Effects of perennial neighbors and nitrogen pulses on growth and nitrogen uptake by Bromus tectorum

An experiment was conducted to determine if growth and biomass responsesof the annual grass Bromus tectorum are affected by themagnitude and timing of nitrogen (N) pulses and if these responses areinfluenced by different perennial neighbor species. Nitrogen(NH₄:NO₃) was applied in three pulse treatments of varyinginterpulse length (3-d, 9-d, or 21-d between N additions). The total amount of Nadded was the same among treatments; hence, both the frequency and magnitude ofN pulses varied (i.e., the longer the interpulse period,the greater the amount of N added for a single pulse).Bromus showed little response to the different N-pulsetreatments. The only characteristic that varied among pulse treatments wasspecific leaf area (SLA), which was significantly greater whenBromus was grown under the 21-d N pulse than when grownunder the 3-d or 9-d N pulses. Bromus height, leaf andtiller numbers, leaf area and aboveground biomass were not affected by theN-pulse treatments nor were tissue-N contents and concentrations. However,Bromus production and tissue-N were significantly differentwhen Bromus was grown with different perennial neighborspecies. Tiller production, aboveground biomass, and seed numbers ofBromus were lowest when the perennial neighbor was thetussock grass Agropyron desertorum, intermediate when theneighbor was the evergreen shrub Artemisia tridentata, andgreatest when the neighbor was the deciduous shrub Chrysothamnusnauseosus. N contents of Bromus leaves were alsolowest when the neighbor was Agropyron. In contrast, root Nuptake capacities were greatest for Agropyron-Bromus rootmixes and lowest for Chrysothamnus-Bromus root mixes. Theseresults suggest that perennial neighbors affect growth, seed production, and Nuptake of Bromus to a greater extent than the timing andmagnitude of N pulses.     

Effects of nitrogen availability on competition between Bromus tectorum and Bouteloua gracilis

Exotic plant invasions are a serious concern for land managers and conservationists. There is evidence that increased nitrogen availability favors exotic species and decreased nitrogen availability favors non-weedy native species. This study was conducted to test the effect of nitrogen availability on competition between two grass species with contrasting life histories, cheatgrass (Bromus tectorum), a North American exotic, and blue grama (Bouteloua gracilis), a North American native. We investigated the effects of nitrogen availability and competition on aboveground biomass, belowground biomass, height, and % nitrogen tissue concentrations by growing the two species in the greenhouse under five levels of nitrogen and six levels of competition. Nitrogen availability affected competition between Bromus tectorum and Bouteloua gracilis. At the lowest level of N availability, neither species was affected by competition. As N availability increased, aboveground biomass gain of Bromus was more negatively affected by intraspecific competition relative to interspecific competition while the opposite occurred for Bouteloua. At the competition level at which each species gained the most aboveground biomass, Bromus had a linear response to increasing N availability while the response of Bouteloua was asymptotic. Our results do provide some support for the theory that fast growing exotic species have a rapid response to nutrient enrichment while native non-weedy species do not, and that low N levels can reduce competitive pressure from the exotic on the native.     

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.     

Carbon fluxes, nitrogen cycling, and soil microbial communities in adjacent urban, native and agricultural ecosystems

Urban ecosystems are expanding globally, and assessing the ecological consequences of urbanization is critical to understanding the biology of local and global change related to land use. We measured carbon (C) fluxes, nitrogen (N) cycling, and soil microbial community structure in a replicated (n=3) field experiment comparing urban lawns to corn, wheat–fallow, and unmanaged shortgrass steppe ecosystems in northern Colorado. The urban and corn sites were irrigated and fertilized. Wheat and shortgrass steppe sites were not fertilized or irrigated. Aboveground net primary productivity (ANPP) in urban ecosystems (383±11 C m^?2 yr^?1) was four to five times greater than wheat or shortgrass steppe but significantly less than corn (537±44 C m^?2 yr^?1). Soil respiration (2777±273 g C m^?2 yr^?1) and total belowground C allocation (2602±269 g C m^?2 yr^?1) in urban ecosystems were both 2.5 to five times greater than any other land‐use type. We estimate that for a large (1578 km²) portion of Larimer County, Colorado, urban lawns occupying 6.4% of the land area account for up to 30% of regional ANPP and 24% of regional soil respiration from land‐use types that we sampled. The rate of N cycling from urban lawn mower clippings to the soil surface was comparable with the rate of N export in harvested corn (both ～12–15 g N m^?2 yr^?1). A one‐time measurement of microbial community structure via phospholipid fatty acid analysis suggested that land‐use type had a large impact on microbial biomass and a small impact on the relative abundance of broad taxonomic groups of microorganisms. Our data are consistent with several other studies suggesting that urbanization of arid and semiarid ecosystems leads to enhanced C cycling rates that alter regional C budgets.     

Stimulation of grassland nitrogen cycling under carbon dioxide enrichment

 Nitrogen (N) limits plant growth in many terrestrial ecosystems, potentially constraining terrestrial ecosystem response to elevated CO₂. In this study, elevated CO₂ stimulated gross N mineralization and plant N uptake in two annual grasslands. In contrast to other studies that have invoked increased C input to soil as the mechanism altering soil N cycling in response to elevated CO₂, increased soil moisture, due to decreased plant transpiration in elevated CO₂, best explains the changes we observed. This study suggests that atmospheric CO₂ concentration may influence ecosystem biogeochemistry through plant control of soil moisture. Received: 18 December 1995 / Accepted: 19 June 1996     

Effects of grassland species on decomposition of litter and soil microbial communities

Decomposition of litter is greatly influenced not only by its chemical composition but also by activities of soil decomposers. By using leaf litter from 15 plant species collected from semi-natural and improved grasslands, we examined (1) how interspecific differences in the chemical composition of litter influence the abundance and composition of soil bacterial and fungal communities and (2) how such changes in microbial communities are related to the processes of decomposition. The litter from each species was incubated in soil of a standard composition for 60 days under controlled conditions. After incubation, the structure of bacterial and fungal communities in the soil was examined using phospholipid fatty-acid analysis and denaturing gradient gel electrophoresis. Species from improved grasslands had significantly higher rates of nitrogen mineralization and decomposition than those from semi-natural grasslands because the former were richer in nitrogen. Litter from improved grasslands was also richer in Gram-positive bacteria, whereas that from semi-natural grasslands was richer in actinomycetes and fungi. Nitrogen content of litter also influenced the composition of the fungal community. Changes in the composition of both bacterial and fungal communities were closely related to the rate of litter decomposition. These results suggest that plant species greatly influence litter decomposition not only through influencing the quality of substrate but also through changing the composition of soil microbial communities.     

Effects of drought and N-fertilization on N cycling in two grassland soils

Changes in frequency and intensity of drought events are anticipated in many areas of the world. In pasture, drought effects on soil nitrogen (N) cycling are spatially and temporally heterogeneous due to N redistribution by grazers. We studied soil N cycling responses to simulated summer drought and N deposition by grazers in a 3-year field experiment replicated in two grasslands differing in climate and management. Cattle urine and NH₄NO₃ application increased soil NH₄ ⁺ and NO₃ ^? concentrations, and more so under drought due to reduced plant uptake and reduced nitrification and denitrification. Drought effects were, however, reflected to a minor extent only in potential nitrification, denitrifying enzyme activity (DEA), and the abundance of functional genes characteristic of nitrifying (bacterial and archaeal amoA) and denitrifying (narG, nirS, nirK, nosZ) micro-organisms. N₂O emissions, however, were much reduced under drought, suggesting that this effect was driven by environmental limitations rather than by changes in the activity potential or the size of the respective microbial communities. Cattle urine stimulated nitrification and, to a lesser extent, also DEA, but more so in the absence of drought. In contrast, NH₄NO₃ reduced the activity of nitrifiers and denitrifiers due to top-soil acidification. In summary, our data demonstrate that complex interactions between drought, mineral N availability, soil acidification, and plant nutrient uptake control soil N cycling and associated N₂O emissions. These interactive effects differed between processes of the soil N cycle, suggesting that the spatial heterogeneity in pastures needs to be taken into account when predicting changes in N cycling and associated N₂O emissions in a changing climate.     

Contribution of root to soil respiration and carbon balance in disturbed and undisturbed grassland communities, northeast China

Changes in the composition of plant species induced by grassland degradation may alter soil respiration rates and decrease carbon sequestration; however, few studies in this area have been conducted. We used net primary productivity (NPP), microbial biomass carbon (MBC), and soil organic carbon (SOC) to examine the changes in soil respiration and carbon balance in two Chinese temperate grassland communities dominated by Leymus chinensis (undisturbed community; Community 1) and Puccinellia tenuiflora (degraded community; Community 2), respectively. Soil respiration varied from 2.5 to 11.9 g CO₂ m⁻² d⁻¹ and from 1.5 to 9.3 g CO₂ m⁻² d⁻¹, and the contribution of root respiration to total soil respiration from 38% to 76% and from 25% to 72% in Communities 1 and 2, respectively. During the growing season (May–September), soil respiration, shoot biomass, live root biomass, MBC and SOC in Community 2 decreased by 28%, 39%, 45%, 55% and 29%, respectively, compared to those in Community 1. The considerably lower net ecosystem productivity in Community 2 than in Community 1 (104.56 vs. 224.73 g C m⁻² yr⁻¹) suggests that the degradation has significantly decreased carbon sequestration of the ecosystems.     

Effects of olive mill wastewater on soil carbon and nitrogen cycling

This study investigated the cycling of C and N following application of olive mill wastewater (OMW) at various rates (0, 42, 84, and 168 m³/ha). OMW stimulated respiration rate throughout the study period, but an increase in soil organic matter was observed only at the highest rate. Soil phenol content decreased rapidly within 2 weeks following application but neither phenol oxidase and peroxidase activity nor laccase gene copies could explain this response. Soil NH₄ ⁺-N content increased in response to OMW application rate, while an opposite trend observed for NO₃ ^?-N, which attributed to immobilization. This decrease was in accordance with amoA gene copies of archaeal and bacterial ammonia oxidizers in the first days following OMW application. Afterwards, although amoA gene copies and potential nitrification rates recovered to values similar to or higher than those in the non-treated soils, NO₃ ^?-N content did not change among the treatments. A corresponding increase in denitrifying gene copies (nirK, nirS, nosZ) during that period indicates that denitrification, stimulated by OMW application rate, was responsible for this effect; a hypothesis consistent with the decrease in total Kjeldahl nitrogen content late in the season. The findings suggest that land application of OMW is a promising practice for OMW management, even at rates approaching the soil water holding capacity.     

Contrasting effects of resource availability and plant mortality on plant community invasion by Bromus tectorum L.

The positive effect of disturbance on plant community invasibility is one of the more consistent results in invasion ecology. It is generally attributed to a coincident increase in available resources (due to the disturbance) that allows non-resident plant species to establish (Davis MA, Grime JP Thompson K, J Ecol 88:528–534, 2000). However, most research addressing this issue has been in artificial or highly modified plant communities. Our goal in this study was to investigate the interactive effects of resource availability and plant mortality disturbance on the invasion of natural plant communities. We conducted a series of experiments that examined the response of Bromus tectorum L., a highly invasive annual grass, to experimentally created gradients of resource availability [nitrogen (N) and water] and resident plant species mortality. We found that B. tectorum biomass was co-limited by N and water. Biomass at the end of the growing season was a saturating function (i.e., increased to a maximum) of water, which determined maximum biomass, and N, which determined the rate at which maximum biomass was attained. Despite that fact that plant mortality increased N availability, it had a negative impact on invasion success. Plant mortality also decreased foliar cover, standing dead biomass, and soil cover by litter. In harsh environments, removing foliar and soil cover may increase germination and seedling stress by increasing soil temperatures and water loss. Across all treatments, B. tectorum success decreased with decreasing foliar cover and standing dead biomass. This, in combination with the strong limitation of B. tectorum biomass by water in this experiment, suggests that our plant mortality disturbance removed soil cover that may have otherwise aided B. tectorum invasion into this semi-arid plant community by reducing water stress.     

Aeolian process effects on vegetation communities in an arid grassland ecosystem

Many arid grassland communities are changing from grass dominance to shrub dominance, but the mechanisms involved in this conversion process are not completely understood. Aeolian processes likely contribute to this conversion from grassland to shrubland. The purpose of this research is to provide information regarding how vegetation changes occur in an arid grassland as a result of aeolian sediment transport. The experimental design included three treatment blocks, each with a 25 × 50 m area where all grasses, semi-shrubs, and perennial forbs were hand removed, a 25 × 50 m control area with no manipulation of vegetation cover, and two 10 × 25 m plots immediately downwind of the grass-removal and control areas in the prevailing wind direction, 19° north of east, for measuring vegetation cover. Aeolian sediment flux, soil nutrients, and soil seed bank were monitored on each treatment area and downwind plot. Grass and shrub cover were measured on each grass-removal, control, and downwind plot along continuous line transects as well as on 5 × 10 m subplots within each downwind area over four years following grass removal. On grass-removal areas, sediment flux increased significantly, soil nutrients and seed bank were depleted, and Prosopis glandulosa shrub cover increased compared to controls. Additionally, differential changes for grass and shrub cover were observed for plots downwind of vegetation-removal and control areas. Grass cover on plots downwind of vegetation-removal areas decreased over time (2004-2007) despite above average rainfall throughout the period of observation, while grass cover increased downwind of control areas; P. glandulosa cover increased on plots downwind of vegetation-removal areas, while decreasing on plots downwind of control areas. The relationships between vegetation changes and aeolian sediment flux were significant and were best described by a logarithmic function, with decreases in grass cover and increases in shrub cover occurring with small increases in aeolian sediment flux.     

Effects of nitrogen addition on vegetation and ecosystem carbon in a semi-arid grassland

To clarify responses of plant and soil carbon (C) and nitrogen (N) pools in grassland ecosystem to N addition, a field experiment was performed in a grassland in Keerqin Sandy Lands, Northeast China. We investigated vegetation composition and C and N pools of plant and soil (0–30 cm) after five consecutive years of N addition at a rate of 20 g N m^?2 y^?1. Vegetation composition and species diversity responded dramatically to N addition, as dominance by C₄ perennials was replaced with C₃ annuals. Carbon in aboveground pool increased significantly (over two-fold), mainly due to the increase of the C in aboveground living plants and surface litter, which increased by 98 and 134%, respectively. Although soil C did not change significantly, the root C pool decreased in response to 5 years of N addition. The total ecosystem C pool was not significantly impacted by N addition because the large soil pool did not respond to N addition, and the increase in aboveground C was offset by the decrease in root C pool. Moreover, N addition significantly increased the aboveground N pool, but had no significant effects on belowground and total ecosystem N pools. Our results suggest that in the mid-term N addition alters the C and N partitioning in above- and belowground pools, but has no significant effects on total ecosystem C and N pools in these N-limited grasslands.     

Bromus tectorum (Poaceae) in midcontinental United States: Population genetic analysis of an ongoing invasion

Biological invasions can be substantially influenced by the genetic sampling associated with a species' introduction. As a result, we assessed the genetic and evolutionary consequences of the entry and spread of the invasive grass Bromus tectorum (cheatgrass) across the United States midcontinent through an analysis of 54 populations, using enzyme electrophoresis. On average, these populations display 1.04 alleles per locus (A), 4.1% percent polymorphic loci per population (%P) and an expected mean heterozygosity (H(exp)) value of 0.009. Heterozygotes, which have been rarely reported for B. tectorum in North America, occur in three populations in the midcontinent and are likely novel multilocus genotypes that arose postimmigration. The midcontinent distribution of multilocus genotypes suggests that plant immigrants came directly from either the native range or the eastern United States, or both. Continued dispersal of preadapted genotypes and the assembly of populations that are genetic admixtures may enhance this invasion by increasing both the genetic diversity within populations and the selection of novel genotypes arising from occasional outcrossing. The potential for postimmigration evolution in most species points to the largely unrecognized need to block the introduction of new, potentially aggressive genotypes of an alien species already in the United States.     

Translocation and turnover of rhizodeposit carbon within soil microbial communities of an extensive grassland ecosystem

Background and Aims

A substantial amount of photosynthesized plant-C is allocated belowground in grassland ecosystems where it influences the structure and function of the soil microbial community with potential implications for C cycling and storage. We applied stable isotope probing of microbial PLFAs and repeated soil sampling in a grassland over a period of 1 year to assess the role of microbial communities in the cycling of rhizodeposit-C.

Methods

Pulse-labeling with ¹³CO₂ was performed in a grassland site near Gent (Belgium). Soil samples were taken 24 h, 1 week, 1 month, 4 months, 9 months and 1 year following labeling and analyzed for ¹³C in soil, roots and microbial PLFAs.

Results

C enrichment of PLFAs occurred rapidly (within 24 h) but temporally varied across microbial groups. PLFAs indicative for fungi and gram-negative bacteria showed a faster ¹³C uptake compared to gram-positive bacteria and actinomycetes. However, the relative ¹³C concentrations of the latter communities increased after 1 week, while those of fungi decreased and those of gram-negative bacteria remained constant. PLFA ¹³C mean residence times were much shorter for fungi compared to bacteria and actinomycetes.

Conclusions

Our results indicate temporally varying rhizodeposit-C uptake by different microbial groups, and faster turnover rates of mycorrhizal versus saprotrophic fungi and fungi versus bacteria. Fungi appeared to play a major role in the initial processing and possible rapid channeling of rhizodeposit-C into the soil microbial community. Actinomycetes and gram-positive bacteria appeared to have a delayed utilization of rhizodeposit-C or to prefer other C sources upon rhizodeposition.     

Genetic variation and local adaptation at a cheatgrass (Bromus tectorum) invasion edge in western Nevada

Cheatgrass ( Bromus tectorum ) is an invasive weed in western North America found primarily growing at elevations less than 2200 m. We asked whether cheatgrass is capable of becoming adapted to a marginal habitat, by investigating a population at a high elevation invasion edge. We used a combination of methods, including reciprocal field transplants, controlled environment studies and molecular analysis. High levels of SSR gene diversity (0.50 vs. 0.43) and comparable variation in phenotypic traits were observed at both the invasion edge and a low elevation, high-density population. Three heterozygotes were observed in the edge population, which is unusual in this predominantly self-pollinating plant. Plants from high elevations germinated more slowly in a growth chamber and had slower seedling growth rates. Survivorship was low at the edge (13%), compared with the low elevation site (55%), but surviving plants were of similar size and had equivalent reproductive output. Seed size positively affected survival and plant performance in the field and this trait was inherited. Emergence timing affected survival at the low elevation site and germination timing was also inherited. Local adaptation was seen in the low, rather than in the high elevation site, because of differential survival. While there was no evidence for local adaptation to the high elevation site observed in the field, family level and genotype-level differences in traits that affected field performance, high genetic diversity at the invasion edge, and evidence of outcrossing in this highly selfing species indicates that the potential for adaptation to a marginal habitat exists within this population.     

氮添加对沙质草地微生物呼吸与根系呼吸的影响

土壤呼吸可以细化为根系呼吸和微生物呼吸,二者对氮添加的响应有所不同.本文以科尔沁沙质草地为研究对象,探讨氮添加对土壤CO₂排放的影响,并细化为微生物呼吸和根系呼吸的响应特征.结果表明: 在观测期(5—10月),土壤呼吸、微生物呼吸月动态均呈先升高后降低的趋势;微生物呼吸是土壤呼吸的主要贡献者,占82.6%;观测期内根系呼吸贡献率随月份而变化,根系呼吸贡献率两个峰值分别出现在5月(占49.4%)和8月(占41.9%),6个月的平均贡献率为17.4%;在10 ℃条件下,根系呼吸较微生物呼吸对氮添加的响应更为敏感,微生物呼吸速率在氮添加后降低了3.9%,而根系呼吸降低了17.7%;氮添加提高了土壤呼吸、微生物呼吸温度敏感性Q₁₀值,也提高了二者对土壤水分变化的敏感程度.     

Species evenness and invasion resistance of experimental grassland communities

Concern for biodiversity loss coupled with the accelerated rate of biological invasions has provoked much interest in assessing how native plant species diversity affects invasibility. Although experimental studies extensively document the effects of species richness on invader performance, the role of species evenness in such studies is rarely examined. Species evenness warrants more attention because the relative abundances of species can account for substantially more of the variance in plant community diversity and tend to change more rapidly and more frequently in response to disturbances than the absolute numbers of species. In this study, we experimentally manipulated species evenness within native prairie grassland mesocosms. We assessed how evenness affected primary productivity, light availability and the resistance of native communities to invasion. The primary productivity of native communities increased significantly with species evenness, and this increase in productivity was accompanied by significant decreases in light availability. However, evenness had no effect on native community resistance to invasion by three common exotic invasive species. In this study, niche complementarity provides a potential mechanism for the effects of evenness on productivity and light availability, but these effects apparently were not strong enough to alter the invasibility of the experimental communities. Our results suggest that species evenness enhances community productivity but provides no benefit to invasion resistance in otherwise functionally diverse communities.