Reindeer influence on ecosystem processes in the tundra

Reindeer have been recorded to increase nutrient cycling rate and primary production in studies from fences almost 40 years old that separate areas with different grazing regimes in northern Fennoscandia. To further understand the mechanism behind the effects of herbivores on primary production, we measured the size of the major C and N pools, soil temperature, litter decomposition rate and N mineralization rate in lightly, moderately and heavily grazed areas along two of these fences.
Plant N found in new biomass, indicative of plant N assimilation, was significantly higher in moderately and heavily grazed areas than in lightly grazed areas, which corresponded to a decreased amount of N in old plant parts. The amount of N found in plant litter or organic soil layer did not differ between the grazing treatments. Together with soil N concentrations and litter decomposition rates, soil temperatures were significantly higher in moderately and heavily grazed areas.
We conclude that the changes in soil temperature are important for the litter decomposition rate and thus on the nutrient availability for plant uptake. However, the changes in plant community composition appear to be more important for the altered N pools and thus the enhanced primary production. The results provide some support for the keystone herbivore hypothesis, which states that intensive grazing can promote a transition from moss-rich tundra heath to productive grasslands. Grazing altered N fluxes and pools, but the total N pools were similar in all grazing treatments. Our study thus indicates that grazing can increase the primary production through enhancing the soil nutrient cycling rate, even in a long term perspective on an ecological timescale.     

Structure of microbial communities of peat soils in two bogs in Siberian tundra and forest zones

The structure and functional activity of microbial complexes of a forest oligo-mesotrophic subshrub- grass-moss bog (OMB, Central Evenkiya) and a subshrub-sedge bog in the polygonal tundra (PB, Lena River Delta Samoylovsky Island) was studied. Soil of the forest bog (OMB) differed from that of the polygonal tundra bog (PB) in higher productivity (C_org, N_total, P, and K reserves), higher biomass of aerobic chemoorganotrophs (2.0 to 2.6 times), and twice the level of available organic matter. The contribution of microorganisms to the carbon pool was different, with the share of C_mic in C_org 1.4 to 2.5 times higher in PB compared to OMB. Qualitative composition of the methane cycle microorganisms in PB and OMB soils differed significantly. Methanogenic archaea (Euryarchaeota) in the shrub-sedge PB of tundra were more numerous and diverse than in the oligo-mesotrophic bog (OMB) and belonged to six families (Methanomassiliicoccaceae, Methanoregulaceae, Methanobacteriaceae, Methanomicrobiaceaee, Methanosarcinaceae, and Methanotrichaceae), while members of only four families (Methanosarcinacea, Methanobacteriaceae, Methanotrichaceae, and Methanomassiliicoccaceae) were revealed in OMB. In both bogs, methane-oxidizing bacteria belonged to Alphaproteobacteria (II) and Gammaproteobacteria (I). Methanotroph diversity was higher in OMB than in PB. Microbial communities of PB soils had higher potential activity of methanogenesis and methanotrophy compared to those of OMB. Methanogenic and methanotrophic activities in PB were 20 and 2.3 times higher, respectively, than in OMB.     

Links between plant community composition,soil organic matter quality and microbial communities in contrasting tundra habitats

Plant communities, soil organic matter and microbial communities are predicted to be interlinked and to exhibit concordant patterns along major environmental gradients. We investigated the relationships between plant functional type composition, soil organic matter quality and decomposer community composition, and how these are related to major environmental variation in non-acid and acid soils derived from calcareous versus siliceous bedrocks, respectively. We analysed vegetation, organic matter and microbial community compositions from five non-acidic and five acidic heath sites in alpine tundra in northern Europe. Sequential organic matter fractionation was used to characterize organic matter quality and phospholipid fatty acid analysis to detect major variation in decomposer communities. Non-acidic and acidic heaths differed substantially in vegetation composition, and these disparities were associated with congruent shifts in soil organic matter and microbial communities. A high proportion of forbs in the vegetation was positively associated with low C:N and high soluble N:phenolics ratios in soil organic matter, and a high proportion of bacteria in the microbial community. On the contrary, dwarf shrub-rich vegetation was associated with high C:N and low soluble N:phenolics ratios, and a high proportion of fungi in the microbial community. Our study demonstrates a strong link between the plant community composition, soil organic matter quality, and microbial community composition, and that differences in one compartment are paralleled by changes in others. Variation in the forb-shrub gradient of vegetation may largely dictate variations in the chemical quality of organic matter and decomposer communities in tundra ecosystems. Soil pH, through its direct and indirect effects on plant and microbial communities, seems to function as an ultimate environmental driver that gives rise to and amplifies the interactions between above- and belowground systems. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The effect and extent of heavy grazing by reindeer in oligotrophic pine heaths in northeastern Fennoscandia

Thirty and fifty years old exclosures established in northeastern Fennoscandia in lichen-rich oligotrophic pine Pinus sylvestris forests on podzolised soil were used to study the effect of reindeer grazing on pine fine roots, microbial activity, and on bryophyte, dwarf shrub and lichen biomasses. There were significantly less lichens, especially Cladina stellaris , at grazed than at ungrazed sites. Coverage of other lichens like C. arbuscula and C. rangiferina and bryophytes, especially Dicranum spp., benefitted from grazing. The biomass of vascular plants, mainly Calluna vulgaris, Empetrum nigrum and Vaccinium vitis-idaea , was reduced at grazed sites, although their coverage was not influenced. Microbial activity was significantly lower at grazed sites. The influence of grazing is most obviously mediated by reduced soil moisture during dry periods at grazed sites. Fine root parameters (per soil and stem volumes) were lower at grazed sites (pPCA = 0.072), the first principal component consisting of a number of fine root tips, length and weight. Grazing decreased all exchangeable nutrients by 30–60% in organic layer. Based on PCA the decrease was significant for exchangeable nutrients, although of individual elements only P and S showed statistically significant difference. The extent of heavy grazing in northeastern Fennoscandia coniferous forest was revealed by remote sensing. It revealed extensive area in which reindeer lichens are reduced in northeastern Finland. The Finnish-Russian border can be clearly distinguished in the satellite image composite.     

Long- and short-term effects of reindeer grazing on tundra wetland vegetation

We studied long-term (50 years) and short-term (4 years) effects of summer grazing of reindeer on subarctic tundra wetland vegetation. The long-term effects of summer grazing were studied by comparing vegetation on Finnish and Norwegian sides of the fence line separating reindeer grazing regimes. The Finnish side was intensively grazed and trampled throughout the year, whereas the Norwegian side was grazed in winter. Experimental fences were erected to examine short-term effects of grazing exclusion. Both in the long- and short-term, summer grazing decreased the height of Salix lapponum whereas the short-term effects on willow cover were less clear than the long-term effects. In contrast, Carex spp. benefited from grazing. Long-term grazing had little effect on total bryophyte cover. Grazing had negligible effects on the nutrient content of leaves of S. lapponum and Eriophorum angustifolium. We conclude that tundra wetlands can withstand moderately high grazing pressure sustained over several decades.     

Effect of wood ash and nitrogen fertilization on soil chemical properties, soil microbial processes, and stand growth in two coniferous stands in Finland

The aim of our study was to investigate long-term effects of wood ash fertilization, given together with nitrogen, on soil chemical properties, soil microbiological processes related to C and N cycling, and tree growth. The study was carried out in a 31-year-old Scots pine stand and in a 45-year-old Norway spruce stand 15 years after application. The treatments were (1) a control with no ash or nutrient addition, (2) wood ash + N (WAN), and (3) a stand-specific fertilization (SSF) formulated on the basis of analyses carried out on needle and soil samples taken from the stand. The SSF treatments included N, Cu and B, and in the spruce stand also P. WAN decreased acidity and increased the extractable Ca, Mg and P concentrations in the organic layer in both stands, but SSF had no effect. The microbial processes reacted more strongly to the treatments in the pine stand, whereas the growth response, although only relatively slight during the third 5-year period after fertilization, was detected only in the spruce stand. WAN increased the NH₄-N concentrations in the organic layer compared to the control and SSF treatments on both sites. In the pine stand, amount of N in microbial biomass and both the C and net N mineralization rates were significantly higher in the WAN treatment than in the SSF treatment. On both sites net nitrification was negligible in all treatments. Soil microbial biomass, microbial respiration and N availability have been used as indices for assessing the biological activity and health of soil, and these parameters either increased or were not affected by the WAN treatment. Hence, with regard to these parameters there are justifiable grounds for applying wood ash.     

Acidobacteria in microbial communities of the bog and tundra lichens

Bacterial communities of the lichens from a Sphagnum bog (Karelia) and tundra (Vorkuta oblast) were investigated. Members of the phylum Acidobacteria were numerous in the thallus of living and decaying lichens (3.8 × 10⁸ cells/g), constituting 6 to 32% of the total bacterial number. Pure cultures of acidobacteria were isolated from the samples of living and decaying lichen thallus. Ten of them were identified and classified as members of subgroup 1 of the Acidobacteria. The hydrolytic activity of two strains isolated from the living and decomposing zones of the thallus was investigated. They were capable of growth on xylan, starch, pectin, laminarin, and lichen extract. Acidobacteria were shown to be a stable and numerous component of microbial communities of the bog and tundra lichens.     

Non-parallel changes in soil microbial carbon and nitrogen dynamics due to reindeer grazing in northern boreal forests

Reindeer grazing has a considerable influence on mineralization processes in northern Fennoscandian boreal forests, but the mechanisms underlying the observed differences between grazed and ungrazed areas are not well understood. We studied the below-ground impacts of reindeer grazing by comparing the carbon and nitrogen mineralization rates inside and outside long-term fenced reindeer exclosure areas in five oligotrophic, lichen-dominated and five mesotrophic, dwarf-shrub dominated forests. The soil C mineralization rates and microbial metabolic activity (qCO₂) were significantly lower in the grazed than the ungrazed areas in both oligotrophic and mesotrophic forests. The reductions occurred irrespective of the impact on soil moisture. We conclude that reindeer grazing causes a reduction in the supply of labile C substrates to microbes, resulting in reduced organic matter decomposition rates through changes in the activity of the microbial biomass. Simultaneously, grazing had no consistent effect on the microbial N dynamics, but the impact ranged from no change to increased or decreased in N mineralization rates at the different study sites. The impact of grazing on the N mineralization potential thus seems to be site-specific and uncoupled from the impact of grazing on soil C mineralization. Reciprocal transplant incubations showed no interactions between N mineralization rates and the reindeer-mediated impact on the soil microclimate. We suggest that plant root damage due to trampling by reindeer may be an important mechanism for the deceleration of soil C cycling. In some cases, however, the impact of grazing on the soil active N pool may be strong enough to outweigh the reduction in soil organic matter decomposition, and by these means uncouple soil N dynamics from soil C quality.     

Effects of grazing and cultivation on soil patterns and processes in the Paulshoek area of Namaqualand

Soils were sampled at three different sites within the Paulshoek area of the Leliefontein District in Namaqualand, South Africa. At one site the effect of heavy grazing on soil characteristics under shrubs and from open positions was compared to a more lightly grazed area. At this site a diverse shrub community was mostly replaced by a single species, Galenia africana, when overgrazed. In another area the effects of cultivation on soil properties was investigated. At the third site the effect of Galenia shrub size on soil properties was measured.Soils from the Paulshoek area of Namaqualand are generally infertile and very low in arbuscular mycorrhizal propagules. Patterns of nitrogen distribution at the landscape level are a result of nutrient enrichment associated with perennial shrubs. Arbuscular mycorrhizal infectivity, soil moisture and pH are also affected by shrubs. Only species capable of forming arbuscular mycorrhizas support populations of mycorrhizal propagules in their rhizosphere. Differences in plant available phosphorus under some shrubs indicate that processes associated with nutrient cycling are also affected by shrubs. Galenia tended to deplete soil moisture and increase soil pH more than other shrubs.Small stock production results in a depletion of soil nutrients at the landscape scale since larger areas of low nutrient soil develop when shrub density decreases following heavy grazing. Heavy grazing also affects soil properties indirectly through a change in shrub composition. Cultivation resulted in a decrease in soil nitrogen and organic matter, a loss which was still detectable after 20 years of fallow. Zones of nutrient enrichment and higher pH were found where Galenia established in the old field. When soil was sampled under different size Galenia shrubs a positive correlation was found between shrub diameter and nitrogen, available phosphorus and pH indicating that Galenia was capable of establishing patches in the soil with different properties rather than selecting such areas for establishment. Thus, Galenia is able to maintain soil patterns typical of Succulent Karoo although it also establishes conditions which probably ensure its success. Individual shrub species also affect soil differently with respect to some components such as mycorrhizas.     

Impact of grazing on soil carbon and microbial biomass in typical steppe and desert steppe of Inner Mongolia

The potential of grazing lands to sequester carbon must be understood to develop effective soil conservation measures and sustain livestock production. Our objective was to evaluate the effects of grazing on soil organic carbon (SOC), total nitrogen (TN), microbial biomass carbon (MBC) in Typical steppe and Desert steppe ecosystems, which are both important grassland resources for animal grazing and ecological conservation in China, and to derive region-specific soil C changes associated with different stocking rates (ungrazed, UG; lightly grazed, LG; moderately grazed, MG; heavily grazed, HG). This study substantiated that significant higher SOC, TN and MBC appeared with the treatment of LG in typical steppe. From 2004 to 2010, grazing treatments increased soil carbon storage in desert steppe, which was partly due to the grazing history. The higher MBC concentration and MBC/SOC suggest a great potential for carbon sequestration in the desert steppe ecosystem. The greater MBC in desert steppe than typical steppe was mainly the result of higher precipitation and temperature, instead of soil substrate. The change of MBC and the strong positive relationships between MBC and SOC indicated that MBC in the soil was a sensitive index to indicate the dynamics of soil organic carbon in both steppes in Inner Mongolia of China.     

Phosphorus availability and microbial respiration across different tundra vegetation types

Phosphorus (P) is an important nutrient in tundra ecosystems that co-limits or in some cases limits primary production. The availability of P is largely driven by soil characteristics, e.g., pH, organic carbon, and abundance of P-sorbing elements such as aluminium (Al) or iron (Fe). We tested how vegetation and soil properties relate to P availability across different tundra vegetation types. The different soil P fractions in the organic horizon were measured and plant foliar nitrogen (N) to P ratio and a plant bioassay was used as indicators of plant nutrient status. Microbial bioassays were used to study microbial respiration kinetics and in response to carbon, N, and P amendments. The distribution of P fractions differed significantly across vegetation types; labile fractions of P were less abundant in meadow sites compared to heath sites. Calcium-phosphates seemed to be an important P-fraction in meadows, but were only found in lower concentrations in the heath. There were only small differences in NaOH–extractable P between the vegetation types and this correlated with the distribution of oxalate-extractable Al. Plant N:P ratios and the plant bioassay indicated decreasing P availability from dry heath to mesic heath to mesic meadow. The microbial bioassay suggests that the heterotrophic microbial community is C-limited with N as a secondary limiting nutrient although there were indications that microbial P availability was lower in the meadow sites. Overall, we suggest that the observed variations in soil P across vegetation types are affecting both plant and microbial function although the differences seem to be relatively small.     

The effects of chronic nitrogen fertilization on alpine tundra soil microbial communities: implications for carbon and nitrogen cycling

Many studies have shown that changes in nitrogen (N) availability affect primary productivity in a variety of terrestrial systems, but less is known about the effects of the changing N cycle on soil organic matter (SOM) decomposition. We used a variety of techniques to examine the effects of chronic N amendments on SOM chemistry and microbial community structure and function in an alpine tundra soil. We collected surface soil (0-5 cm) samples from five control and five long-term N-amended plots established and maintained at the Niwot Ridge Long-term Ecological Research (LTER) site. Samples were bulked by treatment and all analyses were conducted on composite samples. The fungal community shifted in response to N amendments, with a decrease in the relative abundance of basidiomycetes. Bacterial community composition also shifted in the fertilized soil, with increases in the relative abundance of sequences related to the Bacteroidetes and Gemmatimonadetes, and decreases in the relative abundance of the Verrucomicrobia. We did not uncover any bacterial sequences that were closely related to known nitrifiers in either soil, but sequences related to archaeal nitrifiers were found in control soils. The ratio of fungi to bacteria did not change in the N-amended soils, but the ratio of archaea to bacteria dropped from 20% to less than 1% in the N-amended plots. Comparisons of aliphatic and aromatic carbon compounds, two broad categories of soil carbon compounds, revealed no between treatment differences. However, G-lignins were found in higher relative abundance in the fertilized soils, while proteins were detected in lower relative abundance. Finally, the activities of two soil enzymes involved in N cycling changed in response to chronic N amendments. These results suggest that chronic N fertilization induces significant shifts in soil carbon dynamics that correspond to shifts in microbial community structure and function.     

Response of soil microbial activity to grazing, nitrogen deposition, and exotic cover in a serpentine grassland

Background and aims

Exotic species, nitrogen (N) deposition, and grazing are major drivers of change in grasslands. However little is known about the interactive effects of these factors on below-ground microbial communities.

Methods

We simulated realistic N deposition increases with low-level fertilization and manipulated grazing with fencing in a split-plot experiment in California’s largest serpentine grassland. We also monitored grazing intensity using camera traps and measured total available N to assess grazing and nutrient enrichment effects on microbial extracellular enzyme activity (EEA), microbial N mineralization, and respiration rates in soil.

Results

Continuous measures of grazing intensity and N availability showed that increased grazing and N were correlated with increased microbial activity and were stronger predictors than the categorical grazing and fertilization measures. Exotic cover was also generally correlated with increased microbial activity resulting from exotic-driven nutrient cycling alterations. Seasonal effects, on abiotic factors and plant phenology, were also an important factor in EEA with lower activity occurring at peak plant biomass.

Conclusions

In combination with previous studies from this serpentine grassland, our results suggest that grazing intensity and soil N availability may affect the soil microbial community indirectly via effects on exotic cover and associated changes in nutrient cycling while grazing directly impacts soil community function.     

Linking microbial community composition and soil processes in a California annual grassland and mixed-conifer forest

To investigate the potential role of microbial community composition in soil carbon and nitrogen cycling, we transplanted soil cores between a grassland and a conifer ecosystem in the Sierra Nevada California and measured soil process rates (N-mineralization, nitrous oxide and carbondioxide flux, nitrification potential), soil water and temperature, and microbial community parameters (PLFA and substrate utilization profiles) over a 2 year period. Our goal was to assess whether microbial community composition could be related to soil process rates independent of soil temperature and water content. We performed multiple regression analyses using microbial community parameters and soil water and temperature as X-variables and soil process rates and inorganic N concentrations as Y-variables. We found that field soil temperature had the strongest relationship with CO₂ production and soil NH₄⁺ concentration, while microbial community characteristics correlated with N₂O production, nitrification potential, gross N-mineralization, and soil NO₃^– concentration, independent of environmentalcontrollers. We observed a relationship between specific components of the microbial community (as determined by PLFA) and soil processes,particularly processes tightly linked to microbial phylogeny (e.g. nitrification). The most apparent change in microbial community composition in response to the 2 year transplant was a change in relative abundance of fungi (there was only one significant change in PLFA biomarkers for bacteria during 2 years). The relationship between microbial community composition and soil processes suggests that prediction of ecosystem response to environmental change may be improved by recognizing and accounting for changes in microbial community composition and physiological ecology.     

Distinct microbial communities associated with buried soils in the Siberian tundra

Cryoturbation, the burial of topsoil material into deeper soil horizons by repeated freeze–thaw events, is an important storage mechanism for soil organic matter (SOM) in permafrost-affected soils. Besides abiotic conditions, microbial community structure and the accessibility of SOM to the decomposer community are hypothesized to control SOM decomposition and thus have a crucial role in SOM accumulation in buried soils. We surveyed the microbial community structure in cryoturbated soils from nine soil profiles in the northeastern Siberian tundra using high-throughput sequencing and quantification of bacterial, archaeal and fungal marker genes. We found that bacterial abundances in buried topsoils were as high as in unburied topsoils. In contrast, fungal abundances decreased with depth and were significantly lower in buried than in unburied topsoils resulting in remarkably low fungal to bacterial ratios in buried topsoils. Fungal community profiling revealed an associated decrease in presumably ectomycorrhizal (ECM) fungi. The abiotic conditions (low to subzero temperatures, anoxia) and the reduced abundance of fungi likely provide a niche for bacterial, facultative anaerobic decomposers of SOM such as members of the Actinobacteria, which were found in significantly higher relative abundances in buried than in unburied topsoils. Our study expands the knowledge on the microbial community structure in soils of Northern latitude permafrost regions, and attributes the delayed decomposition of SOM in buried soils to specific microbial taxa, and particularly to a decrease in abundance and activity of ECM fungi, and to the extent to which bacterial decomposers are able to act as their functional substitutes.     

Molecular detection and diversity of xylanase genes in alpine tundra soil

Xylan is a major polysaccharide in plant cell walls, and its degradation is mainly conducted by microbial xylanases in nature. To explore the xylanase diversity in the environment, two sets of degenerate primers were designed based on the microbial xylanase sequences in Pfam database of glycosyl hydrolase (GH) family 10 and 11 and were used to amplify objective gene fragments directly from the alpine tundra soil DNA of the Tianshan Mountains, China. Ninety-six distinct GH 10 and 31 GH 11 xylanase gene fragments were retrieved, and most of them have low identities with known sequences in GenBank. Based on phylogenetic analysis, all of the GH 10 xylanase sequences fell into six clusters and were related to xylanases from Actinobacteria, Proteobacteria, Verrucomicrobia, Bacteroidetes, Firmicutes, and Acidobacteria. Three clusters of GH 11 xylanase sequences were established, and two of them were related with enzymes from fungi. These results indicated the diversity of xylanase genes in this cold environment. Four xylanolytic strains were isolated from the soil, and GH 10 xylanase gene fragments were cloned using the same primers. A full-length gene was obtained and expressed in Escherichia coli, and the recombinant enzyme showed some cold-related characteristics. Our study provides an efficient molecular approach to study xylanase in complex environments and casts an insight into the diversity and distribution of xylanases in a cold environment, which is very meaningful to understand their roles in xylan degradation in nature.     

Effects of experimental warming and nitrogen fertilization on soil microbial communities and processes of two subalpine coniferous species in Eastern Tibetan Plateau,China

Aim

This study aimed at predicting how sub-alpine coniferous ecosystems respond to global changes in the Eastern Tibetan Plateau by understanding soil microbial communities and activities, as well as variation in the quality and quantity of soil organic matter.

Methods

An experiment was conducted to examine soil microbial communities and their related soil processes in rhizospheric soil of two coniferous species that were exposed to two levels of temperature (unwarmed and infrared heater warming) and two levels of nitrogen (unfertilized and 25 g N m^?2 a^?1) from April 2007.

Results

Four-year night warming alone slightly affected the phospholipid fatty acid contents of the microbial community. However, the combination of nitrogen addition and soil warming significantly affected soil microbial composition while reducing the biomass of major microbial groups and the activities of most enzymes, especially in Abies faxoniana plots. The combination of warming and nitrogen addition increased soil labile C and N pools in Picea asperata plots and was beneficial for soil recalcitrant C, as well as for labile and total C and N pools in A. faxoniana plots.

Conclusion

Results indicated that future warming will slightly affect soil microbial communities and their related soil processes. However, warming combined with high nitrogen deposition will significantly constrain soil microbial biomass and enzyme activities, consequently increasing soil C and N pools in sub-alpine coniferous forests of this region.