Ectomycorrhizal fungal mycelia turnover in a longleaf pine forest (original) (raw)
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Mycorrhiza, 2007
The rates and controls of ectomycorrhizal fungal production were assessed in a 22-year-old longleaf pine (Pinus palustris Mill.) plantation using a complete factorial design that included two foliar scorching (control and 95% plus needle scorch) and two nitrogen (N) fertilization (control and 5 g N m −2 year −1 ) treatments during an annual assessment. Ectomycorrhizal fungi production comprised of extramatrical mycelia, Hartig nets and mantles on fine root tips, and sporocarps was estimated to be 49 g m −2 year −1 in the control treatment plots. Extramatrical mycelia accounted for approximately 95% of the total mycorrhizal production estimate. Mycorrhizal production rates did not vary significantly among sample periods throughout the annual assessment (p=0.1366). In addition, reduction in foliar leaf area via experimental scorching treatments did not influence mycorrhizal production (p=0.9374), suggesting that stored carbon (C) may decouple the linkage between current photosynthate production and ectomycorrhizal fungi dynamics in this forest type. Nitrogen fertilization had a negative effect, whereas precipitation had a positive effect on mycorrhizal fungi production (p=0.0292; r 2 =0.42). These results support the widely speculated but poorly documented supposition that mycorrhizal fungi are a large and dynamic component of C flow and nutrient cycling dynamics in forest ecosystems.
Soil Biology and Biochemistry, 2013
Mycorrhizal fungi constitute a considerable sink for carbon in most ecosystems. This carbon is used for building extensive mycelial networks in the soil as well as for metabolic activity related to nutrient uptake. A number of methods have been developed recently to quantify production, standing biomass and turnover of extramatrical mycorrhizal mycelia (EMM) in the field. These methods include minirhizotrons, in-growth mesh bags and cores, and indirect measurements of EMM based on classification of ectomycorrhizal fungi into exploration types. Here we review the state of the art of this methodology and discuss how it can be developed and applied most effectively in the field. Furthermore, we also discuss different ways to quantify fungal biomass based on biomarkers such as chitin, ergosterol and PLFAs, as well as molecular methods, such as qPCR. The evidence thus far indicates that mycorrhizal fungi are key components of microbial biomass in many ecosystems. We highlight the need to extend the application of current methods to focus on a greater range of habitats and mycorrhizal types enabling incorporation of mycorrhizal fungal biomass and turnover into biogeochemical cycling models.
Oecologia, 2004
Nutrient uptake by forest trees is dependent on ectomycorrhizal (EM) mycelia that grow out into the soil from the mycorrhizal root tips. We estimated the production of EM mycelia in root free samples of pure spruce and mixed spruce-oak stands in southern Sweden as mycelia grown into sand-filled mesh bags placed at three different soil depths (0-10, 10-20 and 20-30 cm). The mesh bags were collected after 12 months and we found that 590±70 kg ha −1 year −1 of pure mycelia was produced in spruce stands and 420±160 kg ha −1 year −1 in mixed stands. The production of EM mycelia in the mesh bags decreased with soil depth in both stand types but tended to be more concentrated in the top soil in the mixed stands compared to the spruce stands. The fungal biomass was also determined in soil samples taken from different depths by using phospholipid fatty acids as markers for fungal biomass. Subsamples were incubated at 20°C for 5 months and the amount of fungal biomass that degraded during the incubation period was used as an estimate of EM fungal biomass. The EM biomass in the soil profile decreased with soil depth and did not differ significantly between the two stand types. The total EM biomass in the pure spruce stands was estimated to be 4.8 ±0.9×10 3 kg ha −1 and in the mixed stands 5.8 ±1.1×10 3 kg ha −1 down to 70 cm depth. The biomass and production estimates of EM mycelia suggest a very long turnover time or that necromass has been included in the biomass estimates. The amount of N present in EM mycelia was estimated to be 121 kg N ha −1 in spruce stands and 187 kg N ha −1 in mixed stands. The δ 13 C value for mycelia in mesh bags was not influenced by soil depth, indicating that the fungi obtained all their carbon from the tree roots. The δ 13 C values in mycelia collected from mixed stands were intermediate to values from pure spruce and pure oak stands suggesting that the EM mycelia received carbon from both spruce and oak trees in the mixed stands. The δ 15 N value for the EM mycelia and the surrounding soil increased with soil depth suggesting that they obtained their entire N from the surrounding soil.
Biomass of external mycelium of ectomycorrhizal fungi in Norway spruce stands in Poland
Acta Mycologica, 2015
Biomass of extramatrical mycorrhizal mycelium (EMM) was examined under canopies of mature Norway spruce trees grown in different forest stands in Poland. Two mountain forest sites (Brenna and Salmopol), one upland site (Zwierzyniec) and one lowland site (Mirachowo) have been investigated, using sand-filled meshbags method. The in-grow mesh-bags were buried in the soil for 12 months (since October up to the next October) or for 4 months (since June up to October) at four depths at each site: 5, 15, 30 and 45 cm (Brenna and Salmopol) or 60 cm (Zwierzyniec and Mirachowo). The mycelium biomass was estimated from the ergosterol content determined in the mesh-bags. The results indicated significant differences in EMM production and their vertical distribution between the mountain and the upland and lowland forest sites. The lowest EMM biomass was found at the experimental plot in the mountainious site Brenna. Considerable decrease of EMM biomass with the soil depth was recorded after 12 months of the mesh-bags incubation in soil in the upland and lowland sites, while in the mountain forests decrease of the EMM biomass in the lower soil depths diminished more gradually EMM biomass determined in the mesh-bags placed in soil at the upper 5 and 15 cm tended to be higher after 4 months than after 12 months of incubation period. Such results suggest that the time necessary for evaluation of EMM biomass in soil may be limited to the summer-autumn months, when the production of EMM is the highest. Variable stress factors can influence decreased ectomycorrhizal mycelium production and/or their destruction. Further research in different forest types and regions are needed for better understanding factors determining EMM biomass production and surviving in soil.
2012
Aims There is evidence that increased N inputs to boreal forests, via atmospheric deposition or intentional fertilization, may impact negatively on ectomycorrhizal (ECM) fungi leading to a reduced flux of plant-derived carbon (C) back to the atmosphere via ECM. Our aim was to investigate the impact of N fertilization of a Pinus sylvestris (L.) forest stand on the return of recently photoassimilated C via the ECM component of soil respiration. Methods We used an in situ, large-scale, 13 C-CO 2 isotopic pulse labelling approach and monitored the 13 C label return using soil gas efflux chambers placed over three different types of soil collar to distinguish between heterotrophic (R H ), autotrophic (R A ; partitioned further into contributions from ECM hyphae and total R A ) and total (R S ) soil respiration. Results The impact of N fertilization was to significantly reduce R A , particularly respiration via extramatrical ECM hyphae. ECM hyphal flux in control plots showed substantial spatial variability, resulting in mean flux estimates exceeding estimates of total R A , while ECM contributions to R A in N treated plots were estimated at around 30%. Conclusion Significant impacts on soil C cycling may be caused by reduced plant C allocation to ECM fungi in response to increased N inputs to boreal forests; ecosystem models so far lack this detail.
2011
A substantial portion of the carbon (C) fixed by the trees is allocated belowground to ectomycorrhizal (EM) symbionts, but this fraction usually declines after fertilization. The aim of the present study was to estimate the effect of optimal fertilization (including all the necessary nutrients) on the growth of EM fungi in young Norway spruce forests over a three year period. In addition, the amount of carbon sequestered by EM mycelia was estimated using a method based on the difference in d 13 C between C 3 and C 4 plants. Sand-filled ingrowth mesh bags were used to estimate EM growth, and similar bags amended with compost made from maize leaves (a C 4 plant) were used to estimate C sequestration. Fertilizers had been applied either every year or every second year since 2002 and the estimates of EM growth started in 2007. The application of fertilizer reduced EM growth to between 0% and 40% of the growth in the control plots at one site (Ebbegärde), while no significant effect was found at the other three sites studied. The effect of the fertilizer was similar in sand-filled and maize-compost-amended mesh bags, but the total production of EM fungi was 3-4 times higher in maize-compost-amended mesh bags. The fertilizer tended to reduce EM growth more when applied every year than when applied every second year. The amount of C sequestered in maize-compost-amended mesh bags collected from unfertilized treatments was estimated to be between 0.2 and 0.7 mg C g sand À1 at Ebbegärde and between 0.2 and 0.5 mg C g sand À1 at Grängshammar. This corresponds to between 300 and 1100 kg C per ha, assuming a similar production in the soil as in the mesh bags. Fertilization at the Ebbegärde site reduced carbon sequestration, which confirmed the results based on estimates of fungal growth (ergosterol levels). A correlation was found between fungal biomass and d 13 C in mesh bags amended with maize compost. Based on this, it was estimated that a fungal production of 1 lg ergosterol corresponded to 0.33 mg of sequestered carbon. In conclusion, the effect of the fertilizer on EM growth seemed to be dependent on the effect of the fertilizer on tree growth. Thus, at Ebbegärde, were tree growth was less stimulated by the fertilizer, EM growth was reduced upon fertilization. At other sites, where tree growth was more stimulated, the fertilizer did not influence EM growth. The large amounts of carbon sequestered during the experiment may be a result of fungal residues remaining in the soil after the death of the hyphae.
Plant and Soil, 2013
There is growing evidence of the importance of extramatrical mycelium (EMM) of mycorrhizal fungi in carbon (C) cycling in ecosystems. However, our understanding has until recently been mainly based on laboratory experiments, and knowledge of such basic parameters as variations in mycelial production, standing biomass and turnover as well as the regulatory mechanisms behind such variations in forest soils is limited. Presently, the production of EMM by ectomycorrhizal (EM) fungi has been Plant Soil
New Phytologist, 1999
The use of biofuels has been proposed as one possible substitute for fossil fuels, which contribute substantially to the increase in [CO # ] in the atmosphere. However, increased harvesting of forest residues for biofuel might affect the availability of base cations, P and N, as well as the development, community dynamics and function of ectomycorrhizas. This in turn might influence nutrient uptake and tree growth. In this study we investigated the effects of repeated forest residue harvesting on ectomycorrhizal species colonizing spruce roots in the humus layer of a 35-yr-old forest. Harvesting significantly decreased the thickness of the humus layer as well as decreasing the numbers of ectomycorrhizal root tips both per metre root length and per unit humus volume. Changes in mycorrhizal community structure were studied by ITS typing with the use of PCR-RFLP analysis. In total, 19 different ITS types were found on two different sampling occasions (autumn and spring) ; 11 of these were common to both samplings. Nine of the ITS types were identified to at least the genus level by comparison with RFLP patterns of identified fruiting bodies or axenic cultures. Five species, Cortinarius sp. 2, Thelephora terrestris (Ehrenb.) Fr., Lactarius theiogalus (Bull. :Fr.) S. F. Gray s.st. Neuhoff, Tylospora fibrillosa Donk and To$-96-12, occurred on over 5% of the total sampled root tips. Together these five types colonized 63% of the mycorrhizas screened. A similarity index assessment showed no shift in mycorrhizal community structure as a result of harvesting. Our findings suggest that the repeated removal of forest residues might have a strong effect on the quantity and development of ectomycorrhizal roots in the organic horizon, but little effect on the species composition of the community.