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AbstractIn a dry heathland ecosystem we manipulated temperature (warming), precipitation (drought) and atmospheric concentration of CO2in a full-factorial experiment in order to investigate changes in below-ground biodiversity as a result of future climate change. We investigated the responses in community diversity of nematodes, enchytraeids, collembolans and oribatid mites at two and eight years of manipulations. We used a structural equation modelling (SEM) approach analyzing the three manipulations, soil moisture and temperature, and seven soil biological and chemical variables. The analysis revealed a persistent and positive effect of elevated CO2on litter C:N ratio. After two years of treatment, the fungi to bacteria ratio was increased by warming, and the diversities within oribatid mites, collembolans and nematode groups were all affected by elevated CO2mediated through increased litter C:N ratio. After eight years of treatment, however, the CO2-increased litter C:N ratio did not influence the diversity in any of the four fauna groups. The number of significant correlations between treatments, food source quality, and soil biota diversities was reduced from six to three after two and eight years, respectively. These results suggest a remarkable resilience within the soil biota against global climate change treatments in the long term.

Differences in bacterial community composition (BCC) between bulk and rhizosphere soil and between rhizospheres of different plant species are assumed to be strongly governed by quantitative and qualitative rhizodeposit differences. However, data on the relationship between rhizodeposit amounts and BCC are lacking. Other soil microorganisms, e.g. arbuscular mycorrhizal fungi (AMF), may also influence BCC. We simulated foliar herbivory (cutting) to reduce belowground carbon allocation and rhizodeposition of pea plants grown either with or without AMF. This reduced soil respiration, rhizosphere microbial biomass and bacteriovorous protozoan abundance, whereas none of these were affected by AMF. After labelling plants with (13)CO(2), root and rhizosphere soil (13)C enrichment of cut plants were reduced to a higher extent (24-46%) than shoot (13)C enrichment (10-24%). AMF did not affect (13)C enrichment. Despite these clear indications of reduced rhizosphere carbon-input, denaturing gradient gel electrophoresis (DGGE) of 16S rRNA genes PCR-amplified targeting DNA and RNA from rhizosphere soil did not reveal any effects of cutting on banding patterns. In contrast, AMF induced consistent differences in both DNA- and RNA-based DGGE profiles. These results show that a reduction in rhizosphere microbial activity is not necessarily accompanied by changes in BCC, whereas AMF presence inhibits proliferation of some bacterial taxa while stimulating others.

AbstractElevated atmospheric CO2 concentration and climate change may substantially alter soil carbon (C) dynamics, which in turn may impact future climate through feedback cycles. However, only very few field experiments worldwide have combined elevated CO2 (eCO2) with both warming and changes in precipitation in order to study the potential combined effects of changes in these fundamental drivers of C cycling in ecosystems. We exposed a temperate heath/grassland to eCO2, warming, and drought, in all combinations for 8 years. At the end of the study, soil C stocks were on average 0.927 kg C/m2 higher across all treatment combinations with eCO2 compared to ambient CO2 treatments (equal to an increase of 0.120 ± 0.043 kg C m−2 year−1), and showed no sign of slowed accumulation over time. However, if observed pretreatment differences in soil C are taken into account, the annual rate of increase caused by eCO2 may be as high as 0.177 ± 0.070 kg C m−2 year−1. Furthermore, the response to eCO2 was not affected by simultaneous exposure to warming and drought. The robust increase in soil C under eCO2 observed here, even when combined with other climate change factors, suggests that there is continued and strong potential for enhanced soil carbon sequestration in some ecosystems to mitigate increasing atmospheric CO2 concentrations under future climate conditions. The feedback between land C and climate remains one of the largest sources of uncertainty in future climate projections, yet experimental data under simulated future climate, and especially including combined changes, are still scarce. Globally coordinated and distributed experiments with long‐term measurements of changes in soil C in response to the three major climate change‐related global changes, eCO2, warming, and changes in precipitation patterns, are, therefore, urgently needed.

It is vital to understand responses of soil microorganisms to predicted climate changes, as these directly control soil carbon (C) dynamics. The rate of turnover of soil organic carbon is mediated by soil microorganisms whose activity may be affected by climate change. After one year of multifactorial climate change treatments, at an undisturbed temperate heathland, soil microbial community dynamics were investigated by injection of a very small concentration (5.12 µg C g(-1) soil) of (13)C-labeled glycine ((13)C2, 99 atom %) to soils in situ. Plots were treated with elevated temperature (+1°C, T), summer drought (D) and elevated atmospheric carbon dioxide (510 ppm [CO2]), as well as combined treatments (TD, TCO2, DCO2 and TDCO2). The (13)C enrichment of respired CO2 and of phospholipid fatty acids (PLFAs) was determined after 24 h. (13)C-glycine incorporation into the biomarker PLFAs for specific microbial groups (Gram positive bacteria, Gram negative bacteria, actinobacteria and fungi) was quantified using gas chromatography-combustion-stable isotope ratio mass spectrometry (GC-C-IRMS). Gram positive bacteria opportunistically utilized the freshly added glycine substrate, i.e. incorporated (13)C in all treatments, whereas fungi had minor or no glycine derived (13)C-enrichment, hence slowly reacting to a new substrate. The effects of elevated CO2 did suggest increased direct incorporation of glycine in microbial biomass, in particular in G(+) bacteria, in an ecosystem subjected to elevated CO2. Warming decreased the concentration of PLFAs in general. The FACE CO2 was (13)C-depleted (δ(13)C = 12.2‰) compared to ambient (δ(13)C = ∼-8‰), and this enabled observation of the integrated longer term responses of soil microorganisms to the FACE over one year. All together, the bacterial (and not fungal) utilization of glycine indicates substrate preference and resource partitioning in the microbial community, and therefore suggests a diversified response pattern to future changes in substrate availability and climatic factors.

Terrestrial ecosystems are exposed to atmospheric and climatic changes including increases in atmospheric CO2 concentration, temperature and alterations of precipitation patterns, which are predicted to continue with consequences for ecosystem services and functioning in the future. In a field scale experiment on temperate heathland, manipulation of precipitation and temperature was performed with retractable curtains, and atmospheric CO2 concentration was increased by FACE. The combination of elevated CO2 and warming was expected to affect belowground processes additively, through increased belowground sequestration of labile carbohydrates due to elevated CO2 in combination with temperature increased process rates. Together, these changes might increase microbial activity and availability of plant nutrients. Two years after the start of the experiment, belowground processes responded significantly to the treatments. In the combined temperature and CO2 treatment the dissolved organic nitrogen concentration decreased and the ammonium concentration increased, but this release of nutrients was not mirrored by plant parameters. Microbial biomass carbon and microbial enrichment with 13C and 15N (1 year after 13C 2 15 N-glycine was injected into the soil) increased in warmed plots and in elevated CO2 plots, but not when these treatments were combined. Furthermore, drought led to an increase in Calluna biomass and total plant nitrogen pool. The full combination of warming, elevated CO2 and periodic drought did not unambiguously express the ecosystem responses of single factors additively, which complicates predictions of ecosystem responses to multifactor climate change.

AbstractNitrogen (N) dynamic is one of the main controlling factors of responses to climate change in N‐limited terrestrial ecosystems, which rely on nutrient recycling and retention. In this study we investigate the N partitioning in ecosystem compartments of a grassland heath, and the impact of multiple climate change factors on long‐term N retention after 15N pulse labelling. The impacts of elevated carbon dioxide (eCO2), warming and drought and the treatments in combination on ecosystem N retention were investigated in a field scale manipulation experiment. A 6‐year time‐course was assessed by pulse‐labelling with the stable N isotope 15N and by sampling after 1 day, 1 year and 6 years. After 6 years we observed that the total ecosystem retained 42% of the amended 15N across treatments (recovery of the amended 15N in the pool). The fate of the applied 15N was mainly stabilization in soil, with 36% 15N recovery in soil, while the plant compartment and microbial biomass each retained only 1%–2% of the added 15N. This suggests a moderate retention of N, for all treatments, as compared to similar long‐term studies of forest ecosystems. A decreased ammonium and vegetation N pool combined with higher 15N retention in the soil under eCO2 treatments suggests that eCO2 promoted processes that immobilize N in soil, while warming counteracted this when combined with eCO2. Drought treatments contrastingly increased the vegetation N pool. We conclude that as the organic soil layer has the main capacity for N storage in a temperate heathland‐grassland, it is important for buffering nutrient availability and maintaining a resilient ecosystem. However, the full treatment combination of drought, warming and eCO2 did not differ in 15N recovery from the control, suggesting unchanged long‐term consequences of climate change on retention of pulse added N in this ecosystem.

Climatic warming leads to the expansion of deciduous shrubs and trees in the Arctic. This leads to higher leaf litter inputs, which together with warming may alter the rate of carbon and nutrient cycling in the arctic ecosystems. We assessed effects of factorial warming and additional litter on the soil ecosystem of a subarctic heath in a 7-year-long field experiment. Fine root biomass, dissolved organic carbon (DOC) and total C concentration increased in response to warming, which probably was a result of the increased vegetation cover. Litter addition increased the concentration of inorganic P in the uppermost 5 cm soil, while decreasing the pool of total P per unit area of the organic profile and having no significant effects on N concentrations or pools. Microbial biomass C and N were unaffected by the treatments, while the microbial biomass P increased significantly with litter addition. Soil ergosterol concentration was also slightly increased by the added litter in the uppermost soil, although not statistically significantly. According to a principal component analysis of the phospholipid fatty acid profiles, litter addition differed from the other treatments by increasing the relative proportion of biomarkers for Gram-positive bacteria. The combined warming plus litter addition treatment decreased the soil water content in the uppermost 5 cm soil, which was a likely reason for many interactions between the effects of warming and litter addition. The soil organic matter quality of the combined treatment was also clearly different from the control based on a near-infrared reflectance (NIR) spectroscopic analysis, implying that the treatment altered the composition of soil organic matter. However, it appears that the biological processes and the microbial community composition responded more to the soil and litter moisture conditions than to the change in the quality of the organic matter.