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Antibiotics and temperature interact to disrupt soil communities and nutrient cycling

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Abstract

Soils contain immense diversity and support terrestrial ecosystem functions, but they face both anthropogenic and environmental stressors. While many studies have examined the influence of individual stressors on soils, how these perturbations will interact to shape soil communities and their ability to cycle nutrients is far less resolved. Here, we hypothesized that when soils experience multiple stressors their ability to maintain connected and stable communities is disrupted, leading to shifts in C and N pools. To test this, we maintained soils across three temperatures representative of seasonal variability (15, 20 and 30 °C) and introduced high or low doses of the common livestock antibiotic Monensin. We monitored respiration and examined changes to microbial communities through amplicon sequencing and network analyses. We also examined soil C and N pools to understand how temperature and antibiotics shape ecosystem function. We found that antibiotics and rising soil temperatures interacted to disrupt bacterial assemblages and network structure, allowing for a rise in fungal dominance and change in soil nutrient stoichiometry. Antibiotics alone decreased bacterial diversity, abundance, total extractable N, and microbial carbon use efficiency, while increasing bioavailable C. Higher temperatures independently homogenized fungal community composition, decreased dissolved organic C and increased soil respiration rates. These results emphasize that as soils encounter multiple stressors, ecosystem efficiency, stability and resilience may be diminished.

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Tetracyclines are antibiotics commonly used in swine farms to treat disease and promote growth. However, there are growing concerns regarding the discharge of animal feces into the environment owing to the potential for development and dissemination of tetracycline resistance genes (TRGs). In this study, farming wastewater from one Chinese swine farm as well as river water from seven locations downstream of the farm was sampled. Polymerase chain reaction (PCR) showed that 12 TRGs, including six efflux pump genes (tet(B), tet(C), tet(D), tet(E), tet(G) and tet(L)), five ribosomal protection proteins (RPPs) genes (tet(O), tet(M), tet(Q), tet(W) and tet(S)), and one enzymatic modification gene (tet(X)), were present in all wastewater and river water samples. Quantitative real-time PCR (qPCR) showed that the abundance of tet(C), tet(X), tet(O), tet(M), tet(Q) and tet(W) decreased with downstream flow. Among the detected TRGs, tet(C) had the highest abundance, ranging from 459.5 copies/16S rRNA gene copies in wastewater to 33.8 copies/16S rRNA gene copies in river water samples collected from the last location. Furthermore, pig-specific Bacteroidales 16S rRNA genetic marker was quantified by qPCR to determine the level of fecal pollution in the river water. Bivariate correlation analysis confirmed that the total relative abundance of the six TRGs was significantly correlated with the level of swine feces in the aquatic environment (R(2) = 0.63, P < 0.05), suggesting that swine feces mainly contributed to the spread of TRGs in the river water. Supplemental materials are available for this article. Go to the publisher's online edition of Journal of Environmental Science and Health, Part A to view the free supplemental file.
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Interspecies interaction is an essential mechanism for bacterial communities to develop antibiotic resistance via horizontal gene transfer. Nonetheless, how bacterial interactions vary along the environmental transmission of antibiotics and the underpinnings remain unclear. To address it, we explore potential microbial associations by analyzing bacterial networks generated from 16S rRNA gene sequences and functional networks containing a large number of antibiotic-resistance genes (ARGs). Antibiotic concentration decreased by more than 4000-fold along the environmental transmission chain from manure samples of swine farms to aerobic compost, compost-amended agricultural soils, and neighboring agricultural soils. Both bacterial and functional networks became larger in nodes and links with decreasing antibiotic concentrations, likely resulting from lower antibiotics stress. Nonetheless, bacterial networks became less clustered with decreasing antibiotic concentrations, while functional networks became more clustered. Modularity, a key topological property that enhances system resilience to antibiotic stress, remained high for functional networks, but the modularity values of bacterial networks were the lowest when antibiotic concentrations were intermediate. To explain it, we identified a clear shift from deterministic processes, particularly variable selection, to stochastic processes at intermediate antibiotic concentrations as the dominant mechanism in shaping bacterial communities. Collectively, our results revealed microbial network dynamics and suggest that the modularity value of association networks could serve as an important indicator of antibiotic concentrations in the environment.
Article
Biocides have been frequently used to understand the roles of fungi and bacteria on soil carbon (C) and nitrogen (N) cycling. However, addition of biocides to soil can result in unwanted temporary increases in C and N supply to surviving microbes due to a pulse in microbial necromass, and where biocides can directly be used as sources of C and N. We assessed temporary dynamics in microbial biomass C (MBC) and N (MBN), soil respiration and gross N mineralization (GNM) during a 22-day laboratory incubation after addition of fungicides (captan, cycloheximide) and bactericides (bronopol, oxytetracycline) in a grassland soil. We also assessed whether captan and bronopol were a C source for remaining microbes based on δ 13 C measurements in respired CO 2. As expected, fungicides decreased and bactericides increased fungi:bacteria ratios, while all biocides decreased MBC and MBN. However, respiration (except for oxytetracycline) and GNM (particularly for bronopol) increased after biocide addition, likely because of a pulse in necromass causing increased substrate supply to the surviving microbes. We further detected biocide-derived CO 2 up to 10 days for captan, and 22 days for bronopol, suggesting that they can be an important source for C by surviving microbes. However, soil-derived CO 2 remained higher without biocide, indicating that necromass was the most important source for higher soil respiration after biocide addition. We recommend repeated additions of biocides to suppress regrowth of target microbes, and use of 13 C and 15 N labeled substrates, when studying effects of fungi and bacteria on C and N dynamics.
Article
Regions and localities may lose many species to extinction under rapid climate change and may gain other species that colonize from nearby warmer environments. Here, it is argued that warming-induced species losses will generally exceed gains and there will be more net declines than net increases in plant community richness. Declines in richness are especially likely in water-limited climates where intensifying aridity will increasingly exceed plant tolerances, but also in colder temperature-limited climates where steep climatic gradients are lacking, and therefore, large pools of appropriate species are not immediately adjacent. The selectivity of warming-induced losses may lead to declines in functional and phylogenetic diversity as well as in species richness, especially in water-limited climates. Our current understanding of climate-caused diversity trends may be overly influenced by numerous studies coming from north-temperate alpine mountaintops, where conditions are unusually favourable for increases—possibly temporary—in local species richness. This article is part of the theme issue ‘Climate change and ecosystems: threats, opportunities and solutions’.
Article
Soils underpin terrestrial ecosystem functions, but they face numerous anthropogenic pressures. Despite their crucial ecological role, we know little about how soils react to more than two environmental factors at a time. Here, we show experimentally that increasing the number of simultaneous global change factors (up to 10) caused increasing directional changes in soil properties, soil processes, and microbial communities, though there was greater uncertainty in predicting the magnitude of change. Our study provides a blueprint for addressing multifactor change with an efficient, broadly applicable experimental design for studying the impacts of global environmental change.
Article
The soil microbiome governs biogeochemical cycling of macronutrients, micronutrients and other elements vital for the growth of plants and animal life. Understanding and predicting the impact of climate change on soil microbiomes and the ecosystem services they provide present a grand challenge and major opportunity as we direct our research efforts towards one of the most pressing problems facing our planet. In this Review, we explore the current state of knowledge about the impacts of climate change on soil microorganisms in different climate-sensitive soil ecosystems, as well as potential ways that soil microorganisms can be harnessed to help mitigate the negative consequences of climate change.
Article
Predictions of the effects of global change on ecological communities are largely based on single habitats. Yet in nature, habitats are interconnected through the exchange of energy and organisms, and the responses of local communities may not extend to emerging community networks (i.e. metacommunities). Using large mesocosms and meiofauna communities as a model system, we investigated the interactive effects of ocean warming and acidification on the structure of marine metacommunities from three shallow‐water habitats: sandy soft‐bottoms, marine vegetation and rocky reef substrates. Primary producers and detritus – key food sources for meiofauna – increased in biomass under the combined effect of temperature and acidification. The enhanced bottom‐up forcing boosted nematode densities but impoverished the functional and trophic diversity of nematode metacommunities. The combined climate stressors further homogenized meiofauna communities across habitats. Under present‐day conditions metacommunities were structured by habitat type, but under future conditions they showed an unstructured random pattern with fast‐growing generalist species dominating the communities of all habitats. Homogenization was likely driven by local species extinctions, reducing interspecific competition that otherwise could have prevented single species from dominating multiple niches. Our findings reveal that climate change may simplify metacommunity structure and prompt biodiversity loss, which may affect the biological organization and resilience of marine communities.
Article
We examined the impacts of soil moisture, temperature, agricultural management, and habitat type on the degradation dynamics of eDNA in soils. Synthetic eDNA was added to soil microcosms, and its disappearance over time was measured using both high‐throughput sequencing and qPCR. The synthetic eDNA was degraded rapidly, but a small fraction remained detectable throughout the experiments (39‐80 days). The eDNA degradation rate was positively correlated with moisture and temperature, but negatively correlated with soil organic carbon content. End‐point stabilization of eDNA was highest at low moisture and temperature, but exhibited no relationship with soil organic carbon. Tilled soils had higher rates of degradation and less stabilization than no‐till soils. Among different habitats we observed that forest soils had the slowest degradation rate, and meadow soils had the greatest stabilization of eDNA. While eDNA was detectable by qPCR in all treatments across all time‐points, it became inconsistently detectable with high‐throughput gene sequencing in less than one week. We conclude that eDNA degradation and stabilization dynamics vary with moisture, temperature, and habitat characteristics, that small amounts of eDNA may persist in soils indefinitely, and that the ability of persistent eDNA to impact microbial community estimates depends on method sensitivity and experimental objectives. This article is protected by copyright. All rights reserved.
Article
Models representing exchange of carbon between the atmosphere and the terrestrial biosphere include a large variety of processes and mechanisms, and have increased in complexity in the last decades. These models are no exception of the simulation versus understanding conundrum previously articulated for models of the physical climate, which states that increasing detail in process representation in models, and the simulations they produce, hinders understanding of holistic system behavior. However, recent theoretical progress on the mathematical representation of the carbon cycle in ecosystems may help to provide a general framework for the qualitative understanding of models without compromising detail in process representation. Here we (1) briefly review recent ideas on the theory of transient dynamics of the terrestrial carbon cycle and its matrix representation, pointing out issues of interpretation, (2) show that these ideas can be further generalized in the mathematical concept of nonautonomous compartmental systems, and (3) provide thoughts on how this framework can be used to address a new set of questions in carbon cycle science.
Article
The relationship between the spatial variability of soil multifunctionality (i.e. the capacity of soils to conduct multiple functions; SVM) and major climatic drivers, such as temperature and aridity, has never been assessed globally in terrestrial ecosystems. We surveyed 236 dryland ecosystems from six continents to evaluate the relative importance of aridity and mean annual temperature, and of other abiotic (e.g., texture) and biotic (e.g., plant cover) variables as drivers of SVM, calculated as the averaged coefficient of variation for multiple soil variables linked to nutrient stocks and cycling. We found that increases in temperature and aridity were globally correlated to increases in SVM. Some of these climatic effects on SVM were direct, but others were indirectly driven through reductions in the number of vegetation patches and increases in soil sand content. The predictive capacity of our structural equation modelling was clearly higher for the spatial variability of N- than for C- and P- related soil variables. In the case of N cycling, the effects of temperature and aridity were both direct and indirect via changes in soil properties. For C and P, the effect of climate was mainly indirect via changes in plant attributes. These results suggest that future changes in climate may decouple the spatial availability of these elements for plants and microbes in dryland soils. Our findings significantly advance our understanding of the patterns and mechanisms driving SVM in drylands across the globe, which is critical for predicting changes in ecosystem functioning in response to climate change. Key words: Multifunctionality; carbon cycling; nitrogen cycling; phosphorous cycling; spatial heterogeneity; climate change This article is protected by copyright. All rights reserved.
Article
Among the different pharmaceuticals present in soil and water ecosystems as micro-contaminants, considerable attention has been paid to antibiotics, since their increasing use and the consequent development of multi-resistant bacteria pose serious risks to human and veterinary health. Moreover, once they have entered the environment, antibiotics can affect natural microbial communities. The latter play a key role in fundamental ecological processes, most importantly the maintenance of soil and water quality. In fact, they are involved in biogeochemical cycling and organic contaminant degradation thanks to their large reservoir of genetic diversity and metabolic capability. When antibiotics occur in the environment, they can hamper microbial community structure and functioning in different ways and have both direct (short-term) and indirect (long-term) effects on microbial communities. The short-term ones are bactericide and bacteriostatic actions with a consequent disappearance of some microbial populations and their ecological functioning. The indirect impact includes the development of antibiotic resistant bacteria and in some cases bacterial strains able to degrade them by metabolic or co-metabolic processes. Biodegradation makes it possible to completely remove a toxic compound from the environment if it is mineralized.Several factors can influence the significance of such direct and indirect effects, including the antibiotic's concentration, the exposure time, the receiving ecosystem (e.g. soil or water) and the co-occurrence of other antibiotics and/or other contaminants.This review describes the current state of knowledge regarding the effects of antibiotics on natural microbial communities in soil and water ecosystems.
Article
Manure composting has general benefits for production of soil amendment, but the effects of composting on antibiotic persistence and effects of antibiotics on the composting process are not well-characterized, especially for antibiotics commonly used in dairy cattle. This study provides a comprehensive, head-to-head, replicated comparison of the effect of static and turned composting on typical antibiotics used in beef and dairy cattle in their actual excreted form and corresponding influence on composting efficacy. Manure from steers (with or without chlortetracycline, sulfamethazine, and tylosin feeding) and dairy cows (with or without pirlimycin and cephapirin administration) were composted at small scale (wet mass: 20-22 kg) in triplicate under static and turned conditions adapted to represent US Food and Drug Administration guidelines. Thermophilic temperature (>55°C) was attained and maintained for 3 d in all composts, with no measureable effect of compost method on the pattern, rate, or extent of disappearance of the antibiotics examined, except tylosin. Disappearance of all antibiotics, except pirlimycin, followed bi-phasic first-order kinetics. However, individual antibiotics displayed different fate patterns in response to the treatments. Reduction in concentration of chlortetracycline (71-84%) and tetracycline (66-72%) was substantial, while near-complete removal of sulfamethazine (97-98%) and pirlimycin (100%) was achieved. Tylosin removal during composting was relatively poor. Both static and turned composting were generally effective for reducing most beef and dairy antibiotic residuals excreted in manure, with no apparent negative impact of antibiotics on the composting process, but with some antibiotics apparently more recalcitrant than others.
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During the past decade pharmaceuticals have been recognised as a new class of ubiquitously occurring persistent contaminants (Halling-Sørensen et al. 1998; Daughton and Ternes 1999; Kämmerer 2001; Boxall et al. 2003; Diaz-Cruz et al. 2003; Thiele-Bruhn 2003). A large number of drugs are used extensively in both human and veterinary medicine. Antibiotics are one of the most important substance classes with a possible environmental impact. Estimations of the European Federation of Animal Health (FEDESA) revealed that approximately 8500 tons of antibiotics were used in human medicine and 4700 tons in veterinary medicine in the European Union (including Switzerland) in 1999 (Anonymous 2001).
Article
Fungal community composition often shifts in response to warmer temperatures, which might influence decomposition of recalcitrant carbon (C). We hypothesized that evolutionary trade-offs would enable recalcitrant C-using taxa to respond more positively to warming than would labile C-using taxa. Accordingly, we performed a warming experiment in an Alaskan boreal forest and examined changes in the prevalence of fungal taxa. In a complementary field trial, we characterized the ability of fungal taxa to use labile C (glucose), intermediate C (hemicellulose or cellulose), or recalcitrant C (lignin). We also assigned taxa to functional groups (e.g., free-living filamentous fungi, ectomycorrhizal fungi, and yeasts) based on taxonomic identity. We found that response to warming varied most among taxa at the order level, compared to other taxonomic ranks. Among orders, ability to use lignin was significantly related to increases in prevalence in response to warming. However, the relationship was weak, given that lignin use explained only 9% of the variability in warming responses. Functional groups also differed in warming responses. Specifically, free-living filamentous fungi and ectomycorrhizal fungi responded positively to warming, on average, but yeasts responded negatively. Overall, warming-induced shifts in fungal communities might be accompanied by an increased ability to break down recalcitrant C. This change in potential function may reduce soil C storage under global warming.
Article
The human gut harbors a large and complex community of beneficial microbes that remain stable over long periods. This stability is considered critical for good health but is poorly understood. Here we develop a body of ecological theory to help us understand microbiome stability. Although cooperating networks of microbes can be efficient, we find that they are often unstable. Counterintuitively, this finding indicates that hosts can benefit from microbial competition when this competition dampens cooperative networks and increases stability.More generally, stability is promoted by limiting positive feedbacks and weakening ecological interactions. We have analyzed host mechanisms formaintaining stability - including immune suppression, spatial structuring, and feeding of community members - and support our key predictions with recent data.
Article
Soil organisms are considered drivers of soil ecosystem services (primary productivity, nutrient cycling, carbon cycling, water regulation) associated with sustainable agricultural production. Soil biodiversity was highlighted in the soil thematic strategy as a key component of soil quality. The lack of quantitative standardised data at a large scale has resulted in poor understanding of how soil biodiversity could be incorporated into legislation for the protection of soil quality. In 2011, the EcoFINDERS (FP7) project sampled 76 sites across 11 European countries, covering five biogeographical zones (Alpine, Atlantic, Boreal, Continental and Mediterranean) and three land-uses (arable, grass, forestry). Samples collected from across these sites ranged in soil properties; soil organic carbon (SOC), pH and texture. To assess the range in biodiversity and ecosystem function across the sites, fourteen biological methods were applied as proxy indicators for these functions. These methods measured the following: microbial diversity: DNA yields (molecular biomass), archaea, bacteria, total fungi and arbuscular mycorrhizal fungi; micro fauna diversity: nematode trophic groups; meso fauna diversity: enchytraeids and Collembola species; microbial function: nitrification, extracellular enzymes, multiple substrate induced respiration, community level physiological profiling and ammonia oxidiser/nitrification functional genes. Network analysis was used to identify the key connections between organisms under the different land use scenarios. Highest network density was found in forest soils and lowest density occurred in arable soils. Key taxomonic units (TUs) were identified in each land-use type and in relation to SOC and pH categorisations. Top-connected taxonomic units (i.e. displaying the most co-occurrence to other TUs) were identified for each land use type. In arable sites this was dominated by bacteria and fungi, while in grassland sites bacteria and fungi were most connected. In forest soils archaeal, enchytraeid and fungal TUs displayed the largest number of neighbours, reflecting the greatest connectivity. Multiple regression models were applied to assess the potential contribution of soil organisms to carbon cycling and storage and nutrient cycling of specifically nitrogen and phosphorus. Key drivers of carbon cycling were microbial biomass, basal respiration and fungal richness; these three measures have often been associated with carbon cycling in soils. Regression models of nutrient cycling were dependent on the model applied, showing variation in biological indicators.
Chapter
Veterinary medicines are used widely to treat disease and protect the health of animals. Following administration, veterinary medicines may be metabolised and then released along with any metabolites either directly to the environment (as is the case with pasture animals) or indirectly during the application of manure or slurry.
Article
Resolving fungal and bacterial groups within the microbial decomposer community is thought to capture disparate life strategies for soil microbial decomposers, associating bacteria with an r-selected strategy for carbon (C) and nutrient use, and fungi with a K-selected strategy. Additionally, food-web model-based work has established a widely held belief that the bacterial decomposer pathway in soil supports high turnover rates of easily available substrates, while the slower fungal-dominated decomposition pathway supports the decomposition of more complex organic material, thus characterizing the biogeochemistry of the ecosystem. We used a field experiment, the Detritus Input and Removal Treatments, or DIRT, experiment (Harvard Forest Long-Term Ecological Research Site, USA) where litter and root inputs (control, no litter, double litter, or no tree roots) have been experimentally manipulated during 23 years, generating differences in soil C quality. We hypothesized (1) that d13C enrichment would decrease with higher soil C quality and that a higher C quality would favor bacterial decomposers, (2) that the C mineralized in fungal-dominated treatments would be of lower quality and also depleted in d13C relative to bacterial-dominated high-quality soil C treatments, and (3) that higher C mineralization along with higher gross N mineralization rates would occur in bacterialdominated treatments compared with more fungal-dominated treatments. The DIRT treatments resulted in a gradient of soil C quality, as shown by up to 4.5-fold differences between the respiration per soil C between treatments. High-quality C benefited fungal dominance, in direct contrast with our hypothesis. Further, there was no difference between the d13CO2 produced by a fungal compared with a bacterial-dominated decomposer community. There were differences in C and N mineralization between DIRT treatments, but these were not related to the relative dominance of fungal and bacterial decomposers. Thus we find no support for the hypothesized differences between detrital food webs dominated by bacteria compared to those dominated by fungi. Rather, an association between high-quality soil C and fungi emerges from our results. Consequently there is a need to revise our basic understanding for microbial communities and the processes they regulate in soil.
Article
Biofuels will help meet rising demands for energy and, ideally, limit climate change associated with carbon losses from the biosphere to atmosphere. Biofuel management must therefore maximize energy production and maintain ecosystem carbon stocks. Increasingly, there is interest in intercropping biofuels with other crops, partly because biofuel production on arable land might reduce availability and increase the price of food. One intercropping approach involves growing biofuel grasses in forest plantations. Grasses differ from trees in both their organic inputs to soils and microbial associations. These differences are associated with losses of soil carbon when grasses become abundant in forests. We investigated how intercropping switchgrass (Panicum virgatum), a major candidate for cellulosic biomass production, in loblolly pine (Pinus taeda) plantations affects soil carbon, nitrogen, and microbial dynamics. Our design involved four treatments: two pine management regimes where harvest residues (i.e., biomass) were left in place or removed, and two switchgrass regimes where the grass was grown with pine under the same two biomass scenarios (left or removed). Soil variables were measured in four 1-ha replicate plots in the first and second year following switchgrass planting. Under switchgrass intercropping, pools of mineralizable and particulate organic matter carbon were 42% and 33% lower, respectively. These declines translated into a 21% decrease in total soil carbon in the upper 15 cm of the soil profile, during early stand development. The switchgrass effect, however, was isolated to the interbed region where switchgrass is planted. In these regions, switchgrass-induced reductions in soil carbon pools with 29%, 43%, and 24% declines in mineralizable, particulate, and total soil carbon, respectively. Our results support the idea that grass inputs to forests can prime the activity of soil organic carbon degrading microbes, leading to net reductions in stocks of soil carbon. Active microbial biomass, however, is higher under switchgrass, and this microbial biomass is a dominant precursor of soil carbon formation. Future studies need to investigate soil carbon dynamics throughout the lifetime of intercropping rotations to evaluate whether increases in microbial biomass can offset initial declines in soil carbon, and hence, maintain ecosystem carbon stocks.
Article
One of the key questions in climate change research relates to the future dynamics of the large amount of C that is currently stored in soil organic matter. Will the amount of C in this pool increase or decrease with global warming? The future trend in amounts of soil organic C will depend on the relative temperature sensitivities of net primary productivity and soil organic matter decomposition rate. Equations for the temperature dependence of net primary productivity have been widely used, but the temperature dependence of decomposition rate is less clear. The literature was surveyed to obtain the temperature dependencies of soil respiration and N dynamics reported in different studies. Only laboratory-based measurements were used to avoid confounding effects with differences in litter input rates, litter quality, soil moisture or other environmental factors. A considerable range of values has been reported, with the greatest relative sensitivity of decomposition processes to temperature having been observed at low temperatures. A relationship fitted to the literature data indicated that the rate of decomposition increases with temperature at 0°C with a Q10 of almost 8. The temperature sensitivity of organic matter decomposition decreases with increasing temperature, indicated by the Q10 decreasing with temperature to be about 4.5 at 10°C and 2.5 at 20°C. At low temperatures, the temperature sensitivity of decomposition was consequently much greater than the temperature sensitivity of net primary productivity, whereas the temperature sensitivities became more similar at higher temperatures. The much higher temperature sensitivity of decomposition than for net primary productivity has important implications for the store of soil organic C in the soil. The data suggest that a 1°C increase in temperature could ultimately lead to a loss of over 10% of soil organic C in regions of the world with an annual mean temperature of 5°C, whereas the same temperature increase would lead to a loss of only 3% of soil organic C for a soil at 30°C. These differences are even greater in absolute amounts as cooler soils contain greater amounts of soil organic C. This analysis supports the conclusion of previous studies which indicated that soil organic C contents may decrease greatly with global warming and thereby provide a positive feed-back in the global C cycle.
Article
The future of the land carbon sink is a significant uncertainty in global change projections. Here, key controls on global terrestrial carbon storage are examined using a simple model of vegetation and soil. Equilibrium solutions are derived as a function of atmospheric CO2 and global temperature, these environmental variables are then linked in an idealized global change trajectory, and the lag between the dynamic and equilibrium solutions is derived for different linear rates of increase in atmospheric CO2. Terrestrial carbon storage is departing significantly from equilibrium because CO2 and temperature are increasing on a similar timescale to ecosystem change, and the lag is found to be proportional to the rate of forcing. Thus peak sizes of the land carbon sink, and any future land carbon source, are proportional to the rate of increase of CO2. A switch from a land carbon sink to a source occurs at a higher CO2 and temperature under more rapid forcing. The effects of parameter uncertainty in temperature sensitivities of photosynthesis, plant respiration and soil respiration, and structural uncertainty through the effect of fixing the ratio of plant respiration to photosynthesis are explored. In each case, the CO2 fertilization effect on photosynthesis is constrained to reproduce the 1990 atmospheric CO2 concentration within a closed global model. New literature compilations are presented for the temperature sensitivities of plant and soil respiration. A lower limit, Q10=1.29, for soil respiration significantly increases future land carbon storage. An upper limit, Q10=3.63, for soil respiration underpredicts the increase in carbon storage since the Last Glacial Maximum. Fixing the ratio of plant respiration to photosynthesis (R/P) at 0.5 generates the largest and most persistent land carbon sink, followed by the weakest land carbon source.