Plant-soil interactions alter nitrogen and phosphorus dynamics in an advancing subarctic treeline.

Treelines advance due to climate warming. The impacts of this vegetation shift on plant-soil nutrient cycling are still uncertain, yet highly relevant as nutrient availability stimulates tree growth. Here, we investigated nitrogen (N) and phosphorus (P) in plant and soil pools along two tundra-forest transects on Kola Peninsula, Russia, with a documented elevation shift of birch-dominated treeline by 70 m during the last 50 years. Results show that although total N and P stocks in the soil-plant system did not change with elevation, their distribution was significantly altered. With the transition from high-elevation tundra to low-elevation forest, P stocks in stones decreased, possibly reflecting enhanced weathering. In contrast, N and P stocks in plant biomass approximately tripled and available P and N in the soil increased fivefold toward the forest. This was paralleled by decreasing carbon (C)-to-nutrient ratios in foliage and litter, smaller C:N:P ratios in microbial biomass, and lower enzymatic activities related to N and P acquisition in forest soils. An incubation experiment further demonstrated manifold higher N and P net mineralization rates in litter and soil in forest compared to tundra, likely due to smaller C:N:P ratios in decomposing organic matter. Overall, our results show that forest expansion increases the mobilization of available nutrients through enhanced weathering and positive plant-soil feedback, with nutrient-rich forest litter releasing greater amounts of N and P upon decomposition. While the low N and P availability in tundra may retard treeline advances, its improvement toward the forest likely promotes tree growth and forest development.

Controlling biases in targeted plant removal experiments.

Targeted removal experiments are a powerful tool to assess the effects of plant species or (functional) groups on ecosystem functions. However, removing plant biomass in itself can bias the observed responses. This bias is commonly addressed by waiting until ecosystem recovery, but this is inherently based on unverified proxies or anecdotal evidence. Statistical control methods are efficient, but restricted in scope by underlying assumptions. We propose accounting for such biases within the experimental design, using a gradient of biomass removal controls. We demonstrate the relevance of this design by presenting (1) conceptual examples of suspected biases and (2) how to observe and control for these biases. Using data from a mycorrhizal association-based removal experiment, we show that ignoring biomass removal biases (including by assuming ecosystem recovery) can lead to incorrect, or even contrary conclusions (e.g. false positive and false negative). Our gradient design can prevent such incorrect interpretations, regardless of whether aboveground biomass has fully recovered. Our approach provides more objective and quantitative insights, independently assessed for each variable, than using a proxy to assume ecosystem recovery. Our approach circumvents the strict statistical assumptions of, for example, ANCOVA and thus offers greater flexibility in data analysis.

Impacts of snow-farming on alpine soil and vegetation: A case study from the Swiss Alps.

Snow-farming is one of the adaptive strategies used to face the snow deficit in ski resorts. We studied the impact of a shifting snow-farming technique on a pasture slope in Adelboden, Switzerland. Specifically, we compared plots covered by a compressed snow pile for 1.5, 2.5 or 3.5 years, which then recovered from the snow cover for three, two or one vegetation seasons, respectively, with control plots situated around the snow pile. In plots with >1.5 years of compressed snow pile, plant mortality was high, recovery of vegetation was very slow, and few plant species recolonized the bare surface. Soil biological activity decreased persistently under prolonged snow cover, as indicated by reduced soil respiration. The prolonged absence of fresh plant litter and root exudates led to carbon (C) limitation for soil microbial respiration, which resulted in a significant decrease in the ratio of total organic carbon to total nitrogen (TOC/TN) under the snow pile. Microbial C, nitrogen (N) and phosphorus (P) immobilization decreased, while dissolved N concentration increased with compressed snow cover. Longer snow cover and a subsequent shorter recovery period led to higher microbial C/P and N/P but lower microbial C/N. Nitrate and ammonium were released massively once the biological activity resumed after snow clearance and soil aeration. The soil microbial community composition persistently shifted towards oxygen-limited microbes with prolonged compressed snow cover. This shift reflected declines in the abundance of sensitive microorganisms, such as plant-associated symbionts, due to plant mortality or root die-off. In parallel, resistant taxa that benefit from environmental changes increased, including facultative anaerobic bacteria (Bacteroidota, Chloroflexota), obligate anaerobes (Euryarchaeota), and saprophytic plant degraders. We recommend keeping snow piles in the same spot year after year to minimize the area of the impacted soil surface and plan from the beginning soil and ecosystem restoration measures.

Lowland plant arrival in alpine ecosystems facilitates a decrease in soil carbon content under experimental climate warming.

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.

In a terrestrial ecosystem, the carbon cycle primarily represents the balance between plants consuming carbon dioxide from the atmosphere and soil microbes releasing carbon stored in the soil into the atmosphere (mostly as carbon dioxide). Given that carbon dioxide traps heat in the atmosphere, the balance of carbon inputs and outputs from an ecosystem can have important consequences for climate change. Rising temperatures caused by climate warming have led plants from lowland ecosystems to migrate uphill and start growing in alpine ecosystems, where temperatures are lower and most carbon is stored in the soil. Soil microbes use carbon stored in the soil and exuded from plants to grow, and they release this carbon ­ in the form of carbon dioxide ­ into the atmosphere through respiration. Walker et al. wanted to know how the arrival of lowland plants in alpine ecosystems under climate warming would affect carbon stores in the soil. To answer this question, Walker et al. simulated warmer temperatures by moving turfs (plants and soil) from alpine ecosystems to a warmer downhill site and planting lowland plants into the turfs. They compared the concentration of soil carbon in these turfs to that of soil in alpine turfs that had not been moved downhill and had no lowland plants. Their results showed that the warmed turfs containing lowland plants had a lower concentration of soil carbon. This suggests that climate warming will lead to more soil carbon being released into the atmosphere if lowland plants also migrate into alpine ecosystems. Walker et al. also wanted to know the mechanism through which lowland plants were decreasing soil carbon concentration under warming. They find that lowland plants probably release more small molecules into the soil than alpine plants. Soil microbes use the carbon and nutrients in these molecules to break down more complex molecules in the soil, thereby releasing nutrients and carbon that can then be used in respiration. This finding suggests that soil microbes breakdown and respire native soil carbon faster in the presence of lowland plants, releasing more carbon dioxide into the atmosphere and reducing carbon stores in the soil. Walker et al.'s results reveal a new mechanism through which uphill migration of lowland plants could increase the effects of climate change, in a feedback loop. Further research as to whether this mechanism occurs in different regions and ecosystems could help to quantify the magnitude of this feedback and allow scientists to make more accurate predictions about climate change.

Above- and belowground linkages shape responses of mountain vegetation to climate change.

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.

Correction: Long-term in situ permafrost thaw effects on bacterial communities and potential aerobic respiration.

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.

The Legacy Effects of Winter Climate on Microbial Functioning After Snowmelt in a Subarctic Tundra.

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 < 0.1) in the Q10 of microbial respiration and an oxidative EEA. Thus, although warmer winters may induce legacy effects in microbial temperature acclimation, we found no evidence for changes in potential carbon mineralization after spring thaw.

Long-term in situ permafrost thaw effects on bacterial communities and potential aerobic respiration.

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.

Vascular plant-mediated controls on atmospheric carbon assimilation and peat carbon decomposition under climate change.

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-14 C) 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.

Winter ecology of a subalpine grassland: Effects of snow removal on soil respiration, microbial structure and function.

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.

Environmental drivers of carbon and nitrogen isotopic signatures in peatland vascular plants along an altitude gradient.

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 (δ(13)C) and stable nitrogen (N) isotope composition (δ(15)N) 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 (13)C 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 δ(13)C signature, which is independent from the PGFs. Natural abundance of foliar (15)N did not show any seasonal pattern and only in the ericoids showed depletion at lower elevation. This decreasing δ(15)N 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.

Diminished soil functions occur under simulated climate change in a sup-alpine pasture, but heterotrophic temperature sensitivity indicates microbial resilience.

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.

Transplantation of subalpine wood-pasture turfs along a natural climatic gradient reveals lower resistance of unwooded pastures to climate change compared to wooded ones.

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.