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[Advances in study of factors affecting soil N mineralization in grassland ecosystems].

The biological and non-biological factors affecting soil N availability in grassland ecosystems were reviewed in this paper. Nitrogen cycling in grassland ecosystems is one of the focuses widely concerned. Nitrogen mineralization is affected by many factors in grassland ecosystem, which can be classified into biological and non-biological ones. Biological factors include soil animals, soil microorganisms and plants. Soil animals could accelerate the organic matter to degrade. The species, structure and function of soil microorganisms correlate significantly with N degradation and mineralization. Different vegetation has different effects on soil nitrogen mineralization. The non-biological factors include environmental factors and anthropogenic disturbance, which have direct and obvious effects on N mineralization. The effects of soil temperature and moisture on N mineralization are given more attention, but many phenomena, such as the effects of soil type, soil structure and vegetation type on N mineralization still could not be explained clearly, and no general agreements were reached. Anthropogenic disturbance such as grazing, firing and fertilization influence N mineralization evidently. It is of great significance to understand the N cycling pattern and N availability in different grassland ecosystems all around the world.

Animals↗

Response of grassland ecosystem to air pollutants: I-Experimental concept and site of the Braunschweig grassland investigation program.

Between 1984 and 1991 in Braunschweig (South-east Lower Saxony, Germany) experiments concerning the influence of chronic stress by air pollutants to a permanent grassland ecosystem were performed, the conceptual background of which is dealt with in this paper. The site itself, the management practices and treatments applied, and the physical climate are described in detail.

Journal Article↗

Effects of field exposure to diazinon on small mammals inhabiting a semienclosed prairie grassland ecosystem. I. Ecological and reproductive effects.

The widespread use of cholinesterase-inhibiting pesticides in the environment presents increasing concerns about their effects on human, wildlife, and ecosystem health. As a group, these pesticides are generally highly toxic and have great potential for negatively affecting nontarget organisms. Small mammals have proven to be ideal biomonitors of environmental contaminants, and were used here to test for possible effects of a widely used cholinesterase-inhibiting insecticide, diazinon, in a natural field setting. Using 12 0.1-ha terrestrial mesocosms, we examined the effects of low-level diazinon exposure on the small mammal communities inhabiting semienclosed grassland ecosystems. Our primary objective was to test the hypothesis that diazinon, applied at two different recommended label application rates, would not cause any observable adverse ecological or reproductive effects on small mammal populations and communities. Experimental small mammal communities consisting of Sigmodon hispidus, Microtus ochrogaster, Reithrodontomys fulvescens, and Mus musculus were stocked at natural densities and sex ratios inside empty mesocosms. Diazinon 4E was applied at two different maximum recommended label application rates, 0.56 kg a.i./ha (1x) and 4.5 kg a.i./ha (8x), and controls remained unsprayed, with four enclosures (replicates) per treatment. Two 30-d trials were run during peak rodent breeding seasons and enclosures were sampled on days 2, 16, and 30 of each trial. Recovery of small mammals was not significantly different among treatments, although fewer animals were recovered from the diazinon-exposed enclosures in both trials. Analysis of trapping data suggested that the normally strong competitive relationship between Sigmodon and Microtus may be altered by the pesticide, favoring Microtus in the diazinon-exposed enclosures. Incidence of reproductive condition was found to be reduced 20 to 80% and 33 to 100% in diazinon-exposed males and females, respectively. Reproductive productivity, including percentage of pregnant females and of females giving birth, was significantly reduced in diazinon-exposed animals. Percentage of pregnant females ranged from 13.6 to 43.5% in diazinon-exposed animals compared to 40 to 80% for control animals, and percentage of females giving birth ranged from 0 to 17% in diazinon-exposed animals compared to 22 to 50% for control animals. Generally, the effects found in this study suggest that diazinon was relatively persistent in the sprayed enclosures and that oral routes of exposure (consumption of dead and dying arthropods, grooming) may have been important. Ecological relationships and reproduction in both herbivorous and omnivorous mammals were negatively impacted by diazinon exposure. Overall, ecological relationships in the enclosed prairie grassland ecosystem were disrupted by diazinon, probably through a combination of sublethal effects, particularly reproductive effects, impacting individuals and their populations. This suggests that negative impacts on populations and community structure and function may persist longer than diazinon persists in the environment.

Animals↗

Distribution of trace element pollutants in a contaminated grassland ecosystem established on metalliferous fluorspar tailings. 1: lead.

Concentrations of lead in vegetation, invertebrates and small mammals in a grassland ecosystem evolved from a mine waste revegetation scheme indicated the high concentration of lead in the original tailings. Lead levels in invertebrates reflected dietary concentrations and feeding strategy, a pattern also true of the indigenous small mammals. Total body and tissue concentrations of lead in the herbivorous Microtus agrestis L. (field vole) and insectivorous Sorex araneus L. (common shrew) were significantly higher in the contaminated grassland than in an uncontaminated area, but concentration ratios (body:diet) were less than unity and there was no evidence of age-dependent accumulation of lead.

Journal Article↗

Distribution of trace element pollutants in a contaminated grassland ecosystem established on metalliferous fluorspar tailings. 2: Zinc.

Concentrations of zinc in vegetation, invertebrates and small mammals in a grassland ecosystem derived from a mine waste vegetation scheme indicated the high concentration of zinc in the original tailings. However, the considerable differences in soil and vegetation zinc concentrations between the tailings and an uncontaminated site were not reflected in the invertebrate and small mammal communities. Invertebrate zinc levels and total body concentrations in Microtus agrestis L. (field vole) and Sorex araneus L. (common shrew) suggested a notable buffering of zinc absorption and tissue accumulation in the presence of substantially elevated dietary zinc. In the small mammals the skeleton appeared to function as the major storage site for zinc, possibly contributing to homeostasis in the soft tissues.

Journal Article↗

Effect of rainfall patterns on soil surface CO2 efflux, soil moisture, soil temperature and plant growth in a grassland ecosystem of northern Ontario, Canada: implications for climate change.

BACKGROUND: The effect of rainfall patterns on soil surface CO2 efflux, soil moisture, soil temperature and plant growth was investigated in a grassland ecosystem of northern Ontario, Canada, where climatic change is predicted to introduce new precipitation regimes. Rain shelters were established in a fallow field consisting mainly of Trifolium hybridum L., Trifolium pratense L., and Phleum pratense L. Daytime ambient air temperatures within the shelters increased by an average of 1.9 degrees C similar to predicted future increases in air temperatures for this region. To simulate six precipitation regimes which cover the maximum range to be expected under climate change, a portable irrigation system was designed to modify the frequency of monthly rainfall events with a constant delivery rate of water, while maintaining contemporary average precipitation volumes. Controls consisted of blocks irrigated with frequencies and total monthly precipitation consistent with the 25 year average rainfall for this location. RESULTS: Seasonal soil moisture correlated with soil surface CO2 efflux (R = 0.756, P < 0.001) and above ground plant biomass (R = 0.447, P = 0.029). By reducing irrigation frequency, soil surface CO2 efflux decreased by 80%, P < 0.001, while soil moisture content decreased by 42%, P < 0.001. CONCLUSIONS: Manipulating the number of precipitation events and inter-rainfall intervals, while maintaining monthly rainfall averages impacted CO2 efflux and plant growth. Even with monthly rainfall averages that are similar to contemporary monthly precipitation averages, decreasing the number of monthly rainfall events reduced soil surface CO2 efflux and plant growth through soil moisture deficits. Although many have speculated that climate change will increase ecosystem productivity, our results show that a reduction in the number of monthly rainfall events while maintaining monthly averages will limit carbon dynamics.

Biomass↗

Arbuscular mycorrhizal community composition associated with two plant species in a grassland ecosystem.

Arbuscular mycorrhizal (AM) fungi are biotrophic symbionts colonizing about two-thirds of land plant species and found in all ecosystems. They are of major importance in plant nutrient supply and their diversity is suggested to be an important determinant of plant community composition. The diversity of the AM fungal community composition in the roots of two plant species (Agrostis capillaris and Trifolium repens) that co-occurred in the same grassland ecosystem was characterized using molecular techniques. We analysed the small subunit (SSU) ribosomal RNA gene amplified from a total root DNA extract using AM fungal-specific primers. A total of 2001 cloned fragments from 47 root samples obtained on four dates were analysed by restriction fragment length polymorphism, and 121 of them were sequenced. The diversity found was high: a total of 24 different phylotypes (groups of phylogenetically related sequences) colonized the roots of the two host species. Phylogenetic analyses demonstrate that 19 of these phylotypes belonged to the Glomaceae, three to the Acaulosporaceae and two to the Gigasporaceae. Our study reveals clearly that the AM fungal community colonizing T. repens differed from that colonizing A. capillaris, providing evidence for AM fungal host preference. In addition, our results reveal dynamic changes in the AM fungal community through time.

Ecosystem↗

Parasitic plants indirectly regulate below-ground properties in grassland ecosystems.

Parasitic plants are one of the most ubiquitous groups of generalist parasites in both natural and managed ecosystems, with over 3,000 known species worldwide. Although much is known about how parasitic plants influence host performance, their role as drivers of community- and ecosystem-level properties remains largely unexplored. Parasitic plants have the potential to influence directly the productivity and structure of plant communities because they cause harm to particular host plants, indirectly increasing the competitive status of non-host species. Such parasite-driven above-ground effects might also have important indirect consequences through altering the quantity and quality of resources that enter soil, thereby affecting the activity of decomposer organisms. Here we show in model grassland communities that the parasitic plant Rhinanthus minor, which occurs widely throughout Europe and North America, has strong direct effects on above-ground community properties, increasing plant diversity and reducing productivity. We also show that these direct effects of R. minor on the plant community have marked indirect effects on below-ground properties, ultimately increasing rates of nitrogen cycling. Our study provides evidence that parasitic plants act as a major driver of both above-ground and below-ground properties of grassland ecosystems.

Bacteria↗

Phosphorus cycles of forest and upland grassland ecosystems and some effects of land management practices.

The distribution of phosphorus capital and net annual transfers of phosphorus between the major components of two unfertilized phosphorus-deficient UK ecosystems, an oak--ash woodland in the Lake District and an Agrostis-Festuca grassland in Snowdonia (both on acid brown-earth soils), have been estimted in terms of kg P ha--1. In both ecosystems less than 3% of the phosphorus, totalling 1890 kg P ha--1 and 3040 kg P ha--1 for the woodland and grassland, respectively, is contained in the living biomass and half that is below ground level. Nearly all the phosphorus is in the soil matrix. Although the biomass phosphorus is mostly in the vegetation, the soil fauna and vegetation is slower (25%) than in the grassland vegetatation (208%). More than 85% of the net annual vegetation uptake of phosphorus from the soil is returned to the soil, mainly in organic debris, which in the grassland ecosystem is more than twice as rich in phosphorus (0.125% P) as in the woodland ecosystem (0.053% P). These concentrations are related to the rates of turnover (input/P content) of phosphorus in the litter layer on the soil surface; it is faster in the grassland (460%) than in the woodland (144%). In both cycles plant uptake of phosphorus largely depends on the release of phosphorus through decomposition of the organic matter returned to soil. In both the woodland and the grassland, the amount of cycling phosphorus is potentially reduced by its immobilization in tree and sheep production and in undecomposed organic matter accumulating in soil. It is assumed that the reductions are counterbalanced by the replenishment of cycling phosphorus by (i) some mineralization of organically bound phosphorus in the mineral soil, (ii) the income in rainfall and aerosols not being effectively lost in soil drainage waters and (iii) rock weathering. The effects of the growth of conifers and sheep grazing on the balance between decomposition and accumulation of organic matter returned to soil are considered in relation to the rate of phosphorus cycling and the pedogenetic changes in soil phosphorus condition leading to reduced fertility. Although controlled sheep grazing speeds up phosphorus cycling and may reverse the pedogenetic trend in favour of soil improvement, conifers may slow down phosphorus cycling and promote the pedogenetic trend towards infertility.

Agriculture↗

Impacts of soil faunal community composition on model grassland ecosystems.

Human impacts, including global change, may alter the composition of soil faunal communities, but consequences for ecosystem functioning are poorly understood. We constructed model grassland systems in the Ecotron controlled environment facility and manipulated soil community composition through assemblages of different animal body sizes. Plant community composition, microbial and root biomass, decomposition rate, and mycorrhizal colonization were all markedly affected. However, two key ecosystem processes, aboveground net primary productivity and net ecosystem productivity, were surprisingly resistant to these changes. We hypothesize that positive and negative faunal-mediated effects in soil communities cancel each other out, causing no net ecosystem effects.

Animals↗

Drought effects on above- and belowground production of a grazed temperate grassland ecosystem.

The effect of climatic variation on terrestrial aboveground productivity (ANPP) has been well studied. However, little is known about how variable climate, including drought, may influence belowground productivity (BNPP), which constitutes most of the annual primary production of grasslands. The objectives of this study were to (1) examine how a 3-year period of declining moisture, which began as climatically wet to average across Yellowstone National Park (YNP) and ended in drought, affected ANPP and BNPP in grasslands of YNP and (2) how herds of grazing ungulates, which were shown previously to stimulate grassland shoot and root growth in YNP, may have interacted with climatic conditions to influence grassland production. ANPP and 0-20 cm BNPP, representing the bulk of the root dynamics, were measured in grazed and ungrazed (fenced) grassland at nine sites ranging widely in elevation, soil conditions and plant production during the 3-year study. Results revealed that 0-20 cm BNPP was strongly influenced by drought (P = 0.0005) and declined from 1999 to 2001 among ungrazed and grazed grasslands by 39 and 49%, respectively. The greater reduction in 0-20 cm BNPP among grazed grasslands was due, in part, to a decline (P = 0.07) in the stimulatory effect of grazing, i.e., the ratio g BNPP stimulated: g shoot consumed. In contrast, ANPP was unaffected by drought in either type of grassland. Thus, the effect of this drought in YNP was a large reduction in BNPP, which was a function of (1) a direct negative influence of increased moisture stress on root growth and (2) a weak interaction between drought and herbivory that led to a decline in the positive feedback from grazers to BNPP. These findings highlight the need to better understand factors that control root growth and to study the effects of climatic variation on grasslands within an ecosystem framework to include potentially important climate-consumer interactions.

Animals↗

Bacterial community dynamics across a floristic gradient in a temperate upland grassland ecosystem.

Alterations in soil bacterial communities across a transect between a semi natural upland grassland and an agriculturally improved enclosure were assessed using culture-based methods and a nucleic-acid-based method, terminal restriction fragment length polymorphism (TRFLP). While plant diversity decreased across the transect towards the improved area, numbers of 16S rDNA terminal restriction fragments increased, indicating an increase in numbers of bacterial ribotypes. Bacterial numbers, microbial activity, and potential functional diversity also followed a similar trend, increasing with decreasing plant diversity. Alterations in bacterial community structure were coincident with changes in soil physicochemical properties which also changed across the transect. Increases in soil pH, nitrate, phosphorus, potassium, and calcium occurred toward the improved grassland, while organic matter and ammonium declined. The inverse relationship between floristic diversity and bacterial ribotype numbers suggests that soil physicochemical factors may be as influential in determining bacterial diversity in soils of upland grassland communities as floristic diversity.

Bacteria↗

Flux and turnover of fixed carbon in soil microbial biomass of limed and unlimed plots of an upland grassland ecosystem.

The influence of liming on rhizosphere microbial biomass C and incorporation of root exudates was studied in the field by in situ pulse labelling of temperate grassland vegetation with (13)CO(2) for a 3-day period. In plots that had been limed (CaCO(3) amended) annually for 3 years, incorporation into shoots and roots was, respectively, greater and lower than in unlimed plots. Analysis of chloroform-labile C demonstrated lower levels of (13)C incorporation into microbial biomass in limed soils compared to unlimed soils. The turnover of the recently assimilated (13)C compounds was faster in microbial biomass from limed than that from unlimed soils, suggesting that liming increases incorporation by microbial communities of root exudates. An exponential decay model of (13)C in total microbial biomass in limed soils indicated that the half-life of the tracer within this carbon pool was 4.7 days. Results are presented and discussed in relation to the absolute values of (13)C fixed and allocated within the plant-soil system.

Biomass↗

Geographical and interannual variability in biomass partitioning in grassland ecosystems: a synthesis of field data.

Biomass partitioning is an important variable in terrestrial ecosystem carbon modeling. However, geographical and interannual variability in f(BNPP), defined as the fraction of belowground net primary productivity (BNPP) to total NPP, and its relationship with climatic variables, have not been explored. Here we addressed these issues by synthesizing 94 site-year field biomass data at 12 grassland sites around the world from a global NPP database and from the literature. Results showed that f(BNPP) varied from 0.40 to 0.86 across 12 sites. In general, savanna and humid savanna ecosystems had smaller f(BNPP) but larger interannual variability in f(BNPP), and cold desert steppes had larger f(BNPP) but smaller interannual variability. While mean f(BNPP) at a site decreased significantly with increasing mean annual temperature and precipitation across sites, no consistent temporal response of f(BNPP) with annual temperature and precipitation was found within sites. Based on these results, both geographical variability in f(BNPP) and the divergent responses of f(BNPP) with climatic variables at geographical and temporal scales should be considered in global C modeling.

Biomass↗

Bromus tectorum invasion alters nitrogen dynamics in an undisturbed arid grassland ecosystem.

The nonnative annual grass Bromus tectorum has successfully replaced native vegetation in many arid and semiarid ecosystems. Initial introductions accompanied grazing and agriculture, making it difficult to separate the effects of invasion from physical disturbance. This study examined N dynamics in two recently invaded, undisturbed vegetation associations (C3 and C4). The response of these communities was compared to an invaded/ disturbed grassland. The invaded/disturbed communities had higher surface NH4+ input in spring, whereas there were no differences for surface input of NO3-. Soil inorganic N was dominated by NH4+, but invaded sites had greater subsurface soil NO3-. Invaded sites had greater total soil N at the surface four years post-invasion in undisturbed communities, but total N was lower in the invaded/disturbed communities. Soil delta15N increased with depth in the noninvaded and recently invaded communities, whereas the invaded/disturbed communities exhibited the opposite pattern. Enriched foliar delta15N values suggest that Bromus assimilated subsurface NO3-, whereas the native grasses were restricted to surface N. A Rayleigh distillation model accurately described decomposition patterns in the noninvaded communities where soil N loss is accompanied by increasing soil delta15N; however, the invaded/ disturbed communities exhibited the opposite pattern, suggesting redistribution of N within the soil profile. This study suggests that invasion has altered the mechanisms driving nitrogen dynamics. Bromus litter decomposition and soil NO3- concentrations were greater in the invaded communities during periods of ample precipitation, and NO3- leached from the surface litter, where it was assimilated by Bromus. The primary source of N input in these communities is a biological soil crust that is removed with disturbance, and the lack of N input by the biological soil crust did not balance N loss, resulting in reduced total N in the invaded/disturbed communities. Bromus produced a positive feedback loop by leaching NO3- from decomposing Bromus litter to subsurface soil layers, accessing that deepsoil N pool with deep roots and returning that N to the surface as biomass and subsequent litter. Lack of new inputs combined with continued loss will result in lower total soil N, evidenced by the lower total soil N in the invaded/disturbed communities.

Biodiversity↗

[Biogeochemical characteristics of nutrient elements in ungrazed grassland ecosystem in Xilin River Basin].

The biogeochemical characteristics of nutrient elements in Leymus chinensis community and Stipa grandis community, which were both scarcely disturbed by human activities, in Xilin River Basin, Inner Mongolia were studied. The results showed that the storage of N, P, K, S, Ca, Mg, and Si in the soil compartment of soil-plant system was higher than that in the plant subsystem of two communities, and their storage in the live shoots compartment was lower than that in the litter compartment. The disappearing rate of N, P, K and S from the litter compartment was higher than that of Ca, Mg, and Si in the two grassland communities. The decomposition rate of litter in Leymus chinensis community was higher than that in Stipa grandis community. In the grassland soil-plant system, the matter flowing mass kept balance in Leymus chinensis community, but lost balance in Stipa grandis community. The matter participating in biogeochemical cycling in Leymus chinensis community was more than that in Stipa grandis community.

Ecosystem↗

Scale-dependent relationships between the spatial distribution of a limiting resource and plant species diversity in an African grassland ecosystem.

One cornerstone of ecological theory is that nutrient availability limits the number of species that can inhabit a community. However, the relationship between the spatial distribution of limiting nutrients and species diversity is not well established because there is no single scale appropriate for measuring variation in resource distribution. Instead, the correct scale for analyzing resource variation depends on the range of species sizes within the community. To quantify the relationship between nutrient distribution and plant species diversity, we measured NO(3)(-) distribution and plant species diversity in 16 paired, modified Whittaker grassland plots in Serengeti National Park, Tanzania. Semivariograms were used to quantify the spatial structure of NO(3)(-) from scales of 0.4-26 m. Plant species diversity (Shannon-Weiner diversity index; H ') was quantified in 1-m(2) plots, while plant species richness was measured at multiple spatial scales between 1 and 1000 m(2). Small-scale variation in NO(3)(-) (<0.4 m) was positively correlated with 1-m(2) H ', while 1000-m(2) species richness was a log-normal function of average NO(3)(-) patch size. Nine of the 16 grassland plots had a fractal (self-similar across scales) NO(3)(-) spatial distribution; of the nine fractal plots, five were adjacent to plots that had a non-fractal distribution of NO(3)(-). This finding offered the unique opportunity to test predictions of Ritchie and Olff (1999): when the spatial distribution of limiting resources is fractal, communities should display a left-skewed log-size distribution and a log-normal relationship between net primary production and species richness. These predictions were supported by comparisons of plant size distributions and biomass-richness relationships in paired plots, one with a fractal and one with a non-fractal distribution of NO(3)(-). In addition, fractal plots had greater large-scale richness than paired non-fractal plots (1,0-1000 m(2)), but neither species diversity ( H') nor richness was significantly different at small scales (1 m(2)). This result is most likely explained by differences in the scale of resource variation among plots: fractal and non-fractal plots had equivalent NO(3)(-) variation at small scales but differed in NO(3)(-) variation at large scales (as measured by the fractal dimension). We propose that small-scale variation in NO(3)(-) is largely due to the direct effects of plants on soil, while patterns of species richness at large scales is controlled by the patch size and fractal dimension of NO(3)(-) in the landscape. This study provides an important empirical step in understanding the relationship between the spatial distribution of resources and patterns of species diversity across multiple spatial scales.

Africa↗

Response of a grassland ecosystem to air pollutants: II--The chemical climate: Fluxes of sedimenting airborne matter.

Measurement of the deposition of sedimenting particles requires a sampling device, which avoids simultaneous deposition of gases and aerosols to the collection surface. A sampler constructed for the purpose of collecting rain and sedimenting particles is described and characterized in detail, in particular with regard to its collection efficiency for rain. Its collection properties for gases and aerosols are shown to be negligible. From two years of sampling at different heights it was found that resuspension of particles and co-condensation of gases near the plant canopy may lead to a major overestimation of bulk deposition. As a consequence, the extension towards the canopy of the constant flux layer for sedimenting particles has to be determined experimentally. Bulk deposition of sodium, potassium, magnesium, calcium, lead, copper, cadmium, manganese, iron, ammonium, nitrate, phosphate, sulfate, total sulfur and chloride at Braunschweig-Völkenrode, Southeast Lower Saxony, Germany, were recorded for six years. During this period a considerable decrease was observed in the deposition of lead, cadmium, nitrate, sulfate and total sulfur.

Journal Article↗