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 atmospheric CO2 affects the turnover of nitrogen in a European grassland

The availability of nutrients in the soil is key to the potential response of a plant to elevated CO₂ and is central to correctly predicting the response of terrestrial communities to climate change. In order for a plant to fully realise the potential of increased atmospheric CO₂, it must increase its nutrient uptake for the increased production of biomass as well as biochemical compounds. In this study the stable isotope ¹⁵N was used to follow the fate of nitrogen contained in litter in order to determine the effect elevated atmospheric CO₂ had on the loss of nitrogen from decomposing litter and the eventual re-use of this nitrogen. During the decomposition study, on a mass basis more ¹⁵N was transferred from the litter despite the litter grown in elevated CO₂ initially having a lower ¹⁵N signal. This was primarily related to a higher decomposition rate of the elevated CO₂ grown litter. Despite more nitrogen entering the below-ground community under elevated atmospheric CO₂, the additional N did not stay within the terrestrial community and was not exploited by the plants. The results confirm previous suggestions that Lolium perenne plants growing in elevated CO₂ have to derive at least a proportion of their nitrogen from a source external to either added fertiliser or decomposing litter     

大气CO2浓度升高对温带森林土壤微生物活性的影响

Microorganisms play a key role in the response of soil ecosystems to the rising atmospheric carbon dioxide (CO2) as they mineralize organic matter and drive nutrient cycling. To assess the effects of elevated CO2 on soil microbial C and N immobilization and on soil enzyme activities, in years 8 (2006) and 9 (2007) of an open-top chamber experiment that begun in spring of 1999, soil was sampled in summer, and microbial biomass and enzyme activity related to the carbon (C), nitrogen (N) and phosphorus (P) cycling were measured. Although no effects on microbial biomass C were detected, changes in microbial biomass N and metabolic activity involving C, N and P were observed under elevated CO2. Invertase and dehydrogenase activities were significantly enhanced by different degrees of elevated CO2. Nitrifying enzyme activity was significantly (P < 0.01) increased in the August 2006 samples that received the elevated CO2 treatment, as compared to the samples that received the ambient treatment. Denitrifying enzyme activity was significantly (P < 0.04) decreased by elevated CO2 treatments in the August 2006 and June 2007 (P < 0.09) samples. β-N-acetylglucosaminidase activity was increased under elevated CO2 by 7% and 25% in June and August 2006, respectively, compared to those under ambient CO2. The results of June 2006 samples showed that acid phosphatase activity was significantly enhanced under elevated CO2. Overall, these results suggested that elevated CO2 might cause changes in the belowground C, N and P cycling in temperate forest soils.     

Decomposition of C-labeled roots in a pasture soil exposed to 10 years of elevated CO2

The net flux of soil C is determined by the balance between soil C input and microbial decomposition, both of which might be altered under prolonged elevated atmospheric CO₂. In this study, we determined the effect of elevated CO₂ on decomposition of grass root material (Lolium perenne L.). ¹⁴C-labeled root material, produced under ambient (35 Pa pCO₂) or elevated CO₂ (70 Pa pCO₂) was incubated in soil for 64 days. The soils were taken from a pasture ecosystem which had been exposed to ambient (35 Pa pCO₂) or elevated CO₂ (60 Pa pCO₂) under FACE-conditions for 10 years and two fertilizer N rates: 140 and 560 kg N ha⁻¹ year⁻¹. In soil exposed to elevated CO₂, decomposition rates of root material grown at either ambient or elevated CO₂ were always lower than in the control soil exposed to ambient CO₂, demonstrating a change in microbial activity. In the soil that received the high rate of N fertilizer, decomposition of root material grown at elevated CO₂ decreased by approximately 17% after incubation for 64 days compared to root material grown at ambient CO₂. The amount of ¹⁴CO₂ respired per amount of ¹⁴C incorporated in the microbial biomass (q¹⁴CO₂) was significantly lower when roots were grown under high CO₂ compared to roots grown under low CO₂. We hypothesize that this decrease is the result of a shift in the microbial community, causing an increase in metabolic efficiency. Soils exposed to elevated CO₂ tended to respire more native SOC, both with and without the addition of the root material, probably resulting from a higher C supply to the soil during the 10 years of treatment with elevated CO₂. The results show the importance of using soils adapted to elevated CO₂ in studies of decomposition of roots grown under elevated CO₂. Our results further suggest that negative priming effects may obscure CO₂ data in incubation experiments with unlabeled substrates. From the results obtained, we conclude that a slower turnover of root material grown in an ‘elevated-CO₂ world’ may result in a limited net increase in C storage in ryegrass swards.     

Nine years of enriched CO2 changes the function and structural diversity of soil microorganisms in a grassland

To gain insight into microbial function following increased atmospheric CO₂ concentration, we investigated the influence of 9 years of enriched CO₂ (600 μl litre⁻¹) on the function and structural diversity of soil microorganisms in a grassland ecosystem under free air carbon dioxide enrichment (FACE), as affected by plant species (Trifolium repens L. and Lolium perenne L. in monocultures and mixed culture) and nitrogen (N) supply. We measured biomass and activities of enzymes covering cycles of the most important elements (C, N and P). The microbial community was profiled by molecular techniques of phospholipid fatty acid (PLFA) and denaturing gradient gel electrophoresis (DGGE) analysis. The enrichment in CO₂ increased soil microbial biomass (+48.1%) as well as activities of invertase (+36.2%), xylanase (+22.9%), urease (+23.8%), protease (+40.2%) and alkaline phosphomonoesterase (+54.1%) in spring 2002. In autumn, the stimulation of microbial biomass was 25% less and that of enzymes 3–12% less than in spring. Strong correlations between activities of invertase, protease, urease and alkaline phosphomonoesterase and microbial biomass were found. The stimulation of microbial activity in the enriched atmosphere was probably caused by changes in the quantity and kind of root litter and rhizodeposition. The response of soil microorganisms to enriched CO₂ was most pronounced under Trifolium monoculture and under greater N supply. The PLFA analysis revealed that total PLFA contents were greater by 24.7% on average, whereby the proportion of bioindicators representative of Gram‐negative bacteria increased significantly in the enriched CO₂ under less N‐fertilized Lolium culture. Discriminant analysis showed marked differences between the PLFA profiles of the three plant communities. Shannon diversity indices calculated from DGGE patterns were greater (+12.5%) in the enriched CO₂, indicating increased soil bacterial diversity. We conclude that greater microbial biomass and enzyme activity buffer the potential increase in C sequestration occurring from greater C addition in enriched CO₂ due to greater mineralization of soil organic matter.     

Effects of long-term CO2 fumigation on fungal communities in a temperate forest soil

This study aimed at determining the impact of long-time elevated CO₂ fumigation on fungal communities in a temperate forest soil. In addition to the CO₂ concentration, both time and its interaction with the CO₂ affected the activity of 1,4-β-N-acetylglucosaminidase that is mainly of the fungal origin in the soil. No significant change in Shannon's indexes (from 18S rDNA-PCR-DGGE) was observed between the ambient and elevated CO₂ treatments. Analysis of time-course indicated that the succession of soil fungal community was altered by the elevated CO₂ fumigation, and the variations in the soil samples under Pinus koraiensis Sieb. et Zucc were larger than those under the Pinus sylvestriformis (Takenouchi) T. Wang ex Cheng samples. The results suggest that the increase in atmospheric CO₂ concentrations could alter the temporal patterning of soil fungal communities.     

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].     

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.     

Effect of temperature on below-ground N-dynamics in a weedy model ecosystem at ambient and elevated atmospheric CO2 levels

Model multispecies terrestrial communities composed of four trophic levels (plants, herbivores, parasitoids, decomposers) were established in the Ecotron controlled environment facility. Two experimental runs enabled us to investigate the effects of enhanced temperature on below-ground microbial processes (N-mineralisation, urease, arginine deaminase, protease activity and potential denitrification) in both ambient and elevated (ambient +200 ppm) CO₂ atmospheres.The enzyme activities involved in nitrogen cycling showed weak responses to elevated temperature in both experimental runs. In the Ambient CO₂ Run, protease and arginine deaminase values tended to be lower in elevated temperature; on the other hand, N-mineralisation, urease and denitrification enzyme activity (DEA) were higher. In the Elevated CO₂ Run, all microbial variables showed higher activities at elevated temperature, although only the results for DEA and arginine deaminase were statistically significant. The interaction between higher temperature and elevated CO₂ weakly affected root growth and tissue C:N ratio, limiting feedbacks into the microbial community.Besides temperature and CO₂, substrate availability, water stress and successional development regulated the response of the soil microbes. The supply of organic carbon and nitrogen in the soil allowed plant growth and maintenance of the microbial population. Nitrogen competition between vegetation and microbes restricted net microbial growth. The increase of dissolved organic carbon (DOC) at higher CO₂ and temperature levels significantly favoured DEA. The high water regime in the soil also favoured DEA and inhibited oxidation of organic compounds, as indicated by low levels of enzyme activity. Additionally, water stress decreased rooting density in the soil; this resulted in negative feedback into microbial processes. We conclude that water stress and soil nitrogen deficiency caused an early levelling-off of both microbial population growth and activity rates during the early part of the model ecosystem's development.     

Assessment of 10 years of CO2 fumigation on soil microbial communities and function in a sweetgum plantation

Increased vegetative growth and soil carbon (C) storage under elevated carbon dioxide concentration ([CO₂]) has been demonstrated in a number of experiments. However, the ability of ecosystems, either above- or belowground, to maintain increased C storage relies on the response of soil processes, such as those that control nitrogen (N) mineralization, to climatic change. These soil processes are mediated by microbial communities whose activity and structure may also respond to increasing atmospheric [CO₂]. We took advantage of a long-term (ca 10 y) CO₂ enrichment experiment in a sweetgum plantation located in the southeastern United States to test the hypothesis that observed increases in root production in elevated relative to ambient CO₂ plots would alter microbial community structure, increase microbial activity, and increase soil nutrient cycling. We found that elevated [CO₂] had no detectable effect on microbial community structure using 16S rRNA gene clone libraries, on microbial activity measured with extracellular enzyme activity, or on potential soil N mineralization and nitrification rates. These results support findings at other forested Free Air [CO₂] Enrichment (FACE) sites.     

Microbial function in adjacent subtropical forest and agricultural soil

Soil microbial communities and their activities are altered by land use change; however impacts and extent of these alterations are often unclear. We investigated the functional responses of soil microbes in agricultural soil under sugarcane and corresponding native soil under Eucalyptus forest to additions of contrasting plant litter derived from soybean, sugarcane and Eucalyptus in a microcosm system, using a suite of complimentary techniques including enzyme assays and community level physiological profiles (CLPP). Initially agricultural soil had 50% less microbial biomass and lower enzyme activities than forest soil, but significantly higher nitrification rates. In response to litter addition, microbial biomass increased up to 11-fold in agricultural soil, but only 1.8-fold in forest soil, suggesting a prevalence of rapidly proliferating ‘r’ and slower growing ‘K’ strategists in the respective soils. Litter-driven change in microbial biomass and activities were short lived, largely returning to pre-litter addition levels by day 150. Decomposition rates of sugarcane and soybean litter as estimated via CO₂ production were lower in agricultural than in forest soil, but decomposition of more recalcitrant Eucalyptus litter was similar in both soils, contradicting the notion that microbial communities specialise in decomposing litter of the dominant local plant species. Enzyme activities and community level physiological profiles (CLPP) were closely correlated to microbial biomass and overall CO₂ production in the agricultural soil but not the forest soil, suggesting contrasting relationships between microbial population dynamics and activity in the two soils. Activities of enzymes that break down complex biopolymers, such as protease, cellulase and phenol oxidase were similar or higher in the agricultural soil, which suggests that the production of extracellular biopolymer-degrading enzymes was not a factor limiting litter decomposition. Enzyme and CLPP analyses produced contrasting profiles of microbial activity in the two soils; however the combination of both analyses offers additional insights into the changes in microbial function and community dynamics that occur after conversion of forest to agricultural land.     

Acquisition of microbial communities and enhanced availability of soil nutrients by the isopod Porcellio scaber (Latr.) (Isopoda: Oniscidea)

 In microcosm experiments Porcellio scaber increased litter disappearance of oak and alder litter. Alder litter disappeared at more than twice the rate of oak litter. Soil texture did not influence the disappearance of oak litter; however, disappearance of alder litter was enhanced on silt rather than on sand. P. scaber enhanced microbial communities (i.e. microbial respiration and microbial biomass) on both silt and sand when feeding on either alder or oak. Overall, microbial respiration increased 10-fold when isopods fed on oak litter on sand and 20-fold when isopods fed on alder litter on sand. On silt, the initially high microbial respiration remained constant when isopods fed on oak and doubled when they fed on alder litter. In all treatments without P. scaber there was a decrease in microbial respiration over the 12-week experimental period. The availability of macronutrients (C_org, N_tot, P, K, Mg, Ca) in the topsoil was increased when P. scaber fed on alder litter but less pronounced when P. scaber fed on oak litter. Using sand as a substrate, there was an apparent increase only for C_org, Mg and Ca; on silt, increases in C_org, N_tot and P were measured. Under field conditions the contribution of P. scaber to nutrient fluxes will be higher on sand than on silt. Received: 1 July 1999     

Influence of vegetation types and soil properties on microbial biomass carbon and metabolic quotients in temperate volcanic and tropical forest soils

Abstract

There is limited knowledge about the differences in carbon availability and metabolic quotients in temperate volcanic and tropical forest soils, and associated key influencing factors. Forest soils at various depths were sampled under a tropical rainforest and adjacent tea garden after clear-cutting, and under three temperate forests developed on a volcanic soil (e.g. Betula ermanii and Picea jezoensis, and Pinus koraiensis mainly mixed with Tilia amurensis, Fraxinus mandshurica and Quercus mongolica), to study soil microbial biomass carbon (MBC) concentration and metabolic quotients (qCO₂, CO₂-C/biomass-C). Soil MBC concentration and CO₂ evolution were measured over 7-day and 21-day incubation periods, respectively, along with the main properties of the soils. On the basis of soil total C, both CO₂ evolution and MBC concentrations appeared to decrease with increasing soil depth. There was a maximal qCO₂ in the 0–2.5 cm soil under each forest stand. Neither incubation period affected the CO₂ evolution rates, but incubation period did induce a significant difference in MBC concentration and qCO₂ in tea soil and Picea jezoensis forest soil. The conversion of a tropical rainforest to a tea garden reduced the CO₂ evolution and increased the qCO₂ in soil. Comparing temperate and tropical forests, the results show that both Pinus koraiensis mixed with hardwoods and rainforest soil at less than 20 cm depth had a larger MBC concentration relative to soil total C and a lower qCO₂ during both incubation periods, suggesting that microbial communities in both soils were more efficient in carbon use than communities in the other soils. Factor and regression analysis indicated that the 85% variation of the qCO₂ in forest soils could be explained by soil properties such as the C:N ratio and the concentration of water soluble organic C and exchangeable Al (P < 0.001). The qCO₂ values in forest soils, particularly in temperate volcanic forest soils, decreased with an increasing Al/C ratio in water-soluble organic matter. Soil properties, such as exchangeable Ca, Mg and Al and water-soluble organic C:N ratio, were associated with the variation of MBC. Thus, MBC concentrations and qCO₂ of the soils are useful soil parameters for studying soil C availability and microbial utilization efficiency under temperate and tropical forests.     

莲花湖库区几种主要林型枯落物层的持水性能

 研究莲花湖库区红松林、兴安落叶松林、杂木林等几种主要林型和荒草地枯落物储量及其持水特征,结果表明:兴安落叶松林、红松林、杂木林和荒草地枯落物储量分别为20.8、23.9、16.4和5.7t/hm2,以针叶林的枯落物储量最高。兴安落叶松林、红松林、杂木林和荒草地枯落物层最大持水率分别为278.2%、295.3%、260.3%和154.8%。各林型枯落物吸水速度在0.25～3h变化最快,24h达到饱和状态。枯落物持水速度与浸水历时之间呈显著幂函数相关关系。兴安落叶松林、红松林、杂木林和荒草地枯落物对降雨有效拦蓄率依次为53.87%、65.95%、54.50%和53.45%,对降雨有效拦蓄量变化依次为31.20、45.97、23.27和2.94t/hm2。莲花湖库区不同植被类型下枯落物持水能力为红松林最好,荒草地最差。     

Significant changes in soil microfauna in grazed pasture under elevated carbon dioxide

Soil microfauna in 0- to 10-cm soil under grazed pasture on a sand (Mollic Psammaquent) was assessed quarterly in free air CO₂ enrichment (FACE) rings that were at either ambient CO₂ or had been exposed to 475 l l^–1 CO₂ for 4–5 years. There were significant increases in nematode (1.5×) and rotifer (4.1×) abundance in soils subjected to elevated CO₂. Ten nematode taxa were significantly more abundant under elevated CO₂. The greatest increase was 4.3× in root-feeding Longidorus; three other root-feeders showed no increase in population densities at elevated CO₂. Bacterial-feeding Cervidellus was the only nematode with a significant decrease (0.4×). The abundance of all nematode feeding groups increased significantly in soils subjected to elevated CO₂. The relative increases in abundance of feeding groups (bacterial-feeders 1.3×, root-feeders 1.3×, plant-associated 1.5×, fungal-feeders 1.6×, omnivores 2.0×, predators 2.1×) suggest marked increases in fluxes through microbial-feeding nematodes and a multitrophic response among the soil biota to the increase in atmospheric CO₂ above ambient. Data from the site suggest soil microbial biomass C and N pools were unchanged over the sampling period. Of eight nematode indices only total maturity index increased (2.9 to 3.2), reflecting the increased proportion of the large Longidorus. Further work on microbial-microfaunal interactions and their micro-scale relation to roots is needed to better understand the impact of increasing atmospheric CO₂ on soil processes.     

Long term CO2 enrichment stimulates N-mineralisation and enzyme activities in calcareous grassland

Elevated concentration of atmospheric carbon dioxide will affect carbon cycling in terrestrial ecosystems. Possible effects include increased carbon input into the soil through the rhizosphere, altered nutrient concentrations of plant litter and altered soil moisture. Consequently, the ongoing rise in atmospheric carbon dioxide might indirectly influence soil biota, decomposition and nutrient transformations.N-mineralisation and activities of the enzymes invertase, xylanase, urease, protease, arylsulfatase, and alkaline phosphatase were investigated in spring and summer in calcareous grassland, which had been exposed to ambient and elevated CO₂ concentrations (365 and 600 μl l⁻¹) for six growing seasons.In spring, N-mineralisation increased significantly by 30% at elevated CO₂, while there was no significant difference between treatments in summer (+3%). The response of soil enzymes to CO₂ enrichment was also more pronounced in spring, when alkaline phosphatase and urease activities were increased most strongly by 32 and 21%. In summer, differences of activities between CO₂ treatments were greatest in the case of urease and protease (+21 and +17% at elevated CO₂).The stimulation of N-mineralisation and enzyme activities at elevated CO₂ was probably caused by higher soil moisture and/or increased root biomass. We conclude that elevated CO₂ will enhance below-ground C- and N-cycling in grasslands.     

Relative importance of substrate type and previous soil management in synthesis of microbial biomass and substrate mineralization

Our aim was to determine whether the soil microbial biomass, which has developed naturally over many years in a given ecosystem, is specially adapted to metabolize the plant‐derived substrate C of the ecosystem within which it developed or whether the nature of recently added substrate is the more important factor. To examine this, soils from three sites in close proximity (woodland, grassland and arable from the Broadbalk Experiment at Rothamsted Research, Harpenden, UK) were each amended with air‐dried wheat straw (Triticum aestivum), ryegrass leaves (Lolium perenne) or woodland leaf litter (mainly Quercus robur and Fagus sylvatica) in a fully replicated 3 × 3 factorial laboratory experiment. The initial mineralization rates (evolved CO₂‐C) were determined during the first 6.5 hours and again, together with the amount of microbial biomass synthesized (microbial biomass C), at 7, 14, 21, 30 and 49 days of incubation. The hourly rate of CO₂‐C production during the first 6.5 hours was slowest following leaf litter addition, while the added grass gave the fastest rates of CO₂‐C evolution both within and between soils. Ryegrass addition to the arable soil led to approximately four times more CO₂‐C being evolved than when it was added to the woodland soil, at an overall rate in the arable soils of 41 μg C g^?1 soil hour^?1. In each soil, the net amounts of CO₂‐C produced were in the order grass > straw > leaf litter. In each case, the amount produced by the added leaf litter was significantly less (P < 0.05) than either the added grass or straw. Overall, the trend was for much slower rates of mineralization of all substrates in the woodland soil than in either the arable or grassland soils. During 49 days of incubation in the woodland and grassland soils, the net total amounts of CO₂‐C evolved differed significantly (P < 0.01), with grass > straw > leaf litter, respectively. In the arable soil, the amounts of CO₂‐C evolved from added grass and straw were significantly larger (P < 0.01) than from the leaf litter treatment. Our findings indicated that the amounts of CO₂‐C evolved were not related to soil management or to the size of the original biomass but to the substrate type. The amount of biomass C synthesized was also in the order grass > straw > leaf litter, at all stages of incubation in the woodland and grassland soil. In the arable soil, the same effect was observed up to 14 days, and for the rest of the incubation the biomass C synthesized was in the order grass > straw > leaf litter. Up to three times more biomass C was synthesized from the added grass than from the other substrates in all soils throughout the incubation. The maximum biomass synthesis efficiency was obtained with grass (7% of added C). Overall, the woodland soil was most efficient at synthesizing biomass C and the arable soil the least. We conclude that substrate type was the overriding factor that determined the amount of new soil microbial biomass synthesized. Mineralization of substrate C by soil microorganisms was also influenced mainly by substrate type and less by soil management or size of original biomass.     

Palatability trials on hardwood leaf litter grown under elevated CO2: a stable carbon isotope study

The palatability to isopods and microbes of a broad range of hardwood leaf litter, derived from three field CO₂-enrichment experiments in the USA, was investigated, using δ¹³C, to trace the C flow from litter to isopods and to CO₂ respired by microbial decomposition. Leaf litter grown under elevated CO₂ had δ¹³C values ranging from −39 to −45‰, which were significantly different from ambient litter δ¹³C values of around −30‰. Litter palatability to isopods of the Porcellio sp. was tested by incubating ambient- and elevated-CO₂ litter, and a mixture of the two, in the presence of isopods for 14 days, under environmentally controlled conditions; δ¹³C was measured on litter and isopods' body before and after incubation. In an additional experiment, litter was incubated in the absence of fauna for 30 days, and on five occasions the δ¹³C of the CO₂ respired from litter was measured. The ¹³C label was clearly carried from the litter source to the isopods' bodies, and their faeces. For microbial-respired CO₂, δ¹³C was significantly higher than that of the litter source, suggesting preferential degradation of substrates enriched in ¹³C as compared to those in the overall litter. With the exception of Quercus myrtifolia leaf litter, elevated CO₂ did not affect the palatability to isopods nor the microbial degradation of any of the litters, possibly as a result of unaltered litter N concentration. However, significant differences in litter palatability and decay rates were observed among the different species. With this study, the use of isotopically labelled litter material was confirmed as a key methodology that can significantly contribute to the advancement of the understanding of litter decomposition and of the quantification of C fluxes in the process.     

Elevated CO2 differentially alters belowground plant and soil microbial community structure in reed canary grass-invaded experimental wetlands

Several recent studies have indicated that an enriched atmosphere of carbon dioxide (CO₂) could exacerbate the intensity of plant invasions within natural ecosystems, but little is known of how rising CO₂ impacts the belowground characteristics of these invaded systems. In this study, we examined the effects of elevated CO₂ and nitrogen (N) inputs on plant and soil microbial community characteristics of plant communities invaded by reed canary grass, Phalaris arundinacea L. We grew the invasive grass under two levels of invasion: the invader was either dominant (high invasion) at >90% plant cover or sub-dominant (low invasion) at <50% plant cover. Experimental wetland communities were grown for four months in greenhouses that received either 600 or 365 μl l⁻¹ (ambient) CO₂. Within each of three replicate rooms per CO₂ treatment, the plant communities were grown under high (30 mg l⁻¹) or low (5 mg l⁻¹) N. In contrast to what is often predicted under N limitation, we found that elevated CO₂ increased native graminoid biomass at low N, but not at high N. The aboveground biomass of reed canary grass did not respond to elevated CO₂, despite it being a fast-growing C3 species. Although elevated CO₂ had no impact on the plant biomass of heavily invaded communities, the relative abundance of several soil microbial indicators increased. In contrast, the moderately invaded plant communities displayed increased total root biomass under elevated CO_2, while little impact occurred on the relative abundance of soil microbial indicators. Principal components analysis indicated that overall soil microbial community structure was distinct by CO₂ level for the varying N and invasion treatments. This study demonstrates that even when elevated CO₂ does not have visible effects on aboveground plant biomass, it can have large impacts belowground.     

Plant and soil microbial biomasses in Agrostis capillaris and Lathyrus pratensis monocultures exposed to elevated O3 and CO2 for three growing seasons

Elevated CO₂ usually promotes plant growth, whereas elevated O₃ often has a negative effect, especially on root biomass. Thus both these gases can indirectly affect the soil microbial community. We exposed Agrostis capillaris and Lathyrus pratensis to realistic levels of O₃ (40-50 ppb) and CO₂ (ambient air + 100 ppm) in open-top chambers during 2002-2004. The experiment shows negative effects of both O₃ and CO₂, especially on the bulk soil of L. pratensis, in terms of the decreased biomasses of total (25% and 31%), actinobacterial (29% and 31%), bacterial (26% and 33%) and mycorrhizal (AM fungal) (31% and 35%) indicator subgroups, analysed by the PLFA (phospholipid fatty acid) method. The fungal:bacterial PLFA biomass ratio decreased in the bulk soil of A. capillaris, especially with elevated CO₂ alone (38%). These longer-term changes are considered to arise mainly from differences between the plant functional types (i.e. grass cf. N₂-fixing legume) in litter quality and soil C:N ratio. The results also point to interactions and multi-trophic feedbacks between elevated O₃, plant, parasitic rust fungi and soil readily available P, accompanied by a shift in N balance in favour of plants rather than soil microorganisms.