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Terrestrial animals are key elements in the cycling of elements in the Arctic where nutrient availability is low. Waste production by herbivores, in particular urine deposition, has a crucial role for nitrogen (N) recycling, still, it remains largely unexplored. Also, experimental evidence is biased toward short-term studies and Arctic regions under high herbivore pressure. In this study, we aimed to examine the fate of N derived from urine in a nutrient poor tundra heath in West Greenland, with historical low level of herbivory. We performed a pulse labelling with 15N-urea over the plant canopy and explored ecosystem N partition and retention in the short-term (2 weeks and 1 year) and longer-term (5 years). We found that all vascular plants, irrespective of their traits, could rapidly take up N-urea, but mosses and lichens were even more efficient. Total 15N enrichment was severely reduced for all plants 5 years after tracer addition, with the exception of cryptogams, indicating that non-vascular plants constituted a long-term sink of 15N-urea. The 15N recovery was also high in the litter suggesting high N immobilization in this layer, potentially delaying the nutrients from urine entering the soil compartment. Long-term 15N recovery in soil microbial biomass was minimal, but as much as 30% of added 15N remained in the non-microbial fraction after 5 years. Our results demonstrate that tundra plants that have evolved under low herbivory pressure are well adapted to quickly take advantage of labile urea, with urine having only a transient effect on soil nutrient availability. [ABSTRACT FROM AUTHOR]

Increasing temperatures and winter precipitation can influence the carbon (C) exchange rates in arctic ecosystems. Feedbacks can be both positive and negative, but the net effects are unclear and expected to vary strongly across the Arctic. There is a lack of understanding of the combined effects of increased summer warming and winter precipitation on the C balance in these ecosystems. Here we assess the short‐term (1–3 years) and long‐term (5–8 years) effects of increased snow depth (snow fences) (on average + 70 cm) and warming (open top chambers; 1–3°C increase) and the combination in a factorial design on all key components of the daytime carbon dioxide (CO2) fluxes in a wide‐spread heath tundra ecosystem in West Greenland. The warming treatment increased ecosystem respiration (ER) on a short‐ and long‐term basis, while gross ecosystem photosynthesis (GEP) was only increased in the long term. Despite the difference in the timing of responses of ER and GEP to the warming treatment, the net ecosystem exchange (NEE) of CO2 was unaffected in the short term and in the long term. Although the structural equation model (SEM) indicates a direct relationship between seasonal accumulated snow depth and ER and GEP, there were no significant effects of the snow addition treatment on ER or GEP measured over the summer period. The combination of warming and snow addition turned the plots into net daytime CO2 sources during the growing season. Interestingly, despite no significant changes in air temperature during the snow‐free time during the experiment, control plots as well as warming plots revealed significantly higher ER and GEP in the long term compared to the short term. This was in line with the satellite‐derived time‐integrated normalized difference vegetation index of the study area, suggesting that more factors than air temperature are drivers for changes in arctic tundra ecosystems. [ABSTRACT FROM AUTHOR]

As arctic permafrost continues to thaw, previously inaccessible nitrogen (N) becomes available to N‐limited arctic plants. Increased N availability could enhance plant growth and thereby potentially offset climate‐induced carbon release. Arctic plants can take up newly available permafrost‐N locally upon release. However, in a topographically diverse arctic landscape, permafrost‐N may be transported along hillslopes, away from the point‐of‐release. The extent to which topographical N transport can impact arctic vegetation change depends on whether N is retained locally, captured by downslope recipient plant communities, or transported away.We used stable isotope labelling (15N) to simulate upslope release of ammonium (NH4+) and nitrate (NO3−) from thawing permafrost on an arctic hillslope, western Greenland. We tracked the plant species‐specific uptake of simulated permafrost‐released N from the upslope point‐of‐release to the bottom of the slope through 4 years.We found that arctic tundra plants successfully acquired locally released permafrost‐N, even in sloping terrain, and that N was strongly retained in the plant–soil system through multiple years. At the same time, we also importantly demonstrate that permafrost‐N can be transported and taken up by plants up to 30 m downslope from the point‐of‐release. Especially NO3− was more easily redistributed vertically within the soil column compared to NH4+ and therefore potentially more accessible to plants. Specifically, plant species with fast N uptake capacity and deep‐soil foraging strategies may have competitive advantages for capitalising on deep‐soil released and topographically transported permafrost‐N (here exemplified by Equisetum arvense and Salix glauca). Nevertheless, even mosses gained access to permafrost‐N via vertical and lateral redistribution on the slope. Ultimately, the intricate balance between strong local N retention, downslope transport and plant species‐specific uptake strategies may contribute to shaping arctic vegetation change.Synthesis. Across spatially complex arctic ecosystems, arctic plants can take up permafrost‐released N both at the local point‐of‐release and at a considerable distance downslope. The potential for arctic plants to take advantage of topographically transported permafrost‐N could lead to long‐term and landscape‐scale changes in species composition, plant productivity, and ultimately carbon and climate feedbacks across the Arctic. [ABSTRACT FROM AUTHOR]

Background and Aims Climate change is subjecting subarctic ecosystems to elevated temperature, increased nutrient availability and reduced light availability (due to increasing cloud cover). This may affect subarctic vegetation by altering the emissions of biogenic volatile organic compounds (BVOCs) and leaf anatomy. We investigated the effects of increased nutrient availability on BVOC emissions and leaf anatomy of three subarctic dwarf shrub species, Empetrum hermaphroditum , Cassiope tetragona and Betula nana , and if increased nutrient availability modifies the responses to warming and shading. Methods Measurements of BVOCs were performed in situ in long-term field experiments in the Subarctic using a dynamic enclosure system and collection of BVOCs into adsorbent cartridges analysed by gas chromatography–mass spectrometry. Leaf anatomy was studied using light microscopy and scanning electron microscopy. Key Results Increased nutrient availability increased monoterpene emission rates and altered the emission profile of B. nana , and increased sesquiterpene and oxygenated monoterpene emissions of C. tetragona. Increased nutrient availability increased leaf tissue thicknesses of B. nana and C. tetragona , while it caused thinner epidermis and the highest fraction of functional (intact) glandular trichomes for E. hermaphroditum. Increased nutrient availability and warming synergistically increased mesophyll intercellular space of B. nana and glandular trichome density of C. tetragona , while treatments combining increased nutrient availability and shading had an opposite effect in C. tetragona. Conclusions Increased nutrient availability may enhance the protection capacity against biotic and abiotic stresses (especially heat and drought) in subarctic shrubs under future warming conditions as opposed to increased cloudiness, which could lead to decreased resistance. The study emphasizes the importance of changes in nutrient availability in the Subarctic, which can interact with climate warming and increased cloudiness effects. [ABSTRACT FROM AUTHOR]

Understanding N budgets of tundra ecosystems is crucial for projecting future changes in plant community composition, greenhouse gas balances and soil N stocks. Winter warming can lead to higher tundra winter nitrogen (N) mineralization rates, while summer warming may increase both growing season N mineralization and plant N demand. The undulating tundra landscape is inter-connected through water and solute movement on top of and within near-surface soil, but the importance of lateral N fluxes for tundra N budgets is not well known. We studied the size of lateral N fluxes and the fate of lateral N input in the snowmelt period with a shallow thaw layer, and in the late growing season with a deeper thaw layer. We used 15N to trace inorganic lateral N movement in a Low-arctic mesic tundra heath slope in West Greenland and to quantify the fate of N in the receiving area. We found that half of the early-season lateral N input was retained by the receiving ecosystem, whereas half was transported downslope. Plants appear as poor utilizers of early-season N, indicating that higher winter N mineralization may influence plant growth and carbon (C) sequestration less than expected. Still, evergreen plants were better at utilizing early-season N, highlighting how changes in N availability may impact plant community composition. In contrast, later growing season lateral N input was deeper and offered an advantage to deeper-rooted deciduous plants. The measurements suggest that N input driven by future warming at the study site will have no significant impact on the overall N2O emissions. Our work underlines how tundra ecosystem N allocation, C budgets and plant community composition vary in their response to lateral N inputs, which may help us understand future responses in a warmer Arctic. [ABSTRACT FROM AUTHOR]

Traditionally, biogenic volatile organic compound (BVOC) emissions are often considered a unidirectional flux, from the ecosystem to the atmosphere, but recent studies clearly show the potential for bidirectional exchange. Here we aimed to investigate how warming and leaf litter addition affect the bidirectional exchange (flux) of BVOCs in a long‐term field experiment in the Subarctic. We also assessed changes in net BVOC fluxes in relation to the time of day and the influence of different plant phenological stages. The study was conducted in a full factorial experiment with open top chamber warming and annual litter addition treatments in a tundra heath in Abisko, Northern Sweden. After 18 years of treatments, ecosystem‐level net BVOC fluxes were measured in the experimental plots using proton‐transfer‐reaction time‐of‐flight mass spectrometry (PTR–ToF–MS). The warming treatment increased monoterpene and isoprene emissions by ≈50%. Increasing temperature, due to diurnal variations, can both increase BVOC emission and simultaneously, increase ecosystem uptake. For any given treatment, monoterpene, isoprene, and acetone emissions also increased with increasing ambient air temperatures caused by diurnal variability. Acetaldehyde, methanol, and sesquiterpenes decreased likely due to a deposition flux. For litter addition, only a significant indirect effect on isoprene and monoterpene fluxes (decrease by ~50%–75%) was observed. Litter addition may change soil moisture conditions, leading to changes in plant species composition and biomass, which could subsequently result in changes to BVOC emission compositions. Phenological stages significantly affected fluxes of methanol, isoprene and monoterpenes. We suggest that plant phenological stages differ in impacts on BVOC net emissions, but ambient air temperature and photosynthetically active radiation (PAR) also interact and influence BVOC net emissions differently. Our results may also suggest that BVOC fluxes are not only a response to changes in temperature and light intensity, as the circadian clock also affects emission rates. [ABSTRACT FROM AUTHOR]

Arctic and subarctic ecosystems are experiencing substantial changes in hydrology, vegetation, permafrost conditions, and carbon cycling, in response to climatic change and other anthropogenic drivers, and these changes are likely to continue over this century. The total magnitude of these changes results from multiple interactions among these drivers. Field measurements can address the overall responses to different changing drivers, but are less capable of quantifying the interactions among them. Currently, a comprehensive assessment of the drivers of ecosystem changes, and the magnitude of their direct and indirect impacts on subarctic ecosystems, is missing. The Torneträsk area, in the Swedish subarctic, has an unrivalled history of environmental observation over 100 years, and is one of the most studied sites in the Arctic. In this study, we summarize and rank the drivers of ecosystem change in the Torneträsk area, and propose research priorities identified, by expert assessment, to improve predictions of ecosystem changes. The research priorities identified include understanding impacts on ecosystems brought on by altered frequency and intensity of winter warming events, evapotranspiration rates, rainfall, duration of snow cover and lake-ice, changed soil moisture, and droughts. This case study can help us understand the ongoing ecosystem changes occurring in the Torneträsk area, and contribute to improve predictions of future ecosystem changes at a larger scale. This understanding will provide the basis for the future mitigation and adaptation plans needed in a changing climate. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Aims: Plant growth in the Arctic is often nutrient limited due to temperature constraints on decomposition and low atmospheric input of nitrogen (N). Local hotspots of nutrient enrichment found in up to 4000-year-old archaeological deposits can be used to explore the recycling and long-term retention of nutrients in arctic ecosystems. Methods: We investigated old Inuit and Norse deposits (known as middens) and adjacent tundra ecosystems along a wet-dry fjord gradient in western Greenland to explore the isotopic fingerprinting of plant and soil carbon and nitrogen (13C/12C and 15N/14N) derived from human presence. Results: At all locations we observed a significant isotopic fingerprint in soil and plant N related to human deposits. This demonstrates a century-long legacy of past human habitation on plant and soil characteristics and indicates a surprisingly high N retention in these ecosystems. This is consistent with the significantly higher plant biomass in areas with archaeological deposits. Conclusion: Vegetation composition and N in plants and soils displayed marked differences along the wet-dry fjord gradient. Furthermore, the profound nutrient enrichment and organic matter accumulation in archaeological deposits compared to surrounding tundra demonstrates a century-long legacy of past habitation on plant and soil characteristics as well as efficient N cycling with surprisingly limited N loss. [ABSTRACT FROM AUTHOR]

Climate warming enables tree seedling establishment beyond the current alpine treeline, but to achieve this, seedlings have to establish within existing tundra vegetation. In tundra, mosses are a prominent feature, known to regulate soil temperature and moisture through their physical structure and associated water retention capacity. Moss presence and species identity might therefore modify the impact of increases in temperature and precipitation on tree seedling establishment at the arctic‐alpine treeline. We followed Betula pubescens and Pinus sylvestris seedling survival and growth during three growing seasons in the field. Tree seedlings were transplanted along a natural precipitation gradient at the subarctic‐alpine treeline in northern Sweden, into plots dominated by each of three common moss species and exposed to combinations of moss removal and experimental warming by open‐top chambers (OTCs). Independent of climate, the presence of feather moss, but not Sphagnum, strongly supressed survival of both tree species. Positive effects of warming and precipitation on survival and growth of B. pubescens seedlings occurred in the absence of mosses and as expected, this was partly dependent on moss species. P. sylvestris survival was greatest at high precipitation, and this effect was more pronounced in Sphagnum than in feather moss plots irrespective of whether the mosses had been removed or not. Moss presence did not reduce the effects of OTCs on soil temperature. Mosses therefore modified seedling response to climate through other mechanisms, such as altered competition or nutrient availability. We conclude that both moss presence and species identity pose a strong control on seedling establishment at the alpine treeline, and that in some cases mosses weaken climate‐change effects on seedling establishment. Changes in moss abundance and species composition therefore have the potential to hamper treeline expansion induced by climate warming. [ABSTRACT FROM AUTHOR]

Over the past few decades the Arctic has warmed up more than the lower latitudes. Soil organic carbon (SOC) in the Arctic is vulnerable to climate change, and carbon dioxide (CO2) produced via SOC decomposition can amplify atmospheric temperature increase. Although SOC composition is relevant to decomposability, studies on its compositional changes with warming are scarce, particularly in the Arctic. Therefore, we investigated the responses of SOC and the bacterial community to climate manipulation under Cassiope and Salix heath vegetation communities in permafrost‐affected soil in Zackenberg, Greenland. After 8–9 years of experimental warming, we evaluated changes in SOC quantity and quality of three density fractions of soil: free light fraction (FLF), occluded light fraction (OLF) and heavy fraction (HF). The SOC content at 0–5‐cm depth was significantly reduced with warming under Cassiope, and it was accompanied by decreased FLF content, attributed to accelerated decomposition of the FLF by warming. However, SOC molecular composition and bacterial community composition were not affected by warming. By contrast, there was no warming effect on SOC under Salix, which could be partially due to smaller temperature increases caused by higher moisture levels associated with larger silt and clay contents, or to different responses of the dominant plant species to temperature. In both soils, more than 55% of SOC was associated with minerals, and its molecular composition indicated microbial decomposition. Our results suggested that long‐term warming in the high Arctic could induce the loss of SOC, particularly in the FLF; however, the response could vary with vegetation type and/or soil properties, that is, soil texture. Highlights: We show decreased SOC with long‐term (8‐year) warming of heath soils in the high ArcticParticularly, the free light fraction of SOC in topsoil decreased with warming in a Cassiope heath siteMineral‐associated SOC content was more than 55% and showed signs of microbial processingEffects of warming differed according to soil properties and/or plant community composition [ABSTRACT FROM AUTHOR]

Background and Aims Climate change in the Arctic is projected to increase temperature, precipitation and snowfall. This may alter leaf anatomy and gas exchange either directly or indirectly. Our aim was to assess whether increased snow depth and warming modify leaf anatomy and affect biogenic volatile organic compound (BVOC) emissions and CO2 exchange of the widespread arctic shrubs Betula nana and Empetrum nigrum ssp. hermaphroditum. Methods Measurements were conducted in a full-factorial field experiment in Central West Greenland, with passive summer warming by open-top chambers and snow addition using snow fences. Leaf anatomy was assessed using light microscopy and scanning electron microscopy. BVOC emissions were measured using a dynamic enclosure system and collection of BVOCs into adsorbent cartridges analysed by gas chromatography–mass spectrometry. Carbon dioxide exchange was measured using an infrared gas analyser. Key Results Despite a later snowmelt and reduced photosynthesis for B. nana especially, no apparent delays in the BVOC emissions were observed in response to snow addition. Only a few effects of the treatments were seen for the BVOC emissions, with sesquiterpenes being the most responsive compound group. Snow addition affected leaf anatomy by increasing the glandular trichome density in B. nana and modifying the mesophyll of E. hermaphroditum. The open-top chambers thickened the epidermis of B. nana, while increasing the glandular trichome density and reducing the palisade:spongy mesophyll ratio in E. hermaphroditum. Conclusions Leaf anatomy was modified by both treatments already after the first winter and we suggest links between leaf anatomy, CO2 exchange and BVOC emissions. While warming is likely to reduce soil moisture, melt water from a deeper snow pack alleviates water stress in the early growing season. The study emphasizes the ecological importance of changes in winter precipitation in the Arctic, which can interact with climate-warming effects. [ABSTRACT FROM AUTHOR]

How species interact modulate their dynamics, their response to environmental change, and ultimately the functioning and stability of entire communities. Work conducted at Zackenberg, Northeast Greenland, has changed our view on how networks of arctic biotic interactions are structured, how they vary in time, and how they are changing with current environmental change: firstly, the high arctic interaction webs are much more complex than previously envisaged, and with a structure mainly dictated by its arthropod component. Secondly, the dynamics of species within these webs reflect changes in environmental conditions. Thirdly, biotic interactions within a trophic level may affect other trophic levels, in some cases ultimately affecting land-atmosphere feedbacks. Finally, differential responses to environmental change may decouple interacting species. These insights form Zackenberg emphasize that the combination of long-term, ecosystem-based monitoring, and targeted research projects offers the most fruitful basis for understanding and predicting the future of arctic ecosystems. [ABSTRACT FROM AUTHOR]

The muskox ( Ovibos moschatus), a key species in the arctic tundra, is the only large-bodied herbivore in Northeast Greenland. Here, we quantify the biomass removal and fecal deposition by muskoxen during the snow-free period in the years 1996 to 2013 in the high arctic tundra ecosystem at Zackenberg, Northeast Greenland. We show that despite high densities, muskoxen removed only 0.17% and 0.04% of the available forage in graminoid-dominated areas and in Salix snowbeds (including Salix dominated heaths), respectively, during the main plant growing season (from mid-June to end of August). Into the autumn, the biomass removal increased to ∼4.6% and 0.19% in the graminoid-dominated areas and Salix snowbeds, respectively. Muskoxen forage mainly in the graminoid-dominated areas, but defecate primarily in Salix snowbeds, resulting in net nutrient transfers from the nitrogen-rich wet habitats to the nitrogen-poor, drier habitats, corresponding to an addition of 0.016 g m-2 of nitrogen in the Salix snowbeds per year. This nitrogen addition is of same magnitude as the dissolved inorganic nitrogen pool in similar arctic soils. Hence, while the quantitative impact of muskox biomass removal seems negligible, the nitrogen relocation may be important for the arctic vegetation and associated biota in the tundra ecosystem. [ABSTRACT FROM AUTHOR]

Nitrogen (N) mineralization, nutrient availability, and plant growth in the Arctic are often restricted by low temperatures. Predicted increases of cold-season temperatures may be important for plant nutrient availability and growth, given that N mineralization is also taking place during the cold season. Changing nutrient availability may be reflected in plant N and chlorophyll content and lead to increased photosynthetic capacity, plant growth, and ultimately carbon (C) assimilation by plants. In this study, we increased snow depth and thereby cold-season soil temperatures in high Arctic Svalbard in two vegetation types spanning three moisture regimes. We measured growing-season availability of ammonium (NH), nitrate (NO), total dissolved organic carbon (DOC) and nitrogen (TON) in soil; C, N, δN and chlorophyll content in Salix polaris leaves; and leaf sizes of Salix, Bistorta vivipara, and Luzula arcuata at peak season. Nutrient availability was significantly higher with increased snow depth in the two mesic meadow vegetation types, but not in the drier heath vegetation. Nitrogen concentrations and δN values of Salix leaves were significantly higher in all vegetation types, but the leaf sizes were unchanged. Leaves of Bistorta and Luzula were significantly larger but only significantly so in one moist vegetation type. Increased N and chlorophyll concentrations in leaves indicate a potential for increased growth (C uptake), supported by large leaf sizes for some species. Responses to cold-season soil warming are vegetation type- and species-specific, with potentially stronger responses in moister vegetation types. This study therefore highlights the contrasting effect of snow in a tundra landscape and has important implications for projections of whole tundra responses to climate change. [ABSTRACT FROM AUTHOR]

Biogenic volatile organic compounds (BVOCs) emitted from terrestrial vegetation participate in oxidative reactions in the atmosphere, leading to the formation of secondary organic aerosols and longer lifetime of methane. Global models of BVOC emissions have assumed minimal emissions from the high latitudes. However, measurements from this region are lacking, and studies from the high arctic are yet to be published. This study aimed to obtain estimates for BVOC emissions from the high arctic, and hereby to add new knowledge to the understanding of global BVOC emissions. Measurements were conducted in four vegetation types dominated by Cassiope tetragona, Salix arctica, Vaccinium uliginosum and a mixture of Kobresia myosuroides, Dryas spp. and Poa arctica. Emissions were measured by an enclosure technique and collection of volatiles into adsorbent cartridges in August. Volatiles were analyzed by gas chromatography-mass spectrometry following thermal desorption. Isoprene showed highest emissions in S. arctica heath. Monoterpene and sesquiterpene emissions were especially associated with C. tetragona heath. Total observed emissions were comparable in magnitude to emissions previously found in the subarctic, whereas isoprene emissions were lower. This study shows that considerable amounts of BVOCs are emitted from the high arctic. The results are also of importance as the emissions from this region are expected to increase in the future as a result of the predicted climate warming in the high arctic. We suggest further studies to assess the effects of climate changes in the region in order to gain new knowledge and understanding of future global BVOC emissions. [ABSTRACT FROM AUTHOR]

Arctic plants and herbivores are subject to ongoing climatic changes that are more rapid and extreme than elsewhere on the planet, and thus it is pivotal to understand the arctic plant-herbivore interactions in a global change context. We examined how infestation by an eriophyoid gall mite affects the circumpolar shrub Salix arctica, and how the effects vary across vegetation types. Specifically, we compared multiple leaf characteristics (leaf area, biomass, nutrient levels, δ 15N and δ 13C, and stress and performance of the photosynthetic apparatus) of infested leaves to those of un-infested leaves. Furthermore, we examined how altered environmental conditions, here experimentally manipulated levels of temperature, water and nutrients, shading, and UV-B radiation, affect the prevalence, density, and intensity of gall mite infestation and its impacts on S. arctica. Infested leaves were smaller in area and biomass and had lower nitrogen and carbon pools. However, their carbon concentration was higher, possibly because the galls acted as carbon sinks. The smaller photosynthetic area and lower nutrient content caused increased stress on the photosynthetic apparatus in infested leaves. The remaining leaf tissue responded with a higher photosynthetic performance, although there were indications of a general reduction in photosynthesis. Female leaves were more affected than male leaves. The experimental manipulations of environmental conditions did not affect the gall prevalence, density, or intensity on S. arctica leaves. Rather, plants responded positively to the treatments, reducing the effects of the galls to in-significance. This suggests a higher tolerance and defense against gall mites under future climate conditions. [ABSTRACT FROM AUTHOR]

Thawing permafrost represents a poorly understood feedback mechanism of climate change in the Arctic, but with a potential impact owing to stored carbon being mobilized. We have quantified the long-term loss of carbon (C) from thawing permafrost in Northeast Greenland from 1996 to 2008 by combining repeated sediment sampling to assess changes in C stock and >12 years of CO2 production in incubated permafrost samples. Field observations show that the active-layer thickness has increased by >1 cm yr−1 but thawing has not resulted in a detectable decline in C stocks. Laboratory mineralization rates at 5 °C resulted in a C loss between 9 and 75%, depending on drainage, highlighting the potential of fast mobilization of permafrost C under aerobic conditions, but also that C at near-saturated conditions may remain largely immobilized over decades. This is confirmed by a three-pool C dynamics model that projects a potential C loss between 13 and 77% for 50 years of incubation at 5 °C. [ABSTRACT FROM AUTHOR]

Climate change may alter the terrestrial ecosystem carbon balance in the Arctic, and previous studies have emphasized the importance of cold season gas exchange when considering the annual carbon balance. Here, we examined gross ecosystem production (GEP), ecosystem respiration ( R) and net ecosystem exchange (NEE) during autumn at a high arctic dry open heath, over a period where air temperatures decreased from +9.8 to −16.5°C. GEP declined by 95-100% during autumn but GEP significantly different from 0 was measured on October 8 despite sub-zero temperatures. R declined by 90% and dominated NEE throughout the study as the ecosystem on all measurement days was a source of atmospheric CO. We estimated net September carbon losses (NEE) to be 17 g CO m, emphasizing the importance of autumn in relation to annual carbon budgets. The study site has been subjected to 14 summers of water addition, and occasional pulses of nitrogen (N) addition in a fully factorial design. N addition enhanced GEP up to 17-fold during September, although there was no effect in October when GEP was very low. Summer water addition decreased autumn R by up to 25%. Both N amendment and water addition decreased carbon loss, that is, increased NEE; N amendment increased NEE on all dates by 13-64% whereas water addition increased NEE by 20-54% late in September and onward, demonstrating the importance of nutrient and water availability on carbon balance in high arctic tundra, also during the autumn freeze-in. [ABSTRACT FROM AUTHOR]

The Arctic biome is subject to large climatic changes. Pivotal to our understanding of the ecological consequences of these changes is the understanding of how the vegetation responds to the altered abiotic conditions. To examine this, we conducted detailed vegetation composition analyses in permanent, replicated plots in eight widespread, distinct plant communities within the Zackenberg Valley in high Arctic Greenland in 1997 and 2008. This period was characterised by a considerable increase in summer temperature and declining spring snow cover. Across plant communities, we found substantial changes in cover of several functional groups and species, biodiversity and community structure. The fen area in particular had markedly changed, most likely in response to a decreasing supply of melt water. Other vegetation types, however, seemed resistant towards the changed environmental conditions. Hence, while some general patterns emerged, the response differed markedly between plant communities, and often with contrasting responses. Our results stress that although some high Arctic plant communities exhibit a relatively high degree of resistance towards environmental changes, the derived ecological effects of climate change in other plant communities may be substantial even on a decadal scale, and, hence, occur at a fast pace. [ABSTRACT FROM AUTHOR]

The long-term and diurnal responses of photosystem II (PSII) performance to near-ambient UV-B radiation were investigated in High Arctic Betula nana. We conducted an UV exclusion experiment with five replicated blocks consisting of open control (no filter), photosynthetic active radiation and UV-B transparent filter control (Teflon), UV-B-absorbing filter (Mylar) and UV-AB-absorbing filter (Lexan). Ethylenediurea (EDU), a chemical normally used to protect plants against ozone injury, was sprayed on the leaves both in the field and in an additional laboratory study to investigate if EDU mitigated the effects of UV-B. Chlorophyll- a fluorescence induction curves were used for analysis of OJIP test parameters. Near-ambient UV-B radiation reduced across season maximum quantum yield (TRo/ABS = Fv/Fm), approximated number of active PSII reaction center (RC/ABS) and the performance index (PIABS), despite improved leaf screening against UV-B with higher content of UV-B-absorbing compounds and a lower specific leaf area. EDU application counteracted the negative impact of UV-B on TRo/ABS, RC/ABS and PIABS. This indicates that the mechanisms behind UV-B and ozone damage share some common features. The midday depression was present in all treatments, but TRo/ABS and PIABS were persistently lower in near-ambient UV-B compared to UV-B reduction. The recovery phase was particularly impaired in near-ambient UV-B and interactive effects between treatment × hour raised TRo/ABS, RC/ABS and PIABS higher in reduced UV-B compared to near-ambient UV-B. This demonstrates current solar UV-B to reduce the PSII performance both on a daily as well as a seasonal basis in this High Arctic species. [ABSTRACT FROM AUTHOR]

Climate warming will induce changes in Arctic ecosystem carbon balance, but besides climate, nitrogen availability is a critical controlling factor of carbon cycling. It is therefore essential to obtain knowledge on the influence of a changing climate on nitrogen fixation, as this process is the main source of new nitrogen to arctic ecosystems. In order to gain information on future nitrogen fixation rates in a changing climate, we studied the effects of two decades of warming with passive greenhouses, shading with sackcloth, and fertilization with NPK fertilizer on nitrogen fixation rates. To expand the knowledge on species-specific responses, we measured nitrogen fixation associated with two moss species: Hylocomium splendens and Aulacomnium turgidum. Our expectations of decreased nitrogen fixation rates in the fertilizer and shading treatments were met. However, contrary to our expectation of increased nitrogen fixation in the warming treatment, we observed either no change ( Hylocomium) or a decrease ( Aulacomnium) in fixation in the warmed plots. We hypothesize that this could be due to moss-specific responses or to long-term induced effects of the warming. For example, we observed that the soil temperature increase induced by the warming treatment was low and insignificant as vegetation height and total vascular plant cover of the warmed plots increased, and moss cover decreased. Hence, truly long-term studies lasting more than two decades provide insights on changes in key biogeochemical processes, which differ from more transient responses to warming in the Arctic. [ABSTRACT FROM AUTHOR]

The Arctic is extremely vulnerable to projected climate change, and global warming may result in major community reorganizations. The aim of this study was a thorough investigation of plant biomass production throughout an entire growing season in five different high arctic vegetation types: Cassiope, Dryas, and Salix heath, grassland, and fen. The main focus was on the gross ecosystem production (GEP), and the biotic and abiotic factors which may influence GEP. Photosynthesis, aboveground biomass, and carbon, nitrogen, and chlorophyll content were measured weekly during nine weeks. There were large differences in seasonal growth and production within and among vegetation types. Mosses contributed considerably to the total C and N pool in grassland, fen, and Salix heath. Fen, which had the highest pool of leaf N, leaf chlorophyll, and moss N, was the most productive vegetation type in terms of GEP, despite the lowest total biomass. Across vegetation types, leaf biomass, leaf N, and moss N pool size influenced GEP. Within most vegetation types GEP correlated with leaf N, in correspondence with the notion that N may limit plant production in many high arctic ecosystems. The timing of the peaks in C and N pools in leaves did not coincide with that in the mosses and in woody tissues. This emphasizes the importance of sampling throughout the growing season, when using field data from the Arctic to estimate plant biomasses and modeling C and N fluxes and pool sizes. [ABSTRACT FROM AUTHOR]

While substantial cold-season respiration has been documented in most arctic and alpine ecosystems in recent years, the significance of cold-season photosynthesis in these biomes is still believed to be small. In a mesic, subartic heath during both the cold and warm season, we measured in situ ecosystem respiration and photosynthesis with a chamber technique at ambient conditions and at artificially increased frequency of freeze–thaw (FT) cycles during fall and spring. We fitted the measured ecosystem exchange rates to respiration and photosynthesis models with R2-values ranging from 0.81 to 0.85. As expected, estimated cold-season (October, November, April and May) respiration was significant and accounted for at least 22% of the annual respiratory CO2 flux. More surprisingly, estimated photosynthesis during this period accounted for up to 19% of the annual gross CO2 uptake, suggesting that cold-season photosynthesis partly balanced the cold-season respiratory carbon losses and can be significant for the annual cycle of carbon. Still, during the full year the ecosystem was a significant net source of 120 ± 12 g C m−2 to the atmosphere. Neither respiration nor photosynthetic rates were much affected by the extra FT cycles, although the mean rate of net ecosystem loss decreased slightly, but significantly, in May. The results suggest only a small response of net carbon fluxes to increased frequency of FT cycles in this ecosystem. [ABSTRACT FROM AUTHOR]

Recent evidence suggests that biogeochemical processes in the Arctic during late winter and spring-thaw strongly affect the annual cycling of carbon and nutrients, indicating high susceptibility to climate change. We therefore examined the carbon and nutrient dynamics in a sub-arctic heath and a birch forest with high temporal resolution from March until snowmelt at both ambient and experimentally increased snow depths. Ecosystem respiration (ER) from mid-March to snowmelt at ambient snow was high, reaching 99 ± 19 (birch) and 67 ± 1.4 g C m-2 (heath). Enhanced snow depth by about 20-30 cm increased ER by 77-157% during late winter but had no effects during spring-thaw. ER rates at the birch site were poorly described by classic first order exponential models (R² = 0.06-0.10) with temperature as a single variable, but model fit improved considerably by including the supply of dissolved organic carbon (DOC) or nitrogen (DON) in the model (R² = 0.40-0.47). At the heath, model fit with temperature as the single variable was better (R² = 0.38-0.52), yet it improved when the supply of DOC or DON was included (R² = 0.65-0.72). Microbial carbon decreased by 43% within a few days after the first soil freeze-thaw event, while microbial nitrogen and phosphorus decreased more slowly. Because soil inorganic nitrogen and phosphorus concentrations were low, nutrients released from lysed microbial cells may have been sequestered by surviving microbes or by plants resuming growth. The fast change in microbial biomass and the dependence of ER on substrate availability stress the need for high temporal resolution in future research on ecosystem carbon and nutrient dynamics at snowmelt in order to make robust models of their turnover. [ABSTRACT FROM AUTHOR]

Soil microbial biomass in arctic heaths has been shown to be largely unaffected by treatments simulating climate change with temperature, nutrient and light manipulations. Here, we demonstrate that more than 10 years is needed for development of significant responses, and that changes in microbial biomass are accompanied with strong alterations in microbial community composition. In contrast to slight or nonsignificant responses after 5, 6 and 10 treatment years, 15 years of inorganic NPK fertilizer addition to a subarctic heath had strong effects on the microbial community and, as observed for the first time, warming and shading also led to significant responses, often in opposite direction to the fertilization responses. The effects were clearer in the top 5 cm soil than at the 5–10 cm depth. Fertilization increased microbial biomass C and more than doubled microbial biomass P compared to the non-fertilized plots. However, it only increased microbial biomass N at the 5–10 cm depth. Fertilization increased fungal biomass and the relative abundance of phospholipid fatty acid (PLFA) markers of gram-positive bacteria. Warming and shading decreased the relative abundance of fungal PLFAs, and shading also altered the composition of the bacterial community. The long time lag in responses may be associated with indirect effects of the gradual changes in the plant biomass and community composition. The contrasting responses to warming and fertilization treatments show that results from fertilizer addition may not be similar to the effects of increased nutrient mineralization and availability following climatic warming. [ABSTRACT FROM AUTHOR]

It is predicted that Arctic regions may experience an increase in mean temperature in the future. This will affect the frequency of severe climatic events such as summer droughts and freeze–thaw cycles. In order to understand the impact of recurring freezing and thawing on soil organisms and their environment, intact plant–soil samples from the sub-Arctic were subjected to a series of such events. Springtail and mite species composition and abundance were monitored at intervals throughout the experiment. Furthermore, nutrient content and mobilisation in the soil and soil microbial biomass and nutrient content were examined.There was no conclusive evidence that recurring freeze–thaw events had a negative effect on the investigated soil faunal groups, and the treatment even seemed to stimulate the abundance of Acaridida. Respiration of soil subjected to 16 freeze–thaw cycles was low when kept at -2 °C and high when kept at +2 °C, indicating rapid response of microbial activity even after long exposure to low and fluctuating temperatures. Oribatida and Gamasida displayed a higher abundance in controls kept at -2 °C for up to 80 days, compared to controls at +2 °C and the freeze–thaw treatment. The Collembola were unaffected by the temperature treatments, but increased in abundance over time. The microbial C:N ratio increased after 40 days at -2 °C, indicating a higher degree of fungal dominance and lower tolerance of bacteria to constant freezing, but not to freeze–thaw. The decline in inorganic and microbial P during the experiment, and the proportionally stronger decrease of inorganic and microbial P than N in frozen soil compared to +2 °C soil, suggests that P is affected more than N mineralisation by freezing. [Copyright &y& Elsevier]

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 (N 2 O) 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 N 2 O 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 N 2 O 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 N 2 O averaged 1.7 μg N 2 O–N m−2 h−1 (range −3.6 to 10.5 μg N 2 O–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, N 2 O fluxes decreased, likely caused by combined negative effects of low summer temperatures and high soil moisture. Positive correlations between N 2 O and nitrate (NO 3 −) and dissolved organic N (DON) suggested that the availability of N was the main controlling variable along the snowmelt gradient. The maximum N 2 O fluxes were observed in the second half of August associated with high NO 3 − concentrations. The effect of growing season experimental warming on N 2 O surface flux varied along the snowmelt gradient and with time. Generally, the experimental warming stimulated N 2 O fluxes under conditions with increased concentrations of inorganic N. In contrast, warming reduced N 2 O 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 N 2 O 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. [Display omitted] • We studied effects of warming on tundra N 2 O emissions along a snowmelt gradient. • The N 2 O emissions peaked with late snowmelt, likely due to high dissolved soil N. • Areas with highest snow depth (4 m) exhibited low N 2 O emissions (low temperature). • Experimental warming enhanced N 2 O emissions, where soil N was high (late snowmelt). • We conclude that low Arctic heath is a source to N 2 O regulated by dissolved soil N. [ABSTRACT FROM AUTHOR]

[Display omitted] • Meta-analysis reveals Arctic soil labile nitrogen (N) responses to climate change. • Warming shows no overall effect, while snow addition alters soil labile N pools. • Decision tree analysis identifies key determinants of soil labile N changes. • Climate, soil, vegetation, and experimental factors shape soil labile N responses. • Provide insights into the responses of Arctic N dynamics to climate change. The Arctic terrestrial ecosystems are undergoing rapid climate change, causing shifts in the dynamics of soil nitrogen (N), a pivotal but relatively underexplored component. To understand the impacts of climate change on soil labile N pools, we performed meta- and decision-tree analyses of 391 observations from 38 peer-reviewed publications across the Arctic, focusing on experimental warming and snow addition. Soil dissolved organic nitrogen (DON), ammonium (N H 4 +), and nitrate (N O 3 -) pools under experimental warming exhibited overall standard mean differences (SMDs) ranging from −0.08 to 0.02, with no significance (P > 0.05); however, specific conditions led to significant changes. The key determinants of soil labile N responses to warming were experimental duration and mean annual summer temperature for DON; annual precipitation, soil moisture, and sampling timing for N H 4 + ; and soil layer for N O 3 -. Snow addition significantly increased all labile N pools (overall SMD = 0.23–0.36; P < 0.05), influenced by factors such as sampling timing and vegetation type for DON; experimental duration and soil moisture for N H 4 + ; and soil pH for N O 3 -. By consolidating and reprocessing datasets, we not only showed the overall responses of soil labile N pools to climate manipulation experiments in Arctic tundra ecosystems but also identified key determinants for changes in soil N pools among environmental and experimental variables. Our findings demonstrate that warming and snow-cover changes significantly affect soil labile N pools, highlighting how the unique environmental characteristics of different sites influence terrestrial N cycling and underscoring the complexity of Arctic N dynamics under climate change. [ABSTRACT FROM AUTHOR]

Investigates the simultaneous responses in biomass, nitrogen and phosphorous incorporation of plants and microbes undergoing temperature, nutrient and light regimes in the Arctic. Increased biomass of plants as response to fertilizer intake; Increase in microbial response after shading.

Impacts of increased winter snowfall and warmer summer air temperatures on nitrous oxide (N 2 O) dynamics in arctic tundra are uncertain. Here we evaluate surface N 2 O dynamics in both wet and dry tundra in West Greenland, subjected to field manipulations with deepened winter snow and summer warming. The potential denitrification activity (PDA) and potential net N 2 O production (N 2 O net) were measured to assess denitrification and N 2 O consumption potential. The surface N 2 O fluxes averaged 0.49 ± 0.42 and 2.6 ± 0.84 μg N 2 O–N m−2 h−1, and total emissions were 212 ± 151 and 114 ± 63 g N 2 O–N scaled to the entire study area of 0.15 km2, at the dry and wet tundra, respectively. The experimental summer warming, and in combination with deepened snow, significantly increased N 2 O emissions at the dry tundra, but not at the wet tundra. The deepened snow increased winter soil temperatures and growing season soil N availability (DON, NH 4 +-N or NO 3 −-N), but no main effect of deepened snow on N 2 O fluxes was found at either tundra ecosystem. The mean PDA was 5- and 121-fold higher than the N 2 O net at the dry and wet tundra, respectively, suggesting that N 2 O might be reduced and emitted as dinitrogen (N 2). Overall, this study reveals modest but evident surface N 2 O fluxes from tundra ecosystems in Western Greenland, and suggests that projected increases in winter precipitation and summer air temperatures may increase N 2 O emissions, particularly at the dry tundra dominating in this region. • We studied climate change effects on N 2 O fluxes in dry and wet tundra West Greenland. • Generally, N 2 O fluxes were relatively modest in both types of vegetation. • Summer warming combined with deepened snow enhanced N 2 O emissions in dry tundra. • Climate change did not affect N 2 O fluxes in wet tundra. [ABSTRACT FROM AUTHOR]

Examines effects of environmental manipulations simulating climate change on soil microorganisms and nematode population in Arctic terrestrial ecosystems. Influence of high temperature and global warming on the nematode community at high altitudes; Impact of increasing heath temperature on microbial populations.

Arctic and alpine tundra ecosystems are large reservoirs of organic carbon 1,2 . Climate warming may stimulate ecosystem respiration and release carbon into the atmosphere 3,4 . The magnitude and persistency of this stimulation and the environmental mechanisms that drive its variation remain uncertain 5-7 . This hampers the accuracy of global land carbon-climate feedback projections 7,8 . Here we synthesize 136 datasets from 56 open-top chamber in situ warming experiments located at 28 arctic and alpine tundra sites which have been running for less than 1 year up to 25 years. We show that a mean rise of 1.4 °C [confidence interval (CI) 0.9-2.0 °C] in air and 0.4 °C [CI 0.2-0.7 °C] in soil temperature results in an increase in growing season ecosystem respiration by 30% [CI 22-38%] (n = 136). Our findings indicate that the stimulation of ecosystem respiration was due to increases in both plant-related and microbial respiration (n = 9) and continued for at least 25 years (n = 136). The magnitude of the warming effects on respiration was driven by variation in warming-induced changes in local soil conditions, that is, changes in total nitrogen concentration and pH and by context-dependent spatial variation in these conditions, in particular total nitrogen concentration and the carbon:nitrogen ratio. Tundra sites with stronger nitrogen limitations and sites in which warming had stimulated plant and microbial nutrient turnover seemed particularly sensitive in their respiration response to warming. The results highlight the importance of local soil conditions and warming-induced changes therein for future climatic impacts on respiration., (© 2024. The Author(s).)

Plant biomass is a fundamental ecosystem attribute that is sensitive to rapid climatic changes occurring in the Arctic. Nevertheless, measuring plant biomass in the Arctic is logistically challenging and resource intensive. Lack of accessible field data hinders efforts to understand the amount, composition, distribution, and changes in plant biomass in these northern ecosystems. Here, we present The Arctic plant aboveground biomass synthesis dataset, which includes field measurements of lichen, bryophyte, herb, shrub, and/or tree aboveground biomass (g m -2 ) on 2,327 sample plots from 636 field sites in seven countries. We created the synthesis dataset by assembling and harmonizing 32 individual datasets. Aboveground biomass was primarily quantified by harvesting sample plots during mid- to late-summer, though tree and often tall shrub biomass were quantified using surveys and allometric models. Each biomass measurement is associated with metadata including sample date, location, method, data source, and other information. This unique dataset can be leveraged to monitor, map, and model plant biomass across the rapidly warming Arctic., (© 2024. The Author(s).)

Rationale: Analysis of stable isotopes in tissue and excreta may provide information about animal diets and their nutritional state. As body condition may have a major influence on reproduction, linking stable isotope values to animal demographic rates may help unravel the drivers behind animal population dynamics., Methods: We performed sequential analysis of δ 15 N values in guard hair from 21 muskoxen (Ovibos moschatus) from Zackenberg in high arctic Greenland. We were able to reconstruct the dietary history for the population over a 5-year period with contrasting environmental conditions. We examined the linkage between guard hair δ 15 N values in 12 three-month periods and muskox calf recruitment to detect critical periods for muskox reproduction. Finally, we conducted similar analyses of the correlation between environmental conditions (snow depth and air temperature) and calf recruitment., Results: δ 15 N values exhibited a clear seasonal pattern with high levels in summer and low levels in winter. However, large inter-annual variation was found in winter values, suggesting varying levels of catabolism depending on snow conditions. In particular δ 15 N values during January-March were linked to muskox recruitment rates, with higher values coinciding with lower calf recruitment. δ 15 N values were a better predictor of muskox recruitment rates than environmental conditions., Conclusions: Although environmental conditions may ultimately determine the dietary δ 15 N signal in muskox guard hairs, muskox calf recruitment was more strongly correlated with δ 15 N values than ambient snow and temperature. The period January-March, corresponding to late gestation, appears particularly critical for muskox reproduction., (© 2023 The Authors. Rapid Communications in Mass Spectrometry published by John Wiley & Sons Ltd.)

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 PO 4 3- , 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.4 + , 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., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Warming occurs in the Arctic twice as fast as the global average, which in turn leads to a large enhancement in terpenoid emissions from vegetation. Volatile terpenoids are the main class of biogenic volatile organic compounds (VOCs) that play crucial roles in atmospheric chemistry and climate. However, the biochemical mechanisms behind the temperature-dependent increase in VOC emissions from subarctic ecosystems are largely unexplored. Using 13 CO 2 -labeling, we studied the origin of VOCs and the carbon (C) allocation under global warming in the soil-plant-atmosphere system of contrasting subarctic heath tundra vegetation communities characterized by dwarf shrubs of the genera Salix or Betula. The projected temperature rise of the subarctic summer by 5°C was realistically simulated in sophisticated climate chambers. VOC emissions strongly depended on the plant species composition of the heath tundra. Warming caused increased VOC emissions and significant changes in the pattern of volatiles toward more reactive hydrocarbons. The 13 C was incorporated to varying degrees in different monoterpene and sesquiterpene isomers. We found that de novo monoterpene biosynthesis contributed to 40%-44% (Salix) and 60%-68% (Betula) of total monoterpene emissions under the current climate, and that warming increased the contribution to 50%-58% (Salix) and 87%-95% (Betula). Analyses of above- and belowground 12/13 C showed shifts of C allocation in the plant-soil systems and negative effects of warming on C sequestration by lowering net ecosystem exchange of CO 2 and increasing C loss as VOCs. This comprehensive analysis provides the scientific basis for mechanistically understanding the processes controlling terpenoid emissions, required for modeling VOC emissions from terrestrial ecosystems and predicting the future chemistry of the arctic atmosphere. By changing the chemical composition and loads of VOCs into the atmosphere, the current data indicate that global warming in the Arctic may have implications for regional and global climate and for the delicate tundra ecosystems., (© 2020 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Snow may prevent Arctic herbivores from accessing their forage in winter, forcing them to aggregate in the few patches with limited snow. In High Arctic Greenland, Arctic hare and rock ptarmigan often forage in muskox feeding craters. We therefore hypothesized that due to limited availability of forage, the dietary niches of these resident herbivores overlap considerably, and that the overlap increases as winter progresses. To test this, we analyzed fecal samples collected in early and late winter. We used molecular analysis to identify the plant taxa consumed, and stable isotope ratios of carbon and nitrogen to quantify the dietary niche breadth and dietary overlap. The plant taxa found indicated only limited dietary differentiation between the herbivores. As expected, dietary niches exhibited a strong contraction from early to late winter, especially for rock ptarmigan. This may indicate increasing reliance on particular plant resources as winter progresses. In early winter, the diet of rock ptarmigan overlapped slightly with that of muskox and Arctic hare. Contrary to our expectations, no inter-specific dietary niche overlap was observed in late winter. This overall pattern was specifically revealed by combined analysis of molecular data and stable isotope contents. Hence, despite foraging in the same areas and generally feeding on the same plant taxa, the quantitative dietary overlap between the three herbivores was limited. This may be attributable to species-specific consumption rates of plant taxa. Yet, Arctic hare and rock ptarmigan may benefit from muskox opening up the snow pack, thereby allowing them to access the plants.

Tundra regions are projected to warm rapidly during the coming decades. The tundra biome holds the largest terrestrial carbon pool, largely contained in frozen permafrost soils. With warming, these permafrost soils may thaw and become available for microbial decomposition, potentially providing a positive feedback to global warming. Warming may directly stimulate microbial metabolism but may also indirectly stimulate organic matter turnover through increased plant productivity by soil priming from root exudates and accelerated litter turnover rates. Here, we assess the impacts of experimental warming on turnover rates of leaf litter, active layer soil and thawed permafrost sediment in two high-arctic tundra heath sites in NE-Greenland, either dominated by evergreen or deciduous shrubs. We incubated shrub leaf litter on the surface of control and warmed plots for 1 and 2 years. Active layer soil was collected from the plots to assess the effects of 8 years of field warming on soil carbon stocks. Finally, we incubated open cores filled with newly thawed permafrost soil for 2 years in the active layer of the same plots. After field incubation, we measured basal respiration rates of recovered thawed permafrost cores in the lab. Warming significantly reduced litter mass loss by 26% after 1 year incubation, but differences in litter mass loss among treatments disappeared after 2 years incubation. Warming also reduced litter nitrogen mineralization and decreased the litter carbon to nitrogen ratio. Active layer soil carbon stocks were reduced 15% by warming, while soil dissolved nitrogen was reduced by half in warmed plots. Warming had a positive legacy effect on carbon turnover rates in thawed permafrost cores, with 10% higher respiration rates measured in cores from warmed plots. These results demonstrate that warming may have contrasting effects on above- and belowground tundra carbon turnover, possibly governed by microbial resource availability., (© 2017 John Wiley & Sons Ltd.)

The subarctic environment of northernmost Sweden has changed over the past century, particularly elements of climate and cryosphere. This paper presents a unique geo-referenced record of environmental and ecosystem observations from the area since 1913. Abiotic changes have been substantial. Vegetation changes include not only increases in growth and range extension but also counterintuitive decreases, and stability: all three possible responses. Changes in species composition within the major plant communities have ranged between almost no changes to almost a 50 per cent increase in the number of species. Changes in plant species abundance also vary with particularly large increases in trees and shrubs (up to 600%). There has been an increase in abundance of aspen and large changes in other plant communities responding to wetland area increases resulting from permafrost thaw. Populations of herbivores have responded to varying management practices and climate regimes, particularly changing snow conditions. While it is difficult to generalize and scale-up the site-specific changes in ecosystems, this very site-specificity, combined with projections of change, is of immediate relevance to local stakeholders who need to adapt to new opportunities and to respond to challenges. Furthermore, the relatively small area and its unique datasets are a microcosm of the complexity of Arctic landscapes in transition that remains to be documented.

Full recovery of the ozone layer is not expected for several decades and consequently, the incoming level of solar ultraviolet-B (UV-B) will only slowly be reduced. Therefore to investigate the structural and photosynthetic responses to changes in solar UV-B we conducted a 5-year UV-B exclusion study in high arctic Greenland. During the growing season, the gas exchange (H₂O and CO₂) and chlorophyll-a fluorescence were measured in Vaccinium uliginosum. The leaf dry weight, carbon, nitrogen, stable carbon isotope ratio, chlorophyll and carotenoid content were determined from a late season harvest. The net photosynthesis per leaf area was on average 22% higher in 61% reduced UV-B treatment across the season, but per ground area photosynthesis was unchanged. The leaf level increase in photosynthesis was accompanied by increased leaf nitrogen, higher stomatal conductance and F(v)/F(m). There was no change in total leaf biomass, but reduction in total leaf area caused a pronounced reduction of specific leaf area and leaf area index in reduced UV-B. This demonstrates the structural changes to counterbalance the reduced plant carbon uptake seen per leaf area in ambient UV-B as the resulting plant carbon uptake per ground area was not affected. Thus, our understanding of long-term responses to UV-B reduction must take into account both leaf level processes as well as structural changes to understand the apparent robustness of plant carbon uptake per ground area. In this perspective, V. uliginosum seems able to adjust plant carbon uptake to the present amount of solar UV-B radiation in the High Arctic., (Copyright © Physiologia Plantarum 2011.)

Understanding the sensitivity of tundra vegetation to climate warming is critical to forecasting future biodiversity and vegetation feedbacks to climate. In situ warming experiments accelerate climate change on a small scale to forecast responses of local plant communities. Limitations of this approach include the apparent site-specificity of results and uncertainty about the power of short-term studies to anticipate longer term change. We address these issues with a synthesis of 61 experimental warming studies, of up to 20 years duration, in tundra sites worldwide. The response of plant groups to warming often differed with ambient summer temperature, soil moisture and experimental duration. Shrubs increased with warming only where ambient temperature was high, whereas graminoids increased primarily in the coldest study sites. Linear increases in effect size over time were frequently observed. There was little indication of saturating or accelerating effects, as would be predicted if negative or positive vegetation feedbacks were common. These results indicate that tundra vegetation exhibits strong regional variation in response to warming, and that in vulnerable regions, cumulative effects of long-term warming on tundra vegetation - and associated ecosystem consequences - have the potential to be much greater than we have observed to date., (© 2011 Blackwell Publishing Ltd/CNRS.)

Current atmospheric warming due to increase of greenhouse gases will have severe consequences for the structure and functioning of arctic ecosystems with changes that, in turn, may feed back on the global-scale composition of the atmosphere. During more than two decades, environmental controls on biological and biogeochemical processes and possible atmospheric feedbacks have been intensely investigated at Abisko, Sweden, by long-term ecosystem manipulations. The research has addressed questions like environmental regulation of plant and microbial community structure and biomass, carbon and nutrient pools and element cycling, including exchange of greenhouse gases and volatile organic compounds, with focus on fundamental processes in the interface between plants, soil and root-associated and free-living soil microorganisms. The ultimate goal has been to infer from these multi-decadal experiments how subarctic and arctic ecosystems will respond to likely environmental changes in the future. Here we give an overview of some of the experiments and main results.

Long-term responses of ambient solar ultraviolet (UV) radiation were investigated on Salix arctica and Vaccinium uliginosum in a High Arctic heath ecosystem in Zackenberg, northeast Greenland. Over a period of six years, UV exclusion was conducted in the growing season by means of filters: 60% UV-B reduction, 90% UV-B+UV-A reduction, UV transparent filter control, and an open control without filter. Plant responses were evaluated using specific leaf area, leaf content of UV-B absorbing compounds and PSII performance parameters derived from chlorophyll-a fluorescence induction curves. Based on the JIP-test, we calculated the total performance index PI(total), which includes the integrating antennae, the PSII reaction center, intersystem electron transport and reduction of PSI end acceptors-dependent parameters. In both species, UV exclusion significantly decreased the content of UV-B-absorbing compounds. Salix increased its specific leaf area, while Vaccinium decreased it. UV exclusion increased the PI(total) in both species during all six years of experimentation. This response was governed by a significantly decreased RC/ABS, a marginally non-significant increased ET(o)/TR(o) and a significantly increased TR(o)/ABS=F(V)/F(M) and RE(o)/ET(o). These results demonstrate the current level of ambient UV-B to decrease PSII performance significantly in these High Arctic plants. It appears that the two plant species both have improved their UV-screening capacity, but through different strategies, although this did not sufficiently prevent negative effects of the ambient UV radiation. We argue the decreased PSII performance to be part of a response decreasing plant carbon uptake. We speculate the negative effects on PSII performance mediated by ambient UV irradiance to be present in years where warming induces early snowmelt, exposing the vegetation to high spring UV-B, and to be present in the future to the degree the ozone layer is not fully recovered., (Copyright © 2010 Elsevier GmbH. All rights reserved.)

*Biogenic volatile organic compound (BVOC) emissions from arctic ecosystems are important in view of their role in global atmospheric chemistry and unknown feedbacks to global warming. These cold ecosystems are hotspots of climate warming, which will be more severe here than averaged over the globe. We assess the effects of climatic warming on non-methane BVOC emissions from a subarctic heath. *We performed ecosystem-based chamber measurements and gas chromatography-mass spectrometry (GC-MS) analyses of the BVOCs collected on adsorbent over two growing seasons at a wet subarctic tundra heath hosting a long-term warming and mountain birch (Betula pubescens ssp. czerepanovii) litter addition experiment. *The relatively low emissions of monoterpenes and sesquiterpenes were doubled in response to an air temperature increment of only 1.9-2.5 degrees C, while litter addition had a minor influence. BVOC emissions were seasonal, and warming combined with litter addition triggered emissions of specific compounds. *The unexpectedly high rate of release of BVOCs measured in this conservative warming scenario is far above the estimates produced by the current models, which underlines the importance of a focus on BVOC emissions during climate change. The observed changes have implications for ecological interactions and feedback effects on climate change via impacts on aerosol formation and indirect greenhouse effects.

Emissions of isoprene, a reactive hydrocarbon, from Subarctic vegetation are not well documented. However, the Arctic is likely to experience the most pronounced effects of climatic warming, which may increase temperature-dependent isoprene emission. Here, we assessed isoprene emission from a Subarctic heath subjected to a 3-4 degrees C increase in air temperature and mountain birch (Betula pubescens ssp. czerepanovii) litter addition for 7-8 yr, simulating climatic warming and the subsequent expansion of deciduous shrub species and migration of the treeline. The measurements were performed using the dynamic chamber method on a wet heath with a mixture of shrubs, herbs and graminoids. Isoprene emissions averaged across the treatments were 36 +/- 5 microg m(-2) h(-1) in 2006 and 58 +/- 7 microg m(-2) h(-1) in 2007. The experimental warming increased the emissions by 83% in 2007 (P = 0.021) and by 56% in 2006 (P = 0.056), while litter addition had no significant effects. The net ecosystem CO(2) exchange was significantly decreased by warming in 2007. These results show that isoprene emissions from Subarctic heaths are comparable to those from Subarctic peatlands. Climatic warming will increase the emissions, and the amount of carbon lost as isoprene, from Subarctic heath ecosystems.

Environmental changes are likely to alter the chemical composition of plant tissues, including content and concentrations of secondary compounds, and thereby affect the food sources of herbivores. After 10 years of experimental increase of temperature, nutrient levels and light attenuation in a sub-arctic, alpine ecosystem, we investigated the effects on carbon based secondary compounds (CBSC) and nitrogen in one dominant deciduous dwarf shrub, Salix herbacea x polaris and two dominant evergreen dwarf shrubs, Cassiope tetragona and Vaccinium vitis-idaea throughout one growing season. The main aims were to compare the seasonal course and treatment effects on CBSC among the species, life forms and leaf cohorts and to examine whether the responses in different CBSC were consistent across compounds. The changes in leaf chemistry both during the season and in response to the treatments were higher in S. herbacea x polaris than in the corresponding current year's leaf cohort of the evergreen C. tetragona. The changes were also much higher than in the 1-year-old leaves of the two evergreens probably due to differences in dilution and turnover of CBSC in growing and mature leaves paired with different rates of allocation. Most low molecular weight phenolics in the current year's leaves decreased in all treatments. Condensed tannins and the tannin-to-N ratio, however, either increased or decreased, and the strength and even direction of the responses varied among the species and leaf cohorts, supporting views of influential factors additional to resource-based or developmental controls, as e.g. species specific or genetic controls of CBSC. The results indicate that there is no common response to environmental changes across species and substances. However, the pronounced treatment responses imply that the quality of the herbivore forage is likely to be strongly affected in a changing arctic environment, although both the direction and strength of the responses will be different among plant species, tissue types and substances.