Effects of agricultural biostimulants on soil microbial activity and nitrogen dynamics

We investigated the effects of two commercially available soil biostimulants, designated Z93 and W91, on key microbial and nutrient cycling processes in the soil, by conducting short-term (1 week) and longer-term (8 weeks) soil incubations in the laboratory. In the short-term soil incubations, the two compounds differed in their effects on microbial activity: Z93 was effective over a wide range, stimulating substrate-induced respiration (SIR) and dehydrogenase activity (DHA) at remarkably low concentrations (0.5–500 nl/g soil); W91 stimulated SIR at these concentrations, but also inhibited DHA. In longer-term soil incubations, we amended batches of soil with either finely-ground alfalfa leaves, wheat straw, or added no amendments, to alter patterns of soil nitrogen mineralization and immobilization. We treated these soils with Z93 and W91 at two concentrations (0.005 and 0.5 μl/g soil), and incubated them for up to 8 weeks. These extremely low doses of both Z93 and W91 influenced soil SIR, DHA, and cellulase activity significantly (P<0.05). Both compounds also influenced soil nitrogen dynamics significantly; the extent depending upon the quality of the organic amendments. In the alfalfa-amended soil there was a steep increase in NO₃-N concentration during the incubation due to the rapid mineralization of nitrogen-rich alfalfa material. However, in this soil, both Z93 and W91 reduced NO₃-N concentrations greatly after 56 days. In the straw-amended soil, mineral nitrogen concentrations were very low, probably due to rapid immobilization of nitrogen by microbial biomass. In this soil, treatment with both compounds decreased microbial biomass nitrogen and increased dissolved organic nitrogen (DON), relative to that in the controls. Our results suggest that the two biostimulants can stimulate both the breakdown and mineralization of soil organic materials, perhaps by selectively inhibiting or stimulating particular components of the microbial community, leading to lasting (8 weeks or longer) increases in soil nitrogen availability.     

Mineralization of low molecular weight carbon substrates in soil solution under laboratory and field conditions

A more detailed mechanistic understanding of how low molecular weight (MW) carbon (C) substrates are mineralized within the rhizosphere by soil microbial communities is crucial to accurately model terrestrial C fluxes. Currently, most experiments regarding soil C dynamics are conducted ex-situ (laboratory) and can fail to account for key variables (e.g. temperature and soil water content) which vary in-situ. In addition, ex-situ experiments are often highly invasive, e.g. severing root and mycorrhizal networks, changing the input and concentrations of low MW exudates within soil. The aim of this study was to directly compare the mineralization rates of 31 common low MW C substrates under ex- and in-situ conditions. In addition, we also assessed the inter-annual field variability of substrate mineralization rates. We added trace concentrations of 31 individual ¹⁴C-labelled common low MW C substrates into the top soil of an agricultural grassland and monitored the mineralization rates by capturing ¹⁴CO₂ evolved from the soil over 7 d. Our results showed that the contribution of low MW C components to soil respiration was highly reproducible between parallel studies performed either in-situ or ex-situ. We also found that differences in the mineralization of individual compounds were more variable inter-annually in the field than between the laboratory and the field. Our results suggest that laboratory-based C mineralization data can be used to reliably parameterize C models but that multiple experimental measurements should be made over time to reduce uncertainty in model parameter estimation.     

Tree species effects on soil enzyme activities through effects on soil physicochemical and microbial properties in a tropical montane forest on Mt. Kinabalu, Borneo

Tree species influence on the soil mineralization process can regulate overall nutrient cycling in a forest ecosystem, which may occur through their effects on substrate quality, soil physicochemical properties and soil microbial community. We investigated tree species effects on soil enzyme activities in a tropical montane forest on Mt. Kinabalu, Borneo. Specifically, we analyzed C- and P-degrading enzyme activities, as well as the relationships among the enzyme activities, soil physicochemical properties, substrate quality (C, N, and P concentrations), and microbial composition in the top 5 cm soils beneath conifers (Dacrycarpus imbricatus and Dacrydium gracilis) and broadleaves (Lithocarpus clementianus, Palaquium rioence, and Tristaniopsis clementis). Activities of acid phosphatase and β-d-glucosidase were significantly different among the tree species. Soil moisture, total C and N content and microbial lipid abundance (a proxy for microbial composition) could influence the enzyme activities although the relative contributions of microbial composition to the enzyme activities might be smaller. A higher acid phosphatase activity beneath Dacrydium than those beneath the other tree species can compensate for a lower concentration of P in available fractions beneath Dacrydium. This localized mineralization activity could subsequently influence soil nutrient availability in this forest.     

Microbial response time to sugar and amino acid additions to soil

Soil microbial respiration is derived predominantly from the turnover of carbohydrates and proteins in soil. In most agricultural ecosystems, these C compounds enter soil mainly from rhizodeposition (root exudation and turnover). Our aim was to determine how long it takes for the microbial population to reach their maximum mineralization potential after the addition of low-molecular-weight (MW) rhizodeposits to the soil. We added sugar in the form of glucose and amino acids in the form of glycine to an arable, grazed grassland, Eucalyptus forest and boreal forest soil and monitored CO₂ efflux over a 6-h period. Artificial rainwater amended (zero C addition) or unamended soils were used as controls. The Michaelis-Menten substrate utilization profiles showed vastly different patterns of microbial mineralization capacity and substrate affinity between the soils. However, in all soils we showed that activation of the soil microbial community to C addition occurred almost instantaneously (?60 s) with the average time taken to reach half maximal CO₂ production being 14±8 min for glucose and 10±8 min for glycine. After reaching their maximal mineralization potential, the rate of CO₂ evolution remained constant for the remainder of the experiment. Our results showed that while substrate uptake and mineralization within the soil microbial biomass was activated quickly, subsequent adaptation and upregulation of its C processing capacity did not occur at least in the short term. The fast rate of microbial activation and substrate use we partially attribute to the large degree of functional redundancy that exists within the soil microbial community for processing rhizodeposits.     

Free amino sugar reactions in soil in relation to soil carbon and nitrogen cycling

Amino sugars represent a major constituent of microbial cell walls and hydrolyzed soil organic matter. Despite their potential importance in soil nitrogen cycling, comparatively little is known about their dynamics in soil. The aim of this study was therefore to quantify the behaviour of glucosamine in two contrasting grassland soil profiles. Our results show that both free amino sugars and amino acids represented only a small proportion of dissolved organic N and C pool in soil. Based upon our findings we hypothesize that the low concentrations of free amino sugars found in soils is due to rapid removal from the soil solution rather than slow rates of production. Further, we showed that glucosamine removal from solution was a predominantly biotic process and that its half-life in soil solution ranged from 1 to 3 h. The rates of turnover were similar to those of glucose at low substrate concentrations, however, at higher glucosamine concentrations its microbial use was much less than for glucose. We hypothesized that this was due to the lack of expression of a low affinity transport systems in the microbial community. Glucosamine was only weakly sorbed to the soil's solid phase (K_d=6.4±1.0) and our results suggest that this did not limit its bioavailability in soil. Here we showed that glucosamine addition to soil resulted in rapid N mineralization and subsequent NO₃⁻ production. In contrast to some previous reports, our results suggest that free amino sugars turn over rapidly in soil and provide a suitable substrate for both microbial respiration and new biomass formation.     

The carbon we do not see—the impact of low molecular weight compounds on carbon dynamics and respiration in forest soils: a review

Dissolved organic matter (DOM), typically quantified as dissolved organic carbon (DOC), has been hypothesized to play many roles in pedogenesis and soil biogeochemical cycles, however, most research to date concerning forest soils has focussed on the high molecular weight (HMW) components of this DOM. This review aims to assess the role of low molecular weight (LMW) DOM compounds in the C dynamics of temperate and boreal forest soils focussing in particular on organic acids, amino acids and sugars. The current knowledge of concentrations, mineralization kinetics and production rates and sources in soil are summarised. We conclude that although these LMW compounds are typically maintained at very low concentrations in the soil solution (<50 μM), the flux through this pool is extremely rapid (mean residence time 1-10 h) due to continued microbial removal. Due to this rapid flux through the soil solution pool and mineralization to CO₂, we calculate that the turnover of these LMW compounds may contribute substantially to the total CO₂ efflux from the soil. Moreover, the production rates of these soluble transitory compounds could exceed HMW DOM production. The possible impact of climate change on the behaviour of LMW compounds in soil is also discussed.     

Lumbricid macrofauna alter atrazine mineralization and sorption in a silt loam soil

Atrazine is a widely used herbicide and is often a contaminant in terrestrial and freshwater ecosystems. It is uncertain, however, how the activity of soil macrofauna affects atrazine fate and transport. Therefore, we investigated whether earthworms enhance atrazine biodegradation by stimulating herbicide degrading soil microflora, or if they increase atrazine persistence by facilitating herbicide sorption. Short (43 d) and medium term (86 d) effects of the earthworms Lumbricus terrestris and Aporrectodea caliginosa on mineralization, distribution, and sorption of U-ring-¹⁴C atrazine and on soil C mineralization was quantified in packed-soil microcosms using silt loam soil. A priming effect (stimulation of soil C mineralization) caused by atrazine supply was shown that likely lowered the earthworm net effect on soil C mineralization in atrazine-treated soil microcosms. Although earthworms significantly increased soil microbial activity, they reduced atrazine mineralization to ¹⁴CO₂-C from15.2 to 11.7% at 86 d. Earthworms facilitated formation of non-extractable atrazine residues within C-rich soil microsites that they created by burrowing and ingesting soil and organic matter. Atrazine sorption was highest in their gut contents and higher in casts than in burrow linings. Also, gut contents exhibited the highest formation of bound atrazine residues (non-extractable atrazine). Earthworms also promoted a deeper and patchier distribution of atrazine in the soil. This contributed to greater leaching losses of atrazine in microcosms amended with earthworms (3%) than in earthworm-free microcosms (0.003%), although these differences were not significant due to high variability in transport from earthworm-amended microcosms. Our results indicated that earthworms, mainly by casting activity, facilitated atrazine sorption, which increased atrazine persistence. As a consequence, this effect overrode any increase in atrazine biodegradation due to stimulation of microbial activity by earthworms. It is concluded that the affect of earthworms of atrazine mineralization is time-dependent, mineralization being slightly enhanced in the short term and subsequently reduced in the medium term.     

Degradation and binding of atrazine in surface and subsurface soils

Understanding the dissipation rates of chemicals in unsaturated and saturated zones of subsurface soils will help determine if reductions of concentrations to acceptable levels will occur. Chemical properties and microbial biomass and activity were determined for the surface (0-15 cm), lower root (50-105 cm), and vadose (175-220 cm) zones in a Huntington silty clay loam (Fluventic Hapludoll) collected from an agricultural field near Piketon, OH. The rates of sorption, mineralization, and transformation (formation of bound residues and metabolites) of atrazine were determined. Microbial activity was estimated from the mineralization of (14)C-benzoate. We observed decreased levels of nutrients (total organic carbon, N, and P) and microbial biomass with depth, while activity as measured with benzoate metabolism was higher in the vadose zone than in either the surface or the root zones. Sorption coefficients (K(f)) declined from 8.17 in the surface to 3.31 in the vadose zone. Sorption was positively correlated with organic C content. Rates of atrazine mineralization and bound residues formation were, respectively, 12-2.3-fold lower in the vadose than in the surface soil. Estimated half-lives of atrazine ranged from 77 to 101 days in the surface soil, but increased to over 900 days in the subsurface soils. The decreased dissipation of atrazine with increasing depth in the profile is the result of decreased microbial activity toward atrazine, measured either as total biomass or as populations of atrazine-degrading microorganisms. The combination of reduced dissipation and low sorption indicates that there is potential for atrazine movement in the subsurface soils.     

Influence of soil aging on sorption and bioavailability of simazine

Characterization of pesticide bioavailability, particularly in aged soils, is of continued interest because this information is necessary for environmental risk assessment. However, pesticide bioavailability in aged soils has been characterized by a variety of methods with limited success, due in part to methodological limitations. The objective of this study was to use solvent extraction methods to correlate simazine residue bioavailability in aged soils to simazine mineralization using a simazine-mineralizing bacterium. Soils from Brazil, Hawaii, and the midwestern United States were treated with UL-ring-labeled [14C]simazine and incubated for up to 8 weeks. At the end of each incubation period, soils were either incubated further, extracted with 0.01 M CaCl2, or extracted with aqueous methanol (80:20 v/v methanol/water). In a parallel experiment, after each incubation period, soils were inoculated with the bacterium Pseudomonas sp. strain ADP, which is capable of rapidly mineralizing simazine, and 14CO2 was determined. The inoculated soil samples were then extracted with 0.01 N CaCl2 and with aqueous methanol. This allowed for the evaluation of the bioavailability of aged simazine residues, without the contribution of simazine desorption from soil. Results of these studies indicated that simazine sorption to soil increased with aging and that amounts of simazine in aged soils extracted by 0.01 M CaCl2 and aqueous methanol were highly correlated to amounts of simazine mineralized by Pseudomonas sp. strain ADP. Consequently, 0.01 M CaCl2/methanol-extractable simazine in aged soils can be used to estimate bioavailable residues. This technique may be useful in determining the bioavailability of other s-triazine compounds in soils.     

Adenylate energy charge measurements in soil

Adenosine 5'-triphosphate (ATP), adenosine 5'-diphosphatc (ADP) and adenosine 5'-monophosphate (AMP) were extracted from soil with either a solution of trichloroacetic acid, paraquat and phosphate (TCA reagent) or a mixture of chloroform, sodium hydrogen carbonate, phosphate and adenosine (NaHCO₃ reagent). Standard enzymic procedures were used to convert ADP and AMP to ATP, which was measured by the fire-fly luciferin-luciferase system. The measured quantities of nucleotides were corrected for incomplete extraction using the percentage recoveries of added ATP, ADP and AMP. The adenylate energy charge ratio (AEC) was calculated from the formula AEC = ([ATP] + 0.5 [ADP])/([ATP] + [ADP] + [AMP]).Measurements were made on a grassland soil, following a conditioning incubation at 15°C and 50% WHC for 7 days. Additional measurements were made on the same soil after a further 50- or 100-day incubation at 25°C and 50% WHC, with or without an amendment of 1100 μg ryegrass Cg⁻¹ soil, added at the end of the conditioning incubation. Biomass-ATP concentration, measured in TCA extracts, changed little, even on prolonged incubation, and was maintained at a level comparable to that observed in earlier work (about 10 p mol ATP g⁻¹ biomass C). AEC values in TCA soil extracts were high (0.8–0.9) for all soil treatments and independent of substrate addition or length of incubation.In contrast, AEC was low (0.4) in fresh soil extracted with NaHCO₃ reagent, but increased to 0.6 when ryegrass was incubated with the soil for 50 days. Although the total adenine nucleotide pool (i.e. [ATP] + [ADP] + [AMP]) was similar as measured in NaHCO₃ and in TCA soil extracts, both energy charge and ATP content were lower in the NaHCO₃ extracts. It was therefore concluded that the main reason for the lower AECs observed with the NaHCO₃ reagent was that microbial ATPases were still active during extraction and caused appreciable hydrolysis of microbial ATP to ADP and AMP. In contrast, the TCA reagent rapidly inactivates ATPases and is therefore preferable for extracting adenine nucleotides from soil.The results indicate that the soil microbial biomass, although a mainly dormant population, maintains both AEC and ATP at levels characteristic of exponentially growing organisms in vitro, even during prolonged incubation without fresh substrate. It was also concluded that roots make a negligible contribution to total ATP extracted from fresh sieved soil.     

Einfluß variierter Temperatur und Feuchte auf mikrobielle Aktivitäten im Boden unter Laborbedingungen

Influence of varied soil temperature and moisture on microbial activities under laboratory conditions Under laboratory conditions the influence of temperature (10°C, 20°C, fluctuation from 5° to 30°C within 12 h with additional freezing for 3 days) and soil moisture (30%, 60% w.h.c., remoistening to 60% for 1 week) on several microbial activities was investigated. The biomass-related, glucose-induced short-term respiration and the dehydrogenase activity (TTC reduction) were higher at 10°C in most cases as compared to 20°C. Independent of freezing fluctuating temperature caused the lowest activities. The nitrogen mineralization (including nitrification), however, was affected in the opposite way. No marked influences were observed with β-glucosidase, arylsulfatase, and alkaline phosphatase. In the sandy loam nearly no effects of the soil moisture occurred and in the loamy sand especially the dehydrogenase activity was higher at 30% w.h.c., whereas the nitrogen mineralization was lower. From the results it can be concluded, that ecological conditions favouring mineralization without substrate addition may even reduce microbial biomass by decomposition.     

Influence of Cry1Ac toxin on mineralization and bioavailability of glyphosate in soil

The impact of transgenic plants containing Bacillus thuringiensis (Bt) toxin on soil processes has received recent attention. In these studies, we examined the influence of the lepidopterean Bt Cry1Ac toxin on mineralization and bioavailability of the herbicide glyphosate in two different soils. The addition of 0.25-1.0 microg g(-1) soil of purified Cry1Ac toxin did not significantly affect glyphosate mineralization and sorption in either a sandy loam or a sandy soil. In contrast, extractable glyphosate decreased over the 28 day incubation period in both soils. Our findings suggest that the reduction in the bioavailability of glyphosate was not influenced by the presence of Cry1Ac toxin but rather the results of aging or sorption processes. Results from this investigation suggest that the presence of moderate concentrations of Bt-derived Cry1Ac toxin would have no appreciable impact on processes controlling the fate of glyphosate in soils.     

Root exudate effects on the bacterial communities, CO2 evolution, nitrogen transformations and ATP content of rhizosphere and bulk soils

The ATP content, soil respiration, bacterial community composition, and gross N mineralization and immobilization rates were monitored under laboratory condition at 25 °C for 28 d in a model system where low molecular weight root exudates (glucose and oxalic acid) were released by a filter placed on the surface of a forest soil also treated with ¹⁵N, so as to simulate rhizosphere conditions. Periodically, the soil was sampled from two layers, 0-2 and 6-14 mm below the filter's surface, which were indicated as rhizosphere and bulk soils, respectively. The isotope dilution technique was used to determine the effect of these low molecular weight organic compounds (LMWOCs) on gross N mineralization and immobilization rates. From 0 to 3 d both glucose and oxalic acid amended soils showed a rapid evolution of CO₂, more pronunced in the latter treatment together with a decrease in the amount of mineral N of the rhizosphere soil, probably due to N immobilization. Nevertheless, these changes were accompanied by a very small increase in the net ATP content probably because the low C application rate stimulated microbial activity but microbial growth only slightly. A positive ‘priming effect’ probably developed in the oxalic acid amended soil but not in the glucose amended soil. Gross N mineralization and immobilization rates were only observed in the rhizosphere soil, probably due to the greater C and N concentrations and microbial activity, and were a little higher in both amended soils than in the control soil, only between 1 and 7 d. Both glucose and oxalic acid influenced the bacterial communities of the rhizosphere soil, as new bands in the DGGE profiles appeared at 3 and 7 d. Glucose induced lower changes in the bacterial community than oxalic acid, presumably because the former stimulated a larger proportion of soil microorganisms whereas the latter was decomposed by specialized microorganisms. Peaks of net daily soil respiration and net ATP content and the appearence of new dominant bacterial populations were shifted in time, probably because there was less ATP synthesis and DGGE patterns changed after complete substrate mineralization.     

4种湿地植物根系泌氧由根诱导的pH,Eh和Fe(II)变化和根际土壤中铅和锌的组分研究

A rhizobox experiment was conducted to compare iron (Fe) oxidation and changes of pH, redox potential (Eh) and fractions of zinc (Zn) and lead (Pb) in rhizosphere and non-rhizosphere soils of four emergent-rooted wetland plants (Echinodorus macrophyllus, Eleocharis geniculata, Hydrocotyle vulgaris and Veronica serpyllifolia) with different radial oxygen loss (ROL) from roots. The results indicated that all these wetland plants decreased pH and concentration of Fe(Ⅱ) but increased the Eh in the rhizosphere soils. Pb and Zn were transformed from unstable fractions to more stable fractions in the rhizosphere soils, so decreasing their potential metal mobility factors (MF). Among the four plants, E. macrophyllus, with the highest ROL and root biomass, possessed the greatest ability in formation of Fe plaque and in the reduction of heavy metal MFs in the rhizosphere soil. Wetland plants, with higher ROLs and root biomass, may thus be effective in decreasing potential long-term heavy metal bioavailabilities.     

LONG‐TERM LAND USE INFLUENCES SOIL MICROBIAL BIOMASS P AND S,PHOSPHATASE AND ARYLSULFATASE ACTIVITIES,AND S MINERALIZATION IN A BRAZILIAN OXISOL

Land use choices differentially affect soil physical and biological properties. Tillage choices in particular affect soil erosion, the retention of soil organic matter, and the biological activity that organic matter supports. The present study evaluated the consequences of different cropping and tillage systems (undisturbed forest, coffee plantation, conventional, and no‐tillage row cropping) for soil microbial indicators and sulfur mineralization after 24 years of cropping on an Oxisol (Typic Haplorthox) in an experimental area at Londrina, Brazil. Soil samples were taken at 0–5, 5–10, and 10–20 cm depths and evaluated for microbial biomass P and S, S mineralization, and phosphatase and arylsulfatase activities. Land use affected microbial biomass P and S, and enzyme activity at all depths studied. The cultivated sites had lower values of microbial activity than the undisturbed forested site. Although the coffee site was not tilled and had high organic carbon content, there was low microbial activity, probably due to higher soil acidity and Al content. The estimates of pool stock for microbial P and annual P flux through the soil microbial biomass suggest that these pools are large enough to significantly affect plant nutrient availability. The greater microbial biomass and activity under forested and no‐tillage sites may be attributed, at least partially, to higher organic matter content. The soil microbial variables examined proved to be strong indicators of soil sustainability. Copyright © 2013 John Wiley & Sons, Ltd.     

Organic matter, microbial biomass and enzyme activity of soils under different crop rotations in the tropics

Soil organic matter level, soil microbial biomass C, ninhydrin-N, C mineralization, and dehydrogenase and alkaline phosphatase activity were studied in soils under different crop rotations for 6 years. Inclusion of a green manure crop of Sesbania aculeata in the rotation improved soil organic matter status and led to an increase in soil microbial biomass, soil enzyme activity and soil respiratory activity. Microbial biomass C increased from 192 mg kg^–1 soil in a pearl millet-wheat-fallow rotation to 256 mg kg^–1 soil in a pearl millet-wheat-green manure rotation. Inclusion of an oilseed crop such as sunflower or mustard led to a decrease in soil microbial biomass, C mineralization and soil enzyme activity. There was a good correlation between microbial biomass C, ninhydrin-N and dehydrogenase activity. The alkaline phosphatase activity of the soil under different crop rotations was little affected. The results indicate the green manuring improved the organic matter status of the soil and soil microbial activity vital for the nutrient turnover and long-term productivity of the soil. Received: 7 January 1996     

Variation of four phosphorus properties in woodland soils

Variation in total, organic and available-P contents and phosphatase activity of P-deficient soils of some English Lake District woodlands of differing vegetative composition were examined in relation to individual woodlands, two depths in the soil profile, mull, moder and mor humus types, and different times of the year. Depth in the soil profile was a more important source of variation in the P properties than different woodlands. Soils in individual woodlands differed in their degree of variability in the four P properties. Available P contents and phosphatase activities were more variable than total and organic P contents. Available-P and organic-P contents and phosphatase activity showed seasonal variation. Seasonal variation in available-P was almost as great as differences in available-P between woodlands. Total and organic-P contents showed similar patterns of variation with respect to individual woodlands, humus type and soil depth. Differences in degree of variation within woodlands and differences in degree and pattern of variation of the four P-properties may need to be taken into account in soil sampling programmes of studies comparing soils under differing vegetation regimes.Different interpretations of the variation in the soil-P properties were obtained by expressing the data respectively in terms of soil weight (g^?1 soil) or soil volume (cm^?1 soil), due to marked variation in bulk-densities of the woodland soils. It is suggested that where soils vary in bulk-density, soil data should be expressed in terms of soil volume.The P-deficiency of the woodland soils is probably associated with the relatively low total P content per unit volume of soil and the high proportion of it which is organically bound.     

Responses of extracellular enzymes to simple and complex nutrient inputs

Soil microbes produce extracellular enzymes that mineralize organic matter and release carbon and nutrients in forms that can be assimilated. Economic theories of microbial metabolism predict that enzyme production should increase when simple nutrients are scarce and complex nutrients are abundant; however, resource limitation could also constrain enzyme production. We tested these hypotheses by monitoring enzyme activities and nutrient pools in soil incubations with added simple and complex nutrient compounds. Over 28 days of incubation, we found that an enzyme's activity increased when its target nutrient was present in complex but not simple form, and carbon and nitrogen were available. β-Glucosidase and acid phosphatase activities also increased in treatments where only carbon and nitrogen were added. Glycine aminopeptidase and acid phosphatase activities declined in response to ammonium and phosphate additions, respectively. In some cases, mineralization responses paralleled changes in enzyme activity—for example, β-glucosidase activity increased and respiration was 5-fold greater in soil incubations with added cellulose, ammonium, and phosphate. However, a doubling of acid phosphatase activity in response to collagen addition was not associated with any changes in phosphorus mineralization. Our results indicate that microbes produce enzymes according to ‘economic rules’, but a substantial pool of mineral stabilized or constitutive enzymes mediates this response. Enzyme allocation patterns reflect microbial nutrient demands and may allow microbes to acquire limiting nutrients from complex substrates available in the soil.     

Changes in enzyme activities and soil microbial community composition along carbon and nutrient gradients at the Franz Josef chronosequence, New Zealand

During primary succession, the abundance of carbon (C) and nitrogen (N) in soil increases, while phosphorus (P) declines. These changes in nutrient concentrations in organic matter are likely to play an important role in controlling enzyme-mediated nutrient mineralization. We examined how enzyme activity and efficiency changed with successional time in organic and mineral soils taken from the 120 000-year-old Franz Josef soil development sequence, New Zealand, and the relationship between enzyme activity and efficiency and soil nutrient concentrations. We found that the activity of enzymes involved in P mineralization increased with site age across the Franz Josef chronosequence, while the activity of enzymes regulating C and N mineralization declined in organic but not mineral soil. Sulfatase activity peaked at an intermediate-aged site, possibly indicating a transient period of S limitation. The activity of phosphatase enzymes was negatively correlated with the concentration of P in the soil, whereas activity of C-, N- and S-hydrolyzing enzymes was not strongly dependent on nutrient concentrations. When assessed as efficiency (activity per unit microbial biomass), there were strong patterns of increasing efficiency of P-, and decreasing efficiency of C- and N-hydrolyzing enzymes with site age. We suggest that activity patterns for C-, N- and S-hydrolyzing enzymes were obscured by simultaneous and opposing changes in enzyme efficiency and microbial biomass. In mineral soil, efficiency of enzymes was negatively correlated with soil nutrient availability. In contrast, in organic soil, efficiency of C-, N- and S-hydrolyzing enzymes was positively correlated with soil P, while efficiency of P-hydrolyzing enzymes was negatively correlated with soil P. The increase in efficiency of P-hydrolyzing enzymes, and decrease in efficiency of C-, N- and S-hydrolyzing enzymes with site age was accompanied by a shift in microbial community composition towards higher relative abundances of fungi. Changes in enzyme efficiency with site age are likely to be due to both constitutive differences in enzyme production, and down-regulation of enzyme expression.     

Criteria for measurement of microbial growth and activity in soil

Changes in CO₂ evolution, phosphatase and urease activity and ATP contents were related to bacterial and fungal biomass determined microscopically during glucose mineralization at different concentrations of mineral nutrients. Similar results were obtained in a sandy loam and a clay soil except that in the clay the increase in microbial and enzyme activities were delayed. Higher initial rates of CO₂ evolution were noted after the addition of P to a glucose and N amended soil at C:P ratios greater than 30:1. Increases in phosphatase activity coincided with increases in bacterial and fungal populations only in treatments without inorganic P. Peak rates of CO₂ evolution preceded biomass production by 18–24 h, therefore, CO₂ evolution rates did not show a correlation on normal regression analysis with biomass. Soil ATP content was influenced by P concentrations and soil type. ATP was therefore not a specific indicator of biomass in the detailed studies where P concentrations and sequential growth of bacteria and fungi were major factors. Soil urease increased with bacterial and fungal populations. It did not respond to P other than through microbial biomass and was highly correlated with microbial biomass. The results show that no one measurement of microbial biomass or activity is sufficient to interpret microbial growth in the soil system. Each of the criteria measured were sensitive to specific conditions affecting biomass and activity.