Fungal colonization as affected by litter depth and decomposition stage of needle litter

The present study was designated to evaluate the relative effects of litter depth and decomposition stage of needles on fungal colonization of needle litter in field experiments. The experiment was carried out in coniferous temperate forests in central Japan. Needle litter of Chamaecyparis obtusa and Pinus pentaphylla var. himekomatsu at two decomposition stages (recently dead and partly decomposed) were placed into the organic layer at two depths (on the surface of and beneath the litter layer). Fungal colonization of needles after 1 year was examined in terms of hyphal abundance and frequency of fungal species. Total and live hyphal length on needles were affected by the litter depth and (or) the decomposition stage of needles. Length of darkly pigmented hyphae on needles was 1.7-2.6 times greater beneath the litter layer than on the litter surface regardless of the decomposition stage of needles. Length of clamp-bearing hyphae in Pinus pentaphylla was 5.0-5.2 times greater in partly decomposed needles than in recently dead needles regardless of the litter depth. Frequencies of Pestalotiopsis spp. and Cladosporium cladosporioides were higher on recently dead needles than on partly decomposed needles and (or) were higher on the litter surface than beneath the litter layer. Frequencies of Trichoderma, Penicillium, and Umbelopsis species generally were higher on partly decomposed needles than on recently dead needles and were higher beneath the litter layer than on the surface.     

Effects of fungal inocula on the decomposition of lignin and structural polysaccharides in Pinus sylvestris litter

 Litter bags containing sterile Scots pine (Pinus sylvestris) needles (19.8% lignin, 26.5% cellulose and 0.34% N) were inoculated with two species of fungi in the laboratory and then placed in the litter layer of a pine plantation. Marasmius androsaceus, which can degrade lignocellulose, was initially displaced by other fungal colonisers and was not detected in the litter after 2–3 months; but was re-isolated from the needles after 12 months. Trichoderma viride, which is a cellulolytic species and also antagonistic to other fungi, dominated the litter throughout the experiment. The control litter was naturally colonised by litter fungi. After 12 months, mass losses were similar at 52% for M. androsaceus and 48% for T. viride, compared with 36% for the control litter colonised by a more complex fungal community. Lignin concentrations increased with time in control litter and with T. viride because mass losses of carbohydrates were greater than those of lignin. Litter inoculated with M. androsaceus showed significant lignin decomposition throughout the experiment but cellulose concentrations showed a proportional increase in the first 6 months, suggesting that the fungus was preferentially exploiting hemicellulose and non-structural carbohydrates. Analysis of TFA-extractable sugars (mainly from hemicellulose) and CuO-derived phenylpropanoid moieties from lignin confirmed the differential patterns of resource decomposition which were not evident from total mass losses. During the initial stages of decomposition, T. viride was as effective in utilising structural polysaccharides as the complex fungal community in the control litter. Furthermore, M. androsaceus not only exhibited unexpectedly low cellulolytic activity but also facilitated lignin depolymerisation after the fungus was no longer detectable in the litter. The pre-inoculation of litter with these two fungal species therefore affected the overall dynamics of decomposition at a biochemical level. This study illustrates the importance of understanding the effects and interactions of specific fungi, rather than assumptions about the functional competence of diverse communities, on the processes of litter decomposition. Received: 5 July 2000     

Do oribatid mites (Acari: Oribatida) show a higher preference for ubiquitous vs. specialized saprotrophic fungi from pine litter?

Previous studies of oribatid mite feeding preferences for different saprotrophic fungi were limited to ubiquitous fungal species, whereas saprophytes specialized to decompose particular substrates have been neglected. We examined the preference of seven oribatid mite species (Adoristes ovatus, Eniochthonius minutissimus, Eueremaeus silvestris, Nothrus silvestris, Oppiella subpectinata, Porobelba spinosa and Spatiodamaeus verticillipes) for nine autochthonous saprotrophic fungi from Scots pine litter (Pinus sylvestris). Among the fungal species offered were specific coniferous litter colonizers (Allantophomopsis lycopodina, Ceuthospora pinastri, Hormonema dematioides, Scleroconidioma sphagnicola, Verticicladium trifidum, Marasmius androsaceus and Sympodiella acicola) and two ubiquitous species (Cladosporium herbarum and Oidiodendron griseum). The fungi were inoculated on fragments of pine needles and offered simultaneously and separately to the mites. Our main hypothesis, that oribatid mites (usually occurring in more than one type of ecosystems) would prefer the ubiquitous fungal species rather than those specific to pine litter, was supported only partly. The ubiquitous C. herbarum was highly preferred by all studied mites, but most of them preferred one or more of the specialized fungi with similar intensity. The basidiomycete M. androsaceus along with sterile needles were consistently avoided by all mites in all experiments. Our results do not support the hypothesis, that the “true” fungivorous oribatid mites in traditional sense are more selective fungal feeders than are the “unspecialized” panphytophagous ones. We observed no gradation in preference of fungi for oribatid mites as a group, but rather a discontinuous and dynamic mosaic with particular mites preferring particular fungal species. This heterogeneous mosaic shapes the feeding niches occupied by particular oribatid mite species and probably reduces competition for food source among numerous species coexisting in a given habitat and time.     

Immobilization of avian excreta-derived nutrients and reduced lignin decomposition in needle and twig litter in a temperate coniferous forest

The possible effects of excreta of the Great Cormorant Phalacrocorax carbo on decomposition processes and dynamics of nutrients (N, P, Ca, K, Mg) and organic chemical components (lignin, total carbohydrates) were investigated in a temperate evergreen coniferous forest near Lake Biwa in central Japan. Two-year decomposition processes of needles and twigs of Chamaecyparis obtusa were examined at two sites, control site never colonized by the cormorants (site C) and colonizing site (site 2). Mass loss was faster in needles than in twigs. Mass loss of these litter types was faster at site C than at site 2, which was ascribed to the decreased mass loss rate of acid-insoluble ‘lignin’ at site 2. Net immobilization of N, P, and Ca occurred in needles and twigs at site 2; whereas at site C, mass of these elements decreased without immobilization during decomposition. Duration of immobilization phase of these nutrients at site 2 was estimated to be 1.6 to 2.5 years in needles and 19.6 to 23.5 years in twigs. Immobilization potential (maximum amount of exogenous nutrient immobilized per gram initial material) was similar between needles and twigs for N and Ca but was about 10 times higher in twigs than in needles for P. δ¹³C in needles was relatively constant during the first year and then increased during the second year, whereas δ¹³C in twigs was variable during decomposition. Acid-insoluble fraction was depleted in ¹³C compared to whole needles (1.6-2.1‰) and twigs (2.0-2.5‰). δ¹⁵N of needles and twigs and their acid-insoluble fractions approached to δ¹⁵N of excreta during decomposition at site 2. This result demonstrated the immobilization of excreta-derived N into litter due to the formation of acid-insoluble lignin-like substances complexed with excreta-derived N. No immobilization occurred in K and Mg and their mass decreased during decomposition at both sites. Based on these results of nutrient immobilization during decomposition and on the data of litter fall and excreta amount at site 2, we tentatively calculated stand-level immobilization potential of litter fall and its contribution to total amount of N and P deposited as excreta. Thus, the potential maximum amount immobilized into litter fall (needles and twigs) was estimated to account for 5-7% of total excreta-derived N and P.     

Interaction between two decomposer basidiomycetes and a collembolan under Sitka spruce: Grazing and its potential effects on fungal distribution and litter decomposition

The hypothesis that selective grazing by Onychiurus latus Gisin altered the outcome of competition between two basidiomycetes, Mycena galopus (Pers. ex Fr.) Kummer and Marasmius androsaceus (L. ex Fr.) Fr. on Sitka spruce litter was tested. Without grazing M. androsaceus had the higher colonizing ability of litter in the laboratory whereas with grazing the position was reversed. Field exclusion experiments also suggested that higher densities of O. latus resulted in a reduction in the activity of M. androsaceus and an increase in that of M. galopus and vice versa. It is suggested that selective grazing may be an important factor determining the field distribution of these fungi, since M. androsaceus, the preferred food source, was restricted to the L litter horizon where the density of O. latus was not high enough to alter the outcome of competition whereas M. galopus predominated in the F₁ horizon. The rate of decomposition of Sitka spruce litter in the laboratory caused by M. androsaceus (17% dry weight loss over 6 months at 11°C) increased by 4–14% in the presence of grazing by O. latus whereas that of M. galopus (11.5%) decreased by 12–16%.     

Production of lignocellulose-degrading enzymes and degradation of leaf litter by saprotrophic basidiomycetes isolated from a Quercus petraea forest

Due to the production of lignocellulose-degrading enzymes, saprotrophic basidiomycetes can significantly contribute to the turnover of soil organic matter. The production of lignin- and polysaccharide-degrading enzymes and changes of the chemical composition of litter were studied with three isolates from a Quercus petraea forest. These isolates were capable of fresh litter degradation and were identified as Gymnopus sp., Hypholoma fasciculare and Rhodocollybia butyracea. Within 12 weeks of incubation, H. fasciculare decomposed 23%, R. butyracea 32% and Gymnopus sp. 38% of the substrate dry mass. All fungi produced laccase and Mn-peroxidase (MnP) and none of them produced lignin peroxidase or other Mn-independent peroxidases. There was a clear distinction in the enzyme production pattern between R. butyracea or H. fasciculare compared to Gymnopus sp. The two former species caused the fastest mass loss during the initial phase of litter degradation, accompanied by the temporary production of laccase (and MnP in H. fasciculare) and also high production of hydrolytic enzymes that later decreased. In contrast, Gymnopus sp. showed a stable rate of litter mass loss over the whole incubation period with a later onset of ligninolytic enzyme production and a longer lasting production of both lignin and cellulose-degrading enzymes. The activity of endo-cleaving polysaccharide hydrolases in this fungus was relatively low but it produced the most cellobiose hydrolase. All fungi decreased the C/N ratio of the litter from 24 to 15-19 and Gymnopus sp. also caused a substantial decrease in the lignin content. Analytical pyrolysis mass spectrometry of litter decomposed by this fungus showed changes in the litter composition similar to those caused by white-rot fungi during wood decay. These changes were less pronounced in the case of H. fasciculare and R. butyracea. All fungi also changed the mean masses of humic acid and fulvic acid fractions isolated from degraded litter. The humic acid fraction after degradation by all three fungi contained more lignin and less carbohydrates. Compared to the decomposition by saprotrophic basidiomycetes, litter degradation in situ on the site of fungal isolation resulted in the relative enrichment of lignin and differences in lignin composition revealed by analytical pyrolysis. It can most probably be explained by the participation of non-basidiomycetous fungi and bacteria during natural litter decomposition.     

Do same-aged but different height Norway spruce (Picea abies) clones affect soil microbial community?

The influence of individual trees in monocrop forests on soil microbial communities is poorly understood. We measured basal respiration, substrate-induced respiration and phospholipid fatty acids (PLFA), bacterial growth rate with the ³H-thymidine incorporation technique and fungal growth rate as ¹⁴C-acetate incorporation into ergosterol to investigate whether slow- and fast-growing 12-year-old Norway spruce (Picea abies) clones have affected differently on their associated soil microbial communities. Understorey vegetation, soil chemical properties and elemental concentrations of needles were also determined. The slow- and fast-growing spruce clones differed in PLFA profiles, understorey vegetation and elemental concentrations in needles suggesting that spruce clones have directly or indirectly affected soil microbes.     

Differential response of ectomycorrhizal and saprotrophic fungal mycelium from coniferous forest soils to selected monoterpenes

The mycelia of saprotrophic (SP) and ectomycorrhizal (ECM) fungi occur throughout the upper soil horizons in coniferous forests and could therefore be exposed to high concentrations of monoterpenes occurring in the needle litter of some tree species.Monoterpenes are mycotoxic and could potentially affect fungi that are exposed to them in the litter layers. In order to investigate whether monoterpenes typical of coniferous litters could influence fungal communities, we analysed the monoterpene content of freshly fallen needles of Pinus sylvestris, Picea abies and Picea sitchensis. The most abundant monoterpenes were found to be α-pinene, β-pinene and 3-carene. We evaluated the effects of these three monoterpene vapours on the biomass production of 23 SP isolates and 16 ECM isolates. Overall, 75% of ECM isolates and 26% of SP isolates were significantly inhibited by at least one of the monoterpene treatments and both intra- and inter-specific variations in response were observed.Monoterpene concentrations are highest in surface litters. The differential effects on fungal taxa may influence the spatial and temporal distribution of fungal community composition, indirectly affecting decomposition and nutrient cycling, the fundamental ecosystem processes in which fungi have a key role in coniferous forest soils.     

Reduction of fungal growth and lignin decomposition in needle litter by avian excreta

The effects of excessive addition of excreta from the Great Cormorant Phalacrocorax carbo, a colonial piscivorous bird, on the growth and the ability of fungi to decompose needle litter of Chamaecyparis obtusa were examined by a pure-culture test. Colony growth rate, mass loss of needle litter, and utilization patterns of lignin and carbohydrates were investigated and compared for 22 species in basidiomycetes, ascomycetes, and zygomycetes. Colony growth rate of basidiomycetes decreased on medium supplemented with excreta (excreta medium) as compared to control medium without excreta, whereas such a difference was not found for ascomycetes. Mass loss of needle litter caused by basidiomycetes was generally higher than those caused by ascomycetes and zygomycetes. Basidiomycetes decomposed both lignin and carbohydrates in various proportions, whereas ascomycetes and zygomycetes decomposed carbohydrates selectively. Mass loss of litter caused by basidiomycetes and ascomycetes was lower when incubated on excreta medium than on control medium. Mass loss of lignin and nitrogen caused by basidiomycetes was lower on excreta medium than on control medium, whereas such differences were not found for ascomycetes. Mass loss of carbohydrate was not different between the media for basidiomycetes or ascomycetes.     

No evidence that nitrogen enrichment affect fungal communities of Vaccinium roots in two contrasting boreal forest types

In boreal forests ericaceous shrubs often dominate the forest floor vegetation. Nitrogen enrichment has been shown to decrease shrub abundance and in this study we explored whether it also affects the root associated fungal communities. Fine roots of Vaccinium myrtillus were collected in a Norway spruce dominated forest and of Vaccinium vitis-idaea in a Scots pine dominated forest. In both forests, nitrogen enrichment was experimentally induced by adding 12.5 and 50 kg N ha⁻¹ yr⁻¹ for 12 (spruce forest) and four (pine forest) years. Based on terminal restriction fragment length polymorphisms, subcloning and sequencing analyses, the root associated fungal communities were examined. We found 93 fungal species including Asco-, Basidio- and Zygo-mycota. In general, the Rhizoscyphus ericae aggregate was the most dominant and this was followed by Herpotrichiellaceae and Sebacina. Ordination analysis revealed that nitrogen enrichment did not change species composition of the fungal communities in neither the spruce nor the pine forest, while fungal community structures were clearly discriminated between the dominant shrub species in each forest. Similarly, no fungal species showed a significant response to nitrogen enrichment. Therefore, nitrogen enrichment appears to have no effect on root associated fungi of understorey dwarf shrubs in boreal forests, while it is clear that spruce and pine forests harbor distinctive communities of these fungi.     

Litter type, but not plant cover, regulates initial litter decomposition and fungal community structure in a recolonising cutover peatland

Cutover peatlands are often rapidly colonised by pioneer plant species, which have the potential to affect key ecosystem processes such as carbon (C) turnover. The aim of this study was to investigate how plant cover and litter type affect fungal community structure and litter decomposition in a cutover peatland. Intact cores containing Eriophorum vaginatum, Eriophorum angustifolium, Calluna vulgaris and bare soil were removed and a mesh bag with litter from only one of each of these species or fragments of the moss Sphagnum auriculatum was added to each core in a factorial design. The presence or absence of live plants, regardless of the species, had no effect on mass loss, C, nitrogen (N) or phosphorus (P) concentrations of the litter following 12 months of incubation. However, there was a very strong effect of litter type on mass loss and concentrations of C, N and P between most combinations of litter. Similarly, plant species did not affect fungal community structure but litter type had a strong effect, with significant differences between most pairs of litter types. The data suggest that labile C inputs via rhizodeposition from a range of plant functional types that have colonised cutover bogs for 10-15 years have little direct effect on nutrient turnover from plant litter and in shaping litter fungal community structure. In contrast, the chemistry of the litter they produce has much stronger and varied effects on decomposition and fungal community composition. Thus it appears that there is distinct niche differentiation between the fungal communities involved in turnover of litter versus rhizodeposits in the early phases of plant succession on regenerating cutover peatlands.     

Degradation of hemicellulose, cellulose and lignin in decomposing spruce needle litter in relation to N

Decomposing needles from a Norway spruce forest in southern Sweden were studied for 559 days under laboratory conditions. Falling needles were collected in control (Co) plots and plots that had received 100 kg N ha⁻¹ yr⁻¹ as (NH₄)₂SO₄ for 9 years under field conditions. One of the aims was to determine whether the previously documented low decomposition rate of the N fertilized (NS) needles could be explained by a lower degradation degree of lignin. The lignin content was studied using the alkaline CuO oxidation method, the Klason lignin method and CPMAS ¹³C NMR spectroscopy. The amounts of cellulose and hemicellulose were also determined.The fertilized needle litters initially decomposed faster than the unfertilized, but later this reaction reversed, so that at the end the mass loss was 45% initial C in the control and 35% initial C in NS. Klason lignin decreased with time in both treatments and overall, the change of Klason lignin mirrored the litter mass loss. No major difference as regards the decomposition of hemicellulose occurred between the treatments, whereas significantly lower concentrations of cellulose were found in NS needles throughout the incubation. The CuO derived compounds (VSC) were somewhat lower in NS needles throughout the decomposition time. Initially, VSC increased slightly in both treatments, which contradicts the Klason lignin data. There was a weak positive relationship (p>0.05) between VSC and Klason lignin. Both vanillyls compounds (V) and cinnamyl compounds (Ci) increased slightly during decomposition, whereas syringyl compounds (S) vanished entirely. The lignin degradation degree, i.e. the acid-to-aldehyde ratio of the vanillyl compounds expressed as (Ac/Al)_v, showed no significant effect of treatment. The ¹³C NMR analyses of the combined samples showed increased content of aromatic C with increasing decomposition time. The carbohydrate content (O-alkyl C) was lower in the fertilized needle litter throughout the incubation time. The alkyl C content tended to increase with decomposition time and N fertilization. The alkyl C/O-alkyl C ratios increased in both treatments during the incubation. The NMR results were not tested statistically.In conclusion, no major difference concerning lignin degradation could be found between the unfertilized and N fertilized needle litter. Thus, the study contradicts the hypothesis that higher amounts of N reduce lignin degradation. The reduced biological activity is probably due to direct N effects on the microorganisms and their decomposing ability.     

Fungal growth on a common wood substrate across a tropical elevation gradient: Temperature sensitivity, community composition, and potential for above-ground decomposition

Fungal breakdown of plant material rich in lignin and cellulose (i.e. lignocellulose) is of central importance to terrestrial carbon (C) cycling due to the abundance of lignocellulose above and below-ground. Fungal growth on lignocellulose is particularly influential in tropical forests, as woody debris and plant litter contain between 50% and 75% lignocellulose by weight, and can account for 20% of the C stored in these ecosystems. In this study, we evaluated factors affecting fungal growth on a common wood substrate along a wet tropical elevation gradient in the Peruvian Andes. We had three objectives: 1) to determine the temperature sensitivity of fungal growth - i.e. Q₁₀, the factor by which fungal biomass increases given a 10 °C temperature increase; 2) to assess the potential for above-ground fungal colonization and growth on lignocellulose in a wet tropical forest; and 3) to characterize the community composition of fungal wood decomposers across the elevation gradient. We found that fungal growth had a Q₁₀ of 3.93 (95% CI of 2.76-5.61), indicating that fungal biomass accumulation on the wood substrate nearly quadrupled with a 10 °C increase in temperature. The Q₁₀ for fungal growth on wood at our site is higher than Q₁₀ values reported for litter decomposition in other tropical forests. Moreover, we found that above-ground fungal growth on the wood substrate ranged between 37% and 50% of that measured in the soil, suggesting above-ground breakdown of lignocellulose represents an unexplored component of the C cycle in wet tropical forests. Fungal community composition also changed significantly along the elevation gradient, and Ascomycota were the dominant wood decomposers at all elevations. Fungal richness did not change significantly with elevation, directly contrasting with diversity patterns observed for plant and animal taxa across this gradient. Significant variation in fungal community composition across the gradient suggests that the characteristics of fungal decomposer communities are, directly or indirectly, influenced by temperature.     

Influence of urea on the cherry leaf spot pathogen, Blumeriella jaapii, and on microorganisms in decomposing cherry leaves

Growth chamber and orchard experiments were carried out to clarify the response of the cherry leaf spot pathogen, Blumeriella jaapii, and microorganisms in the leaf litter to two levels of urea (2.5% or 5%) applied post leaf fall to sour cherry leaves. In general, urea application reduced the development of B. jaapii measured as biomass by quantitative PTA-ELISA in the leaf litter and spore counting in the spring and increased the overall microbial biomass (measured with biomarker phospholipid fatty acids (PLFAs)) and fungal activity (assessed as β-N-acetylglucosaminidase activity). The biomass increase of all groups of microorganisms in the litter generally began immediately after application of urea and, with the notable excepts of B. jaapii and Gram-negative bacteria containing the biomarker PLFA cyclo17:0, continued after the ammonia and pH levels had stabilized to levels similar to the control leaves approximately 10 days later. Application of 2.5% urea increased the biomass of most groups of saprotrophic microorganisms and accelerated litter decomposition to a higher extent than application of 5% urea and during the first week after treatment applications the 5% urea level inhibited the total microbial biomass. This may be ascribed to ammonia toxicity as 5% urea resulted in a markedly higher ammonia elevation than 2.5% urea, the first week after application. From then onwards in both the 2.5% and 5% treatments the fungal and Gram-positive communities benefited from a lowered C:N ratio, increasing their activities approximately 2 times compared to a water-treated control. Inhibition of B. jaapii coincided with the period of urea breakdown in which elevated levels of ammonia and leaf pH were measured in the treated leaves. This period lasted for approximately 10 days after which the biomass of B. jaapii in the treated leaves continued to decrease at a slow but faster rate than in the untreated leaves. Our results indicate that the urea application caused an elevation in leaf pH and ammonia levels, which together with an acceleration in litter decomposition, adversely affected the saprotrophic growth of B. jaapii, leading to reduced production of ascospores and winter-conidia the following spring.     

Interaction between two decomposer basidiomycetes and a collembolan under Sitka spruce: Distribution,abundance and selective grazing

The fungi Mycena galopus (Pers. ex Fr.) Kummer and Marasmius androsaceus (L. ex Fr.) Fr. produced over 99% of the fruit bodies on a site under Picea sitchensis Carr. In the laboratory M. androsaceus colonized litter from both the L and F₁ horizons more than twice as fast as M. galopus, but in the field its distribution was restricted to the L horizon whereas M. galopus occurred in the F₁ horizon. The collembolan Onychiurus latus Gisin, the most abundant mycophagous arthropod at the field site which consistently had basidiomycete hyphae in its gut, was found throughout the autumn in the litter horizons in densities up to 600 m^?2. This collembolan showed a marked preference for the mycelium of M. androsaceus rather than that of M. galopus in both laboratory and field tests. These results suggest that selective grazing by O. latus may be an important factor in determining the vertical distribution of these two fungi in the field.     

Carbon dioxide emissions of soils under pure and mixed stands of beech and spruce, affected by decomposing foliage litter mixtures

Soil respiration is the largest terrestrial source of CO₂ to the atmosphere. In forests, roughly half of the soil respiration is autotrophic (mainly root respiration) while the remainder is heterotrophic, originating from decomposition of soil organic matter. Decomposition is an important process for cycling of nutrients in forest ecosystems. Hence, tree species induced changes may have a great impact on atmospheric CO₂ concentrations. Since studies on the combined effects of beech-spruce mixtures are very rare, we firstly measured CO₂ emission rates in three adjacent stands of pure spruce (Picea abies), mixed spruce-beech and pure beech (Fagus sylvatica) on three base-rich sites (Flysch) and three base-poor sites (Molasse; yielding a total of 18 stands) during two summer periods using the closed chamber method. CO₂ emissions were higher on the well-aerated sandy soils on Molasse than on the clayey soils on Flysch, characterized by frequent water logging. Mean CO₂ effluxes increased from spruce (41) over the mixed (55) to the beech (59) stands on Molasse, while tree species effects were lower on Flysch (30-35, mixed > beech = spruce; all data in mg CO₂-C m⁻² h⁻¹). Secondly, we studied decomposition after fourfold litter manipulations at the 6 mixed species stands: the O_i - and O_e horizons were removed and replaced by additions of beech -, spruce - and mixed litter of the adjacent pure stands of known chemical quality and one zero addition (blank) in open rings (20 cm inner diameter), which were covered with meshes to exclude fresh litter fall. Mass loss within two years amounted to 61-68% on Flysch and 36-44% on Molasse, indicating non-additive mixed species effects (mixed litter showed highest mass loss). However, base cation release showed a linear response, increasing from the spruce - over the mixed - to the beech litter. The differences in N release (immobilization) resulted in a characteristic converging trend in C/N ratios for all litter compositions on both bedrocks during decomposition. In the summers 2006 and 2007 we measured CO₂ efflux from these manipulated areas (a closed chamber fits exactly over such a ring) as field indicator of the microbial activity. Net fluxes (subtracting the so-called blank values) are considered an indicator of litter induced changes only and increased on both bedrocks from the spruce - over the mixed - to the beech litter. According to these measurements, decomposing litter contributed between 22-32% (Flysch) and 11-28% (Molasse) to total soil respiration, strengthening its role within the global carbon cycle.     

Uptake of Sr from microcline and biotite by ectomycorrhizal fungi in a Norway spruce forest

Ectomycorrhizal (EM) fungi growing in symbiosis with tree seedlings have been found in laboratory experiments to stimulate weathering and the uptake of nutrients from silicate minerals. In the present study, we used the natural abundance of strontium isotopes to confirm that these fungi obtain strontium from biotite and microcline under field conditions. Minerals enriched in radiogenic ⁸⁷Sr were introduced into fungal in-growth mesh bags and placed under a 5-10 cm thick humus layer developed on boulders in a Norway spruce (Picea abies (L.) Karst) forest in south-western Sweden. EM root tips were sampled above the mesh bags and the ⁸⁷Sr/⁸⁶Sr ratio was used to calculate the fraction of Sr in the root tips that originated from the minerals. In the EM root tips sampled above the mesh bags containing the different minerals, 1.5% of the Sr originated from biotite and 6.4% from microcline. The amount of Sr was more than 300 times higher in the mesh bags containing microcline than in those containing biotite, indicating that proportionally more Sr was released from the biotite. This study demonstrates that EM fungi have the potential to take up measurable amounts of nutrients, such as Ca and K, from microcline and biotite in the field.     

Changes in the lignin fraction of spruce and pine needle litter during decomposition as studied by some chemical methods

Changes in the lignin fraction of spruce and pine needle litter were followed by four different methods: Klason lignin, phloroglucinol lignin, dioxane-water-HCl-lignin and alkaline CuO oxidation. The decomposition patterns of the lignins studied were different, the largest differences between the methods being obtained for the spruce needles. Depending on the method used, between 36 and 46% of the original amount of lignin remained in the pine needles and about 30–61% in the spruce needles after 3 yr of decomposition. The decomposition rates of the various lignin pools were highly correlated with the loss in mass of the litter. The phloroglucinol lignin was decomposed significantly faster (P < 0.001) in the spruce needles than in the pine needles for as long as the decomposition process was followed. During decomposition of the litter, the residual amount of Klason lignin was correlated with the residual amounts of the other lignins. The application of the different methods to litter decomposition studies is discussed.     

Low levels of nitrogen addition stimulate decomposition by boreal forest fungi

Climate warming and associated increases in nutrient mineralization may increase the availability of soil nitrogen (N) in high latitude ecosystems, such as boreal forests. These changes in N availability could feed back to affect the decomposition of litter and organic matter by soil microbes. Since fungi are important decomposers in boreal forest ecosystems, we conducted a 69-day incubation study to examine N constraints on fungal decomposition of organic substrates common in boreal ecosystems, including cellulose, lignin, spruce wood, spruce needle litter, and moss litter. We added 0, 20, or 200 μg N to vials containing 200 mg substrate in factorial combination with five fungal species isolated from boreal soil, including an Ascomycete, a Zygomycete, and three Basidiomycetes. We hypothesized that N addition would increase CO₂ mineralization from the substrates, particularly those with low N concentrations. In addition we predicted that Basidiomycetes would be more effective decomposers than the other fungi, but would respond weakly or negatively to N additions. In support of the first hypothesis, cumulative CO₂ mineralization increased from 635 ± 117 to 806 + 108 μg C across all fungal species and substrates in response to 20 μg added N; however, there was no significant increase at the highest level of N addition. The positive effect of N addition was only significant on cellulose and wood substrates which contained very little N. We also observed clear differences in the substrate preferences of the fungal species. The Zygomycete mineralized little CO₂ from any of the substrates, while the Basidiomycetes mineralized all of the substrates except spruce needles. However, the Ascomycete (Penicillium) was surprisingly efficient at mineralizing spruce wood and was the only species that substantially mineralized spruce litter. The activities of β-glucosidase and N-acetyl-glucosaminidase were strongly correlated with cumulative respiration (r = 0.78 and 0.74, respectively), and Penicillium was particularly effective at producing these enzymes. On moss litter, the different fungal species produced enzymes that targeted different chemical components. Overall, our results suggest that fungal species specialize on different organic substrates, and only respond to N addition on low N substrates, such as wood. Furthermore, the response to N addition is non-linear, with the greatest substrate mineralization at intermediate N levels.     

Invertebrate grazing affects nitrogen partitioning in the saprotrophic fungus Phanerochaete velutina

The heterogeneity of nutrients in forest soils is governed by many biotic and abiotic factors. The significance of nutrient patchiness in determining soil processes remains poorly understood. Some saprotrophic basidiomycete fungi influence nutrient heterogeneity by forming large mycelial networks that enable translocation of nutrients between colonized patches of dead organic matter. The effect of mycophagous soil fauna on these networks and subsequent nutrient redistribution has, however, been little studied. We used a soil microcosm system to investigate the potential effects of a mycophagous collembola, Protaphorura armata, on nutrient transfer within, and nutrient loss from, the mycelium of a saprotrophic basidiomycete fungus, Phanerochaete velutina. A ¹⁵N label, added to central mycelium, was used to track nitrogen movement within the microcosms across 32 days. Although collembola grazing had little impact on δ¹⁵N values, it did alter the partitioning of ¹⁵N between different regions of mycelia. Less ¹⁵N was transferred to new mycelial growth in grazed systems than in ungrazed systems, presumably because collembola reduced fungal growth rate and altered mycelial morphology. Surprisingly, collembola grazing did not increase the mineralization of N from mycelium into the bulk soil. Overall, our results suggest that mycophagous soil fauna can alter nutrient flux and partitioning within fungal mycelium; this has the potential to affect the dynamics and spatial heterogeneity of forest floor nutrients.