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Atmospheric dust deposition can be a significant source of phosphorus (P) in some tropical forests, so information on the origins and solubility of atmospheric P is needed to understand and predict patterns of forest productivity under future climate scenarios. We characterized atmospheric dust P across a seasonal cycle in a tropical lowland rain forest on Barro Colorado Nature Monument (BCNM), Republic of Panama. We traced P sources by combining remote sensing imagery with the first measurements of stable oxygen isotopes in soluble inorganic phosphate (δ(18)OP) in dust. In addition, we measured soluble inorganic and organic P concentrations in fine (<1 μm) and coarse (>1 μm) aerosol fractions and used this data to estimate the contribution of P inputs from dust deposition to the forest P budget. Aerosol dry mass was greater in the dry season (December to April, 5.6-15.7 μg m(-3)) than the wet season (May to November, 3.1-7.1 μg m(-3)). In contrast, soluble P concentrations in the aerosols were lower in the dry season (980-1880 μg P g(-1)) than the wet season (1170-3380 μg P g(-1)). The δ(18)OP of dry-season aerosols resembled that of nearby forest soils (∼19.5‰), suggesting a local origin. In the wet season, when the Trans-Atlantic Saharan dust belt moves north close to Panama, the δ(18)OP of aerosols was considerably lower (∼15.5‰), suggesting a significant contribution of long-distance dust P transport. Using satellite retrieved aerosol optical depth (AOD) and the P concentrations in aerosols we sampled in periods when Saharan dust was evident we estimate that the monthly P input from long distance dust transport during the period with highest Saharan dust deposition is 88 ± 31 g P ha(-1) month(-1), equivalent to between 10 and 29% of the P in monthly litter fall in nearby forests. These findings have important implications for our understanding of modern nutrient budgets and the productivity of tropical forests in the region under future climate scenarios.

Foliar nitrogen to phosphorus (N:P) ratios are widely used to indicate soil nutrient availability and limitation, but the foliar ratios of woody plants have proven more complicated to interpret than ratios from whole biomass of herbaceous species. This may be related to tissues in woody species acting as nutrient reservoirs during active growth, allowing maintenance of optimal N:P ratios in recently produced, fully expanded leaves (i.e., "new" leaves, the most commonly sampled tissue). Here we address the hypothesis that N:P ratios of newly expanded leaves are less sensitive indicators of soil nutrient availability than are other tissue types in woody plants. Seedlings of five naturally established tree species were harvested from plots receiving two years of fertilizer treatments in a lowland tropical forest in the Republic of Panama. Nutrient concentrations were determined in new leaves, old leaves, stems, and roots. For stems and roots, N:P ratios increased after N addition and decreased after P addition, and trends were consistent across all five species. Older leaves also showed strong responses to N and P addition, and trends were consistent for four of five species. In comparison, overall N:P ratio responses in new leaves were more variable across species. These results indicate that the N:P ratios of stems, roots, and older leaves are more responsive indicators of soil nutrient availability than are those of new leaves. Testing the generality of this result could improve the use of tissue nutrient ratios as indices of soil nutrient availability in woody plants.

This critical review introduces a template that links phosphorus (P) sources and mobilisation processes to the delivery of P to receiving waters where deleterious impact is of concern. It therefore serves as a key introductory paper in this special issue. The entire process is described in terms of a 'P transfer continuum' to emphasise the interdisciplinary and inter-scale nature of the problem. Most knowledge to date is derived from mechanistic studies on the sources and mobilisation of P using controlled experiments that have formed the basis for mitigation strategies aimed at minimising transfer from agricultural fields. However, our ability to extrapolate this information to larger scales is limited by a poor knowledge base while new conceptual advances in the areas of complex systems and fractal dynamics indicate the limitations of past theoretical frameworks. This is compounded by the conceptual and physical separation of scientists working at different scales within the terrestrial and aquatic sciences. Multi-scaled approaches are urgently required to integrate different disciplines and provide a platform to develop mechanistic modelling frameworks, collect new data and identify critical research questions.

Phosphorus transport from agricultural soils contributes to eutrophication of fresh waters. Computer modeling can help identify agricultural areas with high potential P transport. Most models use a constant extraction coefficient (i.e., the slope of the linear regression between filterable reactive phosphorus [FRP] in runoff and soil P) to predict dissolved P release from soil to runoff, yet it is unclear how variations in soil properties, management practices, or hydrology affect extraction coefficients. We investigated published data from 17 studies that determined extraction coefficients using Mehlich-3 or Bray-1 soil P (mg kg(-1)), water-extractable soil P (mg kg(-1)), or soil P sorption saturation (%) as determined by ammonium oxalate extraction. Studies represented 31 soils with a variety of management conditions. Extraction coefficients from Mehlich-3 or Bray-1 soil P were not significantly different for 26 of 31 soils, with values ranging from 1.2 to 3.0. Extraction coefficients from water-extractable soil P were not significantly different for 17 of 20 soils, with values ranging from 6.0 to 18.3. The relationship between soil P sorption saturation and runoff FRP (microg L(-1)) was the same for all 10 soils investigated, exhibiting a split-line relationship where runoff FRP rapidly increased at P sorption saturation values greater than 12.5%. Overall, a single extraction coefficient (2.0 for Mehlich-3 P data, 11.2 for water-extractable P data, and a split-line relationship for P sorption saturation data) could be used in water quality models to approximate dissolved P release from soil to runoff for the majority of soil, hydrologic, or management conditions. A test for soil P sorption saturation may provide the most universal approximation, but only for noncalcareous soils.

Diet modification to decrease phosphorus (P) concentration in animal feeds and manures can reduce surpluses of manure P in areas of intensive animal production. We generated turkey and broiler litters from two and three flock trials, respectively, using diets that ranged from "high" to "low" in non-phytate phosphorus (NPP) and some of which contained feed additives such as phytase. Phosphorus forms in selected litters were analyzed by sequential chemical fractionation and solution (31)P nuclear magnetic resonance (NMR) spectroscopy. Selected litters were also incubated with four contrasting soils. Reducing dietary NPP and using phytase decreased total P in litters by up to 38%. Water-soluble phosphorus (WSP) in litters was decreased 21 to 44% by feeding NPP closer to animal requirement, but was not affected by phytase addition. Solution (31)P NMR spectroscopy showed that feeding NPP closer to requirement decreased orthophosphate in litters by an average of 38% and that adding phytase to feed did not increase the concentration of orthophosphate in litters. Phytase also decreased phytate P in litters by 25 to 38%, demonstrating that it increases phytate P hydrolysis. Incorporation of litters with soils at the same total P rate increased WSP in soils relative to the control; this increase was correlated to soluble P added with litters at 5 d, but not by 29 d. Changes in soil Mehlich-3 phosphorus (M3-P) were related to total P added in litter, rather than soluble P. We conclude that feeding NPP closer to requirement and using feed additives such as phytase decrease total P concentrations in litters, while having little effect on P solubility in litters and amended soils.

The availability of nitrogen (N) and phosphorus (P) to organisms is a fundamental control on the ecology of uplands in the British Isles, yet there has been relatively little consideration of the role of organic forms of these nutrients. N and P fractions in soil solutions and drainage streams were determined at monthly intervals over 1 year from Widdybank Fell, Upper Teesdale National Nature Reserve, an upland area of northern England. Nitrogen was present in inorganic and organic forms at relatively high concentrations, whereas P was mainly organic, with inorganic P concentrations near the detection limit (1 microg Pl(-1)). The highest concentrations of organic P occurred in 'pulses' during the spring. These pulses may have originated in the soil following drying and rewetting cycles, and appear to demonstrate hydrochemical connectivity between soils and streams. Seasonal changes in N:P ratio were estimated in various ways to assess when the availability of these elements was likely to influence organisms. Based on inorganic nutrients alone, marked P limitation would be expected throughout the year, but inclusion of organic nutrients suggests that some communities may become N-limited during the spring. This agrees with known seasonal changes in the biology of the cyanobacterium Rivularia, which dominates the main drainage streams. Our data highlight the importance of including organic nutrients when assessing biological nutrient limitation.

Aboveground primary productivity is widely considered to be limited by phosphorus (P) availability in lowland tropical forests and by nitrogen (N) availability in montane tropical forests. However, the extent to which this paradigm applies to belowground processes remains unresolved. We measured indices of soil microbial nutrient status in lowland, sub-montane and montane tropical forests along a natural gradient spanning 3400m in elevation in the Peruvian Andes. With increasing elevation there were marked increases in soil concentrations of total N, total P, and readily exchangeable P, but a decrease in N mineralization determined by in situ resin bags. Microbial carbon (C) and N increased with increasing elevation, but microbial C:N:P ratios were relatively constant, suggesting homeostasis. The activity of hydrolytic enzymes, which are rich in N, decreased with increasing elevation, while the ratio of enzymes involved in the acquisition of N and P increased with increasing elevation, further indicating an increase in the relative demand for N compared to P with increasing elevation. We conclude that soil microorganisms shift investment in nutrient acquisition from P to N between lowland and montane tropical forests, suggesting that different nutrients regulate soil microbial metabolism and the soil carbon balance in these ecosystems. [ABSTRACT FROM AUTHOR]

Phosphorus (P) cycles rapidly in lowland tropical forest soils, but the process have been proven difficult to quantify. Recently it was demonstrated that valuable data on soil P transformations can be derived from the natural abundance of stable oxygen isotopes in phosphate (δ18OP). Here, we measured the δ18OP of soils that had received long-term nutrient additions (P, nitrogen, and potassium) or litter manipulations in lowland tropical forest in Panama and performed controlled incubations of fresh soils amended with a single pulse of P. To detect whether δ18OP values measured in the incubations apply also for soils in the field, we examined the δ18OP values after rewetting dry soils. In the incubations, resin-P δ18OP values converged to ∼3.5‰ above the expected isotopic equilibrium with soil water. This contrasts with extra-tropical soils in which the δ18OP of resin-P matches the expected equilibrium with soil water. Identical above-equilibrium resin-P δ18OP values were also found in field soils that did not receive P additions or extra litter. We suggest that the 3.5‰ above-equilibrium δ18OP values reflect a steady state between microbial uptake of phosphate (which enriches the remaining phosphate with the heavier isotopologues) and the release of isotopically equilibrated cell internal phosphate back to the soil. We also found that soil nutrient status affected the microbial turnover rate because in soils that had received chronic P addition, the original δ18OP signature of the fertilizer was preserved for at least eight weeks, indicating that the off-equilibrium δ18OP values produced during microbial phosphate turnover was not imprinted in these soils. Overall, our results demonstrate that ongoing microbial turnover of phosphate mediates its biological availability in lowland tropical soils. [ABSTRACT FROM AUTHOR]

Phosphorus (P) speciation in 21 basaltic and four non-basaltic Irish grassland soils was determined by NaOH–EDTA extraction and 31P NMR spectroscopy. Organic P in basaltic soils ranged between 30 and 697 mg P kg−1 and consisted of phosphate monoesters (84–100%), DNA (0–16%) and phosphonates (0–5%). Inorganic P was mainly phosphate (83–100%) with small concentrations of pyrophosphate (0–17%). Phosphate monoesters were more important as a proportion of extracted P in basaltic soils, probably because of their greater oxalate-extractable Fe and Al contents. Phosphate monoesters appeared to be strongly associated with non-crystalline Al and increased with total soil P concentration, indicating that they do accumulate in grassland soils. In non-basaltic soils myo-inositol hexakisphosphate constituted between 20 and 52% of organic P, while scyllo-inositol hexakisphosphate constituted between 12 and 17%. These compounds were not quantified separately in basaltic soils because of poor NMR resolution in the phosphate monoester region, but appeared to represent a considerable proportion of the organic P in most samples. DNA concentrations were greater in basaltic soils compared with non-basaltic soils and were associated with acidic pH and large total C contents. The inability of the Olsen P test to assess effectively the P status of basaltic soils may result from strong phosphate sorption to Fe and Al oxides, inducing plant utilization of soil organic P. Phosphorus nutrient management should account for this to avoid over-application of P and associated financial and environmental costs. [ABSTRACT FROM AUTHOR]

The extraction of soil organic phosphorus by the NaOH–EDTA procedure was assessed in detail for a tropical forest soil (clay-loam, pH 4.3, total carbon 2.7%). Optimum conditions for the quantification of soil organic phosphorus and characterization of its composition by solution 31P NMR spectroscopy were extraction in a solution containing 0.25 m NaOH and 50 mm Na2EDTA in a 1:20 solid to solution ratio for 4 hours at ambient laboratory temperature. Replicate analyses yielded a coefficient of variation of 3% for organic phosphorus as a proportion of the spectral area. There was no significant difference in total phosphorus extraction from fresh and air-dried soil, although slightly more organic phosphorus and less paramagnetic ions were extracted from dried soil. The procedure was not improved by changing the concentration of NaOH or EDTA, extraction time, or solid to solution ratio. Pre-extraction with HCl or Na2EDTA did not increase subsequent organic phosphorus extraction in NaOH–EDTA or improve spectral resolution in solution 31P NMR spectroscopy. Post-extraction treatment with Chelex resin did not improve spectral resolution, but removed small concentrations of phosphorus from the extracts. Increasing the pH of NaOH–EDTA extracts (up to 1.0 m NaOH) increased the concentration of phosphate monoesters, but decreased DNA to an undetectable level, indicating its hydrolysis in strong alkali. The standardized NaOH–EDTA extraction procedure is therefore recommended for the analysis of organic phosphorus in tropical forest soils. [ABSTRACT FROM AUTHOR]

Phosphorus (P) deficiency is a major limitation to agricultural production in many parts of the world. It is therefore desirable to identify plants with enhanced abilities to utilize P more efficiently. Exudation of phosphatase from roots may improve P availability, yet there is little direct evidence for this. Here we report the dynamics of organic P in the rhizosphere of plants that have enhanced rhizosphere phosphatase activity. Agroforestry species and transgenic subterranean clover (engineered to produce phytase) were compared with crop and wild-type plant controls, respectively. Depletion of organic P was measured in pools defined by chemical extraction, solution 31P NMR spectroscopy, and microbial immobilization of radio-isotopic P. Plants that had greater extracellular phosphatase activity depleted more organic P from P-deficient Oxisols than control plants. Depleted organic P forms were primarily phosphate monoesters. Plants with enhanced extracellular phosphatase activity also had access to a pool of soil P that was less isotopically exchangeable. Transgenic subterranean clover that expresses a microbial phytase gene appeared to have greater access to recently immobilized P, whereas plants expressing endogenous phosphatases utilized the unlabelled portion of soil organic P to a greater extent. Collectively, these results indicate that the enhancement of phosphatase activity in the rhizosphere of plants is implicated in the depletion of organic P forms from soils, most notably orthophosphate monoesters, whilst also suggesting that there is some exclusivity to the pools of organic P utilized by plants and microorganisms. [ABSTRACT FROM AUTHOR]

Phosphorus (P) transfer from agricultural land to surface waters cancontribute to eutrophication, excess algal growth and associated water quality problems. Grasslands have a high potential for P transfer,as they receive P inputs as mineral fertilizer and concentrates cycled through livestock manures. The transfer of P can occur through surface and subsurface pathways, although the capacity of most soils to fix inorganic P has meant that subsurface P transfer by leaching mechanisms has often been perceived as negligible. We investigated this using large-scale monolith lysimeters (135 cm deep, 80 cm diameter) tomonitor leachate P under four grassland soil types. Leachate was collected during the 1997-98 drainage year and analysed for a range of Pfractions. Mean concentrations of total P routinely exceeded 100 mu g l-1 from all soil types and, therefore, exceeded P concentrations above which eutrophication and algal growth can occur. The majority of the leachate P was in algal-available Mo-reactive (inorganic) forms, although a large proportion occurred in unreactive (organic) forms. We suggest that subsurface transfer by leaching can represent a significant mechanism for agricultural P transfer from some soils and must be given greater consideration as a potential source of diffuse P pollution to surface waters. [ABSTRACT FROM AUTHOR]