Variation in root-associated phosphatase activities in wheat contributes to the utilization of organic P substrates in vitro, but does not explain differences in the P-nutrition of plants when grown in soils

To understand whether genotypic variation in root-associated phosphatase activities in wheat impacts on its ability to acquire phosphorus (P), various phosphatase activities of roots were measured in relation to the utilization of organic P substrates in agar, and the P-nutrition of plants was investigated in a range of soils. Root-associated phosphatase activities of plants grown in hydroponics were measured against different organic P substrates. Representative genotypes were then grown in both agar culture and in soils with differing organic P contents and plant biomass and P uptake were determined. Differences in the activities of both root-associated and exuded phosphodiesterase and phosphomonoesterase were observed, and were related to the P content of plants supplied with either ribonucleic acid or glucose 6-phosphate, respectively, as the sole form of P. When the cereal lines were grown in different soils, however, there was little relationship between any root-associated phosphatase activity and plant P uptake. This indicates that despite differences in phosphatase activities of cereal roots, such variability appears to play no significant role in the P-nutrition of the plant grown in soil, and that any benefit derived from the hydrolysis of soil organic P is common to all genotypes.

Diversity in expression of phosphorus (P) responsive genes in Cucumis melo L.

Background: Phosphorus (P) is a major limiting nutrient for plant growth in many soils. Studies in model species have identified genes involved in plant adaptations to low soil P availability. However, little information is available on the genetic bases of these adaptations in vegetable crops. In this respect, sequence data for melon now makes it possible to identify melon orthologues of candidate P responsive genes, and the expression of these genes can be used to explain the diversity in the root system adaptation to low P availability, recently observed in this species. Methodology and Findings: Transcriptional responses to P starvation were studied in nine diverse melon accessions by comparing the expression of eight candidate genes (Cm-PAP10.1, Cm-PAP10.2, Cm-RNS1, Cm-PPCK1, Cm-transferase, Cm-SQD1, Cm-DGD1 and Cm-SPX2) under P replete and P starved conditions. Differences among melon accessions were observed in response to P starvation, including differences in plant morphology, P uptake, P use efficiency (PUE) and gene expression. All studied genes were up regulated under P starvation conditions. Differences in the expression of genes involved in P mobilization and remobilization (Cm-PAP10.1, Cm-PAP10.2 and Cm-RNS1) under P starvation conditions explained part of the differences in P uptake and PUE among melon accessions. The levels of expression of the other studied genes were diverse among melon accessions, but contributed less to the phenotypical response of the accessions. Conclusions: This is the first time that these genes have been described in the context of P starvation responses in melon. There exists significant diversity in gene expression levels and P use efficiency among melon accessions as well as significant correlations between gene expression levels and phenotypical measurements.

Considerations on the use of the p-nitrophenyl phosphomonoesterase assay in the study of the phosphorus nutrition of soil borne fungi

The p-nitrophenyl phosphomonoesterase assay (p NPPase) is commonly used to measure cell-wall-associated and extracellular phosphatase activity of soil fungi. p NPPases are usually assayed in the context of fungal nutrition, where inorganic P supply might be enhanced by the mineralisation of monoester organic P sources in the soil. The importance of the assay to the P nutrition of soil fungi is considered based on the evidence currently available including the consistency of methodological approach. The nature of organic P in the soil and the relevance of the assay to some specific soil substrates is discussed, particularly the chemistry and bioavailability of myo-inositol hexakisphosphate and the lower inositol phosphates. The evidence for the long-term stability of p NPPases in the soil is examined in the light of the persistence of p NPPase in soils. The role of persistent extracellular fungal p NPPases in the soil P cycle is discussed. Conclusions from p NPPase based studies must be based upon an appreciation of the constraints of the assay and the complex chemistry of organic P and p NPPase in the soil.

Spatial variability of flow statistics within regular building arrays

Turbulence statistics obtained by direct numerical simulations are analysed to investigate spatial heterogeneity within regular arrays of building-like cubical obstacles. Two different array layouts are studied, staggered and square, both at a packing density of λp=0.25 . The flow statistics analysed are mean streamwise velocity ( u− ), shear stress ( u′w′−−−− ), turbulent kinetic energy (k) and dispersive stress fraction ( u˜w˜ ). The spatial flow patterns and spatial distribution of these statistics in the two arrays are found to be very different. Local regions of high spatial variability are identified. The overall spatial variances of the statistics are shown to be generally very significant in comparison with their spatial averages within the arrays. Above the arrays the spatial variances as well as dispersive stresses decay rapidly to zero. The heterogeneity is explored further by separately considering six different flow regimes identified within the arrays, described here as: channelling region, constricted region, intersection region, building wake region, canyon region and front-recirculation region. It is found that the flow in the first three regions is relatively homogeneous, but that spatial variances in the latter three regions are large, especially in the building wake and canyon regions. The implication is that, in general, the flow immediately behind (and, to a lesser extent, in front of) a building is much more heterogeneous than elsewhere, even in the relatively dense arrays considered here. Most of the dispersive stress is concentrated in these regions. Considering the experimental difficulties of obtaining enough point measurements to form a representative spatial average, the error incurred by degrading the sampling resolution is investigated. It is found that a good estimate for both area and line averages can be obtained using a relatively small number of strategically located sampling points.

Wind-tunnel modelling of dispersion from a scalar area source in urban-like roughness

A wind-tunnel study was conducted to investigate ventilation of scalars from urban-like geometries at neighbourhood scale by exploring two different geometries a uniform height roughness and a non-uniform height roughness, both with an equal plan and frontal density of λ p = λ f = 25%. In both configurations a sub-unit of the idealized urban surface was coated with a thin layer of naphthalene to represent area sources. The naphthalene sublimation method was used to measure directly total area-averaged transport of scalars out of the complex geometries. At the same time, naphthalene vapour concentrations controlled by the turbulent fluxes were detected using a fast Flame Ionisation Detection (FID) technique. This paper describes the novel use of a naphthalene coated surface as an area source in dispersion studies. Particular emphasis was also given to testing whether the concentration measurements were independent of Reynolds number. For low wind speeds, transfer from the naphthalene surface is determined by a combination of forced and natural convection. Compared with a propane point source release, a 25% higher free stream velocity was needed for the naphthalene area source to yield Reynolds-number-independent concentration fields. Ventilation transfer coefficients w T /U derived from the naphthalene sublimation method showed that, whilst there was enhanced vertical momentum exchange due to obstacle height variability, advection was reduced and dispersion from the source area was not enhanced. Thus, the height variability of a canopy is an important parameter when generalising urban dispersion. Fine resolution concentration measurements in the canopy showed the effect of height variability on dispersion at street scale. Rapid vertical transport in the wake of individual high-rise obstacles was found to generate elevated point-like sources. A Gaussian plume model was used to analyse differences in the downstream plumes. Intensified lateral and vertical plume spread and plume dilution with height was found for the non-uniform height roughness

Sufficiency of Favard's condition for a class of band-dominated operators on the axis

The purpose of this paper is to show that, for a large class of band-dominated operators on $\ell^\infty(Z,U)$, with $U$ being a complex Banach space, the injectivity of all limit operators of $A$ already implies their invertibility and the uniform boundedness of their inverses. The latter property is known to be equivalent to the invertibility at infinity of $A$, which, on the other hand, is often equivalent to the Fredholmness of $A$. As a consequence, for operators $A$ in the Wiener algebra, we can characterize the essential spectrum of $A$ on $\ell^p(Z,U)$, regardless of $p\in[1,\infty]$, as the union of point spectra of its limit operators considered as acting on $\ell^p(Z,U)$.

Shifts in rhizosphere microbial communities and enzyme activity of Poa alpina across an alpine chronosequence

This study quantifies the influence of Poa alpina on the soil microbial community in primary succession of alpine ecosystems, and whether these effects are controlled by the successional stage. Four successional sites representative of four stages of grassland development (initial, 4 years (non-vegetated); pioneer, 20 years; transition, 75 years; mature, 9500 years old) on the Rotmoos glacier foreland, Austria, were sampled. The size, composition and activity of the microbial community in the rhizosphere and bulk soil were characterized using the chloroform-fumigation extraction procedure, phospholipid fatty acid (PLFA) analysis and measurements of the enzymes beta-glucosidase, beta-xylosidase, N-acetyl-beta-glucosaminidase, leucine aminopeptidase, acid phosphatase and sulfatase. The interplay between the host plant and the successional stage was quantified using principal component (PCA) and multidimensional scaling analyses. Correlation analyses were applied to evaluate the relationship between soil factors (C-org, N-t, C/N ratio, pH, ammonium, phosphorus, potassium) and microbial properties in the bulk soil. In the pioneer stage microbial colonization of the rhizosphere of P. alpina was dependent on the reservoir of microbial species in the bulk soil. As a consequence, the rhizosphere and bulk soil were similar in microbial biomass (ninhydrin-reactive nitrogen (NHR-N)), community composition (PLFA), and enzyme activity. In the transition and mature grassland stage, more benign soil conditions stimulated microbial growth (NHR-N, total amount of PLFA, bacterial PLFA, Gram-positive bacteria, Gram-negative bacteria), and microbial diversity (Shannon index H) in the rhizosphere either directly or indirectly through enhanced carbon allocation. In the same period, the rhizosphere microflora shifted from a G(-) to a more G(+), and from a fungal to a more bacteria-dominated community. Rhizosphere beta-xylosidase, N-acetyl-beta-glucosaminidase, and sulfatase activity peaked in the mature grassland soil, whereas rhizosphere leucine aminopeptidase, beta-glucosidase, and phosphatase activity were highest in the transition stage, probably because of enhanced carbon and nutrient allocation into the rhizosphere due to better growth conditions. Soil organic matter appeared to be the most important driver of microbial colonization in the bulk soil. The decrease in soil pH and soil C/N ratio mediated the shifts in the soil microbial community composition (bacPLFA, bacPLFA/fungPLFA, G(-), G(+)/G(-)). The activities of beta-glucosidase, beta-xylosidase and phosphatase were related to soil ammonium and phosphorus, indicating that higher decomposition rates enhanced the nutrient availability in the bulk soil. We conclude that the major determinants of the microllora vary along the successional gradient: in the pioneer stage the rhizosphere microflora was primarily determined by the harsh soil environment; under more favourable environmental conditions, however, the host plant selected for a specific microbial community that was related to the dynamic interplay between soil properties and carbon supply. (C) 2004 Elsevier Ltd. All rights reserved.

Utilisation by upland rice of plant residue- and fertiliser-phosphorus in two tropical acid soils

A dual isotopic technique was used to assess the effects of soil type, and residues of Gliricidia sepium, without and with added fertiliser-P on the utilisation of P. Upland rice (Oryza sativa) was grown for 70 days in two tropical acid soils of different P sorbing capacity and P status. Uniformly P-32-labelled soils were treated with inorganic fertiliser-P tagged with P-33, Gliricidia sepium residue applied at planting and 3 weeks earlier, and in a combination of fertiliser-P and Gliricidia applied at and 3 weeks before planting. There were significant responses of shoot and root weights, and total P uptake to Gliricidia- and/or fertiliser-P addition in the Ultisol (low P status) but not the Oxisol (high P status), suggesting that P in the latter soil was not yield limiting, despite the high standard P requirement. Similarly, incorporation of Gliricidia three weeks before planting further increased shoot weight only in the Ultisol. There were generally higher proportions, quantities and percent utilisations of the Gliricidia- P and fertiliser-P in the Ultisol than in the Oxisol. Gliricidia significantly increased the utilisation of fertiliser-P only in the Ultisol. However, early application of Gliricidia increased Gliricidia- P but not fertiliser-P utilisation in the Ultisol. Added fertiliser-P did not influence Gliricidia- P utilisation.

The Representative Soil Sampling Scheme of England and Wales: the spatial variation of topsoil nutrient status and pH between 1971 and 2001

The Representative Soil Sampling Scheme (RSSS) has monitored the soil of agricultural land in England and Wales since 1969. Here we describe the first spatial analysis of the data from these surveys using geostatistics. Four years of data (1971, 1981, 1991 and 2001) were chosen to examine the nutrient (available K, Mg and P) and pH status of the soil. At each farm, four fields were sampled; however, for the earlier years, coordinates were available for the farm only and not for each field. The averaged data for each farm were used for spatial analysis and the variograms showed spatial structure even with the smaller sample size. These variograms provide a reasonable summary of the larger scale of variation identified from the data of the more intensively sampled National Soil Inventory. Maps of kriged predictions of K generally show larger values in the central and southeastern areas (above 200 mg L-1) and an increase in values in the west over time, whereas Mg is fairly stable over time. The kriged predictions of P show a decline over time, particularly in the east, and those of pH show an increase in the east over time. Disjunctive kriging was used to examine temporal changes in available P using probabilities less than given thresholds of this element. The RSSS was not designed for spatial analysis, but the results show that the data from these surveys are suitable for this purpose. The results of the spatial analysis, together with those of the statistical analyses, provide a comprehensive view of the RSSS database as a basis for monitoring the soil. These data should be taken into account when future national soil monitoring schemes are designed.

Response of organically managed grassland to available phosphorus and potassium in the soil and supplementary fertilization: field trials using grass-clover leys cut for silage

Effective use and recycling of manures together with occasional and judicious use of supplementary fertilizing materials forms the basis for management of phosphorus (P) and potassium (K) within organic farming systems. Replicated field trials were established at three sites across the UK to compare the supply of P and K to grass-clover swards cut for silage from a range of fertilizing materials, and to assess the usefulness of routine soil tests for P and K in organic farming systems. None of the fertilizing materials (farmyard manure, rock phosphate, Kali vinasse, volcanic tuff) significantly increased silage yields, nor was P offtake increased. However, farmyard manure and Kali vinasse proved effective sources of K to grass and clover in the short to medium term. Available P (measured as Olsen-P) showed no clear relationship with crop P offtake in these trials. In contrast, available K (measured by ammonium nitrate extraction) proved a useful measurement to predict K availability to crops and support K management decisions.

Zinc tolerance in wheat cultivars as affected by varying levels of phosphorus

The effect of zinc-phosphorus (Zn-P) interaction on Zn efficiency of six wheat cultivars was studied. The higher dry matter yields were observed when Zn was applied at 5 mu g g(-1) soil than with no Zn application. Phosphorus applications also increased dry matter yield up to the application of 25 mu g P g(-1) soil. The dry matter yield was significantly lower at the P rate of 250 mu g g(-1) soil. At the Zn-deficient level, the Zn-efficient cultivars had higher Zn concentrations in the shoots. Zinc concentrations in all cultivars increased when the P level in the soil was increased from 0 to 25 mu g P g(-1) soil except for the cv. Durati, in which Zn concentrations decreased with increases in P levels. However, when ZnxP interactions were investigated, it was observed that at a Zn-deficient level, Zn concentrations in the plant shoot decreased with each higher level of P, and more severe Zn deficiency was observed at P level of 250 mu g g(-1) soil.

Uncertainties in annual riverine phosphorus load estimation: Impact of load estimation methodology, sampling frequency, baseflow index and catchment population density

Models developed to identify the rates and origins of nutrient export from land to stream require an accurate assessment of the nutrient load present in the water body in order to calibrate model parameters and structure. These data are rarely available at a representative scale and in an appropriate chemical form except in research catchments. Observational errors associated with nutrient load estimates based on these data lead to a high degree of uncertainty in modelling and nutrient budgeting studies. Here, daily paired instantaneous P and flow data for 17 UK research catchments covering a total of 39 water years (WY) have been used to explore the nature and extent of the observational error associated with nutrient flux estimates based on partial fractions and infrequent sampling. The daily records were artificially decimated to create 7 stratified sampling records, 7 weekly records, and 30 monthly records from each WY and catchment. These were used to evaluate the impact of sampling frequency on load estimate uncertainty. The analysis underlines the high uncertainty of load estimates based on monthly data and individual P fractions rather than total P. Catchments with a high baseflow index and/or low population density were found to return a lower RMSE on load estimates when sampled infrequently than those with a tow baseflow index and high population density. Catchment size was not shown to be important, though a limitation of this study is that daily records may fail to capture the full range of P export behaviour in smaller catchments with flashy hydrographs, leading to an underestimate of uncertainty in Load estimates for such catchments. Further analysis of sub-daily records is needed to investigate this fully. Here, recommendations are given on load estimation methodologies for different catchment types sampled at different frequencies, and the ways in which this analysis can be used to identify observational error and uncertainty for model calibration and nutrient budgeting studies. (c) 2006 Elsevier B.V. All rights reserved.

Mitigation of diffuse phosphorus pollution during rewetting of fen peat soils: A trans-European case study

Intensive cultivation of fen peat soils (Eutric Histosols) for agricultural purposes, started in Europe about 250 years ago, resulting in decreased soil fertility, increased oxidation of peat and corresponding CO2-emissions to the atmosphere, nutrient transfer to aquatic ecosystems and losses in the total area of the former native wetlands. To prevent these negative environmental effects set-aside programs and rewetting measures were promoted in recent years. Literature results and practical experiences showed that large scale rewetting of intensively used agricultural Histosols may result in the mobilisation of phosphorus (P), its transport to adjacent surface waters and an accelerated eutrophication risk. The paper summarises results from an international European Community sponsored research project and demonstrates how results obtained at different scales and from different scientific disciplines were compiled to derive a strategy to carry out rewetting measures. A decision support system (DSS) for a hydrologically sensitive area in the Droemling catchment in north-eastern Germany was developed and is presented as a tool to regulate rewetting in order to control P release. It is demonstrated that additional laboratory experiments to identify essential processes of P release during rewetting and the site-specific management of the water table, the involvement of specific knowledge and experience of the stakeholders are necessary to develop an applicable DSS. The presented DSS is practically used to prevent freshwater resources from diffuse P pollution.

Effect of ammonium and nitrate on indigenous mycorrhizal infection, rhizosphere pH change, and phosphorus uptake by sorghum

Sorghum (Sorghum bicolor L.) plants were grown in split pots in three Rothamsted soils with different soil pH values and phosphorus (P) contents. Ammonium addition resulted in higher plant dry weight and P content than comparable nitrate treatments. The pH of soils in the rhizosphere (0.51-mm average thickness) differed from the bulk soil depending on nitrogen (N) form and level. Ammonium application resulted in a pH decrease, but nitrate application slightly increased pH. To examine the effect of rhizosphere acidification on mobilization of phosphate, 0.5 M NaHCO3 extractable phosphate was measured. The lowering rhizosphere pH enhanced the solubility of P in the soil and maybe availability of P to plants. Rhizosphere-P depletion increased with increasing ammonium supply, but when N was supplied as nitrate, P depletion was not related to increasing nitrate supply. Low P status Hoosfield soils developed mycorrhizal infection., and as a result, P inflow was increased. Geescroft soil, which initially had a high P status, did not develop mycorrhizal infection, and P inflow was much smaller and was unaffected by N treatments. Therefore, plant growth and P uptake were influenced by both rhizosphere pH and indigenous mycorrhizal infection.

Effect of mycorrhizae and pH change at the root-soil interface on phosphorus uptake by Sorghum using a rhizocylinder technique

Sorghum (Sorghum bicolor) was grown for 40 days in. rhizocylinder (a growth container which permitted access to rh zosphere and nonrhizosphere soil), in two soils of low P status. Soils were fertilized with different rates of ammonium and nitrate and supplemented with 40 mg phosphorus (P) kg(-1) and inoculated with either Glomus mosseae (Nicol. and Gerd.) or nonmycorrhizal root inoculum.. N-serve (2 mg kg(-1)) was added to prevent nitrification. At harvest, soil from around the roots was collected at distances of 0-5, 5-10, and 10-20 mm from the root core which was 35 mm diameter. Sorghum plants, with and without mycorrhiza, grew larger with NH4+ than with NO3- application. After measuring soil pH, 4 3 suspensions of the same sample were titrated against 0.01 M HCl or 0.01 M NaOH until soil pH reached the nonplanted pH level. The acid or base requirement for each sample was calculated as mmol H+ or OFF kg(-1) soil. The magnitude of liberated acid or base depended on the form and rate of nitrogen and soil type. When the plant root was either uninfected or infected with mycorrhiza., soil pH changes extended up to 5 mm from the root core surface. In both soils, ammonium as an N source resulted in lower soil pH than nitrate. Mycorrhizal (VAM) inoculation did not enhance this difference. In mycorrhizal inoculated soil, P depletion extended tip to 20 mm from the root surface. In non-VAM inoculated soil P depletion extended up to 10 mm from the root surface and remained unchanged at greater distances. In the mycorrhizal inoculated soils, the contribution of the 0-5 mm soil zone to P uptake was greater than the core soil, which reflects the hyphal contribution to P supply. Nitrogen (N) applications that caused acidification increased P uptake because of increased demand; there is no direct evidence that the increased uptake was due to acidity increasing the solubility of P although this may have been a minor effect.