• Effects of warming on root morphology, root mass distribution and microbialactivity were studied in organic and mineral soil layers in two alpine ecosystems over > 10 yr, using open-top chambers, in Swedish Lapland.

• Root mass was estimated using soil cores. Washed roots were scanned and sortedinto four diameter classes, for which variables including root mass (g dry matter(g DM) m –2 ), root length density (RLD; cm cm –3 soil), specific root length (SRL; m gDM –1 ), specific root area (SRA; m 2 kg DM –1 ), and number of root tips m –2 weredetermined. Nitrification (NEA) and denitrification enzyme activity (DEA) in the top10 cm of soil were measured.

• Soil warming shifted the rooting zone towards the upper soil organic layer in bothplant communities. In the dry heath, warming increased SRL and SRA of the finestroots in both soil layers, whereas the dry meadow was unaffected. Neither NEA norDEA exhibited differences attributable to warming.

• Tundra plants may respond to climate change by altering their root morphologyand mass while microbial activity may be unaffected. This suggests that carbon maybe incorporated in tundra soils partly as a result of increases in the mass of the finerroots if temperatures rise.

Summary Climate-warming-induced permafrost thaw exposes large amounts of carbon and nitrogen in soil at considerable depths, below the seasonally thawing active layer. The extent to which plant roots can reach and interact with these hitherto detached, deep carbon and nitrogen stores remains unknown. We aimed to quantify how permafrost thaw affects root dynamics across soil depths and plant functional types compared with above-ground abundance, and potential consequences for plant?soil interactions. A decade of experimental permafrost thaw strongly increased total root length and growth in the active layer, and deep roots invaded the newly thawed permafrost underneath. Root litter input to soil across all depths was 10 times greater with permafrost thaw. Root growth timing was unaffected by experimental permafrost thaw but peaked later in deeper soil, reflecting the seasonally receding thaw front. Deep-rooting species could sequester 15N added at the base of the ambient active layer in October, which was after root growth had ceased. Deep soil organic matter that has long been locked up in permafrost is thus no longer detached from plant processes upon thaw. Whether via nutrient uptake, carbon storage, or rhizosphere priming, plant root interactions with thawing permafrost soils may feed back on our climate both positively and negatively.

There is compelling evidence from experiments and observations that climate warming prolongs the growing season in arctic regions. Until now, the start, peak, and end of the growing season, which are used to model influences of vegetation on biogeochemical cycles, were commonly quantified using above-ground phenological data. Yet, over 80% of the plant biomass in arctic regions can be below ground, and the timing of root growth affects biogeochemical processes by influencing plant water and nutrient uptake, soil carbon input and microbial activity. We measured timing of above- and below-ground production in three plant communities along an arctic elevation gradient over two growing seasons. Below-ground production peaked later in the season and was more temporally uniform than above-ground production. Most importantly, the growing season continued c. 50% longer below than above ground. Our results strongly suggest that traditional above-ground estimates of phenology in arctic regions, including remotely sensed information, are not as complete a representation of whole-plant production intensity or duration, as studies that include root phenology. We therefore argue for explicit consideration of root phenology in studies of carbon and nutrient cycling, in terrestrial biosphere models, and scenarios of how arctic ecosystems will respond to climate warming.

* In northern forests, belowground sequestration of nitrogen (N) in complex organic pools restricts nutrient availability to plants. Oxidative extracellular enzymes produced by ectomycorrhizal fungi may aid plant N acquisition by providing access to N in macromolecular complexes. We test the hypotheses that ectomycorrhizal Cortinarius species produce Mn-dependent peroxidases, and that the activity of these enzymes declines at elevated concentrations of inorganic N. * In a boreal pine forest and a sub-arctic birch forest, Cortinarius DNA was assessed by 454-sequencing of ITS amplicons and related to Mn-peroxidase activity in humus samples with- and without previous N amendment. Transcription of Cortinarius Mn-peroxidase genes was investigated in field samples. Phylogenetic analyses of Cortinarius peroxidase amplicons and genome sequences were performed. * We found a significant co-localization of high peroxidase activity and DNA from Cortinarius species. Peroxidase activity was reduced by high ammonium concentrations. Amplification of mRNA sequences indicated transcription of Cortinarius Mn-peroxidase genes under field conditions. The Cortinarius glaucopus genome encodes 11 peroxidases – a number comparable to many white-rot wood decomposers. * These results support the hypothesis that some ectomycorrhizal fungi – Cortinarius species in particular – may play an important role in decomposition of complex organic matter, linked to their mobilization of organically bound N.

Shrub abundance is expected to increase with enhanced temperature and nutrient availability in the Arctic, and associated changes in abundance of ectomycorrhizal (EM) fungi could be a key link between plant responses and longer-term changes in soil organic matter storage. This study quantifies the response in EM fungal abundance to long-term warming and fertilization in two arctic ecosystems with contrasting responses of the EM shrub Betula nana. Ergosterol was used as a biomarker for living fungal biomass in roots and organic soil and ingrowth bags were used to estimate EM mycelial production. We measured N-15 and C-13 natural abundance to identify the EM-saprotrophic divide in fungal sporocarps and to validate the EM origin of mycelia in the ingrowth bags. Fungal biomass in soil and EM mycelial production increased with fertilization at both tundra sites, and with warming at one site. This was caused partly by increased dominance of EM plants and partly by stimulation of EM mycelial growth. We conclude that cycling of carbon and nitrogen through EM fungi will increase when strongly nutrient-limited arctic ecosystems are exposed to a warmer and more nutrient-rich environment. This has potential consequences for below-ground litter quality and quantity, and for accumulation of organic matter in arctic soils.

Shrub expansion in alpine and arctic areas is a process with possibly profound implications for ecosystem functioning. The recent shrub expansion has been mainly documented by remote sensing techniques, but the drivers for this process largely remain hypotheses.

Here, we outline a dendrochronological method, adapted to shrubs, to address these hypotheses and then present a mechanism for the current shrub expansion by linking recent climate change to shrub growth performance in northern Sweden.

A pronounced increase in radial and vertical growth during recent decades along an elevational gradient from treeline to shrubline indicates an ongoing shrub expansion. Age distribution of the shrub population indicates the new colonization of shrubs at high elevations.

Shrub growth is correlated with warm summers and winter snow cover and suggests the potential for large‐scale ecosystem changes if climate change continues as projected.

* Little of our knowledge about invasibility comes from arctic and alpine ecosystems, despite increasing plant migration and invasion in those regions. Here, we examine how community type, altitude, and small-scale disturbances affect invasibility in a subarctic ecosystem. * Over a period of 4 yr, we studied seedling emergence and establishment in 17 species sown in gaps or undisturbed vegetation in four subarctic community types (Salix scrub, meadow, rich heath, poor heath) along an elevation gradient. * Invasibility was lowest in rich heath and highest in Salix scrub. Small disturbances significantly increased the invasibility in most communities, thereby showing the importance of biotic resistance to invasion in subarctic regions. Unexpectedly, invasibility did not decrease with increasing elevation, and it was also not related to summer temperature. * Our data suggest that biotic resistance might be more important than abiotic stress for invasibility in subarctic tundra and that low temperatures do not necessarily limit seedling establishment at high altitudes. High elevations are therefore potentially more vulnerable to invasion than was originally thought. Changes in community composition as a result of species migration or invasion are most likely to occur in Salix scrub and meadow, whereas Empetrum-dominated rich heath will largely remain unchanged.

Targeted removal experiments are a powerful tool to assess the effects of plant species or (functional) groups on ecosystem functions. However, removing plant biomass in itself can bias the observed responses. This bias is commonly addressed by waiting until ecosystem recovery, but this is inherently based on unverified proxies or anecdotal evidence. Statistical control methods are efficient, but restricted in scope by underlying assumptions.

We propose accounting for such biases within the experimental design, using a gradient of biomass removal controls. We demonstrate the relevance of this design by presenting (1) conceptual examples of suspected biases and (2) how to observe and control for these biases.

Using data from a mycorrhizal association-based removal experiment, we show that ignoring biomass removal biases (including by assuming ecosystem recovery) can lead to incorrect, or even contrary conclusions (e.g. false positive and false negative). Our gradient design can prevent such incorrect interpretations, regardless of whether aboveground biomass has fully recovered.

Our approach provides more objective and quantitative insights, independently assessed for each variable, than using a proxy to assume ecosystem recovery. Our approach circumvents the strict statistical assumptions of, for example, ANCOVA and thus offers greater flexibility in data analysis.

Here, we investigated effects of increased atmospheric CO(2) concentration, increased temperatures, and both factors in combination on ericoid mycorrhizal colonization, mycorrhizal functioning and below-ground carbon allocation in a subarctic forest understorey, to evaluate the hypothesis that photosynthesis is a primary driver for mycorrhizal colonization. Treatment effects on ecosystem processes were investigated using (14)C-pulse labelling and photosynthesis measurements in combination with analysis of ergosterol content in roots. The effects on delta(15)N in leaves were also studied. Ergosterol content in hair roots was positively correlated with ecosystem photosynthesis and was higher in heat- and CO(2)-treated plots. Leaves from CO(2) Plots tended to be more depleted in (15)N compared with controls both for Vaccinium myrtillus and V. vitis-idaea. Our results suggest that changes in ecosystem photosynthesis, plant carbon (C) Allocation may give rise to changing mycorrhizal colonization under elevated CO(2) and temperature. The role of mycorrhizas in ecosystem N-cycling may change on a long-term basis as inorganic N availability declines with increasing levels of atmospheric CO(2).

In arctic ecosystems, climate change has increased plant productivity. As arctic carbon (C) stocks predominantly are located belowground, the effects of greater plant productivity on soil C storage will significantly determine the net sink/source potential of these ecosystems, but vegetation controls on soil CO2 efflux remain poorly resolved.

In order to identify the role of canopy‐forming species in belowground C dynamics, we conducted a girdling experiment with plots distributed across 1 km2 of treeline birch (Betula pubescens ) forest and willow (Salix lapponum ) patches in northern Sweden and quantified the contribution of canopy vegetation to soil CO2 fluxes and belowground productivity.

Girdling birches reduced total soil CO2 efflux in the peak growing season by 53%, which is double the expected amount, given that trees contribute only half of the total leaf area in the forest. Root and mycorrhizal mycelial production also decreased substantially. At peak season, willow shrubs contributed 38% to soil CO2 efflux in their patches.

Our findings indicate that C, recently fixed by trees and tall shrubs, makes a substantial contribution to soil respiration. It is critically important that these processes are taken into consideration in the context of a greening arctic because productivity and ecosystem C sequestration are not synonymous.

Rising atmospheric CO2 and temperatures are probably altering ecosystem carbon cycling, causing both positive and negative feedbacks to climate. Below-ground processes play a key role in the global carbon (C) cycle because they regulate storage of large quantities of C, and are potentially very sensitive to direct and indirect effects of elevated CO2 and temperature. Soil organic matter pools, roots and associated rhizosphere organisms all have distinct responses to environmental change drivers, although availability of C substrates will regulate all the responses. Elevated CO2 increases C supply below-ground, whereas warming is likely to increase respiration and decomposition rates, leading to speculation that these effects will moderate one another. However, indirect effects on soil moisture availability and nutrient supply may alter processes in unexpected directions. Detailed, mechanistic understanding and modelling of below-ground flux components, pool sizes and turnover rates is needed to adequately predict long-term, net C storage in ecosystems. In this synthesis, we discuss the current status of below-ground responses to elevated CO2 and temperature and potential feedback effects, methodological challenges, and approaches to integrating models and measurements.

* Biological nitrogen fixation (BNF) performed by moss-associated cyanobacteria is one of the main sources of new nitrogen (N) input in pristine, high-latitude ecosystems. Yet, the nutrients that limit BNF remain elusive. Here, we tested whether this important ecosystem function is limited by the availability of molybdenum (Mo), phosphorus (P), or both. * BNF in dominant mosses was measured with the acetylene reduction assay (ARA) at different time intervals following Mo and P additions, in both laboratory microcosms with mosses from a boreal spruce forest and field plots in subarctic tundra. We further used a 15N2 tracer technique to assess the ARA to N2 fixation conversion ratios at our subarctic site. * BNF was up to four-fold higher shortly after the addition of Mo, in both the laboratory and field experiments. A similar positive response to Mo was found in moss colonizing cyanobacterial biomass. As the growing season progressed, nitrogenase activity became progressively more P limited. The ARA : 15N2 ratios increased with increasing Mo additions. * These findings show that N2 fixation activity as well as cyanobacterial biomass in dominant feather mosses from boreal forests and subarctic tundra are limited by Mo availability.

• Interactions between vascular plants and bryophytes determine plant community composition in many ecosystems. Yet, little is known about the importance ofinterspecific differences between bryophytes with respect to their effects on vascular plants. We compared the extent to which species-specific bryophyte effects onvascular plant generative recruitment depend on the following underlying mechanisms: allelopathy, mechanical obstruction, soil moisture and temperature control.

• We sowed 10 vascular plant species into monospecific mats of six chemicallyand structurally diverse bryophytes, and examined 1-yr seedling recruitment.Allelopathic effects were also assessed in a laboratory phyto-assay.

• Although all bryophytes suppressed vascular plant regeneration, there weresignificant differences between the bryophyte species. The lack of interactionsindicated the absence of species-specific adaptations of vascular plants for recruitment in bryophyte mats. Differences between bryophyte species were bestexplained by alterations in temperature regime under bryophyte mats, mostly byreduced temperature amplitudes during germination. The temperature regimeunder bryophyte mats was well predicted by species-specific bryophyte cushionthickness. The fitness of established seedlings was not affected by the presence ofbryophytes.

• Our results suggest that climatically or anthropogenically driven changes in thespecies’ composition of bryophyte communities have knock-on effects on vascularplant populations via generative reproduction.

Fine roots, and their functional traits, influence associated rhizosphere microorganisms via root exudation and root litter quality. However, little information is known about their relationship with rhizosphere microbial taxa and functional guilds. We investigated the relationships of 11 fine root traits of 20 sub-arctic tundra meadow plant species and soil microbial community composition, using phospholipid fatty acids (PLFAs) and high-throughput sequencing. We primarily focused on the root economics spectrum, as it provides a useful framework to examine plant strategies by integrating the co-ordination of belowground root traits along a resource acquisition?conservation trade-off axis. We found that the chemical axis of the fine root economics spectrum was positively related to fungal to bacterial ratios, but negatively to Gram-positive to Gram-negative bacterial ratios. However, this spectrum was unrelated to the relative abundance of functional guilds of soil fungi. Nevertheless, the relative abundance of arbuscular mycorrhizal fungi was positively correlated to root carbon content, but negatively to the numbers of root forks per root length. Our results suggest that the fine root economics spectrum is important for predicting broader groups of soil microorganisms (i.e. fungi and bacteria), while individual root traits may be more important for predicting soil microbial taxa and functional guilds.

* Information about the incidence and magnitude of local adaptation can help to predict the response of natural populations to a changing environment, and should be of particular interest in arctic and alpine environments where the effects of climate change are expected to be severe. * To quantify adaptive differentiation in the arctic-alpine perennial herb Arabis alpina, we conducted reciprocal transplant experiments for 3 yr between Spanish and Scandinavian populations. At the sites of one Spanish and one Scandinavian population, we planted seedlings representing two Spanish and four Scandinavian populations, and recorded survival, flowering propensity and fecundity. The experiment was replicated in two subsequent years. * The results demonstrate strong adaptive differentiation between A. alpina populations from the two regions. At the field site in Spain, survival and fruit production of Spanish populations were higher than those of Scandinavian populations, while the opposite was true at the site in Scandinavia, and these differences were consistent across years. By comparison, fitness varied little among populations from the same region. * The results suggest that the magnitude and geographical scale of local adaptation need to be considered in predictions of the effects of global change on the dynamics of arctic and alpine plant populations.

Considerable progress has been made in ecological and evolutionary genetics with studies demonstrating how genes underlying plant and microbial traits can influence adaptation and even 'extend' to influence community structure and ecosystem level processes. Progress in this area is limited to model systems with deep genetic and genomic resources that often have negligible ecological impact or interest. Thus, important linkages between genetic adaptations and their consequences at organismal and ecological scales are often lacking. Here we introduce the Sphagnome Project, which incorporates genomics into a long-running history of Sphagnum research that has documented unparalleled contributions to peatland ecology, carbon sequestration, biogeochemistry, microbiome research, niche construction, and ecosystem engineering. The Sphagnome Project encompasses a genus-level sequencing effort that represents a new type of model system driven not only by genetic tractability, but by ecologically relevant questions and hypotheses.