Spatial changes of soil fungal and bacterial biomass from a sub-alpine coniferous forest to grassland in a humid, sub-tropical region

 Fungal and bacterial biomass were determined across a gradient from a forest to grassland in a sub-alpine region in central Taiwan. The respiration-inhibition and ergosterol methods for the evaluation of the microbial biomass were compared. Soil fungal and bacterial biomass both significantly decreased (P<0.05) with the shift of vegetation from forest to grassland. Fungal and bacterial respiration rates (evolved CO₂) were, respectively, 89.1 μl CO₂ g^–1 soil h^–1 and 55.1 μl CO₂ g^–1 soil h^–1 in the forest and 36.7 μl CO₂ g^–1 soil h^–1 and 35.7 μl CO₂ g^–1 soil h^–1 in the grassland surface soils (0–10 cm). The fungal ergosterol content in the surface soil decreased from the forest zone (108 μg g^–1) to the grassland zone (15.9 μg g^–1). A good correlation (R ²=0.90) was exhibited between the soil fungal ergosterol content and soil fungal CO₂ production (respiration) for all sampling sites. For the forest and grassland soil profiles, microbial biomass (respiration and ergosterol) declined dramatically with depth, ten- to 100-fold from the surface organic horizon to the deepest mineral horizon. With respect to fungal to bacterial ratios for the surface soil (0–10 cm), the forest zone had a significantly (P<0.05) higher ratio (1.65) than the grassland zone (1.05). However, there was no fungal to bacterial ratio trend from the surface horizon to the deeper mineral horizons of the soil profiles. Received: 30 March 2000     

Earthworms increase the ratio of bacteria to fungi in northern hardwood forest soils, primarily by eliminating the organic horizon

The exotic earthworm invasion in hardwood forests of the northern United States is associated with many ecosystem-level changes. However, less is known about the effects of the invasion on the composition of the soil microbial community through which ecosystem-level changes are mediated. Further, earthworm effects on soil microbial community composition have not been well studied in the field. To evaluate changes in bacterial and fungal abundance associated with the earthworm invasion we quantified bacterial and fungal biomass by microscopic counts in paired earthworm-invaded (earthworm) and earthworm-free (reference) plots in five forest stands in central New York (USA). Earthworms significantly increased the ratio of bacteria to fungi on an area basis (per m²), by more than two times in mid-summer and early autumn. While this effect was associated primarily with the lack of the fungal-dominated organic horizon in earthworm plots, a higher ratio of bacteria to fungi in the surface 5 cm mineral soil also contributed as it developed between spring and mid-summer. Earthworm reduction of fungal biomass was confirmed by substantially lower growth of fungal hyphae into mesh sand bags in earthworm compared to reference plots. Burrowing activity by the earthworm Lumbricus terrestris increased the ratio of bacteria to fungi over the short-term within earthworm plots, introducing small-scale spatial heterogeneity associated with burrows. Our study suggests that the exotic earthworm invasion in these northern hardwood forests markedly increased the ratio of bacteria to fungi by eliminating the fungal-rich organic horizon, and was associated localized increases in bacterial vs. fungal abundance in mineral soil, setting the stage for future research into linkages between the earthworm invasion, bacterial and fungal abundance, and ecosystem processes.     

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).     

Influence of nitrogen on cellulose and lignin mineralization in blackwater and redwater forested wetland soils

 Microcosms were used to determine the influence of N additions on active bacterial and active fungal biomass, cellulose degradation and lignin degradation at 5, 10 and 15 weeks in soils from blackwater and redwater wetlands in the northern Florida panhandle. Blackwater streams contain a high dissolved organic C concentration which imparts a dark color to the water and contain low concentrations of nutrients. Redwater streams contain high concentrations of suspended clays and inorganic nutrients, such as N and P, compared to blackwater streams. Active bacterial and fungal biomass was determined by direct microscopy; cellulose and lignin degradation were measured radiometrically. The experimental design was a randomized block. Treatments were: soil type (blackwater or redwater forested wetlands) and N additions (soils amended with the equivalent of 0, 200 or 400 kg N ha^–1 as NH₄NO₃). Redwater soils contained higher concentrations of C, total N, P, K, Ca, Mn, Fe, B and Zn than blackwater soils. After N addition and 15 weeks of incubation, the active bacterial biomass in redwater soils was lower than in blackwater soils; the active bacterial biomass in blackwater soils was lower when 400 kg N ha^–1, but not when 200 kg N ha^–1, was added. The active fungal biomass in blackwater soils was higher when 400 kg N ha^–1, but not when 200 kg N ha^–1, was added. The active fungal biomass in redwater wetland soils was lower when 200 kg N ha^–1, but not when 400 kg N ha^–1, was added. Cellulose and lignin degradation was higher in redwater than in blackwater soils. After 10 and 15 weeks of incubation, the addition of 200 or 400 kg N as NH₄NO₃ ha^–1 decreased cellulose and lignin degradation in both wetland soils to similar levels. This study indicated that the addition of N may slow organic matter degradation and nutrient mineralization, thereby creating deficiencies of other plant-essential nutrients in wetland forest soils. Received: 7 April 1999     

Influence of soil properties on microbial populations, activity and biomass in humid subtropical mountainous ecosystems of India

 Microbial populations, biomass, soil respiration and enzyme activities were determined in slightly acid organic soils of major mountainous humid subtropical terrestrial ecosystems, along a soil fertility gradient, in order to evaluate the influence of soil properties on microbial populations, activity and biomass and to understand the dynamics of the microbial biomass in degraded ecosystems and mature forest. Although the population of fungi was highest in the undisturbed forest (Sacred Grove), soil respiration was lowest in the 7-year-old regrowth and in natural grassland (approximately 373 μg g^–1 h^–1). Dehydrogenase and urease activities were high in "jhum" fallow, and among the forest stands they were highest in the 7-year-old regrowth. Microbial biomass C (MBC) depended mainly on the organic C status of the soil. The MBC values were generally higher in mature forest than in natural grassland, 1-year-old jhum fallow and the 4-year-old alder plantation. The MBC values obtained by the chloroform-fumigation-incubation technique (330–1656 μg g^–1) did not vary significantly from those obtained by the chloroform-fumigation-extraction technique (408–1684 μg g^–1), however, the values correlated positively (P<0.001). The enzyme activities, soil respiration, bacterial and fungal populations and microbial biomass was greatly influenced by several soil properties, particularly the levels of nutrients. The soil nutrient status, microbial populations, soil respiration and dehydrogenase activity were greater in Sacred Grove, while urease activity was greater in grassland. Received: 14 October 1998     

Evaluation of the selective respiratory inhibition method for measuring the ratio of fungal:bacterial activity in acid agricultural soils

A procedure for the measurement of the fungal and bacterial contribution to substrate-induced respiration was tested in three arable soils. Glucose and different amounts of cycloheximide (eukaryote inhibitor) and streptomycin sulfate (prokaryote inhibitor) were added to soil suspensions, and respiration (CO₂ evolution) was measured. Streptomycin sulfate concentrations from 10 to 120 mg ml^–1 soil solution caused a stable inhibition of respiration. Amounts of cycloheximide ranging from 5 to 35 mg ml^–1 showed an increasing inhibition. In a test with separate and combined addition of the antibiotics at maximum inhibitory concentrations, inhibition by streptomycin was completely overlapped by cycloheximide. This indicated non-target inhibition which may lead to overestimation of fungal respiration. Experiments with sterilized soils inoculated with either fungi or bacteria confirmed that streptomycin selectively inhibited bacteria. Cycloheximide, however, did not only inhibit fungal respiration already at 2 mg ml^–1, but also increasingly inhibited bacterial respiration at increasing concentrations. Only at less than 5 mg cycloheximide ml^–1 was the condition of selective fungal inhibition fulfilled. When 2 mg cycloheximide and 10 mg streptomycin sulfate ml^–1 were applied, the sum of the separate inhibitions almost equalled the combined inhibition by the mix of both inhibitors in field samples. This method yielded fungal:bacterial respiration ratios of 0.50 to 0.60, and confirmed the dominance of bacteria in Dutch arable soils. The ratios obtained by the selective inhibitors were not correlated with, and were higher than, ratios of fungal:bacterial biovolume (0.19 to 0.46) as determined by microscopy and image analysis. Similar measurements in a forest soil (A-horizon) raised doubts on the reliability of the fungal inhibition by cycloheximide in this soil. It is concluded that the separate:combined inhibition ratio should always be checked, and comparison with other approaches is recommended. Received: 17 September 1996     

Abundance, production and stabilization of microbial biomass under conventional and reduced tillage

Soil tillage practices affect the soil microbial community in various ways, with possible consequences for nitrogen (N) losses, plant growth and soil organic carbon (C) sequestration. As microbes affect soil organic matter (SOM) dynamics largely through their activity, their impact may not be deduced from biomass measurements alone. Moreover, residual microbial tissue is thought to facilitate SOM stabilization, and to provide a long term integrated measure of effects on the microorganisms. In this study, we therefore compared the effect of reduced (RT) and conventional tillage (CT) on the biomass, growth rate and residues of the major microbial decomposer groups fungi and bacteria. Soil samples were collected at two depths (0-5 cm and 5-20 cm) from plots in an Irish winter wheat field that were exposed to either conventional or shallow non-inversion tillage for 7 growing seasons. Total soil fungal and bacterial biomasses were estimated using epifluorescence microscopy. To separate between biomass of saprophytic fungi and arbuscular mycorrhizae, samples were analyzed for ergosterol and phospholipid fatty acid (PLFA) biomarkers. Growth rates of saprophytic fungi were determined by [¹⁴C]acetate-in-ergosterol incorporation, whereas bacterial growth rates were determined by the incorporation of ³H-leucine in bacterial proteins. Finally, soil contents of fungal and bacterial residues were estimated by quantifying microbial derived amino sugars. Reduced tillage increased the total biomass of both bacteria and fungi in the 0-5 cm soil layer to a similar extent. Both ergosterol and PLFA analyses indicated that RT increased biomass of saprophytic fungi in the 0-5 cm soil layer. In contrast, RT increased the biomass of arbuscular mycorrhizae as well as its contribution to the total fungal biomass across the whole plough layer. Growth rates of both saprotrophic fungi and bacteria on the other hand were not affected by soil tillage, possibly indicating a decreased turnover rate of soil microbial biomass under RT. Moreover, RT did not affect the proportion of microbial residues that were derived from fungi. In summary, our results suggest that RT can promote soil C storage without increasing the role of saprophytic fungi in SOM dynamics relative to that of bacteria.     

Microbial colonisation of roots as a function of plant species

Fifteen plants species were grown in the greenhouse on the same soil and sampled at flowering to obtain rhizosphere soil and root material. In both fractions, the data on fungal and bacterial tissue obtained by amino sugar analysis were compared with the total microbial biomass based on fumigation-extraction and ergosterol data. The available literature on glucosamine concentrations in fungi and on muramic acid concentrations in bacteria was reviewed to prove the possibility of generating conversion values for general use in root material. All microbial properties analysed revealed strong species-specific differences in microbial colonisation of plant roots. The root material contained considerable amounts of microbial biomass C and biomass N, reaching mean levels of 10.9 and 1.4 mg g⁻¹ dry weight, respectively. However, the majority of CHCl₃ labile C and N, i.e. 89 and 55% was root derived. The average amount of ergosterol was 13 μg g⁻¹ dry weight and varied between 0.0 for Phacelia roots and 45.5 μg g⁻¹ dry weight for Vicia roots. The ergosterol content in root material of mycorrhizal and non-mycorrhizal plant species did not differ significantly. Fungal glucosamine was converted to fungal C by multiplication by 9 giving a range of 7.1-25.9 mg g⁻¹ dry weight in the root material. Fungal C and ergosterol were significantly correlated. Bacterial C was calculated by multiplying muramic acid by 45 giving a range from 1.7 to 21.6 mg g⁻¹ dry weight in the root material. In the root material of the 15 plant species, the ratio of fungal C-to-bacterial C ranged from 1.0 in mycorrhizal Trifolium roots to 9.5 in non-mycorrhizal Lupinus roots and it was on average 3.1. These figures mean that the microbial tissue in the root material consists on average of 76% fungal C and 24% bacterial C. The differences in microbial colonisation of the roots were reflected by differences in microbial indices found in the rhizosphere soil, most strongly for microbial biomass C and ergosterol, but to some extent also for glucosamine and muramic acid.     

Influence of nitrogen on atrazine and 2, 4 dichlorophenoxyacetic acid mineralization in blackwater and redwater forested wetland soils

 Microcosms were used to determine the influence of N additions on active bacterial and fungal biomass, atrazine and dichlorophenoxyacetic acid (2,4-D) mineralization at 5, 10 and 15 weeks in soils from blackwater and redwater wetland forest ecosystems in the northern Florida Panhandle. Active bacterial and fungal biomass was determined by staining techniques combined with direct microscopy. Atrazine and 2,4-D mineralization were measured radiometrically. Treatments were: soil type, (blackwater or redwater forested wetland soils) and N additions (soils amended with the equivalent of 0, 200 or 400 kg N ha^–1 as NH₄NO₃). Redwater soils contained higher concentrations of C, total N, P, K, Ca, Mn, Fe, B and Zn than blackwater soils. After N addition and 15 weeks of incubation, active bacterial biomass in redwater soils was lower when N was added. Active bacterial biomass in blackwater soils was lower when 400 kg N ha^–1, but not when 200 kg N ha^–1, was added. Active fungal biomass in blackwater soils was higher when 400 kg N ha^–1, but not when 200 kg N ha^–1, was added. Active fungal biomass in redwater soils was lower when 200 kg N ha^–1, but not when 400 kg N ha^–1, was added. After 15 weeks of incubation 2,4-D degradation was higher in redwater wetland soils than in blackwater soils. After 10 and 15 weeks of incubation the addition of 200 or 400 kg N ha^–1 decreased both atrazine and 2,4-D degradation in redwater soils. The addition of 400 kg N ha^–1 decreased 2,4-D degradation but not atrazine degradation in blackwater soils after 10 and 15 weeks of incubation. High concentrations of N in surface runoff and groundwater resulting from agricultural operations may have resulted in the accumulation of N in many wetland soils. Large amounts of N accumulating in wetlands may decrease mineralization of toxic agricultural pesticides. Received: 26 June 1998     

The fungal and bacterial biomass (selective inhibition) and the production of CO<Subscript>2</Subscript> and N<Subscript>2</Subscript>O by soddy-podzolic soils of postagrogenic biogeocenoses

In the humus horizon of soddy-podzolic soils of postagrogenic cenoses and primary forests, the contributions of the fungi and bacteria were determined by the selective inhibition of the substrate-induced respiration (SIR) by antibiotics; the basal (microbial) respiration and the net-produced nitrous oxide (N₂O) were also determined. The procedure of the SIR separation using antibiotics (cycloheximide and streptomycin) into the fungal and bacterial components was optimized. It was shown that the fungi: bacteria ratio was 1.58, 2.04, 1.55, 1.39, 2.09, and 1.86 for the cropland, fallow, and different-aged forests (20, 45, 90, and 450 years), respectively. The fungal and bacterial production of CO₂ in the primary forest soil was higher than in the cropland by 6.3 and 11.4 times, respectively. The production of N₂O in the soils of the primary and secondary (90-year-old) forests (3 and 7 ng N-N₂O/g soil per hour, respectively) was 2–13 times lower than in the postagrogenic cenoses, where low values were also found for the microbial biomass carbon (C_mic), its components (the C_mic-bacteria and C_mic-fungi), and the portion of C_mic in the organic carbon of the soil. A conclusion was drawn about the misbalance of the microbial processes in the overgrown cropland accompanied by the increased production of N₂O by the soil during its enrichment with an organic substrate (glucose).     

Comparison of chloroform fumigation-extraction, phospholipid fatty acid, and DNA methods to determine microbial biomass in forest humus

Techniques developed to measure microbial biomass in mineral soils may not give reliable results in humus. We evaluated the relationships between three techniques to estimate microbial biomass in forest humus: chloroform fumigation-extraction (CFE), total extractable phospholipid fatty acids (PLFA), and extractable DNA. There was a good relationship between PLFA and CFE (R²=0.96), with a slope slightly different from that previously reported for mineral soils (1 nmol PLFA corresponded to a flush of 3.2 μg C released by fumigation in humus cf. 2.4 μg C in mineral soil). There was no relationship between DNA concentration and the other two measurements of microbial biomass. This may be due, in part, to the high fungal biomass in forest humus, as DNA concentration per unit biomass is much more variable for fungi than bacteria.     

Impact of land-use change on nitrogen mineralization in subalpine grasslands in the Southern Alps

 A field study was conducted to investigate the effect of abandonment of management on net N mineralization (NNM) in subalpine meadows. NNM, soil microbial biomass N (SMBN), fungal biomass and physicochemical characteristics (total C, total N, dissolved organic carbon (DOC) and pH) were determined in surface (0–10 cm) soil from May to October 1997 in an intensively managed and an abandoned meadow at 1770 m a.s.l.. The cumulative NNM was lower in the abandoned area and ranged from 150 to 373 and from 25 to 85 μg N g^–1 soil in the intensively managed and the abandoned areas, respectively. The total organic C increased in the abandoned area, while total N showed no difference between abandoned and managed meadow. SMBN showed no difference between the investigated sites, whereas ergosterol, a measure of fungal biomass, increased significantly at the abandoned site. The cumulative NNM was negatively correlated with total organic C, C : N ratio, DOC and ergosterol content, and positively correlated with the NH₄ ⁺-N content of soil. The decrease in NNM at the abandoned site may be explained by the lower availability of N in substrates characterized by a high C : N ratio which, together with a decrease in pH in the litter layer, may have increased fungal biomass. Received: 8 January 1999     

Quantitative assessment of the fungal contribution to microbial tissue in soil

The fungi-to-bacteria ratio in soil ecological concepts and its application to explain the effects of land use changes have gained increasing attention over the past decade. Four different main approaches for quantifying the fungal and bacterial contribution to microbial tissue can be distinguished: (1) microscopic methods, (2) selective inhibition, (3) specific cell membrane components and (4) specific cell wall components. In this review, the different methods were compared and we hypothesized that all these approaches result in similar values for the fungal and bacterial contribution to total microbial biomass, activity, and residues (dead microbial tissue) if these methods are evenly reliable for the estimation of fungal biomass. The fungal contribution to the microbial biomass or respiration varied widely between 2 and 95% in different data sets published over the past three decades. However, the majority of the literature data indicated that fungi dominated microbial biomass, respiration or non-biomass microbial residues, with mean percentages obtained by the different methodological approaches varying between 35 and 76% in different soil groups, i.e. arable, grassland, and forest soils and litter layers. Microscopic methods generally gave the lowest average values, especially in arable and grasslands soils. Very low ratios in fungal biomass C-to-ergosterol obtained by microscopic methods suggest a severe underestimation of fungal biomass by certain stains. Relatively consistent ratios of ergosterol to linoleic acid (18:2ω6,9) indicate that both cell membrane components are useful indicators for saprotrophic and ectomycorrhizal fungi. More quantitative information on the PLFA content of soil bacteria and the 16:1ω5 content of arbuscular mycorrhizal fungi is urgently required to fully exploit the great potential of PLFA measurements. The most consistent results have been obtained from the analysis of fungal glucosamine and bacterial muramic acid in microbial residues. Component-specific δ¹³C analyses of PLFA and amino sugars are a promising prospect for the near future.     

Soil enzyme activities,microbial community composition and function after 47 years of continuous green manuring

Green manuring practices can influence soil microbial community composition and function and there is a need to investigate the influence compared with other types of organic amendment. This study reports long-term effects of green manure amendments on soil microbial properties, based on a field experiment started in 1956. In the experiment, various organic amendments, including green manure, have been applied at a rate of 4 t C ha⁻¹ every second year. Phospholipid fatty acid analysis (PLFA) indicated that the biomass of bacteria, fungi and total microbial biomass, but not arbuscular mycorrhizal (AM) fungi, generally increased due to green manuring compared with soils receiving no organic amendments. Some differences in abundance of different microbial groups were also found compared with other organic amendments (farmyard manure and sawdust) such as a higher fungal biomass and consequently a higher fungal/bacterial ratio compared with amendment with farmyard manure. The microbial community composition (PLFA profile) in the green manure treatment differed from the other treatments, but there was no effect on microbial substrate-utilization potential, determined using the Biolog EcoPlate. Protease and arylsulphatase activities in the green manure treatment were comparable to a mineral fertilized treatment receiving no additional C, whereas acid phosphatase activity increased. It can be concluded that green manuring had a beneficial impact on soil microbial properties, but differed in some aspects to other organic amendments which might be attributed to differences in quality of the amendments.     

Reciprocal transfer of carbon and nitrogen by decomposer fungi at the soil-litter interface

We have investigated whether decomposer fungi translocate litter-derived C into the underlying soil while simultaneously translocating soil-derived inorganic N up into the litter layer. We also located and quantified where the translocated C is deposited within the soil aggregate structure. When ¹³C-labeled wheat straw was decomposed on the surface of soil amended with ¹⁵N-labeled inorganic N, we found that C and N were reciprocally transferred by fungi, with a significant quantity (121-151 μg C g⁻¹ whole soil) of litter-derived C being deposited into newly formed macroaggregates (>250 μm sized aggregates). Fungal inhibition reduced fungal biomass and the bidirectional C and N flux by approximately 50%. The amount of litter-derived C found in macroaggregates was positively correlated with litter-associated fungal biomass. This fungal-mediated litter-to-soil C transfer, which to our knowledge has not been demonstrated before for saprophytic fungi, may represent an important mechanism by which litter C enters the soil and becomes stabilized as soil organic matter within the macroaggregate structure.     

Fungi,bacteria and protozoa in soil from four arable cropping systems

Summary The effects of four cropping systems on soil microorganisms were investigated during 3 years. The cropping systems were B0, barley without nitrogen fertilizers; B120, barley with 120 kg N ha^–1 year^–1; GL, grass ley receiving 200 kg N ha^–1 year^–1; and LL, lucerne ley without nitrogen fertilizer additions. At samplings in September during three consecutive years no differences were found between treatments. Total fungal lengths ranged between 0.7 and 2.0 × 10³ m and bacterial numbers between 3.5 and 7.2 × 10⁹ g^–1 dry wt. soil.Twenty samplings over 3 years in B120 and in GL indicated higher numbers of bacteria and protozoa during the growing season, except for periods with moisture stress. No clear seasonal trends were found for the fungi. When comparing mean values for the 20 samplings, the grass ley contained significantly (P < 0.05) higher numbers of amoebae. Means of the bacterial numbers and biomass, total and FDA-active hyphal lengths were also higher or equal (FDA-active hyphae) but not significantly so.Seventy-nine per cent of the bacterial biomass and 73% of the total fungal lengths were found in the top soil, where also 85% of the oxygen was consumed.     

Annual population dynamics and production ecology of Testacea (Protozoa,Rhizopoda) in an aspen woodland soil

The population studies of Testacea in an aspen woodland soil revealed 28 taxa of living Testacea, 14 of which were considered constant. The 14 species, while comprising 98 and 96% of the mean annual density and total annual production numbers respectively, accounted for only 80% of the mean annual biomass and 86% of the total annual production biomass. All 14 species had a peak in abundance in autumn, immediately or soon after leaf litter fall. While most species maintained small, active and reproducing populations over the winter period, some maintained higher than normal densities, had seasonal peaks in abundance and biomass, or higher than normal rates of production and turnover. All species had an increase in reproduction and usually in abundance also during the late winter-early spring period before spring thaw. The annual mean weekly density and biomass and the annual production totals tor numbers and biomass were highest in the H layer and lowest in the L. The F layer produced the highest numbers of generations per year and the highest annual mean weekly biomass turnover rate for most of the species. The majority of the periods of high intrinsic rate of natural increase, biomass production and turnover occurred during or after an increase, a decrease or a stabilization in soil moisture content. Mean annual biomass and total annual production were estimated as 0.72 and 206 g wet weight m^?2 respectively. The aspen woodland soil Testacea (in a mor humus) had a much higher production and number of generations per year than Testacea from mull and moder humus forms.Total annual ingestion, respiration losses and egestion losses for testate amoebae in an aspen woodland soil were calculated as 1377, 344 and 826 g wet weight of biomass m^?2, respectively. The annual secondary production of Testacea was about 250–300 times the standing crop of Testacea in all the soil layers. The dry weight of carbon respired per year by the Testacea was estimated as 16.2 g m^?2, which amounted to about 6% of the total carbon input.     

Catclaw (Mimosa buincifiera): a pest or a means to restore soil fertility in heavily eroded soil from the central highlands of Mexico?

 In the central highlands of Mexico, heavily eroded soils are often colonized by catclaw (Mimosa buincifiera): an N₂-fixing shrub. An experiment was carried out to investigate how this shrub affected characteristics of the soil and its biological functioning. Soil was sampled from outside and under the canopy of catclaw at three sites characterized by different degrees of erosion and an increase in plant density. The soil microbial biomass C, total amounts of bacteria, fungi, actinomycetes and free-living N₂-fixing micro-organisms were measured, while production of CO₂ and dynamics of nitrate (NO₃ ^–), nitrite (NO₂ ^–) and ammonium (NH₄ ⁺) were monitored in an aerobic incubation at 22±1  °C for 35 days. The C content was 1.6 times greater in the area with the largest density of plants and the least erosion (RECUP) compared with the site with the lowest density and greatest erosion (DEGR), while it was 1.2 times greater under the canopy of the catclaw than outside it (average of the three sites). The incorporation of N into the soil organic matter was greater under the canopy of the catclaw than outside it as the C:N ratio was on average 8.4 and 9. 1, respectively. The microbial biomass C, as a percentage of soil organic matter, was 1.5 times greater in the RECUP than in the DEGR site. Greatest total number of colony-forming bacteria and fungi (mean of organisms found under and outside the canopy) were found in the RECUP treatment and lowest in the DEGR treatment. Free-living N₂-fixing organisms and actinomycetes showed opposite trends. Greater total numbers of colony-forming bacteria, fungi, actinomycetes and free-living N₂-fixing organisms (mean of the three treatments) were found under the canopy of catclaw than outside of it, Production of CO₂ was 1.8 times greater in the RECUP than in the DEGR and 1.6 times greater under the canopy of catclaw than outside. Production of NO₃ ^– was 1.3 times greater in the RECUP than in the DEGR and 3.5 times greater under the canopy of catclaw than outside. There was no significant effect of location or canopy on NO₂ ^– and NH₄ ⁺ concentrations. It is concluded that the natural vegetation of catclaw increased microbial biomass and soil organic matter content under, but also outside its canopy, and preserved N better, releasing greater amounts of inorganic N upon mineralization. Catclaw can serve as a first colonizer of heavily eroded soil and be replaced by other vegetation, natural or crops, when fertility is restored. Received: 4 November 1999     

Evaluation of soil quality parameters in a tropical paddy soil amended with rice residues and tree litters

Laboratory and greenhouse experiments were conducted to study the effects of applications of rice residue and Pongamia pinnata and Azadirachta indica leaf litters on biochemical properties (extraction yield of humus, composition of humus, microbial biomass carbon, activities of urease and acid phosphatase) of a lowland rice soil under flooded conditions. Bulk soil sample collected from the Mandya paddy fields was used for the green house trials and the laboratory incubation studies. The organic materials were added at three rates – zero, 25.0 g carbon kg⁻¹ (2.5% C) and 50.0 g carbon kg⁻¹ dry soil (5.0% C). Results showed that tree leaf litter and rice residue at 5.0% C rate decreased instantaneous decay constant (k), there by retarded the rate of C mineralization. Carbon contents of HA increased with the rate of C added. Study of delta–log K values and C contents of humic acids revealed that greatest molecular weight of HA was in the pongamia litter treatment, followed by neem litter and rice residue. Grain and straw yields of rice crop in the pot culture study were statistically correlated to the soil quality parameters. Neem and pongamia tree litter incorporation at 2.5% C could be considered for improving soil health and crop yields of rice under flooded conditions; however, application at higher rates significantly (P ≤ 0.05) lowered total dry matter production in rice, despite favorable soil health parameters such as humic yields, microbial biomass – C content and acid phosphatase and urease activity. Among different soil health parameters, microbial quotient was found to be more sensitive indicator of decline in soil quality.     

Impact of inorganic nitrogen additions on microbes in biological soil crusts

Many studies have shown that changes in nitrogen (N) availability affect the diversity and composition of soil microbial community in a variety of terrestrial systems, but less is known about the responses of microbes specific to biological soil crusts (BSCs) to increasing N additions. After seven years of field experiment, the bacterial diversity in lichen-dominated crusts decreased linearly with increasing inorganic N additions (ambient N deposition; low N addition, 3.5 g N m⁻² y⁻¹; medium N addition, 7.0 g N m⁻² y⁻¹; high N addition, 14.0 g N m⁻² y⁻¹), whereas the fungal diversity exhibited a distinctive pattern, with the low N-added crust containing a higher diversity than the other crusts. Pyrosequencing data revealed that the bacterial community shifted to more Cyanobacteria with modest N additions (low N and medium N) and to more Actinobacteria and Proteobacteria and much less Cyanobacteria with excess N addition (high N). Our results suggest that soil pH, together with soil organic carbon (C), structures the bacterial communities with N additions. Among the fungal communities, the relative abundance of Ascomycota increased with modest N but decreased with excess N. However, increasing N additions favored Basidiomycota, which may be ascribed to increases in substrate availability with low lignin and high cellulose contents under elevated N conditions. Bacteria/fungi ratios were higher in the N-added samples than in the control, suggesting that the bacterial biomass tends to dominate over that of fungi in lichen-dominated crusts after N additions, which is especially evident in the excess N condition. Because bacteria and fungi are important components and important decomposers in BSCs, the alterations of the bacterial and fungal communities may have implications in the formation and persistence of BSCs and the cycling and storage of C in desert ecosystems.