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.     

Fungal endophyte diversity in tundra grasses increases by grazing

Fungal endophytes are species rich and ubiquitous, yet, apart from the genus Epichloë, their ecology is largely unknown. Here we explore how herbivores affect the diversity of fungal endophytes in tundra grasslands. We assess both hyphal morphological and taxonomic diversity in grass individuals.By microscopic examination we identified endophytes to be present in all sampled grass individuals whereas identification to taxonomic units were only achieved in a subset of the individuals using laser micro dissection pressure catapulting and culturing for endophyte isolation. Hyphal morphological diversity was significantly higher in grasses exposed to grazing, along with 45 % more taxonomic units achieved.Our results suggest that grazing is an important mediator of fungal endophyte diversity in tundra grasslands. Furthermore, we suggest laser dissection of stained endophytes as a method for further exploring the ecological role of fungal endophytes.     

土壤氮素转化的关键微生物过程及机制

微生物是驱动土壤元素生物地球化学循环的引擎.氮循环是土壤生态系统元素循环的核心之一,其四个主要过程,即生物固氮作用、氨化作用、硝化作用、反硝化作用,均由微生物所驱动.近10年来,随着免培养的分子生态学技术和高通量测序技术等的发展,在硝化微生物多样性及其作用机理、厌氧氨氧化过程和机理等研究方面取得了突破性进展.本文重点阐述了我国有关土壤硝化微生物方面的研究进展,在此基础上,简要介绍了反硝化微生物和厌氧氨氧化及硝酸盐异化还原成铵作用的研究进展,并对今后的研究工作提出了展望.今后土壤氮素转化微生物生态学的研究,应瞄准国际微生生态学发展的前沿,加强新技术新方法的应用,结合我国农业可持续发展、资源环境保护和全球变化研究的重大需求,重点开展以下几方面的工作:(1)开展大尺度上土壤硝化作用及氨氧化微生物分布的时空演变特征及驱动因子的研究;(2)加强氮素转化关键微生物过程与机理的研究,并与相关过程的通量(如氨挥发、N2O释放)和反应速率(如矿化速率、硝化速率)关联起来;(3)在特定生态系统中系统研究各个氮转化过程的耦合关系,构建相关氮素转化和氮素平衡模型,为定向调控土壤氮素转化过程,提高氮素利用效率并减少其负面效应提供科学依据.     

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.     

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.     

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.     

青藏高原东缘高山森林-苔原交错带土壤微生物生物量碳、氮和可培养微生物数量的季节动态

为了了解青藏高原东缘高山森林-苔原交错带土壤微生物的特征和季节变化, 研究了米亚罗鹧鸪山原始针叶林、林线、树线、密灌丛、疏灌丛和高山草甸土壤微生物生物量碳(MBC)、氮(MBN)和可培养微生物数量的季节动态。结果表明, 植被类型和季节动态对MBC、MBN和微生物数量都有显著影响。不同时期的微生物在各植被类型间分布有差异, 植物生长季初期和生长季中期, 树线以上群落的MBC高于树线下的群落, 而到生长季末期恰恰相反, 暗针叶林、林线和树线的MBC显著升高, 各植被之间MBC的差异减小; 微生物数量基本上也是以树线为界, 树线以下群落土壤微生物数量显著低于树线以上群落, 其中密灌丛的细菌数量最高; 可培养微生物数量为生长季末期>生长季初期>生长季中期。生长季末期真菌数量显著增加, 且MBC/MBN最高。统计分析表明, MBN与细菌、真菌、放线菌数量存在显著的相关关系, 而MBC仅与真菌数量存在显著相关关系( p < 0.05)。植物生长季末期大量的凋落物输入和雪被覆盖可能是微生物季节变异的外在因素, 而土壤微生物和高山植物对有效氮的竞争可能是微生物季节变异的内在因素。植物生长季初期对氮的吸收和土壤微生物在植物生长季末期对氮的固定加强了高山生态系统对氮的利用。气候变暖可能会延长高山植物的生长季, 增加高山土壤微生物生物量, 加速土壤有机质的分解, 进而改变高山土壤碳的固存速率。     

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.     

基于高通量测序的禁牧对土壤微生物群落结构的影响

麋鹿的采食、躺卧和践踏行为均会对栖息地土壤环境造成影响,进而影响土壤微生物群落结构。利用高通量测序技术,分析江苏大丰麋鹿国家级自然保护区禁牧点和补饲点土壤细菌和真菌群落结构差异,并结合土壤理化性质探究禁牧对土壤微生物群落结构的影响。结果表明细菌群落的优势菌门为变形菌门,真菌群落的优势菌门为子囊菌门。禁牧改变了土壤微生物群落结构,在门水平上提高了变形菌门、放线菌门和担子菌门的相对丰度,降低了绿弯菌门、厚壁菌门和子囊菌门的相对丰度,禁牧点与补饲点土壤微生物群落多样性的相似性较低。冗余分析中,细菌受土壤环境因子的影响大于真菌,其中土壤pH是影响细菌和真菌群落最大的土壤环境因子。研究揭示了禁牧对土壤微生物群落结构的影响,为保护区制定麋鹿生境恢复方案提供参考。     

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.