Soil solution and extractable soil nitrogen response to climate change in two boreal forest ecosystems

Several studies show that increases in soil temperature result in higher N mineralization rates in soils. It is, however, unclear if additional N is taken up by the vegetation or accumulates in the soil. To address this question two small, forested catchments in southern Norway were experimentally manipulated by increasing air temperature (+3°C in summer to +5°C in winter) and CO₂ concentrations (+200 ppmv) in one catchment (CO₂T-T) and soil temperature (+3°C in summer to +5°C in winter) using heating cables in a second catchment (T-T). During the first treatment year, the climate treatments caused significant increases in soil extractable NH₄ under Vaccinium in CO₂T-T. In the second treatment year extractable NH₄ in CO₂T-T and NO₃ in T-T significantly increased. Soil solution NH₄ concentrations did not follow patterns in extractable NH₄ but changes in soil NO₃ pools were reflected by changes in dissolved NO₃. The anomalous behavior of soil solution NH₄ compared to NO₃ was most likely due to the higher NH₄ adsorption capacity of the soil. The data from this study showed that after 2 years of treatment soil inorganic N pools increased indicating that increases in mineralization, as observed in previous studies, exceeded plant demand and leaching losses.     

Increased moisture and methanogenesis contribute to reduced methane oxidation in elevated CO2 soils

Awareness of global warming has stimulated research on environmental controls of soil methane (CH₄) consumption and the effects of increasing atmospheric carbon dioxide (CO₂) on the terrestrial CH₄ sink. In this study, factors impacting soil CH₄ consumption were investigated using laboratory incubations of soils collected at the Free Air Carbon Transfer and Storage I site in the Duke Forest, NC, where plots have been exposed to ambient (370 μL L⁻¹) or elevated (ambient + 200 μL L⁻¹) CO₂ since August 1996. Over 1 year, nearly 90% of the 360 incubations showed net CH₄ consumption, confirming that CH₄-oxidizing (methanotrophic) bacteria were active. Soil moisture was significantly (p < 0.01) higher in the 25–30 cm layer of elevated CO₂ soils over the length of the study, but soil moisture was equal between CO₂ treatments in shallower soils. The increased soil moisture corresponded to decreased net CH₄ oxidation, as elevated CO₂ soils also oxidized 70% less CH₄ at the 25–30 cm depth compared to ambient CO₂ soils, while CH₄ consumption was equal between treatments in shallower soils. Soil moisture content predicted (p < 0.05) CH₄ consumption in upper layers of ambient CO₂ soils, but this relationship was not significant in elevated CO₂ soils at any depth, suggesting that environmental factors in addition to moisture were influencing net CH₄ oxidation under elevated CO₂. More than 6% of the activity assays showed net CH₄ production, and of these, 80% contained soils from elevated CO₂ plots. In addition, more than 50% of the CH₄-producing flasks from elevated CO₂ sites contained deeper (25–30 cm) soils. These results indicate that subsurface (25 cm+) CH₄ production contributes to decreased net CH₄ consumption under elevated CO₂ in otherwise aerobic soils.     

Effects of Aporrectodea caliginosa (Savigny) on nitrogen mobilization and decomposition of elevated‐CO2 Charlock mustard litter

This study was conducted to improve our understanding of how earthworms and microorganisms interact in the decomposition of litter of low quality (high C : N ratio) grown under elevated atmospheric [CO₂]. A microcosm approach was used to investigate the influence of endogeic earthworm (Aporrectodea caliginosa Savigny) activity on the decomposition of senescent Charlock mustard (Sinapis arvensis L.) litter produced under ambient and elevated [CO₂]. Earthworms and microorganisms were exposed to litter which had changed in quality (C : N ratio) while growing under elevated [CO₂]. After 50 d of incubation in microcosms, C mineralization (CO₂ production) in the treatment with elevated‐[CO₂] litter was significantly lower in comparison to the ambient‐[CO₂] litter treatment. The input of Charlock mustard litter into the soil generally induced N immobilization and reduced N₂O‐emission rates from soil. Earthworm activity enhanced CO₂ production, but there was no relationship to litter quality. Although earthworm biomass was not affected by the lower quality of the elevated‐[CO₂] litter, soil microbial biomass (C_mic, N_mic) was significantly decreased. Earthworms reduced C_mic and fungal biomass, the latter only in treatments without litter. Our study clearly showed that A. caliginosa used the litter grown under different [CO₂] independent of its quality and that their effect on the litter‐decomposition process was also independent of litter quality. Soil microorganisms were shown to negatively react to small changes in Charlock mustard litter quality; therefore we expect that microbially mediated C and N cycling may change under future atmospheric [CO₂].     

Elevated CO<Subscript>2</Subscript> concentration affected pine and oak litter chemistry and the respiration and microbial biomass of soils amended with these litters

Elevated atmospheric CO₂ concentration ([CO₂]) may change litter chemistry which affects litter decomposability. This study investigated respiration and microbial biomass of soils amended with litter of Pinus densiflora (a coniferous species; pine) and Quercus variabilis (a deciduous species; oak) that were grown under different atmospheric [CO₂] and thus had different chemistry. Elevated [CO₂] increased lignin/N through increased lignin concentration and decreased N concentration. The CO₂ emission from the soils amended with litter produced under the same [CO₂] regime was greater for oak than pine litter, confirming that broadleaf litter with lower lignin decomposes faster than needle leaf litter. Within each species, however, soils amended with high lignin/N litter grown under elevated [CO₂] emitted more CO₂ than those with low lignin/N litter grown under ambient [CO₂]. Such contrasting effects of lignin/N on inter- and intra-species variations in litter decomposition should be ascribed to the effects of other litter chemistry variables including nonstructural carbohydrate, calcium and manganese as well as inhibitory effect of N on lignin decomposition. The microbial biomass was also higher in the soils amended with high lignin/N litter than those with low lignin/N litter probably due to low substrate use efficiency of lignin by microbes. Our study suggests that elevated [CO₂] increases lignin/N for both species, but increased lignin/N does not always reduce soil respiration and microbial biomass. Further study investigating a variety of tree species is required for more comprehensive understanding of inter- and intra-species variations of litter decomposition under elevated [CO₂].     

Impacts of extreme winter warming events on litter decomposition in a sub-Arctic heathland

Arctic climate change is expected to lead to a greater frequency of extreme winter warming events. During these events, temperatures rapidly increase to well above 0 °C for a number of days, which can lead to snow melt at the landscape scale, loss of insulating snow cover and warming of soils. However, upon return of cold ambient temperatures, soils can freeze deeper and may experience more freeze-thaw cycles due to the absence of a buffering snow layer. Such loss of snow cover and changes in soil temperatures may be critical for litter decomposition since a stable soil microclimate during winter (facilitated by snow cover) allows activity of soil organisms. Indeed, a substantial part of fresh litter decomposition may occur in winter. However, the impacts of extreme winter warming events on soil processes such as decomposition have never before been investigated. With this study we quantify the impacts of winter warming events on fresh litter decomposition using field simulations and lab studies.Winter warming events were simulated in sub-Arctic heathland using infrared heating lamps and soil warming cables during March (typically the period of maximum snow depth) in three consecutive years of 2007, 2008, and 2009. During the winters of 2008 and 2009, simulations were also run in January (typically a period of shallow snow cover) on separate plots. The lab study included soil cores with and without fresh litter subjected to winter-warming simulations in climate chambers.Litter decomposition of common plant species was unaffected by winter warming events simulated either in the lab (litter of Betula pubescens ssp. czerepanovii), or field (litter of Vaccinium vitis-idaea, and B. pubescens ssp. czerepanovii) with the exception of Vaccinium myrtillus (a common deciduous dwarf shrub) that showed less mass loss in response to winter warming events. Soil CO₂ efflux measured in the lab study was (as expected) highly responsive to winter warming events but surprisingly fresh litter decomposition was not. Most fresh litter mass loss in the lab occurred during the first 3-4 weeks (simulating the period after litter fall).In contrast to past understanding, this suggests that winter decomposition of fresh litter is almost non-existent and observations of substantial mass loss across the cold season seen here and in other studies may result from leaching in autumn, prior to the onset of “true” winter. Further, our findings surprisingly suggest that extreme winter warming events do not affect fresh litter decomposition.     

Immediate and adaptational temperature effects on nitric oxide production and nitrous oxide release from nitrification and denitrification in two soils

 Nitrification and denitrification are, like all biological processes, influenced by temperature. We investigated temperature effects on N trace gas turnover by nitrification and denitrification in two soils under two experimental conditions. In the first approach ("temperature shift experiment") soil samples were preincubated at 25  °C and then exposed to gradually increasing temperatures (starting at 4  °C and finishing at 40–45  °C). Under these conditions the immediate effect of temperature change was assessed. In the second approach ("discrete temperature experiment") the soil samples were preincubated at different temperatures (4–35  °C) for 5 days and then tested at the same temperatures. The different experimental conditions affected the results of the study. In the temperature shift experiment the NO release increased steadily with increasing temperature in both soils. In the discrete temperature experiment, however, the production rates of NO and N₂O showed a minimum at intermediate temperatures (13–25  °C). In one of the soils (soil B9), the percent contribution of nitrification to NO production in the discrete temperature experiment reached a maximum (>95% contribution) at 25  °C. In the temperature shift experiment nitrification was always the dominant process for NO release and showed no systematic temperature dependency. In the second soil (soil B14), the percent contribution of nitrification to NO release decreased from 50 to 10% as the temperature was increased from 4  °C to 45  °C, but no differences were evident in the discrete temperature experiment. The N₂O production rates were measured in the discrete temperature experiment only. The contribution of nitrification to N₂O production in soil B9 was considerably higher at 25–35  °C (60–80% contribution) than at 4–13  °C (15–20% contribution). In soil B14 the contribution of nitrification to N₂O production was lowest at 4  °C. The effects of temperature on N trace gas turnover differed between the two soils and incubation conditions. The experimental set-up allowed us to distinguish between immediate effects of short-term changes in temperature on the process rates, and longer-term effects by which preincubation at a particular temperature presumably resulted in the adaptation of the soil microorganisms to this temperature. Both types of effects were important in regulating the release of NO and N₂O from soil. Received: 20 October 1998     

Quantifying microbial biomass phosphorus in acid soils

 This study aimed to validate the fumigation-extraction method for measuring microbial biomass P in acid soils. Extractions with the Olsen (0.5 M NaHCO₃, pH 8.5) and Bray-1 (0.03 M NH₄F–0.025 M HCl) extractants at two soil:solution ratios (1 : 20 and 1 : 4, w/v) were compared using eight acid soils (pH 3.6–5.9). The data indicated that the flushes (increases following CHCl₃-fumigation) of total P (P_t) and inorganic P (P_i) determined by Olsen extraction provided little useful information for estimating the amount of microbial biomass P in the soils. Using the Bray-1 extractant at a soil:solution ratio of 1 : 4, and analysing P_i instead of P_t, improves the reproducibility (statistical significance and CV) of the P flush in these soils. In all the approaches studied, the P_i flush determined using the Bray-1 extractant at 1 : 4 provided the best estimate of soil microbial biomass P. Furthermore, the recovery of cultured bacterial and fungal biomass P added to the soils and extracted using the Bray-1 extractant at 1 : 4 was relatively constant (24.1–36.7% and 15.7–25.7%, respectively) with only one exception, and showed no relationship with soil pH, indicating that it behaved differently from added P_i (recovery decreased from 86% at pH 4.6 to 13% at pH 3.6). Thus, correcting for the incomplete recovery of biomass P using added P_i is inappropriate for acid soils. Although microbial biomass P in soil is generally estimated using the P_i flush and a conversion factor (k _P) of 0.4, more reliable estimates require that k _P values are best determined independently for each soil. Received: 3 February 2000     

Effect of temperature on reduction of iron and production of carbon dioxide and methane in anoxic wetland rice soils

 Wetland rice soils from Italy (Pavia) and the Philippines (Bugallon, Luisiana, Maligaya) were incubated under anoxic conditions at 31 different temperatures ranging from 4.7  °C to 49.5  °C. Production of CO₂ was most intensive at the beginning of the incubation (0–4 days) and was predominantly coupled to the reduction of free Fe(III). The optimum temperature for these processes was between 32  °C and 41  °C. After 9–16 days, CO₂ production rates had decreased and the available Fe(III) had been completely reduced at the optimum temperatures. However, Fe(III) was still available at temperatures below and above the optimum. Maximum CH₄ production rates were observed after 4–16 days (except in soil from Maligaya) with temperature optima between 32  °C and 41  °C, similar to those for CO₂ production and Fe reduction. Since ongoing Fe reduction is known to suppress CH₄ production, the temperature range of optimum CH₄ production was restricted to those temperatures at which Fe(III) had already been depleted. Nevertheless, the temperature characteristics of both CO₂ and CH₄ production often exhibited two temperature optima at some time during the incubation, suggesting a complex pattern of adaptation of the methanogenic microbial community to temperature. When available Fe(III) was completely depleted by anoxic pre-incubation at 30  °C, CH₄ was produced at a constant rate (steady state conditions) which increased with increasing temperature. Steady state CH₄ production reached a first maximum at about 40  °C, but increased further up to at least 50  °C, suggesting the presence of thermophilic microorganisms whose activity was apparently masked when Fe had not been completely reduced. The apparent activation energy of CH₄ production at steady state ranged between 48 kJ mol^–1 and 65 kJ mol^–1. Received: 26 August 1999     

Potential contribution of Lumbricus terrestris L. to carbon dioxide, methane and nitrous oxide fluxes from a forest soil

 Potential effects of earthworms (Lumbricus terrestris L.) inoculated into soil on fluxes of CO₂, CH₄ and N₂O were investigated for an untreated and a limed soil under beech in open topsoil columns under field conditions for 120 days. Gas fluxes from L. terrestris, beech litter and mineral soil from soil columns were measured separately in jars at 17  °C. The inoculation with L. terrestris and the application of lime had no effect on cumulative CO₂ emissions from soil. During the first 3–4 weeks earthworms significantly (P<0.05) increased CO₂ emissions by 16% to 28%. In contrast, significantly lower (P<0.05) CO₂ emission rates were measured after 11 weeks. The data suggest that earthworm activity was high during the first weeks due to the creation of burrows and incorporation of beech litter into the mineral soil. Low cumulative CH₄ oxidation rates were found in all soil columns as a result of CH₄ production and oxidation processes. L. terrestris with fresh feces and the beech litter produced CH₄ during the laboratory incubation, whereas the mineral soil oxidised atmospheric CH₄. Inoculation with L. terrestris led to a significant reduction (P<0.02) in the CH₄ oxidation rate of soil, i.e. 53% reduction. Liming had no effect on cumulative CH₄ oxidation rates of soil columns and on CH₄ fluxes during the laboratory incubation. L. terrestris significantly increased (P<0.001) cumulative N₂O emissions of unlimed soil columns by 57%. The separate incubation of L. terrestris with fresh feces resulted in rather high N₂O emissions, but the rate strongly decreased from 54 to 2 μg N kg^–1 (dry weight) h^–1 during the 100 h of incubation. Liming had a marked effect on N₂O formation and significantly (P<0.001) reduced cumulative N₂O emissions by 34%. Although the interaction of liming and L. terrestris was not significant, N₂O emissions of limed soil columns with L. terrestris were 8% lower than those of the control. Received: 2 September 1999     

Effects of elevated temperature, elevated CO2 and fertilization on quality and subsequent decomposition of silver birch leaf litter

We examined the quality and decomposition of naturally abscised leaves of silver birch (Betula pendula) seedlings subjected to three different levels of fertilization under ambient and elevated levels of temperature and CO₂. At the end of the second growing season, the chemical composition of the litter collected from the seedlings was analyzed. Whole-leaf samples from pooled litter from each of the four replicates from each treatment were put in mesh bags and transferred to ambient climate in the field. The remaining mass of litter was measured by sampling bags in May and October throughout the four-year incubation period. Fertilization with all nutrients decreased the initial carbon and tannin contents of litter, and increased the proportion of the fast-decomposing fraction, but still fertilization slowed down the decomposition of this fraction. Initially, the estimated proportion of the fast-decomposing fraction was smallest in elevated CO₂ + temperature, and largest in ambient climate. During decomposition, elevated growth-temperature slowed down decomposition of the fast fraction under ambient CO₂ but increased it under elevated CO₂. The changes in litter decomposition rates found over four years were not very large. However, we conclude that the interactions of different factors lead to different results than if the factors had been studied separately, and future studies should take interactions into account.     

Warming and elevated CO2 combine to increase microbial mineralisation of soil organic matter

The net annual exchange of carbon between the atmosphere and terrestrial ecosystems is of prime importance in determining the concentration of CO₂ ([CO₂]) in the atmosphere and consequently future climate. Carbon loss occurs primarily through soil respiration; it is known that respiration is sensitive to the global changes in [CO₂] and temperature, suggesting that the net carbon balance may change in the future. However, field manipulations of temperature and [CO₂] alter many important environmental factors so it is unclear how much of the observed alterations in soil respiration is due to changes of microbial function itself instead of changes to the physical and chemical environment. Here we focus on resolving the importance of changes in the microbial community in response to warming and elevated [CO₂] on carbon mineralisation, something not possible in field measurements. We took plant material and soil inocula from a long running experiment where native grassland had been exposed to both warming and elevated CO₂ and constructed a reciprocal transplant experiment. We found that the rate of decomposition (heterotrophic respiration) was strongly determined by the origin of the microbial community. The combined warming + elevated CO₂ treatment produced a soil community that gave respiration rates 30% higher when provided with shoot litter and 70% for root litter than elevated CO₂ treatment alone, with the treatment source of the litter being unimportant. Warming, especially in the presence of elevated CO₂, increased the size of the apparent labile carbon pool when either C₃ or C₄ litter was added. Thus, the metabolic activity of the soil community was affected by the combination of warming and elevated CO₂ such that it had an increased ability to mineralise added organic matter, regardless of its source. Therefore, soil C efflux may be substantially increased in a warmer, high CO₂ world. Current ecosystem models mostly drive heterotrophic respiration from plant litter quality, soil moisture and temperature but our findings suggest equal attention will need to be paid to capturing microbial processes if we are to accurately project the future C balance of terrestrial ecosystems and quantify the feedback effect on atmospheric concentrations of CO₂.     

Microwave irradiation of soil for routine measurement of microbial biomass carbon

 Microwave irradiation was evaluated as a non-toxic alternate to chloroform fumigation for routine measurement of soil microbial biomass C. Microwave energy was applied to moist soil to disrupt microbial cells. The flush of C released was then measured after extraction or incubation. Microwave irradiation at 800 J g^–1 soil was optimal because this level resulted in an almost instantaneous rise in soil temperature (≥80  °C), an abrupt reduction in microbial activity, maximal release of biomass C, and minimal solubilization of humic substances. Both incubation-CO₂ titration and extraction-colorimetry methods were used on separate 20-g subsamples to compare the labile C in the microwave-treated and untreated soil samples. The incubation-titration method was also used to measure C in chloroform-fumigated soil samples. Averaged across soils, the chloroform fumigation yielded 123.3±5.1 mg CO₂-C kg^–1. Microwave irradiation yielded 93.6±3.9 mg CO₂-C kg^–1 soil determined by incubation and 52.4±2.4 mg C kg^–1 soil determined by extraction, accounting for 76% and 42% of the net flush of C measured by the chloroform fumigation. Microwave-stimulated net flushes of C were correlated closely (r ²=0.974 for incubation or 0.908 for extraction) with microbial biomass C measured by the chloroform fumigation. Little correlation was found with the total soil organic C (r ²=0.241 for incubation or for 0.166 extraction). Mean efficiency factors for incubation (K _MI) or extraction (K _ME) were used to calculate microbial biomass C from net flushes of C between microwaved and unmicrowaved soils. Values of K _MI and K _ME were not affected by soil pH, bulk density or clay contents. Extraction of microwaved soil by 0.5M K₂SO₄ proved to be a simple, fast, precise, reliable, and safe method to measure soil microbial biomass C. Received: 12 September 1997     

Responses of nitrous oxide, dinitrogen and carbon dioxide production and oxygen consumption to temperature in forest and agricultural light-textured soils determined by model experiment

 The experiment, carried out on a forest and arable light-textured soil, was designed to study the temperature response of autotrophic and heterotrophic N₂O production and investigate how the N₂O flux relates to soil respiration and O₂ consumption. Although N₂O production seemed to be stimulated by a temperature increase in both soils, the relationship between production rate and temperature was different in the two soils. This seemed to depend on the different contribution of nitrification and denitrification to the overall N₂O flux. In the forest soil, almost all N₂O was derived from nitrification, and its production rate rose linearly from 2  °C to 40  °C. A stronger effect of temperature on N₂O production was observed in the arable soil, apparently as a result of an incremental contribution of denitrification to the overall N₂O flux with rising temperature. The soil respiration rate increased exponentially with temperature and was significantly correlated with N₂O production. O₂ consumption stimulated denitrification in both soils. In the arable soil, N₂O and N₂ production increased exponentially with decreasing O₂ concentration, though N₂O was the main gas produced at any temperature. In the forest soil, only the N₂ flux was related exponentially to O₂ consumption and it outweighed the rate of N₂O production only at >34  °C. Thus, it appears that in the forest soil, where nitrification was the main source of N₂O, temperature affected the N₂O flux less dramatically than in the arable soil, where a temperature increase strongly stimulated N₂O production by enhancing favourable conditions for denitrification. Received: 26 August 1998     

Carbon dioxide emissions of soils under pure and mixed stands of beech and spruce, affected by decomposing foliage litter mixtures

Soil respiration is the largest terrestrial source of CO₂ to the atmosphere. In forests, roughly half of the soil respiration is autotrophic (mainly root respiration) while the remainder is heterotrophic, originating from decomposition of soil organic matter. Decomposition is an important process for cycling of nutrients in forest ecosystems. Hence, tree species induced changes may have a great impact on atmospheric CO₂ concentrations. Since studies on the combined effects of beech-spruce mixtures are very rare, we firstly measured CO₂ emission rates in three adjacent stands of pure spruce (Picea abies), mixed spruce-beech and pure beech (Fagus sylvatica) on three base-rich sites (Flysch) and three base-poor sites (Molasse; yielding a total of 18 stands) during two summer periods using the closed chamber method. CO₂ emissions were higher on the well-aerated sandy soils on Molasse than on the clayey soils on Flysch, characterized by frequent water logging. Mean CO₂ effluxes increased from spruce (41) over the mixed (55) to the beech (59) stands on Molasse, while tree species effects were lower on Flysch (30-35, mixed > beech = spruce; all data in mg CO₂-C m⁻² h⁻¹). Secondly, we studied decomposition after fourfold litter manipulations at the 6 mixed species stands: the O_i - and O_e horizons were removed and replaced by additions of beech -, spruce - and mixed litter of the adjacent pure stands of known chemical quality and one zero addition (blank) in open rings (20 cm inner diameter), which were covered with meshes to exclude fresh litter fall. Mass loss within two years amounted to 61-68% on Flysch and 36-44% on Molasse, indicating non-additive mixed species effects (mixed litter showed highest mass loss). However, base cation release showed a linear response, increasing from the spruce - over the mixed - to the beech litter. The differences in N release (immobilization) resulted in a characteristic converging trend in C/N ratios for all litter compositions on both bedrocks during decomposition. In the summers 2006 and 2007 we measured CO₂ efflux from these manipulated areas (a closed chamber fits exactly over such a ring) as field indicator of the microbial activity. Net fluxes (subtracting the so-called blank values) are considered an indicator of litter induced changes only and increased on both bedrocks from the spruce - over the mixed - to the beech litter. According to these measurements, decomposing litter contributed between 22-32% (Flysch) and 11-28% (Molasse) to total soil respiration, strengthening its role within the global carbon cycle.     

Changes in soil microbial biomass, metabolic quotient, and organic matter turnover under Hieracium (H. pilosella L.)

 In New Zealand Hieracium is an opportunistic plant that invades high country sites more or less depleted of indigenous vegetation. To understand the invasive nature of this weed we assessed the changes in soil C, N and P, soil microbial biomass C, N and P contents, microbial C : N and C : P ratios, the metabolic quotient, and turnover of organic matter in soils beneath Hieracium and its adjacent herbfield resulting from the depletion of tussock vegetation. The amounts of soil organic C and total N were higher under Hieracium by 25 and 11%, respectively, compared to soil under herbfield. This change reflects an improvement in both the quantity and quality of organic matter input to mineral soil under Hieracium, with higher percentage organic C and a lower C : N ratio. The microbial biomass C, N and P contents were also higher under Hieracium. The amount of C respired during the 34-week incubation indicated differences in the nature of soil organic matter under Hieracium, the unvegetated "halo" zone surrounding Hieracium patches, and herbfield (depleted tussock grassland). Decomposition of organic matter in these zones showed that the Hieracium soil had the greatest rate of CO₂ respired, and the halo soil had the lowest. We relate the enhanced organic C turnover to the invasive nature of Hieracium. Net N mineralization was significantly lower from the Hieracium soil (57 mg N g^–1 soil N) than from herbfield and halo soils (74 and 71 mg N g^–1 soil N, respectively), confirming that the nature of organic N in Hieracium soil is different from adjoining halo and herbfield soils. It seems plausible that specific compounds such as polyphenols and lignins released by Hieracium are not only responsible for increased organic N, but also control the form and amount of N released during organic matter transformations. We conclude that the key to the success of Hieracium in the N-deficient South Island high country of New Zealand lies in its ability to control and sequester N supply through modifying the soil organic matter cycle. Received: 1 December 1998     

The microbial biomass and its activity and structure in the soils of old forests in the European Russia

The humus-accumulative layer of soils (podzolic, gray, rzhavozem, burozem, and karbolitozem) of old-age forests (>60–450 years old) localized in various vegetation subzones (middle-taiga, southern taiga, subtaiga, dark coniferous forests outside the boreal region, and mountain forests) of the European part of Russia (22 sites of soil sampling of them, 13 in nature reserves and specially protected territories) was studied. The carbon content of the microbial biomass (C_mic) in the soil was determined by the substrate-induced respiration method. The fungal to bacterial ratio was determined by the selective inhibition technique with antibiotics. The basal respiration (BR) was also measured. The BR/C_mic = qCO₂ ratio and the portion of C_mic in the total organic soil carbon was determined. It was shown that the C_mic and BR in the soils of a separate vegetation subzone varied significantly; however, their values increased from the middle-taiga to dark coniferous subzone and decreased in the mountain-forest zone (348 ± 44, 670 ± 66, 1000 ± 86, 1142 ± 49, 789 ± 79 μkg C/g soil and from 0.68 ± 0.23, 1.85 ± 0.10, 2.13 ± 0.15, 1.56 ± 0.14, 0.92 ± 0.07 μkg CO₂-C/soil h, respectively). The fungal component in the humus-accumulative layer of soils is 53–99% of the total Cmic; however, its absolute values increase from the middle subzone to the southern one. The Cmic pool and the total BR in the profile of some soils (mineral horizons and forest litter) were calculated.     

Soil microbial and extractable C and N after wildfire

 The effect of wildfire on soil microbes and extractable C (C_ext) and N (N_ext) changed with respect to the time from burning and soil depth. Initially, microbial biomass C (C_mic) and N (N_mic) were drastically reduced in the soil surface layer (0–5 cm) and reduced by 50% in the subsurface (5–10 cm), whereas C_ext increased by 62% in the surface layer and did not significantly change in the subsurface. These parameters were affected for the following 4 years, during which the average reductions in the soil surface and subsurface layers were, respectively, 60% and 50% for C_mic, 70% and 45% for N_mic, 60% and 40% for the ratio C_mic: organic C (C_org) and 70% and 30% for the ratio N_mic: total N (N_tot), while for C_ext the surface layer was the only zone consistently affected and C_ext decreased by up to 59%. Immediately after a fire, the C_ext : C_org ratio increased by 3.5-fold and 2-fold in the surface and subsurface layers, respectively; thereafter for 2 years, it decreased in the surface layer (by up to 45%) while the effect on the subsurface layer was not consistent. The effect of burning on N_ext lasted 1 year, in which N_ext increased by up to 7- and 3-fold in the surface and subsurface layers, respectively, while the average N_ext : N_tot ratio doubled in the surface layer and increased by 34% in the subsurface. During the time in which each parameter was affected by burning, the soil factor explained a high percentage of variance in the fluctuations of C_mic, N_mic, C_mic : C_org and N_mic : N_tot, while those of N_ext and N_ext : N_tot, but not those of C_ext and C_ext : C_org depended on both the soil and its depth. In the burned soils similar patterns of response were found between the following parameters listed in pairs: C_mic and N_mic; C_mic : C_org and N_mic : N_tot; C_ext and N_ext; and C_ext : C_org and N_ext : N_tot. However, after the fire relationships found previously between the parameters studied and many other soils properties were either no longer evident, or were inverted. Although the addition of cellulose to the burned soil favoured fungal mycelium development and increased C_mic and C_ext contents, the negative effect of burning on the microbial biomass and the C_ext was not counteracted even under incubation conditions suitable for both microbial growth and C mineralization. Received: 28 May 1997     

Structural and functional diversity of soil microbes is affected by elevated [CO<Subscript>2</Subscript>] and N addition in a poplar plantation

Background, Aims, and Scope  The genetic structure and the functionality of soil microbes are both important when studying the role of soil in the C cycle in elevated CO₂ scenarios. The aim of this work was to investigate the genetic composition of the fungal community by means of PCR-DGGE and the functional diversity of soil micro-organisms in general with MicroResp-based community level physiological profiling (CLPP) in a poplar plantation (POPFACE) grown under elevated [CO₂] with and without nitrogen fertilization. Materials and Methods  The POPFACE experimental plantation and FACE facility are located in central Italy, Tuscania (VT). Clones of Populus alba, Populus nigra and Populus x euramericana were grown, from 1999 to 2004, in six 314 m² plots treated either with atmospheric (control) or enriched (550 μmol mol⁻¹) CO₂ with FACE (Free Air CO₂ Enrichment) technology in each growing season. Each plot is divided into six triangular sectors, with two sectors per poplar genotype: three species × two nitrogen levels. After removal of the litter layer one soil core per genotype (10 cm wide, 20 cm depth) was taken inside each of the three sectors in each plot, for a total of 36 soil cores (3 replicates × 2 [CO₂] × 2 fertilization × 3 species) in October 2004 and in July 2005. DNA was extracted with a bead beating procedure. 18S rDNA gene fragments were amplified with PCR using fungal primers (FR1 GC and FF390). Analysis of CLPP was performed using the MicroResp method. Carbon substrates were selected depending on their ecological relevance to soil and their solubility in water. In particular rhizospheric C sources (carboxylic acids and carbohydrates) were chosen considering the importance of root inputs for microbial metabolism. Results  The fertilization treatment differentiated the fungal community composition regardless of elevated [CO₂] or the poplar species; moreover the number of fungal species was lower in fertilized soil. The effect of elevated [CO₂] on the fungal community composition was evident only as interaction with the fertilization treatment as, in N-sufficient soils, the elevated [CO₂] selected a different microbial community. For CLPP, the differ ent poplar species were the main factors of variation. The FACE treatment, on average, resulted in lower C utilization rates in un-fertilized soils and higher in fertilized soils. Discussion  Fungal biomass and fungal composition depend on different factors: from previous studies we know that the greater quantity and the higher C/N ratio of organic inputs under elevated [CO₂] influenced positively the fungal biomass both in fertilized and in un-fertilized soil, whereas nitrogen availability resulted to be the main determinant of fungal community composition in this work. Whole active microbial community was directly influenced by the soil nutrient availability and the poplar species. Under elevated CO₂ the competition for N with plants strongly affected the microbial communities, which were not able to benefit from added rhizospheric substrates. Under Nsufficient conditions, the increase of microbial activity due to [CO₂] enrichment was related to a more active microbial community, favoured by the current availability of C and N. Conclusions  Different factors influenced the microbial community at different levels: poplar species and root exudates affected the functional properties of the microbial community, while the fungal specific composition (as seen with DGGE) remained unaffected. On the other hand, factors such as N and C availability had a strong impact on the community functionality and composition. Fungal community structure reflected the availability of N in soils and the effect of elevated [CO₂] on community structure and function was evident only in N-sufficient soils. The simultaneous availability of C and N was therefore the main driving force for microbial structure and function in this plantation. Recommendations and Perspectives  Using the soil instead of soil extracts for CLPP determination provides a direct measurement of substrate catabolism by microbial communities and reflects activity rather than growth because more immediate responses to substrates are measured. Further applications of this approach could include selective inhibition of different microbial functional groups to investigate specific CLPPs. To combine the structural analysis and the catabolic responses of specific microbial communities (i.e. fungi or bacteria) could provide new outlooks on the role of microbes on SOM decomposition. ESS-Submission Editor: Dr. Kirk Semple (k.semple@lancaster.ac.uk)     

Microbial biomass carbon and the microbial carbon dioxide production by soddy-podzolic soils in postagrogenic biogeocenoses and in native spruce forests of the southern taiga (Kostroma oblast)

In two layers of the humus horizons in soddy-podzolic soils of different biogeocenoses (Kostroma oblast) representing a succession series, the carbon content in the microbial biomass (C_mic) was determined using the method of substrate-induced respiration and the rate of microbial CO₂ production (basal respiration, BR). The C_mic content was from 110 to 755 μg/g soil, and the BR was from 0.40 to 2.52 μg CO₂-C/g/h. A gradual increase in the C_mic content and BR was found in the following sequence: cropland—fallow (7-year-old)—young (20- and 45-year-old) forests—secondary and native (primary) forests (90- and 450-year-old, respectively). In the litter, the C_mic content was higher in the 45-year-old forest than in the secondary and native forests: 10423, 6459, and 4258 μg C/g of substrate, respectively. The portion of C_mic in the soil organic carbon content in the upper layer of the soils studied varied from 1.3 to 5.4%; its highest value was in the soils under the secondary and native forests. The pool of microbial biomass carbon and the microbial CO₂ production in the upper 25-cm layer of the soils were calculated.     

Soil respiration, nitrogen mineralization and uptake in barley following cultivation of grazed grasslands

 Soil tillage was studied as a strategy to synchronize N mineralization with plant demand following ploughing of two types of grazed pastures [ryegrass/white clover (Lolium perenne/Trifolium repens) and pure ryegrass]. The swards were either rotovated and ploughed or ploughed only. Soil respiration, as determined by a dynamic chamber method, was related to net N mineralization and to plant N uptake in a subsequent spring barley crop (Hordeum vulgare). Diurnal variations in temperature were important for the CO₂ flux and care must be taken that temperatures during measuring periods are representative of the daily mean. Soil tillage increased the CO₂ flux considerably compared with untilled soil with total emissions of 2.6 and 1.4 t C ha^–1, respectively, from start of April to end of June. Sward type or rotovation did not markedly influence accumulated emissions. Rotovation significantly increased the content of nitrate in the soil until 43 days after rotovation, showing that net N mineralization occurred rapidly during this period, in spite of low soil temperatures (5–10  °C). Rotovation increased barley grain yield by 10–12% and N-uptake by 14%. For both sward types, rotovation caused an extra N-uptake in harvested plant material of about 12 kg ha^–1. The availability of soil inorganic N at the early stages of barley was important for the final yield and N-uptake. The results indicated that soil biological activity was not enhanced by rotovation and that the yield effect of rotovation was mainly caused by quicker availability and better synchrony between N mineralization and plant uptake due to earlier start of decomposition. Received: 3 May 2000