| 1. |
- Björk, Robert G., 1974, et al.
(författare)
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Climate-related soil changes in tundra ecosystems at Latnjajaure, northern Sweden – an ITEX-IPY project
- 2010
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Ingår i: International Polar Year Oslo Science Conference.
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Konferensbidrag (refereegranskat)abstract
- During the 90'ies, the International Tundra Experiment (ITEX) was established as a leading project in arctic and alpine ecology, and has become a model for many later network establishments. The present study capitalizes on the early efforts of ITEX and aims at assessing ecosystem changes in the alpine areas of northern Sweden above timberline, i.e. the tundra, in relation to global change. By using the "old" ITEX plots established during the early years of the program we have measured ecosystem respiration (ER), the Normalized Difference Vegetation Index, and nitrogen (N) mineralization over the growing season. In addition, have soil samples been taken to quantify changes in the carbon (C) and N pool, including 13C and 15N. After 12 to 15 years of open top chamber (OTC) treatment no statistical effect was found on the soil temperature (10 cm soil depth), although the was an overall increase in all OTC by +0.2°C. However, the soil moisture decreased significantly by 3-14%, depending on plant community, in the OTCs compared to ambient conditions. Preliminary, there was a 20-37% non-significant higher mean ER in the OTC compared to the ambient plots over the growing season. Furthermore, the OTC treatment did not affect the growing season mineralization of inorganic N, or total C and N content of the soil. The stable isotope data showed both enrichment and depletion as a consequence of the OTC treatment, but no general pattern was discerned. Thus, this non-significant higher ER is most likely of plant origin than soil, as the plant standing biomass has increased in the OTCs. This study does not support the current consensus that tundra soils will alter their C and N dynamics in response to climate change.
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| 2. |
- Björk, Robert G., 1974, et al.
(författare)
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A Climate Change aspect on root dynamics and nitrogen partitioning in a tundra landscape
- 2005
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Ingår i: Sediment budgets and rates of sediment transfer across cold environments in Europe. 3rd Workshop of the ESF Network SEDIFLUX, Durham, UK, 15 – 19 December 2005..
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Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract
- The Arctic Climate Impact Assessment (ACIA) recently reported that Arctic is rapidly changing due to Climate Change. Likewise, the mountains of Europe are going to experience large shifts in plant composition and 41-56% of the alpine species might be on the edge of extinction according to the 1st synthesis of the Global Observation Research Initiative in Alpine Environments (GLORIA). Although the tundra ecosystems are subjected to dramatical changes as a result of Climate Change, there is little knowledge of the effect on root dynamics and its implication on the nitrifying and denitrifying microbial community. Here, we compare nitrification enzyme activity (NEA) and denitrification enzyme activity (DEA) rates along an altitudinal gradient with the effects of climatic warming using Open Top Chambers (OTCs) in consideration with root dynamics and architecture. This study was conducted at Latnjajaure Field Station (LFS) located in the midalpine region in northern Sweden. LFS is the Swedish field site for the International Tundra Experiment (ITEX), established in 1993. This gives an opportunity to investigate long-term effects of climatic warming by OTCs and an altitudinal gradient (1000m to 1365m), both within a very small geographical range. The OTCs used at LFS increases the soil surface temperature by approximately 1.5ºC whereas air temperatures normally falls with 0.6ºC with every hundred meter of increased altitude. To analyse the NEA and DEA we used an anaerobic incubation technique, based on acetylene inhibition technique, resulting in N2O as the only end product, which then were analysed by gas chromatography. Soil cores were additionally sampled in the OTCs to study the effects of climatic warming on the root system. The specific root length, root length density and root biomass were determined for the different root fractions. The results from NEA and DEA contradict each other. The gradient study show decreased NEA and DEA rates with falling altitude, whereas the warming experiment show a slight non significant increase due to the temperature enhancement by OTCs. The preliminary results from the root sampling indicate that there is a plant community specific response in root architecture, which has an output on root biomass and particularly in the fraction of fine roots, although, climatic warming did not have any significant affect on the root biomass. The fact that altitudinal temperature decline did not reduce NEA and DEA rates might in part be explained of the variables measured here, although they are not conclusive.
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| 3. |
- Björk, Robert G., 1974, et al.
(författare)
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Biocomplexity and biogeochemical cycling in terrestrial ecosystems
- 2008
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Ingår i: 1st Workshop and planning meeting ‘Winter processes in arctic tundra ecosystems’, Longyearbyen, Svalbard, 9-11 June 2008..
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Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract
- The activities concerted within Tellus are aiming at adding essential knowledge to the system scale aspects of fluxes and transformation of carbon (C) in the terrestrial domain. The systems studied are the Arctic, the alpine area of the Scandes, and organic soils in southern Sweden (including both forest and agricultural systems). The main research sites are Skogaryd, in the boreal forest, and Latnjajaure Field Station in the western Abisko Mountains. The overall aim with the present work at Skogaryd is to increase our fundamental understanding of process involved in cycling of C and nitrogen (N) in forest ecosystems, and generate high quality data on C/N cycle from drained forested organic soils using micrometeorological methods, laser and automatic chambers techniques. Skogaryd is also included in NitroEurope, an EU project focusing on modelling and up-scaling of greenhouse gas fluxes, and is incorporated in two interdisciplinary research centres at University of Gothenburg, Tellus (http://www.tellus.science.gu.se/english/) and Gothenburg Atmospheric Science Centre (GAC; http://www.chalmers.se/gmv/gac-en/). Furthermore, our research group has been instrumental in the establishment of the International Tundra Experiment (ITEX) network by Professor Ulf Molau who chaired ITEX 1992-1996 and Latnjajaure Field Station have been a master site within the network since 1992. As an outcome, much of the research in Latnjajaure during the 90ies was focused plant responsiveness to global change. However, since 2002 the research has expanded and projects are now running that are dealing with snow-vegetation-soil interactions. Recently, a collaboration with Dr. Elisabeth J. Cooper, of the University Centre in Svalbard (UNIS), and Prof. Bo Elberling, University of Copenhagen, was initiated focusing on winter soil respiration and comparing sub-arctic and high-arctic trace gas fluxes. Currently, we have these projects running in tundra ecosystems: 1.Climate-related changes in tundra ecosystems – An IPY project (PIs Björk and Molau with others). 2.Temporal pattern of CO2, CH4 and N2O fluxes and soil microbial structure in snow-covered ecosystems (PIs Björk, Elberling, Klemedtsson, and Cooper). 3.The responsiveness of tundra ecosystems to warming: linking above- and below-ground components (PI Björk). There is also a project under evaluation by the Swedish Research Council, which are entitled ‘The fate of carbon in high-arctic tundra ecosystems under changing snow cover conditions’. The research questions that we want to address within this winter ecology network are 1) to increase our understand of the C dynamics (thus explicitly linking ecosystem C sequestration pattern and soil microbial dynamics) in tundra ecosystems, and, particularly, (2) to improve the winter resolution of carbon dioxide (and other greenhouse gases) effluxes to the atmosphere. Furthermore, (3) to understand the influence of changing snow cover, both in depth and duration, for the C dynamics in tundra ecosystems.
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| 4. |
- Björk, Robert G., 1974, et al.
(författare)
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Contrasting effects of wood ash application on microbial community structure, biomass and processes in drained forested peatlands
- 2010
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Ingår i: FEMS Microbiology Ecology. - : Oxford University Press (OUP). - 1574-6941 .- 0168-6496. ; 73:3, s. 550-562
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Tidskriftsartikel (refereegranskat)abstract
- The effects of wood ash application on soil microbial processes were investigated in three drained forested peatlands, which differed in nutrient status and time since application. Measured variables included concentrations of soil elements and phospholipid fatty acids (PLFAs), net nitrogen mineralisation, nitrification and denitrification enzyme activity, potential methane oxidation, methane production and microbial respiration kinetics. Wood ash application had a considerable influence on soil element concentrations. This mirrored a decrease in the majority of the microbial biomarkers by more than one-third in the two oligotrophic peatlands, although microbial community composition was not altered. The decreases in PLFAs coincided with reduced net ammonification and net nitrogen mineralisation. Other measured variables did not change systematically as a result of wood ash application. No significant changes in microbial biomass or processes were found in the mesotrophic peatland, possibly because too little time (1 year) had elapsed since the wood ash application. This study suggests that oligotrophic peatlands can be substantially affected by wood ash for a period of at least four years after application. However, within 25 years of the wood ash application, the microbial biomass seemed to have recovered or adapted to enhanced element concentrations in the soil.
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| 5. |
- Björk, Robert G., 1974, et al.
(författare)
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Effect of reduced below-ground C sequestration on greenhouse gas fluxes within dry tundra ecosystems along an altitudinal gradient
- 2008
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Ingår i: Mountain soils under a changing climate and land-use, Birmensdorf, Switzerland, 6–8 March 2008..
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Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract
- It has been suggested that global climate change will have a great impact on arctic and alpine areas, affecting the carbon and nitrogen dynamics in these ecosystems. Temperature are widely thought to be the main limiting factors for plants and microorganisms in these tundra ecosystem, and warming the soil in high latitude tundra have been shown to change trace gas (CO2, CH4, and N2O) exchange rates and increase N availability. However, little attention has been paid, to date, to variations in trace gas fluxes with altitude, although it is a key determinant of temperature and should therefore be strongly correlated with these fluxes if temperature is the main variable affecting these processes. The objectives of this study were, therefore, to measure growing season variation in carbon dioxide (CO2), methane (CH4), and nitrous oxide (N2O) fluxes from heath plant communities along an altitudinal gradient ranging from mid alpine (~950 m a.s.l) to high alpine (~1,365 m a.s.l) zones within an alpine tundra landscape in the subarctic region of Sweden. Furthermore, by reducing the C supply to the root system and mycorrhizal fungi (achieved by clipping the above-ground plant parts) we wanted to decrease the rhizosphere priming effect and thereby change the C sequestration pattern within the ecosystem to be able to separate heterotrophic and root respiration. The study was conducted on the slopes of Mt. Latnjatjårro (1,447 m; 68°21’N, 18°31’E), near Latnjajaure Field Station, 16 km west of Abisko in Northern Sweden. Flux measurements of CO2 were analysed using a portable infra red gas analyser (IRGA) based on the SBA-4 OEM CO2 Analyzer (PP System). Fluxes of CH4 and N2O were sampled using a closed chamber system, where chambers were placed at collars, which were gently pressed into the ground. Air from the chamber was circulated into a headspace bottle and analysed by gas chromatograph. A two-step incubation technique was also used to determine Nitrification Enzyme Activity (NEA) for analysing nitrification in acid soils with low activities, and for Denitrification Enzyme Activity (DEA) an anaerobic incubation technique, based on acetylene inhibition of the N2O-reductase, was used. Preliminary, our results show a decrease in average growing season CO2 efflux with altitude, but not constantly, and although soil temperature in general decreased with altitude there were no perfect fit between soil temperature and average growing season CO2 efflux. Furthermore, the clipping of the above-ground plant parts reduced the CO2 efflux at all altitudes, except at 1,225 m a.s.l., and in August the reduction in CO2 efflux was largest at 950 m a.s.l. (231 mg CO2 m-2 h-1) and decreased with altitude (to 33 mg CO2 m-2 h-1 at 1,365 m a.s.l). However, the proportion of the reduced CO2 efflux, corresponding to root respiration, was relatively constant with altitude (28-43% of total respiration), except at 1,365 m a.s.l where the root respiration only contributed with 12%. The fluxes of CH4 and N2O was very low with poor resolution of the fluxes due to many samples had a flux lower than the limit of detection for the gas chromatograph, thus no particularly pattern was discerned. However, to try to improve the resolution along the altitudinal gradient the NEA and DEA was used and give a potential measure on the nitrification and denitrification rates, which goes back to the actual populations of nitrifiers and denitrifiers in the soil. The results shows that there were a substantial increases with altitude in the activities of nitrifying and denitrifying microbes, this is contrary to expectations and the average growing season CO2 efflux if the decline in mean annual temperature with altitude is the main driver for nitrification and denitrification. Thus, our results are just indicative for the complex interaction that may occur along altitudinal gradients. But, clearly, there is a need for further studies to assess the effects of altitude and temperature on carbon and nitrogen dynamics in high alpine and arctic ecosystems across wide altitudinal ranges.
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| 6. |
- Björk, Robert G., 1974, et al.
(författare)
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Effect of reduced below-ground C sequestration on greenhouse gas fluxes within dry tundra ecosystems along an altitudinal gradient
- 2008
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Ingår i: The 15th ITEX workshop, Reykjavik, Iceland, 9–12 October 2008..
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Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract
- It has been suggested that global climate change will have a great impact on arctic and alpine areas, affecting the carbon and nitrogen dynamics in these ecosystems. Temperature is widely thought to be the main limiting factor for plants and microorganisms in these tundra ecosystems, and warming the soil in high latitude tundra has been shown to change trace gas (CO2, CH4, and N2O) exchange rates and increase N availability. However, little attention has been paid, to date, to variations in trace gas fluxes with altitude, although altitude is a key determinant of temperature and should therefore be strongly correlated with these fluxes if temperature is a major variable affecting these processes. The objectives of this study were, therefore, to measure growing season variation in carbon dioxide (CO2), methane (CH4), and nitrous oxide (N2O) fluxes from heath plant communities along an altitudinal gradient ranging from mid alpine (~950 m a.s.l) to high alpine (~1,365 m a.s.l) zones within an alpine tundra landscape in the subarctic region of Sweden. Furthermore, by reducing the C supply to the root system and mycorrhizal fungi (achieved by clipping the above-ground plant parts) we wanted to decrease the rhizosphere priming effect and thereby change the C sequestration pattern within the ecosystem to be able to separate heterotrophic and autotrophic respiration. The study was conducted on the slopes of Mt. Latnjatjårro (1,447 m a.s.l.; 68°21’N, 18°31’E), near Latnjajaure Field Station, 16 km west of Abisko in Northern Sweden. Flux measurements of CO2 were analysed using a portable infra red gas analyser (IRGA) based on the SBA-4 OEM CO2 Analyzer (PP System). Fluxes of CH4 and N2O were sampled using a closed chamber system, where chambers were placed on collars, which were gently pressed into the ground. Air from the chamber was circulated into a headspace bottle and analysed by gas chromatograph. A two-step incubation technique was also used to determine Nitrification Enzyme Activity (NEA) for analysing nitrification in acid soils with low activities, and for Denitrification Enzyme Activity (DEA) an anaerobic incubation technique, based on acetylene inhibition of the N2O-reductase, was used. Our results show a decrease in average growing season CO2 efflux with altitude, but not consistently, and although soil temperature in general decreased with altitude there was only a loose association between soil temperature and average growing season CO2 efflux. Furthermore, the clipping of the above-ground plant parts reduced the CO2 efflux at all altitudes, except at 1,225 m a.s.l., and in August the reduction in CO2 efflux was largest at 950 m a.s.l. (231 mg CO2 m-2 h-1) and decreased with altitude (to 33 mg CO2 m-2 h-1 at 1,365 m a.s.l.). However, the proportion of the reduced CO2 efflux, corresponding to autotrophic respiration, was relatively constant with altitude (28-43% of total respiration), except at 1,365 m a.s.l. where the autotrophic respiration only contributed 12%. The fluxes of CH4 and N2O were very low, and resolution was constrained by the large number of samples with apparent fluxes below the limit of detection for the gas chromatograph, thus no particular pattern could be identified. However, to try to improve the resolution along the altitudinal gradient, the NEA and DEA were used, and give a potential measure of the nitrification and denitrification rates, which goes back to the actual populations of nitrifiers and denitrifiers in the soil. The results show that there were a substantial increases with altitude in the activities of nitrifying and denitrifying microbes; this is contrary to expectations and the average growing season CO2 efflux if the decline in mean annual temperature with altitude is the main driver for nitrification and denitrification. Thus, our results are indicative of the complex interaction that may occur along altitudinal gradients. But, clearly, there is a need for further studies to assess the effects of altitude and temperature on carbon and nitrogen dynamics in high alpine and arctic ecosystems across wide altitudinal ranges.
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| 7. |
- Björk, Robert G., 1974, et al.
(författare)
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Linkages between N turnover and plant community structure in a tundra landscape
- 2007
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Ingår i: Plant and Soil. - : Springer Science and Business Media LLC. - 0032-079X .- 1573-5036. ; 294:1-2, s. 247-261
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Tidskriftsartikel (refereegranskat)abstract
- The spatial distribution of organic soil nitrogen (N) in alpine tundra was studied along a natural environmental gradient, covering five plant communities, at the Latnjajaure Field Station, northern Swedish Lapland. The five communities (mesic meadow, meadow snowbed, dry heath, mesic heath, and heath snowbed) are the dominant types in this region and are differentiated by soil pH. Net N mineralization, net ammonification, and net nitrification were measured using 40-day laboratory incubations based on extractable NH4+ and NO3-. Nitrification enzyme activity (NEA), denitrification enzyme activity (DEA), amino acid concentrations, and microbial respiration were measured for soils from each plant community. The results show that net N mineralization rates were more than three times higher in the meadow ecosystems (mesic meadow 0.7 mu g N g(-1) OM day(-1) and meadow snowbed 0.6 mu g N g(-1) OM day(-1)) than the heath ecosystems (dry heath 0.2 mu g N g(-1) OM day(-1), mesic heath 0.1 mu g N g(-1) OM day(-1) and heath snowbed 0.2 mu g N g(-1) OM day(-1)). The net N mineralization rates were negatively correlated to organic soil C/N ratio (r = -0.652, P < 0.001) and positively correlated to soil pH (r = 0.701, P < 0.001). Net nitrification, inorganic N concentrations, and NEA rates also differed between plant communities; the values for the mesic meadow were at least four times higher than the other plant communities, and the snowbeds formed an intermediate group. Moreover, the results show a different pattern of distribution for individual amino acids across the plant communities, with snowbeds tending to have the highest amino acid N concentrations. The differences between plant communities along this natural gradient also illustrate variations between the dominant mycorrhizal associations in facilitating N capture by the characteristic functional groups of plants.
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| 8. |
- Björk, Robert G., 1974, et al.
(författare)
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Long-term warming effects on carbon and nitrogen dynamics in tundra soils
- 2012
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Ingår i: 20th Anniversary ITEX Workshop, El Paso, USA, 17–21 January 2012.
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Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract
- During IPY 2008 we used the ITEX experiment in Latnjajaure (northern Sweden), established during the early years of the program, to investigate long-term warming effects on ecosystem respiration (ER), carbon (C) and nitrogen (N) pool (including d13C and d15N), soil organic C (SOC) chemical composition, and N mineralization among plant communities. After 12 to 15 years of open top chamber (OTC) treatment no statistical effect was found on the soil temperature (10 cm soil depth), although the was an overall increase in all OTC by +0.2°C. However, the soil moisture decreased significantly by 3-14%, depending on plant community, in the OTCs compared to ambient conditions. Preliminary, there was a 19-61% non-significant increase in annual growing season ER in the OTC compared to the ambient plots over the growing season. The were distinct differences in the SOM functional composition among plant communities with c 10% more O-alkyls stored in tussock tundra than in dry meadow. The OTCs did not consistently alter the SOM composition among the vegetation types but clearly showed a trend for reduced aliphatic and O-alkyl C in the OTCs suggesting increased decomposition (or reduced inputs) of these compounds. Thus, the non-significantly higher ER may in some communities be of plant origin linked to greater plant biomass in the OTCs, and in other (e.g. tussock tundra) from increased decomposition rates. In conclusion, this study showed that 12-15 years of OTC treatment had a modest effects impact C and N dynamics in tundra soils specific to distinct plant communities.
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| 9. |
- Björk, Robert G., 1974, et al.
(författare)
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Long-term warming effects on root morphology, root mass distribution, and microbial activity in two dry tundra plant communities in northern Sweden
- 2007
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Ingår i: New Phytologist. - : Wiley. - 0028-646X .- 1469-8137. ; 176:4, s. 862-873
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Tidskriftsartikel (refereegranskat)abstract
- center dot Effects of warming on root morphology, root mass distribution and microbial activity were studied in organic and mineral soil layers in two alpine ecosystems over > 10 yr, using open-top chambers, in Swedish Lapland. center dot Root mass was estimated using soil cores. Washed roots were scanned and sorted into 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 g DM-1), specific root area (SRA; m(2) kg DM-1), and number of root tips m(-2) were determined. Nitrification (NEA) and denitrification enzyme activity (DEA) in the top 10 cm of soil were measured. center dot Soil warming shifted the rooting zone towards the upper soil organic layer in both plant communities. In the dry heath, warming increased SRL and SRA of the finest roots in both soil layers, whereas the dry meadow was unaffected. Neither NEA nor DEA exhibited differences attributable to warming. center dot Tundra plants may respond to climate change by altering their root morphology and mass while microbial activity may be unaffected. This suggests that carbon may be incorporated in tundra soils partly as a result of increases in the mass of the finer roots if temperatures rise.
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| 10. |
- Björk, Robert G., 1974, et al.
(författare)
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Nitrification and Denitrification Enzyme Activity: a successful tool in Arctic and Alpine soil ecology
- 2007
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Ingår i: The 14th ITEX workshop, Falls Creek, Victoria, Australia, 2–6 February 2007..
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Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract
- Nitrogen is consideration to be a limiting factor for plants and microbes in arctic and alpine ecosystems and the rates of mineralization, nitrification, and denitrifi¬cation are known to be generally low. However, Climate Change is expected to alter the nitrogen availability and dynamics and, as a consequence, affect plant community composition and production. The general consensus today is that increased temperature will lead to greater microbial activity and more plant-available nitrogen. Nevertheless, nitrification and denitrification are restricted by a number of environmental factors such as low tem¬perature and low pH. The C/N ratio and the water content of the soils also play an important role in determining the rates of nitrification and denitrification. Since 2002 microbial studies has been undertaken at Latnjajure, and comprise several microbial techniques, e.g. Nitrification Enzyme Activity (NEA), Denitrification Enzyme Activity (DEA), Phospholipid fatty acid analysis (PFLA), and Temperature Gradient Gel Electrophoresis (TGGE). These studies focuses on the interaction between plants and microbes along natural environmental gradients, both within plant communities and within the landscape, but also entails the OTCs used in the ITEX studies at Latnjajaure. Here we present the techniques NEA and DEA and give some brief results from how these have been successfully applied at Latnjajaure. In ecosystems with low nitrification activity, small amounts of NO3-/NO2- will be formed and it is thus difficult to measure low fluxes. However, NO3-/NO2- can be converted to N2O and then analysed by gas chromatography, whereby the detection limit is increased at least 1000 times compared to the spectroscopical technique. These techniques are referred to Nitrification (NEA) and Denitrification Enzyme Activity (DEA) and give a potential measure on the nitrification and denitrification rates, which goes back to the actual populations of nitrifiers and denitrifiers in the soil. For instance, NEA has been proved to better correlate with extractable NH4+ concentration than net nitrification does, and still after twelve weeks show a strong correlation with the initial extractable NH4+ concentration. Therefore, these variables, in particular NEA, have the advantage of being a much more stable variable than, for instance, extractable N concentrations and net nitrification, and NEA and DEA are therefore suitable when working in fringe environments with restricted logistics like the Latnjajaure catchment. NEA is measured using a two-step incubation technique; first by incubate the soil with a nutrient solution for 24 hours in darkness, at room temperature on a rotary shaker. Sub-samples are then withdrawn after a specified time schedule. The second step allows NO3- to be reduced to N2O by adding a modified denitrifying bacterium, Pseudomonas chlororaphis ATCC 43928, together with a carbon source. This strain of bacteria lacks the enzyme to reduce N2O to N2. The samples are then again incubated in darkness, at room temperature for 24 hours, and analysed by gas chromatography. This method was first used by Lensi et al. (1985, 1986), to study nitrification potentials in forest soils. Furthermore, the method has been developed for soils with low pH and small amounts of NO3- and the analysis makes the quantification without interference of organic matter, which makes it suitable for arctic and alpine ecosystems. To analyse DEA an anaerobic incubation technique is used, based on acetylene inhibition of the N2O-reductase resulting in N2O as the only end product. The soil sample is evacuated and flushed with N2. Thereafter acetylene is inserted to a final acetylene concentration of 10%, and the samples are shaken continuously and gas samples are withdrawn after a specified time schedule, which is then analysed by gas chromatography. This provides an estimate of the maximum concentration of functional denitrifying enzymes in the soil. Denitrifiers, in contrast to nitrifiers, are heterotrophs and can switch from using NO3- as an alternative electron acceptor to O2 under aerobic conditions. This makes other factors in the soil important determinants of DEA, e.g. availability of oxygen and C. Hence, the presence of denitrifiers is rarely a limitation for denitrification and they usually make up a reasonably large fraction of the soil bacteria. At Latnjajaure NEA shows a larger differentiation across plant communities than DEA. However, the spatial variability in the landscape, at the meso-scale, was in the same range in both variables and increased with altitude from 1000 to 1365 m a.s.l, particularly in heath plant communities. This result suggests that the decrease in mean annual temperature with altitude (0.6ºC with every one hundred meters) did not reduce nitrification and denitrification rates, as one might have expected. None of the other variable studied could explain the altitudinal increase in all cases, and the factors controlling the nitrification and denitrification rates seem to vary with the vegetation type. Furthermore, neither NEA nor DEA exhibited any changes between the ambient and warmed plots in the warming experiments. However, the warming experiment in the dry heath exhibited a change in root morphology via increased specific root length (SRL; m gDM-1) and specific root area (SRA; m2 kgDM -1). As both heterotrophic microbes and plants out-compete nitrifiers for NH4+, a change in root morphology, as seen in the warming experiment, may also explain the increased activity of nitrifying and denitrifying microbes with altitude.
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