Sven Jonasson

Global Warming and Terrestrial Ecosystems: A Conceptual Framework for Analysis

raise global mean temperature over the next century by 1.0–3.5 °C (Houghton et al. 1995, 1996). Ecologists from around the world have begun experiments to investigate the effects of global warming on terrestrial ecosystems, the aspect of global climate change that attracts the most public attention (Woodwell and McKenzie 1995, Walker and Steffen 1999). The effort to understand response to warming builds on a history of investigations of the effects of elevated CO 2 on plants and ecosystems (Koch and Mooney 1996, Schulze et al. 1999). There are important differences, however, between increases in atmospheric CO 2 and temperature change, both in the temporal and spatial patterns of change and in how they affect ecosystems. The scientists involved in temperature change research have had to face new technical and conceptual challenges in designing and interpreting their experiments (Schulze et al. 1999). In this paper we describe these challenges and present a conceptual framework for interpreting experimental results and predicting effects of warming on ecosystems.

RESPONSES IN MICROBES AND PLANTS TO CHANGED TEMPERATURE, NUTRIENT, AND LIGHT REGIMES IN THE ARCTIC

Previous research has shown that experimental perturbations of arctic ecosystems simulating direct and indirect effects of predicted environmental changes have led to strong responses in the plant communities, mostly associated with increased plant nutrient availability. Similarly, changes in decomposition and nutrient mineralization are likely to occur if the soil warms and the soil moisture conditions are altered. Plant and microbial responses have usually been investigated separately, and few, if any, studies have addressed simultaneous responses to environmental changes in plants and soil microorganisms, except in models. We measured simultaneous responses in biomass, nitrogen (N), and phosphorus (P) incorporation in plants and microorganisms after five years of factorial fertilizer addition, air warming, and shading. We expected increased N and P uptake by microorganisms after fertilizer addition and also after warming, due to increases in mineralization rates in warmer soils. Plant productivity and N and P uptake were expected to increase after fertilizer addition but less after warming, because microbes were expected to absorb most of the extra released nutrients. Shading was expected to decrease plant production and also microbial biomass, due to the reduced production of labile carbon (C) in plant root exudates associated with reduced photosynthesis. We found that the plants responded strongly to fertilizer addition by increased biomass accumulation and N and P uptake. They responded less to warming, but more than expected, showing a decline in N and P concentrations in many cases. There were few significant responses to shading. The strongest response was found in combined fertilizer addition and warming treatments. All functional vascular plant groups responded similarly. However, mosses declined under those conditions when vascular plant growth was most pronounced. Contrary to our expectation, microbial C, N, and P did not increase after warming, but microbial N and P increased after shading. As expected, fertilizer addition led to increased microbial P content, whereas microbial N either increased or did not change. In general, microbial C did not change in any treatment. The microbes accumulated extra N and P only when soil inorganic N or P levels increased, suggesting that the soil microorganisms absorbed extra nutrients only in cases of declining N and P sink strength in plants.

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 NOinf3sup-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 NHinf4sup+-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: NHinf4sup+, NOinf3sup-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.

Evaluation of the point intercept method for the estimation of plant biomass

With the point intercept method, contacts are registered between plants and the tips of narrow pins passed into the vegetation. The summed number of contacts over a large number of pin positions has been used to estimate plant cover and leaf areas. This study shows that number of point intercepts also correlates highly with biomass. The intercepts can, therefore, be used as a regression variable to predict the mass of both stems and leaves in plant stands. Doubly In-transformed and untransformed simple regression models yielded high explained variance for biomass of graminoids and dwarf shrubs when regressed on number of pin contacts in quadrat samples. For transformed variables, r2-values ranged from 0.81 to 0.93 in quadrats sampled with 200 pins each, and between 0.67 and 0.80 when sampling intensity was reduced to 50 pins per quadrat. Time studies combined with calculations of how to optimize the sampling effort between 1) number of quadrats to sample and 2) number of pin positions per quadrat, revealed that the time needed for obtaining a regression equation of high precision (including the time for both registrations of pin contacts and destructive biomass estimation) is relatively limited. Subsequent estimation of biomass, from number of pin contacts only, are rapid in comparison with, e.g., destructive estimations, particularly if samples need to be fractionated into different components.

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.

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

Abstract 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.

Mineralization and microbial immobilization of N and P in arctic soils in relation to season, temperature and nutrient amendment

Abstract In situ summer, winter and annual net N and P mineralization and microbial immobilization of mineralized nutrients were measured in a low-altitude subarctic/alpine dwarf shrub heath and in a high-altitude fellfield. Net mineralization was determined by using the buried-bag technique, which was combined with fumigation–extraction to recover microbial nutrients and estimate microbial nutrient immobilization. The measurements were carried out in unperturbed plots and in plots, which had been subjected to elevated temperature and fertilizer addition for 5 years before the experiment started. During the growing season, the microbes in the unperturbed plots immobilized the major part of the mineralized nutrients, which resulted in low net mineralization. This occurred also during winter, when we assumed that nutrients should be released during microbial die-back. However, we found no evidence for a large microbial winter die-back and the surviving microbial biomass even immobilized extra nutrients. Soil temperature enhancement by ca. 2°C in general increased, or tended to increase, net mineralization. However, there was not necessarily any strong correlation between net mineralization and temperature because temperature-induced increase in gross mineralized nutrients could either lead to nutrient immobilization in the microbes or the nutrients could be released to the soil inorganic pool. Fertilizer additions had no major effect on net nutrient mineralization or nutrient immobilization. However, the ligno-cellulose index, which has been used as a predictor of substrate quality and usually correlates negatively with decomposition rate, also appeared to be a good predictor of gross mineralization, but a poor predictor of net mineralization.

Plant responses to fertilization and species removal in tundra related to community structure and clonality

A range of tundra plant communities were experimentally perturbed in 1984 by 1) application of NPK-fertilizer to three stable and two frost-heaved sites and 2) by removal of the dominant deciduous shrubs from the three stable tundra sites. Effects on vertically projected plant area (PPA) on perturbed plots and on unperturbed controls were measured by point intercepts in 1985, 1986 and 1988. There were no significant changes of the PPA of vascular plants and bryophytes on the control and removal plots during 1984-1988, and few changes among lichens. Fertilization increased the PPA of vascular plants strongly, reduced the PPA of lichens and increased or reduced the PPA of bryophytes. The PPA of only two vascular species declined, whereas that of 21 species increased (...)

Coupling of nutrient cycling and carbon dynamics in the Arctic, integration of soil microbial and plant processes

Abstract Most studies of nutrient cycling in arctic ecosystems have either addressed questions of plant nutrient acquisition or of decomposition and mineralization processes, while few studies have integrated processes in both the soil and plant compartments. Here, we synthesize information on nutrient cycling within, and between, the soil/microbial and the plant compartments of the ecosystems and integrate the cycling of nutrients with the turnover of organic matter and the carbon balance in tundra ecosystems. Based on this compilation and integration, we discuss implications for ecosystem function in response to predicted climatic changes. Many arctic ecosystems have high amounts of nutrients in the microbial biomass compared to the pools in the plant biomass both due to large nutrient-containing organic deposits in the soil and low plant biomass. The microbial pools of N and P, which are the most commonly limiting nutrients for plant production, may approach (N) or even exceed (P) the plant pools. Net nutrient mineralization is low, the residence time of nutrients in the soil is long and the nutrients are strongly immobilized in the soil microorganisms. This contributes to pronounced nutrient limitation for plant productivity, implies that the microbial sink strength for nutrients is strong and that the microbes may compete with plants for nutrients, but also that they are a potential source of plant nutrients during periods of declining microbial populations. The extent of this competition is poorly explored and it is uncertain whether plants mainly take up nutrients continuously during the summer when the microbial activity and, presumably, also the microbial sink strength is high, or whether the main nutrient uptake occurs during pulses of nutrient release when the microbial sink strength declines. Improved knowledge of mechanisms for plant-microbial interactions in these nutrient-limited systems is important, because it will form a basis also for our understanding of the C exchange between the ecosystems and the atmosphere under the predicted, future climatic change. High microbial nutrient immobilization, i.e. low release of plant-available nutrients, paired with high microbial decomposition of soil organic matter will lead to a loss of C from the soil to the atmosphere, which may not be compensated fully by increased plant C fixation. Hence, the system will be a net source of atmospheric C. Conversely, if plants are able to sequester extra nutrients efficiently, their productivity will increase and the systems may accumulate more C and turn into a C sink, particularly if nutrients are allocated to woody tissues of low nutrient concentrations.

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.