Nitrogen retention efficiency and nitrogen losses of a managed and phytodiverse temperate grassland

Maintaining nitrogen retention efficiency (NRE) is crucial in minimizing N losses when intensifying management of temperate grasslands. Our aim was to evaluate how grassland management practices and sward compositions affect NRE (1 − N losses/soil available N), defined as the efficiency with which soil available N is retained in an ecosystem. A three-factorial grassland management experiment was established with two fertilization treatments (without and combined N, phosphorus and potassium fertilization), two mowing frequencies (cut once and thrice per year) and three sward compositions (control, monocot- and dicot-enhanced swards). We measured N losses as leaching and nitrous oxide emissions, and soil available N as gross N mineralization rates. Fertilization increased N losses due to increased nitrification and decreased microbial N immobilization, and consequently decreased NRE. Intensive mowing partly dampened high N losses following fertilization. Sward compositions influenced NRE but not N losses: control swards that developed for decades under extensive management had the highest NRE, whereas monocot-enhanced sward had the lowest NRE. NRE was highly correlated with microbial NH₄⁺ immobilization and microbial biomass and only marginally correlated with plant N uptake, underlining the importance of microbial N retention in the soil-plant system. Microbial N retention is reflected in NRE but not in indices commonly used to reflect plant response. NRE was able to capture the effects of sward composition and fertilization whereas N losses were only sensitive to fertilization; thus, NRE is a better index when evaluating environmental sustainability of sward compositions and management practices of grasslands.     

Nitrogen relationships in intensively managed temperate grasslands

Summary Most studies of N relationships in grassland have used cut swards. These have shown that for annual inputs of 200 to 400 kg N/ha from fertilizer or fixation, 55 to 80% of the N is recovered in harvested herbage. Generally, no more than 5 to 15% is lost through leaching and denitrification with most of the remaining N incorporated into soil organic matter. The relatively high efficiency of N use by cut swards reflects rapid uptake of N and the removal of a large part of the input in herbage. Inclusion of the grazing ruminant alters the efficiency of N use; only 5–20% of the input is recovered in meat or milk, and 75 to 90% of the N ingested is excreted, mainly as urea in urine. Application of N in urine ranges from 30–100 g/m². Too much N is voided for effective recovery by the sward whilst soils usually contain insufficient C to allow appreciable immobilization. The surfeit is lost. Hydrolysis of urea is usually complete within 24 h of urine deposition. For urine-treated pasture in New Zealand (NZ) losses by NH₃ volatilization of up to 66% of applied N are found during warm dry weather, with an average of 28% for a range of seasonal conditions. In the UK, the average rate of NH₃ loss from an intensively grazed ryegrass sward was 0.75 kg N/ha/day during a 6-month season. NH ₄ ⁺ remaining in the soil may be nitrified, nitrification being complete within 3 to 6 weeks. Although some NO ₃ ^– is recovered by plants, a substantial portion is leached and/or denitrified. On average such losses were 42%, with only 30% of the added N recovered by plants in urine-treated pasture in NZ. In the UK annual leaching of 150 to 190 kg N/ha has been observed for grazed swards receiving 420 kg N/ha/yr. Low retention of N by grazing ruminants results in a breakdown of N relationships in intensively managed grasslands. The substantial losses through NH₃ volatilization, leaching and denitrification have serious agronomic, economic and environmental implications.     

Large-scale manipulation of plant litter and fertilizer in a managed successional temperate grassland

Plant litter may play an important role in herbaceous plant communities by limiting primary production and influencing plant species richness. However, it is not known how the effect of litter interacts with fertilization. We tested for the role of litter and fertilization in a large-scale experiment to investigate effects on diversity and biomass of plant species, growth forms, native vs. non-native groups, and abiotic ecosystem components (e.g., soil moisture, PAR). We manipulated plant litter (removed vs. left in situ) and nutrient availability (NPK-fertilized vs. unfertilized) for 4 years in 314-m² plots, replicated six times, in an old-field grassland. While many of our species-level results supported previously published studies and theory, our plant group results generally did not. Specifically, grass species richness and forb biomass was not affected by either fertilization or plant litter. Moreover, plant litter removal significantly increased non-native plant species richness. Relative to native plant species, all of our experimental manipulations significantly increased both the biomass and the species richness of non-native plant species. Thus, this grassland system was sensitive to management treatments through the facilitation of non-native plant species. We coupled biotic and abiotic components within a nonmetric multidimensional scaling (NMS) analysis to investigate treatment effects, which revealed that specific treatments altered ecosystem development. These results suggest that fertilization and plant litter may have larger impacts on plant communities and on ecosystem properties than previously understood, underscoring the need for larger-scale and longer-term experiments.     

Nitrogen deposition promotes phosphorus uptake of plants in a semi-arid temperate grassland

Plant and Soil - Nitrogen (N) deposition greatly influences ecosystem processes through the alteration of plant nutrition; however, there is limited understanding about the effects of phosphorus...     

Nitrogen competition between three dominant plant species and microbes in a temperate grassland

Background and aims

To test the hypothesis that dominant plant species could acquire different nitrogen (N) forms over a spatial scale and they also have the ability to compete for available N with microbes.

Methods

A short-term ¹⁵N labeling experiment was conducted in the temperate grassland ecosystem of North China in July of 2013. Three N forms (NO₃ ^? _, NH₄ ⁺ and glycine) labeled with ¹⁵N were injected into the two soil depths (0–5 and 5–15 cm) surrounding each plant to explore N acquisition by plants and microbes. Three dominant plant species (Artemisia frigida, Cleistogenes squarrosa and Artemisia capillaris) were investigated.

Results

Two hours after ¹⁵N labeling, all three dominant plant species absorbed both organic and inorganic N, but different patterns were observed at two soil depths. Uptake of NO₃ ^? was significantly higher at 0–5 cm than at 5–15 cm soil depth among all the dominant plant species. ¹⁵N recovery by microbes was significantly higher than plants. However, ¹⁵N recovery by plants showed different patterns over soil depths.

Conclusions

Dominant plant species in the temperate grassland have different patterns in acquisition of N added to soil in organic form and absorption of inorganic N, and microbes were more effectively than plants at competing for N in a short-term period.

     

Nitrogen enrichment weakens ecosystem stability through decreased species asynchrony and population stability in a temperate grassland

Biodiversity generally promotes ecosystem stability. To assess whether the diversity–stability relationship observed under ambient nitrogen (N) conditions still holds under N enriched conditions, we designed a 6‐year field experiment to test whether the magnitude and frequency of N enrichment affects ecosystem stability and its relationship with species diversity in a temperate grassland. Results of this experiment showed that the frequency of N addition had no effect on either the temporal stability of ecosystem and population or the relationship between diversity and stability. Nitrogen addition decreased ecosystem stability significantly through decreases in species asynchrony and population stability. Species richness was positively associated with ecosystem stability, but no significant relationship between diversity and the residuals of ecosystem stability was detected after controlling for the effects of the magnitude of N addition, suggesting collinearity between the effects of N addition and species richness on ecosystem stability, with the former prevailing over the latter. Both population stability and the residuals of population stability after controlling for the effects of the magnitude of N addition were positively associated with ecosystem stability, indicating that the stabilizing effects of component populations were still present after N enrichment. Our study supports the theory predicting that the effects of environmental factors on ecosystem functioning are stronger than those of biodiversity. Understanding such mechanisms is important and urgent to protect biodiversity in mediating ecosystem functioning and services in the face of global changes.     

Nitrogen response efficiency increased monotonically with decreasing soil resource availability: a case study from a semiarid grassland in northern China

The concept of nutrient use efficiency is central to understanding ecosystem functioning because it is the step in which plants can influence the return of nutrients to the soil pool and the quality of the litter. Theory suggests that nutrient efficiency increases unimodally with declining soil resources, but this has not been tested empirically for N and water in grassland ecosystems, where plant growth in these ecosystems is generally thought to be limited by soil N and moisture. In this paper, we tested the N uptake and the N use efficiency (NUE) of two Stipa species (S. grandis and S. krylovii) from 20 sites in the Inner Mongolia grassland by measuring the N content of net primary productivity (NPP). NUE is defined as the total net primary production per unit N absorbed. We further distinguished NUE from N response efficiency (NRE; production per unit N available). We found that NPP increased with soil N and water availability. Efficiency of whole-plant N use, uptake, and response increased monotonically with decreasing soil N and water, being higher on infertile (dry) habitats than on fertile (wet) habitats. We further considered NUE as the product of the N productivity (NP the rate of biomass increase per unit N in the plant) and the mean residence time (MRT; the ratio between the average N pool and the annual N uptake or loss). The NP and NUE of S. grandis growing usually in dry and N-poor habitats exceeded those of S. krylovii abundant in wet and N-rich habitats. NUE differed among sites, and was often affected by the evolutionary trade-off between NP and MRT, where plants and communities had adapted in a way to maximize either NP or MRT, but not both concurrently. Soil N availability and moisture influenced the community-level N uptake efficiency and ultimately the NRE, though the response to N was dependent on the plant community examined. These results show that soil N and water had exerted a great impact on the N efficiency in Stipa species. The intraspecific differences in N efficiency within both Stipa species along soil resource availability gradient may explain the differences in plant productivity on various soils, which will be conducive to our general understanding of the N cycling and vegetation dynamics in northern Chinese grasslands.     

Root trait responses of six temperate grassland species to intensive mowing and NPK fertilisation: a field study in a temperate grassland

Background and aims

Plant traits may characterize functional ecosystem properties and help to predict community responses to environmental change. Since most traits used relate to aboveground plant organs we aim to explore the indicative value of root traits.

Methods

We examined the response of root traits (specific root length [SRL], specific root surface area [SRA], root diameter [RD], root tissue mass density [TMD], root N concentration) in six grassland species (3 grasses, 3 herbs) to four management regimes (low vs. high mowing frequency; no fertilization vs. high NPK fertilization). The replicated experiment in temperate grassland with long continuity simulated the increase in grassland management intensity in the past 50 years in Central Europe.

Results

Increasing mowing frequency (one vs. three cuts per year) led to no significant root trait changes. NPK fertilization resulted in considerable trait shifts with all species responding in the same direction (higher SRL, SRA and N concentration, lower TMD) but at different magnitude. Fertilization-driven increases in SRA were mainly caused by lowered tissue density while root diameter reduction was the main driver of SRL increases.

Conclusion

We conclude that root morphological traits may be used as valuable indicators of environmental change and increasing fertilization in grasslands.     

Precipitation regulates plant gas exchange and its long-term response to climate change in a temperate grassland

Aims Climate change largely impacts ecosystem carbon and water cycles by changing plant gas exchange, which may further cause positive or negative feedback to global climate change. However, long-term global change manipulative experiments are seldom conducted to reveal plant ecophysiological responses to climatic warming and altered precipitation regimes.Methods An 8-year field experiment with both warming and increased precipitation was conducted in a temperate grassland in northern China. We measured leaf gas exchange rates (including plant photosynthesis, transpiration and instantaneous water use efficiency [WUE]) of two dominant plant species (Stipa sareptana var. krylovii and Agropyron cristatum) from 2005 to 2012 (except 2006 and 2010) and those of other six species from 2011 to 2012.Important findings Increased precipitation significantly stimulated plant photosynthetic rates (A) by 29.5% and 19.9% and transpiration rates (E) by 42.2% and 51.2% for both dominant species S. sareptana var. krylovii and A. cristatum, respectively, across the 8 years. Similarly, A and E of the six plant functional types were all stimulated by increased precipitation in 2011 and 2012. As the balance of A and E, the instantaneous WUEs of different plant species had species-specific responses to increased precipitation. In contrast, neither warming nor its interaction with increased precipitation significantly affected plant leaf gas exchange rates. Furthermore, A and E of the two dominant species and their response magnitudes to water treatments positively correlated with rainfall amount in July across years. We did not find any significant difference between the short-term versus long-term responses of plant photosynthesis, suggesting the flexibility of leaf gas exchange under climate change. The results suggest that changing precipitation rather than global warming plays a prominent role in determining production of this grassland in the context of climate change.     

Low assimilate partitioning to root biomass is associated with carbon losses at an intensively managed temperate grassland

Plant and Soil - This study aimed to investigate how efficiently assimilated carbon (C) is incorporated in plant biomass at an intensively managed old permanent grassland, how C is partitioned...     

Carbon dioxide and water vapor exchange in a warm temperate grassland

Grasslands cover about 40% of the ice-free global terrestrial surface, but their contribution to local and regional water and carbon fluxes and sensitivity to climatic perturbations such as drought remains uncertain. Here, we assess the direction and magnitude of net ecosystem carbon exchange (NEE) and its components, ecosystem carbon assimilation (A _c) and ecosystem respiration (R _E), in a southeastern United States grassland ecosystem subject to periodic drought and harvest using a combination of eddy-covariance measurements and model calculations. We modeled A _c and evapotranspiration (ET) using a big-leaf canopy scheme in conjunction with ecophysiological and radiative transfer principles, and applied the model to assess the sensitivity of NEE and ET to soil moisture dynamics and rapid excursions in leaf area index (LAI) following grass harvesting. Model results closely match eddy-covariance flux estimations on daily, and longer, time steps. Both model calculations and eddy-covariance estimates suggest that the grassland became a net source of carbon to the atmosphere immediately following the harvest, but a rapid recovery in LAI maintained a marginal carbon sink during summer. However, when integrated over the year, this grassland ecosystem was a net C source (97 g C m^–2 a^–1) due to a minor imbalance between large A _c (–1,202 g C m^–2 a^–1) and R _E (1,299 g C m^–2 a^–1) fluxes. Mild drought conditions during the measurement period resulted in many instances of low soil moisture (<0.2 m³m^–3), which influenced A _c and thereby NEE by decreasing stomatal conductance. For this experiment, low had minor impact on R _E. Thus, stomatal limitations to A _c were the primary reason that this grassland was a net C source. In the absence of soil moisture limitations, model calculations suggest a net C sink of –65 g C m^–2 a^–1 assuming the LAI dynamics and physiological properties are unaltered. These results, and the results of other studies, suggest that perturbations to the hydrologic cycle are key determinants of C cycling in grassland ecosystems.     

Carbon dioxide fluxes in a spatially and temporally heterogeneous temperate grassland

Landscape position, grazing, and seasonal variation in precipitation and temperature create spatial and temporal variability in soil processes, and plant biomass and composition in grasslands. However, it is unclear how this variation in plant and soil properties affects carbon dioxide (CO₂) fluxes. The aim of this study is to explore the effect of grazing, topographic position, and seasonal variation in soil moisture and temperature on plant assimilation, shoot and soil respiration, and net ecosystem CO₂ exchange (NEE). Carbon dioxide fluxes, vegetation, and environmental variables were measured once a month inside and outside long-term ungulate exclosures in hilltop (dry) to slope bottom (mesic) grassland throughout the 2004 growing season in Yellowstone National Park. There was no difference in vegetation properties and CO₂ fluxes between the grazed and the ungrazed sites. The spatial and temporal variability in CO₂ fluxes were related to differences in aboveground biomass and total shoot nitrogen content, which were both related to variability in soil moisture. All sites were CO₂ sinks (NEE>0) for all our measurments taken throughout the growing season; but CO₂ fluxes were four- to fivefold higher at sites supporting the most aboveground biomass located at slope bottoms, compared to the sites with low biomass located at hilltops or slopes. The dry sites assimilated more CO₂ per gram aboveground biomass and stored proportionally more of the gross-assimilated CO₂ in the soil, compared to wet sites. These results indicate large spatio-temporal variability of CO₂ fluxes and suggest factors that control the variability in Yellowstone National Park.     

Arthropod corpses in a temperate grassland: a limited supply?

The disappearance of arthropod corpses in a temperate grassland of the Mediterranean coast (Barcelona, Spain) has been analyzed in this study. The mean time in which the corpses stayed out in the field is less than five minutes (= 285 ± 282 s), in a range of 2-2228 s. This holds true in cleared areas as well as in high vegetation areas. Ants are the principal scavengers of this type of remains, although the species that collect them vary according to the time of day and season of year, with the corpses'disappearance being the most rapid during the middle hours of the day, precisely when ants'density on the ground is less. The reason for this was the peak daytime abundance of insectivore, opportunisms ants. Anyway, the short time lapse that the prey stay in the field - at any time of the day - can be considered as an indirect estimate of the scarcity of these resources.     

Dung beetles as secondary seed dispersers in a temperate grassland

The two-phase dispersal event in which dung beetles move seeds after endozoochory is often assumed to be advantageous for plant regeneration. Because seeds are expected to end up in favourable and safe germination sites, it is considered as an example of directed dispersal. However, literature so far is restricted to tropical rain forest ecosystems, while data for temperate regions are lacking. In this study, the effect of dung beetles on seedling establishment of endozoochorically dispersed seeds is evaluated for a temperate grassland ecosystem. We performed a field experiment in which cages excluded dung beetles from horse and cattle dung samples with mixed-in grass seeds. Seed germination from these samples was significantly higher than that from samples which were accessible to dung beetles. This indicates that the effect of dung beetles on short-term seedling establishment was negative, which contrasts with the patterns found for large-seeded species used in tropical studies. This is most likely attributed to the lack of roller species and the larger depth at which tunneling Geotrupes species bury seeds.     

Physiognomic changes in response to herbivory increase carbon allocation to roots in a temperate grassland of central Argentina

Plant Ecology - In most temperate grasslands, cattle grazing can promote physiognomic changes on plant communities, as well as changes in species growth patterns. Through these changes in...     

Soil seed bank composition and diversity in a managed temperate deciduous forest

Little is known about the influence of forest management on the interaction between seed bank and aboveground vegetation. We surveyed seed banks and vegetation in 10 forest stands under similar abiotic conditions but submitted either to a coppice-with-standards treatment (n=5) or to a selective-cutting system (n=5). We analyzed species composition and diversity, community ecological profile, and distribution of taxa among different life forms, strategy, morphology and functional type categories. A total of 2085 seedlings (8296 seedsm^–2) germinated-corresponding to 28 species, among which Juncus effusus was the most abundant. Fifty-seven percent of the species were also recorded in the aboveground vegetation, the dominant species being Rubus fruticosus agg., but only 28% of the aboveground species were present in the seed bank. Our results suggest that (1) vernal geophytes and shade-tolerant perennials, which group most true forest species, are not incorporated in the seed bank, (2) parent plants of most seeds were present either in the stand in an earlier dynamic stage or apart from the stand and long-distance dispersed, (3) as expected, early-successional species are well represented in the seed bank, (4) forestry vehicles seem to be a major means of dispersion for stress-tolerant species normally found in forest lanes and wheel tracks. We conclude that seed banks contain species that have a potentially negative impact on the true forest flora and, thus, forest management should minimize soil disturbance and retain remnants of old-coppice woods to conserve disturbance-sensitive true forest species.     

Living roots magnify the response of soil organic carbon decomposition to temperature in temperate grassland

Increasing atmospheric carbon dioxide (CO₂) concentration is both a strong driver of primary productivity and widely believed to be the principal cause of recent increases in global temperature. Soils are the largest store of the world's terrestrial C. Consequently, many investigations have attempted to mechanistically understand how microbial mineralisation of soil organic carbon (SOC) to CO₂ will be affected by projected increases in temperature. Most have attempted this in the absence of plants as the flux of CO₂ from root and rhizomicrobial respiration in intact plant‐soil systems confounds interpretation of measurements. We compared the effect of a small increase in temperature on respiration from soils without recent plant C with the effect on intact grass swards. We found that for 48 weeks, before acclimation occurred, an experimental 3 °C increase in sward temperature gave rise to a 50% increase in below ground respiration (ca. 0.4 kg C m^?2; Q₁₀ = 3.5), whereas mineralisation of older SOC without plants increased with a Q₁₀ of only 1.7 when subject to increases in ambient soil temperature. Subsequent ¹⁴C dating of respired CO₂ indicated that the presence of plants in swards more than doubled the effect of warming on the rate of mineralisation of SOC with an estimated mean C age of ca. 8 years or older relative to incubated soils without recent plant inputs. These results not only illustrate the formidable complexity of mechanisms controlling C fluxes in soils but also suggest that the dual biological and physical effects of CO₂ on primary productivity and global temperature have the potential to synergistically increase the mineralisation of existing soil C.