The aim of this study was to investigate possible interactions between minerals and ectomycorrhizal (EM) mycelia. Fungal rhizomorphs growing in association with apatite and/or biotite were sampled both from a laboratory experimental system (Rhizopogon sp. and Pinus muricata) and from mesh bags buried in forest soil in the field. The elemental composition of the samples was analyzed with particle-induced X-ray emission (PIXE). Many EM rhizomorphs associated with apatite in laboratory systems and in mesh bags contained larger amounts of Ca (mean ranges between 12 and 31 mg Ca g(-1)) than similar rhizomorphs connected to acid-washed sand (range 0.3-3.5 mg Ca g(-1)). Ca originating from apatite was deposited as calcium oxalate crystals on the surface of the rhizomorphs. EM mycelium produced in mesh bags had a capacity to mobilize 0.6 mg P kg(-1) year(-1) from apatite-amended sand (which is 0.04% of the added apatite). A high concentration of K in some rhizomorphs (up to 11 mg K g(-1)) suggests that these fungi are good accumulators of K and may have a significant role in transporting K to trees.
In Sweden application of granulated wood ash has been suggested as a method to supplement nutrient loss resulting from harvesting of forest residues for bioenergy production. Mycelia of two ectomycorrhizal fungi Piloderma sp. 1 and Ha-96-3, were commonly found to colonise ash granules in a wood ash fertilised spruce forest. Thirty-eight fungal isolates were selected from 10 taxa to investigate the possible role of different ectomycorrhizal fungi in nutrient mobilisation from ash. The taxa were Cenococcum geophilum Fr., Piloderma croceum Erikss. and Hjortst., Piloderma sp. 1, Thelephora terrestris (Ehrenb.) Fr., Tylospora fibrillosa Donk, and five unidentified species, all originating from a wood ash fertilised spruce forest. The isolates were tested for their ability to solubilise tricalcium phosphate (TCP) or hardened wood ash (HWA) in vitro. Ha-96-3, P. croceum and Piloderma sp. 1 were the only taxa which solubilised TCP. Abundant calcium oxalate crystals were formed in TCP and HWA plates with Piloderma sp. 1. Ha-96-3 and two isolates of P. croceum produced intermediate amounts of crystals. Ha-96-1 and T. fibrillosa produced low amounts of crystal but no crystal formation was observed by any of the other isolates. Piloderma sp. 1 from HWA plates had significantly higher concentrations of P, compared to P. croceum or Ha-96-3. Piloderma sp. 1 and P. croceum were further tested for their ability to colonise wood ash in microcosms containing intact mycorrhizal associations. After 7 months Piloderma sp. 1 colonised ash amended patches with a dense, mat like mycelium, whereas P. croceum mycelia avoided the ash patches. Possible differences between these fungi in patterns of carbon allocation were investigated by labelling seedlings with 14CO(2). Piloderma sp. 1 mycelia allocated significantly more 14C to ash patches than P. croceum. P. croceum allocated relatively more 14C to control patches than to the ash patches. The possible role of ectomycorrhizal fungi in mobilisation of nutrients from wood ash is discussed.
The influence of ectomycorrhizal fungi on the soil bacterial community was studied by growing pine seedlings in artificial soils consisting of a peat/sand mixture amended with microcline, biotite or apatite. In the microcline‐amended and unamended soils both Suillus variegatus and Paxillus involutus reduced bacterial activity as measured by thymidine incorporation. S. variegatus grew best in the biotite soil, where it increased both bacterial activity and biomass as measured by microscopic counts and specific bacterial fatty acids. Further, the positive influence of S. variegatus on the bacteria in the biotite soil modified the bacterial community, as reflected in the bacteria‐specific phospholipid fatty acid composition. The increases in bacterial biomass and activity and changes in the bacterial community induced by S. variegatus may be due to the production of organic substances by this fungus, as indicated by an 10‐fold increase in soil‐solution citric acid. Two isolates of S. variegatus and an unidentified ectomycorrhizal fungus all tended to stimulate bacterial activity in the apatite‐amended soil in compartments isolated from roots by a mesh. We conclude that the same ectomycorrhizal fungus may stimulate bacterial growth under certain conditions and inhibit bacterial growth under other conditions.
Plant growth, nutrient uptake, microbial biomass and activity were studied in pot systems containing spruce seedlings colonised with different ectomycorrhizal fungi from an ash-fertilised forest. The seedling root systems were enclosed in mesh bags inside an outer compartment containing crushed, hardened wood ash. Three different species of mycorrhizal fungi and a non-mycorrhizal control were exposed to factorial combinations of ash and N addition. Ash treatment had a highly significant, positive effect on plant growth and on shoot and root concentrations of K, Ca and P, irrespective of mycorrhizal status. Mycorrhizal inoculation had a significant effect on plant growth, which was proportionally greater in the absence of ash. N addition had a significant positive effect on plant biomass in mycorrhizal treatments with ash, but no effect in non-mycorrhizal treatments or most of the mycorrhizal treatments without ash. Piloderma sp. 1, which was earlier found to colonise wood ash granules in field studies, appeared to accumulate Ca from ash in the mycorrhizal roots. 5-6.7% of the total P in the ash was solubilised, with 0.9-1.5% in solution, 3.6-4.6% in the plants and 0.5-1.5% in microbial biomass. Bacterial activity as determined by [(3)H]-thymidine and [(14)C]-leucine incorporation was significantly greater in ash treatments than in controls with no ash addition. Principal component analysis (PCA) of phospholipid fatty acids (PLFAs) showed a clear difference in bacterial community structure between samples collected from ash-treated pots and controls without ash.
Intensive harvesting of forest residues for energy production may lead to the depletion of organic matter and mineral nutrients in the forest floor. In order to restore nutrient content wood ash has been suggested as a fertiliser. Ectomycorrhizal (EM) fungi are involved in the nutrient uptake of forest trees and this study investigates the influence of intensive harvesting and wood ash fertilisation on the external EM mycelium in forest soil. Nylon mesh bags filled with sand were buried in September 1997 in field plots which had or had not been intensively harvested. The effect of wood ash on the production of external EM mycelium was studied in mesh bags amended with wood ash. Mesh bags were retrieved in May and October 1998. The relative amount of fungal mycelia in the mesh bags was estimated with phospholipid fatty acid analysis. The fungi colonising the mesh bags were mainly (>90%) ectomycorrhizal. Fungal biomass in the mesh bags was low in the spring but high in the autumn. No significant effect on EM fungal biomass was observed in the mesh bags collected from intensively harvested plots compared with those from control plots, but wood ash amendment resulted in 2.4 times more EM fungal biomass (P<0.05). The effect of external EM mycelium on the dissolution of wood ash was studied in mesh bags filled with wood ash, using mesh bags buried in soil isolated from roots as EM-free controls. The external EM mycelium had no effect on the dissolution rate of the wood ash. 80% of the potassium was lost from the wood ash within a month, whereas no phosphorus was lost during the experimental period (up to 13 months).
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