Short rotation forestry – Earthworm interactions: A field based mesocosm experiment

Short rotation forestry (SRF) which consists of planting rapidly growing native and non-native tree species has been introduced to the UK to increase woody biomass production. A largely unknown aspect of SRF species is their interaction with soil fauna, of which the earthworm community is a major component. Earthworms have a pronounced impact on litter decomposition, nutrient cycling and tree growth. Conversely, tree litter and root chemistry can impact on the associated earthworm community development. The aim of this study was to determine direct interactions between SRF species and earthworms. A field-based mesocosm experiment was conducted using Betula pendula (birch) and Eucalyptus nitens (eucalyptus) with two earthworm species Lumbricus terrestris and Allolobophora chlorotica. The one year experiment revealed that native birch and non-native eucalyptus had a similar influence on L. terrestris population development. However, birch had a positive impact on A. chlorotica population establishment compared with eucalyptus. In the presence of earthworms, total tree biomass and leaf nitrogen concentration of eucalyptus were increased respectively by 25% and 27% compared with an earthworm-free control. In the presence of earthworms, surface litter incorporation was greater for both tree species (almost 5 times for birch and 3 times for eucalyptus) compared with controls. This work showed direct SRF-earthworm interactions which differed for tree species.     

Burrowing activity of the earthworms Lumbricus terrestris and Aporrectodea giardi and consequences on C transfers in soil

Two earthworms species, Lumbricus terrestris (epianecic) and Aporrectodea giardi (anecic) were incubated in microcosms with an epigeic ¹³C-labelled litter for 246 d. At the end of the experiment, different soil compartments (surface casts, walls and peripheries of burrows, and surrounding soil) were sampled for ¹³C analysis. Two-dimensional images acquired using X-ray computed tomography allowed to estimate the weight of the ‘burrow wall’ and ‘burrow periphery’ compartments which are required to establish C balance. In the case of L. terrestris, the formed structures were more C litter enriched compared to the other species. The permanent character of the burrow system could lead to a high and constant enrichment of the entire burrow system. As consequence, the percentage of C litter in the ‘burrow wall’ and ‘burrow periphery’ compartments was important in spite of their low volume. The denser system developed by A. giardi resulted in C litter dilution in the whole formed structures. The C litter enrichment decreased with the soil depth, but owing to the intensity of the burrowing activity, the C litter transfers into the ‘burrow walls’ and ‘burrow periphery’ were important and the C litter was homogeneously distributed throughout the whole column.     

The interactions between soil type and earthworm species determine the properties of earthworm casts

Earthworms are recognized to increase soil porosity, reorganize soil structure, and stimulate soil microflora and nutrient mineralization. The properties of earthworm casts should depend both on earthworm species or ecological group and on soil properties. Interactions between earthworm species and soil types have been suggested, but only poorly demonstrated. In order to better understand those interactions, two hypotheses led our study: (1) Soil type has a greater influence on cast properties than earthworm; (2) Earthworms from different species influence cast properties differently; (3) The intensity and direction of the impact of each earthworm species on cast properties vary with soil properties. Fifteen physical and chemical variables (N–NH₄⁺, N–NO₃⁻, total organic C and N, C/N ratio, CaCO₃, pH, P, K⁺, Mg²⁺, Mn²⁺, Na⁺, CEC, moisture, wettability) were measured in casts of three earthworm species (Lumbricus terrestris, Allolobophora chlorotica and Aporrectodea rosea) produced in three temperate soils. Univariate and multivariate analyses showed that earthworm species and soil types significantly impacted cast properties. pH, N_t, K and Mg contents were interactively altered by both factors. Multivariate analysis showed that a difference of soil type had a major impact on casts properties (62%) compared to the impact of a difference of earthworm species (10%). Cast properties were most impacted by L. terrestris, then by A. chlorotica and last by A. rosea. The response ratio (ratio of the properties of the casts to the properties of the bulk soil) was used to quantify the effect of earthworm species compared to the control soil. It showed a higher response of variables in casts in nutrient-rich soils, especially in casts of L. terrestris. The interactions between earthworm species and soil types on cast properties were discussed with regards to earthworm ecology, properties of the soil, and earthworm modifications of cast microflora.     

Effects of liming on survival and reproduction of two potentially invasive earthworm species in a northern forest Podzol

During the last several decades, colonization of soil by exotic earthworms and their effects on soil properties and biodiversity have been reported in forests of North America. In some northern hardwood stands, acid soils or harsh climate may have prevented earthworm colonization. However, climatic change and the increasing use of liming to restore the vigor of declining sugar maple (Acer saccharum Marsh.) stands, situated on base-poor soils in USA and Canada, could make many of these sites more suitable for earthworm colonization. We tested survival and reproduction of two exotic earthworm species (Lumbricus terrestris and Amynthas hawayanus) in unlimed and limed soils at the northern limit of the northern hardwood forest distribution in Canada. Improving soil parameters of base-poor, acidic soils by liming positively influenced activity, survivability and reproductive output of L. terrestris in this northern hardwood forest. In contrast, the high mortality and low vigor of L. terrestris observed in the unlimed plots show that soils in this area with a pH of 4.3 are not favorable to this species. Our results suggest that A. hawayanus was very active prior to winter at both soil pHs, but was not able to complete its life cycle during one year at this latitude. Both earthworm species significantly reduced organic C and total N, and increased the C/N ratio of the forest floor. Given that forest liming activities are increasing in proximity to human activities, there is high probability that some earthworm species, such as L. terrestris, will invade limed northern hardwood forests in the next decades, with possible consequences for soil organic matter turnover, nutrient cycling and forest biodiversity and dynamics.     

Assessing daily egestion rates in earthworms: using fungal spores as a natural soil marker to estimate gut transit time

Earthworms have an important role in ‘bioturbation’—the mixing of soil due to biological processes. Quantification of earthworm bioturbation relies on estimating earthworm egestion rates which in turn depend on two parameters: the gut content of the worms and the gut transit time (GTT). Gut content can be determined relatively easily, but determining GTT is problematic. The present study aimed at estimating daily soil egestion rates of Aporrectodea caliginosa and Lumbricus terrestris, refining the most common approach for estimating GTT by using fungal spores as natural markers in ingested soil. This approach avoids the use of artificial markers that may adversely affect the earthworms. Gut transit time was estimated by tracking the passage of marked soil through the gut by the appearance of the spores in the egested faeces. Gut transit time was estimated to be 9.6?±?0.3 h for A. caliginosa and 11.6?±?0.5 h for L. terrestris. Gut content averaged 465?±?40(± standard error (SE))?mg dw g^?1 dw worm for A. caliginosa and 265?±?80 mg dw g^?1 dw worm for L. terrestris. From these values, daily egestion rates of 1.16 and 0.66 g dw faeces g^?1 dw worm d^?1 were calculated for A. caliginosa and L. terrestris, respectively. Both values compare well to literature values for each species. The presented method for GTT estimation is inexpensive, rapid and easy to evaluate, with spores being a good alternative to existing markers.     

Earthworm species composition affects the soil bacterial community and net nitrogen mineralization

Knowledge of the effects of species diversity within taxonomic groups on nutrient cycling is important for understanding the role of soil biota in sustainable agriculture. We hypothesized that earthworm species specifically affect nitrogen mineralization, characteristically for their ecological group classifications, and that earthworm species interactions would affect mineralization through competition and facilitation effects. A mesocosm experiment was conducted to investigate the effect of three earthworm species, representative of different ecological groups (epigeic: Lumbricus rubellus; endogeic: Aporrectodea caliginosa tuberculata; and anecic: Lumbricus terrestris), and their interactions on the bacterial community, and on nitrogen mineralization from ¹⁵N-labelled crop residue and from soil organic matter.Our results indicate that L. rubellus and L. terrestris enhanced mineralization of the applied crop residue whereas A. caliginosa had no effect. On the other hand, L. rubellus and A. caliginosa enhanced mineralization of the soil organic matter, whereas L. terrestris had no effect. The interactions between different earthworm species affected the bacterial community and the net mineralization of soil organic matter. The two-species interactions between L. rubellus and A. caliginosa, and L. rubellus and L. terrestris, resulted in reduced mineral N concentrations derived from soil organic matter, probably through increased immobilization in the bacterial biomass. In contrast, the interaction between A. caliginosa and L. terrestris resulted in increased bacterial growth rate and reduced total soil C. When all three species were combined, the interaction between A. caliginosa and L. terrestris was dominant. We conclude that the effects of earthworms on nitrogen mineralization depend on the ecological traits of the earthworm species present, and can be modified by species interactions. Knowledge of these effects can be made useful in the prevention of nutrient losses and increased soil fertility in agricultural systems, that typically have a low earthworm diversity.     

Invasion of a deciduous forest by earthworms: Changes in soil chemistry, microflora, microarthropods and vegetation

Ecosystems of northern North America existed without earthworm fauna until European settlers arrived and introduced European species. The current extent of invasion by some of these species, Lumbricus terrestris L., Octolasion tyrtaeum Savigny and Dendrobaena octaedra Savigny, into an aspen forest in the Canadian Rocky Mountains and the effects of the invasion on soil chemistry, microflora, soil microarthropods and vegetation were investigated. Densities of earthworm species, soil structure, plant coverage and abundance were determined along three transects starting at the edge of the forest. At locations with L. terrestris, litter was incorporated into the soil, and where O. tyrtaeum was present, organic layers were mixed with mineral soil layers. Organic layers disappeared almost entirely when both species occurred together. Carbon and nitrogen concentrations were reduced in organic layers in the presence of L. terrestris and O. tyrtaeum. Microbial biomass and basal respiration were reduced when L. terrestris and O. tyrtaeum were present, presumably due to resource competition and habitat destruction. Microarthropod densities and the number of microarthropod species were strongly reduced in the presence of O. tyrtaeum (−75% and −22%, respectively), probably through mechanical disturbances, increasing compactness of the soil and resource competition. The coverage of some plant species was correlated with earthworm abundance, but the coverage of others was not. Despite harsh climatic conditions, the invasion of boreal forest ecosystems by mineral soil dwelling earthworm species is proceeding and strongly impacts soil structure, soil chemistry, microorganisms, soil microarthropods and vegetation.     

Organic matter characteristics and C and N transformations in the humus layer under two tree species, Betula pendula and Picea abies

The aim of this study was to compare the effects of silver birch (Betula pendula Roth) and Norway spruce (Picea abies (L.) Karst.) on soil C and N transformations and on the characteristics of organic matter. Soil samples were taken from the humus layer of a replicated 35-year-old birch-spruce field experiment growing on Vaccinium myrtillus site type in middle-eastern Finland. The soil was a podzol and humus type mor. Soil pH was higher under birch (4.7) than under spruce (4.1). The C-to-N ratio was lower under birch (17) than under spruce (23). Per unit organic matter, microbial biomass C and N, net N mineralization and net nitrification were all higher in birch soil than in spruce soil. The rate of C mineralization (CO₂ production) was, however, the same regardless of tree species. Water-extracts were analyzed for the concentrations of dissolved organic C (DOC) and N (DON) and characterized according to molecular size distribution by ultrafiltration and according to chemical composition using a resin fractionation technique. The concentration of DON, in particular, was higher in birch soil than in spruce soil. The distribution of DOC and DON into different fractions based on molecular size or chemical composition was rather similar in both soils. The concentration of total phenolics, expressed as tannic acid equivalents, was higher in the humus layer under birch than in the humus layer under spruce, because the birch humus layer contained significantly more low-molecular weight (about <0.5 kD) phenolics than the spruce humus layer did. The concentration of proanthocyanidins (condensed tannins) was higher in spruce soil than in birch soil. The concentrations of the five most abundant phenolic acids showed that ferulic and p-coumaric acids were more abundant in spruce soil. Birch soil tended to contain slightly more nonvolatile sesquiterpenes than the spruce soil. The concentration of diterpenes was similar in both soils; but birch soil contained significantly more triterpenes, mainly sterols, than spruce soil did.     

Introduced earthworm species exhibited unique patterns of seasonal activity and vertical distribution,and <Emphasis Type="Italic">Lumbricus terrestris</Emphasis> burrows remained usable for at least 7 years in hardwood and pine stands

It is difficult to obtain non-destructive information on the seasonal dynamics of earthworms in northern forest soils. To overcome this, we used a Rhizotron facility to compile 7 years of data on the activity of anecic (Lumbricus terrestris) and endogeic (Aporrectodea caliginosa complex) earthworms in two contrasting soil/plant community types. We hypothesized that L. terrestris burrows would be used for longer than a typical L. terrestris lifetime, and that the distribution and activity pattern of the two earthworm species would respond differently to changes in soil moisture and temperature. For 7 years we recorded earthworm distribution and activity state bi-weekly to a depth of 1.5 m, tracked L. terrestris burrows using images captured annually, and measured soil temperature and moisture. Activity and vertical distribution of earthworms was closely linked to earthworm species and soil temperature in the fall, winter and spring. Lumbricus terrestris typically remained active through the winter, whereas the A. caliginosa complex was more likely to enter an aestivation period. Activity of all earthworms decreased substantially in July and August when soil temperature was at its highest and soil moisture at its lowest for the year. Most L. terrestris burrows were used continuously and moved very little during the 7-year study, likely creating spatiotemporally stable hotspots of soil resources. The different patterns of response of these species to soil temperature and moisture suggests that endogeic earthworms are more likely than anecic earthworms to adjust activity states in response to climate change mediated shifts in soil moisture and temperature.     

Influence of organic matter on earthworm production and behaviour: a laboratory-based approach with applications for soil restoration

Restored soils are often low in organic matter (OM). As a result, OM is applied to increase soil fertility. Earthworms can assist soil incorporation of OM, but feeding behaviours, and as a result their roles in this process, differ between species. Laboratory experiments examined the effects of OM availability on the growth and behaviour of Aporrectodea longa and Lumbricus terrestris. Hatchling growth was recorded from 1 l vessels containing soil and OM (separated cattle solids), either surface applied (SUR) or mixed into the soil (MIX). Growth of L. terrestris was also recorded where separated cattle solids (SCS) were concentrated in bands at the surface (SUR) or at depths of 0.03 (UPP) or 0.08 m (LOW). In a further experiment, L. terrestris and L. rubellus were kept in mono-species cultures and paired combination, in treatments with selected SCS placements, which allowed assessment of OM position on species interactions. After 28 weeks, both L. terrestris and A. longa exhibited significantly greater growth rates (P < 0.05) in SUR treatments. The behaviour of both species changed with time. L. terrestris were located in areas where OM was concentrated. After 24 weeks, mean masses of 3.1, 2.2 and 1.6 g were recorded for SUR, UPP and LOW treatments, respectively. Where L. rubellus and L. terrestris were co-cultured, the former had significant (P < 0.05) negative effects on the growth of the latter, but only when surface OM was limited. At restored sites, application of OM may be important in the development of sustainable earthworm populations and hence their role in soil amelioration.     

Metal Accumulation by Woody Species on Contaminated Sites in the North of France

Metal accumulation was investigated in a range of woody species that were planted on Cd-, Zn- and Pb- polluted sites in North of France. The study is unique in that we directly compare a large number of woody species (25). The highest accumulation of Zn and Cd was found in the Salicaceae family members with up to 950 mg Zn kg^?1 dry weight (DW) and 44 mg Cd kg^?1 DW in leaves of Populus tremula × Populus tremuloides. Zn content was positively correlated with Cd content, both in leaves and stems. Pb concentration was generally low and was species-independent. Oak and birch species accumulated more Mn as compared to other woody species. A seasonal variation in metal accumulation could be found. Although soil compositions and metal bioavailabilities differed amongst the experimental sites chosen in this study, variation of metal concentrations within a given species was small. High bioconcentration factors for poplar and willow suggested the high potential of these species over other woody species for metal accumulation. Taken together, these data suggest that poplar and willow species are good candidates for phytoremediation programmes.     

Soil and vegetation changes in experimental birch plots on a Calluna podzol

A long-term experiment set up by G. W. Dimbleby to test the effect of birch as a soil improver on a heather moor podzol is described. After 30 yr, an acid woodland type of vegetation developed under the birch trees with Deschampsia flexuosa and Vaccinium myrtillus locally dominant. The mean annual dry matter production of stems and branches was approximately 2.9 t ha^?1. The raw humus layer removed as an experimental treatment had re-formed. Its pH, about 3.4, had remained unchanged for 15 yr. The mean extractable Ca content of the upper 3 cm horizon of the birch plots and of the control plots under heather had increased and on some of the birch plots the concentrations of exchangeable Ca were somewhat higher than those under heather. The N content of the Ol + Of, Oh and Ea horizons was higher under the birch than under heather but there was no evidence of movement of Fe out of the ironpan under the influence of birch. The profile under birch remained a fully differentiated podzol.     

Volatile monoterpenes in soil atmosphere under birch and conifers: Effects on soil N transformations

The aim of this study was to examine the occurrence and concentrations of volatile organic compounds (VOCs), in particular, volatile monoterpenes, in soil atmosphere under silver birch (Betula pendula L.) and two conifers, Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.), and to determine the effects of the most relevant monoterpenes on transformations of soil N. The study site was a 70-year-old tree species experiment in Kivalo, northern Finland. VOCs were collected using two methods, passive air samplers and a chamber method. In soil atmosphere under spruce and especially under pine, the concentrations of monoterpenes were high, α- and β-pinene, Δ-3-carene and myrcene being the most abundant compounds, whereas concentrations of monoterpenes in soil atmosphere under birch were negligible. Samples of humus layer from the birch stand incubated in vitro and exposed to vapors from monoterpenes typical of coniferous forest soil showed decreased rates of net N mineralization but simultaneously increased rates of C mineralization. The response of soil microbial biomass C and N to different monoterpenes varied, but some monoterpenes considerably decreased soil microbial biomass. Altogether these results suggest that these compounds have negative effects on soil N transformations, but may serve as carbon and energy source for part of soil microbes.     

Earthworm communities in birch stands with different origin in central Finland

The aim of the study was to compare earthworm communities in anthropogenous birch stands with different origin in Finland. A total of nine forest sites were investigated: three birch stands (Betula pendula) planted ca. 30 years prior to the study after clear-cutting of spruce stands (“Birch after Spruce”, BS), three birch stands planted ca. 30 years earlier on arable soil that had been under normal cultivation until forestation (“Birch after Field”, BF), and three “Natural Deciduous” forests (D). Earthworms were sampled in May and October 1999 using a combination of formaline extraction and modified wet funnels. There were conspicuous differences between replicates of similarly managed forests. Earthworms were totally lacking in one of the D sites, while another had an abundant and diverse community. Only Dendrobaena octaedra was present in one BS site, while the two others harboured also Aporrectodea caliginosa and three Lumbricus species. All these species were also present in the BF sites, where their total biomass (ranging from 70 to 138 g (f.w.)/m²) was 2.6 times the average in BS, and of the same magnitude as the average in natural deciduous stands. A separate experiment revealed that L. terresris and A. caliginosa, which are not found in the surrounding coniferous forest, are able to live and reproduce in the soil of the D site where they were absent. It was concluded that earthworm species survive and reproduce in birch stands established on arable soil, where they have invaded during the long cultivation. On the other hand, their possibilities to disperse from cultural landscapes determine their presence not only in birch stands established in earlier coniferous forests, but also in “natural deciduous” forests where source populations are not present in the surroundings.     

Microbial biomass and activity in soil and litter underPinus sylvestris, Picea abies andBetula pendula at originally similar field afforestation sites

Microbial biomass C and N, and activities related to C and N cycles, were compared in needle and leaf litter, and in the uppermost 10 cm of soil under the litter layer in Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies L.) and silver birch (Betula pendula L.) stands, planted on originally similar field afforestation sites 23–24 years ago. The ground vegetation was differentiated under different tree species, consisting of grasses and herbs under birch and pine, and mosses or no vegetation with a thick layer of needles under spruce. The C:N ratio of the soils was 13–21 and the soil pH_{CaCl 2} 3.8–5.2. Both showed little variation under different tree species. Microbial biomass C and N, C mineralization, net ammonification, reduction) did not differ significantly in soil under different tree species either. Birch leaf litter had a higher pH_{CaCl 2} (5.9) than spruce and pine needle litter (pH 5.0 and 4.8, respectively). The C:N ratio of spruce needles was 30, and was considerably higher in pine needles (69) and birch leaves (54). Birch leaves tended to have the highest microbial biomass C and C mineralization. Spruce needles appeared to have the highest microbial biomass N and net formation of mineral N, whereas formation of mineral N in pine needles and birch leaves was negligible. Microbial biomass C and N were of the same order of magnitude in the soil and litter samples but C mineralization was tenfold higher in the litter samples.     

Comparison of Nitrate Leaching under Silver Birch (Betula Pendula) and Corsican Pine (Pinus Nigra SSP. Laricio) in Flanders (Belgium)

In a forest in Flanders (Belgium), situated in a region of intensive livestock production, comparable stands of Corsican pine and silver birch were studied for (1) NH₄ ⁺ and NO₃ ^- concentrations in throughfall water and soil solution and (2) depositions and leaching of NH₄ ⁺ and NO₃ ^- to groundwater. In each stand, throughfall collectors and porous cup lysimeters at three depths (0.1m, 0.5m and 1m) were installed in three replicated sets. Throughfall concentrations of ammonium and nitrate were significantly different for both species as well as soil solution concentrations of nitrate at all depths. Under pine, nitrate concentrations of the soil solution at 1m depth clearly exceeded the Belgian critical level for drinking water (50 mg.1^-1). Under birch, this level was only sporadically exceeded. During the sampling period, the depositions of NH₄ ⁺-N and NO₃ ^--N reached respectively 21.6 kg/ha and 6.3 kg/ha under birch and 81.3 and 15.2 kg/ha under Corsican pine. Nitrate-N leaching under silver birch amounted 25.4 kg/ha whereas 56.4 kg/ha was measured under Corsican pine.     

The effect of feeding behavior on Hg accumulation in the ecophysiologically different earthworms Lumbricus terrestris and Octolaseon cyaneum: A microcosm experiment

 A soil microcosm experiment was performed to assess the uptake of Hg from various Hg-spiked food sources (soil, leaf litter and root litter of Trifolium alexandrinum) by two earthworm species, Lumbricus terrestris (anecic) and Octolaseon cyaneum (endogeic). Treatments were applied in which one of the three food sources was Hg spiked and the other two were not. Additional treatments in which all or none of the food sources were Hg spiked were used as controls. Uptake of Hg from soil into tissues of both earthworm species was significantly higher than uptake of Hg from leaf litter or root litter, indicating that soil may be the most important pool for the uptake of Hg into earthworms. In addition, the anecic L. terrestris significantly accumulated Hg from all Hg-spiked food sources (leaf litter, root litter and soil), whereas the endogeic O. cyaneum took up Hg mainly from soil particles. Interestingly, there was no further increase in Hg in L. terrestris when all food sources were Hg spiked compared to the single Hg-spiked sources. This may be attributed to the relatively high Hg content in the soil, which may have influenced the feeding behavior of the earthworms, although their biomass did not significantly decline. We suggest that, in addition to the physiological differences, feeding behavior may also play a role in the contrasting uptake of Hg by the two earthworm species.     

Introducing deep burrowing earthworms (Lumbricus terrestris L.) into arable heavy clay under boreal conditions

An inoculation of the deep burrowing earthworm Lumbricus terrestris L. was carried out for soil amelioration purposes at an arable field on heavy clay in southern Finland. L. terrestris, whilst entirely absent from the field and its immediate surrounding, was present in many other local soils. Prior to inoculation, field management was changed in favour of L. terrestris by improving the subdrainage and by implementation of reduced tillage. In Autumn 1996, 82 earthworm inoculation units (EIUs) were introduced in three transects at one end of the field, one transect lying within a permanent grass strip outside the cultivated area. In Autumn 1998, L. terrestris had persisted outside the cultivated area with maximum density of 28 ind. m⁻² (median: 0 ind. m⁻²). No individuals were found inside the field. In Autumn 2003, maximum density was 43 ind. m⁻² (median: 9 ind. m⁻²) outside the cultivated area, with evidence for 8 m dispersal from inoculation points (1.1 m year⁻¹). In 2003 individuals were also found in low densities inside the field, close to subdrains, with a maximum density of 4 ind. m⁻². Results call for critical evaluation of this inoculation practice.     

Short-term uptake of <Superscript>15</Superscript>NH<Subscript>4</Subscript><Superscript>+</Superscript> into soil microbes and seedlings of pine,spruce and birch in potted soils

Short-term competition between soil microbes and seedlings of Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies (L.) Karst.) and silver birch (Betula pendula Roth) for N was assessed in a pot study using (¹⁵NH₄)₂SO₄ as a tracer. Seedlings were grown in organic and mineral soil, collected from a podsol soil; 3.18 mg (¹⁵NH₄)₂SO₄ per pot were injected into the soil, corresponding to 4 µg ¹⁵N g^-1 d.m. (dry matter) mineral soil and 17 µg ¹⁵N g^-1 d.m. organic soil. The amounts of N and ¹⁵N in the seedlings and in microbial biomass derived from fumigation-extraction were measured 48 h after addition of ¹⁵N. In the mineral soil, 19–30% of the added ¹⁵N was found in the plants and 14–20% in the microbial biomass. There were no statistically significant differences between the tree species. In the organic soil, 74% of the added ¹⁵N was recovered in the microbial biomass in birch soil, compared to 26% and 17% in pine and spruce soils, respectively. Correspondingly, about 70% of the ¹⁵N was recovered in pine and spruce seedlings, and only 23% in birch seedlings. In conclusion, plants generally competed more successfully for added ¹⁵NH₄ ⁺ than soil microbes did. An exception was birch growing in organic soil, where the greater amount of available C from birch root exudates perhaps enabled micro-organisms to utilise more N.     

Interactive effects of tree species and soil moisture on methane consumption

Methane consumption by temperate forest soils is a major sink for this important greenhouse gas, but little is known about how tree species influence CH₄ uptake by soils. Here, we show that six common tree species in Siberian boreal and temperate forests significantly affect potential CH₄ consumption in laboratory microcosms. Overall, soils under hardwood species (aspen and birch) consumed CH₄ at higher rates than soils under coniferous species and grassland. While NH₄⁺ addition often reduces CH₄ uptake, we found no effect of NH₄⁺ addition, possibly because of the relatively high ratio of CH₄-to-NH₄⁺ in our incubations. The effects of soil moisture strongly depended on plant species. An increase in soil moisture enhanced CH₄ consumption in soils under spruce but had the opposite effect under Scots pine and larch. Under other species, soil moisture did not affect CH₄ consumption. These results could be explained by specific responses of different groups of CH₄-oxidizing bacteria to elevated moisture.