Soil N dynamics in a natural calcareous grassland under a changing climate

 This paper reports the results from a medium-term field scale investigation into the effects of simulated climate change on soil N mineralisation in a semi-natural calcareous grassland in southern England. The experiment utilised soil warming cables, automatic rainshelters and a watering system to examine two climate change scenarios: warmer winters with summer drought and warmer winters with enhanced summer rainfall. Gross N mineralisation rates in treated plots were determined, using ¹⁵N pool dilution techniques, at 6-weekly intervals over a 3-year period. Results from control plots showed a strong seasonality of mineralisation with highest rates in autumn and winter and lowest rates in summer. They suggest that water availability is the main constraint on microbial processes and plant growth. Unexpectedly, additional summer rainfall had no direct effect on N mineralisation at the time of application (summer). The treatment did, however, significantly (<0.05%) reduce rates in subsequent autumn and winter months. In contrast, summer drought significantly increased N mineralisation rates in autumn and winter. Winter warming similarly had no direct effect on N mineralisation in winter but decreased rates in spring. We hypothesise that the observed treatment effects result from changes in organic C and N input, in plant litter, resulting from the direct impact of climatic manipulation on perennial plant growth, death and senescence. This paper compares and contrasts the response to climate manipulation in the grassland system with results from other ecosystem types such as northern forests. Received: 1 December 1997     

Short-term effects of different management regimes on the response of calcareous grassland vegetation to increased nitrogen

The short-term (4-year period) effects of different management regimes on the response of calcareous grassland vegetation to increased nitrogen were investigated. Four levels of nitrogen and two levels of management were applied in a factorial design. Besides, the effects of management practices alone on species diversity were also studied. Species diversity was investigated at different levels of sub-sampling using a nested plot design. In this way, both species saturation and accumulation curves could be established. The effect of community structure, as reflected by the light regimes at various heights in the vegetation, on species diversity was also investigated. Species diversity at all levels of sub-sampling decreased significantly with management regimes and nitrogen supply rates. Percent light penetration differed significantly among management regimes and nitrogen supply rates. Grazing proved to be most efficient in countering the negative effects of nitrogen supply, but could not prevent competitors from becoming dominant. In the absence of any form of disturbance, species diversity decreased relatively rapidly. Decreased light availability, the loss of gap formation, a decrease in small-scale environmental heterogeneity and proliferation of strong competitors, all may have contributed to the observed decrease in species richness. The results of this study highlight the importance of management in maintaining high species diversity in the studied calcareous grassland.     

Does seed dispersal by sheep affect the population genetic structure of the calcareous grassland species Bromus erectus?

This study investigates the contribution of the traditional land-use system “sheep herding” (transhumance) to the gene flow among populations. Three sheep trails with altogether 12 calcareous grassland sites were chosen for the analysis of the tussock grass Bromus erectus, which is known to be the species with the highest rate of dispersal via sheep fleece. Each trail links two to four isolated populations. Allozymes were chosen as genetic markers. Six loci were used, five of which were polymorphic. While genetic variability within populations was found to be quite high (averaged h=0.352), only little between-population differentiation was measured (G_ST=5.5%). Despite this, the UPGMA-dendrogram of Nei's genetic distance shows genetic similarity relations largely congruent with the sheep herding trails. However, a (more) detailed comparison reveals several differences between genetic patterns and trail system. Hence, additional factors other than dispersal by sheep are regarded as being responsible for exchange of genes, and sheep herding is considered to have only minor importance for the genetic patterns of the investigated species. The comparison of Bromus erectus to species with the same life history traits (i.e. pollination by wind, a perennial life cycle, and autopolyploid constitution) shows similar values, thus providing further evidence for the relatively small contribution of dispersal via sheep fleece.     

The effect of pig slurry on exchangeable potassium in calcareous soils

Pig slurry in quantities of 200, 400, 500, 600, 800, and 1000 m³ ha^-1 year^-1 was added to two calcareous soils, which differed in concentration and type of clay minerals. The soils were cultivated with two successive crops of pepper and tomatoes grown in containers. A control was given no slurry. The soils were analysed after harvesting for exchangeable K⁺. Differences in exchangeable K⁺ were studied in relation to the concentration and type of clay minerals. The soil with the higher clay content and of the illite type retained K in the exchangeable form to a much greater extent than the soil with the low clay content. In the soil with the lower clay content, of the interstratified illitic — montmorillonite type, very little K was incorporated into the exchange complex. The exchange capacity being low, the amount of K added had little effect on the level of exchangeable K⁺.     

Effect of PGPR,Phosphate sources and vermicompost on growth and nutrients uptake by lettuce in a calcareous soil

In order to study the effect of plant growth promoting rhizobacteria (PGPR), vermicompost and phosphate sources on the growth and nutrients uptake by lettuce, a greenhouse experiments was conducted. Treatments consisted of PGPR (Pseudomonas fluorescens) (with and without inoculation), vermicompost (0 and 1% w/w) and phosphate sources (control, rock phosphate (RP), tricalcium phosphate and triple super phosphate (TSP) at 25 mg P kg^?1 level). Biological fertilizers, RP and TSP significantly increased shoot dry matter (SDM) and some measured nutrients uptake. Co-application of PGPR and RP, in non-vermicompost treatments significantly increased SDM, shoot nitrogen, phosphorus (P), potassium, zinc and manganese uptake rates. Shoot P uptake had no significant difference between TSP and RP treatments. Co-application of PGPR, vermicompost and TSP significantly decreased SDM, which may be due to the P toxic levels in the plant aerial parts and/or the inhibition of the bacterial activities in the rhizosphere soil.     

The electro-ultrafiltration method for controlling the effect of Bacillus cereus on phosphorus mobilization in a calcareous soil

Summary The effects of phosphate-solubilizing bacteria on the phosphorus mobilization and P utake by Lolium multiflorum from a calcareous soil were investigated using a greenhouse pot experiment. The soil was inoculated with Bacillus cereus and the pots were either planted with Lolium multiflorum or left fallow. Treatments were designed to study the effect of inoculation of phosphate-solubilizing bacteria on phosphorus dynamics throughout the experiment. The dynamics were studied by the electro-ultrafiltration method, the results of which were used to interpret phosphorus mobilization in the soil. Phosphate uptake by L. multiflorum from the inoculated soil showed a 40% increase over the control soil, and an increase in total dry matter of 50%.     

The efect of Zinc (Zn) application on the available Iron (Fe) contents of calcareous soils in thrace region

This research was carried out in order to determine the effect of zinc (Zn) application on the available iron (Fe) contents of calcareous soils in Thrace Region. For this purpose, an experiment was carried out in greenhouse conditions by maize growing. The effect of increasing rates of Zn application on the biological indices of maize plant was determined. Fe concentration of maize plant decreased with Zn application and dry matter yield and Fe uptake of maize plant increased with Zn application. These decreases and increases were determined to be statistically significant (P&lt;0.01).     

The effect of vesicular-arbuscular mycorrhizae in decreasing Ca acquisition by alfalfa plants in calcareous soils

Summary The legume Medicago sativa L. was grown in three calcareous soils supplied with increasing amounts of soluble phosphate, or a vesicular-arbuscular mycorrhizal (VAM) inoculum. The three test soils had high concentrations of extractable Ca. Analyses of dry-matter production and of the concentrations and content of the nutrients N, P, K, Ca, and Mg in plant tissues showed that, for each soil, a particular level of P application was able to match the VAM effects on N, P, and K levels. The Ca concentration and content in the VAM inoculated plants were, however, significantly lower than those in the P-supplied non-mycorrhizal treatments that matched the VAM effects. The N:P and the K:P ratios were about the same for mycorrhizal and non-mycorrhizal P-supplied control plants in all the three soils, but VAM inoculation lowered the Ca:P ratio in all soils. The mycorrhizae decreased Mg uptake in one of the soils, where non-mycorrhizal plants had high Mg concentrations in tissues. It is concluded that VAM depress the excessive acquisition of Ca by plants in calcareous soils.     

Effects of acid rain on leaf-litter decomposition in a beech forest on calcareous soil

Summary The effects of simulated acid rain on litter decomposition in a calcareous soil (pH_{H 2 O} 5.8) were studied. Litterbags (45 m and 1 mm mesh size) containing freshly fallen beech leaf litter were exposed to different concentrations of acid in a beech forest on limestone (Göttinger Wald. Germany) for 1 year. Loss of C, the ash content, and CO₂–C production were measured at the end of the experiment. Further tests measured the ability of the litter-colonizing microflora to metabolize ¹⁴C-labelled beech leaf litter and hyphae. The simulated acid rain strongly reduced CO₂–C and ¹⁴CO₂–C production in the litter. This depression in production was very strong when the input of protons was 1.5 times greater than the normal acid deposition, but comparatively low when the input was 32 times greater. acid deposition may thus cause a very strong accumulation of primary and secondary C compounds in the litter layer of base-rich soils, even with a moderate increase in proton input. The presence of mesofauna significantly reduced the ability of the acid rain to inhibit C mineralization. The ash content to the 1-mm litterbags indicated that this was largely due to transport of base-rich mineral soil into the litter.     

A landscape model for quantifying the trade-off between conservation needs and economic constraints in the management of a semi-natural grassland community

We present a landscape model to investigate the ecological consequences and costs of different management regimes in semi-natural grasslands. The model integrates dynamic abiotic conditions, management (i.e. disturbance) regime and response of more than 50 characteristic plant and insect species by modelling the dynamics of relevant niche parameters as predictors for species distribution models. We compare our results for exemplary scenarios differing in spatial and temporal scales and exemplary species belonging to different functional groups through several steps of aggregation.Our analysis aims at the question whether an infrequent massive disturbance by rototilling can serve as a less expensive alternative to annual mowing for preserving the characteristic species composition of open dry grasslands in Southern Germany. Rototilling results in a shifting mosaic determining the habitat quality for plant and animal species that may reduce the survival of local or regional populations.For some meadow species as well as the encroaching shrub species, rototilling has a detrimental effect on regional habitat quality. Other species, e.g. weeds and annual pioneers, strongly benefit or show only negligible reaction. Since this is a multi-objective problem, there is a no magic bullet in selecting an optimum scenario of measures. But by visualising the trade-off between ecological consequences and costs, our model is a valuable tool for conservation managers providing a sound scientific basis for management decisions relying on available ecological knowledge. It is also an interesting example for a model describing complex communities in a relatively simple way, simultaneously considering the main driving factors.     

Spatial distribution of fungal communities in a coastal grassland soil

In grasslands, saprotrophic fungi, including basidiomycetes, are major decomposers of dead organic matter, although spatial distributions of their mycelial assemblages are little described. The aim of this study was to characterise the scale and distribution of saprotrophic fungal communities in a coastal grassland soil using terminal restriction fragment length polymorphism (T-RFLP).Soil fungi were sampled at Point Reyes, California, USA, by taking forty-five 26 mm diam. cores in a spatially defined manner. Within each sampled core, complete core sections at 1-2 cm and 14-15 cm depths were removed and sub-sampled for DNA extraction and amplification using the primer pairs ITS1F-FAM/ITS4 (general fungi) or ITS1F-FAM/ITS4B (basidiomycete-specific).Nonmetric Multidimensional Scaling showed that general fungal communities could be clearly separated by depth, although basidiomycete communities could not. There were no strong patterns of community similarity or dissimilarity for general or basidiomycete fungal communities at horizontal geographical distances from 25 cm to 96 m in the upper horizon. These results show considerable vertical, but little horizontal, variability in fungal community structure in a semi-natural grassland at the spatial scales measured here.     

Microbial biomass P,labile P,and acid phosphatase activity in the humus layer of a spruce forest,after repeated additions of fertilizers

A 20-year-old forest fertilization trial was used to investigate the effects of repeated P additions on P availability in the humus layer of a Norway spruce forest soil. N was supplied annually, and P, K, and micronutrients were supplied every 4th year. The last P application was made 2 years before the investigation started. Microbial P concentrations in the P+NK+micro-amended plots were about half as high as those in the control and the N-only treatment. In plots without P amendments, around 50% of the total P in the humus layer was found in microorganisms, whereas in P-amended plots the figure was around 25%. The block supporting more rapid tree growth, situated on the middle of a slope, showed a significantly higher microbial biomass P concentration than the less productive block at the bottom of the slope. Labile P concentrations did not vary between treatments and thus could not have directly contributed to the treatment-related differences in total microbial biomass P. Acid phosphatase activities were around three times lower in the sites treated with P+NK+micro-nutrients. Two sources are suggested for the acid phosphatases, active excretion by living roots and fungi and passive release from ruptured cells. For all eight plots investigated, there was a positive correlation (R=0.83) between acid phosphatase activity and the microbial P concentration. The P concentration in current-year needles was the lowest in the N-only treatment at 1,13 mg g^-1 dry weight, and the highest in the P+NK + micronutrients + lime treatment at 1.92 mg g^-1 dry weight. The P:N ratio in needles varied from 0.115 in the P+NK + micronutrients + lime plots to 0.068 in the N-only plots. The latter value is at the level where P is considered to be the growth rate-determining nutrient.     

Estimating the three-dimensional structure of canopy foliage based on the light measurements in a Betula ermanii stand

In order to more accurately evaluate the functional activity of forest stands by canopy production and evapotranspiration, we improved the methods for field measurements and statistical modeling to estimate foliage configuration (spatial distribution of leaves) while simultaneously reconstructing the three-dimensional photosynthetically active photon flux density (PPFD) distribution (PPFD pattern) in a forest canopy. By using a sensor (photodiode) array, a PPFD pattern was observed in summer 2002 under the canopy in an even-aged, pure stand of Japanese mountain birch Betula ermanii Cham. (17-years old) in Hokkaido, northern Japan. A Markov chain Monte Carlo (MCMC) sampling technique is developed such that a set of foliage configurations generated by the model referred to as the Gibbs foliage canopy (GFC) approximates the field-measured PPFD pattern. The posterior distribution of the foliage configurations is generated by the parallel tempering MCMC of eight independent series of foliage configurations. The GFC model generated the posterior distribution of the LAI estimates (mean 4.56) that appeared to be appropriate in comparison to other LAI estimates of the B. ermanii stand based on the indirect and nondestructive methods by LAI-2000 (LAI = 3.43) and litterfall traps (LAI = 5.56) because they could be under- and overestimated, respectively. Our evaluations of the canopy production and evapotranspiration rates suggest that the relationship between LAI and canopy functions was not very simple because it depended on the nonlinear functional forms of the leaf responses of photosynthesis and transpiration to PPFD. The current study demonstrates an application of MCMC techniques that can generate a set of possible structures of unobserved/unobservable objects based on the high-resolution dataset obtained by some indirect (or remote-sensing) methods.     

Response of microbial community composition and activity in agricultural and grassland soils after a simulated rainfall

Rainfall in Mediterranean climates may affect soil microbial processes and communities differently in agricultural vs. grassland soils. We explored the hypothesis that land use intensification decreases the resistance of microbial community composition and activity to perturbation. Soil carbon (C) and nitrogen (N) dynamics and microbial responses to a simulated Spring rainfall were measured in grassland and agricultural ecosystems. The California ecosystems consisted of two paired sets: annual vegetable crops and annual grassland in Salinas Valley, and perennial grass agriculture and native perennial grassland in Carmel Valley. Soil types of the respective ecosystem pairs were derived from granitic parent material and had sandy loam textures. Intact cores (30 cm deep) were collected in March 1999. After equilibration, dry soil cores (approx. −1 to −2 MPa) were exposed to a simulated Spring rainfall of 2.4 cm, and then were measured at 0, 6, 24, and 120 h after rewetting. Microbial biomass C (MBC) and inorganic N did not respond to rewetting. N₂O and CO₂ efflux and respiration increased after rewetting in all soils, with larger responses in the grassland than in the agricultural soils. Phospholipid fatty acid (PLFA) profiles indicated that changes in microbial community composition after rewetting were most pronounced in intensive vegetable production, followed by the relict perennial grassland. Changes in specific PLFA markers were not consistent across all sites. There were more similarities among microbial groups associated with PLFA markers in agricultural ecosystems than grassland ecosystems. Differences in responses of microbial communities may be related to the different plant species composition of the grasslands. Agricultural intensification appeared to decrease microbial diversity, as estimated from numbers of individual PLFA identified for each ecosystem, and reduce resistance to change in microbial community composition after rewetting. In the agricultural systems, reductions in both the measures of microbial diversity and the resistance of the microbial community composition to change after a perturbation were associated with lower ecosystem function, i.e. lower microbial responses to increased moisture availability.     

The response of a high-yielding canola hybrid to sulfur fertilization in three contrasting Saskatchewan soils

Abstract

Canola (Brassica napus) is the primary oilseed crop in western Canada; however, it is often grown on sulfur (S)-deficient soils. Moreover, canola has a high S demand compared to cereals and, therefore, is particularly sensitive to S deficiency. This study examined the growth and nutrient uptake responses of a high-yielding canola hybrid cultivar to S fertilization when grown on three contrasting soils differing in S fertility, with and without the addition of fertilizer S. The soils were collected from three soil-climatic zones within Saskatchewan (Brown, Black, and Gray) and three different fertilizer S forms were used: ammonium sulfate (AS); ammonium thiosulfate (ATS); and a composite fertilizer containing nitrogen (N), phosphorus (P), and S (NPS; 50-50 blend of sulfate (SO₄) salt and elemental S). Sulfur fertilization increased the canola biomass, along with plant uptake of N, P, and S on all three soils. Fertilizer S use efficiency (i.e. recovery) ranged from 11-75%. For all three soils, the general trend among fertilizer S forms for biomass, nutrient uptake, and fertilizer use efficiency was AS?>?ATS?>?NPS. The greatest differences were observed with the Gray soil, which had the poorest S fertility. Residual soil SO₄ after harvest was greater for ATS and NPS; reflecting continued oxidation of thiosulfate and elemental S to SO₄. Principal component analysis demonstrated the importance of tissue N:S ratio as a key diagnostic measurement related to canola growth and nutrient uptake in S-deficient soils.     

Microbial reaction in activity, biomass, and community structure after long-term continuous mixing of a grassland soil

Long-term continuous mixing at 40% water holding capacity (WHC) or as slurry at 400% WHC should result in increased soil organic matter decomposition rates in comparison to a control treatment at 40% WHC, but may have strong impacts on soil microbial indices for activity, biomass, and community structure. The amount of extractable inorganic N (NO₃-N+NH₄-N) accumulated in the soil solution after 40 weeks of incubation at 25 °C was 3% of total N in the control treatment and 4% in the two continuous mixing treatments. However, in the treatment mixing at 40% WHC, this 33% increase compared to the control treatment might be explained solely by the decrease in microbial biomass N. In the control treatment, microbial indices decreased in the order microbial biomass C (−10%), microbial biomass N (−40%), ergosterol (−45%) and ATP (−60%). In the treatment mixing at 40% WHC, all four microbial biomass indices were significantly lower than the respective index in the control treatment. This was especially true for microbial biomass N. In the treatment mixing as slurry, only the contents of microbial biomass C and ATP were significantly lower in comparison to the control treatment. The correspondence analysis ordination biplot of the phospholipid fatty acid (PLFA) profiles showed distinct clusters for the three treatments at the end of the incubation. The strongest relative decline of 64% was observed for the fungi-specific PLFA 18:3ω6 in the treatment mixing as slurry in comparison to the control treatment. The content of total bacterial PLFA decreased only by 23%. The differences between the control treatment and the treatment mixing at 40% WHC were less apparent. Fungi represent on average 21% of total microbial biomass C at the end of the incubation if the ergosterol content is recalculated into fungal biomass C. In accordance with this percentage, 22% of the group-specific PLFA could be attributed to fungi.     

Changes of lignin phenols and neutral sugars in different soil types of a high-elevation forest ecosystem 25 years after forest dieback

Long-term effects of forest disturbance 25 yr ago on lignin and non-cellulosic polysaccharide pools in an unmanaged high-elevation Norway spruce (Picea abies L. [Karst.]) forest were investigated by comparing three dieback sites with three adjacent control sites with non-infested spruce on identical soils. Samples were taken from the forest floor and the mineral soil; one Ah horizon sample per site was physically fractionated into density and particle size fractions. Additionally, changes in the above- and belowground input of lignin and non-cellulosic polysaccharides after forest dieback were quantified. Lignin and its degree of structural alteration in plant and soil samples were assessed by CuO oxidation and subsequent analysis of the lignin phenols. Non-cellulosic polysaccharides were determined after hydrolysis with trifluoroacetic acid (TFA), derivatisation of their neutral sugar monomers by reduction to alditols, and subsequent acetylation. The total plant-derived input of lignin and non-cellulosic polysaccharides to the soil was similar for the dieback and the control sites. The chemical composition of the input has changed considerably after forest dieback, as shown by significantly higher syringyl/vanillyl (S/V) ratios and significantly lower (galactose+mannose)/(arabinose+xylose) (GM/AX) ratios. This indicates a changed plant input and a higher contribution of microbial sugars. Contents of lignin phenols in the forest floor and coarse particle size fractions of the A horizons were significantly smaller at the dieback sites (p<0.01). Moreover, larger acid-to-aldehyde ratios of vanillyl units (Ac/Al)_v indicated an increased degree of lignin phenol alteration. Also contents of neutral sugars were significantly (p<0.01) smaller in the forest floor, but not in the A horizons of the dieback sites. The GM/AX mass ratios as well as the (rhamnose+fucose)/(arabinose+xylose) (RF/AX) ratios in the forest floor and coarse particle size fractions of the mineral topsoil were significantly (p<0.01) larger after forest dieback, indicating a larger relative contribution of microbial sugars. In general, the lignin phenol and neutral sugar pools of all three soil types exhibited similar response patterns to the changed site conditions. Our results demonstrate that the lignin and neutral sugar pools of humic topsoil horizons are highly sensitive to forest disturbances. However, the two compounds show different patterns in the mineral soil, with the major neutral sugar pool being stabilized against changes whereas the lignin phenol pool decreases significantly.     

The effect of petrocalcic horizons on the content and distribution of organic carbon in a Mediterranean semiarid landscape

Petrocalcic horizons are frequent in soils of semiarid landscapes. A survey of SIC and SOC contents made in Southern Spain in a pilot area with well defined geomorphological surfaces showed that topsoils overlying petrocalcic horizons are almost twice as rich in SOC as soil of similar depth without petrocalcic horizons. This could be due to impedance to root penetration, changes in redox potential and soil water availability caused by the presence of indurated crust. Soil age, on the contrary, seems not be an essential factor, since only a short time is required to reach a steady state in SOC in comparison to the time span available for soil formation on the different geomorphic surfaces.     

Sites and processes of nutrient accumulation in the subsoil through water percolation from the plow layer in a submerged paddy soil microcosm

The leaching of nutrients from the plow layer by water percolation and their accumulation in the subsoil observed in a Japanese paddy field (Katoh et al. 2004: Soil Sci. Plant Nutr., 50, 721-729) were determined semi-quantitatively in a soil column experiment. Ca²⁺, Mg²⁺, K⁺, Mn²⁺, Fe²⁺, and phosphate in percolating water from the plow layer soil column were retained in the subsoil columns that were connected to the plow layer soil column. Fe²⁺, K⁺, and phosphate accumulated in the uppermost part of the subsoil. Accumulation of Fe²⁺ in the uppermost part of the subsoil was presmnably due to the cation exchange process with concomitant desorption of Ca²⁺. In contrast, Ca²⁺ and Mg²⁺ in percolating water from the plow layer soil colmnn accumulated once in the subsoil, and translocated downwards slowly with successive water percolation. Considerable amounts of inorganic carbon (IC) and dissolved organic carbon (DOC) in percolating water from the plow layer soil column were also retained in the subsoil columns. IC did not accumulate a gaseous form.     

The classification, distribution, and extent of soils with a xeric moisture regime in the United States

Soils with cool, moist winters and relatively warm, dry summers, a Mediterranean climate, are recognized as having a xeric moisture regime in Soil Taxonomy. These soils are classified mostly in taxa that use the formative element “xer” in the name. Soil series with either a xeric moisture regime or an aridic regime bordering on a xeric moisture regime make up more than 48,640,000 hectares in the western part of the United States. They are classified in the orders of Mollisols (20,080,000 ha), Aridisols (11,200,000 ha), Alfisols (5,320,000 ha), Inceptisols (4,800,000 ha), Entisols (4,400,000 ha), Vertisols (1,520,000 ha), Andisols (960,000 ha), and Ultisols (960,000 ha).