magnesium and Ca concentrations were only slightly affected by N fertilization; however, K, K/(Ca+Mg), total N, C-A, and aconitate were increased. Reclipping increased Mg, N, K, N/TWSC, C-A, and aconitate. Estimates of blood-plasma Mg concentrations were obtained by using the data for plant N, K, and Mg. These estimates did not indicate increased tetany hazard as a result of reclipping, but did indicate increased tetany hazard from N fertilization. Forage C-A and aconitate concentrations were decreased by fertilization with KCl which seemed to have been caused by the increased Cl concentrations in the forage. Estimates of quantities of Mg, arriving at the root surfaces from the soil by mass flow, far exceeded amounts of Mg in the forage. Mass flow seemed to be the principal mechanism by which Mg and Ca arrived at root surfaces but this mechanism was much less important for K.This study indicated an increased potential tetany hazard resulting primarily from N fertilization with either NH₄-N or NO₃-N sources. However, the potential for increased forage and livestock-carrying capacity with N fertilization is very large. Therefore, management practices corroborated by livestock data are vitally needed to minimize tetany hazard while increasing bromegrass yields by N fertilization.Contribution from Soil, Water, and Air Sciences, North Central and Northeastern Regions, ARS-USDA.Follett, Power, and Grunes are soil scientists and Kleinis a biological laboratory technician. Follett is now National Program Staff Scientist, ARS, BARC-West, Beltsville, MD 20705. Power and Klein are at the USDA Northern Great Plains Research Center, Mandan, ND 58554, as formerly was Follett. Grunes is at the U.S. Plant, Soil and Nutrition Laboratory, Ithaca, NY 14853.     

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

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

Soil conditions and regeneration after clear felling of a Pinus sylvestris L. stand in a nitrogen experiment,Central Sweden

Forest N fertilization is a common practice in areas of Sweden that are not affected by high levels of N deposition. The environmental consequences of high N input to closed forests are fairly well known, but the long-term effects following clear-felling are a lot less well known. Thus, residual effects on soil and planted seedlings of previous N additions at an experimental N gradient 11 years after clear-felling were studied at a naturally nutrient-poor forest site in central Sweden. The experimental N gradient had been established by three repeated applications (in 1967, 1974 and 1981) of six dosages of NH₄NO₃ with increments of 120 kg N ha^–1. Thus, in total, the applied N dose ranged between 0 and 1800 kg N ha^–1. The study examined extractable base cations and P, soil pH, total-N, total-C, net N-mineralization and potential nitrification in four soil horizons (the humus layer, and 0–5, 5–10 and 10–20 cm in the mineral soil). We also measured the survival and growth of planted Pinus sylvestris L. seedlings. The applied N had no effect on the amounts of extractable-P or base cations in the soil. The soil pH decreased with increasing N dose in the deeper soil horizons, while in the humus the pH showed a weak but statistically significant increase due to the N application. Both total-C and total-N increased as a result of the N application, while the C/N ratio decreased. In the humus layer and the uppermost mineral soil layer NH₄ ⁺ was the major inorganic N source, in contrast to the lowest mineral soil horizon where NO₃ ^– dominated. For most of the studied horizons, there was a positive linear relationship between applied N dose and amount of inorganic N. Both net N-mineralization and potential nitrification showed increases with increasing N dose. As for the plants, no difference in survival or growth was found between the different N treatments. For doses generally applied in forest fertilization no significant differences in any of the studied properties were found.     

Tree species (Picea abies and Fagus sylvatica) effects on soil water acidification and aluminium chemistry at sites subjected to long-term acidification in the Ore Mts., Czech Republic

The effect of European beech (Fagus sylvatica) and Norway spruce (Picea abies) on acid deposition and soil water chemistry was studied at a site in the Ore Mts., Czech Republic, that has been subjected to decades of elevated acidic deposition. Dry deposition onto the spruce canopy significantly increased acid input to the soil in comparison to the beech canopy. As a result soil waters were more acidic; Al, SO4(2-), and NO3- concentrations were significantly higher; and Ca and K concentrations were lower in the spruce stand than in the beech stand. The concentrations of potentially toxic inorganic aluminium (Al(in)) were, on average, three times higher in the spruce stand than in the beech stand. Thus, Al played a major role in neutralizing acid inputs to mineral soils in the spruce stand. Despite the higher dissolved organic carbon (DOC) concentrations in spruce organic soil solutions, organic Al (Al(org)) accounted for only 30% of total Al (Al(tot)), whereas in beech organic soil solutions Al(org) was 60% of Al(tot). Soil waters in the beech stand exhibited Al(in) concentrations close to solubility with jurbanite (Al(SO4)OH.5H2O). The more acidic soil waters in the spruce stand were oversaturated with respect to jurbanite. The Bc/Al(in) ratio (Bc = Ca + Mg + K) in O horizon leachate was 4.6 and 70 in spruce and beech stands, respectively. In beech mineral soil solutions, the Bc/Al(in) ratio declined significantly to about 2. In the spruce stand, mineral soil solutions had Bc/Al(in) values below the critical value of 1. The observed Bc/Al(in) value of 0.4 at 30 cm depth in the spruce stand suggests significant stress for spruce rooting systems. A more favourable value of 31 was observed for the same depth in the beech stand. The efficiency of the spruce canopy in capturing acidic aerosols, particulates, and cloud water has resulted in the long-term degradation of underlying soils as a medium for sustainable forest growth.     

Cation concentrations of short roots of Norway spruce as affected by acid irrigation and liming

The longterm effect of acid irrigation and liming (dolomitic limestone) on the mineral element content of roots of Norway spruce (Picea abies [L.] Karst.) was investigated in an 80-year-old Norway spruce stand in South Germany (Bavaria). Soil cores of four soil depths (humic layer, 0–10, 10–20 and 20–30 cm) were taken over 2 years (August 1991 and August 1992) from six plots with different treatments (control, normal irrigation, acid irrigation solely or in combination with liming) and living short roots selected and analysed for calcium (Ca), magnesium (Mg), manganese (Mn) and aluminium (Al).On the acid irrigated plot, the Ca and Mg contents of roots were decreased in 1991, but by 1992, 2 years after the irrigation had been terminated, no difference could be found. The Al content of the roots was not increased by acid irrigation but rose with increasing soil depth, regardless of treatment. Liming increased root contents of Ca and Mg and reduced contents of Mn and Al. This effect was especially distinct in the humic layer and decreased with increasing soil depth. Even though the molar Ca/Al-ratio in the roots in the mineral soil was generally low (0.09–0.52), no evidence of Al toxicity could be found. The formation of Al complexes is discussed as a reason for this behaviour.     

Effects of ammonium sulphate application on the chemistry of bulk soil,rhizosphere, fine roots and fine-root distribution in a Picea abies (L.) karst. stand

The effect of ammonium sulphate application on the bulk and rhizosphere soil chemistry, elemental concentration of living fine roots (<2 mm in diameter), amounts of living and dead fine roots, root length density and specific root length density were investigated in a 28 year old Norway spruce stand in SW Sweden. The treatments started in 1988. Core samples of the LFH layer and mineral soil layers were sampled in control (C) and ammonium sulphate (NS) treatment plots in 1988, 1989 and 1990. Soil pH and NO₃-S and SO₄-S, Al, Ca, Mg, Mn and K concentrations were measured for both the bulk soil and rhizosphere soil.The pH-values of the bulk and rhizosphere soil decreased in 1989 and 1990 in NS plots compared to control plots, while the SO₄-S concentration increased. The Ca, Mg and K concentration increased in the NS treatment in almost all layers in the bulk and the rhizosphere soil. Ammonium ions may have replaced these elements in the soil organic matter. The NS treatment reduced Mg concentration in fine roots in all layers in 1990. The Al concentrations in the rhizosphere and bulk soil were higher in NS plots in all layers, except at 0–10 cm depth, both in 1989 and 1990. The Al content of living fine roots was higher in NS plots than C plots but the differences were not significant. The NS addition did not affect the P and K contents of fine roots in any soil layer, but the S concentrations of fine roots were significantly higher in NS plots in 1989 and 1990. The fine root necromass was higher in NS than in C in 1990, in the LFH layer, indicating a gradual decrease in the vitality of the fine roots. It was suggested that the NS treatment resulted in displacement of Mg and K from exchange sites in the LFH layer leading to leaching of these cations to the mineral soil. Further application of ammonium sulphate may damage the fine roots and consequently adversely affect the water and nutrient uptake of root systems.     

Does forest liming impact the enzymatic profiles of ectomycorrhizal communities through specialized fungal symbionts?

Liming (Ca–Mg soil amendment) is a forestry practice used to correct soil acidification and restore health and productivity in declining stands. Liming is known to modify tree mineral nutrition beyond the sole Ca and Mg. We hypothesized that liming also modifies the very functioning of the tree absorbing system (that is the ectomycorrhizal fine roots) in a way that facilitates the mobilization of mineral nutrients, particularly those entrapped in soil organic matter. This hypothesis has been tested here in beech and Norway spruce stands in North-Eastern France. In autumn, we compared the ectomycorrhizal community structure and the enzymatic profiles of ectomycorrhizal root tips in limed and untreated plots by measuring the activities of eight enzymes related to the degradation of soil organic matter. The results show that the ectomycorrhizal community responds to the Ca–Mg amendment and to the resulting soil modifications by modified enzyme activity profiles and ability to mobilize nutrients from soil organic matter. The effects of liming on the belowground functioning of the tree stands result essentially from specialized ECM fungal species such as Clavulina cristata (with strong glucuronidase activity), Lactarius subdulcis (with strong laccase activity) or Xerocomus pruinatus (with strong leucine aminopeptidase activity).     

Land use intensity,rather than plant species richness,affects the leaching risk of multiple nutrients from permanent grasslands

The intensification of land use constitutes one of the main drivers of global change and alters nutrient fluxes on all spatial scales, causing landscape‐level eutrophication and contamination of natural resources. Changes in soil nutrient concentrations are thus indicative for crucial environmental issues associated with intensive land use. We measured concentrations of NO₃–N, NH₄–N, P, K, Mg, and Ca using 1,326 ion‐exchange resin bags buried in 20 cm depth beneath the main root zone in 150 temperate grasslands. Nutrient concentrations were related to land use intensity, that is, fertilization, mowing, grazing intensities, and plant diversity by structural equation modeling. Furthermore, we assessed the response of soil nutrients to mechanical sward disturbance and subsequent reseeding, a common practice for grassland renewal. Land use intensity, especially fertilization, significantly increased the concentrations of NO₃–N, NH₄–N, K, P, and also Mg. Besides fertilization (and tightly correlated mowing) intensity, grazing strongly increased NO₃–N and K concentrations. Plant species richness decreased P and NO₃–N concentrations in soil when grassland productivity of the actual year was statistically taken into account, but not when long‐term averages of productivity were used. Thus, we assume that, in the actual study year, a distinct drought period might have caused the observed decoupling of productivity from fertilization and soil nutrients. Breaking up the grassland sward drastically increased NO₃–N concentrations (+146%) but reduced NH₄–N, P, and K concentrations, unbalancing soil nutrient stoichiometry and boosting the risk of N leaching. Reseeding the sward after disturbance did not have a short‐term effect on nutrient concentrations. We conclude that renewal of permanent grassland should be avoided as far as possible and future grassland management has to strongly rise the effectiveness of fertilization. Additionally, grassland management might have to increasingly taking care of periods of drought, in which nutrient additions might not increase plant growth but potentially only facilitate leaching.     

Long term effect of liming and fertilization on ectomycorrhizal colonization and tree growth in old Scots pine (Pinus sylvestris L.) stands

In this study, we surveyed the long term effects of liming and fertilizing in old Scots pine stands on the ectomycorrhiza (ECM) colonization, tree growth and needle nutrient concentration 35 years later. Four mature stands of Scots pine on low productive mineral soil were limed in 1959 and 1964 with total doses of limestone ranging from 3 to 15 Mg ha^?1 and fertilized with nitrogen (N) in 1970. Thirty-five years after the first liming treatment, all stands were analysed for tree growth and needle nutrient concentrations and two of the stands were also analysed for ECM colonization. ECM colonization increased significantly with liming from 61.5% in the control plots to 88% in the plot with the highest limestone dose. ECM colonization increased with increasing pH in the humus layer from 62% colonization at pH?=?3.5 to 90% at pH?=?6.5 and decreased with increasing amount of extractable phosphorus (P) in the humus. Liming did not affect the frequencies of different ECM morphotypes or dead short root tips, the fine root biomass or necromass. ECM colonization was uncorrelated with needle nutrient concentrations or tree increment. Liming did not significantly affect tree growth. However, nutrient concentrations of current-year needles were affected by prior liming. Ca concentrations in current-year needles increased from approximately 15 mg g^?1 in control treatments to more than 30 mg g^?1 in limed plots, whereas concentrations of Mn, Al, Fe, and in two stands, B, decreased due to liming. In conclusion, liming with doses up to 15 Mg ha^?1 was detectable in stands 35 years after treatment. The liming significantly increased the ECM colonization of Scots pine fine roots, increased the needle nutrient concentration of Ca and decreased the needle concentrations of Mn, Al, and Fe.     

Influence of soil acidification and liming on selected enzymes of the carbohydrate metabolism and the contents of two major organic acids of mycorrhizal roots of Norway spruce (Picea abies [L.] Karst.)

Effects of soil acidification and liming on the activities of three enzymes of the carbohydrate metabolism and the quantities of two of the major organic acids of mycorrhizal roots of Norway spruce (Picea abies [L.) Karst.) were studied at Höglwald Forest in southern Germany. The enzymes investigated were glucosephosphate isomerase, pyruvate kinase and 6-phosphogluconate dehydrogenase. The organic acids studied were citric acid and malic acid.Annual mean activities of the three enzymes were equal in mycorrhizal roots of the humus and the upper mineral soil. But in autumn and winter the activities of each of the three enzymes were higher than in summer. Of the various soil treatments only soil acidification affected the activities of the three enzymes. It stimulated activities by a factor of 1.5 in mycorrhizal roots of the humus but had no effect on mycorrhizal roots from the upper mineral soil.Mycorrhizal roots in the humus contained approximately 10 times more citrate and two times more malate than mycorrhizal roots from the upper mineral soil (0–5 cm). In mycorrhizal roots from the humus citrate and malate were of similar concentrations. In mycorrhizal roots from the upper mineral soil malate was approximately four times more concentrated than citrate. In the humus the citric acid concentration of mycorrhizal roots decreased under soil acidification by a factor of 1.4 while it increased under liming and compensatory liming (acid irrigation after liming) by a factor of 1.5. Malic acid concentrations increased exclusively under liming in mycorrhizal roots of the humus by a factor of 1.3.     

Effects of ammonium sulphate application on the rhizosphere,fine-root and needle chemistry in aPicea abies (L.) Karst. stand

Rhizosphere, fine-root and needle chemistry were investigated in a 28 year old Norway spruce stand in SW Sweden. The uptake and allocation pattern of plant nutrients and aluminium in control plots (C) and plots repeatedly treated with ammonium sulphate (NS) were compared. Treatments started in 1988. Current year needles, one-year-old needles and cylindrical core samples of the LFH-layer and the mineral soil layers were sampled in 1988, 1989 and 1990. Compared to the control plots, pH decreased significantly in the rhizosphere soil in the NS plots in 1989 and 1990 while the SO₄-S concentration increased significantly. Aluminium concentration in the rhizosphere soil was generally higher in the NS plots in all soil layers, except at 0–10 cm depths, both in 1989 and 1990. Calcium, Mg and K concentrations also increased after treatment with ammonium sulphate. Ammonium ions may have replaced these elements in the soil organic matter. The NS treatment significantly reduced Mg concentrations in fine roots in all layers in 1990. A similar trend was found in the needles. Ca concentrations in fine roots were significantly lower in the NS plots in the LFH layer in 1990 and the same pattern was found in the current needles. The N and S concentrations of both fine roots and needles were significantly higher in the NS plots. It was suggested that NS treatment resulted in displacement of Mg, Ca and K from exchange sites in the LFH layer leading to leaching of these cations to the mineral soil. Further application of ammonium sulphate may damage the fine roots and consequently adversely affect the water and nutrient uptake of root systems.     

Effects of understory removal, N fertilization, and litter layer removal on soil N cycling in a 13-year-old white spruce plantation infested with Canada bluejoint grass

Canada bluejoint grass [Calamagrostis canadensis (Michx.) Beauv., referred to as bluejoint below] is a competitive understory species widely distributed in the boreal region in North America and builds up a thick litter layer that alters the soil surface microclimate in heavily infested sites. This study examined the effects of understory removal, N fertilization, and litter layer removal on litter decomposition, soil microbial biomass N (MBN), and net N mineralization and nitrification rates in LFH (the sum of organic horizons of litter, partially decomposed litter and humus on the soil surface) and mineral soil (0–10 cm) in a 13-year-old white spruce [Picea glauca (Moench.) Voss] plantation infested with bluejoint in Alberta, Canada. Removal of the understory vegetation and the litter layer together significantly increased soil temperature at 10 cm below the mineral soil surface by 1.7 and 1.3°C in summer 2003 and 2004, respectively, resulting in increased net N mineralization (by 1.09 and 0.14 mg N kg⁻¹ day⁻¹ in LFH and mineral soil, respectively, in 2004) and net nitrification rates (by 0.10 and 0.20 mg N kg⁻¹ day⁻¹ in LFH and mineral soil, respectively, in 2004). When the understory vegetation was intact, nitrification might have been limited by NH₄ ⁺ availability due to competition for N from bluejoint and other understory species. Litter layer removal increased litter decomposition rate (percentage mass loss per month) from 2.6 to 3.0% after 15 months of incubation. Nitrogen fertilization did not show consistent effects on soil MBN, but increased net N mineralization and nitrification rates as well as available N concentrations in the soil. Clearly, understory removal combined with N fertilization was most effective in increasing rates of litter decomposition, net N mineralization and nitrification, and soil N availability. The management of understory vegetation dominated by bluejoint in the boreal region should consider the strong effects of understory competition and the accumulated litter layer on soil N cycling and the implications for forest management.     

Effects of liming on soil chemical characteristics and grass growth in laboratory and long-term field-amended soils

Summary The influence of liming on soil solution composition was compared in two laboratory amended soils and one field amended soil. In the laboratory study, soil solutions were sampled by miscible displacement at intervals of 1 and 10 weeks after liming. In addition to increases in pH and Ca, there were large reductions in the concentrations of Mg, K, Na, Si and Mn. Solution concentration of free Al decreased with liming; however, organically complexed Al increased, as did soluble organic matter. Liming also stimulated mineralization of N as indicated by increased solution NO₃ levels. The field amended soils were obtained from a long-term cutting trial investigating the effects of lime on pasture. Despite the passage of a 16-year interval since application, the effects of lime on soil solution characteristics were still clearly evident and generally consistent with those observed in the laboratory study. Estimated leaching losses of Ca from limed soil were relatively low, amounting to 12%, 27% and 44% of the 4.2, 8.4 and 12.5 t lime ha⁻¹ applied, respectively. The results suggest that, in Eastern Ireland, a lime treatment would maintain and elevated pH and would influence the avialability and mobility of plant nutrients for some decades following application.     

Effects of inorganic nitrogen application on the dynamics of the soil solution composition in the root zone of maize

The effect of inorganic nitrogen (N) fertilizer on the ionic composition of the soil solution under maize (Zea mays L.) was studied. A pot experiment was carried out with two treatments combined factorially, with or without N application (Ca(NO₃)₂; +N and –N treatments, respectively), and with or without plants. Three looped hollow fiber samplers were installed in each pot to sample soil solutions nondestructively from the root zone, seven times during the 50-day growth period. Plants were harvested on the 50th day, and their nutrient contents determined.Effects of N fertilizer on the soil solutions were observed by the first sampling, 2 days after sowing. The concentrations of Ca and NO₃ ^– and electrical conductivity (EC) increased significantly in the +N treatments as direct effects of fertilizer application. In addition, the concentrations of Mg, K, Na and H⁺ also increased and that of P decreased significantly as indirect effects caused by the re-establishment of chemical equilibria. This suggested the greater supply as well as the greater possibility of leaching loss not only of NO₃ ^– but also of Ca, Mg and K. In the treatments with plants, the concentrations of NO₃ ^–, Ca, Mg and K decreased with time and pH increased significantly compared with the unplanted soil. The depletion of N in the soil solution roughly agreed with the amount of N taken up by the plant. The depletions of K from the soil solution amounted to less than 10% of the amount of the K taken up, suggesting intensive replenishment of K from exchange sites in the soil. Depletions of Ca and Mg were several times higher than the amounts taken up, indicating that the depletions resulted from the adsorption of the divalent cations by the soil rather than uptake by plants. Because NO₃ ^– is hardly absorbed by exchange sites in soil and was the dominant anion in solution, it was concluded that NO₃ ^– had a major role in controlling cation concentrations in the soil solution and, consequently, on their availability for uptake by plants as well as their possible leaching loss. ei]H Marschner