A thorough understanding of the role of microbes in C cycling in relation to fire is important for estimation of C emissions and for development of guidelines for sustainable management of dry ...ecosystems. We investigated the seasonal changes and spatial distribution of soil total, dissolved organic C (DOC) and microbial biomass C during 18 months, quantified the soil CO2 emission in the beginning of the rainy season, and related these variables to the fire frequency in important dry vegetation types grassland, woodland and dry forest in Ethiopia. The soil C isotope ratios (δ13C) reflected the 15-fold decrease in the grass biomass along the vegetation gradient and the 12-fold increase in woody biomass in the opposite direction. Changes in δ13C down the soil profiles also suggested that in two of the grass-dominated sites woody plants were more frequent in the past. The soil C stock ranged from being 2.5 (dry forest) to 48 times (grassland) higher than the C stock in the aboveground plant biomass. The influence of fire in frequently burnt wooded grassland was evident as an unchanged or increasing total C content down the soil profile. DOC and microbial biomass measured with the fumigation–extraction method (Cmic) reflected the vertical distribution of soil organic matter (SOM). However, although SOM was stable throughout the year, seasonal fluctuations in Cmic and substrate-induced respiration (SIR) were large. In woodland and woodland–wooded grassland Cmic and SIR increased in the dry season, and gradually decreased during the following rainy season, confirming previous suggestions that microbes may play an important role in nutrient retention in the dry season. However, in dry forest and two wooded grasslands Cmic and SIR was stable throughout the rainy season, or even increased in this period, which could lead to enhanced competition with plants for nutrients. Both the range and the seasonal changes in soil microbial biomass C in dry tropical ecosystems may be wider than previously assumed. Neither SIR nor Cmic were good predictors of in situ soil respiration. The soil respiration was relatively high in infrequently burnt forest and woodland, while frequently burnt grasslands had lower rates, presumably because most C is released through dry season burning and not through decomposition in fire-prone systems. Shifts in the relative importance of the two pathways for C release from organic matter may have strong implications for C and nutrient cycling in seasonally dry tropical ecosystems.
Aims
This study aimed at elucidating divergent effects of two dominant plant functional types (PFTs) in tundra heath, dwarf shrubs and mosses, on soil microbial processes and soil carbon (C) and ...nutrient availability, and thereby to enhance our understanding of the complex interactions between PFTs, soil microbes and soil functioning.
Methods
Samples of organic soil were collected under three dwarf shrub species (of distinct mycorrhizal association and life form) and three moss species in early and late growing season. We analysed soil C and nutrient pools, extracellular enzyme activities and phospholipid fatty acid profiles, together with a range of plant traits, soil and abiotic site characteristics.
Results
Shrub soils were characterised by high microbial biomass C and phosphorus and phosphatase activity, which was linked with a fungal-dominated microbial community, while moss soils were characterised by high soil nitrogen availability, peptidase and peroxidase activity associated with a bacterial-dominated microbial community. The variation in soil microbial community structure was explained by mycorrhizal association, root morphology, litter and soil organic matter quality and soil pH-value. Furthermore, we found that the seasonal variation in microbial biomass and enzyme activities over the growing season, likely driven by plant belowground C allocation, was most pronounced under the tallest shrub
Betula nana
.
Conclusion
Our study demonstrates a close coupling of PFTs with soil microbial communities, microbial decomposition processes and soil nutrient availability in tundra heath, which suggests potential strong impacts of global change-induced shifts in plant community composition on carbon and nutrient cycling in high-latitude ecosystems.
In high‐latitude ecosystems bryophytes are important drivers of ecosystem functions. Alterations in abundance of mosses due to global change may thus strongly influence carbon (C) and nitrogen (N) ...cycling and hence cause feedback on climate. The effects of mosses on soil microbial activity are, however, still poorly understood. Our study aims at elucidating how and by which mechanisms bryophytes influence microbial decomposition processes of soil organic matter and thus soil nutrient availability.
We present results from a field experiment in a subarctic birch forest in northern Sweden, where we partly removed the moss cover and replaced it with an artificial soil cover for simulating moss effects on soil temperature and moisture. We combined this with a fertilization experiment with 15N‐labelled N for analysing the effects of moss N sequestration on soil processes.
Our results demonstrate the capacity of mosses to reduce soil N availability and retard N cycling. The comparison with artificial soil cover plots suggests that the effect of mosses on N cycling is linked to the thermal insulation capacity of mosses causing low average soil temperature in summer and strongly reduced soil temperature fluctuations, the latter also leading to a decreased frequency of freeze‐thaw events in autumn and spring. Our results also showed, however, that the negative temperature effect of mosses on soil microbial activity was in part compensated by stimulatory effects of the moss layer, possibly linked to leaching of labile substrates from the moss. Furthermore, our results revealed that bryophytes efficiently sequester added N from wet deposition and thus prevent effects of increased atmospheric N deposition on soil N availability and soil processes.
Synthesis. Our study emphasizes the important role of mosses in carbon and nutrient cycling in high‐latitude ecosystems and the potential strong impacts of reductions in moss abundance on microbial decomposition processes and nutrient availability in subarctic and boreal forests.
We performed a moss removal/artificial soil cover experiment combined with a nitrogen fertilization experiment in a subarctic birch forest in order to investigate the mechanisms by which mosses influence microbial decomposition processes of soil organic matter and soil nutrient availability.
Climate change is profound in the Arctic where increased snowfall during winter and warmer growing season temperatures may accelerate soil nitrogen (N) turnover and increase inorganic N availability. ...Nitrous oxide (N2O) is a potent greenhouse gas formed by soil microbes and in the Arctic, the production is seen as limited mainly by low inorganic N availability. Hence, it can be hypothesized that climate change in the Arctic may increase total N2O emissions, yet this topic remains understudied. We investigated the combined effects of variable snow depths and experimental warming on soil N cycling in a factorial field study established along a natural snowmelt gradient in a low Arctic heath ecosystem. The study assessed N2O surface fluxes, gross N mineralization and nitrification rates, potential denitrification activity, and the pools of soil microbial, soil organic and soil inorganic N, carbon (C) and phosphorus (P) during two growing seasons. The net fluxes of N2O averaged 1.7 μg N2O–N m−2 h−1 (range −3.6 to 10.5 μg N2O–N m−2 h−1), and generally increased from ambient (1 m) to moderate (2–3 m) snow depths. At the greatest snow depth (4 m) where snowmelt was profoundly later, N2O fluxes decreased, likely caused by combined negative effects of low summer temperatures and high soil moisture. Positive correlations between N2O and nitrate (NO3−) and dissolved organic N (DON) suggested that the availability of N was the main controlling variable along the snowmelt gradient. The maximum N2O fluxes were observed in the second half of August associated with high NO3− concentrations. The effect of growing season experimental warming on N2O surface flux varied along the snowmelt gradient and with time. Generally, the experimental warming stimulated N2O fluxes under conditions with increased concentrations of inorganic N. In contrast, warming reduced N2O fluxes when inorganic N was low. Experimental warming had no clear effects on soil inorganic N. The study suggests that if increased winter precipitation leads to a deeper snow cover and a later snowmelt, total emissions of N2O from low Arctic heath ecosystems may be enhanced in the future and, dependent on dissolved N availability, summer warming may stimulate or reduce total emissions.
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•We studied effects of warming on tundra N2O emissions along a snowmelt gradient.•The N2O emissions peaked with late snowmelt, likely due to high dissolved soil N.•Areas with highest snow depth (4 m) exhibited low N2O emissions (low temperature).•Experimental warming enhanced N2O emissions, where soil N was high (late snowmelt).•We conclude that low Arctic heath is a source to N2O regulated by dissolved soil N.
The Arctic climate is projected to change during the coming century, with expected higher air temperatures and increased winter snowfall. These climatic changes might alter litter decomposition ...rates, which in turn could affect carbon (C) and nitrogen (N) cycling rates in tundra ecosystems. However, little is known of seasonal climate change effects on plant litter decomposition rates and N dynamics, hampering predictions of future arctic vegetation composition and the tundra C balance. We tested the effects of snow addition (snow fences), warming (open top chambers), and shrub removal (clipping), using a full-factorial experiment, on mass loss and N dynamics of two shrub tissue types with contrasting quality: deciduous shrub leaf litter (Salix glauca) and evergreen shrub shoots (Cassiope tetragona). We performed a 10.5-month decomposition experiment in a low-arctic shrub tundra heath in West-Greenland. Field incubations started in late fall, with harvests made after 249, 273, and 319 days of field incubation during early spring, summer and fall of the next year, respectively. We observed a positive effect of deeper snow on winter mass loss which is considered a result of observed higher soil winter temperatures and corresponding increased winter microbial litter decomposition in deep-snow plots. In contrast, warming reduced litter mass loss during spring, possibly because the dry spring conditions might have dried out the litter layer and thereby limited microbial litter decomposition. Shrub removal had a small positive effect on litter mass loss for c. tetragona during summer, but not for S. glauca. Nitrogen dynamics in decomposing leaves and shoots were not affected by the treatments but did show differences in temporal patterns between tissue types: there was a net immobilization of N by C. tetragona shoots after the winter incubation, while S. glauca leaf N-pools were unaltered over time. Our results support the widely hypothesized positive linkage between winter snow depth and litter decomposition rates in tundra ecosystems, but our results do not reveal changes in N dynamics during initial decomposition stages. Our study also shows contrasting impacts of spring warming and snow addition on shrub decomposition rates that might have important consequences for plant community composition and vegetation-climate feedbacks in rapidly changing tundra ecosystems.
Northern latitude tundra heaths have accumulated large amounts of organic carbon (C) in the soil. Changes in climatic conditions such as temperature and winter precipitation might affect the C ...balance and potentially change these tundra ecosystems from being C sinks to sources of CO2 emitted to the atmosphere. However, studies on C fluxes with single and combined winter snow and summer warming effects are scarce. This study investigates gross ecosystem production (GEP), ecosystem respiration (ER), net ecosystem production (NEP) and carbon isotopic composition of CO2 emitted from a dry heath in arctic Greenland one and two years following field manipulations of summer temperature, shrub abundance and winter snow depth. Our aims were to quantify climatic change effects on CO2 fluxes and the growing season carbon balance of the ecosystem and to investigate shifts in δ13C of emitted CO2 potentially due changes in emission from old soil C versus recently fixed carbon. Ecosystem CO2 fluxes and δ13C-CO2 were measured using closed chambers, and soil CO2 concentrations and δ13C were measured depth-specifically using gas probes.
We found a significant increase of CO2 emissions in all treatments during both years. Growing season NEP increased by 38 and 73% with 1 m enhanced winter snow depth, by 113 and 144% with summer warming and by 61 and 320% with total shrub removal in 2013 and 2014, respectively. The snow effect can be explained by the delay in the onset of growth as indicated by early season reduced vegetation greenness. The effect of warming was a result of an increase of ER by 39 and 32%, and the effect of shrub removal was mainly due to a reduction in GEP by 34 and 48%, in 2013 and 2014, respectively. Furthermore, the δ13C of the carbon source of CO2 emitted from warmed plots increased significantly two years after the experiment was initiated. This might indicate increased decomposition of 13C enriched soil organic matter and hence increased mineralization of the old carbon stock in the soil under warmed conditions. The increase of NEP, the additive response of all treatments, and the indications of increased emission of carbon from old stocks due to warming (or warming-induced drying), demonstrate the risk of a relatively fast feedback to climate warming during the snow-free season.
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•More snow, summer warming and shrub removal increase arctic tundra net CO2 emissions.•Snow effect on carbon balance due to delayed leafing and low spring photosynthesis.•Warming effect on carbon balance due to increased ER and unchanged GEP.•Carbon isotope composition may indicate increased decomposition of old SOM by warming.•Shrub removal effects on carbon balance mainly due to reduced photosynthesis.
Warming in the Arctic accelerates top‐soil decomposition and deep‐soil permafrost thaw. This may lead to an increase in plant‐available nutrients throughout the active layer soil and near the ...permafrost thaw front. For nitrogen (N) limited high arctic plants, increased N availability may enhance growth and alter community composition, importantly affecting the ecosystem carbon balance. However, the extent to which plants can take advantage of this newly available N may be constrained by the following three factors: vertical distribution of N within the soil profile, timing of N‐release, and competition with other plants and microorganisms. Therefore, we investigated species‐ and depth‐specific plant N uptake in a high arctic tundra, northeastern Greenland. Using stable isotopic labelling (15N‐NH4+), we simulated autumn N‐release at three depths within the active layer: top (10 cm), mid (45 cm) and deep‐soil near the permafrost thaw front (90 cm). We measured plant species‐specific N uptake immediately after N‐release (autumn) and after 1 year, and assessed depth‐specific microbial N uptake and resource partitioning between above‐ and below‐ground plant parts, microorganisms and soil. We found that high arctic plants actively foraged for N past the peak growing season, notably the graminoid Kobresia myosuroides. While most plant species (Carex rupestris, Dryas octopetala, K. myosuroides) preferred top‐soil N, the shrub Salix arctica also effectively acquired N from deeper soil layers. All plants were able to obtain N from the permafrost thaw front, both in autumn and during the following growing season, demonstrating the importance of permafrost‐released N as a new N source for arctic plants. Finally, microbial N uptake markedly declined with depth, hence, plant access to deep‐soil N pools is a competitive strength. In conclusion, plant species‐specific competitive advantages with respect to both time‐ and depth‐specific N‐release may dictate short‐ and long‐term plant community changes in the Arctic and consequently, larger‐scale climate feedbacks.
Climate warming in the High Arctic may increase soil nitrogen (N) availability, both in shallow soil layers and near the permafrost thaw front. Increased N availability could enhance plant growth and change species composition, importantly affecting the ecosystem carbon balance. We show that high arctic plants actively forage for N in the autumn, and that species differ in terms of where in the soil profile they take up N, including at the permafrost thaw front and in competition with microorganisms. Plant species‐specific competitive advantages with respect to both time‐ and depth‐specific N uptake may dictate long‐term plant community change in the Arctic.
Nitrogen fixation in the High Arctic Rousk, Kathrin; Sorensen, Pernille Laerkedal; Michelsen, Anders
Biogeochemistry,
11/2017, Letnik:
136, Številka:
2
Journal Article
Recenzirano
Biological nitrogen (N₂) fixation performed by diazotrophs (N₂ fixing bacteria) is thought to be one of the main sources of plant available N in pristine ecosystems like arctic tundra. However, ...direct evidence of a transfer of fixed N₂ to non-diazotroph associated plants is lacking to date. Here, we present results from an in situ ¹⁵N–N₂ labelling study in the High Arctic. Three dominant vegetation types (organic crust composed of free-living cyanobacteria, mosses, cotton grass) were subjected to acetylene reduction assays (ARA) performed regularly throughout the growing season, as well as ¹⁵N–N₂ incubations. The ¹⁵N-label was followed into the dominant N₂ fixer associations, soil, soil microbial biomass and nondiazotroph associated plants three days and three weeks after labelling. Mosses contributed most to habitat N₂ fixation throughout the measuring campaigns, and N₂ fixation activity was highest at the beginning of the growing season in all plots. Fixed ¹⁵N–N₂ became quickly (within 3 days) available to non-diazotroph associated plants in all investigated vegetation types, proving that N₂ fixation is an actual source of available N in pristine ecosystems.
Previous research has shown that experimental perturbations of arctic ecosystems simulating direct and indirect effects of predicted environmental changes have led to strong responses in the plant ...communities, mostly associated with increased plant nutrient availability. Similarly, changes in decomposition and nutrient mineralization are likely to occur if the soil warms and the soil moisture conditions are altered. Plant and microbial responses have usually been investigated separately, and few, if any, studies have addressed simultaneous responses to environmental changes in plants and soil microorganisms, except in models. We measured simultaneous responses in biomass, nitrogen (N), and phosphorus (P) incorporation in plants and microorganisms after five years of factorial fertilizer addition, air warming, and shading. We expected increased N and P uptake by microorganisms after fertilizer addition and also after warming, due to increases in mineralization rates in warmer soils. Plant productivity and N and P uptake were expected to increase after fertilizer addition but less after warming, because microbes were expected to absorb most of the extra released nutrients. Shading was expected to decrease plant production and also microbial biomass, due to the reduced production of labile carbon (C) in plant root exudates associated with reduced photosynthesis. We found that the plants responded strongly to fertilizer addition by increased biomass accumulation and N and P uptake. They responded less to warming, but more than expected, showing a decline in N and P concentrations in many cases. There were few significant responses to shading. The strongest response was found in combined fertilizer addition and warming treatments. All functional vascular plant groups responded similarly. However, mosses declined under those conditions when vascular plant growth was most pronounced. Contrary to our expectation, microbial C, N, and P did not increase after warming, but microbial N and P increased after shading. As expected, fertilizer addition led to increased microbial P content, whereas microbial N either increased or did not change. In general, microbial C did not change in any treatment. The microbes accumulated extra N and P only when soil inorganic N or P levels increased, suggesting that the soil microorganisms absorbed extra nutrients only in cases of declining N and P sink strength in plants.
Global warming and increased nutrient availability in the Arctic have attracted wide attention. However, it is unknown how an increased supply of nitrogen (N), phosphorus (P) and/or labile carbon (C) ...– alone and in combinations – affects the concentrations and pools of C and nutrients in plants, soil and soil microorganisms, and whether the cessation of these additions allows the ecosystem to recover from amendments. Six treatments (control, C, N, P, NP and C + NP) were applied at a subarctic heath for 8–10 years. After being untreated for two years, amendments were re-applied to one half of the plots for four years while the other plot half received no amendments. When the plots were harvested, we could therefore compare responses in plots with nearly continuous 14–16-year amendments to those with six years with discontinued treatments. The responses to individual and combined nutrient additions were mostly similar in re-initiated and discontinued plots. Individual N addition strongly increased the C and N pools in the graminoids, thereby also increasing the C and N pools in litter and fine roots compared to the plots without added N. This contributed to the increased microbial biomass C and total C in soil. P addition alone increased C and N pools in vascular cryptogams, as well as PO43−, NH4+, dissolved organic carbon and dissolved organic nitrogen concentrations in soil compared to the plots without added P. Hence, plant functional groups showed differential responses to long-term N and P amendment, and after the initial nutrient additions for 8–10 years, the investigated subarctic tundra ecosystem had reached a new steady state that was resilient to further changes still six years after cessation of additions.
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•Similar effects for recent and old nutrient additions in a subarctic tundra ecosystem.•N addition alone increased C and N pools in graminoids, litter and fine roots.•P addition alone increased C and N pools in vascular cryptogams.•Individual and combined additions of C, N, and P increased microbial biomass C.•After long-term nutrient additions, the tundra ecosystem reached a new steady state.
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