Comparison of vertical and horizontal flow planted and unplanted subsurface flow wetlands treating municipal wastewater.

In the search for design criteria for constructed wetlands (CWs) in Nepal a semi-scale experimental setup including horizontal flow (HF) and vertical flow (VF) CWs was developed. This paper compares the performance of HF and VF wetlands, and planted with unplanted beds. The experimental setup consists of two units of HF and VF beds of size 6 m × 2 m × 0.6 m and 6 m × 2 m × 0.8 m (length × width × depth) respectively. For both HF and VF systems, one unit was planted with Phragmites karka (local reed) and one was not planted. The systems were fed with wastewater drawn from the grit chamber of a municipal wastewater treatment plant. The media consisted of river gravel. In the first phase of the experiment the hydraulic loading rate (HLR) was varied in steps; 0.2, 0.08, 0.04 m(3)/m(2)/d and the percent removal increase with decrease in HLR for all beds and parameters except for total phosphorus. In the second phase the loading rate of 0.04 m(3)/m(2)/d was run for 7 months. In both parts of the experiment the planted beds performed better than the unplanted beds and the VF better than the HF beds. To meet Nepalese discharge standards HF beds are sufficient, but to meet stricter requirements a combination of HF and VF beds are recommended.

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.

Root mechanical properties related to disturbed and stressed habitats in the Arctic.

Various mechanical and architectural properties of roots were measured on plants characteristic of different levels of soil disturbance associated with frost-heave cycles on sorted polygons in Swedish Lapland: one of these measures, resilience, has not, apparently, been recorded previously in literature. Some roots were sampled from species which occurred on both disturbed and stable soils. Root length, root angle, numbers of roots and root diameter varied greatly between species but the degree of branching of roots was generally low. Among all species, and particularly forbs and graminoids which are the most common life forms on polygons, there was a clear trend of decreasing root diameter with increasing soil disturbance, while the species growing on both disturbed and stable ground showed no clear trend associated with soil disturbance. The small root diameter of species growing on the polygons could result from the different species and life forms found there together with a younger age class distribution of plants, younger plants having smaller roots. Only one third of the species growing on the most disturbed soils had rhizomes, whereas twice as many species possessed rhizomes in areas of lower disturbance. The tensile strength, breaking strain (a measure of how much a root is deformed by stretching), breaking stress (a measure of root strength corrected for the cross-sectional area) and the resilience (a measure of the elastic-recovery of stretched roots) of the roots varied highly significantly among the species. The tensile strength varied 6.5-fold, the breaking strain varied by up to almost 8-fold and the breaking stress 13-fold. The modulus of resilience varied least, by a factor of only two. There was no clear trend in the way mechanical properties of the roots of the three species occurring on both stable and frost-heaved soils varied between the two regimes. However, among all species, those from the frost-heaved soils tended to have the weakest roots, because of their small diameter, but the strongest roots per cross-sectional area (i.e. they had the greatest breaking stress). Among life-forms, graminoids toler̀Gated a significantly greater strain than forbs and dwarf shrubs and much higher stress before they broke. Dwarf shrubs and forbs had significantly higher resilience than graminoids and the highest tensile strengths. Forbs showed the lowest values for all of the measured variables except for resilience, which fell between the values for graminoids and dwarf shrubs. There were no significant differences in any of the mechanical measures of root resistance among species of different frost-heaving regimes, indicating that there are few architectural or mechanical properties which enable roots to survive in the most disturbed soil. Indeed, the absence of a complete plant canopy and an increase in abundance of young plants on the polygon area compared with the stable areas suggest that if there are any adaptations to strong disturbance, they are less than adequate and the hypothesis that arctic plants avoid, rather than adapt, to severe environment is supported.

Seasonal uptake and allocation of phosphorus in Eriophorum vaginatum L measured by labelling with 32 P.

Seasonal measurements of phosphorus (P) pools in the tussock-forming sedge Eriophorum vaginatum L. indicated low annual P uptake, but pronounced fluxes of P to growing biomass from below-ground stores in spring, and a return in autumn. Additions of 32 P to tussocks in early spring and mid summer confirmed this deduction. Addition of 32 P caused a rapid incorporation of about 10% of the labelled P in the tillers within a few days. Subsequent uptake was negligible, presumably because the 32 P not immediately absorbed was fixed in the soil and unavailable to plants. This demonstrates that nutrient pulses were more important than steady-state mineralization in supporting P uptake by Eriophorum vaginatum. After the 32 P additions, roots initially retained 60% of the total 32 P taken up. During the following weeks, part of the 32 P was transported slowly to stem bases and leaf sheaths, the principal storage organs. A lower proportion was transported to the leaf blades. We suggest that storage and recirculation of nutrients are the main sources of the annual nutrient supply to growth of E. vaginatum under normal field conditions, where P fixation in the soil and low rate of P diffusion towards the roots limit P uptake from soil except during periods of pulsed release.

Environmental control and intersite variations of phenolics in Betula nana in tundra ecosystems.

• Secondary metabolites make leaves unpalatable for herbivores and influence decomposition. Site-specific differences are presented in phenolics and nitrogen in Betula nana leaves from dwarf shrub tundra at Abisko, northern Sweden, and from tussock tundra at Toolik Lake, Alaska, subjected to a decade of warming, fertilization and shading. • Nitrogen and a number of phenolics, including condensed and hydrolysable tannins, flavonoids, phenolic glucosides and chlorogenic acids, were analysed in B. nana leaves. • Phenolic concentrations showed marked between-site differences (e.g. condensed tannins were 50% higher at Abisko than at Toolik); responses to the environmental manipulations were more pronounced at Toolik compared with Abisko. Warming increased condensed tannins and decreased hydrolysable tannins at Toolik, but had no effect at Abisko, whereas fertilization and shading generally decreased concentrations. • Betula invests less carbon in phenolics at Toolik than at Abisko and shows a greater response to environmental changes by investing more carbon in growth and less to phenolic production. Hence, the Toolik population has a lower herbivore-defense level, which declines further if nutrient availability increases. By contrast, under warmer conditions, the increase in bulk phenolics and decrease in leaf palatability are greater at Toolik than at Abisko.

Differential responses of grass and a dwarf shrub to long-term changes in soil microbial biomass C, N and P following factorial addition of NPK fertilizer, fungicide and labile carbon to a heath.

Microbial immobilization may decrease the inorganic nutrient concentrations of the soil to the extent of affecting plant nutrient uptake and growth. We have hypothesized that graminoids with opportunistic nutrient-acquisition strategies are strongly influenced by nutrient limitation imposed by microbes, whereas growth forms such as dwarf shrubs are less affected by the mobilization-immobilization cycles in microbes. By adding NPK fertilizer, labile C (sugar) and fungicide (benomyl) over a 5 yr period in a fully factorial design, we aimed to manipulate the sink-source potential for nutrients in a non-acidic heath tundra soil. After 2 yr, N and P accumulated in the microbial biomass after fertilization with no change in microbial C, which suggests that nutrients did not limit microbial biomass growth. After 5 yr, microbial C was enhanced by 60% in plots with addition of labile C, which points to C-limitation of the microbial biomass. Microbial biomass N and P tended to increase following addition of labile C, by 10 and 25%, respectively. This caused decreased availability of NH4 + and P, showing close microbial control of nutrient availability. The most common graminoid, Festuca ovina, responded to fertilizer addition with a strong increase, and to labile C addition with a strong decrease in cover, providing the first direct field evidence that nutrient limitation imposed by immobilizing microbes can affect the growth of tundra plants. Also in support of our hypothesis, following addition of labile C the concentrations of N and K in leaves and that of N in roots of F. ovina decreased, whilst the demand of roots for P increased. In contrast, the most common dwarf shrub, Vaccinium uliginosum, was only slightly sensitive to changes in resource availability, showing no cover change after 4 yr addition of labile C and fertilizer, and little change in leaf nutrient concentrations. We suggest that the differential responses of the two growth forms are due to differences in storage and nutrient uptake pathways, with the dwarf shrub having large nutrient storage capacity and access to organic forms of N through its mycorrhizal association. While the fungicide had no effect on ericoid mycorrhizal colonization of roots or symbiotic function inferred from plant 15 N natural abundance, it decreased microbial biomass C and N after 2 yr. Throughout the fifth season, the availability of soil NO3 - and inorganic P was decreased with no change in microbial biomass C, N or P, suggesting a negative impact of benomyl on N and P mineralization.

Significance of sequential leaf development for nutrient balance of the cotton sedge,Eriophorum vaginatum L.

The sedgeEriophorum vaginatum in an interior Alaskan muskeg site produced leaves sequentially at about 1.5-month intervals. Each leaf remained active for two growing seasons. Young leaves (even those initiated late in the season) always had high concentrations of N, P, K and Mg and were low in Ca. Stems had high concentrations of nutrients, sugar, amino acid N and soluble organic P in autumn and spring but low concentrations in summer. Growth of leaves in spring was strongly supported by translocation from storage. Leaves approached their maximum nutrient pool before nutrient uptake began in late spring, one month before maximum biomass. Retranslocation of nutrients from aging leaves could support nutrient input into new, actively growing leaves as a consequence of the sequential leaf development. For instance retranslocation from aging leaves accounted for more than 90 and 85% of P and N input to new leaves appearing in early summer and 100% to leaves that appeared later. Leaching losses were negligible. Half time for decay of standing dead litter was 10 years. We suggest that sequential leaf development paired with highly efficient remobilization of nutrients from senescing leaves enables plants to recycle nutrients within the shoot and minimize dependence upon soil nutrients. This may be an important mechanism enablingEriophorum vaginatum to dominate nutrient-poor sites. This may also explain why graminoids with sequential leaf production cooccur with evergreen shrubs and dominate over forbs and deciduous shrubs in nutrient-poor sites in the boreal forest (e.g., in bogs) and at the northern limit of the tundra zone.

Effects on plant production after addition of labile carbon to arctic/alpine soils.

Biomass production was analysed in Festuca vivipara, grown for 3 months in pots with non-sterilized or sterilized soil after factorial addition of three levels of labile carbon combined with high and low levels of N and P. The soil was a nutrient-poor subarctic heath soil. In the non-sterilized soil plant biomass production increased strongly only in the treatment with high levels of both N and P, which suggests that both nutrients limited plant growth. In the sterilized soil addition of a high level of N without P addition gave almost the same growth response as in the combined NP treatment. This was because of a more than 30-fold increase of inorganic phosphorus in the soil as P was released from the killed microbial biomass after sterilization. Sugar addition reduced plant growth in all treatments. The reduction in plant growth was dose dependent within the range of 0-450 µg C g-1 soil added to the non-sterilized soil, but the response levelled off at 233 µg C g-1 soil in the soil that had been sterilized at the start of the experiment. The plant response, together with observed depletion of soil inorganic N and P, indicated that the microbial biomass immobilized nutrients efficiently and reduced plant growth when extra labile carbon was added. The inhibition of growth was lower, however, in the soil which had been sterilized, probably because of a slow recovery of the microbial populations in it. Two of the nutrient-carbon solutions closely matched the N, P and C concentrations in a solution containing leaf extracts of Cassiope tetragona and Betula tortuosa that had been used previously to test for possible allelopathic effects of compounds in the leaf extracts. These extracts also reduced plant growth. The growth reduction was equally large or larger after nutrient-sugar addition than after addition of leaf extracts in three out of the four possible combinations of species and sterilized or non-sterilized soil. In the fourth case (Betula extract added to sterilized soil), the effect was larger when leaf extract was added than after addition of the nutrient-carbon solution. This could be due to a low rate of microbial degradation of phytotoxic substances in this soil because of a slow recovery of the microbial populations after sterilization. The generally stronger or equal effect of the nutrient-sugar addition compared to the leaf extract addition leads to the conclusion that microbial nutrient immobilization and microbial competition for nutrients increased as a function of labile carbon addition with the extract. Hence, it appears that enhanced microbial activity and microbial nutrient immobilization rather than phytotoxic effects was the primary reasons for the reduced biomass production in F. vivipara even after addition of the leaf extracts.

Effects of labile soil carbon on nutrient partitioning between an arctic graminoid and microbes.

We measured partitioning of N and P uptake between soil microorganisms and potted Festuca vivipara in soil from a subarctic heath in response to factorial addition of three levels of labile carbon (glucose) combined with two levels of inorganic N and P. The glucose was added to either non-sterilized or sterilized (autoclaved) soils in quantities which were within the range of reported, naturally occurring amounts of C released periodically from the plant canopy. The aims were, firstly, to examine whether the glucose stimulated microbial nutrient uptake to the extent of reducing plant nutrient uptake. This is expected in nutrient-deficient soils if microbes and plants compete for the same nutrients. Secondly, we wanted to test our earlier␣interpretation that growth reduction observed in graminoids after addition of leaf extracts could be caused directly by labile carbon addition, rather than by phytotoxins in the extracts. Addition of high amounts of N did not affect the microbial N pool, whereas high amounts of added P significantly increased the microbial P pool, indicating a luxury P uptake in the microbes. Both plant N and in particular P uptake increased strongly in response to soil sterilization and to addition of extra N or P. The increased␣uptake led to enhanced plant growth when both elements were applied in high amounts, but only led to increased tissue concentrations without growth responses when the nutrients were added separately. Glucose had strong and contrasting effects on plant and microbial N and P uptake. Microbial N and P uptake increased, soil inorganic N and P concentrations were reduced and plant N and P uptake declined when glucose was added. The responses were dose-dependent within the range of 0-450 µg C g-1 soil added to the non-sterilized soil. The opposite responses of plants and microbes showed that plant acquisition of limiting nutrients is dependent on release of nutrients from the soil microbes, which is under strong regulation by the availability and microbial uptake of labile C. Hence, we conclude, firstly, that the microbial populations can compete efficiently with plants for nutrients to an extent of affecting plant growth when the microbial access to labile carbon is high in nutrient deficient soils. We also conclude that reduced growth of plants after addition of leaf extracts to soil can be caused by carbon-induced shifts in nutrient partitioning between plants and microbes, and not necessarily by phytotoxins added with the extracts as suggested by some experiments.

Vascular plant 15N natural abundance in heath and forest tundra ecosystems is closely correlated with presence and type of mycorrhizal fungi in roots.

In this study we show that the natural abundance of the nitrogen isotope 15, δ15N, of plants in heath tundra and at the tundra-forest ecocline is closely correlated with the presence and type of mycorrhizal association in the plant roots. A total of 56 vascular plant species, 7 moss species, 2 lichens and 6 species of fungi from four heath and forest tundra sites in Greenland, Siberia and Sweden were analysed for δ15N and N concentration. Roots of vascular plants were examined for mycorrhizal colonization, and the soil organic matter was analysed for δ15N, N concentration and soil inorganic, dissolved organic and microbial N. No arbuscular mycorrhizal (AM) colonizations were found although potential host plants were present in all sites. The dominant species were either ectomycorrhizal (ECM) or ericoid mycorrhizal (ERI). The δ15N of ECM or ERI plants was 3.5-7.7‰ lower than that of non-mycorrhizal (NON) species in three of the four sites. This corresponds to the results in our earlier study of mycorrhiza and plant δ15N which was limited to one heath and one fellfield in N Sweden. Hence, our data suggest that the δ15N pattern: NON/AM plants > ECM plants ≥ ERI plants is a general phenomenon in ecosystems with nutrient-deficient organogenic soils. In the fourth site, a␣birch forest with a lush herb/shrub understorey, the differences between functional groups were considerably smaller, and only the ERI species differed (by 1.1‰) from the NON species. Plants of all functional groups from this site had nearly twice the leaf N concentration as that found in the same species at the other three sites. It is likely that low inorganic N availability is a prerequisite for strong δ15N separation among functional groups. Both ECM roots and fruitbodies were 15N enriched compared to leaves which suggests that the difference in δ15N between plants with different kinds of mycorrhiza could be due to isotopic fractionation at the␣fungal-plant interface. However, differences in δ15N between soil N forms absorbed by the plants could also contribute to the wide differences in plant δ15N found in most heath and forest tundra ecosystems. We hypothesize that during microbial immobilization of soil ammonium the microbial N pool could become 15N-depleted and the remaining, plant-available soil ammonium 15N-enriched. The latter could be a main source of N for NON/AM plants which usually have high δ15N. In contrast, amino acids and other soil organic N compounds presumably are 15N-depleted, similar to plant litter, and ECM and ERI plants with high uptake of these N forms hence have low leaf δ15N. Further indications come from the δ15N of mosses and lichens which was similar to that of ECM plants. Tundra cryptogams (and ECM and ERI plants) have previously been shown to have higher uptake of amino acid than ammonium N; their low δ15N might therefore reflect the δ15N of free amino acids in the soil. The concentration of dissolved organic N was 3-16 times higher than that of inorganic N in the sites. Organic nitrogen could be an important N source for ECM and, in particular, ERI plants in heath and forest tundra ecosystems with low release rate of inorganic N from the soil organic matter.

Shoot biomass, δ13C, nitrogen and chlorophyll responses of two arctic dwarf shrubs to in situ shading, nutrient application and warming simulating climatic change.

As climatic change might induce ecophysiological changes in plants which affect their long-term performance, we investigated responses in above-ground biomass, δ13C, nitrogen and chlorophyll of two evergreen arctic dwarf shrubs, Cassiope tetragona and Empetrum hermaphroditum, to 5 (biomass, N) or 6 years of shading, nutrient application and air/soil warming at a dwarf shrub dominated tree-line heath (450 m a.s.l) and a high altitude fellfield (1100 m a.s.l.) in Swedish Lapland. Warming enhanced the green biomass (equivalent to the last 3-4 years of leaf production) and the ratio of green to brown biomass of C. tetragona at the fellfield, and diluted the shoot N concentration. Fertilizer application led to higher shoot N concentration and larger green-to-brown biomass ratio at both sites, and fertilizer application and warming generally had an additive effect on the green biomass. We conclude that both warming and increased soil nutrient availability stimulated the growth of C. tetragona at the fellfield whereas at the heath there was a clear increase in production only if enhanced temperature was combined with nutrient application. Across treatments C. tetragona at the fellfield had 0.6‰ higher δ13C and 1.4 mg g-1 more leaf N, and the soil organic matter δ13C was 1.0‰ higher at the fellfield than at the heath. However, an increase in shoot N concentration with altitude does not necessarily lead to higher δ13C as no differences in δ13C were observed when leaf N of the two dwarf shrubs was increased by fertilizer application c. tetragona in non-warmed plots had higher δ13C values than those from warmed plots at the same altitude, which provides the first in situ experimental validation of the theory that temperature partly is responsible for altitudinal trends in plant carbon isotope discrimination. Increased biomass and chlorophyll concentration of C. tetragona in warmed plots points to increased assimilation, at least at the fellfield. As the δ13C-based and, therefore, time-integrated estimate of the ratio of CO2 concentration in the leaf intercellular spaces to that in the atmosphere (C i/C a) also increased, warming probably enhanced the stomatal conductance relatively more than the C assimilation, which may be harmful if climatic change leads to reduced soil moisture content and increased plant competition for water. At both sites C. tetragona and E. hermaphroditum responded to shade by increasing the concentration of shoot N and photosynthetic pigments whereas biomass production (and therefore also net photosynthesis) did not decline. Shade was accompanied by a 0.6-1.3‰ (E. hermaphroditum) or 1.2-2.2‰ (C. tetragona) decrease in δ13C. This could be due to enhanced stomatal conductance with shading, and perhaps to shade reducing the ericoid mycorrhizal uptake of soil organic C, a factor which has been overlooked as an influence on plant δ13C.

Leaf 15N abundance of subarctic plants provides field evidence that ericoid, ectomycorrhizal and non-and arbuscular mycorrhizal species access different sources of soil nitrogen.

The natural abundance of the nitrogen isotope 15, δ15N, was analysed in leaves of 23 subarctic vascular plant species and two lichens from a tree-line heath at 450 m altitude and a fellfield at 1150 m altitude close to Abisko in N. Sweden, as well as in soil, rain and snow. The aim was to reveal if plant species with different types of mycorrhizal fungi also differ in their use of the various soil N sources. The dwarf shrubs and the shrubs, which in combination formed more than 65% of the total above-ground biomass at both sites, were colonized by ericoid or ectomycorrhizal fungi. Their leaf δ15N was between-8.8 and-5.5‰ at the heath and between-6.1 and -3.3‰ at the fellfield. The leaf δ15N of non- or arbuscular mycorrhizal species was markedly different, ranging from -4.1 to -0.4‰ at the heath, and from -3.4 to+2.2‰ at the fellfield. We conclude that ericoid and ectomycorrhizal dwarf shrubs and shrubs utilize a distinct N source, most likely a fraction of the organic N in fresh litter, and not complexed N in recalcitrant organic matter. The latter is the largest component of soil total N, which had a δ15N of -0.7‰ at the heath and +0.5‰ at the fellfield. Our field-based data thus support earlier controlled-environment studies and studies on the N uptake of excised roots, which have demonstrated protease activity and amino acid uptake by ericoid and ectomycorrhizal tundra species. The leaves of ectomycorrhizal plants had slightly higher δ15N (fellfield) and N concentration than leaves of the ericoids, and Betula nana, Dryas octopetala and Salix spp. also showed NO inf3sup- reductase activity. These species may depend more on soil inorganic N than the ericoids. The δ15N of non- or arbuscular mycorrhizal species indicates that the δ15N of inorganic N available to these plants was higher than that of average fresh litter, probably due to high microbial immobilization of inorganic N. The δ15N of NH inf4sup+ -N was +12.3‰ in winter snow and +1.9‰ in summer rain. Precipitation N might be a major contributer in species with poorly developed root systems, e.g. Lycopodium selago. Our results show that coexisting plant species under severe nutrient limitation may tap several different N sources: NH inf4sup+ , NO inf3sup- and organic N from the soil, atmospheric N2, and N in precipitation. Ericoid and ectomycorrhizal fungi are of major importance for plant N uptake in tundra ecosystems, and mycorrhizal fungi probably exert a major control on plant δ15N in organic soils.

In situ mineralization of nitorgen and phosphorus of arctic soils after perturbations simulating climate change.

Seasonal net nitrogen (N) and phosphorus (P) mineralization was investigated at Abisko, Swedish Lapland in soils of a subarctic heath and in soils of a colder (by about 4° C), high altitude fellfield by (a) using in situ soil incubation in soils which had been shaded or subjected to two levels of increased temperature, combined with (b) reciprocal transplantation of soils between the two sites. Proportionally large and significant net seasonal mineralization of N, in contrast to non-significant P mineralization, was found in untransplanted and transplanted fellfield soil. In contrast, P was mineralized in proportionally large amounts, in contrast to low N mineralization, in the transplanted and untransplanted heath soil. The differences indicate that P was strongly immobilized in relation to N at the fellfield and that N was more strongly immobilized than P in the heath soil. The immobilization in both soils remained high even after a temperature change of 4-5° C experienced by transplanted soils. Air temperature increases of up to 4-5° C in greenhouses resulted in a soil temperature increase of 1-2° C and did not cause any extra increase of net N and P mineralization. The results suggest that soil temperature increases of up to 2° C, which are likely to occur by the end of the next century as an effect of a predicted 4-5° C rise in air temperature, have only small effects on net mineralization in at least two characteristic tundra soils. These effects are probably smaller than the natural fluctuation of plant available nutrients from site to site, even within the same plant community. A further soil temperature increase of up to 4-5° C may enhance decomposition and gross mineralization, but the rate of net mineralization, and hence the change of nutrient availability to the plants, depends on the extent of microbial immobilization of the extra nutrients released.

Microbial biomass C, N and P in two arctic soils and responses to addition of NPK fertilizer and sugar: implications for plant nutrient uptake.

The soil microbial carbon (C), nitrogen (N) and phosphorus (P) pools were quantified in the organic horizon of soils from an arctic/alpine low-altitude heath and a high-altitude fellfield by the fumigation-extraction method before and after factorial addition of sugar, NPK fertilizer and benomyl, a fungicide. In unamended soil, microbial C, N and P made up 3.3-3.6%, 6.1-7.3% and 34.7% of the total soil C, N and P content, respectively. The inorganic extractable N pool was below 0.1% and the inorganic extractable P content slightly less than 1% of the total soil pool sizes. Benomyl addition in spring and summer did not affect microbial C or nutrient content analysed in the autumn. Sugar amendments increased microbial C by 15 and 37% in the two soils, respectively, but did not affect the microbial nutrient content, whereas inorganic N and P either declined significantly or tended to decline. The increased microbial C indicates that the microbial biomass also increased but without a proportional enhancement of N and P uptake. NPK addition did not affect the amount of microbial C but almost doubled the microbial N pool and more than doubled the P pool. A separate study has shown that CO2 evolution increased by more than 50% after sugar amendment and by about 30% after NPK and NK additions to one of the soils. Hence, the microbial biomass did not increase in response to NPK addition, but the microbes immobilized large amounts of the added nutrients and, judging by the increased CO2 evolution, their activity increased. We conclude: (1) that microbial biomass production in these soils is stimulated by labile carbon and that the microbial activity is stimulated by both labile C and by nutrients (N); (2) that the microbial biomass is a strong sink for nutrients and that the microbial community probably can withdraw substantial amounts of nutrients from the inorganic, plant-available pool, at least periodically; (3) that temporary declines in microbial populations are likely to release a flush of inorganic nutrients to the soil, particularly P of which the microbial biomass contained more than one third of the total soil pool; and (4) that the mobilization-immobilization cycles of nutrients coupled to the population dynamics of soil organisms can be a significant regulating factor for the nutrient supply to the primary producers, which are usually strongly nutrient-limited in arctic ecosystems.

Inhibition of growth, and effects on nutrient uptake of arctic graminoids by leaf extracts - allelopathy or resource competition between plants and microbes?

Previous research has shown that plant extracts, e.g. from boreal dwarf shrubs and trees, can cause reduced growth of neighbouring plants: an effect known as allelopathy. To examine whether arctic and subarctic plants could also be affected by leaching of phytochemicals, we added extracts from the commonly occurring arctic dwarf shrubs Cassiope tetragona and Empetrum hermaphroditum, and from mountain birch, Betula pubescens ssp. tortuosa to three graminoid species, Carex bigelowii, Festuca vivipara and Luzula arcuata, grown in previously sterilized or non-sterilized arctic soils. The graminoids in non-sterilized soil grew more slowly than those in sterilized soil. Excised roots of the plants in non-sterilized soil had higher uptake rate of labelled P than those in sterilized soil, demonstrating larger nutrient deficiency. The difference in growth rate was probably caused by higher nutrient availability for plants in soils in which the microbial biomass was killed after soil sterilization. The dwarf shrub extracts contained low amounts of inorganic N and P and medium high amounts of carbohydrates. Betula extracts contained somewhat higher levels of N and much higher levels of P and carbohydrates. Addition of leaf extracts to the strongly nutrient limited graminoids in non-sterilized soil tended to reduce growth, whereas in the less nutrient limited sterilized soil it caused strong growth decline. Furthermore, the N and P uptake by excised roots of plants grown in both types of soil was high if extracts from the dwarf shrubs (with low P and N concentrations) had been added, whereas the P uptake declined but the N uptake increased after addition of the P-rich Betula extract. In contrast to the adverse extract effects on plants, soil microbial respiration and soil fungal biomass (ergosterol) was generally stimulated, most strongly after addition of the Betula extract. Although we cannot exclude the possibility that the reduced plant growth and the concomitant stimulation of microbial activity were caused by phytochemicals, we believe that this was more likely due to labile carbon in the extracts which stimulated microbial biomass and activity. As a result microbial uptake increased, thereby depleting the plant available pool of N and P, or, for the P-rich Betula extract, depleting soil inorganic N alone, to the extent of reducing plant growth. This chain of events is supported by the negative correlation between plant growth and sugar content in the three added extracts, and the positive correlation between microbial activity, fungal biomass production and sugar content, and are known reactions when labile carbon is added to nutrient deficient soils.

Past changes in Arctic terrestrial ecosystems, climate and UV radiation.

At the last glacial maximum, vast ice sheets covered many continental areas. The beds of some shallow seas were exposed thereby connecting previously separated landmasses. Although some areas were ice-free and supported a flora and fauna, mean annual temperatures were 10-13 degrees C colder than during the Holocene. Within a few millennia of the glacial maximum, deglaciation started, characterized by a series of climatic fluctuations between about 18,000 and 11,400 years ago. Following the general thermal maximum in the Holocene, there has been a modest overall cooling trend, superimposed upon which have been a series of millennial and centennial fluctuations in climate such as the "Little Ice Age spanning approximately the late 13th to early 19th centuries. Throughout the climatic fluctuations of the last 150,000 years, Arctic ecosystems and biota have been close to their minimum extent within the most recent 10,000 years. They suffered loss of diversity as a result of extinctions during the most recent large-magnitude rapid global warming at the end of the last glacial stage. Consequently, Arctic ecosystems and biota such as large vertebrates are already under pressure and are particularly vulnerable to current and projected future global warming. Evidence from the past indicates that the treeline will very probably advance, perhaps rapidly, into tundra areas, as it did during the early Holocene, reducing the extent of tundra and increasing the risk of species extinction. Species will very probably extend their ranges northwards, displacing Arctic species as in the past. However, unlike the early Holocene, when lower relative sea level allowed a belt of tundra to persist around at least some parts of the Arctic basin when treelines advanced to the present coast, sea level is very likely to rise in future, further restricting the area of tundra and other treeless Arctic ecosystems. The negative response of current Arctic ecosystems to global climatic conditions that are apparently without precedent during the Pleistocene is likely to be considerable, particularly as their exposure to co-occurring environmental changes (such as enhanced levels of UV-B, deposition of nitrogen compounds from the atmosphere, heavy metal and acidic pollution, radioactive contamination, increased habitat fragmentation) is also without precedent.

Effects on the function of Arctic ecosystems in the short- and long-term perspectives.

Historically, the function of Arctic ecosystems in terms of cycles of nutrients and carbon has led to low levels of primary production and exchanges of energy, water and greenhouse gases have led to low local and regional cooling. Sequestration of carbon from atmospheric CO2, in extensive, cold organic soils and the high albedo from low, snow-covered vegetation have had impacts on regional climate. However, many aspects of the functioning of Arctic ecosystems are sensitive to changes in climate and its impacts on biodiversity. The current Arctic climate results in slow rates of organic matter decomposition. Arctic ecosystems therefore tend to accumulate organic matter and elements despite low inputs. As a result, soil-available elements like nitrogen and phosphorus are key limitations to increases in carbon fixation and further biomass and organic matter accumulation. Climate warming is expected to increase carbon and element turnover, particularly in soils, which may lead to initial losses of elements but eventual, slow recovery. Individual species and species diversity have clear impacts on element inputs and retention in Arctic ecosystems. Effects of increased CO2 and UV-B on whole ecosystems, on the other hand, are likely to be small although effects on plant tissue chemisty, decomposition and nitrogen fixation may become important in the long-term. Cycling of carbon in trace gas form is mainly as CO2 and CH4. Most carbon loss is in the form of CO2, produced by both plants and soil biota. Carbon emissions as methane from wet and moist tundra ecosystems are about 5% of emissions as CO2 and are responsive to warming in the absence of any other changes. Winter processes and vegetation type also affect CH4 emissions as well as exchanges of energy between biosphere and atmosphere. Arctic ecosystems exhibit the largest seasonal changes in energy exchange of any terrestrial ecosystem because of the large changes in albedo from late winter, when snow reflects most incoming radiation, to summer when the ecosystem absorbs most incoming radiation. Vegetation profoundly influences the water and energy exchange of Arctic ecosystems. Albedo during the period of snow cover declines from tundra to forest tundra to deciduous forest to evergreen forest. Shrubs and trees increase snow depth which in turn increases winter soil temperatures. Future changes in vegetation driven by climate change are therefore, very likely to profoundly alter regional climate.

Uncertainties and recommendations.

An assessment of the impacts of changes in climate and UV-B radiation on Arctic terrestrial ecosystems, made within the Arctic Climate Impacts Assessment (ACIA), highlighted the profound implications of projected warming in particular for future ecosystem services, biodiversity and feedbacks to climate. However, although our current understanding of ecological processes and changes driven by climate and UV-B is strong in some geographical areas and in some disciplines, it is weak in others. Even though recently the strength of our predictions has increased dramatically with increased research effort in the Arctic and the introduction of new technologies, our current understanding is still constrained by various uncertainties. The assessment is based on a range of approaches that each have uncertainties, and on data sets that are often far from complete. Uncertainties arise from methodologies and conceptual frameworks, from unpredictable surprises, from lack of validation of models, and from the use of particular scenarios, rather than predictions, of future greenhouse gas emissions and climates. Recommendations to reduce the uncertainties are wide-ranging and relate to all disciplines within the assessment. However, a repeated theme is the critical importance of achieving an adequate spatial and long-term coverage of experiments, observations and monitoring of environmental changes and their impacts throughout the sparsely populated and remote region that is the Arctic.

Biodiversity, distributions and adaptations of Arctic species in the context of environmental change.

The individual of a species is the basic unit which responds to climate and UV-B changes, and it responds over a wide range of time scales. The diversity of animal, plant and microbial species appears to be low in the Arctic, and decreases from the boreal forests to the polar deserts of the extreme North but primitive species are particularly abundant. This latitudinal decline is associated with an increase in super-dominant species that occupy a wide range of habitats. Climate warming is expected to reduce the abundance and restrict the ranges of such species and to affect species at their northern range boundaries more than in the South: some Arctic animal and plant specialists could face extinction. Species most likely to expand into tundra are boreal species that currently exist as outlier populations in the Arctic. Many plant species have characteristics that allow them to survive short snow-free growing seasons, low solar angles, permafrost and low soil temperatures, low nutrient availability and physical disturbance. Many of these characteristics are likely to limit species' responses to climate warming, but mainly because of poor competitive ability compared with potential immigrant species. Terrestrial Arctic animals possess many adaptations that enable them to persist under a wide range of temperatures in the Arctic. Many escape unfavorable weather and resource shortage by winter dormancy or by migration. The biotic environment of Arctic animal species is relatively simple with few enemies, competitors, diseases, parasites and available food resources. Terrestrial Arctic animals are likely to be most vulnerable to warmer and drier summers, climatic changes that interfere with migration routes and staging areas, altered snow conditions and freeze-thaw cycles in winter, climate-induced disruption of the seasonal timing of reproduction and development, and influx of new competitors, predators, parasites and diseases. Arctic microorganisms are also well adapted to the Arctic's climate: some can metabolize at temperatures down to -39 degrees C. Cyanobacteria and algae have a wide range of adaptive strategies that allow them to avoid, or at least minimize UV injury. Microorganisms can tolerate most environmental conditions and they have short generation times which can facilitate rapid adaptation to new environments. In contrast, Arctic plant and animal species are very likely to change their distributions rather than evolve significantly in response to warming.

Effects on the structure of Arctic ecosystems in the short- and long-term perspectives.

Species individualistic responses to warming and increased UV-B radiation are moderated by the responses of neighbors within communities, and trophic interactions within ecosystems. All of these responses lead to changes in ecosystem structure. Experimental manipulation of environmental factors expected to change at high latitudes showed that summer warming of tundra vegetation has generally led to smaller changes than fertilizer addition. Some of the factors manipulated have strong effects on the structure of Arctic ecosystems but the effects vary regionally, with the greatest response of plant and invertebrate communities being observed at the coldest locations. Arctic invertebrate communities are very likely to respond rapidly to warming whereas microbial biomass and nutrient stocks are more stable. Experimentally enhanced UV-B radiation altered the community composition of gram-negative bacteria and fungi, but not that of plants. Increased plant productivity due to warmer summers may dominate food-web dynamics. Trophic interactions of tundra and sub-Arctic forest plant-based food webs are centered on a few dominant animal species which often have cyclic population fluctuations that lead to extremely high peak abundances in some years. Population cycles of small rodents and insect defoliators such as the autumn moth affect the structure and diversity of tundra and forest-tundra vegetation and the viability of a number of specialist predators and parasites. Ice crusting in warmer winters is likely to reduce the accessibility of plant food to lemmings, while deep snow may protect them from snow-surface predators. In Fennoscandia, there is evidence already for a pronounced shift in small rodent community structure and dynamics that have resulted in a decline of predators that specialize in feeding on small rodents. Climate is also likely to alter the role of insect pests in the birch forest system: warmer winters may increase survival of eggs and expand the range of the insects. Insects that harass reindeer in the summer are also likely to become more widespread, abundant and active during warmer summers while refuges for reindeer/caribou on glaciers and late snow patches will probably disappear.