Particle size fractionation of fungal and bacterial biomass in subalpine grassland and forest soils

Characterization of soil aggregates according to particle size fractions is a useful tool in process-oriented research into soil organic matter and biological properties. Substrate-induced respiration (SIR) inhibition was used to quantify microbial, fungal and bacterial biomass in particle size fractions of soils ranging from forest to grassland in a subalpine region of central Taiwan. In addition, ergosterol content was determined in the same samples to verify fungal biomass measured by SIR inhibition technique. Surface soil (0–10 cm) was fractionated into four particle size fractions: coarse sand (250–2000 μm), fine sand (53–250 μm), silt (2–53 μm) and clay (0.2–2 μm). The larger sized fractions (>250 μm and 53–250 μm) contained higher levels of fungal ergosterol than the smaller sized ones (2–53 μm and 0.2–2 μm). The largest particle size fraction (250–2000 μm) from all studied habitats showed the highest level of microbial biomass, with no clear trend in microbial biomass level among the other size fractions. SIR-calculated fungal biomass level and ergosterol converted fungal biomass content were positively correlated (r=0.71, p<0.05), and such correlation decreased as biomass levels were high. Ratios of fungi to bacteria ranged between 0.6 and 1.3 in fractions obtained in this study. This study indicates a high variability of microbial (fungal and bacterial) biomass level among particle size fractions in soil, and that the large-sized fractions tend to contain a high level of microbial biomass in a given ecosystem.     

Does soil ergosterol concentration provide a reliable estimate of soil fungal biomass?

Our aim was to determine if soil ergosterol concentration provides a quantitative estimate of the soil fungal biomass concentration, as is usually assumed. This was done by comparing soil ergosterol measurements with soil fungal biomass (fungal biomass C) concentrations estimated by microscopic measurements and by the selective inhibition technique linked to substrate-induced respiration (SIR). The measurements were compared in a silty-clay loam soil given a range of previous treatments designed to increase or decrease the soil fungal biomass and so also to change the soil ergosterol concentration. The treatments used were ryegrass amendment, to increase the total and fungal biomass, and CHCl₃-fumigation and the addition of the biocides, captan, bronopol and dinoseb, to decrease both ergosterol and fungal biomass C concentrations. The mineralization of ergosterol following addition to sand innoculated with soil extract, and to a sandy loam soil, was also determined. The added ergosterol was little, if at all, degraded following addition to either sand or the unfumigated or fumigated soil during a 10 d aerobic incubation. Similarly, pesticide addition did not significantly change soil ergosterol concentrations yet the soil fungal biomass C concentration decreased significantly. Thus, the ratio: (soil ergosterol concentration/soil fungal biomass C concentration) was much higher in the pesticide-treated soils than the control soil. Following ryegrass amendment, soil ergosterol concentration increased from about 6-12 μg⁻¹ soil within 5 d and then decreased gradually to about 7 μg g⁻¹ soil by 20 d incubation. Changes in fungal biomass C (measured by direct microscopy) closely mirrored changes in soil ergosterol over this period. However, when the amended soil was fumigated and then incubated for a further 5 d, the initial ergosterol concentration declined from 7 to 5 μg g⁻¹ soil by 20 d incubation (a decline of about 0.4). The comparable decline in fungal biomass C was about eight-fold. Thus the ratio of ergosterol to fungal biomass C increased from 0.005 to about 0.01. There was a significant correlation (r>0.84, P<0.001) between soil ergosterol concentration and fungal biomass measured by either SIR or microscopy. However, three data points played a vital role in the correlation. When these points were excluded the relationship was very poor (r<0.4). Our results therefore suggest that substantial amounts of ergosterol may exist, other than in living cells, for considerable periods, with little, if any mineralization. Thus, these results indicate that ergosterol and fungal biomass C concentrations are not always closely correlated, due to the slow metabolism of ergosterol in recently dead fugal biomass and/or the existence of exocellular ergosterol in soil.     

The use of phospholipid fatty acid analysis to estimate bacterial and fungal biomass in soil

The cell content of 12 bacterial phospholipid fatty acids (PLFA) was determined in bacteria extracted from soil by homogenization/centrifugation. The bacteria were enumerated using acridine orange direct counts. An average of 1.40×10^-17 mol bacterial PLFA cell^-1 was found in bacteria extracted from 15 soils covering a wide range of pH and organic matter contents. With this factor, the bacterial biomass based on PLFA analyses of whole soil samples was calculated as 1.0–4.8 mg bacterial C g^-1 soil C. The corresponding range based on microscopical counts was 0.3–3.0 mg bacterial C g^-1 soil C. The recovery of bacteria from the soils using homogenization/centrifugation was 2.6–16% (mean 8.7%) measured by PLFA analysis, and 12–61% (mean 26%) measured as microscopical counts. The soil content of the PLFA 18:26 was correlated with the ergosterol content (r=0.92), which supports the use of this PLFA as an indicator of fungal biomass. The ratio 18:26 to bacterial PLFA is therefore suggested as an index of the fungal:bacterial biomass ratio in soil. An advantage with the method based on PLFA analyses is that the same technique and even the same sample is used to determine both fungi and bacteria. The fungal:bacterial biomass ratio calculated in this way was positively correlated with the organic matter content of the soils (r=0.94).     

Comparison of soil fungal/bacterial ratios in a pH gradient using physiological and PLFA-based techniques

We have compared the total microbial biomass and the fungal/bacterial ratio estimated using substrate-induced respiration (SIR) in combination with the selective inhibition technique and using the phospholipid fatty acid (PLFA) technique in a pH gradient (3.0-7.2) consisting of 53 mature broad-leaved forest soils. A fungal/bacterial biomass index using the PLFA technique was calculated using the PLFA 18:2ω6,9 as an indicator of fungal biomass and the sum of 13 bacterial specific PLFAs as indicator of the bacterial biomass. Good linear correlation (p<0.001) was found between the total microbial biomass estimated with SIR and total PLFAs (totPLFA), indicating that 1 mg biomass-C was equivalent to 130 nmol totPLFA. Both biomass estimates were positively correlated to soil pH. The fungal/bacterial ratio measured using the selective inhibition technique decreased significantly with increasing pH from about 9 at pH 3 to approximately 2 at pH 7, while the fungal/bacterial biomass index using PLFA measurements tended to increase slightly with increasing soil pH. Good correlation between the soil content of ergosterol and of the PLFA 18:2ω6,9 indicated that the lack of congruency between the two methods in estimating fungal/bacterial ratios was not due to PLFA 18:2ω6,9-related non-fungal structures to any significant degree. Several PLFAs were strongly correlated to soil pH (R² values >0.8); for example the PLFAs 16:1ω5 and 16:1ω7c increased with increasing soil pH, while i16:0 and cy19:0 decreased. A principal component analysis of the total PLFA pattern gave a first component that was strongly correlated to soil pH (R²=0.85, p<0.001) indicating that the microbial community composition in these beech/beech-oak forest soils was to a large extent determined by soil pH.     

Changes in soil fungal:bacterial biomass ratios following reductions in the intensity of management of an upland grassland

In this study we examined the effect on soil fungal:bacterial biomass ratios of withholding fertiliser, lime, and sheep-grazing from reseeded upland grassland. The cessation of fertiliser applications on limed and grazed grassland resulted in a reduction in soil pH from 5.4 to 5.1. The cessation of fertiliser applications and liming on grazed grassland resulted in a fall in pH from 5.4 to 4.7, whereas withholding fertiliser and lime and the removal of grazing resulted in a further reduction to pH 4.5. Substrate-induced respiration was reduced in the unfertilised grazed (21%; P<0.01) and unfertilised ungrazed (36%; P<0.001) treatments. Bacterial substrate-induced respiration and bacterial fatty acids were unaffected by the treatments. The relative abundance of the fungal fatty acid 18:26 increased by 39 and 72% (P<0.05) in the limed grazed and unfertilised grazed treatments, respectively. Fungal substrate-induced respiration increased in the limed grazed (18%) and unfertilised grazed (65%; P<0.05) treatments. The ratio of 18:26: bacterial fatty acids was correlated with the ratio of fungal:bacterial substrate-induced respiration (r=0.69; P<0.001).     

Effects of coniferous resin on fungal biomass and mineralisation processes in wood ant nest materials

 Wood ants (Formica rufa group) often bring large quantities of conifer resin to their mounds. The aim of this study was to test the hypothesis that the resin acts as a fungicide and thereby reduces C and N mineralisation. Two laboratory incubation experiments were carried out using two different materials: F/H layer from a Scots pine (Pinus sylvestris) stand and mixed litter from Scots pine and Norway spruce (Picea abies) stands. We estimated the effects of resin addition on fungal biomass and on the rates of C and N mineralisation. Addition of resin to the F/H material caused an increase in fungal biomass and C mineralisation, whereas N mineralisation decreased. Addition of resin to litter material did not significantly affect fungal biomass or C and N mineralisation. The results indicate that rather than having a fungicidal effect, resin acts as a C source that increases C mineralisation (mainly from the resin itself) and decreases net N mineralisation. The latter factor might be important in preventing plants dependent on inorganic N from invading and covering the ant mounds. Received: 17 December 1998     

Can the extent of degradation of soil fungal mycelium during soil incubation be used to estimate ectomycorrhizal biomass in soil?

The extent of degradation of the fungal biomass in forest soil during laboratory incubation was investigated as a measure of ectomycorrhizal (EM) biomass. The method simulates the disappearance of fungal mycelium after root trenching, where the EM fungi, deprived of its energy source (the tree), will start to die off. Incubating a forest humus soil at 25 °C resulted in a decrease in the relative proportion (mol%) of the phospholipid fatty acid 18:2ω6,9 (a fungal marker molecule) within 3-6 months, indicating that fungal biomass was disappearing. Incubation at 5 °C resulted in essentially no change in the amount of 18:2ω6,9. The measurement of ergosterol, another fungal marker molecule, gave similar results. Incubation of different forest soils (pine, spruce and spruce/oak), and assuming that the disappearance of fungal biomass during this period of time was entirely due to EM fungi, resulted in an estimation of EM biomass of between 47 and 84% of the total fungal biomass in these soils. The humus layer had more EM biomass than deeper mineral layers.     

Spatial distribution of fungal communities in a coastal grassland soil

In grasslands, saprotrophic fungi, including basidiomycetes, are major decomposers of dead organic matter, although spatial distributions of their mycelial assemblages are little described. The aim of this study was to characterise the scale and distribution of saprotrophic fungal communities in a coastal grassland soil using terminal restriction fragment length polymorphism (T-RFLP).Soil fungi were sampled at Point Reyes, California, USA, by taking forty-five 26 mm diam. cores in a spatially defined manner. Within each sampled core, complete core sections at 1-2 cm and 14-15 cm depths were removed and sub-sampled for DNA extraction and amplification using the primer pairs ITS1F-FAM/ITS4 (general fungi) or ITS1F-FAM/ITS4B (basidiomycete-specific).Nonmetric Multidimensional Scaling showed that general fungal communities could be clearly separated by depth, although basidiomycete communities could not. There were no strong patterns of community similarity or dissimilarity for general or basidiomycete fungal communities at horizontal geographical distances from 25 cm to 96 m in the upper horizon. These results show considerable vertical, but little horizontal, variability in fungal community structure in a semi-natural grassland at the spatial scales measured here.     

Influence of polychlorinated biphenyls on microbial biomass and its activity in grassland soil

Microbial biomass content, soil respiration and biomass specific respiration rate were measured in two parts of an area polluted by a municipal waste incinerator [polychlorinated biphenyls (PCBs) from combustion processes]. The soils in the studied parts differed significantly only in their levels of PCBs. The concentration of PCBs found in a control plot (4.4 ng g^-1 soil) can be regarded as a background value while the polluted plot contained an increased amount of PCBs (14.0 ng g^-1 soil). A significantly lower microbial biomass (decreased by 23%, based on the chloroform-fumigation extraction technique) and a lower specific respiration rate (decreased by 14%) were observed in the polluted plot in comparison with the control plot at the end of experimental period (1992–1994). Furthermore, a lower ability of microorganisms in the polluted plot to convert available C_org into new biomass was found in laboratory incubations with glucose-amended samples.     

Ergosterol and microbial biomass relationship in soil

Ergosterol and microbial biomass C were measured in 26 arable, 16 grassland and 30 forest soils. The ergosterol content ranged from 0.75 to 12.94 g g^-1 soil. The geometric mean ergosterol content of grassland and forest soils was around 5.5 g g^-1, that of the arable soils 2.14 g g^-1. The ergosterol was significantly correlated with biomass C in the entire group of soils, but not in the subgroups of grassland and forest soils. The geometric mean of the ergosterol: microbial biomass C ratio was 6.0 mg g^-1, increasing in the order grassland (5.1), arable land (5.4) and woodland (7.2). The ergosterol:microbial biomass C ratio had a strong negative relationship with the decreasing cation exchange capacity and soil pH, indicating that the fungal part of the total microbial biomass in soils increased when the buffer capacity decreased. The average ergosterol concentration calculated from literature data was 5.1 mg g^-1 fungal dry weight. Assuming that fungi contain 46% C, the conversion factor from micrograms ergosterol to micrograms fungal biomass C is 90. For soil samples, neither saponification of the extract nor the more effective direct saponification during extraction seems to be really necessary.     

Response of the soil fungal community to multi-factor environmental changes in a temperate forest

Both environmental and climatic changes are known to influence soil microbial biomes in terrestrial ecosystems. However, there are limited data defining the interactive effects of multi-factor environmental disturbances, including N-deposition, precipitation, and air temperature, on soil fungal communities in temperate forests. A 3-year outdoor pot experiment was conducted to examine the temporal shifts of soil fungal communities in a temperate forest following N-addition, precipitation and air temperature changes. The shifts in the structure and composition of soil fungal communities were characterized by denaturing gradient gel electrophoresis and DNA sequencing. N-addition regimen induced significant alterations in the composition of soil fungal communities, and this effect was different at both higher and lower altitudes. The response of the soil fungal community to N-addition was much stronger in precipitation-reduced soils compared to soils experiencing enhanced precipitation. The combined treatment of N-addition and reduced precipitation caused more pronounced changes in the lower altitude versus those in the higher one. Certain fungal species in the subphylum Pezizomycotina and Saccharomycotina distinctively responded to N fertilization and soil water control at both altitudes. Redundancy discrimination analysis showed that changes in environmental factors and soil physicochemical properties explained 43.7% of the total variability in the soil fungal community at this forest ecosystem. Variations in the soil fungal community were significantly related to the altitude, soil temperature, total soil N content (TN) and pH value (P < 0.05). We present evidence for the interactive effects of N-addition, water manipulation and air temperature to reshape soil fungal communities in the temperate forest. Our data could provide new insights into predicting the response of soil micro-ecosystem to climatic changes.     

Dynamics of soil biomass C,N, and P in a dry tropical forest in India

Summary Three dry tropical forest soils along a topographic sequence were examined to determine the seasonal dynamics of microbial C, N, and P. The lowest microbial biomass was found in forest soils at the foot of the hill followed by midslope forest soils. The hilltop soil, which had the most fine particles, water-holding capacity, organic C, and total N, reflected the presence of greater amounts of microbial C, N, and P. Mean annual microbial C, N, and P ranges were 466–662, 48–72 to 21–30 g g^-1, respectively. The seasonal pattern of microbial biomass, C, N, and P was similar at all sites, the values being greatest during the dry season and lowest during the wet season. The seasonal values for microbial biomass C, N, and P were positively correlated with each other and a negative correlation was found between microbial biomass and the fine root mass in these forest soils.     

Determination of microbial biomass and fungal and bacterial distribution in cattle faeces

As an important component of organic fertilizers, animal faeces require methods for determining diet effects on their microbial quality to improve nutrient use efficiency in soil and to decrease gaseous greenhouse emissions to the environment. The objectives of the present study were (i) to apply the chloroform fumigation extraction (CFE) method for determining microbial biomass in cattle faeces, (ii) to determine the fungal cell-membrane component ergosterol, and (iii) to measure the cell-wall components fungal glucosamine and bacterial muramic acid as indices for the microbial community structure. Additionally, ergosterol and amino sugar data provide independent control values for the reliability of the microbial biomass range obtained by the CFE method. A variety of extractant solutions were tested for the CFE method to obtain stable extracts and reproducible microbial biomass C and N values, leading to the replacement of the original 0.5 M K₂SO₄ extractant for 0.05 M CuSO₄. The plausibility of the data was assessed in a 28-day incubation study at 25 °C with cattle faeces of one heifer, where microbial biomass C and N were repeatedly measured together with ergosterol. Here, the microbial biomass indices showed dynamic characteristics and possible shifts in the microbial community. In faeces of five different heifers, the mean microbial biomass C/N ratio was 5.6, the mean microbial biomass to organic C ratio was 2.2%, and the mean ergosterol to microbial biomass C ratio was 1.1‰. Ergosterol and amino sugar analysis revealed a significant contribution of fungi, with a percentage of more than 40% to the microbial community. All three methods are expected to be suitable tools for analysing the quality of cattle faeces.     

Microbial reaction in activity, biomass, and community structure after long-term continuous mixing of a grassland soil

Long-term continuous mixing at 40% water holding capacity (WHC) or as slurry at 400% WHC should result in increased soil organic matter decomposition rates in comparison to a control treatment at 40% WHC, but may have strong impacts on soil microbial indices for activity, biomass, and community structure. The amount of extractable inorganic N (NO₃-N+NH₄-N) accumulated in the soil solution after 40 weeks of incubation at 25 °C was 3% of total N in the control treatment and 4% in the two continuous mixing treatments. However, in the treatment mixing at 40% WHC, this 33% increase compared to the control treatment might be explained solely by the decrease in microbial biomass N. In the control treatment, microbial indices decreased in the order microbial biomass C (−10%), microbial biomass N (−40%), ergosterol (−45%) and ATP (−60%). In the treatment mixing at 40% WHC, all four microbial biomass indices were significantly lower than the respective index in the control treatment. This was especially true for microbial biomass N. In the treatment mixing as slurry, only the contents of microbial biomass C and ATP were significantly lower in comparison to the control treatment. The correspondence analysis ordination biplot of the phospholipid fatty acid (PLFA) profiles showed distinct clusters for the three treatments at the end of the incubation. The strongest relative decline of 64% was observed for the fungi-specific PLFA 18:3ω6 in the treatment mixing as slurry in comparison to the control treatment. The content of total bacterial PLFA decreased only by 23%. The differences between the control treatment and the treatment mixing at 40% WHC were less apparent. Fungi represent on average 21% of total microbial biomass C at the end of the incubation if the ergosterol content is recalculated into fungal biomass C. In accordance with this percentage, 22% of the group-specific PLFA could be attributed to fungi.     

Response of ecosystem respiration to soil water and plant biomass in a semiarid grassland

Abstract

The Mongolian steppe zone constitutes a major part of East Asian grasslands. The objective of this study was to evaluate the quantitative dependence of ecosystem respiration (R_eco) on the environmental variables of soil water and plant biomass in a semiarid grassland ecosystem. We determined R_eco using opaque, closed chambers in a Mongolian grassland dominated by graminaceous perennial grasses during six periods: July 2004, May 2005, July 2005, September 2005, June 2006, and August 2009. Using the data collected when soil water content and aboveground biomass were relatively constant, values of R_eco were fitted to an exponential temperature function, and the standardized rate of R_eco at 20°C (R₂₀) and temperature sensitivity (Q₁₀) of R_eco were calculated for each measurement plot and period. The results indicate that aboveground biomass significantly affected the variation in R₂₀, and the relationship was expressed with a linear model. The R₂₀ residuals of the linear biomass model were highly correlated with soil water content by a quadratic function. The Q₁₀ values showed a weak positive relationship with soil water content. Temporal and spatial variations in R_eco were well predicted by the exponential temperature model with R₂₀, which relates to aboveground biomass and soil water content, and with Q₁₀, which relates to soil water content.     

Antagonistic and synergistic effects of fungal and bacterial growth in soil after adding different carbon and nitrogen sources

The effect of adding easily available and more complex carbon sources, with and without nitrogen, on fungal and bacterial growth and activity in soil were studied in the laboratory. Total microbial activity was estimated by measuring respiration, fungal growth with the acetate-in-ergosterol incorporation technique and bacterial growth with the thymidine and leucine incorporation techniques. The substrate additions consisted of glucose and cellulose, with and without nitrogen (as ammonium nitrate), and gelatine. The microbial development was followed over a 2-month period. The respiration rate increased within a few days after adding glucose, with and without nitrogen, and gelatine, initially by more than 10 times, but after 2 months no differences were seen compared with the control. Bacterial growth estimated with the thymidine and leucine incorporation techniques gave similar results. Adding glucose with nitrogen, or gelatine, increased bacterial growth within a few days up to 10 times, but even after 2 months of incubation bacterial growth rates were still about 5 times higher than in the control. Adding only glucose increased bacterial growth rates by about twice over the whole incubation period. Fungal growth rates especially increased after adding cellulose and nitrogen, although a minor increase was found after adding cellulose alone. Fungal growth rates started to increase after 10 days of incubation with cellulose. There were indications of synergistic effects in that bacterial growth increased after the fungi had started to grow after adding cellulose. Treatments resulting in high bacterial growth rates (adding easily available carbon sources) led to decreased fungal growth rates compared with the control, indicating antagonistic effects of bacteria.     

Responses of the bacterial and fungal biomass in a grassland soil to multi-year applications of dairy manure slurry and fertilizer

Plots of a tall fescue (Festuca arundinacea) sward in the south coastal region of BC, Canada, were treated with dairy manure slurry or fertilizer at 50 or 100 kg NH₄-N ha⁻¹ up to four times per year for six consecutive years; control plots received no manure or fertilizer. The length of fungal hyphae and abundance of bacterial cells were determined by direct counting at 19 sample dates during the fourth (1997), fifth (1998) and sixth (1999) application years. Bacterial abundance was significantly greater in manured soil than in fertilized and untreated soils. In contrast, hyphal length was significantly greater in untreated soil than in manured and fertilized soils. In subplots that ceased to receive manure in 1998, bacterial abundance remained greater through 1998 and 1999 than in previously fertilized plots, indicating that the 4 year cumulative effect of manure was detectable for at least two growing seasons after applications cease. The apparently negative effect of manure and fertilizer on fungal hyphae also appeared to persist through 2 years after applications ceased. Bacterial abundance increased after an initial application of manure for 1 year to previously untreated plots, but not to levels comparable to plots treated with manure continuously from 1994 to 1998.Increases in bacterial abundance, during the one to three week intervals immediately following individual applications of manure, were inconsistent and other factors, such as soil moisture, temperature and perhaps crop phenology appear to have had strong effects on the timing of these microbial responses. Annual means for bacterial abundance and total microbial biomass in the continuous manure treatment were similar for all 3 years. This suggested that the manure-induced increase in microbial biomass probably reached a plateau between one and 3 years after applications commenced. The large bacterial populations along with abundant carbon substrates in manured soil, relative to fertilized soil, were probably capable of immobilizing influxes of mineral N, explaining the observations that less leaching occured from manured than from fertilized soils.     

Carbon stocks, soil respiration and microbial biomass in fire-prone tropical grassland, woodland and forest ecosystems

A thorough understanding of the role of microbes in C cycling in relation to fire is important for estimation of C emissions and for development of guidelines for sustainable management of dry ecosystems. We investigated the seasonal changes and spatial distribution of soil total, dissolved organic C (DOC) and microbial biomass C during 18 months, quantified the soil CO₂ emission in the beginning of the rainy season, and related these variables to the fire frequency in important dry vegetation types grassland, woodland and dry forest in Ethiopia. The soil C isotope ratios (δ¹³C) reflected the 15-fold decrease in the grass biomass along the vegetation gradient and the 12-fold increase in woody biomass in the opposite direction. Changes in δ¹³C down the soil profiles also suggested that in two of the grass-dominated sites woody plants were more frequent in the past. The soil C stock ranged from being 2.5 (dry forest) to 48 times (grassland) higher than the C stock in the aboveground plant biomass. The influence of fire in frequently burnt wooded grassland was evident as an unchanged or increasing total C content down the soil profile. DOC and microbial biomass measured with the fumigation-extraction method (C_mic) reflected the vertical distribution of soil organic matter (SOM). However, although SOM was stable throughout the year, seasonal fluctuations in C_mic and substrate-induced respiration (SIR) were large. In woodland and woodland-wooded grassland C_mic and SIR increased in the dry season, and gradually decreased during the following rainy season, confirming previous suggestions that microbes may play an important role in nutrient retention in the dry season. However, in dry forest and two wooded grasslands C_mic and SIR was stable throughout the rainy season, or even increased in this period, which could lead to enhanced competition with plants for nutrients. Both the range and the seasonal changes in soil microbial biomass C in dry tropical ecosystems may be wider than previously assumed. Neither SIR nor C_mic were good predictors of in situ soil respiration. The soil respiration was relatively high in infrequently burnt forest and woodland, while frequently burnt grasslands had lower rates, presumably because most C is released through dry season burning and not through decomposition in fire-prone systems. Shifts in the relative importance of the two pathways for C release from organic matter may have strong implications for C and nutrient cycling in seasonally dry tropical ecosystems.     

Soil carbon sequestration and changes in fungal and bacterial biomass following incorporation of forest residues

Sequestering carbon (C) in forest soils can benefit site fertility and help offset greenhouse gas emissions. However, identifying soil conditions and forest management practices which best promote C accumulation remains a challenging task. We tested whether soil incorporation of masticated woody residues alters short-term C storage at forested sites in western and southeastern USA. Our hypothesis was that woody residues would preferentially stimulate soil fungal biomass, resulting in improved C use efficiency and greater soil C storage. Harvest slash at loblolly pine sites in South Carolina was masticated (chipped) and either (1) retained on the soil surface, (2) tilled to a soil depth of 40 cm, or (3) tilled using at least twice the mass of organics. At comparative sites in California, live woody fuels in ponderosa pine stands were (1) masticated and surface applied, (2) masticated and tilled, or (3) left untreated. Sites with clayey and sandy soils were compared in each region, with residue additions ranging from 20 to 207 Mg ha⁻¹. Total and active fungal biomass were not strongly affected by residue incorporation despite the high input of organics. Limited response was also found for total and active bacterial biomass. As a consequence, fungal:bacterial (F:B) biomass ratios were similar among treatments at each site. Total soil C was elevated at one California site following residue incorporation, yet was significantly lower compared to surface-applied residues at both loblolly pine sites, presumably due to the oxidative effects of tilling on soil organic matter. The findings demonstrated an inconsequential effect of residue incorporation on fungal and bacterial biomass and suggest a limited potential of such practices to enhance long-term soil C storage in these forests.