Sources and bioavailability of phosphorus fractions in freshwaters: a British perspective

This paper seeks a perspective on the forms of phosphorus which promote aquatic eutrophication, with the particular quest of establishing their sources. A short background traces the development of understanding of nutrient enrichment and the suppositions about the relative contributions of agriculture, sewage and detergent residues. Most aquatic systems, and their primary producers, are naturally deficient in biologically-available phosphorus. Aquatic plants have evolved very efficient phosphorus uptake mechanisms. The biomass responses to an increase in the supply of phosphorus are stoichiometrically predictable. The most bioavailable forms of phosphorus are in solution, as orthophosphate ions, or are readily soluble or elutable from loose combinations. Ready bioavailability coincides well with what is measurable as molybdate-reactive (MRP) or soluble-reactive phosphorus (SRP). Most other forms, including phosphates of the alkaline earth metals, aluminium and iron are scarcely available at all. Orthophosphate ions sorbed to metal oxides and hydroxides are normally not biologically available either, except through weak dissociation ('desorption'). The production of alkaline phosphatase provides organisms with an additional mechanism for accelerating the sequestration of phosphate from organic compounds. Bioavailable phosphate is liberated when redox- or alkali-sensitive metal hydroxides dissolve but these processes are minor contributors to the biological responses to nutrient enrichment. Most of the familiar eutrophication is attributable to the widespread application of secondary sewage treatment methods to the wastes emanating from a burgeoning and increasingly urbanised human population. The use of polyphosphate-based detergents, now in decline, has contributed to the problem. In aquatic systems, the additional phosphorus raises the biological supportive capacity, sometimes to the capacity of the next limiting factor (carbon, light, hydraulic retention or of another nutrient). At high orthophosphate loadings, the straight stoichiometric yield relationship between biomass yield and phosphorus a vailability is lost. Movements of phosphorus and its recycling within aquatic systems do not prevent the slow gravitation of phosphorus to the bottom substrata. The phosphorus retentivity of sediments depends upon their chemical composition. While oxide-hydroxide binding capacity in the surface sediments persists, they act as a sink for phosphorus and a control on further cycling. Iron-rich and clay-rich sediments perform best in these conditions; calcareous sediments least so. Eutrophication may lead to the exhaustion of sediment P-binding capacity. Non-sorbed phosphate is readily recyclable if primary producers have access to it. Recycling is most rapid in shallow waters (where sediment disturbance, by flow, by wind action and through bioturbation, is frequent and least in deep ventilated sediments. The contributions of phosphorus from catchments are assessed. The slow rate of weathering of (mostly apatitic) minerals, the role of chemical binding in soils and the incorporation and retentivity bv forested terrestrial ecosystems each contribute to the minimisation of phosphorus leakage to drainage waters. Palaeolimnological and experimental evidence confirms that clearance of land and ploughing its surface weakens the phosphorus retentivity of catchments. The phosphorus transferred from arable land to drainage remains dominated by sorbed fractions which are scarcely bioavailable. Some forms of intensive market gardening or concentrated stock rearing may mobilise phosphates to drainage but it is deduced that drainage from agricultural land is not commonly a major source of readily bioavailable phosphorus in water. Careful budgeting of the phosphates in run-off from over-fertilised soils may nevertheless show that a proportionately small loss of bioavailable phosphorus can still be highly significant in promoting aquatic plant production. The bioavailable-phosphorus (BAP) load achieving the OECD threshold of lake eutrophy (35 mg P m(-3)) is calculated to be equivalent to a terrestrial loss rate of approximately 17.5 kg BAP km(-2) year(-1)), or only 1-2% of a typical fertiliser application. The output is shown to be comparable with the P yield from secondary treatment of the sewage produced by a resident population of 30-44 persons km(-2). With tertiary treatment, the equivalence is with approximately 200 persons km(-2).     

Maize root system growth and development as influenced by phosphorus deficiency

Effects on leaf growth, biomass accumulation and root morphogenesis associated with the establishment of phosphorus (P) deficiency were studied on maize in order to test the hypothesis that the root system response can be accounted for by the effect of P deficiency on the carbon budget of the plant. P deprivation had a large and rapid negative effect on leaf expansion. For 7 d after P deprivation, the total dry matter production per plant was almost fully accounted for by the effect of P starvation on leaf growth and its subsequent effect on photosynthetically active radiation (PAR) interception. No strong effect of P deficiency was observed on the radiation use efficiency during this first period, although it was reduced thereafter. Root growth was slightly enhanced a few days after P starvation, but strongly reduced thereafter. The elongation rate of axile roots was maintained throughout the experiment, whereas emergence of new axile roots and elongation of first-order laterals were drastically reduced. The density of first-order laterals was not severely affected. These morphological responses are very similar to what is observed when root growth is limited by the availability in carbohydrates. The results are therefore compatible with the hypothesis that P deficiency mainly affects the root system morphology through its effect on the carbon budget of the plant with no additional specific effect of P deficiency on root morphogenesis. The drastic and early reduction of shoot growth after P deprivation may explain that more carbohydrates were available for root growth which was observed a few days after P starvation and reported by several authors. Later on, however, because of the reduced leaf area of P-deprived plants, their capacity to intercept light was severely reduced so that root growth was finally reduced.Keywords: Zea mays L., maize, phosphorus, root, root morphogenesis.      

Soil phosphorus fractions and adsorption as affected by organic and inorganic sources

The effect of organic and inorganic sources of phosphorus (P) on soil P fractions and P adsorption was studied in a field without plant growth on a Kandiudalf in western Kenya. A high-quality organic source, Tithonia diversifolia (Hemsley) A. Gray leaves, and a low-quality source, maize (Zea mays L.) stover, were applied alone or in combination with triple superphosphate (TSP). The P rate was kept constant at 15 kg P ha^-1. Soil extractable P (resin, bicarbonate and sodium hydroxide), microbial biomass P and C and P adsorption isotherms were determined during 16 weeks after application of treatments. Application of tithonia either alone or with TSP increased resin P, bicarbonate P, microbial P, and sodium hydroxide inorganic P. Tithonia alone reduced P adsorption at 2–16 weeks. Maize stover had no effect on any of the P fractions or P adsorption. At 8 weeks, the application of tithonia reduced microbial C-to-P ratio (20) as compared to maize stover, TSP and the control (31–34). The reduction in P adsorption by tithonia was accompanied by increases in all measured P fractions, the sum of P in those fractions (resin, bicarbonate and sodium hydroxide) being larger than the P added. The reduction in P adsorption apparently resulted from competition for adsorption sites, probably by organic anions produced during decomposition of the high quality tithonia. Integration of inorganic P (TSP) with organic materials had little added benefit compared to sole application of TSP, except that combination of tithonia with TSP increased microbial biomass. The results indicate that a high quality organic input can be comparable to or more effective than inorganic P in increasing P availability in the soil.     

Growth and nutrient status of citrus plants as influenced by mycorrhiza and phosphorus application

To test the hypothesis that high levels of soluble phosphate applied in combination with VAM fungi, to citrus plants, can cause growth depression even in the absence of other limiting factors, and also to test if rock phosphate, under these conditions, may be a satisfactory P source, a greenhouse experiment was conducted using sterilized soil with four levels of phosphate (0, 50, 100 and 200 ppm P) supplied either as soluble P or as rock phosphate. Citrus seedlings were either inoculated with the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus etunicatum or left uninoculated. Six months after the start of the experiment, the plants were harvested and shoot dry weight, P and K uptake, root colonization and the number of spores in 50 cm³ of soil were determined. Significant increases were found in dry matter yields and in P and K contents, due to VAM fungus inoculation, at the zero and 50 ppm soluble P levels and at all rock phosphate levels. At 100 ppm soluble P, the development of VAM plants was equilvalent to that of non-VAM plants, and at 200 ppm, growth was significantly less than that of non-VAM plants. Root colonization and sporulation were reduced at higher P levels. The absolute growth depression of VAM plants at the higher P level was likely due to P toxicity. In addition, high leaf P and K concentrations may have interfered with carbohydrate distribution and utilization in these symbioses. Rock phosphate may be used with VAM citrus to substitute for medium amounts of soluble phosphate.     

Functional groups' performances as influenced by nitrogen,phosphorus and nodule inhibition of legumes

Aims Nitrogen (N) and phosphorus (P) constitute essential elements for plant growth and their availability influence species diversity in herbaceous plant communities. Legumes exhibit relatively high abundance in N-limited soils. Moreover, the legumes' N:P ratios are much higher than those of the other plant species grown in the same site, probably because they are able to fix atmospheric N 2. The objective of this study was to determine how the relative proportion in N and P availability and the restriction of legumes to fix atmospheric N 2 affect: (i) the primary productivity of plant species, (ii) species composition and (iii) N and P concentrations of species.Methods In an outdoor experiment, mixtures containing grasses, legumes and non-legume forbs were established in 32 containers under four soil treatments (control, N addition, P addition and disinfected soil), in a completely randomized design with eight replicates. Plant growth was examined when N and P were limited in the control soil:sand mixture, in a pot experiment sown with Plantago lanceolata .Important findings The pot experiment indicated that both N and P were limiting for the growth of P. lanceolata. Soil treatments affected primary productivity and species composition. Legumes had a relatively high abundance in the control and their growth was favoured, especially that of Medicago sativa, by P addition. Grasses' growth was increased by the addition of N. Inhibition of rhizobia resulted in poor growth of legumes and concomitant higher growth of grasses, in comparison to the control. The N:P ratios of non-legume species differed between treatments and were always higher in the legume species, even in the disinfected soil. The latter provides evidence that the high N concentrations found in legumes are a physiological characteristic of this specific group of plants.     

Recovery of adsorbed zinc in acid soils as influenced by prior phosphorus applications

Summary Five acid soils of Hawaii, having histories of heavy P applications were equilibrated with graded quantities of Zn. Amounts of adsorbed Zn were extracted with a single extraction of 0.005M DTPA. The data indicated that most of the added Zn was in available form. Prior P applications either had no effect on recovery or slightly increased it. The results substantiated the earlier findings that P-induced Zn deficiency could not be due to precipitation of Zn as insoluble Zn–P compounds in the soils.     

Potassium and phosphorus uptake by corn genotypes grown in the field as influenced by root characteristics

Summary Root parameters of three corn (Zea mays L.) genotypes influencing P and K uptake were investigated in solution culture and field experiments. The data for these parameters were used to simulate P and K uptake by plants grown in the field using the Claassen-Barber model⁵. Root characteristics for ion influx, maximum rate of influx,Imax; Michaelis-Menten constant,Km; and minimum concentration of solution below which no further net influx occurs,Cmin were determined in solution culture. These kinetic parameters varied 2 to 3 fold among genotypes. Variations among genotypes were different for K than for P.Three corn genotypes were grown in the field and harvested 47, 54 and 68 days after emergence. Yield and root surface per plant increased about 3 fold during this time. At 47 days, 2/3 of the total root surface was in the top soil whereas 3 weeks later, it was less than 50%. Genotypes differed in distribution of roots between the topsoil and subsoil as well as in root surface per unit of shoot.K uptake predicted by the Claassen-Barber model was 2 to 3 times the observed. The overprediction could be related to high root density (length of root per unit soil volume) which indicated that competition between roots occurred that was not considered in the simulation model. The predicted P uptake (y) was correlated (r=0.91) to observed uptake (x) byy=0.98+0.67x, indicating underprediction of P uptake. The presence of root hairs may have been the cause of the underprediction. The calculated contribution of the subsoil to the observed uptake was 10% for K and 1% in the case of P. It was concluded that the plant parameters used to simulate nutrient uptake were rated accurately when allowance was made for root competition and presence of root hairs.Journal Paper No. 7608. Purdue University, Agric. Exp. Station, West Lafayette, IN 47907. Contribution from the Department of Agronomy. This research was supported in part by the Tennessee Valley Authority and the Deutsche Forschungsgemeinschaft.     

Balance sheet of soil phosphorus and potassium as influenced by intensive cropping and fertilizer use

Summary The balance sheet of phosphorus and potassium were worked out from a long term manure and fertilizer experiment conducted for eight years and is still continuing at Ranchi Agricultural College, Kanke, Ranchi, India. Increasing levels of fertilizer combination with organic manure and lime give the highest yield as well as removed the highest amount of phosphorus and potassium from the soil and gave the positive gain of soil phosphorus and potassium in intensive cropping. The highest gains of 59.0 and 278.0 kg/ha of phosphorus and potassium respectively were recorded in 150% NPK application.     

Modulation of dissolved oxygen levels in a hypertidal estuary by sediment resuspension

Hyperconcentrated benthic layers, which form during neap tides, recruit much of the fine sediment population of the turbidity maximum of a hypertidal estuary. Measurements of tidal amplitude and suspended solids concentration reveal that resuspension of the hyperconcentrated layers occurs between three and eight tides after neap tides rather than during spring tides (12 to 15 tides after neaps). During these resuspension events, dissolved oxygen levels are reduced but recover by spring tides. Peak solids concentrations and critically depressed dissolved oxygen levels are out of phase with tidal current amplitude. Thus observations close to neap and spring tides do not necessaraly capture the extremes of the envelope of water quality conditions.     

Ozone effects on growth of radish plants as influenced by nitrogen and phosphorus nutrition and by temperature

Summary Raphanus sativus L. (radish) plants were grown in sand culture at two temperatures and fed with nutrient solutions containing relatively low or high levels of either N or P. At the 4-leaf stage, the plants were exposed to ozone at a concentration of 25 pphm for 4 h. Ozone treatments resulted in decreased dry weight of low- and high-N plants at both temperatures and of low and high P plants only at the lower temperature. The study showed that air pollutant growth reduction is not necessarily accentuated by luxuriant growth resulting from high nutritional status. Responses to the nutrition of specific mineral nutrients depend on the modifying effect of temperature. re]19720524     

The physics of sediment transport,resuspension, and deposition

The physics of sediment transport, resuspension and deposition is reviewed. First, the general problem is described and then emphasis is given to recent quantitative research in Lakes Ontario, Michigan and Erie. In this discussion, some original calculations are presented to show that Ekman layer sediment transport is important in determining the deposition areas in deep lakes.GLERL Contribution No. 436     

Salt hardiness and dye reduction by potato tissue and mitochondrial fractions as influenced by previous storage of the tubers

Oxygen uptake and tetrazolium reduction occurred at higher rates in discs from potato tubers (Solanum tuberosum L.) stored at 0° than in discs from tubers stored at 12.8°. Tetrazolium reduction was at a higher rate in mitochondrial fractions from tubers stored at 0° than in mitochondrial fractions from tubers stored at 12.8°. These physiological activities were more resistant to hypertonic KCl treatments in tissue and mitochondrial fractions from tubers stored at 0° than in tissue and mitochondrial fractions from tubers stored at 12.8°. Inhibition of O₂ uptake and tetrazolium reduction progressively increased with increasing concentrations of KCl for tissue and mitochondrial fractions from tubers stored at 0 and 12.8°, but inhibition was more severe and occurred at lower concentrations of KCl for the material from tubers stored at 12.8°. Tissue from tubers stored at 0° was at the same time more sensitive to hypotonic solutions and more resistant to hypertonic solutions than corresponding tissue from tubers stored at 12.8°. Adaptive changes brought on in the tubers by the stress of cold storage were demonstrated in the discs and mitochondrial fractions prepared from cold-stored tubers.     

Possibilities and limitations of sediment traps to measure sedimentation and resuspension

Settling flux of particulate matter measured by sediment traps in shallow and turbulent waters considerably exceeds settling flux calculated from mass balances. Theoretical consideration on sinking distance and particle settling leads to the conclusion that traps overtrap particles in even moderately turbulent waters. The disappearance of turbulent diffusion and bottom shear stress within the traps causes the high rates of trapping, whereas their presence in natural water bodies hampers sedimentation.The use of sediment traps may be restricted to the identification of rapidly sinking particles, measurement of sinking velocities of well defined types of particles, and estimation of resuspension.     

外源磷对苗期小麦和水稻根际砷形态及其生物有效性的影响

采用土(中度砷污染土)-土根袋培养的方法研究了两个浓度的外源磷(P)对苗期小麦和水稻根际砷(As)形态分布及其生物有效性的影响.结果表明:(1)两种作物生长的土壤中各砷形态的分配比例依次为:结晶铁锰或铁铝水化氧化物结合态(45%～52%)＞无定形和弱结晶铁铝或铁锰水化氧化物结合态(26%～34%)＞专性吸附态(12%～14%)＞残渣态(4%～7%)＞非专性吸附态(0.09%～0.25%).(2)添加外源磷浓度为100 mg·kg-1与不施磷处理相比显著提高了两种作物地上部的生物量(p＜0.01).(3)苗期小麦在添加100 mg·kg-1外源磷时,不仅促进了作物生长而且抑制根中砷向地上部的转运.(4)任何磷处理下,水稻对砷的吸收能力以及由根系向地上部转移能力均高于小麦.因此,在轻中度砷污染土壤上与水稻相比更适宜种植小麦(或其他旱作植物);而在水稻种植季,可以通过添加适量磷肥(100 mg·kg-1)来减弱砷在水稻体内的累积.     

Shear stress and sediment resuspension in relation to submersed macrophyte biomass

We examined the impacts of macrophyte beds dominated by a canopy-forming (Myriophyllum sibiricum) and a meadow-forming (Chara canescens) species on bottom shear stress (τ) and resuspension in shallow Lake Christina, Minnesota (U.S.A.). Studies were conducted in late summer, 1998, when macrophyte biomass levels exceeded 200 g m^?2, and in early summer, 2000, when biomass was greatly reduced (<20 g m^?2) in both plant beds. The critical shear stress (τ_c) of sediments, measured experimentally in the laboratory, was low (1.4 dynes cm^?2) indicating potential for resuspension in the absence of macrophytes. During 1998, turbidity was low at the M. sibiricum and Chara station, rarely increasing when calculated bottom τ (calculated from wave theory assuming no biomass obstruction) exceeded τsub c sub, indicating that both beds reduced sediment resuspension at high biomass levels. In situτ (estimated τ), measured via gypsum sphere dissolution, did not exceed τ_c above the sediment interface in either bed during 1998. In contrast, sediment resuspension occurred in both beds during similar high winds in 2000. However, estimated τ was lower than calculated bottom τ, suggesting that at low biomass, macrophytes were having some impact on τ.     

Tolerance of cotton to Meloidogyne incognita as influenced by phosphorus fertiliser and a vesicular arbuscular mycorrhizal fungus

Cotton was grown in soils of differing phosphorus levels with and without a mycorrhizal fungus and with a range (0 to 3200 eggs/ 100 c^-3 soil) of Meloidogyne incognita eggs added. Tolerance of cotton to M. incognita did not differ between plants grown in different soils with or without the mycorrhizal fungus, though plant size could differ markedly. Plant tolerance was increased by a delay in adding nematodes.