Phosphate,Fe and Mn uptake of N2 fixing red clover and ryegrass from an Oxisol as affected by P and model humic substances application. 1. Plant parameters and soil solution composition

The effect of humic substances on P-availability in soil is still debated. Therefore, the effect of model humic substances synthesized from hydroquinone on P, Fe, and Al solubility in a strong P fixing Oxisol and on P acquisition by red clover and ryegrass was investigated. After 4 months of incubation, P concentration of soil solution had increased by a factor of > 10 at the highest humic level (50 g humics kg^?1 soil), accompanied by a similar increase in Fe and Al concentrations. Soil samples with 0, 10, 30, 50 g humics kg^?1 soil were planted with red clover and ryegrass. Red clover showed a small increase of shoot yield and a moderate increase of P uptake after humics addition. High humics levels increased slightly Fe concentration in the shoots but strongly that of Mn leading to Mn toxicity. Ryegrass showed a strong increase in shoot yield after humics addition of about 150 % at the highest humics level compared to the control without humics. At each humic level, P application (100 mg kg^?1 soil) had no effect on P uptake of red clover and a small effect on P uptake by ryegrass. The relatively small effect of humics and P application on shoot yield of clover compared to grass can be explained by chemical P mobilization of red clover via exudation of citrate (about 12 μmol citrate g^?1 soil). This agrees with the finding that P solubility increased in the soil under red clover but not under ryegrass from the first to the second harvest, indicating that red clover mobilized P.     

Phosphate aquisition by red clover and black mustard on a humic podzol

Recent investigations have shown that phosphate (P) mobilization by root exudates is an important feature of genotypes to acquire P even in soils of low‐P availability. We, therefore, investigated P mobilization processes in the rhizosphere of red clover (Trifolium pratense L.) and black mustard (Brassica nigra L.) on a humic podzol. As measured by the Kuchenbuch‐Jungk method (Kuchenbuch and Jungk, 1982), both species accumulated similar quantities of citrate (12 μmol/g soil) in the rhizosphere in about 1 mm distance from the soil‐root interface. Despite of similar concentrations of P‐mobilizing citrate in the rhizosphere of both species, red clover took up nearly the two‐fold of P compared to black mustard. Differences in rhizosphere pH were determined between both species. Black mustard did not acidify the rhizosphere, whereas red clover decreased the pH in the rhizosphere from 5.8 to about 4.0 (in 0.01M CaCl₂). The simultaneous acidification and excretion of citrate compared to citrate excretion alone had consequences for P mobilization processes in the rhizosphere. Phosphate mobilization from the soil solid phase was higher at higher pH. Thus, the citrate‐induced P desorption was not the limiting step in P acquisition by red clover and black mustard. Calculations of P distribution in the soil solution between free ortho‐P and humic‐associated P showed that at higher pH most of the P was associated with dissolved humic substances, whereas at pH < 5, most of the P was present as free ortho‐P. These P species can readily be taken up by the roots whereas humic‐associated P must probably be desorbed from the humic surface before uptake. Phosphate species calculations, therefore, explained the higher P uptake of red clover compared to black mustard. Aluminum species distribution calculations in the soil solution further show that even at pH < 5.0 in the soil solution, citrate strongly complex Al and thereby reduce the activity of monomeric Al species. The excretion of citrate can, therefore, counteract the root induced acidification of the rhizosphere with respect to Al toxicity.     

Aluminum and iron(III) species in the soil solution including organic complexes with citrate and humic substances

The solubility of Al and Fe in soil is of relevance for their toxicity and availability, respectively, to plant roots. Humic substances as the main part of stable soil organic matter and citrate which is often excreted by P deficient plants are strong complexants of Al and Fe(III). Therefore, equations were developed to calculate the Al and Fe(III) species distribution in the soil solution in the presence of humic substances and citrate as organic ligands. Calculations in the pH range 4.0–7.0 showed that at higher pH humic-Al complexes were the most important species whereas AlOH-citrate^? dominated between pH 4.0 and 5.4. Free monomeric Al and AlSO₄⁺ were of minor relevance. Iron(III) species calculations showed that humic-Fe complexes were the main species in the pH range 4.0–7.0. But if mugineic acid, a Fe complexing phytosiderophore released into the rhizosphere by graminaceous plant species, was present in the soil solution (10^?6 M), Fe-mugineic acid complexes accounted for most of the Fe in solution. Fe-citrate^? was relevant at lower pH but contributed little to Fe(III) species at pH > 6.0. The results demonstrate the strong importance of the considered organic ligands for Fe and Al in the soil solution.     

Phosphate,aluminium and iron in the soil solution of three different soils in relation to varying concentrations of citric acid

Several plant species excrete di- and tricarboxylic acids as a reaction to P and Fe deficiency. Therefore, the effect of citric acid on P, Al, and Fe soil solution concentrations of three soils was investigated in an incubation experiment. Citric acid additions of 10–50 μmole/g soil increased P, Al, and Fe solubility. The effect was strongly pronounced in the podzol with humic-P complexes as the main P fraction. The maximum increase was about 80 μmole P/L in the luvisol, 500 μmole P/L oxisol, and 3000 μmole P/L in the podzol. The Fe and Al solubility increased parallel to that of P with a ratio of Fe+Al_solubized/P_solubized of 10 or more. The solubization effect by citric acid was strong even after 90 days. Possible solubization reactions are discussed.     

Productivity parameters and soil health dynamics under long-term 2-year potato rotations in Atlantic Canada

Crop rotation effects, due to emerging soil ecological interactions and processes that occur with time, can have a major impact on crop productivity and soil health. 2-year potato (Solanum tuberosum L.) rotation studies were conducted during an 11-year period on a fine sandy loam (Orthic Podzol) under a cool, humid climate in Prince Edward Island in eastern Canada. Rotation partners included Italian ryegrass (Lolium multiflorum Lam.), red clover (Trifolium pratense L.), and barley (Hordeum vulgare L.). The study objective was to assess trends in yield parameters, populations and diversity in nematodes and soil-borne pathogens, and soil structural stability and organic matter retention with time. Italian ryegrass had the largest biomass of both herbage (6.5 Mg ha⁻¹) and roots (4.8 Mg ha⁻¹), compared to red clover or barley. Averaged across 9 years, potato tuber yield was highest under Italian ryegrass (35.0 Mg ha⁻¹) and lowest under red clover (31.1 Mg ha⁻¹) rotations. Differences in tuber yield among rotations was associated with soil N limitation and nematode activity, but was not evident until Year 6. Population densities of specific plant parasitic nematodes recovered from roots and soil were generally low and varied according to host plant. Root lesion (Pratylenchus spp.) and root-knot (Meloidogyne spp.) nematodes were more prevalent under red clover compared to other rotational crops. Stunt (Merlinius spp. and Tylenchorhynchus spp.) nematodes were more common under Italian ryegrass. The main organisms associated with tuber-borne disease were Rhizoctonia solani, Streptomyces spp., Fusarium spp., and Helminthosporium solani. The greatest reduction in tuber-borne diseases occurred with the potato–red clover rotation but significant differences were infrequent. Soil structural form throughout the soil profile (to 30 cm depth) was not adversely affected by the rotational treatments during the duration of the study, while soil structural stability at the soil surface (0–10 cm) was significantly improved. Losses of soil organic C, during the 11-year period ranged from marginal (4%) for rotations with Italian ryegrass, to significant (16%) under the barley rotation. Computer simulations, using the Century model, illustrated the importance of C inputs to maintain soil organic matter levels. Overall, potato crop productivity and soil organic C were generally maintained in rotations that contained Italian ryegrass, but declined under rotations with red clover and barley.     

Differences in aluminium tolerance of ryegrass and white clover plants grown from seed or propagated vegetatively

The effect of aluminium (Al) on the relative yield of plants grown from seeds of ryegrass (Lolium perenne L.) or white clover (Trifolium repens L.) and either tillers (ryegrass) or stolon tip cuttings (white clover) were investigated using a low ionic strength (2.7 x 10^‐3 M) solution culture technique. In ryegrass, plants grown from tillers had higher relative yields than plants grown from seedlings in the tops when solution Al concentrations were greater than 16 μM and in the roots when solution Al concentrations were greater than 7 μM. In white clover, relative yields in the tops and roots plants were higher in plants grown from stolon tip cuttings than from seedlings when solution Al concentrations were greater than 10 μM. There were no significant cultivar effects. The results indicate that plants used in Al‐tolerance experiments can be grown from seed or vegetatively propagated, provided solution Al rates are adjusted to reflect differences in Al tolerance.     

Rhizodeposition of nitrogen by red clover,white clover and ryegrass leys

Correct assessment of the rhizodeposition of N in grassland is essential for the evaluation of biological N₂-fixation of legumes, for the total N balance of agro-ecosystems, and for the pre-cropping value of grasslands. Using a leaf-feeding technique by which plants were ¹⁵N labelled while growing in mezotrons in the field, the rhizodeposition of N by unfertilised red clover, white clover and perennial ryegrass growing in pure stands was shown to amount to 64, 71 and 9 g N m⁻², respectively, over two complete growing seasons. The corresponding values for red clover and white clover growing in mixtures with ryegrass were 89 and 32 g N m⁻², respectively. The rhizodeposited N compounds, including fine roots, constituted more than 80% of the total plant-derived N in the soil, and in all cases exceeded the amount of N present in stubble. In the mixtures of red clover–ryegrass and white clover–ryegrass and the pure stands of red clover, white clover and ryegrass, respectively, the rhizodeposition constituted a 1.05, 1.52, 1.26, 2.21 and 2.77 fold increase over the total N in the shoots harvested during the two production years. In pure stands and mixtures of clover, 84 and 92%, respectively, of this N derived from biological N₂ fixation. It is concluded that rhizodeposition provides a very substantial input of N to the legume-based grassland systems with great consequences for ecosystem N balance and turnover. Furthermore, the amount of atmospheric-derived N in the rhizodeposits may exceed that in the harvested shoots.     

Photodegradation of the surfactants Na-dodecylbenzenesulfonate and dodecylpyridinium-chloride as affected by humic substances

Surfactants account for a great proportion of organic xenobiotica released into the environment by anthropogenic action. In natural waters and sewage sludge they can be removed from solution by different reactions, including adsorption to solid phases, biodegradation and abiotic photolytic degradation. In the present investigation we found that photodegradation of Na- dodecylbenzenesulfonate (LAS) was reduced by humic substances by a factor of 2 or more. In contrast, photodegradation of dodecylpyridinium-chloride (LPC) was increased in the presence of humic substances by a factor of 1.16–1.43. These differences were explained by the strong bonding of LPC but not of LAS to the humics. Photolytic degradation of LAS begins at the aliphatic side chains followed by aromatic ring cleavage. Photodegradation of LPC mainly associated with the humics showed similarity to that of pyridine. The experimental data demonstrate strong effects of photodegradation on the persistence of surfactants and the relevance of humic-surfactants associations for photolytic processes.     

土壤腐殖物质组分结合态磷素有效性的研究

以棕色针叶林土、暗棕壤和黄棕壤为供试土壤,采用常规的碱(0.1 mol L-1 NaOH+ 0.1 mol L-1Na4P2O7)提取液酸化法分离出胡敏酸(HA),碱不溶性的粗胡敏素(CHU)再经10％HF-HC1处理得到去除矿质灰分后纯化的胡敏素(PHU);利用酶水解和紫外照射方法,分析了上述各腐殖物质组分结合态磷素...     

Solubilization of Fe (III) from humic-Fe complexes,humic/Fe-oxide mixtures and from poorly ordered Fe-oxide by organic acids - consequences for P adsorption

The effect of organic acids on the solubility of Fe from humic Fe complexes, poorly ordered Fe-oxides and humic/Fe-oxide mixtures was investigated at pH 4 and 7. After reaction, the suspensions were filtered through a 20 000 D membrane and Fe and humic concentrations were determined in the filtrate. Among four organic acids, citric acid had the strongest effect on Fe-solubility and solubilized up to 25% of total Fe from the humic complexes at pH 4. Humic substances were solubilized by citrate too, the proportions being higher at pH 7 than at pH 4. Malic and tartaric acid solubilized substantial but lower amounts of Fe from humic complexes than citric acid, whereas phthalic acid did not solubilize Fe and humics. Citrate added to poorly ordered Fe-oxide and its mixtures with humics, aged for 65 days, increased 20 000 D filtrable Fe, the effect being higher for humic/Fe-oxide complexes. Adding citrate to humic/Fe-oxide mixtures and to Fe-oxide strongly decreased the P sorption, the extent, being higher for the former adsorbent.     

Some structural and electronic features of the interaction of phosphate with metal-humic complexes

Previous studies demonstrated the formation of stable phosphate-metal-humic complexes in solution. These studies, however, indicated that the proportion of complexed metal that intervenes in phosphate fixation is rather low. In this study we investigate the possible structural and electronic features of the binding site involved in phosphate fixation in metal-humic complexes that could explain this fact. To this end, we have studied phosphate-metal-humic complexes involving Fe(III), Al(III), and Zn(II) using three complementary techniques: infrared spectroscopy (FTIR), fluorescence, and molecular modeling. The FTIR study indicated that, in the case of those complexes involving Fe and Zn phosphate, fixation is associated with a stabilization of the metal-carboxylate bond. In the case of Al this effect is less clear. This effect of phosphate fixation on the characteristics of the metal-humic binding site was also supported by the results obtained in the Fluorescence study, which showed significant changes in the quenching effect normally associated with metal complexation in humic substances upon phosphate fixation. Finally, the molecular modeling study revealed that the stability of phosphate-metal-humic complexes is inversely related to the stability of the metal-humic interaction. This result could explain why only a relatively low proportion of humic complexed metal is involved in phosphate fixation.     

Wurzelstudien und Phosphat-Aufnahme von Weidelgras und Rotklee unter Feldbedingungen

Root studies and phosphorus uptake of rye-grass and red clover under field conditions Root parameters (fresh weight, density, surface, length, cation exchange capacity) and phosphate uptake were studied with rye grass and red clover, grown in the field on a brown podsolic soil. In all root parameters, ry grass was superior to red clover. Also, phosphate uptake of rye grass was higher than that of red clover. The greatest difference between both species was found in root length, that of rye grass being about five times longer than that of red clover. Rye grass had longer root hairs than red clover; whereas root diameter of clover was about twice as thick the average rye grass. Significant correlations were observed between root parameters and phosphate uptake in the plants studied. The highest correlation coefficients were obtained for the relationship P-uptake versus root length (clover 0.91***, grass 0.87***) and P-uptake versus root fresh weight (clover 0.92***, grass 0.88***). The phosphate uptake per unit root parameters was significantly higher in red clover, compared with rye grass, for the parameters root fresh weight, cation exchange capacity and root length. Because of this high P-uptake rate for clover it is assumed that clover also requires a higher P-concentration in soil solution as compared with grass. Thus grass may still grow with low P concentrations in the soil solution without P deficiency at which clover cannot grow. It is for this reason that in mixed swards clover is depressed by grass, if the available P in the soil is low.     

Fixation and transfer of nitrogen by white clover to ryegrass

Abstract. ¹⁵N₂ was used in a sealed controlled environment chamber to investigate the transfer of fixed nitrogen from white clover to perennial ryegrass growing in soil in pots. There was no difference in the ¹⁵N content of roots and shoots of clover plants after exposure to ¹⁵N. No labelled fixed nitrogen was detected in ryegrass plants growing with the clover plants for a period of 129 days. There was therefore no evidence of rapid direct transfer (excretion) of fixed nitrogen from clover to ryegrass.     

红三叶草丛枝菌根对有机磷的吸收

The capacities of two arbuscular mycorrhizal (AM) fungi, Glomus mosseae and Glomus versderme, tomineralize added organic P were studied in a sterilized calcareous soil. Mycorrhizal (inoculated with either of the AM fungi) and non-mycorrhizal red clover (Trghlium pmtense L.) plants were grown for eight weeksin pots with upper root, central hyphal and lower soil compartments. The hyphal and soil compartmentsreceived either organic P (as Na-phytate) or inorganic P (as KH₂PO₄) at tbe rate of 50 mg P kg^-1. No P wasadded to the root compartments. Control pots received no added P. Yields were higher in mycorrhizal than innon-mycorrhizal clover. Mycorrhizal inoculation doubled shoot P concentration and more than doubled total P uptake of plaflts in P-amended soil, irrespective of the form of applied P. The mycorrhizal contributionto inorganic P uptake was 80% or 76% in plants inoculated with G. mosseae or G. verefforme, respectively. Corresponding values were 74% and 82% when Na-phytate was applied. In the root compartments of the mycorrhizal treatments, the proportion of root length infected, hyphal length density and phosphatase activity were all higher when organic P was applied than when inorganic P was added.     

The amount,but not the proportion,of N2 fixation and transfers to neighboring plants varies across grassland soils

ABSTRACT

Biological nitrogen fixation (BNF) is an important nitrogen source for both N₂-fixers and their neighboring plants in natural and managed ecosystems. Biological N fixation can vary considerably depending on soil conditions, yet there is a lack of knowledge on the impact of varying soils on the contribution of N from N₂-fixers in mixed swards. In this study, the amount and proportion of BNF from red clover were assessed using three grassland soils. Three soil samples, Hallsworth (HH), Crediton (CN), and Halstow (HW) series, were collected from three grassland sites in Devon, UK. A pot experiment with ¹⁵N natural abundance was conducted to estimate BNF from red clover, and the proportion of N transferred from red clover to the non-N₂ fixing grass in a grass-clover system. The results showed that BNF in red clover sourced from atmosphere in the HH soil was 2.92 mg N plant^?1, which was significantly lower than that of the CN (6.18 mg N plant^?1) and HW (8.01 mg N plant^?1) soils. Nitrogen in grass sourced from BNF via belowground was 0.46 mg N plant^?1 in the HH soil, which was significantly greater than that in CN and HW soils. However, proportionally there were no significant differences in the percentage N content of both red clover and grass sourced from BNF via belowground among soils, at 65%, 67%, 65% and 35%, 27%, 31% in HH, CN, and HW, respectively. Our observations indicate that the amount of BNF by red clover varies among grassland soils, as does the amount of N sourced from BNF that is transferred to neighboring plants, which is linked to biomass production. Proportionally there was no difference among soils in N sourced from BNF in both the red clover plants and transferred to neighboring plants.     

Effects of soil moisture and plant interactions on the soil microbial community structure

A greenhouse pot experiment was conducted to investigate the influence of soil moisture content and plant species on soil microbial community structure using cultivation-independent methods. White clover and ryegrass were grown individually or in a mixture. Plants were subjected to soil moisture content corresponding to 60% field capacity (FC) and 80% FC. Total plant biomass of white clover and ryegrass increased with increasing soil moisture contents. At a given soil moisture content, total biomass of white clover was lower in the ryegrass–clover (RC) mixture compared with those grown individually, while total biomass of ryegrass was higher. Microbial community structure assessed by phospholipid fatty acid analysis (PLFA) was more affected by plant species than soil moisture. Community level physiological profiles (CLPP), in terms of diversity of substrate utilization and average well colour development (AWCD) were affected by plant species and soil moisture. Soil moisture effects were more pronounced in clover than in ryegrass. AWCD and diversity of substrate utilization in the ryegrass–clover mix were similar to those of sole clover while they differed from that of ryegrass suggesting a dominant effect of clover in the mix.     

Mobilization of dissolved organic matter, aluminium and iron in podzol eluvial horizons as affected by formation of metal-organic complexes and interactions with solid soil material

Interactions with dissolved organic matter (DOM) are generally believed to play a crucial role in the translocation of Al and Fe in acid sandy soils. Binding of Al and Fe to DOM affects their mobility in soils by altering sorption equilibria of charged sites on solid soil material, inducing precipitation of organo‐metallic complexes and preventing the formation of inorganic Al and Fe phases. The relative importance of the different processes, especially with respect to the translocation of Al, Fe and organic matter in podzols, remains unresolved. We determined the effect of the presence of solid soil material from the eluvial (AhE and AE, respectively) horizons of a Fimic Anthrosol and a Haplic Podzol on the metal‐to‐organic carbon (M/C) ratio in solution and the formation of dissolved organic Al and Fe complexes. Furthermore, we assessed the resulting influence on the mobilization of Al, Fe and DOM. Even under considerable metal loading, the M/C ratios and ‘free’ metal fractions in solution remained low and relatively constant, due to an apparent buffering by the solid phase and the formation of organo‐metal complexes in solution. The M/C ratios remained so low that significant precipitation of organo‐metal complexes due to saturation with metals was not found. The apparent buffering by the solid phase can be explained by a strong release of organic matter from solid soil material and adsorption of non‐complexed Al and Fe on solid organic matter upon metal addition. Adsorption of organo‐metal complexes most likely played only a minor role. The observations confirm the expected mobilization of Al, Fe and DOM in eluvial horizons and seem to indicate that even under fluctuating input of Al, Fe and DOM the soil solution will have a constant composition with respect to M/C ratios and percentage of Al and Fe present in dissolved organo‐metal complexes.     

Aspects of the phosphorus nutrition of white clover populations II. Root exocellular acid phosphatase activity

The effect of phosphorus (P) level, solution pH, aluminium (Al) level on root exocellular acid phosphatase activity of white clover (Trifolium repens L.) populations was investigated in three experiments. Lotus pedunculatus Cav. cv. Maku was also included in one experiment.

Phosphatase activity increased with decreasing P and increasing Al supply, but was unaffected by the solution pH at which populations had been grown.

There was usually a significant negative correlation between acid phosphatase activity and parameters measuring plant size, for plants grown at adequate P levels, but in only one of the three experiments were there significant differences between white clover populations for phosphatase activity. There was no clear correlation between the acid phosphatase activity of populations and the P status of soil from which they were collected.

We conclude that root exocellular acid phosphatase activity is unlikely to be a useful character in screening for tolerance of low‐P conditions.     

腐殖酸和植物促生芽孢杆菌对黑麦草抗逆性及砒砂岩土壤性状的影响

[目的]探究微生物在砒砂岩土壤修复工程中的作用,为微生物强化植物改良砒砂岩土壤的科学设想提供新思路.[方法]以黑麦草作为供试作物,通过室内盆栽模拟试验的方法,研究接种植物促生芽孢杆菌Bacillus halotolerans P75,施用腐殖酸肥料以及不同的菌肥搭配对水土流失严重的砒砂岩土壤理化性质及黑麦草生长和抗逆性...     

Utilisation by white clover and ryegrass of sulphur from 35s‐Labelled Gypsum

Abstract

This investigation reports the uptake of S from a surface application of ³⁵S‐labelled gypsum by a ryegrass‐white clover mixture sward and by a pure ryegrass stand, each growing at three levels of N in the field. Nitrogen stimulated ryegrass growth, reducing the contribution of white clover to the total yield, whereas S did not influence the yield of either species. Gypsum, while not increasing the total S in the white clover, contributed 23 to 50% of the total S concentration. In contrast, gypsum increased the total S in the ryegrass. The level of N nutrition did not alter the fertiliser S in white clover, but depressed the total S in the ryegrass. Nitrogen enhanced the fertiliser S in ryegrass at the first harvest, however, at the second harvest N depressed the fertiliser S.

Recovery of applied S was increased by N, reaching a maximum value of 19.8% by two harvest, and was decreased with increasing rate of gypsum. Without ‐N the white clover accounted for 50% and 27% of the S recovery by the mixture at the 1st and 2nd harvests respectively, the proportion dropping to less than 20% for each harvest at a high level of N.

There was no apparent competitive advantage of ryegrass over white clover when grown in association although the data indicated a greater ability by ryegrass to absorb S from a surface application. Under conditions of incipient S deficiency the reduction in the contribution of white clover to production with increasing N supply was considered to be due to factors other than the availability of S in the environment.