The fate of carbon in a mature forest under carbon dioxide enrichment.

Atmospheric carbon dioxide enrichment (eCO2) can enhance plant carbon uptake and growth1-5, thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO2 concentration6. Although evidence gathered from young aggrading forests has generally indicated a strong CO2 fertilization effect on biomass growth3-5, it is unclear whether mature forests respond to eCO2 in a similar way. In mature trees and forest stands7-10, photosynthetic uptake has been found to increase under eCO2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO2 unclear4,5,7-11. Here using data from the first ecosystem-scale Free-Air CO2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responded to four years of eCO2 exposure. We show that, although the eCO2 treatment of +150 parts per million (+38 per cent) above ambient levels induced a 12 per cent (+247 grams of carbon per square metre per year) increase in carbon uptake through gross primary production, this additional carbon uptake did not lead to increased carbon sequestration at the ecosystem level. Instead, the majority of the extra carbon was emitted back into the atmosphere via several respiratory fluxes, with increased soil respiration alone accounting for half of the total uptake surplus. Our results call into question the predominant thinking that the capacity of forests to act as carbon sinks will be generally enhanced under eCO2, and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO2 fertilization as a driver of increased carbon sinks in global forests.

Strong isoprene emission response to temperature in tundra vegetation.

Emissions of biogenic volatile organic compounds (BVOCs) are a crucial component of biosphere-atmosphere interactions. In northern latitudes, climate change is amplified by feedback processes in which BVOCs have a recognized, yet poorly quantified role, mainly due to a lack of measurements and concomitant modeling gaps. Hence, current Earth system models mostly rely on temperature responses measured on vegetation from lower latitudes, rendering their predictions highly uncertain. Here, we show how tundra isoprene emissions respond vigorously to temperature increases, compared to model results. Our unique dataset of direct eddy covariance ecosystem-level isoprene measurements in two contrasting ecosystems exhibited Q10 (the factor by which the emission rate increases with a 10 °C rise in temperature) temperature coefficients of up to 20.8, that is, 3.5 times the Q10 of 5.9 derived from the equivalent model calculations. Crude estimates using the observed temperature responses indicate that tundra vegetation could enhance their isoprene emissions by up to 41% (87%)-that is, 46% (55%) more than estimated by models-with a 2 °C (4 °C) warming. Our results demonstrate that tundra vegetation possesses the potential to substantially boost its isoprene emissions in response to future rising temperatures, at rates that exceed the current Earth system model predictions.

Separating direct and indirect effects of rising temperatures on biogenic volatile emissions in the Arctic.

Volatile organic compounds (VOCs) are released from biogenic sources in a temperature-dependent manner. Consequently, Arctic ecosystems are expected to greatly increase their VOC emissions with ongoing climate warming, which is proceeding at twice the rate of global temperature rise. Here, we show that ongoing warming has strong, increasing effects on Arctic VOC emissions. Using a combination of statistical modeling on data from several warming experiments in the Arctic tundra and dynamic ecosystem modeling, we separate the impacts of temperature and soil moisture into direct effects and indirect effects through vegetation composition and biomass alterations. The indirect effects of warming on VOC emissions were significant but smaller than the direct effects, during the 14-y model simulation period. Furthermore, vegetation changes also cause shifts in the chemical speciation of emissions. Both direct and indirect effects result in large geographic differences in VOC emission responses in the warming Arctic, depending on the local vegetation cover and the climate dynamics. Our results outline complex links between local climate, vegetation, and ecosystem-atmosphere interactions, with likely local-to-regional impacts on the atmospheric composition.

Influence of increased nutrient availability on biogenic volatile organic compound (BVOC) emissions and leaf anatomy of subarctic dwarf shrubs under climate warming and increased cloudiness.

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.

The multidimensional nutritional niche of fungus-cultivar provisioning in free-ranging colonies of a neotropical leafcutter ant.

Foraging trails of leafcutter colonies are iconic scenes in the Neotropics, with ants collecting freshly cut plant fragments to provision a fungal food crop. We hypothesised that the fungus-cultivar's requirements for macronutrients and minerals govern the foraging niche breadth of Atta colombica leafcutter ants. Analyses of plant fragments carried by foragers showed how nutrients from fruits, flowers and leaves combine to maximise cultivar performance. While the most commonly foraged leaves delivered excess protein relative to the cultivar's needs, in vitro experiments showed that the minerals P, Al and Fe may expand the leafcutter foraging niche by enhancing the cultivar's tolerance to protein-biased substrates. A suite of other minerals reduces cultivar performance in ways that may render plant fragments with optimal macronutrient blends unsuitable for provisioning. Our approach highlights how the nutritional challenges of provisioning a mutualist can govern the multidimensional realised niche available to a generalist insect herbivore.

Synergistic effects of insect herbivory and changing climate on plant volatile emissions in the subarctic tundra.

Climate change increases the insect abundance, especially in Arctic ecosystems. Insect herbivory also significantly increases plant emissions of volatile organic compounds (VOCs), which are highly reactive in the atmosphere and play a crucial role in atmospheric chemistry and physics. However, it is unclear how the effects of insect herbivory on VOC emissions interact with climatic changes, such as warming and increased cloudiness. We assessed how experimental manipulations of temperature and light availability in subarctic tundra, that had been maintained for 30 years at the time of the measurements, affect the VOC emissions from a widespread dwarf birch (Betula nana) when subjected to herbivory by local geometrid moth larvae, the autumnal moth (Epirrita autumnata) and the winter moth (Operophtera brumata). Warming and insect herbivory on B. nana stimulated VOC emission rates and altered the VOC blend. The herbivory-induced increase in sesquiterpene and homoterpene emissions were climate-treatment-dependent. Many herbivory-associated VOCs were more strongly induced in the shading treatment than in other treatments. We showed generally enhanced tundra VOC emissions upon insect herbivory and synergistic effects on the emissions of some VOC groups in a changing climate, which can have positive feedbacks on cloud formation. Furthermore, the acclimation of plants to long-term climate treatments affects VOC emissions and strongly interacts with plant responses to herbivory. Such acclimation complicates predictions of how climate change, together with interacting biotic stresses, affects VOC emissions in the high latitudes.

Phenological stage of tundra vegetation controls bidirectional exchange of BVOCs in a climate change experiment on a subarctic heath.

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.

Volatile organic compound emission in tundra shrubs - Dependence on species characteristics and the near-surface environment.

Temperature is one of the key abiotic factors during the life of plants, especially in the Arctic region which is currently experiencing rapid climate change. We evaluated plant traits and environmental variables determining leaf temperature in tundra shrubs and volatile organic compound (VOC) emissions with field measurements on deciduous tundra shrubs, Salix myrsinites and Betula nana, and evergreen Cassiope tetragona and Rhododendron lapponicum. Higher leaf-to-air temperature difference was observed in evergreen, compared to deciduous shrubs. Evergreen shrubs also showed continuously increasing photosynthesis with increasing temperature, suggesting high thermal tolerance. For the deciduous species, the optimum temperature for net photosynthesis was between our measurement temperatures of 24 °C and 38 °C. Air temperature and vapor pressure deficit were the most important variables influencing leaf temperature and VOC emissions in all the studied plants, along with stomatal density and specific leaf area in the deciduous shrubs. Using climate data and emission factors from our measurements, we modelled total seasonal tundra shrub VOC emissions of 0.3-2.3 g m-2 over the main growing season. Our results showed higher-than-expected temperature optima for photosynthesis and VOC emission and demonstrated the relative importance of plant traits and local environments in determining leaf temperature and VOC emissions in a subarctic tundra.

Origin of volatile organic compound emissions from subarctic tundra under global warming.

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 CO2 -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 CO2 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.

Long-term effects of elevated CO2, nighttime warming and drought on plant secondary metabolites in a temperate heath ecosystem.

BACKGROUND AND AIMS: Plant secondary metabolites play critical roles in plant stress tolerance and adaptation, and are known to be influenced by the environment and climate changes, yet the impacts and interactions of multiple climate change components are poorly understood, particularly under natural conditions. METHODS: Accumulation of phenolics and emissions of volatile organic compounds (VOCs) were assessed on heather, Calluna vulgaris, an abundant evergreen dwarf shrub in European heathlands, after 6 years of exposure to elevated CO2, summer drought and nighttime warming. KEY RESULTS: Drought alone had the strongest effects on phenolic concentrations and compositions, with moderate effects of elevated CO2 and temperature. Elevated CO2 exerted the greatest impact on VOC emissions, mainly by increasing monoterpene emissions. The response magnitudes varied among plant tissue types and chemical constituents, and across time. With respect to interactive effects of the studied climate change components, the interaction between drought and elevated CO2 was most apparent. Drought mainly reduced phenolic accumulation and VOC emissions, while elevated CO2 mitigated such effects. CONCLUSIONS: In natural ecosystems, co-occurring climate factors can exert complex impacts on plant secondary metabolite profiles, which may in turn alter ecosystem processes.

Volatile emissions from thawing permafrost soils are influenced by meltwater drainage conditions.

Vast amounts of carbon are bound in both active layer and permafrost soils in the Arctic. As a consequence of climate warming, the depth of the active layer is increasing in size and permafrost soils are thawing. We hypothesize that pulses of biogenic volatile organic compounds are released from the near-surface active layer during spring, and during late summer season from thawing permafrost, while the subsequent biogeochemical processes occurring in thawed soils also lead to emissions. Biogenic volatile organic compounds are reactive gases that have both negative and positive climate forcing impacts when introduced to the Arctic atmosphere, and the knowledge of their emission magnitude and pattern is necessary to construct reliable climate models. However, it is unclear how different ecosystems and environmental factors such as drainage conditions upon permafrost thaw affect the emission and compound composition. Here we show that incubations of frozen B horizon of the active layer and permafrost soils collected from a High Arctic heath and fen release a range of biogenic volatile organic compounds upon thaw and during subsequent incubation experiments at temperatures of 10°C and 20°C. Meltwater drainage in the fen soils increased emission rates nine times, while having no effect in the drier heath soils. Emissions generally increased with temperature, and emission profiles for the fen soils were dominated by benzenoids and alkanes, while benzenoids, ketones, and alcohols dominated in heath soils. Our results emphasize that future changes affecting the drainage conditions of the Arctic tundra will have a large influence on volatile emissions from thawing permafrost soils - particularly in wetland/fen areas.

Contrasting above- and belowground organic matter decomposition and carbon and nitrogen dynamics in response to warming in High Arctic tundra.

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.

Leaf anatomy, BVOC emission and CO2 exchange of arctic plants following snow addition and summer warming.

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.

Climate change alters leaf anatomy, but has no effects on volatile emissions from Arctic plants.

Biogenic volatile organic compound (BVOC) emissions are expected to change substantially because of the rapid advancement of climate change in the Arctic. BVOC emission changes can feed back both positively and negatively on climate warming. We investigated the effects of elevated temperature and shading on BVOC emissions from arctic plant species Empetrum hermaphroditum, Cassiope tetragona, Betula nana and Salix arctica. Measurements were performed in situ in long-term field experiments in subarctic and high Arctic using a dynamic enclosure system and collection of BVOCs into adsorbent cartridges analysed by gas chromatography-mass spectrometry. In order to assess whether the treatments had resulted in anatomical adaptations, we additionally examined leaf anatomy using light microscopy and scanning electron microscopy. Against expectations based on the known temperature and light-dependency of BVOC emissions, the emissions were barely affected by the treatments. In contrast, leaf anatomy of the studied plants was significantly altered in response to the treatments, and these responses appear to differ from species found at lower latitudes. We suggest that leaf anatomical acclimation may partially explain the lacking treatment effects on BVOC emissions at plant shoot-level. However, more studies are needed to unravel why BVOC emission responses in arctic plants differ from temperate species.

Climate change-induced vegetation change as a driver of increased subarctic biogenic volatile organic compound emissions.

Emissions of biogenic volatile organic compounds (BVOCs) have been earlier shown to be highly temperature sensitive in subarctic ecosystems. As these ecosystems experience rapidly advancing pronounced climate warming, we aimed to investigate how warming affects the BVOC emissions in the long term (up to 13 treatment years). We also aimed to assess whether the increased litterfall resulting from the vegetation changes in the warming subarctic would affect the emissions. The study was conducted in a field experiment with factorial open-top chamber warming and annual litter addition treatments on subarctic heath in Abisko, northern Sweden. After 11 and 13 treatment years, BVOCs were sampled from plant communities in the experimental plots using a push-pull enclosure technique and collection into adsorbent cartridges during the growing season and analyzed with gas chromatography-mass spectrometry. Plant species coverage in the plots was analyzed by the point intercept method. Warming by 2 °C caused a 2-fold increase in monoterpene and 5-fold increase in sesquiterpene emissions, averaged over all measurements. When the momentary effect of temperature was diminished by standardization of emissions to a fixed temperature, warming still had a significant effect suggesting that emissions were also indirectly increased. This indirect increase appeared to result from increased plant coverage and changes in vegetation composition. The litter addition treatment also caused significant increases in the emission rates of some BVOC groups, especially when combined with warming. The combined treatment had both the largest vegetation changes and the highest BVOC emissions. The increased emissions under litter addition were probably a result of a changed vegetation composition due to alleviated nutrient limitation and stimulated microbial production of BVOCs. We suggest that the changes in the subarctic vegetation composition induced by climate warming will be the major factor indirectly affecting the BVOC emission potentials and composition.

Plant volatiles in extreme terrestrial and marine environments.

This review summarizes the current understanding on plant and algal volatile organic compound (VOC) production and emission in extreme environments, where temperature, water availability, salinity or other environmental factors pose stress on vegetation. Here, the extreme environments include terrestrial systems, such as arctic tundra, deserts, CO2 springs and wetlands, and marine systems such as sea ice, tidal rock pools and hypersaline environments, with mangroves and salt marshes at the land-sea interface. The emission potentials at fixed temperature and light level or actual emission rates for phototrophs in extreme environments are frequently higher than for organisms from less stressful environments. For example, plants from the arctic tundra appear to have higher emission potentials for isoprenoids than temperate species, and hypersaline marine habitats contribute to global dimethyl sulphide (DMS) emissions in significant amounts. DMS emissions are more widespread than previously considered, for example, in salt marshes and some desert plants. The reason for widespread VOC, especially isoprenoid, emissions from different extreme environments deserves further attention, as these compounds may have important roles in stress resistance and adaptation to extremes. Climate warming is likely to significantly increase VOC emissions from extreme environments both by direct effects on VOC production and volatility, and indirectly by altering the composition of the vegetation.

Acclimation of subarctic vegetation to warming and increased cloudiness.

Subarctic ecosystems are exposed to elevated temperatures and increased cloudiness in a changing climate with potentially important effects on vegetation structure, composition, and ecosystem functioning. We investigated the individual and combined effects of warming and increased cloudiness on vegetation greenness and cover in mesocosms from two tundra and one palsa mire ecosystems kept under strict environmental control in climate chambers. We also investigated leaf anatomical and biochemical traits of four dominant vascular plant species (Empetrum hermaphroditum, Vaccinium myrtillus, Vaccinium vitis-idaea, and Rubus chamaemorus). Vegetation greenness increased in response to warming in all sites and in response to increased cloudiness in the tundra sites but without associated increases in vegetation cover or biomass, except that E. hermaphroditum biomass increased under warming. The combined warming and increased cloudiness treatment had an additive effect on vegetation greenness in all sites. It also increased the cover of graminoids and forbs in one of the tundra sites. Warming increased leaf dry mass per area of V. myrtillus and R. chamaemorus, and glandular trichome density of V. myrtillus and decreased spongy intercellular space of E. hermaphroditum and V. vitis-idaea. Increased cloudiness decreased leaf dry mass per area of V. myrtillus, palisade thickness of E. hermaphroditum, and stomata density of E. hermaphroditum and V. vitis-idaea, and increased leaf area and epidermis thickness of V. myrtillus, leaf shape index and nitrogen of E. hermaphroditum, and palisade intercellular space of V. vitis-idaea. The combined treatment caused thinner leaves and decreased leaf carbon for V. myrtillus, and increased leaf chlorophyll of E. hermaphroditum. We show that under future warmer increased cloudiness conditions in the Subarctic (as simulated in our experiment), vegetation composition and distribution will change, mostly dominated by graminoids and forbs. These changes will depend on the responses of leaf anatomical and biochemical traits and will likely impact carbon gain and primary productivity and abiotic and biotic stress tolerance.

Close coupling of plant functional types with soil microbial community composition drives soil carbon and nutrient cycling in tundra heath.

Aims: This study aimed at elucidating divergent effects of two dominant plant functional types (PFTs) in tundra heath, dwarf shrubs and mosses, on soil microbial processes and soil carbon (C) and nutrient availability, and thereby to enhance our understanding of the complex interactions between PFTs, soil microbes and soil functioning. Methods: Samples of organic soil were collected under three dwarf shrub species (of distinct mycorrhizal association and life form) and three moss species in early and late growing season. We analysed soil C and nutrient pools, extracellular enzyme activities and phospholipid fatty acid profiles, together with a range of plant traits, soil and abiotic site characteristics. Results: Shrub soils were characterised by high microbial biomass C and phosphorus and phosphatase activity, which was linked with a fungal-dominated microbial community, while moss soils were characterised by high soil nitrogen availability, peptidase and peroxidase activity associated with a bacterial-dominated microbial community. The variation in soil microbial community structure was explained by mycorrhizal association, root morphology, litter and soil organic matter quality and soil pH-value. Furthermore, we found that the seasonal variation in microbial biomass and enzyme activities over the growing season, likely driven by plant belowground C allocation, was most pronounced under the tallest shrub Betula nana. Conclusion: Our study demonstrates a close coupling of PFTs with soil microbial communities, microbial decomposition processes and soil nutrient availability in tundra heath, which suggests potential strong impacts of global change-induced shifts in plant community composition on carbon and nutrient cycling in high-latitude ecosystems. Supplementary Information: The online version contains supplementary material available at 10.1007/s11104-023-05993-w.

Volatile responses of dwarf birch to mimicked insect herbivory and experimental warming at two elevations in Greenlandic tundra.

Plants release a complex blend of volatile organic compounds (VOCs) in response to stressors. VOC emissions vary between contrasting environments and increase with insect herbivory and rising temperatures. However, the joint effects of herbivory and warming on plant VOC emissions are understudied, particularly in high latitudes, which are warming fast and facing increasing herbivore pressure. We assessed the individual and combined effects of chemically mimicked insect herbivory, warming, and elevation on dwarf birch (Betula glandulosa) VOC emissions in high-latitude tundra ecosystems in Narsarsuaq, South Greenland. We hypothesized that VOC emissions and compositions would respond synergistically to warming and herbivory, with the magnitude differing between elevations. Warming increased emissions of green leaf volatiles (GLVs) and isoprene. Herbivory increased the homoterpene, (E)-4,8-dimethyl-1,3,7-nonatriene, emissions, and the response was stronger at high elevation. Warming and herbivory had synergistic effects on GLV emissions. Dwarf birch emitted VOCs at similar rates at both elevations, but the VOC blends differed between elevations. Several herbivory-associated VOC groups did not respond to herbivory. Harsher abiotic conditions at high elevations might not limit VOC emissions from dwarf birch, and high-elevation plants might be better at herbivory defense than assumed. The complexity of VOC responses to experimental warming, elevation, and herbivory are challenging our understanding and predictions of future VOC emissions from dwarf birch-dominated ecosystems.

Two decades of experimental manipulations of heaths and forest understory in the subarctic.

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.