Extracellular secretion of Aspergillus phytase from Arabidopsis roots enables plants to obtain phosphorus from phytate

Phosphorus (P) deficiency in soil is a major constraint for agricultural production worldwide. Despite this, most soils contain significant amounts of total soil P that occurs in inorganic and organic fractions and accumulates with phosphorus fertilization. A major component of soil organic phosphorus occurs as phytate. We show that when grown in agar under sterile conditions, Arabidopsis thaliana plants are able to obtain phosphorus from a range of organic phosphorus substrates that would be expected to occur in soil, but have only limited ability to obtain phosphorus directly from phytate. In wild-type plants, phytase constituted less than 0.8% of the total acid phosphomonoesterase activity of root extracts and was not detectable as an extracellular enzyme. By comparison, the growth and phosphorus nutrition of Arabidopsis plants supplied with phytate was improved significantly when the phytase gene (phyA) from Aspergillus niger was introduced. The Aspergillus phytase was only effective when secreted as an extracellular enzyme by inclusion of the signal peptide sequence from the carrot extensin (ex) gene. A 20-fold increase in total root phytase activity in transgenic lines expressing ex::phyA resulted in improved phosphorus nutrition, such that the growth and phosphorus content of the plants was equivalent to control plants supplied with inorganic phosphate. These results show that extracellular phytase activity of plant roots is a significant factor in the utilization of phosphorus from phytate and indicate that opportunity exists for using gene technology to improve the ability of plants to utilize accumulated forms of soil organic phosphorus.     

An examination of some possible sources of soil inositol phosphates

Summary Examination of extracts prepared from soil fungi and yeast cells (S. cavlsbergensis) showed that phytate phosphorus was not present. Phytin was successfully extracted from the acorns ofQuercus robur andQ. hodgkinsonii but was apparently absent from acorns ofQ. ilex. Acorn phytin was shown to consist only of phosphate esters ofmyo-inositol although freescyllo-inositol is present in this plant tissue. Small amounts of phytate phosphorus were isolated from sand/clay cultures, after a period of incubation, but onlymyo-inositol derivatives appeared to be present.     

Expression of a fungal phytase gene in Nicotiana tabacum improves phosphorus nutrition of plants grown in amended soils

Transgenic Nicotiana tabacum plants expressing a chimeric phytase gene (ex::phyA) from the soil fungus Aspergillus niger were generated. Three independently transformed lines showed increased extracellular phytase activity compared with a vector control and wild-type plants, both of which had no detectable extracellular phytase. Transgenic N. tabacum plants grown in sterile agar supplied with phosphorus (P) as phytate accumulated 3.7-fold more P than vector control plants. Despite this, the expression of ex::phyA in plants did not lead to an improved accumulation of P from two unamended P-deficient soils. However, when soils were amended with either phytate or phosphate and lime, transgenic plants accumulated up to 52% more P than controls. Positive responses by transgenic plants were, in some instances, coincident with a putative increase in soil phytate. We conclude that the development of plants that exude phytase to the soil may not ensure improved plant P nutrition, as the availability of phytate in the soil also appears to be critical. Nevertheless, if plants that express ex::phyA are combined with soil amendments that promote the availability of phytate, there is the potential to enhance the P nutrition of crop plants and to improve the efficiency of P fertilizer use in agricultural systems.     

Assessment of functional diversity and structure of phytate-hydrolysing bacterial community in <Emphasis Type="Italic">Lolium perenne</Emphasis> rhizosphere

Background and aims

Plant growth is frequently limited by the availability of inorganic phosphorus (P) in the soil. In most soils, a considerable amount of the soil P is bound to organic molecules. Of these, phytate is the most abundant identifiable organic P form, but is not readily available to plants. In contrast, microorganisms have been shown to degrade phytate with high efficiency. The current study aims to characterize the members of the phytate-hydrolysing bacterial community in rhizosphere, and the molecular and enzymatic ability of these bacteria to degrade phytate.

Methods and results

The phytate-hydrolysing bacterial community was characterized from the rhizosphere of plants cultivated in the presence or absence of phytate supplementation. Major changes in the bacterial community structure were observed with both culture-dependent and -independent methods, which highlighted the predominance of Proteobacteria and Actinobacteria. Phytase activity was detected for a range of rhizobacterial isolates as well as the presence of, β-propeller phytases (BPP) for both isolates and directly in a soil sample.

Conclusion

A wide taxonomic range of functional phytate utilizers have been discovered, in soil bacterial taxa that were previously not well known for their ability to utilise phytate as P or C sources. This study provides new insights into microbial carbon and phosphorus cycling in soil.

     

Effect of vesicular-arbuscular mycorrhizal and phosphatase-producing fungal inoculation on growth and nutrition of white clover supplied with organic phosphorus

In a pot experiment white clover was grown for 50 d in two sterilized low phosphorus soils supplied with organic phosphorus as sodium phytate. Seed inoculation withAspergillus fumigatus, a phosphatase-producing fungus (PPF), and soil inoculation withGlomus mossece, a vesicular-arbuscular mycorrhizal (VAM) fungus resulted in increased shoot and root dry mass and root length, phosphatase activity in soil and shoot concentrations of P and to a lesser extent of Mg. The effects on these parameters were most distinct upon combined inoculation treatment (PPF+VAM). A significant increase in hyphal length density and decrease in Mn concentration was observed only upon PPF+VAM treatment. Shoot concentrations of Cu and Zn were highly enhanced by VAM but not by PPF. Of the total P uptake the PPF contribution accounted for 39–41 %, VAM contributed 41–50 % and their combination resulted in 55–58 %, in comparison to only 38–40 % contribution due to soil microorganisms under unsterilized conditions. The depletion of organic phosphorus in the rhizosphere soil increased in the order (PPF+VAM)>VAM>PPF> unsterilized soil>sterilized soil. The results demonstrate the efficient use of phytate phosphorus byA. fumigatus andG. mosseœ and suggest that dual inoculation is superior to a single one.     

Ability of <Emphasis Type="Italic">Emericella rugulosa</Emphasis> to mobilize unavailable P compounds during Pearl millet [<Emphasis Type="Italic">Pennisetum glaucum</Emphasis> (L.) R. Br.] crop under arid condition

Phosphate solubilizing microorganisms are ubiquitous in soils and could play an important role in supplying P to plants where plant unavailable P content in soil was more. A phosphatase and phytase producing fungus Emericella rugulosa was isolated and tested under field condition (Pearl millet as a test crop) in a loamy sand soil. In the experimental soil 68% organic phosphorous was present as phytin; less than 1% of phosphorous was present in a plant available form. The maximum effect of inoculation on different enzyme activities (acid phosphatase, alkaline phosphatase, phytase, and dehydrogenase) was observed between 5 and 8 weeks of plant age. The depletion of organic P was much higher than mineral and phytin P. The microbial contribution was significantly higher than the plant contribution to the hydrolysis of the different P fractions. A significant improvement in plant biomass, root length, seed and straw yield and P concentration of root and shoot resulted from inoculation. The results suggest that Emericella rugulosa produces phosphatases and phytase, which mobilize P and enhance the production of pearl millet.     

Novel Glucose-1-Phosphatase with High Phytase Activity and Unusual Metal Ion Activation from Soil Bacterium Pantoea sp. Strain 3.5.1

Phosphorus is an important macronutrient, but its availability in soil is limited. Many soil microorganisms improve the bioavailability of phosphate by releasing it from various organic compounds, including phytate. To investigate the diversity of phytate-hydrolyzing bacteria in soil, we sampled soils of various ecological habitats, including forest, private homesteads, large agricultural complexes, and urban landscapes. Bacterial isolate Pantoea sp. strain 3.5.1 with the highest level of phytase activity was isolated from forest soil and investigated further. The Pantoea sp. 3.5.1 agpP gene encoding a novel glucose-1-phosphatase with high phytase activity was identified, and the corresponding protein was purified to apparent homogeneity, sequenced by mass spectroscopy, and biochemically characterized. The AgpP enzyme exhibits maximum activity and stability at pH 4.5 and at 37°C. The enzyme belongs to a group of histidine acid phosphatases and has the lowest K_m values toward phytate, glucose-6-phosphate, and glucose-1-phosphate. Unexpectedly, stimulation of enzymatic activity by several divalent metal ions was observed for the AgpP enzyme. High-performance liquid chromatography (HPLC) and high-performance ion chromatography (HPIC) analyses of phytate hydrolysis products identify dl-myo-inositol 1,2,4,5,6-pentakisphosphate as the final product of the reaction, indicating that the Pantoea sp. AgpP glucose-1-phosphatase can be classified as a 3-phytase. The identification of the Pantoea sp. AgpP phytase and its unusual regulation by metal ions highlight the remarkable diversity of phosphorus metabolism regulation in soil bacteria. Furthermore, our data indicate that natural forest soils harbor rich reservoirs of novel phytate-hydrolyzing enzymes with unique biochemical features.     

Phytate utilization of maize mediated by different nitrogen forms in a plant–arbuscular mycorrhizal fungus–phosphate-solubilizing bacterium system

A pot experiment was conducted to investigate the organic phosphorus (P) (phytate) utilization of Zea mays L. with different nitrogen (N) forms (NH₄⁺ and NO₃^?) when both arbuscular mycorrhizal (AM) fungus (Funelliformis mosseae) and phosphate-solubilizing bacterium (PSB, Pseudomonas alcaligenes) are present. The soil was supplied with either KNO₃ or (NH₄)₂SO₄ (200 mg kg^?1 N) with or without phytin (75 mg P kg^?1). Results showed that the application of NH₄⁺ to the soil in a plant–AM fungus–PSB system decreased rhizosphere pH and increased phosphatase activity. It also enhanced the mineralization rate of phytin, which resulted in the release of more inorganic P. The application of NO₃^? promoted mycorrhizal colonization and hyphal length density in the soil. The inorganic P in the hyphosphere decreased, but more P was transferred to the plant through the mycorrhizal hyphae. Hence, in addition, the application of the two different N forms did not significantly alter the content of plant P. The plant supplied with different N fertilizers acquired P through different mechanisms associated with other microbes. NH₄⁺ application promoted phytin mineralization by decreasing soil pH, whereas NO₃^? application increased inorganic P uptake by strengthening the mycorrhizal pathway.     

The effect of phytase on the availability of P from myo-inositol hexaphosphate (phytate) for maize roots

The effect of adding phytase to the root medium of maize plants on the P-availability of added myo-inositol hexaphosphate (phytin) has been studied in pot experiments. When 40 mM phytin-P in nutrient solution was incubated in quartz-sand for 15 days in the absence of plants, 80% of it could be recovered from the solution as soluble organic P. Maize plants growing on this mixture assimilated P from phytin at rates comparable to those from inorganic phosphate (Pi). At a lower addition rate (2 mM phytin-P) only 10% was recovered in the soil solution, and plant growth was severely limited by P. At this low phytin level, the addition of phytase (10 enzyme units per kg sand) increased the plants' dry weight yield by 32%. The relative increases of the Pi concentration in the solution and of the amount of P in the plants were even higher, indicating that the observed growth stimulation was due to an increased rate of phytin hydrolysis. The enzyme-induced growth stimulation was also observed with plants growing in pots filled with soil low in P, when phytin was added. However, on three different soils the addition rates of phytin and phytase necessary for obtaining a significant phytase effect were both about 10 times higher than those required in quartzsand. It is concluded that the P-availability from organic sources can be limited by the rate of their hydrolytic cleavage.Abbreviation Pi inorganic phosphate     

Protein Body Composition in Cuburbita maxima Cotyledons as Determined by Energy Dispersive X-Ray Analysis

Energy dispersive x-ray analysis was used to study the composition of certain protein body components in Cucurbita maxima cotyledons. The globoid crystal was rich in phosphorus, potassium, and magnesium. This elemental composition provides further evidence that the globoid crystal in squash cotyledon protein bodies is composed of phytin, a myoinositol hexaphosphoric acid salt of potassium and magnesium. Calcium, a common component of phytin in many species, was absent or present in only trace amounts in the globoid crystals of squash. Results of analyses of globoid crystals from seeds produced in different parts of North America suggest that there is definite specificity for the cations used in phytin deposition. Variations in soil types and other environmental factors seem not to have influenced the type of cation stored. Energy dispersive x-ray analysis of the proteinaceous regions revealed the presence of phosphorus, sulfur, and a trace of chlorine. Sulfur was expected, due to the presence of some sulfur containing amino acids in the protein.     

Assimilation of Phytate-phosphorus by the Extracellular Phytase Activity of Tobacco (Nicotiana tabacum) is Affected by the Availability of Soluble Phytate

Phytate, the major organic phosphorus in soil, is not readily available to plants as a source of phosphorus (P). It is either complexed with cations or adsorbed to various soil components. The present study was carried out to investigate the extracellular phytase activities of tobacco (Nicotiana tabacum variety GeXin No.1) and its ability to assimilate external phytate-P. Whereas phytase activities in roots, shoots and growth media of P_i-fed 14-day-old seedlings were only 1.3–4.9% of total acid phosphatase (APase) activities, P starvation triggered an increase in phytase secretion up to 914.9 mU mg⁻¹ protein, equivalent to 18.2% of total APase activities. Much of the extracellular phytase activities were found to be root-associated than root-released. The plants were not able to utilize phytate adsorbed to sand, except when insoluble phytate salts were preformed with Mg²⁺ and Ca²⁺ ions for supplementation. Tobacco grew better in sand supplemented with Mg-phytate salts (31.9 mg dry weight plant⁻¹; 0.68% w/w P concentration) than that with Ca-phytate salts (9.5 mg plant⁻¹; 0.42%), presumably due to its higher solubility. We conclude that insolubility of soil phytate is the major constrain for its assimilation. Improving solubility of soil phytate, for example, by enhancement of citrate secretion, may be a feasible approach to improve soil phytate assimilation.     

Effect of phytase supplementation on phosphorus digestibility in low-phytate barley fed to finishing pigs

Forty crossbred barrows (Camborough 15 Line female x Canabred sire) weighing an average of 79.6 +/- 8.0 kg were used in a factorial design experiment (5 barleys x 2 enzyme levels) conducted to determine the effects of phytase supplementation on nutrient digestibility in low-phytate barleys fed to finishing pigs. The pigs were assigned to one of 10 dietary treatments comprised of a normal 2-rowed, hulled variety of barley (CDC Fleet, 0.26% phytate) or 2 low-phytate hulled genotypes designated as LP422 (0.14% phytate) and LP635 (0.09% phytate). A normal, hulless barley (CDC Dawn, 0.26% phytate) and a hulless genotype designated as LP422H (0.14% phytate) were also included. All barleys were fed with and without phytase (Natuphos 5000 FTU/kg). The diets fed contained 98% barley, 0.5% vitamin premix, 0.5% trace mineral premix, 0.5% NaCl and 0.5% chromic oxide but no supplemental phosphorus. The marked feed was provided for a 7-day acclimatization period, followed by a 3-day faecal collection. In the absence of phytase, phosphorus digestibility increased substantially (P < 0.05) as the level of phytate in the barley declined. For the hulled varieties, phosphorus digestibility increased from 12.9% for the normal barley (0.26% phytate) to 35.3 and 39.8% for the two low-phytate genotypes (0.14 and 0.09% phytate respectively). For the hulless varieties, phosphorus digestibility increased from 9.2% for the normal barley (0.26% phytate) to 34.7% for the hulless variety with 54% of the normal level of phytate (0.14% phytate). In contrast, when phytase was added to the diet, there was little difference in phosphorus digestibility between pigs fed normal barley and those fed the low-phytate genotypes (significant barley x enzyme interaction, P = 0.01). For the hulled varieties, phosphorus digestibility was 50.1% for the barley with the normal level of phytate (0.26% phytate) compared with 51.1 and 52.4% for the varieties with 54 and 35% of the normal level of phytate (0.14 and 0.09% phytate respectively). For the hulless varieties, phosphorus digestibility increased from 47.1% for the normal barley (0.26% phytate) to 54.4% for the hulless variety with 54% of the normal level of phytate (0.14% phytate). In conclusion, both supplementation with phytase and selection for low-phytate genotypes of barley were successful in increasing the digestibility of phosphorus for pigs. Unfortunately, the effects did not appear to be additive. Whether or not swine producers will choose low-phytate barley or supplementation with phytase as a means to improve phosphorus utilization, will likely depend on the yield potential of low-phytate barley and the additional costs associated with supplementation with phytase.     

Utilisation of phytate phosphorus by rumen bacteria in a semi-continuous culture system (Rusitec) in lactating goats fed on different forage to concentrate ratios

Experimental data on phytate phosphorus utilisation by ruminants are scarce. The aim of this study was to estimate the phytase activity of rumen micro-organisms when phytate phosphorus supply is high. A semi-continuous culture system fermentor (RUSITEC) was used. The inoculum was obtained from eight goats fed on either high or low forage level diets. Experimental buffers only differed by the nature of phosphorus monosodium phosphate vs. corn sodium phytate. The nylon bags containing 15 g DM of substrate were removed after a 48-hour incubation period. The system was maintained for 15 days: 5 days for adaptation, in order to obtain a steady state, and 10 days for sampling and recording. No significant differences were observed for DM digestibility, gas production, pH, N-NH3, and SCFA for the different treatments. Bacterial efficiency of phytate phosphorus utilisation was significantly higher (p < 0.001) with organic P, but remained lower than the data usually reported in the literature. These results may be explained by the relative saturation of bacterial phytase activity when the buffer contains a high level of phytate phosphorus.     

Effect of phytase supplementation on phosphorus digestibility in low-phytate barley fed to finishing pigs

Forty crossbred barrows (Camborough 15 Line female×Canabred sire) weighing an average of 79.6±8.0?kg were used in a factorial design experiment (5 barleys×2 enzyme levels) conducted to determine the effects of phytase supplementation on nutrient digestibility in low-phytate barleys fed to finishing pigs. The pigs were assigned to one of 10 dietary treatments comprised of a normal 2-rowed, hulled variety of barley (CDC Fleet, 0.26% phytate) or 2 low-phytate hulled genotypes designated as LP422 (0.14% phytate) and LP635 (0.09% phytate). A normal, hulless barley (CDC Dawn, 0.26% phytate) and a hulless genotype designated as LP422H (0.14% phytate) were also included. All barleys were fed with and without phytase (Natuphos 5000 FTU/kg). The diets fed contained 98% barley, 0.5% vitamin premix, 0.5% trace mineral premix, 0.5% NaCl and 0.5% chromic oxide but no supplemental phosphorus. The marked feed was provided for a 7-day acclimatization period, followed by a 3-day faecal collection. In the absence of phytase, phosphorus digestibility increased substantially (P<0.05) as the level of phytate in the barley declined. For the hulled varieties, phosphorus digestibility increased from 12.9% for the normal barley (0.26% phytate) to 35.3 and 39.8% for the two low-phytate genotypes (0.14 and 0.09% phytate respectively). For the hulless varieties, phosphorus digestibility increased from 9.2% for the normal barley (0.26% phytate) to 34.7% for the hulless variety with 54% of the normal level of phytate (0.14% phytate). In contrast, when phytase was added to the diet, there was little difference in phosphorus digestibility between pigs fed normal barley and those fed the low-phytate genotypes (significant barley×enzyme interaction, P=0.01). For the hulled varieties, phosphorus digestibility was 50.1% for the barley with the normal level of phytate (0.26% phytate) compared with 51.1 and 52.4% for the varieties with 54 and 35% of the normal level of phytate (0.14 and 0.09% phytate respectively). For the hulless varieties, phosphorus digestibility increased from 47.1% for the normal barley (0.26% phytate) to 54.4% for the hulless variety with 54% of the normal level of phytate (0.14% phytate). In conclusion, both supplementation with phytase and selection for low-phytate genotypes of barley were successful in increasing the digestibility of phosphorus for pigs. Unfortunately, the effects did not appear to be additive. Whether or not swine producers will choose low-phytate barley or supplementation with phytase as a means to improve phosphorus utilization, will likely depend on the yield potential of low-phytate barley and the additional costs associated with supplementation with phytase.     

菌根真菌磷酸酶活性对红三叶草生境中土壤有机磷亏缺的影响

利用PVC分室培养装置研究了菌根际和菌丝际磷酸酶活性变化与土壤有机磷亏缺间的关系,结果表明,施用有机磷（植酸钠）能促进菌根根系侵染、提高土壤磷酸酶尤其是酸性磷酸酶的活性,使菌丝际范围变宽。菌丝际的存在使土壤有机磷亏缺范围加大,与非菌根植物相比,由于菌根真菌的作用,植物能更容易地从有机磷中获得磷营养以满足植物生长的需要,从而使其干物重和磷吸收量更高。     

Distribution of phosphorus compounds in corn processing

Distillers dried grains with solubles (DDGS) and corn gluten feed (CGF) are major coproducts of ethanol production from corn dry grind and wet milling facilities, respectively. These coproducts contain important nutrients, nevertheless, high levels of phosphorus (P). About 50-80% of the P in these products is in an organically bound form known as phytate. The phytate P in these products cannot be digested by nonruminant animals. Consequently, large quantities of phytate are deposited into the soil with the animal wastes which potentially could cause P pollution in soil and underground water resources. As regulations on the concentration of P material in ethanol production coproducts become more restrictive, measures need to be taken for effective extraction of phytate P from the coproducts to make these processes more environmentally compatible. Proper marketing of coproducts is critical to the overall economy of ethanol production facilities. In this study, distribution of P compounds in different streams of dry grind and wet milling operations was determined. In the dry grind process, the highest P concentration was found to be in the condensed distillers solubles (CDS) at about 1.34 wt.% (db). About 59% of P in this stream was in phosphates form. The highest concentration of P in the wet milling process was found in the light steep water at about 3.4 wt.% (db). In this stream, about 22% of P was attributed to phosphates.     

Influence on plant growth of the breakdown of organic phosphorus compounds by micro-organisms

Summary Micro-organisms which break down lecithin or phytin have been isolated from soil by enrichment cultures. Only those organisms were investigated which on an agar medium containing lecithin or phytin as the sole source of P (in the case of lecithin, also as a source of N and energy), produced a clear area around their colonies. Certain of these organisms were inoculated into sterile cultures of radish plants grown on sand or agar substrates containing lecithin or phytin as the source of P, and the growth and N- and P-uptakes of the plants were compared with those of similar plants grown in non-sterile sand cultures or supplied with KH₂PO₄. Evidence was obtained that both lecithin and phytin can serve as P-sources for higher plants even under sterile conditions, phytin producing a greater effect than lecithin. Inoculation, however, had only in the phytin-containing medium sometimes a slight effect on the P-nutrition of the plants and further work in this field is therefore necessary.     

Phytase activity in tobacco (Nicotiana tabacum) root exudates is exhibited by a purple acid phosphatase

Phytases are enzymes that catalyze liberation of inorganic phosphates from phytate, the major organic phosphorus in soil. Tobacco (Nicotiana tabacum) responds to phosphorus starvation with an increase in extracellular phytase activity. By a three-step purification scheme, a phosphatase with phytase activity was purified 486-fold from tobacco root exudates to a specific activity of 6,028 nkat mg(-1) and an overall yield of 3%. SDS-PAGE revealed a single polypeptide of 64 kDa, thus indicating apparent homogeneity of the final enzyme preparation. Gel filtration chromatography suggested that the enzyme was a ca. 56 kDa monomeric protein. De novo sequencing by tandem mass spectrometry resulted in a tryptic peptide sequence that shares high homology with several plant purple acid phosphatases. The identity of the enzyme was further confirmed by molybdate-inhibition assay and cDNA cloning. The purified enzyme exhibited pH and temperature optima at 5.0-5.5 and 45 degrees C, respectively, and were found to have high affinities for both p-nitrophenyl phosphate (pNPP; K(m)=13.9 microM) and phytate (K(m)=14.7 microM), but a higher kcat for pNPP (2,056 s(-1)) than phytate (908 s(-1)). Although a broad specificity of the enzyme was observed for a range of physiological substrates in soil, maximum activity was achieved using mononucleotides as substrates. We conclude that the phytase activity in tobacco root exudates is exhibited by a purple acid phosphatase and its catalytic properties are pertinent to its role in mobilizing organic P in soil.     

不同磷源对磷高效利用野生大麦根际土壤磷组分的影响

在土培盆栽条件下,以野生大麦磷高效利用基因型IS-22-30、IS-22-25和低效基因型IS-07-07为材料,研究不施磷(CK)、无机磷(KH₂PO₄,Pi)、有机磷(phytate,Po)及二者混合(KH₂PO₄+phytate,Pi+Po)的方式施磷30 mg·kg^-1时,磷高效基因型野生大麦对磷素吸收利用能力及土壤磷组分特征.结果表明： Pi处理野生大麦干物质量和磷素积累量最大,Pi+Po处理其次,Po处理最小,均显著高于CK处理,且磷高效基因型物质生产和磷素吸收能力显著高于磷低效基因型.土壤有效磷在不同磷源处理间差异显著,Pi处理时含量最高,Pi+Po处理次之,且磷高效基因型野生大麦根际有效磷含量显著高于磷低效基因型.磷高效基因型野生大麦根际有效磷呈现亏缺现象,在Pi和Pi+Po处理时亏缺程度较大.根际与非根际土壤无机磷组分含量为Ca₁₀-P>O-P>Fe-P>Al-P>Ca₂-P>Ca₈-P,且其含量随着Pi的增加而增加.各磷源处理下,磷高效基因型野生大麦根际土壤Ca₂-P、Ca₈-P出现亏缺;Pi处理磷高效基因型野生大麦根际土壤Al-P、Fe-P出现富集.土壤中有机磷各组分含量为中活性有机磷>中稳性有机磷、高稳性有机磷>活性有机磷.野生大麦根际土壤活性有机磷和中活性有机磷呈现富集,其富集量在Pi处理时最大;中稳性有机磷和高稳性有机磷呈现亏缺.各磷源处理下,磷高效基因型野生大麦根际土壤活性有机磷含量显著高于磷低效基因型,中稳性有机磷和高稳性有机磷在基因型间差异不显著.Pi缺乏时,磷高效基因型野生大麦活化吸收Ca₂-P、Ca₈-P、Al-P和活性有机磷的能力较强.     

Phytin is synthesized in the cotyledons of germinated castor-bean seeds in response to exogenously supplied phosphate

Following germination of the castor bean (Ricinus communis L.) seed, levels of phytin decline in both the endosperm and the embryo. However, as seedling growth continues, phytin increase in the latter to a level exceeding that present in the mature dry embryo, while phytin declines concomitantly in the endosperm. It is likely that phosphate mobilized from phytin in the endosperm acts as a substrate for phytin synthesis in the embryo. This is supported by the observation that isolated embryos supplied with phosphate accumulate phytin, particularly in the cotyledons. This increase is enhanced whenmyo-inositol is provided concurrently as a carbon source. Phytin synthesis in the cotyledons of the isolated embryos can occur without the attached axis. Whether initially exposed to exogenous phosphate or not, the isolated cotyledons remain competent in their ability to synthesize phytin for an extended post-germinative period, even though the major reserves are being mobilized at this time.