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Mountain ecosystems are particularly threatened by ongoing climate change and the species composition of high elevation grasslands is already changing. An open research question is how these ecosystems will adapt to changes in their key environmental constraints. The responses of wooded pastures to experimental climate forcing were analysed in a transplantation experiment conducted downslope, along an elevational temperature and precipitation gradient on the lee side of Jura Mountains, Switzerland (up to +4.17 degrees C and -35% precipitation). To improve mechanistic understanding of biodiversity and biomass decreases in response to transplantation, changes in functional traits within foliage and roots of one ubiquitous (Taraxacum officinale) and one montane (Alchemilla monticola) perennial forb species were investigated. In consequence of transplantation, the two studied species raised their temperature optimum for CO2 assimilation and net photosynthesis yield from 20 to 30 degrees C. During cool periods, the highest rates of leaf gas exchanges were measured at the lower recipient sites. However, an opposite trend was observed during a spring drought and summer warm spell. Regarding the more integrative morpho-anatomical traits, Alchemilla primarily acclimated to warmer temperatures at the recipient sites with increased leaf and foliage rosette size. Missing xeromorphic and/or hydraulic adjustments in foliage and roots, its susceptibility to higher vapour pressure deficits and lower soil moisture availability was thus enhanced. Taraxacum showed adjustments to both warmer temperature and lower moisture availability, including reduced leaf size, lower hydraulic diameter of xylem vessels and theoretical specific hydraulic conductivity. The anticipated shift in the environmental conditions at high elevation, with reduced coldness limitation but increasingly constraining water economy, could thus become particularly demanding for montane species of wooded pastures. It may favour perennials with large phenotypic plasticity but leads to maladjustments and loss of the species which are more specifically adapted to montane conditions. Read the free Plain Language Summary for this article on the Journal blog.

Soil organic matter (SOM) plays an important role in the global carbon cycle, especially in alpine ecosystems. However, ongoing forest expansion in high elevation systems potentially alters SOM storage through changes in organic matter (OM) inputs and microclimate. In this study we investigated the effects of an Picea abies L. afforestation chrono-sequence (40–130 years) of a former subalpine pasture in Switzerland on soil organic carbon (SOC) stocks and SOM dynamics. We found that SOC stocks remained relatively constant throughout the chrono-sequence, with comparable SOC stocks in the mineral soils after afforestation and previous pasture (SOC40-year-old forest = 11.6 ± 1.1 kg m−2, SOC130-year-old forest = 11.0 ± 0.3 kg m−2, and SOC pasture = 11.5 ± 0.5 kg m−2). However, including the additional carbon of the organic horizons in the forest, reaching up to 1.7 kg m−2 in the 55-year-old forest, resulted in a slight in-crease in overall SOC stocks following afforestation. We found that the soil C:N ratio in the mineral soil increased in the topsoil (0–5cm) with increasing forest stand age, from 11.9 ± 1.3 in the grassland to 14.3 ± 1.8 in the 130-year-old forest. In turn, we observed a decrease in soil C:N ratio with increasing depth in all forest stand ages. This suggests that litter-derived organic matter (C:N from 35.1 ± 1.9 to 42.4 ± 10.8) is likely incorporated and translocated from the organic horizon to the mineral topsoil (0–10 cm) of the profiles. As roots had very high C:N ratios (pasture 63.5 ± 2.8 and forests between 54.7 ± 3.9 and 61.2 ± 2.9), particulate root-derived organic matter seems to have a minor influence on forest soil C:N ratio and thereby on SOC stock accumulation in the mineral soil. These results suggest that, although the afforestation only moderately affected the SOC stock, there is an apparent alteration in the SOC dynamics through changes of the litter composition caused by the vegetation shift. We conclude that spruce afforestation on a former subalpine pasture does not necessarily change the total SOC stock and that consequently there is no SOC sequestration on a decadal to centennial scale.

Warming-induced microbial decomposition of organic matter in permafrost soils constitutes a climate-change feedback of uncertain magnitude. While physicochemical constraints on soil functioning are relatively well understood, the constraints attributable to microbial community composition remain unclear. Here we show that biogeochemical processes in permafrost can be impaired by missing functions in the microbial community-functional limitations-probably due to environmental filtering of the microbial community over millennia-long freezing. We inoculated Yedoma permafrost with a functionally diverse exogenous microbial community to test this mechanism by introducing potentially missing microbial functions. This initiated nitrification activity and increased CO2 production by 38% over 161 days. The changes in soil functioning were strongly associated with an altered microbial community composition, rather than with changes in soil chemistry or microbial biomass. The present permafrost microbial community composition thus constrains carbon and nitrogen biogeochemical processes, but microbial colonization, likely to occur upon permafrost thaw in situ, can alleviate such functional limitations. Accounting for functional limitations and their alleviation could strongly increase our estimate of the vulnerability of permafrost soil organic matter to decomposition and the resulting global climate feedback. Carbon dioxide emissions from permafrost thaw are substantially enhanced by relieving microbial functional limitations, according to incubation experiments on Yedoma permafrost.

In alpine areas of the European Alps, many of the pastures that are no longer economically profitable are being converted into forests (Bolli et al., 2007). Afforestation of former pasture has been acknowledged as a contribution to mitigate CO2 emissions by an increased storage of soil carbon in soil and biomass (Smal et al., 2019). So far, several studies indicated that afforestation of former pastures does not always lead to an increase or decrease of soil organic carbon stocks after 30 to 40 years of afforestation. The loss of soil organic carbon in the mineral soil, however, can be rebalanced by the increased accumulation of soil organic carbon in the organic forest floor (Thuille and Schulze, 2006). Nevertheless, studies concerning the changes as well as restoration of SOM following afforestation are limited.In this study, we aimed to trace the source and transformation of SOM in a subalpine afforestation sequence (0-130 years) with Norway spruce (Picea abies L.) on a former pasture in Jaun, Switzerland. Soil and root samples were taken with volumetry cylinders to a depth of 45cm at 5cm increments. To trace the source and transformation of SOM, soil samples were analysed for plant-and microorganism-derived SOM by combining multiple compound classes as free extractable lipids, such as n-alkanes and free fatty acids.Preliminary results show a higher (p-2) compared to forested areas. The highest (p=0.92) fine root biomass was observed in the youngest forest (40yr; 2.3±0.7gm-2), followed by the 130yr (0.7±0.2gm-2) and 55yr (0.6±0.2gm-2) old forest. Highest carbon stocks (14.0±0.8 kgm-2) were observed in the youngest forest followed by the 130yr (11.0±0.3 kgm-2) old and 55yr (9.6±1.1kgm-2) old forest. In summary, afforestation of former pasture (11.2±0.0 kgm-2) does not result in changes (p=0.37) in the total (0-45cm) organic carbon stock over a period of decades. However, there is a significant (pReferencesBolli, J. C., Rigling, A., and Bugmann, H. (2007). The influence of changes in climate and land-use on regeneration dynamics of Norway spruce at the treeline in the Swiss Alps. Silva Fennica, 41, 55.Smal, H., Ligęza, S., Pranagal, J., Urban, D., and Pietruczyk-Popławska, D. (2019). Changes in the stocks of soil organic carbon, total nitrogen and phosphorus following afforestation of post-arable soils: A chronosequence study. Forest Ecology and Management, 451, 117536.Thuille, A., and Schulze, E. D. (2006). Carbon dynamics in successional and afforested spruce stands in Thuringia and the Alps. Global Change Biology, 12, 325-342

Tundra ecosystems hold large stocks of soil organic matter (SOM), likely due to low temperatures limiting rates of microbial SOM decomposition more than those of SOM accumulation from plant primary productivity and microbial necromass inputs. Here we test the hypotheses that distinct tundra vegetation types and their carbon supply to characteristic rhizosphere microbes determine SOM cycling independent of temperature. In the subarctic Scandes, we used a three-way factorial design with paired heath and meadow vegetation at each of two elevations, and with each combination of vegetation type and elevation subjected during one growing season to either ambient light (i.e., ambient plant productivity), or 95% shading (i.e., reduced plant productivity). We assessed potential above- and belowground ecosystem linkages by uni- and multivariate analyses of variance, and structural equation modelling. We observed direct coupling between tundra vegetation type and microbial community composition and function, which underpinned the ecosystem's potential for SOM storage. Greater primary productivity at low elevation and ambient light supported higher microbial biomass and nitrogen immobilisation, with lower microbial mass-specific enzymatic activity and SOM humification. Congruently, larger SOM at lower elevation and in heath sustained fungal-dominated microbial communities, which were less substrate-limited, and invested less into enzymatic SOM mineralisation, owing to a greater carbon-use efficiency (CUE). Our results highlight the importance of tundra plant community characteristics (i.e., productivity and vegetation type), via their effects on soil microbial community size, structure and physiology, as essential drivers of SOM turnover. The here documented concerted patterns in above- and belowground ecosystem functioning is strongly supportive of using plant community characteristics as surrogates for assessing tundra carbon storage potential and its evolution under climate and vegetation changes. Published version The study benefitted from internal CIRC funding for the use of the Abisko Scientific Research Station, kindly operated by the Swedish Polar Research Secretariat. K.G. was further supported by funding from the Swiss National Science Foundation (grant no. PZ00P2_174047).

Climate warming is releasing carbon from soils around the world, constituting a positive climate feedback. Warming is also causing species to expand their ranges into new ecosystems. Yet, in most ecosystems, whether range expanding species will amplify or buffer expected soil carbon loss is unknown. Here, we used two whole-community transplant experiments and a follow-up glasshouse experiment to determine whether the establishment of herbaceous lowland plants in alpine ecosystems influences soil carbon content under warming. We found that warming (transplantation to low elevation) led to a negligible decrease in alpine soil carbon content, but its effects became significant and 52% ± 31% (mean ± 95% confidence intervals) larger after lowland plants were introduced at low density into the ecosystem. We present evidence that decreases in soil carbon content likely occurred via lowland plants increasing rates of root exudation, soil microbial respiration, and CO2 release under warming. Our findings suggest that warming-induced range expansions of herbaceous plants have the potential to alter climate feedbacks from this system, and that plant range expansions among herbaceous communities may be an overlooked mediator of warming effects on carbon dynamics., eLife, 11, ISSN:2050-084X

The conversion of tropical forest for cassava cultivation is widely known to decrease the soil organic matter (OM) and nutrient contents of highly weathered soils in the tropics. Amazonian Dark Earth (ADE) might be more resistant to this process due to their historical anthropogenic amelioration with e.g. charcoal, ceramics and bones, leading to higher soil OM and nutrient concentrations. In this study, we analyzed the effect of land use change on the OM dynamics under tropical conditions and how this is related with P distribution at the microscale, using ADE and an adjacent Acrisol (ACR) as model systems. Soil samples were obtained south of Manaus (Brazil), from a secondary forest and an adjacently located 40-year-old cassava plantation. The land use change induced a severe decrease of organic carbon (OC) concentrations in ADE (from 35 to 15 g OC kg‑1) while OC in the adjacent ACR was less affected (18 to 16 g OC kg‑1). The analysis by 13C NMR spectroscopy showed that the conversion of secondary forest to cassava changed the chemical composition of OM to a more decomposed state (increase of alkyl:O/N-alkyl ratio) in the ADE whereas the OM in ACR changed to a less decomposed state (decrease of alkyl:O/N-alkyl ratio). According to neutral sugar and lipid extraction analyses, land use change led to a larger impact on the microbial-derived and plant-derived compounds in the ADE compared to the ACR. In order to analyze the interactions of OC and P at the microscale, we conducted an incubation experiment with 13C glucose for the analysis with Scanning X-ray Microscopy (SXM) and Nano scale Secondary Ion Mass Spectrometry (NanoSIMS). In both soil types ADE and ACR, land use change caused a reduction of the total 13C glucose respiration by approximately one third in a 7-days incubation, implying lower microbial activity. Microorganisms in both soil types appear to be more readily active in soils under forest, since we observed a distinct lag time between 13C glucose addition and respiration under cassava planation. This indicated differences in microbial community structure, which we will be assessed further by determining the 13C label uptake by the microbial biomass and the microbial community structure using 13C PLFA analysis. Preliminary results from synchrotron-based STXM demonstrate a distinct arrangement of OM at fine-sized charcoal-particle interfaces. From ongoing NanoSIMS analyses, we expect further insights on the co-localization of P and 13C-labelled spots at the microscale. Despite the high loss of OC in the ameliorated ADE through land use change, the remaining OM might foster nutrient dynamics at the microscale thanks to charcoal interactions compared to the ACR. Our results contribute to a better understanding of the C and P interactions and how these respond to land use change in highly weathered tropical soils.

European forests are facing higher frequencies of extreme droughts, potentially impairing tree growth and ecosystem functioning. Drought limits the metabolic activity of plants and soil organisms, either directly or through reduced belowground carbon (C) allocation of recent assimilates, thus affecting C cycling in the plant-soil system. However, the net effect on belowground soil C storage is still unclear, as drought suppresses both C inputs from plants and outputs from soils. Moreover, understanding the underlying mechanisms is complicated due to long-term acclimation and adaptation of plant and soil organisms to water limitation.We investigated the impact of repeated summer droughts in a Scots pine (Pinus sylvestris L.) forest on soil C storage and C cycling, taking advantage of a large-scale irrigation experiment running since 2003 in a dry inner-Alpine valley in Switzerland (Pfynwald, Valais), which removed the “natural” water limitation. We assessed the responses of soil organic carbon (SOC) stocks and C fluxes by measuring litter fall and decomposition, fine root biomass and production, soil CO2 effluxes, C-mineralization, and 13C-labelled glucose utilization by soil microorganisms. After 16 years of irrigation, the organic layers lost significant amounts of C (-1000 g m-2), despite a 50% increase in litter fall. This C loss was almost compensated by a C gain in the mineral soil (+870 g m-2) under irrigation. The decrease in C storage in the organic layers can be related to a three-fold increase in litter decomposition mainly through soil macrofauna as indicated by a litter-bag experiment. In parallel, the C gain in the mineral soil can be attributed mainly to increased incorporation of litter by soil fauna, together with greater C input from the rhizosphere (+70% fine root biomass for Scots pine in mineral soil). Furthermore, irrigation stimulated soil CO2 efflux as well as microbial C-mineralization of organic and mineral soil, indicating enhanced soil C cycling. Addition of 13C-enriched glucose to mineral soils revealed a stronger utilization of this easily available C substrate in the drought than in the irrigated soils, together with a negative priming of soil organic matter (SOM) decomposition shortly after substrate addition. These results suggest that the altered quantity and quality of C inputs under irrigation has increased the availability of easily degradable C in soil.This study reveals that long-term summer irrigation in a drought-prone pine forest has strong impacts on multiple interlinked processes of the soil C cycle. The removal of water limitation strongly altered vertical soil C distribution, accelerated soil C cycling and altered the substrate use by soil organisms, but had only a small net effect on the whole-profile SOC stocks.

Environmental changes feedback to climate through their impact on soil functions such as carbon (C) and nutrient sequestration. Abiotic conditions and the interactions between above- and belowground biota drive soil responses to environmental change but these (a)biotic interactions are challenging to study. Nonetheless, better understanding of these interactions would improve predictions of future soil functioning and the soil-climate feedback and, in this context, permafrost soils are of particular interest due to their vast soil C-stores. We need new tools to isolate abiotic (microclimate, chemistry) and biotic (roots, fauna, microorganisms) components and to identify their respective roles in soil processes. We developed a new experimental setup, in which we mimic thermokarst (permafrost thaw-induced soil subsidence) by fitting thawed permafrost and vegetated active layer sods side by side into mesocosms deployed in a subarctic tundra over two growing seasons. In each mesocosm, the two sods were separated from each other by barriers with different mesh sizes to allow varying degrees of physical connection and, consequently, (a)biotic exchange between active layer and permafrost. We demonstrate that our mesh-approach succeeded in controlling 1) lateral exchange of solutes between the two soil types, 2) colonization of permafrost by microbes but not by soil fauna, and 3) ingrowth of roots into permafrost. In particular, experimental thermokarst induced a ~60% decline in permafrost nitrogen (N) content, a shift in soil bacteria and a rapid buildup of root biomass (+33.2 g roots m−2 soil). This indicates that cascading plant-soil-microbe linkages are at the heart of biogeochemical cycling in thermokarst events. We propose that this novel setup can be used to explore the effects of (a)biotic ecosystem components on focal biogeochemical processes in permafrost soils and beyond.

The conversion of tropical forest for cassava cultivation is widely known to decrease the soil organic matter (OM) and nutrient contents of highly weathered soils in the tropics. Amazonian Dark Earth (ADE) might be affected less due to their historical anthropogenic amelioration with e.g. charcoal, ceramics and bones, leading to higher soil OM and nutrient concentrations. In this study, we analysed the effect of land use change on the OM dynamics and its composition under tropical conditions, using ADE and an adjacent Acrisol (ACR) as model systems. Soil samples were obtained south of Manaus (Brazil), from a secondary forest and an adjacently located 40-year-old cassava plantation. The land use change induced a severe decrease of organic carbon (OC) concentrations in ADE (from 35 to 15 g OC kg‑1) while OC in the adjacent ACR was less affected (18 to 16 g OC kg‑1). Soils were analysed by 13C NMR spectroscopy to obtain information on how the conversion of secondary forest to cassava affected the chemical composition of OM. Our results show that land use change induces differences in the OM composition: The OM in ADE changes to a more decomposed state (increase of alkyl:O/N-alkyl ratio) whereas the OM in ACR changes to a less decomposed state (decrease of alkyl:O/N-alkyl ratio). According to a molecular mixing model, land use change influenced mostly the proportion of lipids, which might be related with a change of the plant input. The incubation of the soils with 13C glucose enabled resolving how soil microorganisms were affected by land use change. In both soil types ADE and ACR, land use change caused a reduction of the total 13C glucose respiration by approximately one third in a 7-days incubation, implying lower microbial activity. Microorganisms in both soil types appear to be more readily active in soils under forest, since we observed a distinct lag time between 13C glucose addition and respiration under cassava planation. This indicated differences in microbial community structure, which we will assess further by determining the 13C label uptake by the microbial biomass and the microbial community structure using 13C PLFA analysis. Preliminary results from synchrotron-based STXM demonstrate a distinct arrangement of OM at fine-sized charcoal-particle interfaces. Samples of soils receiving 13C label will be further analysed by NanoSIMS with the hypothesis that charcoal interfaces foster nutrient dynamics at the microscale. Despite the high loss of OC in the ameliorated ADE through land use change, the remaining OM might improve the nutrient availability thanks to charcoal interactions compared to the ACR. Our results contribute to a better understanding of the sensitivity of OM upon land use change and how the microbial community is responding to land use change in highly weathered tropical soils.

Forest soils are storing large quantities of carbon, but their quantitative role in sequestering C is less certain. In principal, soils developed over millennia are assumed to be ‘in equilibrium’ with minimal C stock changes. This concept is challenged by forest soil inventories (in Germany and France) indicate a substantial increase in soil C storage. However, soil organic matter (SOM) storage is susceptible to recent changes in forests - climate warming and droughts, increasing forest disturbances, and a more intensive forest management are all potentially increasing SOM turnover which may turn forest soils into C sources. Here, I will critically discuss the role in Swiss forest soils as C sinks by presenting data from 1000 soil profiles across environmental gradients and from flux measurements in large scale ecosystem manipulation experiments.Swiss forests soils are among the C-richest soils in Europe storing on average 140 t C/ha. Analysis of 1000 forest soils show that these SOM stocks are caused by their high contents in potential SOM sorbents (pH, Al+Fe-oxides, Ca, clay), but also by the cool temperatures and high amounts of precipitation. Climate manipulation experiments suggest Swiss forest soils are vulnerable to loose C with expected climatic changes. A six year long soil warming experiment at treeline revealed soil C losses, while a 15 year long irrigation experiment in a dry forest induced C gains in the mineral soil, implying that a warmer and more frequent droughts will lead to C losses.Switzerland - as other European mountainous areas – is currently experiencing a major change in land-use due to land abandonment, with the forests expanding by 3 to 4% per decade. Forest expansion affects a multitude of factors driving SOM cycling and storage, including the quantity and quality of organic matter inputs above and below the ground, a cooler and drier microclimate, and change in microbial diversity and activity. In contrast to the intuitive assumption that forests expansion leads to C gains in soils, measurements along an afforestation chronosequence of alpine grassland show that forest expansion leads to minimal changes in SOM stocks but a strong change in SOM quality. Soils gains in particulate organic matter with increasing forest age but lose C in mineral-associated organic matter. In support, reconstructing forest cover ages of 850 soil profiles showed that forest age and hence time since conversion into forest (predominantly from grasslands) did not significantly affect total SOM stocks, while other factors, especially physico-chemical soil characteristics and climate were more important. Overall, these results show that the inherently C rich forest soils in Switzerland are unlikely to gain additional C but rather loose it in response to the ongoing changes in climate and land-use.

Seasonal snow cover provides essential insulation for mountain ecosystems, but expected changes in precipitation patterns and snow cover duration due to global warming can influence the activity of soil microbial communities. In turn, these changes have the potential to create new dynamics of soil organic matter cycling. To assess the effects of experimental snow removal and advanced spring conditions on soil carbon (C) and nitrogen (N) dynamics, and on the biomass and structure of soil microbial communities, we performed an in situ study in a subalpine grassland in the Austrian Alps, in conjunction with soil incubations under controlled conditions. We found substantial winter C-mineralisation and high accumulation of inorganic and organic N in the topsoil, peaking at snowmelt. Soil microbial biomass doubled under the snow, paralleled by a fivefold increase in its C:N ratio, but no apparent change in its bacteria-dominated community structure. Snow removal led to a series of mild freeze-thaw cycles, which had minor effects on in situ soil CO2 production and N mineralisation. Incubated soil under advanced spring conditions, however, revealed an impaired microbial metabolism shortly after snow removal, characterised by a limited capacity for C-mineralisation of both fresh plant-derived substrates and existing soil organic matter (SOM), leading to reduced priming effects. This effect was transient and the observed recovery in microbial respiration and SOM priming towards the end of the winter season indicated microbial resilience to short-lived freeze-thaw disturbance under field conditions. Bacteria showed a higher potential for uptake of plant-derived C substrates during this recovery phase. The observed temporary loss in microbial C-mineralisation capacity and the promotion of bacteria over fungi can likely impede winter SOM cycling in mountain grasslands under recurrent winter climate change events, with plausible implications for soil nutrient availability and plant-soil interactions. (C) 2017 Elsevier B.V. All rights reserved.

Upward shifts of mountain vegetation lag behind rates of climate warming, partly related to interconnected changes belowground. Here, we unravel above- and belowground linkages by drawing insights from short-term experimental manipulations and elevation gradient studies. Soils will likely gain carbon in early successional ecosystems, while losing carbon as forest expands upward, and the slow, high-elevation soil development will constrain warming-induced vegetation shifts. Current approaches fail to predict the pace of these changes and how much they will be modified by interactions among plants and soil biota. Integrating mountain soils and their biota into monitoring programs, combined with innovative comparative and experimental approaches, will be crucial to overcome the paucity of belowground data and to better understand mountain ecosystem dynamics and their feedbacks to climate.

Since the publication of the original article, the authors noticed some errors in reference citation had been introduced throughout the paper. The following text contains excerpts from the original article and how they should appear with correct referencing. The publisher apologises for any inconvenience this has caused readers.

Severe constraints on grasslands productivity, ecosystem functions, goods and services are expected to result from projected warming and drought scenarios under climate change. Negative effects on vegetation can be mediated via soil fertility and water holding capacity, though specific mechanisms are fairly complex to generalise. In field drought experiments, it can be difficult to disentangle a drought effect per se from potential confounding effects related to vegetation or soil type, both varying along with climate. Furthermore, there is the need to distinguish the long-term responses of vegetation and soil to gradual climate shift from responses to extreme and stochastic climatic events. Here we address these limitations by means of a factorial experiment using a single dominant grassland species (the perennial ryegrass Lolium perenne L.) grown as a phytometer on two soils types with contrasted physicochemical characteristics, placed at two elevation sites along a climatic gradient, and exposed to early or late-season drought during the plant growing season. Warmer site conditions and reduced precipitation along the elevational gradient affected biogeochemistry and plant productivity more than the drought treatments alone, despite the similar magnitude in volumetric soil moisture reduction. Soil type, as defined here by its organic matter content (SOM), modulated the drought response in relation to local site climatic conditions and, through changes in microbial biomass and activity, determined the seasonal above and belowground productivity of L. perenne. More specifically, our combined uni- and multivariate analyses demonstrate that microbes in a loamy soil with low SOM are strongly responsive to change in climate, as indicated by a simultaneous increase in their C,N,P pools at high elevation with cooler temperatures and wetter soils. Contrastingly, microbes in a clay-loam soil with high SOM are mainly sensitive to temperature, as indicated by a strong increase in microbial biomass under warmer temperatures at low elevation and a concomitant increase in C:N, C:P and N:P ratios. High SOM promoted a better annual yield of the phytometer grass under warmer climate and the effect of drought on productivity was transient. In contrast, low SOM reduced cumulative yield under warmer conditions and root production strongly decreased, enduring a lasting drought effect. Microbes in soils with high organic matter remained more active during warmer and drier conditions, ensuring soil fertility and stimulating a higher overall plant nutrient availability and productivity. Our study highlights the important role of soil type for grassland responses to both stochastic climatic extremes and long-term climate change. Management practices enhancing SOM accumulation via organic residue incorporation seem a promising way to mitigate the effects of increased temperature and drought on plants and soil microbes alike promoting thereby a sustainable ecosystem functioning.

The decomposition of large stocks of soil organic carbon in thawing permafrost might depend on more than climate change-induced temperature increases: indirect effects of thawing via altered bacterial community structure (BCS) or rooting patterns are largely unexplored. We used a 10-year in situ permafrost thaw experiment and aerobic incubations to investigate alterations in BCS and potential respiration at different depths, and the extent to which they are related with each other and with root density. Active layer and permafrost BCS strongly differed, and the BCS in formerly frozen soils (below the natural thawfront) converged under induced deep thaw to strongly resemble the active layer BCS, possibly as a result of colonization by overlying microorganisms. Overall, respiration rates decreased with depth and soils showed lower potential respiration when subjected to deeper thaw, which we attributed to gradual labile carbon pool depletion. Despite deeper rooting under induced deep thaw, root density measurements did not improve soil chemistry-based models of potential respiration. However, BCS explained an additional unique portion of variation in respiration, particularly when accounting for differences in organic matter content. Our results suggest that by measuring bacterial community composition, we can improve both our understanding and the modeling of the permafrost carbon feedback. A correction to this article has been published. DOI: 10.1038/s41396-019-0384-1

International audience; In mountain ecosystems with marked seasonality, climate change can affect various processes in soils, potentially modifying long-term key soil services via change in soil organic carbon (C) storage. Based on a four-year soil transplantation experiment in Swiss subalpine grasslands, we investigated how imposed climate warming and reduced precipitation modified the drivers of soil carbon enzyme potential activities across winter and summer seasons. Specifically, we used structural equation models (SEMs) to identify biotic (microbial community structure, abundance and activity) and abiotic (quantity and quality of organic matter resources) drivers of soil C-enzymes (hydrolase and oxidase) in two seasons under two different climate scenarios. We found contrasting impacts of the climate manipulation on the drivers of C-enzymes between winter and summer. In winter, no direct effect of climate manipulation (reduced rainfall and warming) on enzyme activity was observed. Yet, climate indirectly down-regulated enzyme activity through a decrease in the availability of water extractable organic carbon (WEOC) labile resources. During summer, reduced soil moisture -induced by the climate manipulation- directly reduced soil microbial biomass, which led to a decrease in C-enzyme activity. In general, across both seasons, neither microbial community structure, nor organic matter quality were strong determinants of enzymatic activity. In particular organic matter recalcitrance (aromaticity) was not found as a general driver of either hydrolase or oxidase C-enzyme potential activities, though we did observe higher C-enzyme activities led to an increase of particulate organic matter recalcitrance in the summer season. Overall, our results highlight the seasonality of climate change effects on soil organic matter enzymatic decomposition, providing a comprehensive picture of seasonal potential cause and effect relationships governing C mineralization in subalpine grassland.

Dinitrogen fixation by cyanobacteria is of particular importance for the nutrient economy of cold biomes, constituting the main pathway for new N supplies to tundra ecosystems. It is prevalent in cyanobacterial colonies on bryophytes and in obligate associations within cyanolichens. Recent studies, applying interspecific variation in plant functional traits to upscale species effects on ecosystems, have all but neglected cryptogams and their association with cyanobacteria. Here we looked for species-specific patterns that determine cryptogam-mediated rates of N-2 fixation in the Subarctic. We hypothesised a contrast in N-2 fixation rates (1) between the structurally and physiologically different lichens and bryophytes, and (2) within bryophytes based on their respective plant functional types. Throughout the survey we supplied N-15-labelled N-2 gas to quantify fixation rates for monospecific moss, liverwort and lichen turfs. We sampled fifteen species in a design that captures spatial and temporal variations during the growing season in Abisko region, Sweden. We measured N-2 fixation potential of each turf in a common environment and in its field sampling site, in order to embrace both comparativeness and realism. Cyanolichens and bryophytes differed significantly in their cyanobacterial N-2 fixation capacity, which was not driven by microhabitat characteristics, but rather by morphology and physiology. Cyanolichens were much more prominent fixers than bryophytes per unit dry weight, but not per unit area due to their low specific thallus weight. Mosses did not exhibit consistent differences in N-2 fixation rates across species and functional types. Liverworts did not fix detectable amounts of N-2. Despite the very high rates of N-2 fixation associated with cyanolichens, large cover of mosses per unit area at the landscape scale compensates for their lower fixation rates, thereby probably making them the primary regional atmospheric nitrogen sink.

Warming-induced increases in microbial CO2 release in northern tundra may positively feedback to climate change. However, shifts in microbial extracellular enzyme activities (EEAs) may alter the impacts of warming over the longer term. We investigated the in situ effects of 3 years of winter warming in combination with the in vitro effects of a rapid warming (6 days) on microbial CO2 release and EEAs in a subarctic tundra heath after snowmelt in spring. Winter warming did not change microbial CO2 release at ambient (10 °C) or at rapidly increased temperatures, i.e., a warm spell (18 °C) but induced changes (P

Microbial communities drive soil organic matter (SOM) decomposition through the production of a variety of extracellular enzymes. Climate change impact on soil microbial communities and soil enzymatic activities can therefore strongly affect SOM turnover, and thereby determine the fate of ecosystems and their role as carbon sinks or sources. To simulate projected impacts of climate change on Swiss Jura subalpine grassland soils, an altitudinal soil transplantation experiment was set up in October 2009. On the fourth year of this experiment, we measured microbial biomass (MB), microbial community structure (MCS), and soil extracellular enzymatic activities (EEA) of nine hydrolytic and oxidative extracellular enzymes in the transplanted soils on a seasonal basis. We found a strong sampling date effect and a smaller but significant effect of the climate manipulation (soil transplantation) on EEA. Overall EEA was higher in winter and spring but enzymes linked to N and P cycles showed higher potential activities in autumn, suggesting that other factors than soil microclimate controlled their pool size, such as substrate availability. The climate warming manipulation decreased EEA in most cases, with oxidative enzymes more concerned than hydrolytic enzymes. In contrast to EEA, soil MB was more affected by the climate manipulation than by the seasons. Transplanting soils to lower altitudes caused a significant decrease in soil MB, but did not affect soil MCS. Conversely, a clear shift in soil MCS was observed between winter and summer. Mass-specific soil EEA (EEA normalized by MB) showed a systematic seasonal trend, with a higher ratio in winter than in summer, suggesting that the seasonal shift in MCS is accompanied by a change in their activities. Surprisingly, we observed a significant decrease in soil organic carbon (SOC) concentration after four years of soil transplantation, as compared to the control site, which could not be linked to any microbial data. We conclude that medium term (four years) warming and decreased precipitation strongly affected MB and EEA but not MCS in subalpine grassland soils, and that those shifts cannot be readily linked to the dynamics of soil carbon concentration under climate change. (C) 2014 Elsevier Ltd. All rights reserved.

Climate change can alter peatland plant community composition by promoting the growth of vascular plants. How such vegetation change affects peatland carbon dynamics remains, however, unclear. In order to assess the effect of vegetation change on carbon uptake and release, we performed a vascular plant-removal experiment in two Sphagnum-dominated peatlands that represent contrasting stages of natural vegetation succession along a climatic gradient. Periodic measurements of net ecosystem CO2 exchange revealed that vascular plants play a crucial role in assuring the potential for net carbon uptake, particularly with a warmer climate. The presence of vascular plants, however, also increased ecosystem respiration, and by using the seasonal variation of respired CO2 radiocarbon (bomb-14C) signature we demonstrate an enhanced heterotrophic decomposition of peat carbon due to rhizosphere priming. The observed rhizosphere priming of peat carbon decomposition was matched by more advanced humification of dissolved organic matter, which remained apparent beyond the plant growing season. Our results underline the relevance of rhizosphere priming in peatlands, especially when assessing the future carbon sink function of peatlands undergoing a shift in vegetation community composition in association with climate change.

International audience; Mountain soils stock large quantities of carbon as particulate organic matter that may be highly vulnerable to climate change. To explore potential shifts in soil organic matter (SOM) form and stability under climate change (warming and reduced precipitations), we studied the dynamics of SOM pools of a mountain grassland in the Swiss Jura as part of a climate manipulation experiment. The climate manipulation (elevational soil transplantation) was set up in October 2009 and simulated two realistic climate change scenarios. After 4 years of manipulation, we performed SOM physical fractionation to extract SOM fractions corresponding to specific turnover rates, in winter and in summer. Soil organic matter fraction chemistry was studied with ultraviolet, 3D fluorescence, and mid-infrared spectroscopies. The most labile SOM fractions showed high intra-annual dynamics (amounts and chemistry) mediated via the seasonal changes of fresh plant debris inputs and confirming their high contribution to the microbial loop. Our climate change manipulation modified the chemical differences between free and intra-aggregate organic matter, suggesting a modification of soil macro-aggregates dynamics. Interestingly, the 4-year climate manipulation affected directly the SOM dynamics, with a decrease in organic C bulk soil content, resulting from significant C-losses in the mineral-associated SOM fraction (MAOM), the most stable form of SOM. This SOC decrease was associated with a decrease in clay content, above- and belowground plants biomass, soil microbial biomass and activity. The combination of these climate changes effects on the plant-soil system could have led to increase C-losses from the MAOM fraction through clay-SOM washing out and DOC leaching in this subalpine grassland.

International audience; Climate change can affect the process of carbon cycling and leaf litter decomposition in multiple ways, both directly and indirectly, though the strength and direction of this relationship is often context dependent. In this experiment, we followed decomposition of a standard litter type-senescent leaves of Fagus sylvatica collected from a single location-along a 1000 m altitudinal gradient of four sites over 2.5 years. To control the edaphic conditions, we transplanted intact turf mesocosms from three different land-use types [densely wooded, sparsely wooded, and unwooded (UW) pastures] from the highest altitude site into UW pastures along the altitudinal gradient from the moist, cool high-elevation site to the dry, warm low-elevation site, using shade cloth to mimic the light conditions in the original habitats. Decomposition in the drier UW pasture mesocosms increased with altitude, likely because of higher moisture at the highest sites. Decomposition in the more mesic mesocosms from sparsely and densely wooded sites was insensitive to altitude, suggesting an overriding moisture, rather than temperature, constraint on decomposition across these sites. The functional composition of decomposer microbial communities (fungal/bacterial ratio) was similarly insensitive to altitude. Our findings bring substantial evidence for the controlling role of soil moisture on litter decomposition, as well as for the indirect effects of climate through changes in the decomposer community.

Climate change could impact strongly on cold-adapted mountain ecosystems, but little is known about its interaction with traditional land-use practices. We used an altitudinal gradient to simulate a year-round warmer and drier climate for semi-natural subalpine grasslands across a landscape of contrasting land-use management. Turf mesocosms from three pasture-woodland land-use types-unwooded pasture, sparsely wooded pasture, and densely wooded pasture-spanning a gradient from high to low management intensity were transplanted downslope to test their resistance to two intensities of climate change. We found strong overall effects of intensive (+4 K) experimental climate change (i.e., warming and reduced precipitation) on plant community structure and function, while moderate (+2 K) climate change did not substantially affect the studied land-use types, thus indicating an ecosystem response threshold to moderate climate perturbation. The individual land-use types were affected differently under the +4 K scenario, with a 60 % decrease in aboveground biomass (AGB) in unwooded pasture turfs, a 40 % decrease in sparsely wooded pasture turfs, and none in densely wooded ones. Similarly, unwooded pasture turfs experienced a 30 % loss of species, advanced (by 30 days) phenological development, and a mid-season senescence due to drought stress, while no such effects were recorded for the other land-use types. The observed contrasting effects of climate change across the pasture-woodland landscape have important implications for future decades. The reduced impact of climate change on wooded pastures as compared to unwooded ones should promote the sustainable land use of wooded pastures by maintaining low management intensity and a sparse forest canopy, which buffer the immediate impacts of climate change on herbaceous vegetation.

In agro-ecosystems, fertilization practices are crucial for sustaining crop productivity. Here, based on a 50-year long-term experiment, we studied the influence of fertilization practices (inorganic and/or organic) and nitrogen (N) application rates on (i) soil physicochemical properties, (ii) microbial and earthworm communities and (iii) crop production. Our results showed that soil organic carbon content was increased by incorporation of crop residues (+2.45%) and farmyard manure application (+6.40%) in comparison to the use of mineral fertilizer alone. In contrast, soil carbon stock was not significantly affected by these fertilization practices. Overall, only farmyard manure application improved soil physicochemical properties compared to mineral fertilization alone. Soil microbial population was enhanced by the application of organic amendments as indicated by microbial biomass and phospholipid-derived fatty acids contents. The fertilization practices and the N application rates affected significantly both the biomass and composition of earthworm populations, especially the epigeic and endogeic species. Finally, farmyard manure application significantly increased crop yield (+3.5%) in comparison to mineral fertilization alone. Crop residue incorporation rendered variable but similar crop yields over the 50-year period. The results of this long-term experiment indicate that the use of organic amendments not only reduces the need for higher amount of mineral N fertilizer but also improves the soil biological properties with direct effects on crop yield. (C) 2016 Agroscope. Published by Elsevier B.V.

The frequency of freezing events during the early growing season and the vulnerability to freezing of plants in European high-altitude environments could increase under future atmospheric and climate change. We tested early growing season freezing sensitivity in 10 species, from four plant functional types (PFTs) spanning three plant growth forms (PGFs), from a long-term in situ CO2 enrichment (566 vs. 370ppm) and 2-year soil warming (14K) experiment at treeline in the Swiss Alps (Stillberg, Davos). By additionally tracking plant phenology, we distinguished indirect phenology-driven CO2 and warming effects from direct physiology-related effects on freezing sensitivity. The freezing damage threshold (lethal temperature 50) under ambient conditions of the 10 treeline species spanned from � 6.7 � 0.31 C( Larix decidua )t o� 9.9 � 0.61 C( Vaccinium gaultherioides). PFT, but not PGF, explained a significant amount of this interspecific variation. Long-term exposure to elevated CO2 led to greater freezing sensitivity in multiple species but did not influence phenology, implying that physiological changes caused by CO2 enrichment were responsible for the effect. The elevated CO2 effect on freezing resistance was significant in leaves of Larix, Vaccinium myrtillus, and Gentiana punctata and marginally significant in leaves of Homogyne alpina and Avenella flexuosa. No significant CO2 effect was found in new shoots of Empetrum hermaphroditum or in leaves of Pinus uncinata, Leontodon helveticus, Melampyrum pratense, and V. gaultherioides. Soil warming led to advanced leaf expansion and reduced freezing resistance in V. myrtillus only, whereas Avenella showed greater freezing resistance when exposed to warming. No effect of soil warming was found in any of the other species. Effects of elevated CO2 and soil warming on freezing sensitivity were not consistent within PFTs or PGFs, suggesting that any future shifts in plant community composition due to increased damage from freezing events will likely occur at the individual species level.

The pressure of climate change is disproportionately high in mountainous regions, and small changes may push ecosystem processes beyond sensitivity thresholds, creating new dynamics of carbon and nutrient cycling. Given that the rate of organic matter decomposition is strongly dependent upon temperature and soil moisture, the sensitivity of soil respiration to both metrics is highly relevant when considering soil–atmosphere feedbacks under a changing climate. To assess the effects of changing climate in a mountain pasture system, we transplanted turfs along an elevation gradient, monitored in situ soil respiration, incubated collected top-soils to determine legacy effects on temperature sensitivity, and analysed soil organic matter (SOM) to detect changes in quality and quantity of SOM fractions. In situ transplantation down-slope reduced soil moisture and increased soil temperature, with concurrent reductions in soil respiration. Soil moisture acted as an overriding constraint to soil respiration, and significantly reduced the sensitivity to temperature. Under controlled laboratory conditions, removal of the moisture constraint to heterotrophic respiration led to a significant respiration-temperature response. However, despite lower respiration rates down-slope, the response function was comparable among sites, and therefore unaffected by antecedent conditions. We found shifts in the SOM quality, especially of the light fraction, indicating changes to the dynamics of decomposition of recently deposited material. Our findings highlighted the resilience of the microbial community to severe climatic perturbations, but also that soil moisture stress during the growing season can significantly reduce soil function in addition to direct effects on plant productivity. This demonstrated the sensitivity of subalpine pastures under climate change, and possible implications for sustainable use given reductions in organic matter turnover and consequent feedbacks to nutrient cycling., Science of The Total Environment, 473-474, ISSN:0048-9697, ISSN:1879-1026

International audience; Silvopastoral systems of the Swiss Jura Mountains serve as a traditional source of forage and timber in the subalpine vegetation belt, but their vulnerability to land use and climate change puts their future sustainability at stake. We coupled experimental and modeling approaches to assess the impact of climate change on the pasture-woodland landscape. We drew conclusions on the resistance potential of wooded pastures with different management intensities by sampling along a canopy cover gradient. This gradient spanned from unwooded pastures associated with intensive farming to densely wooded pastures associated with extensive farming. Transplanted mesocosms of these ecosystems placed at warmer and drier conditions provided experimental evidence that climate change reduced herbaceous biomass production in unwooded pastures but had no effect in sparsely wooded pastures, and even stimulated productivity in densely wooded pastures. Through modeling these results with a spatially explicit model of wooded pastures (WoodPaM) modified for the current application, results were extrapolated to the local landscape under two regionalized Intergovernmental Panel on Climate Change scenarios for climate change. This led to the suggestion that within the Jura pasture-woodlands, forage production in the near future (2000–2050 AD) would be affected disproportionately throughout the landscape. A stable forage supply in hot, dry years would be provided only by extensive and moderate farming, which allows the development of an insulating tree cover within grazed pastures. We conclude that such structural landscape diversity would grant wood-pastures with a buffering potential in the face of climate change in the forthcoming decades.

National audience; Les pâturages boisés du Jura servent principalement à la production de fourrage et de bois, mais ils sont aussi importants pour leur biodiversité et les espaces de récréation, de même que la qualité du paysage qu’ils offrent à proximité des grandes zones urbaines de l’Arc lémanique. Cet article montre, à l’aide d’une expérience de transplantation de mésocosmes et de modèles de simulation, l’effet des changements climatiques sur la production de biomasse herbacée et, dans ce contexte, les conséquences de la nouvelle réforme agraire (PA 14–17) sur l’évolution du paysage. Les pâturages boisés pourraient mieux résister à l’augmentation des températures et à la diminution des précipitations estivales que les pâturages sans arbres, en gardant une production fourragère plus stable. Le couplage du modèle de la végétation avec un modèle socio-économique indique qu’avec la nouvelle politique agricole (PA 14–17), la pression de pâture va diminuer, et que dans le cadre d’un scénario de réchauffement modéré, cela conduira à la fermeture des paysages. La nouvelle politique agricole devrait permettre de prendre des mesures ciblées pour conserver les pâturages boisés.

openaire: EC/H2020/819202/EU//SOS.aquaterra As global temperatures continue to rise, a key uncertainty of climate projections is the microbial decomposition of vast organic carbon stocks in thawing permafrost soils. Decomposition rates can accelerate up to fourfold in the presence of plant roots, and this mechanism—termed the rhizosphere priming effect—may be especially relevant to thawing permafrost soils as rising temperatures also stimulate plant productivity in the Arctic. However, priming is currently not explicitly included in any model projections of future carbon losses from the permafrost area. Here, we combine high-resolution spatial and depth-resolved datasets of key plant and permafrost properties with empirical relationships of priming effects from living plants on microbial respiration. We show that rhizosphere priming amplifies overall soil respiration in permafrost-affected ecosystems by ~12%, which translates to a priming-induced absolute loss of ~40 Pg soil carbon from the northern permafrost area by 2100. Our findings highlight the need to include fine-scale ecological interactions in order to accurately predict large-scale greenhouse gas emissions, and suggest even tighter restrictions on the estimated 200 Pg anthropogenic carbon emission budget to keep global warming below 1.5 °C.

Climatic changes resulting from anthropogenic activities over the passed century are repeatedly reported to alter the functioning of pristine ecosystems worldwide, and especially those in cold biomes. Available literature on the process of plant leaf litter decomposition in the temperate Alpine zone is reviewed here, with emphasis on both direct and indirect effects of climate change phenomena on rates of litter decay. Weighing the impact of biotic and abiotic processes governing litter mass loss, it appears that an immediate intensification of decomposition rates due to temperature rise can be retarded by decreased soil moisture, insufficient snow cover insulation, and shrub expansion in the Alpine zone. This tentative conclusion, remains speculative unless empirically tested, but it has profound implications for understanding the biogeochemical cycling in the Alpine vegetation belt, and its potential role as a buffering mechanism to climate change.

Peatlands are important sinks of atmospheric carbon (C) that, in response to climate warming, are undergoing dynamic vegetation succession. Here we examined the hypothesis that the uptake of nutrients by different plant growth forms (PGFs) is one key mechanism driving changes in species abundance in peatlands. Along an altitude gradient representing a natural climate experiment, we compared the variability of the stable C isotope composition (delta C-13) and stable nitrogen (N) isotope composition (delta N-15) in current-year leaves of two major PGFs, i.e. ericoids and graminoids. The climate gradient was associated with a gradient of vascular plant cover, which was parallelled by different concentrations of organic and inorganic N as well as the fungal/bacterial ratio in peat. In both PGFs the C-13 natural abundance showed a marginal spatial decrease with altitude and a temporal decrease with progression of the growing season. Our data highlight a primary physical control of foliar delta C-13 signature, which is independent from the PGFs. Natural abundance of foliar N-15 did not show any seasonal pattern and only in the ericoids showed depletion at lower elevation. This decreasing delta N-15 pattern was primarily controlled by the higher relative availability of organic versus inorganic N and, only for the ericoids, by an increased proportion of fungi to bacteria in soil. Our space-for-time approach demonstrates that a change in abundance of PGFs is associated with a different strategy of nutrient acquisition (i.e. transfer via mycorrhizal symbiosis versus direct fine-root uptake), which could likely promote observed and predicted dwarf shrub expansion under climate change.